June 1 - 3 , 1978 - University of Hawaii at Manoa

COOPERATIVE NATIONAL PARK RESOURCES STUDIES UNIT 

UNIVERSITY OF HAWAII AT MANOA 

Department of Botany 

Honolulu, Hawaii 96822 

(808) 948-8218 

Clifford W. Smith, Unit Director 

Associate Professor of Botany 

PROCEEDINGS 

SECOND CONFERENCE IN NATURAL SCIENCES 

HAWAII VOLCANOES NATIONAL PARK 

held at 

Hawaii Field Research Center 

Hawaii Volcanoes National Park 

on 

June 1 - 3, 1978 

Edited by 

C. W. Smith, Director, CPSU/UH

PREFACE 

The Second Conference in Natural Sciences was held from 

June 1-3, 1978, at the Hawaii Field Research Center in Hawaii 

Volcanoes National Park. The Research Center has expanded quite 

dramatically since the first Conference in 1976. First the Mauna 

Loa Field Station was moved into the area at which time the U. S. 

Fish and Wildlife Service erected a separate laboratory and 

office building. Shortly thereafter the U. S. Forest Service 

also built their own facility. In the meantime, the large green- 

house was erected and the Avian Disease Laboratory was set up in 

one of the old YCC buildings. The dormitory building opposite 

Magma House was converted into offices and laboratories while 

some small dormitories were retained and renovated. The her- 

barium is located in this building. The facilities, probably the 

best field station in Hawai'i, are available to the public. 

Dormitory fees are $4.00 a night and should be booked in advance 

by writing to the Research Biologist, Hawaii Volcanoes National 

Park, Hawaii 96718. 

During the Conference a questionnaire was distributed con- 

cerning future meetings. There was a unanimous vote to continue 

the conferences. Fifty-two percent of the respondents preferred 

annual meetings, 40% biennual meetings with the remainder 

undecided about the frequency. Sixty percent of the participants 

prefer June meetings, 30% had no opinion, and 10% liked August. 

Three-quarters of those attending felt that three days was suffi- 

cient, the remainder had a number of other options. Concerning 

the length of papers, half the respondents felt 15 minutes was 

adequate, 25% thought 10 minutes were enough, the remainder 

wanted longer presentations. Three-quarters of those responding 

felt 5 minutes was sufficient time for questions. Fifty-five 

percent of the participants would prefer topic-oriented sessions, 

30% preferred contributed papers. 

In the comments and suggestion section numerous complaints 

about the Conference room were aired. Other common suggestions 

included restricting each participant to a single paper, broad- 

ening the scope of disciplines able to participate, and sending 

out the abstracts and time table before the Conference. 

We will endeavor to correct all the deficiencies noted. We 

appreciate the time and effort put into your responses which we 

think illustrates your interest in these meetings. 

C. W. Smith 

Director, CPSU/UH

PREFACE 

TABLE OF CONTENTS 

Apple, Russell A. 

THE WHITNEY LABORATORY OF SEISMOLOGY (1912-1963) 

Baker, James K., and Melinda S. Allen 

ROOF RAT DEPREDATIONS ON HIBISCADELPHUS (MALVACEAE) 

TREES 

Banko, Paul C. 

NENE REINTRODUCTION PROGRAM AND RESEARCH 

IN HAWAIIAN NATIONAL PARKS 

Banko, Winston E. 

SOME LIMITING FACTORS AND RESEARCH NEEDS 

OF ENDANGERED HAWAIIAN FOREST BIRDS 

Beardsley, John W., Jr . 

BIOLOGICAL CONTROL OF WILDLAND WEED PESTS IN HAWAII-- 

IS IT A FEASIBLE SOLUTION? 2 6 

Beardsley, J. W., R. Burkhart, M. L. Goff, 

A. Hara, and G. Teves 

HALEAKALA NATIONAL PARK CRATER DISTRICT 

RESOURCES BASIC INVENTORY: 

INSECTS AND OTHER TERRESTRIAL ARTHROPODS 

Bridges, Kent W. 

HAWAII IBP SYNTHESIS: 5. SHORT-TERM TEMPORAL PATTERNS 

AMONG ISLAND BIOTA 34 

Callaham, Robert Z. 

SECOND CONFERENCE ON NATURAL RESOURCES 

REMARKS BY ROGER SKOLMEN 

AT DEDICATION OF THE HAWAII FIELD RESEARCH CENTER 

ON BEHALF OF STATION DIRECTOR CALLAHAM 

Carr, Gerald D. 

HYBRIDIZATION IN THE HAWAIIAN SILVERSWORD COMPLEX 

Carson, Hampton L. 

HAWAII IBP SYNTHESIS: 6. GENETIC VARIATION 

AND POPULATION STRUCTURE IN ISLAND SPECIES 

Chan, John G. 

SOME ASPECTS OF A SHELL DISEASE IN THE HAWAIIAN 

FRESHWATER SHRIMP, BISULCATA (RANDALL)

iii 

Clarke, Garvin 

THE DISTRIBUTION OF MYRICA FAYA AND OTHER SELECTED 

PROBLEM EXOTICS WITHIN HAWAII VOLCANOES NATIONAL PARK 51 

Collins, Mark S., and Robert J. Shallenberger 

FOREST BIRD POPULATIONS ON O'AHU 

Conant, Patrick 

LEK BEHAVIOR AND ECOLOGY OF TWO HOMOSEQUENTIAL - 

SYMPATRIC HAWAIIAN DROSOPHILA : 

DROSOPHILA HETERONEURA AND DROSOPHILA SILVESTRIS 

Conant, Sheila 

HAWAII IBP SYNTHESIS: 3. THE KILAUEA RAIN FOREST 

ECOSYSTEM 58 

Conant, Sheila 

BIRDS OF THE KALAPANA EXTENSION 

Conant, Sheila, and Maile Stemmermann 



BIRDS OF THE CRATER DISTRICT 

Corn, Carolyn A. 

EXPERIMENTAL HYBRIDIZATIONS IN HAWAIIAN METROSIDEROS 7 7 

Croft, Lisa K., and Paul K. Higashino 

THE RARE AND THREATENED PLANTS IN THE AHUPUA'A OF 

MANUKA, KAULANAMAUNA, AND KAPU'A, SOUTH KONA, HAWAI'I 8 6 

Davis, Bertell D. 

HUMAN SETTLEMENT AND ENVIRONMENTAL CHANGE 

AT BARBERS POINT, O'AHU 

Davis, C. J. 

ESTABLISHMENT OF SOME RECENT IMMIGRANT INSECTS 

IN HAWAII VOLCANOES NATIONAL PARK 98 

Evenson, William E. 

A MATHEMATICAL MODEL OF 'OHI'A DIEBACK 

AS A NATURAL PHENOMENON 

Gardner, Donald E. 

EVALUATION OF A NEW TECHNIQUE FOR HERBICIDAL TREATMENT 

OF MYRICA FAYA TREES 114 

Gerrish., Grant 

FACTORS CONTROLLING THE DISTRIBUTION OF EXOTIC PLANTS 

IN THE KO'OLAU MOUNTAINS, O'AHU 120 

Goff, M. Lee 

RESOURCE TRACKING PATTERNS IN ACARI ASSOCIATED WITH 

BIRDS IN HAWAII VOLCANOES NATIONAL PARK: 

A PRELIMINARY REPORT 125

Gon, Samuel M., 111 

ALTITUDINAL EFFECTS ON THE GENERAL DIVERSITY OF ENDEMIC 

INSECT COMMUNITIES IN A LEEWARD HAWAIIAN FOREST SYSTEM, 

MANUKA FOREST RESERVE, SOUTH KONA, HAWAI'I 134 

Hoe, William J. 



MOSSES OF THE CRATER DISTRICT 

Howarth, F. G. 

HAWAII IBP SYNTHESIS: 4. THE HAWAIIAN LAVA TUBE 

ECOSYSTEM 

Jacob i , James D. 

DESCRIPTION OF A NEW LARGE-SCALE VEGETATION MAPPING 

PROJECT IN HAWAI'I 

Kaschko, Michael W., and Melinda S. Allen 

THE IMPACT OF THE SWEET POTATO ON PREHISTORIC 

HAWAIIAN CULTURAL DEVELOPMENT 

Kilgore, Bruce M. 

DEDICATION ADDRESS FOR HAWAII FIELD RESEARCH CENTER 184 

Kjargaard, John I. 

THE STATUS OF THE HAWAIIAN DARK-RUMPED PETREL 

AT HALEAKALA 193 

Lamoureux , C harles H. 

HAWAII IBP SYNTHESIS: 7. IMPACT OF EXOTIC PLANTS 

AND ANIMALS IN HAWAI'I 198 

Lee, Ah Fat 

POHAKULOA PROPAGATION PROJECT: 

A CONTINUING SUCCESS STORY 

Lee, Barbara 

STUDIES IN THE LIFE HISTORY OF THE 'ALALA IN CAPTIVITY 207 

Merlin, Mark David 

HUMAN PERCEPTION OF THE HAWAIIAN ENDANGERED SPECIES: 

A PRELIMINARY REPORT ON A THREE-YEAR RANDOM SURVEY 208 

Miller, John M., and Alan M. Yoshinaga 

ACID RAIN IN HAWAI'I 

Moore, Richard B., Daniel Dzurisin, Gordon P. Eaton, 

Robert Y. Koyanagi, Peter W. Lipman, John P. Lockwood, 

Gary S. Puniwai, and Rosalind Tuthill Helz 

THE SEPTEMBER 1977 ERUPTION OF KILAUEA VOLCANO, HAWAI'I 218 

Mueller-Dombois, Dieter 

HAWAII IBP SYNTHESIS : 1. BRIEF INTRODUCTORY SURVEY 219

Mueller-Dombois, Dieter 

HAWAII IBP SYNTHESIS: 2. THE MAUNA LOA TRANSECT 

ANALYSIS 

Mueller-Dombois, ~ieter' 

HAWAII IBP SYNTHESIS: 8. ISLAND ECOSYSTEMS: 

WHAT IS UNIQUE ABOUT THEIR ECOLOGY? 

Mull, William P. 

VARIABILITY IN DORSAL PATTERNING AMONG POPULATIONS OF 

HAWAIIAN "HAPPY-FACE" SPIDERS (THERIDION SP. OR SPP.) 

ON THE BIG ISLAND 

..-. ~ 

Pam. Richard P. ~ 

THE ROLE OF THE HAWAIIAN TWO-LINED 'OHI'A BORER, 

PLAGITHMYSUS BILINEATUS SHARP, IN THE DECLINE OF 

'OHI'A-LEHUA FORESTS ON THE ISLAND OF HAWAI'I 

Radovsky, Frank J., JoAnn M. Tenorio, P. Quentin Tomich, 

and James D. Jacobi 

ACARI ON MURINE RODENTS ON MAUNA LOA, HAWAI'I 

Ralph, C. John 

HABITAT UTILIZATION AND NICHE COMPONENTS IN SOME 

HAWAIIAN ENDANGERED FOREST BIRDS 

Reeser, Don 

PLANTING, A TOOL FOR NATIVE ECOSYSTEM RESTORATION 

Scott, J. Michael 

FOREST BIRD SURVEY OF THE HAWAIIAN ISLANDS 

Scowcroft, Paul G. 

DIRECT SOWING OF TREATED MAMANE SEEDS: 

AN INEFFECTIVE REGENERATION TECHNIQUE 

Shimoda, Jerry Y. 

COMMUNICATIONS TECHNIQUES AND THE SCIENTIST 

Skolmen, Roger G. 

VEGETATIVE PROPAGATION OF ACACIA KOA GRAY 

Smathers, Garrett A., and Donald E. Gardner 

STAND ANALYSIS OF AN INVADING FIRETREE (MYRICA - FAYA) 

POPULATION, HAWAI ' I 

Smith, Clifford W. 



INTRODUCTION AND GENERAL OVERVIEW 

Smith, Clifford W. 


RESOURCES BASIC INVENTORY: THE LICHEN FLORA

Stemmermann, Lani 



THE VASCULAR FLORA OF HALEAKALA 

Tabor, Kim0 

THE ACQUISITION OF NATURAL AREAS IN HAWAI'I 

Tomich, P. Quentin 

STUDIES OF LEPTOSPIROSIS IN NATURAL HOST POPULATIONS: 

I. SMALL MAMMALS OF WAIPI'O VALLEY, ISLAND OF HAWAI'I 308 

van Riper, Charles, 111, and Sandra Guest van Riper 

A NECROPSY PROCEDURE FOR SAMPLING DISEASE 

IN WILD BIRD POPULATIONS 309 

Walters, Gerald A. 

BRINGING BACK THE MONARCH OF HAWAIIAN FORESTS-- 

ACACIA KOA 

- 

Whiteaker, Louis D. 



VEGETATION MAP OF THE CRATER DISTRICT 

Wolfe, Claire M., C. John Ralph, and Paul K. Higashino 

FOREST BIRD POPULATION VARIATION AS RELATED TO 

HABITAT TYPES 345 

Yoshinaga, Alvin Y. 

VEGETATION OF THE HANA RAIN FOREST, 

HALEAKALA NATIONAL PARK 

Ziegler, Alan C. 

PREHISTORIC HAWAIIAN BIRDS 

LIST OF PARTICIPANTS 350 

SUBJECT INDEX 354 

333

THE WHITNEY LABORATORY OF SEISMOLOGY (1912-1963)* 

Russell A. Apple 

State Director's Office 

National Park Service 


The Whitney vault was built in 1912 on the rim of Kilauea 

crater to begin the resident study of the volcanic and seismic 

activity of Kilauea and Mauna Loa volcanoes. In 1912 and 1913, 

standard seismometers were imported from Japan and Germany. By 

1928, these instruments had been modified and new ones designed 

and built by the Hawaiian Volcano Observatory to deal with three 

local volcanic phenomena: near quick-period earthquakes, har- 

monic tremor, and ground tilting. Hawaiian-type seismometers, 

based on designs evolved in the Whitney laboratory, were manu- 

factured in the Observatory's machine shop and installed in a 

network of stations on the Big Island of Hawai'i. Hawaiian-type 

seismometers and trained personnel were also sent to Lassen 

volcano in California and to the Aleutian Islands to institute 

seismic studies in these regions. 

The structural history of the vault and its seismometers and 

other instruments are given. Abandonment of vault and instru- 

ments came in 1963. By this time, its mechanical seismometers 

were technologically obsolete and replaced by electronic instru- 

ments whose ground-movement magnification capabilities were 

hundreds of thousands of times greater. The Whitney laboratory 

and its instrumentation ca. 1950 are being rehabilitated as a 

historical exhibit by the National Park Service and the U. S. 

Geological Survey's Hawaiian Volcano Observatory. 

* Abstract

ROOF RAT DEPREDATIONS 

ON HIBISCADELPHUS (MALVACEAE) TREES 

James K. Baker and Melinda S. Allen 

Mauna Loa Field Station 


Hawaii 96718 

INTRODUCTION 

The genus Hibiscadelphus Rock (Malvaceae) is endemic to the 

Hawaiian Islands. It is one of the world's rarest (genera) 

groups of trees. Of six described taxa, H. bombycinus Forbes 

(1920), from the island of Hawai'i, azd H. wilderianus Rock 

(lgll), from the island of Maui, are believea to be extinct. 

- H. hualalaiensis Rock (1911), from the island of Hawai'i, and 

H. - distans Bishop and Herbst (1973), from the island of Kaua'i, 

still survive in the wild but both species are few in numbers. 

H. giffardianus Rock (lgll), from the island of Hawai'i, the 

type species for the genus, is extinct in the wild but four trees 

are growing under cultivation in arboreta, and seven others are 

growing in the type locality in Kipuka Puaulu (Bird Park) in 

Hawaii Volcanoes National Park. 

A hybrid, H. X puakuahiwi Baker and Allen (1976, 1977) 

originated in Kipuka Puaulu where its parent species, 

- H. giffardianus and H. hualalaiensis, grow in close proximity. 

The hybrid has been-cultivated widely in arboreta and in private 

gardens around Hawai' i. 

It was during observations on the damage to a number of 

Hibiscadelphus trees by roof rats, Rattus rattus L., that the 

hybrid trees were discovered in 1973. This series of observa- 

tions on rat utilization of bark, buds, flowers, nectar, and seed 

pods followed that initial study. 

Feeding on bark 

Roof rats were sporadically observed feeding on bark on 

three trees in close proximity to one another in Kipuka Puaulu. 

Two of these trees are H. giffardianus and the third a hybrid. 

Bark feeding on trees was first noticed in the late 1960's when a 

number of major limbs were girdled and killed on the oldest of 

the living Hibiscadelphus trees. Efforts were made at that time 

to control the problem by placing rat guards around some of the 

limbs; by poisoning the rats with warfarin; and by catching them 

in snap traps. Warfarin was the most successful control 

technique.

Bark consumption in ensuing years seems to have occurred 

principally during summer dry seasons when it is believed that 

the soft, succulent bark was consumed largely for its moisture 

content. 

Feeding on necta'r 

Nectar feeding was found to occur largely on flowers of 

H. giffardidnus and hybrid trees, prob,ably because of their 

Targer flowers and quantities of nectar. Nectar feeding was not 

observed on any of the smaller blossoms of - H. hualalaiensis and 

- H. distans. 

In order to reach nectar deep within the tubular corollas of 

the larger flowered species, rats chew holes through the base of 

the calyxes. The feeding on flower nectar occurs in all parts of 

the trees, even out on twigs the diameters of match sticks in the 

upper reaches of the canopy, demonstrating the agility of roof 

rats. 

In April and May 1976, we examined 317 flowers from a F1 

hybrid tree for evidence of nectar feeding. A total of 239 (75%) 

were rat damaged. We also examined 1967 flowers in June through 

August of which 538 (27%) were rat damaged. Then in March 

through May 1977, we determined that of 1525 flowers observed 977 

(64%) were damaged. In June, 412 of 590 (69%) flowers examined 

were also damaged. In total, 4399 flowers were examined and 2164 

(49%) were found to have been fed upon by rats. 

Feeding upon buds and flower parts 

Roof rats also eat the staminal column which includes the 

anthers and pollen. They not only eat the stamina1 column of 

mature blossoms, but they will also chew open buds to reach the 

anthers and pollen inside. Most of what we observed of this 

feeding behavior occurred in April and May 1976, when we noticed 

that 68 (21%) out of 317 blossoms had missing staminal columns. 

It appears that rats are after the relatively large amounts 

of pollen present. Individual pollen grains in Hibiscadelphus 

are large, up to 220p, which can be seen easily with the naked 

eye. We smeared and dried quantities of pollen and then stained 

them with ninhydrin producing a deep, purple-colored response 

indicating the presence of relatively large amounts of amino- 

acids (after the staining techniques of Baker and Baker, 1973). 

The nectar also stains a deep purple, more so than nectars of any 

other native Hawaiian flower we have analyzed so far. This may 

suggest that the pollen and nectar of Hibiscadelphus is 

especially nutritious both to rats and bird utilizers.

Feeding upon seed pods 

Immature seeds are normally consumed. Each of the five car- 

pels is opened and the two to five seeds present removed. Only 

the endosperm of the seeds is eaten; the outer husk is discarded. 

In 1976, we placed a 1 rn2 seed catching tray under the canopy of 

a hybrid tree for 30 days and collected the husks of approx- 

imately 150 seeds. A total of 21 uneaten seeds were present, 

obviously dropped by the rats, indicating an 88% destruction of 

the seed crop during this one period of observation. 

There is an obvious'difference in the seed feeding habits 

between two particular rat populations feeding on two different 

but nearby trees. On one tree the empty pods are left dangling 

by their peduncles, indicating the rats feeding in that tree are 

eating the seeds, in situ. On the other tree the rats chew 

through the peduncle and carry the pod down the tree to a nearby 

feeding station. A cache of 136 empty seed pods was found in 

this feeding area in 1976, and another 120 pods were found in 

1977. 

We estimated that the total 256 fruits destroyed represented 

about 90% of the total seed production, and that about 3000 seeds 

were eaten in the two seasons of seed production. 

Also, in 1976, we estimated that there were about 300 empty 

pods in a large cache found near the base of one of the two 

living H. hualalaiensis trees on Mt. Hualalai. It appeared that 

all of-the remaining pods would be taken for a total destruction 

of that year's seed crop. 

SUMMARY 

All of our observations indicate that roof rats cause 

serious damage to Hibiscpdelphus trees by eating bark, buds, 

flowers, nectar, and seeds,'and one has to consider the possi- 

bility that introduction(s) of roof rats into Hawai'i sometime 

around the mid-1800's may have been an additional reason, among 

several, for the present rarity of these trees. Indications are 

that as much as 50% or more of the flowers on a tree, and as much 

as 90% of the seed crop, may be destroyed. 

The climbing agility of roof rats in Hibiscadelphus trees 

suggests that these non-native animals are capable of foraging in 

a similar manner through the canopies of any native tree, and 

that the nectar feeding habits of the rats probably compete with 

native nectar feeding birds.

LITERATURE CITED 

Baker, J. K., and S. Allen. 1976. Hybrid Hibiscadelphus 

(Malvaceae) from Hawaii. Phyt. 33(4): 276. 

. 1977. Hybrid Hibiscadelphus (Malvaceae) in the 

Hawaiian Islands. Pac. Sci. 31(3): 285-291 

Baker, H. G., and I. Baker. 1973. Amino-acids in nectar and 

their' evolutionary significance. Nature 241: 543-545. 

Bishop, L. E., and D. Herbst. 1973. A new Hibiscadelphus 

(Malvaceae) from Kauai. Brittonia 25(3): 290-293. 

Forbes, C. N. 1920. New Hawaiian plants - VII. Occ. Pap. B. P. 

Bishop Mus. VII(3): 3-4. 

Rock, J. F. 1911. Hibiscadelphus. In L. Radlkoffer and 

7. 

J. F. Rock. New and noteworthy Hawallan plants. Hawaiian 

Bd. Agric. For. Bot. Bull. 1: 8-14.

NENE REINTRODUCTION PROGRAM AND RESEARCH 

IN HAWAIIAN NATIONAL PARKS 

Paul C. Banko 

Cooperative Park Studies Unit 

College of Forest Resources 

University of Washington 

Seattle, Washington 98195 

and 

National Park Service 



The reduction of range and numbers of Nene (Branta 

sandvicensis), the Hawaiian Goose, within historic times has been 

an international concern of conservationists since P. H. Baldwin 

published the results of his literature and field survey 33 years 

ago (Baldwin 1945). At that time the wild population was esti- 

mated at about 50 individuals, but by 1951 only 30 were thought 

to exist in the wild (Smith 1952). A portion of this remnant 

population was breeding immediately adjacent to Hawaii Volcanoes 

National Park (HAVO) at high elevation--about 1980 m (6500 ft) 

(Elder & Woodside 1958). Nene have historically occupied high 

and low elevation habitats within HAVO (Baldwin 1945). However, 

traditional lowland nesting grounds have never been positively 

identified within the Park. The existence of Nene on Maui within 

Haleakala National Park (HALE) is less well documented than in 

HAVO, but the species was believed to have been breeding within 

Haleakala Crater prior to 1890 (Henshaw 1902; Perkins 1903). 

Long before the need for conservation measures became 

apparent, Nene had been bred in captivity. The first record of 

successful captive propagation of the species was in 1834 when 

Lord Stanley, Earl of Derby, reared one gosling out of a clutch 

of four eggs laid at Knowsley, England (Stanley 1834). Young 

Nene were soon distributed to other private collections and zoos 

throughout Europe and eventually became fairly common in cap- 

tivity (Delacour 1954). However, Nene had become rare in 

European collections after 1900 (Blaauw 1904) and finally disap- 

peared when the last specimen, a 42-year old gander, vanished 

during the German invasion of France in 1940 (Delacour 1954). 

The Hawaii Board of Agriculture and Forestry (now Department 

of Land and Natural Resources) began a captive breeding program 

in 1927 at the territorial Game Farm, Mokapu, O'ahu, having 

received a pair of Nene from Mr. Leighton Hind of Pu'u Waawaa 

Ranch (Smith 1952). Mr. Hind and Mr. Herbert Shipman of Hilo, 

who had been maintaining and breeding Nene since 1918, con- 

tributed additional birds to the territorial program and by 1935

the captive flock had grown to 42. In 1935, the flock was dis- 

banded, and the birds were distributed to various individuals who 

were to continue captive propagation efforts. However, within 15 

years, all but one had died, vanished, or had been released to 

the wild (Smith 1952). 

The Board of Agriculture and Forestry began a new captive 

propagation project in 1949 at Pohakuloa which has eliminated 

the immediate threat of extinction and bolstered the number of 

Nene living in the wild through the release of captive-reared 

birds (DLNR 1966, 1968, 1970, 1974, l974a). That the present 

population can increase or remain stzble without additional 

annual releases has yet to be determined. 

In fulfilling its mission to protect, manage, and restore 

native wildlife presently or historically found within Park 

boundaries, the National Park Service (NPS) began Nene propaga- 

tion and release programs in HAVO and HALE in 1972. The goals of 

the HAVO program are to reestablish breeding populations in suit- 

able low and mid-elevation habitats, to maintain and manage 

habitat at all elevations, and to control or reduce factors which 

are inimical to Nene survival. At HALE the objectives are sim- 

ilar, but the habitat available is restricted to high elevations. 

Research begun in 1976 is being integrated with NPS propagation- 

release programs to guide management decisions and to answer 

basic questions regarding Nene life history and ecology. Since I 

have been more directly involved in the HAVO research and 

management program, I will confine my remarks to this park. 

The primary justification for directing propagation and 

release efforts at low and mid-elevations in HAVO has its basis 

in the observations of the naturalists of the 1890's. H. W. 

Henshaw (1902) commented on the lowland breeding range of Nene as 

follows: 

It has been stated and seems to be the general impres- 

sion that the nene rears its young in the uplands where 

it is found in summer, but such is not the fact. The 

greater number, probably all, leave the upper grounds 

beginning early in the fall, and resort to lower alti- 

tudes, from about 1,200 feet downwards. There are 

barren lava flats near the sea in Puna, Kona, Kau and 

Kohala, rarely indeed visited by man, and it is to 

these deserted solitudes that the nene resorts at the 

beginning of the love season. 

The cause of the desertion of the uplands by the geese 

for the low-lying lava flats near the sea is doubtless 

the failure of the food supply in the former, at least 

of such as is adapted to the wants of the young. At 

high altitudes there is but a scanty crop of berries in 

winter, and most of the pualele dies; whereas near the 

sea there is an abundance of this plant and of freshly 

sprouted grasses during the winter and spring months.

R. C. L. Perkins (1903) also believed that Nene bred 

primarily in the lowlands: 

As is well-known the Goose, like many other native 

birds, changes its abode at different seasons of the 

year, being no doubt chiefly influenced by the food 

SUPPI Y - In the summer months it affects the open 

upland region, which is covered with a scrubby vegeta- 

tion and traversed by many lava flows, such for 

instance as parts of the plateau between the three 

great mountains of Hawaii, at an elevation of four or 

five thousand feet above the sea. Near the crater of 

Kilauea about two miles from the Volcano House hotel 

flocks of some size may be occasionally seen in the 

later summer. In such situations it feeds on the abun- 

dant Ohelo berries (Vaccinium), on the wild strawberry 

(Fragaria chilensis) where the cattle still allow this 

to exist, and still more commonlv on the black berries 

of the .creeping Coprosma, one of the commonest plants 

in some of its favourite localities. In the winter 

months large numbers of these upland geese resort to 

the lowlands and remain there for such time as the 

vegetation is fresh and green, and they are said to 

breed during this season. 

G. C. Munro (1944) provides further documentation of lowland 

breeding, as follows: 

It [Nene] had become accustomed to semiarid waterless 

country where it obtained the moisture it needed from 

the upland berries on which it fed in the summer and 

the rich soft plants of the lowland lava flows where it 

wintered and raised its young . . . The sparse vege- 

tation on the open lava flows is rich, especially on 

the lowlands in the wet season, hence the birds 

migrated to the lowlands to breed. These we collected 

there were much fatter than the specimens we took at 

about 2,000 feet elevation. 

We hunted this goose in December 1891 on the rough lava 

flow of 1801, down nearly to sea level, and up the side 

of the mountain on the Huehue ranch to about 2,200 feet 

elevation. It was open shooting season and a party of 

hunters went over ground at the higher elevation where 

we had taken specimens a few days before. They found a 

nest with four eggs, caught two very young chicks and 

shot a young bird nearly full grown. 

When the following record, reported in Baldwin (l945), is 

considered in the context of the observations of Henshaw, 

Perkins, and Munro, there is little doubt that the Nene was once 

a breeding resident of low and mid-elevations in HAVO:

An old Hawaiian resident of Puna, Sam Konanui, who 

traveled across the Puna lowlands to Kau around 1894, 

says Nene were plentiful above the inshore clifEs 

around 1,500 to 2,000 feet but not on the flats whish 

line the shore. He saw them as far to the east as 

Panau. 

In light of what was known about Nene in the 18901s, there 

is ample justification for attempts to reestablish a breeding 

population at lower elevations in HAVO. Further, since Hawaii 

State Division of Fish and Game (HSDFG) has operated its 

restocking program exclusively at high elevations--above 1525 m 

(5000 ft)--there is even a greater need for the HAVO program to 

concentrate on lowland population restoration. The overall 

effect of both state and federal programs will hopefully be to 

complimentarily repopulate the full altitudinal range of Nene 

habitat, at least in Puna and Ka'u. 

Besides focusing management efforts at different elevations, 

the HSDFG and HAVO programs differ fundamentally in the methods 

of propagation and release. HSDFG employs a "game farm" 

approach. Captive birds are intensively bred by incubating eggs 

artificially and naturally (by the goose) and by removing eggs of 

the first clutch so that a second or third clutch w i l l be laid. 

Goslings are brooded artificially and naturally and are kept at 

Pohakuloa until their distribution to release pens prior to 

fledging. Once their wing feathers have fully emerged, the young 

are free to leave the release pen and begin their life in the 

wild. 

At HAVO Nene are bred in pens which are already situated in 

habitats where wild populations are to be reestablished. The 

eight pens so far in use are located from 213 m to 1219 m (700- 

4000 ft) elevation and range in size from about 0.11 ha to 1.47 

ha (0.27-3.63 acres). Pens 1, 2, and 3 are stationed in upper 

'Ainahou where Mr. Herbert Shipman temporarily maintained his 

flock of semi-domesticated Nene from which the original stock was 

obtained for the HSDFG program in 1949. Pen 4 is situated near 

Kilauea Crater, 5 is stationed at Pu'u Kaone, 6 is found at 

Kukalau'ula Pali, and 7 and 8 are situated along the top of 

Hilina Pali in lower 'Ainahou. 

Breeding inside HAVO is unmanipulated in that eggs are not 

removed for artificial incubation nor for obtaining a second or 

third clutch. Parents brood and rear their own goslings during 

the entire period prior to fledging. Once the young are capable 

of flight they depart from and return to the pen many times be- 

fore becoming fully independent. By employing this method of 

propagation and release, young Nene have full contact with their 

parents and siblings, as well as free-living birds which fre- 

quently visit the pens. The young inside the pens also become 

quickly acquainted with the habitat which they w i l l inhabit as 

fledglings and adults. In addition, the young are thoroughly 

acclimatized to the temperature and rainfall regimes with which 

they must cope in the wild.

Although Nene in HAVO pens are exposed to conditions similar 

to those in the wild they are not totally self-sufficient. Pro- 

longed occupancy in the pens results in depletion of food plants, 

SO a commercially available game bird ration is supplied regu- 

larly. Fresh drinking water is continually available, but Nene 

are prevented from bathing in the water and fouling it. Poten- 

tial predators such as mongooses and feral cats, are trapped out- 

side of the pens and are further discouraged from entering the 

pens by small-mesh, buried wire along the bottom portion of the 

pens. However, rats and mice occur in all of the pens and Barn 

Owls are present regularly at Pu'u Kaone. 

HAVO breeding stock has been obtained from two sources, 

HSDFG and U. S. Fish and Wildlife Service (USFWS). HSDFG has 

contributed three pairs, one each in 1972, 1973, and 1976, plus 

three unpaired females and a male in 1976. All were raised at 

Pohakuloa. USFWS has contributed eight females and four males 

reared at Patuxent Wildlife Research Center, Patuxent, Maryland, 

and three females from Northern Prairie Wildlife Center, 

Jamestown, North Dakota. All USFWS birds were less than one year 

old when received in 1974. USFWS has since distributed its 

breeding flock to various U. S. zoos. 

In any program involving penned birds there are deaths and 

escapes which deplete the stock. Opportunities for losses are 

probably higher in the type of program operating in HAVO because 

birds are unattended most of the time and predators, disease, or 

other factors may not be detected or prevented from occurring 

before damage is done. Eight captives have died and one has 

escaped from pens since 1972. In 1973, the only pair of Nene in 

HAVO died, apparently as the result of predation by feral cats 

which were trapped at the pen soon after the incident. One of 

the females obtained from Jamestown died of exhaustion in 

December 1974 while trying to free herself from a piece of fence 

wire which had become caught on her leg band. A pair of Patuxent 

birds died with one of their juveniles in August 1976. A pr-ed- 

ator may have been involved but the actual cause of death was 

impossible to determine since the carcasses were already decom- 

posing when found. Two other juveniles were found unharmed in 

the pen. A Pohakuloa male died in a pen accident in January 

1978. Again in January a Pohakuloa male and Patuxent female died 

in their pen. Both were emaciated although food was plentiful in 

the pen and the intestines and gizzards were nearly full. 

Results of autopsy have revealed no clues as to causes of death. 

To detemine if coccidia might be present in the captive 

birds, fecal samples were sent to Hawaii State Depertment of 

Agriculture laboratories for analysis in 1976, but results were 

negative. 

One Pohakuloa female escaped from her pen in November 1976 

and was never resighted. Pohakuloa and Jamestown birds have tem- 

porarily escaped in the past despite clipped primary feathers. 

When strong winds are blowing, as frequently occurs at Pu'u Kaone 

and Kukalau'ula Pali, even wing-clipped birds are sometimes able 

to clear the pen fence which stands nearly 2 m (6 ft). They are

usually recaptured near the pen and returned. Patuxent Nene have 

not escaped, because they have been tenectomized or tenotomized 

as youngsters and are consequently unable to fully extend one 

wing. Their primary feathers do not require clipping. 

Breeding results at HAVO are summarized in Tables 1 and 2. 

Of 21 nesting attempts since 1974, one was initiated in October, 

four in November, nine in December, two in January, one in 

February, and four in March. The one February and two of the 

March nests represent renest efforts. A total of 77 eggs have 

been laid in the pens, 56 (73%) of which have hatched. The mean 

number of eggs laid per female, including renests, is 4.28 (range 

2 to 121, while the mean clutch size is 3.67 (range 2 to 6). 

Of the 56 eggs which have hatched, 21 goslings (37%) have 

died, almost all within four weeks after hatching. Causes of 

gosling mortality have been difficult to assess; however, preda- 

tion cannot be implicated in at least 13 cases where carcasses 

which showed no wounds were recovered. Disease or nutritional 

deficiencies may have been involved in some fatalities. 

Of the 35 goslings which fledged successfully only three are 

known to have died. As mentioned during the discussion of adult 

mortality, one juvenile died with its parents inside a pen while 

its two siblings survived. Another juvenile died of injuries 

sustained when it flew into the water catchment system at the 

pen. The third known fatality occurred when a juvenile was 

struck by a car on Hilina Pali Road at night. 

Twenty-six of the 32 surviving juveniles have been sighted 

since November 1977 and the majority of these have been seen very 

recently. Only two pen-reared birds have not been sighted since 

July 1977. Most fledglings remain with their parents until the 

following breeding season when finally they appear to leave the 

vicinity of the pen and begin wandering. By the time they are 

one-and-a-half to two years old, pen-reared birds usually reap- 

pear at or near their natal pen. They may be accompanied by a 

mate from another pen, or by one or more siblings, or they may be 

alone. Females have proven to be more faithful than males in 

returning to the vicinity of their natal pen. Only one indi- 

vidual, a male, has been sighted more than a few kilometers from 

his natal pen. This bird was accompanying an unbanded female at 

about 2010 m (6600 ft) elevation near the terminus of Mauna Loa 

Strip Road. All pen-reared Nene are sexed and banded before 

fledging, using unique combinations of colored plastic and seri- 

ally numbered aluminum leg bands. Birds released by HSDFG are 

banded using plastic leg bands only. 

One of the problems affecting the early stage of the HAVO 

propagation-release program has been the lack of mates available 

to birds produced during the first three years. The problem has 

been compounded by the great distance between most pens. Conse- 

quently, the initial broods produced in the pens have been widely 

separated from other broods and have not had the opportunity to 

contact previously released birds which would be expected to be 

found in the vicinity. As a result, independent, pen-reared

irds have been forced to mate with siblings and even captive 

individuals inside the pens. Three free-living, pen-reared 

females have returned to pens and mated with captive ganders. 

Two of these nests resulted in goslings while the eggs failed to 

hatch in the third nest. One sibling mating occurred in the 

wild in 1977, resulting in two offspring, but this pair bond 

dissolved and the female nested with another male in 1978, pro- 

ducing one fledgling. Ironically, this new male partner is a 

brother of the female which was hatched in a different pen. 

Captive stock is rotated annually in the pens so that different 

localities will be seeded by more than one pair. In any case, 

sibling pair bonds w i l l prevail for a few more years until 

fledglings have been produced to provide mates for newly released 

birds. 

To obtain information concerning the breeding biology of 

wild populations in and near HAVO, field work was begun during 

the 1977-8 breeding season at high elevations--1675 m to 2134 m 

(5500-7000 ft)--with the objective of locating and monitoring 

nesting success. Six nests were found between 1860 m and 1980 m 

(6100 & 6500 ft) elevation in the state Keauhou Nene sanctuary, 

while five nests were found between 1920 m and 1950 m (6300 & 

6400 ft) elevation in the adjacent lands of Kapapala within 

HAVO. One wild nest was located at 1160 m (3800 ft) elevation 

near Kilauea Crater. Eight of these nests were active when found 

and the remaining four had been vacated not long before their 

discovery. The progress of the active nests was followed for as 

long as possible, but after the eggs hatched, broods were diffi- 

cult to locate. Four of six pairs that were known to have pro- 

duced viable offspring lost one or more of their young within 

four weeks after hatching. In addition, three pairs were 

observed with young, but nests were not located. 

At least 28 pairs were identified which did not breed suc- 

cessfully, although some apparently made nestlng attempts, 

judglng from the brood patches evident on a few females. Poor 

nesting results in the wild and in HAVO pens was perhaps partly 

due to extraordinarily dry weather occurring throughout the 

breeding season. 

Additional research is being conducted on breeding biology, 

food habits, habitat utilization, behavior, population dynamics, 

and inimical factors of wild and captive Nene in conjunction with 

Park breeding-release programs. The objectives of this research 

are to contribute to the restoration of viable, wild populations 

of Nene and to explore the biology of this unique species of 

goose.


Baldwin, P. H. 1945. The Hawaiian Goose, its distribution and 

reduction in numbers. Condor 47: 27-37. 

Blaauw, F. E. 1904. On the breeding of some of the waterfowl at 

Gooilust in the year 1903. Ibis, 8th Ser., 4: 67-75. 

Delacour, J. 1954. The waterfowl of the world, vol. 1. Country 

Life, London. 284 pp. 

Department of Land and Natural Resources. 1966. Nene 

restoration project report. Elepaio 26: 96-100. 

-- . 1968. Nene restoration project report. Elepaio 

28: 103-106. 

. 1970. Nene restoration project report. Elepaio 

31: 1-7. 

. 1974. 1972 report of Nene restoration program, 1st 

installment. Elepaio 34: 123-127. 

. 1974a. 1972 report of Nene restoration program, final 

installment. Elepaio 34: 135-142. 

Elder, W. H., and D. H. Woodside. 1958. Biology and management 

of the Hawaiian Goose. Trans. North American Wildlife Conf. 

23: 198-215. 

Henshaw, H. W. 1902. Birds of the Hawaiian Islands. Thomas G. 

Thrum, Honolulu. 146 pp. 

Munro, G. C. 1944. Birds of Hawaii. Tongg Publishing Co., 

Honolulu. 192 pp. 

Perkins, R. C. L. 1903. Vertebrata. Pages 365-466 in D. Sharp, 

ed. Fauna Hawaiiensis. Vol. 1, Part 4. Univerzty Press, 

Cambridge, England. 

Smith, J. D. 1952. The Hawaiian Goose (Nene) restoration 

program. J. Wildl. Mgmt. 16: 1-9. 

Stanley, E. S. 1834. (Note on a specimen of a young Sandwich 

Island goose). Proc. Zool. Soc. London, Part 2. Pp. 41-43.

TABLE 1. Individual results of 6nl breedillg in WIVO pens (* = wild-release female which bred 

with captive male; + = died). 

Approx.lBte Eggs Wgs Gosling Fledglings Fledgling 

Breed illg 1st Egg Laid Prcduced Hatched Mrtality Produced Mrtality 

Season Male Female Pen Clutch Clutch Clutch Clutch Clutch Clutch 

1 2 1 2 1 2 1 2 1 2 1 2 

1974-75 962 917 2 29 NOV - 4 - 4 - 0 - 4 - 0 - 

285 3 24 WC 

+8081 4 7 Mar 

8092 4 8 Mar 

917 2 - 

8083 4 - 

- 4 - 

- 4 - 

- 4 -

TABLE 1--Continued . 

Approx. Date Eggs Eggs Gosling Fledglings Fledgling 

Breed irg 1st JQg Laid Produced Hatched Mortality Produced hbrtality 

Season Male Female Fen Clutch Clutch Clutch Clutch Clutch Clutch 

1 2 1 2 1 2 1 2 1 2 1 2 

8088 283 5 19 Nov 1 Feb 4 3 4 3 4 0 0 3 - 0 

962 917 8 12 Dec - 4 - 4 - 0 - 4 - 1 - 

8089 8092 1 13 Dec 7 Mar 6 6 4 6 4 0 0 6 - 0 

8087 8093 6 29 Dec - 3 - 2 - 2 - 0 - - - 

8090 *403 3 30 Dec - 3 - 3 - 0 - 3 - 0 - 

1976-77 8090 285 3 24 Jan - 3 - 0 - - - - - - - 

576 8083 3 - - 0 - - - - - - - - - 

8080 929 7 - - 0 - - - - - - - - - 

8084 912 2 - - 0 - - - - - - - - - 

511 922 5 - - 0 - - - - - - - - - 

8082 - 6 - - - - - - - - - - - - 

32 26 10 16 1 

8088 283 6 30 kt 10 Mar 

8082 *406 4 4 Nov - 

8087 8093 5 30 NW - 

962 917 1 4 Dec - 

8090 285 3 12 Dec - 

1977-78 8084 *401 2 24 Dec - 

8084 912 2 - - 

8089 8092 8 - - 

8080 929 7 - - 

+576 +8083 3 - - 

+511 922 4 - -

TABLE 2. Combined results of NEnS breeding in HAVO pens. 

Breeding 

Season 1974-75 1975-76 1976-77 1977-78 Total 

' NO. Pairs 

Available 

No. Pairs Bred 1 5 6 6 18 

Eggs Produced 4 17 3 2 2 4 77 

Me an No. 

Eggs/Female 4.0 3.4 5.33 4.0 '.4 . 2 8 

(rangel (3-4) (3-12) (2-8) (2-12) 

Me an 

Clutch Size 

(range) 

Eggs Hatched ( % ) 4(1.0) 15( .88) 26( .87) 11( .46) 56( .75) 

Gosling 

Mortality (%) o(0) 4(.27) lo( -38) 7(.64) 21( .37) 

Fledglings 

Produced (%) 4(1.0) 11( .73) 16( .62) 4( .36) 35( .63) 

Fledgling 

Mortality (%) o(0) 2(.18) 1(.06) o(0) 3( .08) 

Includes free-living females which voluntarily occupy and breed inside pens 

with captive males. 

18 females nesting, 3 of which were involved in renesting efforts. 

21 clutches

SOME LIMITING FACTORS AND RESEARCH NEEDS OF 

ENDANGERED HAWAIIAN FOREST BIRDS 

Winston E. Eanko 

U. S. Fish & Wildlife Service 



It is well known that Hawaiian birds are particularly sus- 

ceptible to depopulation and extinction. Twenty-three of 69 

endemic species or races have disappeared since discovery of 

Hawai'i by Europeans 200 years ago. 

Except for Warner (1968) and Atkinson (l977), only super- 

ficial inquiries have been made into historical aspects and 

underlying factors of the Hawaiian forest bird decline. After 

several years of field and laboratory investigation, Warner 

explained the disappearance of forest birds as being caused 

primarily by disease. Atkinson advanced a theory based on 

historical evidence that arboreal predation by rats was a leading 

factor. 

The object of my long-term historical investigation is to 

document and compare the salient facts on the geography and chro- 

nology of Hawaiian bird loss, species by species; to chronicle 

what is known about all factors of depopulation--predation, 

disease, habitat alteration, and food competition; and to draw 

such conclusions as seem warranted. 

At the First Conference in Natural Sciences two years ago, 

Banko and Banko (1976) reported on the potential significance of 

food depletion in the decline of Hawailan forest birds. The role 

played by the Big-headed ant (Pheidole megacephala) in destroying 

much of the endemic insect fauna at elevations generally less 

than 3000 feet (914 m) before 1890 was sketched at that time. 

(The term "insect" w i l l be used hereafter as including other 

arthropods as well). 

I now wish to elaborate on the possible impact of foreign 

parasitic flies and wasps in depleting native insect foods impor- 

tant to the small Hawaiian forest birds at higher elevations. It 

is acknowledged at the outset that all of the information in the 

historical literature dealing with habitat alteration, predation, 

disease, and food competition as they relate to bird decline, has 

not yet been fully extracted or analyzed. However, the effects 

of habitat destruction are in evidence almost everywhere around 

us, while claims that disease and predation were leading factors 

in bird depopulation are now being systematically studied by 

Charles van Riper and James K. Baker, respectively. It may

therefore be of interest to review some historical facts indi- 

cating parasitism of bird food-insects by foreign flies and wasps 

played a prominent role in depopulating birds through depletion 

of food supplies. If this theory is correct, foreign parasites 

may be a major factor limiting population size and range of 

forest birds today. 

In considering the significance of insect foods to forest 

birds, it is first necessary to point out that insects are impor- 

tant, if not essential, in the diets of the young of all species, 

irregardless of food preferences of adults. Though specific 

foods of nestling and newly-fledged forest birds have not been 

studied extensively, judging from what has been recorded in the 

literature, their diets appear to consist principally if not 

entirely of insects. 

The history of events in a forest ecosystem above 3000 feet 

(914 m) elevation illustrates the destructive impact of foreign 

parasites on native insect foods and indirectly on populations of 

endemic forest birds, thereby serving as a model of typical 

effects which have occurred in Hawaiian National Parks and else- 

where. A suitable case history is provided by the Acacia koa 

forest which extends some 40 miles (64 km) along the Kona Coast, 

at a'pproximately 4000 to 7000 feet (1219-2134 m) elevation. 

Eiahtv-five vears aao this Kona Koa forest suooorted 

substantiai popuiations a of the 'Oma'o (Phaeornis ob'icurus 

obscurus), 'O'u (Psittirostra psittacea) , and G r x r Koa Finch 

(Psittirostra palmeri), to mention but a few of the dozon or so 

species of small forest birds which formed the avifauna of that 

p&ticular region. Perkins (1903: 375, 433, 435) noted the 

'Oma'o to be "almost ubiquitous throughout the forest . . . . 

from the lower limits to the upper"; the 'O'u in Kona "in count- 

less numbers," moving seasonally upwards into the Koa woods; and 

the Greater Koa Finch seen in "hundreds" between 4000 and 5000 

feet (1219 and 1524 m) in the Koa belt. 

Today, judging from a review of the literature and from per- 

sonal observation, populations of the 'Oma'o, 'O'u, and Greater 

Koa Finch are either absent or in low numbers in the Kona Koa 

forest, exemplifying what has happened to other native Hawaiian 

forest birds in Kona and elsewhere. 

Consider the following ecological relationships and history 

of events. Acacia &, especially old decadent stands, is of 

paramount importance to Hawaiian forest birds as a source of 

insect food. Beginning in 1887 a long procession of naturalists 

and ornithologists found more kinds of birds in Koa woods than in 

any other type of Hawaiian forest. In retrospect, the most obvious 

rationale for avian diversity in Koa forests is that Acacia 

- koa harbored a greater variety of endemic insects than any other 

generic group of Hawaiian trees (Swezey 1954: 1).

Of all the insects hosted by Acacia koa none were apparently 

as important as food to more kinds of forest birds than the 

endemic moth genus Scotorythra spp. (Geometridae) . Af ter years 

of observing feeding behavior and investigating stomach contents 

of many species of Hawaiian forest birds, Perkins (1913: clii) 

concluded that Scotor thra spp. "form a most important part of 

the food supply o 

--f41-, 

endemlc birds, and are supplied by the parents 

to the young of nearly all the species, while they are a favorite 

food of many adult birds as well." The historical record is 

replete with references to the prominence of Scotorythra spp., 

and other caterpillars, in diets of Hawaiian forest birds. 

While Scotorythra spp. moths and larvae were found on 

Hawaiian trees and shrubs other than Koa, it was the species that 

fed on Koa that were in such demand. Some species of Scotorythra 

hosted by Koa were very abundant in the 18901s, and occasionally 

erupted, defoliating entire forests. 

In Kona, the 'Oma'o, 'O'u, and Greater Koa Finch had special 

affinities for Scotorythra spp., or looper caterpillars as they 

were commonly called. Perkins (1903: 374) mentions several 

instances of the 'Oma'o feeding on Koa loopers and adds that 

these birds "also continue to feed the young on these for some 

time after they have left the nest." Perkins (1903: 433-434) 

also documented the seasonal dependence of 'O'u on caterpillars 

after fruiting of its normal food, 'Ie'ie (Freycinetia arborea), 

had terminated. On several occasions, Perkins observed excur- 

sions of the 'O'u out of its usual haunts into Koa woods for the 

purpose of obtaining (Scotorythra spp.) caterpillars. Likewise, 

Perkins (1903: 437) noted that the Greater Koa Finch displayed an 

affinity for Koa looper caterpillars. 

Unfortunately, Scotorythra caterpillars are also preyed upon 

by foreign parasitic insects which began arriving in the Islands 

about 1890 ancl continued to become established until at least 

1942. While the impact of foreign organisms on insect foods of 

native birds has never been systematically studied in the field, 

at least four kinds of exotic flies and wasps are known to attack 

Scotorythra spp. What little is known of the arrival and status 

of foreign parasites in the Islands, and their relationships to 

Scotorythra spp., is condensed in Tables 1 and 2. 

According to various authorities quoted in Table 1, two for- 

eign tachinid flies arrived before 1900 and, one at least, was 

universal in mountain forests about 1892. An ichneumonid wasp 

was first discovered on O'ahu in 1925 and by 1931 had become very 

numerous at times. A braconid wasp was introduced in 1942 and 

not long thereafter was found well into the native forests of the 

six major Hawaiian Islands. Some foreign fly and wasp parasites 

'apparently have high reproduction and dispersal characteristics. 

By the 1940's it was known from impromptu investigations by 

authorities cited in Table 1, that the two tachinid flies at- 

tacked one and three species of Scotorythra spp., respectively. 

The ichneumonid wasp was discovered to hit seven species of 

Scotorythra spp., four host-specific to Acacia koa. The braconid

wasp was found to parasitize the larvae of one species of 

Scotorythra spp. moth hosted by Acacia - koa. 

Most, if not all of the prey-parasite relationships cited by 

E. C. Zimmerman in Table 2 were discovered by an economic ento- 

mologist, 0. H. Swezey. During a long life of work in the field 

and laboratory, from about 1904 to the late 19301s, Swezey spent 

what time could be spared from his official duties in the forest 

studying Hawaiian. insects. 

In the 40 years since Swezey ceased to be active in the 

field, other potential parasites of Scotorythra spp. have 

arrived, creating a need for revision and possible expansion of 

host relationships shown in Table 2. For example, Bianchi (1959: 

993) documented the introduction to Hawai'i of three braconid 

wasps, including Apanteles marginiventris and Meteorus laphygmae, 

in addition to the apparently accidental arrival of another 

tachinid fly, ~ucelatoiia armigera. Bianchi reported that it was 

not long after arrival of these parasites in 1942 that they 

became abundant in the lowlands and then spread to the upland 

grass ranges and even well into native forests where he theorized 

they might parasitize native caterpillars unrecorded as hosts. 

Whitesell (1964) stated, from a forester's viewpoint, that 

Scotorythra spp. appeared to be under good 

and seldom built up to "damaging levels." 

control biologically 

Reduction of Scotorythra spp. populations by continental 

flies and wasps is not the only example which might be cited of 

depletion of a valuable insect food by foreign parasites. 

Another large group of endemic moths extensively parasitized by 

foreign insects is the family Pyralidae. Pyralids were major 

foods of the Palila (Psittirostra bailleui) and Greater Koa Finch 

(Psittirostra palmeri), and a general food of the Hawaii Creeper 

(Loxops maculatus mana), according to Perkins (1903: 436, 437; 

1913: clx). In Kona, Perkins (1903: 414, 435) found the Hawaii 

Creeper "extremely common" at about-3500 feet (1067 m) upwards, 

and the Palila "extremely numerous" from below 4000 to at least 

6000 feet (1219-1829 m). 

Commenting on the biological control of species in one genus 

of Pyraustidae, Hedylepta, Zimmerman (1958b[8]: - 66, 68-69) 

states: 

. . . eight foreign wasps and three foreign flies are 

extremely active parasites, and parasitism now commonly 

exceeds 90 per cent . . . Two of the species of this 

genus (Hedylepta) break the general rule that endemic 

Hawaiian insects are not pests of economic importance, 

because (H.) accepta is the well-known sugarcane leafroller, 

and (H.) blackburni is the common coconut leafroller. 

The Tntroduction of parasites to control these 

species, especially the sugarcane leafroller, have 

resulted in mass destruction of the endemic Lepidoptera 

and have greatly altered the composition of the 

fauna of the islands. 

insect

Today the Palila is not known to inhabit the Mamane (Sophora 

chrysophylla) zone of the Kona Koa forest and the Hawaii Creeper 

appears to be much less common than in the past. 

Foreign flies and wasps were not the only organisms de- 

pleting foods of forest birds. Other continental and Hawaiian 

organisms also compete with endemic forest birds for food. But 

continental parasites appear to have played a dominant role in 

many areas. 

It is clear from the historical record that endemic 

Lepidoptera were the most heavily utilized food resources of the 

small Hawaiian forest birds in the 1890's. Concerning their gen- 

eral depletion in the past 80 years, Zimmerman (1958g171: 28) has 

this to say: 

When Dr. Swezey laid down his pen and left Hawaii (in 

1952) a golden era of Hawaiian entomology closed . . . 

Swezey was the last of the entomologists to have seen 

many of the endemic Hawaiian Lepidoptera in a semblance 

of their natural abundance. The importation of parasites 

to control various moths of economic importance, 

together with the accidental importation of other parasites 

has resulted in wholesale slaughter and near or 

complete extermination of countless species. It is now 

impossible to see the Hawaiian Lepidoptera in the 

natural proliferation of species and individuals of 

Perkins day. Many are lost forever. 

From an analysis of the historical literature, it may there- 

fore be argued that depletion of insect foods by waves of foreign 

parasites was a significant factor in depopulation of endemic 

Hawaiian forest birds. If true, such a theory would have serious 

contemporary implications. Many continental ants, flies, wasps, 

and other foreign organisms are well established in native 

forests, and others continue to arrive. Wherever endemic birds 

are constrained to share food with continental fauna, superior 

competition by foreign species possess perhaps overlooked poten- 

tial to significantly restrict population sizes and ranges of 

Hawaiian birds. 

It is my contention that the overall predicament of endan- 

gered forest birds is primarily the result of dissolution of the 

natural Hawaiian ecosystem by continental fauna, and that the 

challenge of preserving what remains can best be accomplished by 

efforts to learn more about, maintain, and reestablish long 

neglected but vital interdependencies. Viewed in this light, 

research to determine the role of food supply in first reducing 

and then limiting population size of endemic Hawaiian forest 

birds is seen as an urgent conservation necessity.

In line with this thinking, immediate inquiry is needed to 

shed light on the following questions: 

1. What is the present distributional relationships between 

availability of insect foods and endemic forest bird 

populations, especially during and after the critical 

nesting and fledging periods? 

2. What proportions of diets of the various forest bird 

species are composed of foods of foreign origin? 

3. Are foreign organisms limiting the quantity and quality 

of food presently available to forest birds? If so, 

which exotics are the most influential and where are 

they exerting the most competitive pressure? 

4. Are Scotorythra spp. populatians under biological 

control by foreign organisms as has been asserted? 

5. What is the contemporary role of predatory ants in 

reducing food supplies of forest birds? 

These and many other high priority questions face contem- 

porary research ecologists charged with preserving endangered 

birds and maintaining native Hawaiian ecosystems.


Atkinson, I. A. E. 1977. A reassessemnt of factors, particu- 

larly Rattus rattus L., that influenced the decline of 

endemic forest birds in the Hawaiian Islands. Pacific Sci. 

31(2): 109-133. 

Banko, W. E., and P. C. Banko. 1976. Role of food depletion by 

foreign organisms in historical decline of Hawaiian forest 

birds. Pages 29-34 - in C. W. Smith, ed. Proceedings, First 

Conf. in Natural Science, Hawaii Volcanoes National Park. 

CPSU/UH (University of Hawaii, Botany Dept.) 

Beardsley, J. W. 1961. A review of the Hawaiian Braconidae 

(Hymenoptera). Proc. Hawaii Ent. Soc. 17(3): 333-366. 

Bianchi, F. A. 1959. Entomological changes in the sugarcane 

fields of Hawaii. Proc. 10th Cong. Int. Soc. Sugar Cane 

Tech., pp. 989-994. 

Perkins, R. C. L. 1903. Fauna Hawaiiensis. Vol. l(1V). 

Vertebrata. Cambridge At The University Press. PP - 

365-466. 

. 1913. Fauna Hawaiiensis. Vol. l(V1). Introductory 

essay on the fauna. Cambridge At The University Press. 

Pp. 1-ccxxviii and 16 plates. 

Swezey, 0. H. 1954. Forest entomology in Hawaii. B. P. Bishop 

Museum Speci.al Publication 44. Bishop Museum Press, 

Honolulu. 

Warner, R. E. 1968. The role of introduced diseases in the 

extinction of the endemic Hawaiian avifauna. Condor 70: 

101-120. 

Williams, F. X. (compiler). 1931. Handbook of the insects and 

other invertebrates of Hawaiian sugar-cane fields. Exp. 

Sta. Haw. Sugar Planter's Assoc., Honolulu. 400 pp. 

Whitesell, C. D. 1964. Silivical characteristics of koa (Acacia 

- koa Gray). U. S. Forest Service Res. Paper. PSW-16. 

12 PP. 

Zimmerman, E. C. 1958a. 

- Insects of Hawaii. Vol. 7. 

Macrolepidoptera. University of Hawaii Press, Honolulu. 

542 pp. 

. 1958b. Insects of Hawaii. Vol. 8. Lepidoptera: 

~yraloidea. University of Hawaii Press, Honolulu. 456 pp.

aIal 

5 ." 

U) 

0 a, 

U u 

C 0 

4 w 

rlw 

rl 

aIw 

3 0 

N 

w 

m 

rl 

C 

.rl 

'& 

L1 

U 

C 

.rl

TABU 2. Foreign parasites of Scotorythra spp. (Zimnerman 1958a[7]: - 42-147). 

IZLCHINID FLIES 

Spec if ic 

Host Caterpillar Host Plant 

-- 

Chaetogaedia monticola - S. corticea kacia koa 

Frontina archipivora - S. paludicola Acacia - koa 

-- 

-- 

ICHNEXMONID WASP S. rara 

Acacia koa 

Hpsoter exiguae - S. caryopis Acacia - koa 

BRACONID WASP - S. paratactis Dodonaea spp. 

Apanteles marginiventris - S. trapezias mdonaea spp. 

- S. sp.? 

Dubautia spp.

BIOLOGICAL CONTROL OF WILDLAND WEED PESTS IN 

HAWAI'I--IS IT A FEASIBLE SOLUTION? 

John W. Beardsley, Jr. 

Department of Entomology 

University of Hawaii at Manoa 


The problem of aggressive exotic weed species invading 

native Hawaiian ecoysystems and overwhelming or out-competing 

endemic plants has been a recurrent one since man's arrival in 

these islands. The Polynesians brought with them the hau and the 

kukui, among others, which have since become widespread and abundant 

elements of the Hawaiian flora. These at least are plants 

which at the time were useful. Since the arrival of the Europeans, 

we have seen Hawai'i's forests invaded in successive waves 

by lantana (Lantana camara), the guavas (Psidium guajava and 

- P. cattleianum), rose myrtle (Rhodomyrtus tomentosa), the firetree 

(Myrica 

blackberries (Rubus spp. ) , melastoma 

(Melastoma mala P athricum), banana pok~ssiflora mollissima), 

Koster's curse (Clidemia hirta), and New Zealand tea (Leptos 

ermum sco arium), to mention a few of the more obnoxious 

-. -77 

species Severa of these are still rapidly extending their 

ranges and some, such as banana poka and Koster's curse, appear 

to be causing the rapid decline and disappearance of elements of 

the endemic flora in those areas which they have invaded. I am 

sure that the botanists could name additional species which 

invaded wildland ecosystems within the past few decades, and I am 

almost certain that we will be seeing other species, which are 

not yet considered to be problems, developing into serious pests 

in the future. 

Biologists concerned with the preservation of native 

Hawaiian ecosystems and the individual elements thereof, are 

faced with a serious dilemma. The cost of physically or chem- 

ically removing or killing invading weed species which threaten 

native ecosystems is generally prohibitive, given the budgetary 

limitations under which most of us must operate, and excepting 

incipient infestations which involve relatively small and acces- 

sible areas. Furthermore, physical and chemical methods often 

have undesirable side effects such as the inadvertent destruction 

of native plants. Also, such methods are rarely 100% effective, 

which means that within a few years the treated area, in all 

probability, will have been reinvaded from adjacent untreated 

1 Published with the approval of the Director of the Hawaii Agri- 

cultural Experiment Station as Journal Series No. 2249.

lands; or a few surviving plants, or their seeds, will have 

reestablished the weed infestation. In most situations, it seems 

to me, the application of physical methods or herbicides to 

combat well-established aggressive weed species in Hawaiian wild- 

land ecosystems is, in the long run, doomed to failure. I be- 

li,eve that the biological method of control offers a practical 

alternative to physical and chemical methods which can be suc- 

cessfully utilized against many of the weed pests in Hawai'i 

which compete with native plants. 

It has been repeatedly observed that in those areas in which 

they are endemic, plant species which have become aggressive 

weeds in Hawai'i are, in general, relatively minor and innocuous 

elements of the floras in which they occur. Thus, in Mexico, the 

exploratory entomologist Albert Koebele found Lantana camara 

occurring only sparingly as scattered shrubs, but not in continuous 

stands (Perkins & Swezey 1924). Similarly, it has been 

stated that in tropical America Passiflora species, such as 

- 

P. mollissima, occur primarily as scattered individuals in forest 

environments, not as overwhelming canopies (Gilbert, pers. 

comm. ) . 

There is an increasing body of evidence that, in many 

instances, the distribution and abundance of a particular plant 

species is determined not only by parameters of the physical 

environment and by competition from other plants, but also by the 

predators and parasites which feed upon it e . , herbivorous 

animals and pathogenic microorganisms). In the case of tropical 

passion vines, for example, Gilbert (1975) has shown that heavy 

herbivore pressure from the larvae of Heliconius butterflies has 

resulted in the hyperdispersion of Passiflora populations in 

Central American forests. Furthermore, in the case of many, 

perhaps most, phytophagous arthropods and plant diseases, long 

coevolution between the plant and its natural enemies has 

resulted in highly specific ~ host/herbivore 

--- ~ ~- ~ and host/parasite 

~ ~ ~ ~ ~~~~. - . 

relationships. 

When a potential weed species is brought to Hawai'i, usually 

in the form of seed, it leaves behind virtually all of these spe- 

cific types of associated arthropods and disease organisms. 

Thus, freed from the constraints exercised by these specific 

natural enemies, it is able to flourish and reproduce far beyond 

what would be possible in those areas where it is endemic, out- 

competing and overwhelming native species which bear their own 

burdens of specific native herbivores and parasites. 

The classical biological control strategy for combating an 

introduced pest organism, be it arthropod or weed, involves 

seeking out natural enemies of the pest in those areas where it 

is endemic, and establishing these in areas which the pest has 

invaded. The method has worked extremely well against several 

very serious range and pasture weeds (e.g., Opuntia Spp. in 

Australia and Hawai' i; Hypericum perforaturn in Australia and 

California; Lantana camara in several tropical areas, including 

Hawai'i) . Lantana, although it cannot be said to have been com- 

pletely controlled in all situations in Hawai'i, is today, with

15 species of introduced insects established on it, under a great 

deal of herbivore pressure which did not exist prior to these 

introductions, and is far less prevalent than it was at the turn 

of the century (Perkins & Swezey 1924). 

I believe that the reason why the majority of the serious 

weeds which affect native ecosystems in Hawai'i have not been 

brought under biological control is simply that, for most of 

them, little or no effort has yet been expended. Work which has 

been done on controlling Lantana and Clidemia, was directed at 

these species primarily as range pests; hence the natural enemies 

best suited to control these species in non-rangeland ecosystems 

may still be undiscovered. 

A major concern, often expressed by biologists and non- 

biologists alike, is that organisms which are imported for 

biological control of weeds will themselves become pests by 

attacking economic plants, ornamentals, or elements of the native 

flora. Careful research and testing carried out in the areas of 

origin, and under quarantine at the destination, can almost com- 

pletely eliminate this possibility. About 40 species of phyto- 

phagous insects have been successfully introduced into Hawai'i to 

combat weeds since this phase of biological control was initiated 

in 1902. Of these, one of the earliest introductions, made 

before adequate procedures for testing candidates for intro- 

duction had been developed, became a very minor pest of eggplant, 

and another, also among the first introductions, has twice been 

reported feeding on a native tree (Myoporum). These are the only 

'exceptions I know of to an otherwise unblemished record. The 

generally high degree of host specificity which is characteristic 

of many phytophagous arthropods and disease organisms, plus the 

fact that many of the important forest weeds have no close rela- 

tives among the endemic flora, reduces the chance of unforeseen 

host transfer by well selected biological control organisms to 

the realm of a remote possibility. Even in the case of a weed 

such as banana poka, where a member of the same genus is a food 

plant of minor economic importance, there is still a good possi- 

bility of achieving biological control without materially 

affecting commercial passion fruit production. Species-specific 

insects or diseases may exist which w i l l not affect the culti- 

vated passion. Furthermore, ecological isolating mechanisms may 

exist which would prevent species introduced to combat a wildland 

weed from attacking a related crop plant growing in an agricul- 

tural or urban environment. Thus, among Heliconius the specific 

ecological and host requirements (i.e., ovipositional stimuli) of 

the adult butterflies limits their oviposition to specific 

Passiflora species within certain forest environments, even 

though the larvae themselves may be capable of feeding on other 

species of Passiflora. 

In the case of weeds such as the introduced grasses which 

grow in environments similar to those of native grasses, or the 

introduced Rubus species which may occupy habitat similar to that 

of the endemic R. hawaiiensis, there is perhaps less chance of 

achieving satisfactory biological control without some damage to

the native flora, although the possibility of finding phyto- 

phagous forms with a sufficiently high degree of host specificity 

still exists. For example, the various species of smut fungi 

which attack grasses usually are highly host specific. 

The biological method of weed suppression is, of course, no 

panacea. Even in the most successful programs the target weed 

remains present in the environment, although reduced to the 

status of a relatively minor element of the flora, limited to 

those special sites where it can survive and compete successfully 

despite the pressure of its introduced natural enemies. However, 

barring some major ecological upset, control, once achieved, is 

permanent and self-perpetuating. In achieving control we w i l l 

have added some additional elements to the total biota, even 

though these elements are restricted to close association with 

the target weed. These consequences must be accepted if a 

biological control program is to be undertaken. 

To me, the choice, with respect to many of our more aggres- 

sive wildland weeds, is obvious. Either we opt for biological 

control, or we accept the fact that there is no economically 

feasible control available. I believe that biological control is 

an acceptable, and perhaps the only practical alternative for 

controlling many of the more serious wildland weed pests in 

Hawai' i. 


Gilbert, L. E. 1975. Ecological consequences of a coevolved 

mutualism between butterflies and plants. Pages 210-240 in 

L. E. Gilbert and P. H. Raven, eds. Coevolution of animai? 

and plants. Univ. of Texas Press, Austin. 

Perkins, R. C. L., and 0. H. Swezey. 1924. The introduction 

into Hawaii of insects that attack Lantana. Bul. Expt. Stn. 

Hawaiian Sugar Planters Assoc., Entomol. Series no. 16, 

83 PP.



INSECTS AND OTHER TERRESTRIAL ARTHROPODS~ 

J. W. Beardsley, Jr., R. Burkhart, 

M. L. Goff, A. Hara, and G. Teves 




During the past three summers (1975-77) RBI collections of 

insects and other terrestrial arthropods were made at numerous 

sites within Haleakala National Park, primarily along established 

RBI transects. The most intensive collecting was done in the 

relatively dry higher elevation areas of the Park above 1800 m, 

within the crater and on the western rim between Hosmer Grove and 

the summit, and in Kaupo Gap at 1500 m elevation and above. 

These collections contain approximately 20,000 specimens, most of 

which have been mounted and labeled for study. Much of the 

material has been identified by the authors and collaborators, 

and a preliminary checklist has been prepared. The ultimate 

objective of this study is to produce, in as complete a form as 

possible, a checklist of the terrestrial arthropods of Haleakala 

National Park, annotated with data on host relationships, distri- 

bution, and other pertinent ecological information. However, 

several important major taxa are as yet either unstudied or only 

partly identified, and it is anticipated that the checklist will 

require several years for completion. 

On the basis of material which has been identified to date, 

we estimate that our Haleakala collections contain well over 400 

species of insects and other arthropods, of which approximately 

250 species are Hawaiian Island endemics (greater than 60% of the 

total fauna), and at least 50 species are known only from 

Haleakala. The collections contain a substantial number of 

undescribed endemic insects, several of which had not been col- 

lected previously (e.g. , a new flower-infesting tephritid fly 

associated with the Maui wormwood, Artemesia mauiensis). On the 

other hand, several endemic species previously described from 

Haleakala are not represented in our material. 

We estimate that our collections contain around 70-80% of 

the terrestrial arthropod species which occur in the highlands of 

1 Published with the approval of the Director of the Hawaii Agri- 

cultural Experiment Station as Journal Series No. 2253.

Haleakala. However, the wetter environments found within the 

Park are relatively poorly represented in our material. Collec- 

tions from Paliku and vicinity contain a number of rain forest 

associated elements which were not taken elsewhere within the 

Park, but the Paliku fauna appears to be a relatively depauperate 

segment of the richer wet forest insect faunas which occur out- 

side the Park boundaries, and, probably, within the Kipahulu 

extension. Should collections from the Kipahulu area become 

available for study, we anticipate that the present checklist 

would have to be greatly expanded. 

Collecting Methods and Results 

The beating net proved to be the most productive tool for 

sampling the insect faunas associated with various species of 

shrubs and small trees which dominate the flora of the higher 

elevation ecosystems of Haleakala. Hand picking, while sorting 

through leaf litter and bunch grasses, and searching beneath 

loose stones were most effective for sampling the litter and soil 

associated forms, many of which are flightless. Pitfall traps 

proved useful in some situations for sampling nocturnally active 

ground dwellers such as carabid beetles. Aerial nets were used 

to some extent to sample diurnal flying insects such as the 

Odonata, Diptera, Aculeate Hymenoptera, and diurnal Lepidoptera. 

A malaise trap proved to be productive of flying forms in some 

areas where suitable sites for erection of the trap were avail- 

able. A battery-powered ultraviolet light trap was operated at a 

number of sites, both within the crater and on the west rim, with 

generally good results. The light trap yielded primarily noctuid 

moths, both endemic and introduced species, frequently in large 

numbers. In comparison to similar traps which have been operated 

elsewhere, relatively small numbers were obtained of species 

belonging to such moth families as the Geometr idae and Pyral idae, 

which normally are attracted to light in large numbers, except at 

Paliku where some night-flying rain forest-associated elements 

occurred. It appeared that, except for the Noctuidae, there was 

relatively little nocturnal flight activity in most of the areas 

sampled. This may have been a consequence of the relatively low 

night temperatures which prevail on Haleakala at elevations above 

1800 m. Some species which belong to groups that are usually 

night fliers, appear to be primarily day fliers on Haleakala. 

For example, Eupithecia scoriodes (Meyrick), a geometrid moth 

endemic to Haleakala, was often taken while flying in daylight, 

but never taken in our light trap 

The endemic insect fauna of the Hawaiian Islands is charac- 

teristically disharmonic, with many of the major taxa very poorly 

represented or completely absent (Perkins, 1913; Zimmerman, 

1948). The fauna endemic to the higher elevations of Haleakala 

is particularly depauperate as some elements which are better 

represented in endemic faunas of lower elevations (e.g., many of 

the endemic weevil genera; the orthopteroid families Gryllidae 

and Tettigoniidae) are virtually or completely absent in the 

higher and drier ecosystems of Haleakala. Some of the more suc- 

cessful groups which have been able to occupy these high altitude

areas include the planthoppers (Delphacidae) and mealybugs 

(Pseudococcidae) , the seed bugs (Lygaeidae) , plant bugs 

(Miridae), and the predaceous Nabidae, among the Hemiptera; the 

Hemerobiidae (Neuroptera) ; the Carabidae (Coleoptera) ; the 

Tephritidae (Diptera); certain elements of the Lepidoptera, 

particularly the Noctuidae but also some of the Geometridae, 

Pyralidae, Xylorictidae, and Cosmopterygidae; and the Aculeate 

Hymenoptera, represented by the Eumenidae (Odyneurus) and 

Sphecidae (Ectemnius) and the Hylaeidae (Nesoprosopis). 

It is possible to make some further generalizations con- 

cerning the endemic arthropods of the high elevation ecosystems 

of Haleakala, based upon collections and associated ecological 

data obtained during the RBI survey, plus information derived 

from earlier collections and publications. 

1) The non-native insect fauna (recent adventives) of 

Haleakala includes a number of temperate climate (Holarctic 

or Nearctic) species which do not occur in lowland areas of 

the state, although many are found at the higher elevations 

on other islands (e.g., the syrphid fly, Eristalis tenax 

L.; the vespid wasp, Vespula vulgaris (L.); the brown 

lacewing, Hemerobius pacificus Banks; and the ensign scale, 

Arctorthezia occidentalis (Douglas), and several species of 

aphids. 

2) Among the endemic phytophagous insects, particularly the 

Hemiptera, most groups exhibit a high degree of host speci- 

ficity. Thus, among the mealybugs (Pseudococcidae) , plant- 

hoppers (Delphacidae), plant bugs (Miridae), and seed bugs 

(Lygaeidae) most species appear to be restricted largely to 

one, or a few closely related species of native hosts. 

There are exceptions, such as the mealybug Pseudococcus 

nudus Ferris, which, although restricted in distribution to 

elevations above 1800 m, infests several unrelated native 

plants (i.e., Dubautia, Styphelia, and Vaccinium). 

3) Many of the most precinctive endemics are flightless 

insect species which belong to groups that are usually 

capable of flight. For example, the flightless endemic 

ground beetle genus Mecyclothorax Sharp (Carabidae) attains 

its greatest diversity on East Maui with 36 described species 

on Haleakala (out of 86 known species), only one of 

which is known elsewhere (West Maui). Other unusual flightless 

elements of the Haleakala insect fauna include the 

xylorictid moth, Hodegia apatella Walsingham; the flightless 

lacewings (Hemerobiidae) Pseudospectra lobipennis Perkins, 

- P. cookeorum Zimmerman, and Nesothauma haleakalae Perkins; a 

flightless dolichopodid fly, Campsicnemis haleakalaae 

Zimmerman; and a flightless reduviid bug, Saicella smithi 

Usinaer. It should be ~ointed out that most of the aenera 

mentioned contain-other-flightless species which occur2elsewhere 

in Hawai'i.

4) There are many similarities between the high altitude 

endemic insect fauna of Haleakala and those of Mauna Loa, 

Mauna Kea, and Hualalai on Hawai'i. Many species are common 

to both islands. Other elements of the Haleakala fauna are 

represented on Hawai'i by closely related species which 

occur in similar environments (e.g., the large endemic 

Haleakala ground beetle, Barypristus rupicola [Blackburnl 

and its Hawai'i counterpart, B. in~endarius [Blackburn] ). 

However, there are some elements of the Haleakala fauna 

which appear to be unique to that mountain, and for which 

homologues apparently do not exist on Hawai'i; for example, 

the laige daylflying geometrid moth Megalotica (Megalotica) 

holombra (Meyrick). The high altitude insect fauna of 

Haleakala seems to be slightly more diverse than that of 

Hawai'i, although this may be due, in part, to more thorough 

collecting. 


Perkins, R. C. L. 1913. Introduction. Pages 1:XV-CCXXVII 

D. Sharp, ed. Fauna Hawaiiensis. Cambridge Univ. Press, 

Cambridge. 

Zimmerman, E. C. 1948. Insects of Hawaii. Vol. 1. Intro- 

duction. Univ. of Hawaii Press, Honolulu. XVII + 206 pp.

HAWAII IBP SYNTHESIS : 

5. SHORT-TERM TEMPORAL PATTERNS AMONG ISLAND BIOTA* 

Kent W. Bridges 




Analyses have been undertaken to determine the extent to 

which temporal patterns can be seen in the biota sampled in the 

IBP studies. Particular attention has been given to the quanti- 

tative analysis of these trends to test their correspondence to 

temporal and general climate patterns and specific climatic 

events. These patterns will be discussed relative to specific 

organism groups. 

Conclusions will be drawn as they relate interspecific rela- 

tionships and temporal phasing to problems of island ecosystem 

stability. 

* Abstract

SECOND CONFERENCE ON NATURAL RESOURCES 

REMARKS BY ROGER SKOLMEN 

AT DEDICATION OF THE HAWAII FIELD RESEARCH CENTER 

ON BEHALF OF STATION DIRECTOR CALLAHAM 

Robert 2. Callaham 

Pacific Southwest Forest and Range Experiment Station 

Berkeley, California 

I am delighted to be here this morning representing Director 

Callaham, our Station Director. He regrets being unable to be 

here but sends his greetings and enthusiastic endorsement of the 

development of this Center, which includes the new field labora- 

tory for our Experiment Station. Everyone in the Forest Service 

had planned and hoped to have our building in place by the time 

of this dedication. But our building is now on a boat somewhere 

this side of Oakland. Ordered from the factory in Oakland last 

March, it is expected to arrive at any moment. 

Ours w i l l be a carbon copy of the building purchased by the 

Fish and Wildlife Service. Attractive siting was planned by Tom 

Fake of the National Park Service. Only by piggybacking on the 

order for a building by the Fish and Wildlife Service and having 

strong support by the Park Service were we able to get this new 

facility. Their cooperation and help made it all possible. 

This new field lab will provide space for eight of our sci- 

entists and technicians who will work here most of the year. 

They belong to one of the four research projects of our Institute 

of Pacific Islands Forestry. The aim of this project is to 

provide technology for maintaining native Hawaiian forest ecosys- 

tems. Their particular purpose is to learn more about the hab- 

itat requirements of endangered forest birds on the Big Island. 

Emphasis so far has been in the Kilauea Forest Reserve--Keauhou 

land area. We are starting new work on the Hawaiian Crow and 

endangered plants. C. J. Ralph, leader of this research project, 

and his associates will be speaking on these subjects today. 

This new field laboratory certainly will expedite our 

research. Most of the researchers' time is spent in the field, 

establishing plots, capturing and examining birds, and carefully 

monitoring them as they feed, rest, seek cover, and nest. The 

lab w i l l give our research team a place to work on equipment, 

analyze data, and do some bench-related research. Most impor- 

tant, its location here in this Research Center w i l l enable our 

people to share experiences, ideas, and resources with others. 

Such a place is vital to the success of our research program.

We in the Forest Service are excited about joining our 

research forces with others who will work here at the Hawaii 

Field Research Center. The National Park Service is to be com- 

mended for its initiative and hospitality which brought us all 

together. The environment is bound to be synergistic, causing 

each to produce more than if we were working alone. Such syner- 

gism is essential, for there is a greater need for knowledge than 

there is a research capacity to produce it. We expect and know 

that this Center will foster cooperation, coordination, and 

mutual concern, enabling all of us to find the answers we seek.

HYBRIDIZATION IN THE HAWAIIAN SILVERSWORD COMPLEX 

Gerald D. Carr 




The Haleakala silversword (Arqyroxiphium macrocephalum) Gray 

is one of Hawai'i's most well-known and publicized endemic 

plants. Each year it is in fact sought out by hundreds of 

tourists including many botanists from all parts of the world. 

However, the 'ahinahina . macrocephalum) is only one of six 

species in the genus Argyroxiphium which includes plants called 

greenswords as well as those called silverswords. The silver- 

sword ordinarily produces a basal rosette of leaves for a number 

of years before it finally produces a single massive, elongated 

capitulescence and dies. The floral heads of sword plants are 

characteristically large and provided with rays. They are found 

on Hawai'i and Maui, growing in cinder or, as in the case of 

A. virescens Hbd., in boggy areas like greensword meadow in the 

upper Hana rain forest on Maui. 

Wilkesia, the iliau, is a related bitypic genus which has a 

growth pattern similar to that of sword plants. It is endemic to 

dry slopes in and near Waimea Canyon, Kaua'i. 

In marked contrast to Argyroxiphium the related Hawaiian 

endemic genus Dubautia ( including Railliardia) is comprised 

largely of woody, branching shrubs that produce small, rayless 

heads year after year. However, the genus shows a truly remarkable 

spectrum of variation from largely herbaceous, low-growing 

forms like - D. scabra (DC.) Keck in dry, pioneer habitats as well 

as rain forest situations through woody shrubs or small trees 

like - D. linearis (Gaud.) Keck and D. arborea (Gray) Keck in dry 

sites; and - D. raillardioides Hbd. ana D. knudsenii Hbd. in wet 

sites to larger trees li'ke - D. reticulata (Sherff) Keck in the 

vicinity of Pu'u Nianiau. One striking species (g. latifolia 

lGravl Keck) from Kaua'i is a larae liana with a basal diameter 

if up'to three inches. Altogether there are about 25 species of 

Dubautia including those called na'ena'e or kupaoa 

tively they are distributed from Kaua'i to Hawai'i. 

and collec- 

Field Observations 

Dubautia scabra is a widespread species with considerable 

ecological amplitude and thus comes into contact with several 

other species of Dubautia. Often the result is spontaneous 

hybridization as is the case in the upper Hana rain forest. Here 

the diminutive - D. scabra is sympatric with an undescribed large

shrubby Dubautia and numerous hybrids between the two have become 

established. These hybrids have been given at least four names 

(Railliardia ternifolia Sherff, R. thyrsiflora Sherff var. cernua 

Sherff, R. coriacea Sherff, an3 R. demissifolia Sherff var. 

dolichopriylla St. John). The hybrid plants have an intermediate 

morphology and pale lemon-yellow flowers, a color that readily 

distinguishes all known hybrids involving the white-flowered 

- D. scabra. 

A second hybrid combination occurring in large numbers also 

involves D. scabra. In several areas on Hawai1 i, including 

Hawaii volcanoes National Park, D. ciliolata (DC.) Keck and 

- D. scabra are sympatric. In these instances hybrids between the 

twp have invariably been found. The hybrid morphology is intermediate 

between the shrubby D. ciliolata and the subherbaceous 

- D. scabra and again the flower color is pale lemon-yellow. These 

plants have been referred to as - D. ciliolata var. laxiflora (DC.) 

Keck. 

A third instance of hybridization involving D. scabra occurs 

in the vicinity of Putu Nianiau, Maui, where one Fybrid with the 

large tree, - D. reticulata, has been detected. The hybrid is a 

very diffuse, spreading plant about 2 m tall. As one would 

expect it also has pale lemon-yellow flowers. 

In the same area, within 50 m of the previous hybrid occur 

two hybrids between D. scabra and D. plantaginea Gaud., a large 

shrubby, wide-leaved species. The hybrids are somewhat pendulous 

and viney with ascending shoot tips and lemon-yellow corollas. 

They have been given the name Railliardia lonchophylla Sherff 

var. stipitata (Sherff) Sherff and I suspect that plants of this 

hybrid combination also occur elsewhere and have been ascribed 

other names. 

Only one hybTii3 Dubautia combination has been documented 

from O'ahu. Two individuals of the combination D. sherffiana 

Fosb. x D. lanta inea have been detected in wideiy separated 

locations -i1anae Mts. Both parents are shrubs with 

orange-yellow flowers and as one might expect, the hybrids are 

not as morphologically distinct as in the previous cases. 

The most spectacular instance of hybridization in this com- 

plex occurs in Haleakala on Maui. There, Argyroxiphium macro- 

cephalum, an essentially monocarpic rosette plant with large, 

radiate heads hybridizes spontaneously with the scrubby, woody 

Dubautia menziesii (Gray) Keck having small rayless heads that 

are produced annually. The hybrid is somewhat intermediate in 

appearance (cf. Kobayashi, 1973). It produces tufts of leaves 

usually on three or more branches, each of which eventually 

flowers and dies independently. The hybrid has heads of inter- 

mediate size with small yellowish distorted rays.

Cytology 

The difference between this and other studies of hybridization 

involving Hawaiian taxa is that in these instances hybrids 

can be identified with certainty through an analysis of chromosome 

pairing during meiosis in floral buds. In each of these 

cases the parents are differentiated chromosomally and these 

differences can be positively detected at meiosis. In every 

hybrid except D. scabra x g. plantaginea the diploid chromosome 

number is 2n 27, indicating in these instances that one parent 

furnished 13 chromosomes and the other parent furnished 14. In 

all cases the chromosome numbers of the parents are consistent 

with this argument (cf. Carr, 1978). Although D. plantaginea and 

- 

D. scabra both have 14 pairs of chromosomes, hyErids between the 

two can be readily recognized at meiosis by virtue of the fact 

that their genomes are differentiated by two reciprocal chromo- 

some translocations resulting in the appearance of two chains of 

4 chromosomes each (cf. Table 1). 

These meiotic perturbations result in the inviability of 

some of the gametes formed. This depression of fertility can be 

assessed by the staining reaction of pollen grains in certain 

dyes like cotton blue. Genetic differentiation between parents 

can also cause low viability and thus low pollen stainability in 

hybrid plants. Pollen stainability in the hybrids discussed 

herein ranges from 6 to 86% (Table 1). 

Conclusion 

In spite of the spectacular morphological and ecological 

diversity exhibited by these genera, the occurrence of many 

intergeneric, intersubgeneric, and interspecific hybrid combi- 

nations under field conditions attests to the fact that they form 

a thoroughly natural, genetically cohesive group that has in all 

probability resulted from rapid evolutionary differentiation of a 

single colonizing progenitor. Collectively, these plants consti- 

tute what may be considered an unparalleled example of adaptive 

radiation and as such are exceedingly interesting to students of 

evolutionary phenomena. 


Carr, G. D. 1978. Chromosome numbers of Hawaiian flowering 

plants and the significance of cytology in selected taxa. 

Amer. J. Bot. 65: 236-242. 

Kobayashi, H. K. 1973. Putative generic hybrids of 

Haleakala' s silversword and kupaoa (Argyroxiphium 

sandwicense x Dubautia menziesii) Compositae. Pac. Sci. 

27: 207-208.

TABLE 1. Dubautia and Argyroxiphium spontaneous hybrids. 

Combination 

Number 

ExaminedfSeen 

- D. scabra x - D. n. sp. 

5 /many 5 7 

- D. scabra x - D. ciliolata 

12/many 76 

D. scabra x D. reticulata 

- - 111 86 

- D. scabra x - D. plantaginea 

112 35 

D. plantaginea x D. sherffiana 

- - 212 4 1 

- A. macrocephalum x - D. menziesii 2/14 6 

Pollen Diploid (2n) 

Meiotic 

Stainability Chromosome Location 

Configuration 

% 

Number 

27 1211 + Ch3 Maui 

27 lZII + Ch3 Hawai ' i 

2 7 lZII + Ch3 Maui 

28 loII + 2 Ch4 Maui 

27 loII + Ch4 + Ch3 O'ahu 

27 gII + 3 Ch3 Maui

HAWAII IBP SYNTHESIS: 

6. GENETIC VARIATION AND POPULATION STRUCTURE IN ISLAND SPECIES* 

Hampton L. Carson 

Department of Genetics 



Evolution is an extremely active process on islands, even in 

highly isolated archipelagos like Hawai'i and Galapagos. Species 

formation is frequently exuberant and, in many cases, wholly 

novel adaptations have developed. At first glance, it seems 

remarkable that this should be true because it is clear that in 

many relevant instances the lineage which evolves actively in the 

islands can be traced ultimately to one or only a few continental 

founder individuals. Even more founder effects appear to occur 

as new species are formed within an archipelago. On the surface, 

this appears to be a system which might deplete genetic vari- 

ability and thus reduce evolutionary potential. Precise data on 

genetic variation within island species of Drosophila flies have 

been accumulating now for about 15 years. Modern methods of 

analysis of genetic variability (biochemical, chromosomal, and 

polygenic) have been employed to assay genetic variability in 

various endemic and introduced species. Virtually without excep- 

tion, these species have local populations which are fully as 

polymorphic genetically as those of widespread continental 

species. Accordingly, such populations are highly competent for 

adaptive evolution. Most island species, however, have small 

total populations. Thus, even though the local populations may 

be rich in genetic variability, the total variability sequestered 

within continental species is certainly much larger, but this 

does not appear to be a crucial difference. In high-altitude 

archipelagos like Hawai'i, many factors promote isolation. The 

serial isolations to which island populations are subjected have 

a profound effect on their genetic structure. This is true not 

only for populations between islands (species formation is by 

interisland founder effects) but also for local populations 

within an island or even a volcano. 

As long ago as 1932, Sewall Wright proposed that the condi- 

tions most favorable for rapid evolutionary change exist within a 

species which is subdivided into local semi-isolated populations, 

some of which have quite small effective sizes. Many island 

species appear to reflect precisely this dispersive type of popu- 

lation structure and this may explain their observed evolutionary 

momentum. 

In Addition to IBP support, this work has been supported by 

grants GB 27586, 29288, and BMS 22532 from the National Science 

Foundation. 

Abstract

SOME ASPECTS OF A SHELL DISEASE IN THE 

HAWAIIAN FRESHWATER SHRIMP, ATYA BISULCATA (RANDALL) 

John G. Chan 

Department of Biology 

University of Hawaii at Hilo 

Hilo, Hawaii 96720 

INTRODUCTION 

A disease syndrome characterized by dark necrotic areas of 

the exoskeleton has been reported in a number of aquatic crusta- 

ceans (Rosen 1970). The syndrome has been called black spot 

disease, brown spot disease, rust disease, burnt disease, and 

shell disease. Shell disease was reported in the American 

lobster, Homarus americanus, by Hess (1937). The disease in this 

species was thought to be promoted by crowded condition of the 

lobster holding pens. Similarly, Alaskan king crabs, Para- 

lithodes sp., often developed necrotic "rust" spots when h e l d n 

captivity. The "rust" appeared to be superficial infections 

occurring most commonly on the ventral body area at natural 

breaks and abraded surfaces (Bright, Durham, & Knudsen 1960). 

Rosen (1967) described a necrotic shell disease of American blue 

crab, Callinectes sapidus, collected from shedding pens in 

Crisfield Harbor, Maryland. The crabs had been crowded into 

enclosures until ecdysis yielded the more desirable "soft-shell" 

crabs. While the shell disease syndrome is most discernible 

among crustaceans held in captivity, its occurrence has been 

reported among crustaceans in their nat-ur-a1 .habitat-. More ~ ~ - (1969) 

~ -~ 

reported severely eroded carapaces among blue crabs, - C. sapidus, 

caught in Galveston Bay, Texas. And recently Iversen and 

Beardsley (1976) reported shell disease among marine crustaceans 

of South Florida. They found pitted, darkened carapaces among 

the commercially important stone crab Menippe mercenaria, and 

several other species. 

Shell disease has also been reported in a freshwater prawn, 

Macrobrachium rosenbergii (Sindermann 1974) and is believed to 

occur among pond reared specimens in Hawai'i (R. Nakamura, pers. 

comm. ) . The presence of shell disease in M. rosenbergii is not 

surprising since Kubota (1972) reported a severe necrotic shell 

disease in a closely related species, the Tahitian prawn, - M. - lar 

in Kahana Stream on O'ahu. 

. , 

This study describes the shell disease syndrome in the 

Hawaiian freshwater atyid shrimp, Atya bisulcata, with emphasis 

on the frequency of occurrence, the nature of the lesion, and its 

etiology.

MATERIALS AND METHOD 

Collection and Examination of Shrimp 

Shrimp specimens were captured from various coastal streams 

in an area bounded by Hilo to the south and Waikamalo Stream to 

the north. In this area, characterized by high annual rainfall, 

streams run throughout the year and support some of the largest 

populations of atyid shrimps on the island. The majority of our 

specimens were collected from the Wailuku River, which borders 

Hilo, and Pukihae Stream a few miles to the north. 

In the Wailuku River with its deep pools, shrimps were 

captured by diving with hand-held scoop nets of 0.25-inch mesh. 

In smaller, shallow streams, the net was simply held underwater 

and rocks immediately upstream were overturned. The dislodged 

shrimps were then swept into the nets and captured. Specimens 

were transported back to the laboratory in Styrofoam chests for 

examination before returning them to the stream. In some cases, 

specimens were preserved in 2% formalin for later examination. 

Examination of Specimens for Lesions 

All specimens brought into the laboratory were initially 

examined with a Nikon SMZ Stereomicroscope at 10X magnification 

with an attached ring lamp for illumination. Live specimens had 

to be cooled in ice water at 5-7OC to facilitate handling of the 

otherwise very active shrimp. The numbers, size, and location of 

lesions were noted along with the size and sex of each shrimp. 

Specimens selected for scanning electron microscopy (SEM) were 

dissected, dehydrated in an alcohol/acetone series and followed 

by critical-point drying in amyl acetate or acetone to minimize 

shrinkage and distortion (Hayat 1978). Specimens were then 

mounted on aluminum stubs with colloidal silver paint and coated 

with evaporated gold. Examination was made with an ETEC Autoscan 

Model U-1 at 20 kv accelerating voltage. 

Isolation of Chitinoclastic Bacteria 

Lesions selected from newly captured shrimp were aseptically 

excised and macerated in a sterile mortar and pestle to form a 

fine slurry. The resultant slurry was diluted 10-fold in sterile 

water and a 0.1 ml aliquot inoculated onto chitin overlay agar by 

the spread plate method. Inoculated plates were incubated at 

25OC for a minimum of a week and observed for signs of chitin 

digestion. Chitinoclastic colonies showed a clear halo around 

the colony as a result of the disolution of the opaque chitin 

particles (Skerman 1959). Colonies showing distinct chitino- 

clastic activity were isolated, purified, and maintained on 

nutrient agar.

Induction of Lesions 

To induce lesions, specimens were abraded carefully to 

remove just the epicuticle. This was accomplished by using a 

high speed hand drill fitted with a fine abrasive bit. To facil- 

itate handling, the shrimps were first cooled in ice water at 

5-7 OC for 10 min and strapped onto a polyethylene foam sheet 

(Nalgene) with nichrome wire hoops. Just prior to abrading, the 

shell was swabbed with 70% ethanol. 

Abraded animals were then rinsed in tap water and trans- 

ferred to 500 ml of sterile stream water in 1-liter beakers and 

covered with aluminum foil. Usually, 3 to 5 animals were con- 

tained in each beaker. A buffered washed suspension of a 48-hour 

culture of chitinoclastic bacteria was added to each beaker to 

achieve an initial cell density of about 105 cells per ml. As a 

control, an identical beaker was used without inoculated bac- 

teria. Also, a control was set up with M thimerosal added 

as a bacterial inhibitor. 

To determine whether cuticle abrasion was necessary for 

lesion formation, the wounds of some specimens were sealed imme- 

diately after abrasion. These specimens, sealed with Cutex clear 

nail polish, were subjected to the same incubation conditions as 

specimens with fresh abrasions. 

The test animals were maintained in their containers at ca. 

25OC for up to a month with inspection for lesions initiating 

after one week. 

RESULTS AND DISCUSSION 

A total of 3423 specimens were collected from various Hilo 

coast streams. Lesions were found in 582 (17%) of the specimens. 

Table 1 is representative of lesion incidence found in three 

streams during March 1976. The Wailuku River site yielded the 

highest incidence of lesions. Ka'ie'ie Stream, the smallest of 

the three, had the lowest incidence. Of the three streams, 

Wailuku River sites consistently yielded the highest incidence of 

shell lesions. This may be related, in part, to the physical- 

chemical characteristics of the stream. The greatest volume, 

water velocity, and silt load are found in the Wailuku River. 

High silt load and lesion incidence might be related since it has 

been suggested that abrasive action in the environment could lead 

to lesion formation (Bright et al. 1960). 

The data in Table 1 also shows a significantly higher pro- 

portion of female shrimp developing lesions. Not only did female 

shrimp develop lesions more frequently, they also had more 

lesions per individual. In this sample period, some females had 

as many as 30 lesions. Almost all had more than one. By 

contrast, males often had but a single lesion.

This difference may be explained in part by the observation 

that most of the females were bearing eggs. During tneir 

ovigerous condition, the females do not molt. Hence, lesions 

have a longer period in which to develop and to become visible. 

By contrast, the males have no such constraint and molt more fre- 

quently. Upon ecdysis, the animal is generally freed of its 

lesions. In some individuals, however, previous infection is 

evidenced by a "scar" or a deformation of the new exoskeleton at 

the site of the former lesion. This deformed cuticle is often 

the site of subsequent infection. 

Extensive sampling of Wailuku River sites revealed a defi- 

nite seasonal trend in lesion incidence. While lesions occurred 

among specimens throughout the year, the highest incidences were 

found in the winter months. A peak of 92% lesion occurrence was 

reached in February when the water temperature was 14OC. The 

lowest incidence of 4.7% was recorded in July when water temper- 

ature was 24OC. It appeared that the trend in lesion occurrences 

was inversely related to stream temperature. This unexpected 

observation may be explained by the fact that lower water temper- 

ature would retard the rate of molting, i .e., the intermolt 

period was lengthened. Again, longer retention of the exo- 

skeleton allows for greater development of the necrotic lesions. 

The distribution of lesions on the body surface of 

A. - bisulcata is shown in Table 2. The areas most frequently 

affected were the cephalothorax and the abdominal segments. This 

is not surprising since these two body parts offer the greatest 

exposed surface areas. What is surprising is that the vast 

majority of lesions occurred on the dorsal or lateral surfaces of 

the shrimp. Rosen (1967) had shown the ventral surfaces of the 

blue crab to be the site of intense necrotic lesions. Similarly, 

Bright et al. (1960) attributed the intensity of lesions occurring 

on the ventral surfaces of Alaskan king crab to mechanical 

abrasion by the substrate. It appears that the ventral surfaces 

and appendages of A. bisulcata are relatively protected from 

abrasive water-borne silt particles and are, therefore, less 

susceptible to damage and lesion formation. 

Current opinion indicates that exoskeletal lesions in 

crustacea are in part due to chitin-digesting microorganisms-- 

particularly bacteria. Rosen (1970) and Cook and Lofton (1973) 

have suggested a causal role for chitinoclastic bacteria. In 

their studies, the bacteria with chitin-digesting ability had 

been isolated from diseased hosts. However, controlled reinfec- 

tion experiments were lacking. In this study, chitinoclastic 

bacteria were consistently isolated from lesions of A. bisulcata 

(Table 3). For comparison, lesions of M. lar from the same 

stream were also examined and found to consistently yield chitin- 

oclastic bacteria. It was found that even the "normal" cuticle 

Of the atyid shrimp would occasionally yield chitinoclastic 

bacteria. The chitinoclastic bacteria isolated from normal 

cuticle may have been associated with undetected early stage

lesions. Or they may be part of the normal microflora. Chitino- 

clastic bacteria are anything but rare in the stream environment. 

At times they reached over 50,000 per ml of stream water and made 

up a substantial proportion of all the bacteria present in 

streams we sampled. 

The chitinoclastic bacteria isolated from A. bisulcata were 

of two types. The majority of isolates were gram negative rods, 

motile by polar flagella, facultatively anaerobic, and glucose 

fermentors. These isolates appear to fit the description of the 

genus Beneckea as proposed by Baumann, Baumann, and Mandel 

(1971). The other isolate was characterized by bright orange 

colonies on agar plates and consisted of gram negative, slender 

rods with gliding motility. The gliding motility is typical of 

members of the genus Cytophaga. Both the Beneckea and Cytophaga 

type of isolates showed stronq chitinoclastic activity under 

aerobic condition, but none when incubated anaerobically. 

To establish the etiology of the necrotic lesions, pure cul- 

tures of chitinoclastic bacteria were used in reinfection experi- 

ments. A particularly active Beneckea type, designated WCh-1, 

was used for the induction of lesions. Table 4 shows that 

A. bisulcata, abraded to damage the outer cuticle surface, always 

xeveloped necrotic lesions when confined in a system with abun- 

dant chitinoclastic bacteria. Likewise, abraded shrimp confined 

in raw stream water also consistently developed lesions. Water 

as taken directly from the stream always contained numerous 

chitinoclastic bacteria. It was believed that these autoch- 

thonous bacteria serve as an infectious reservoir in the stream. 

When stream water was sterilized ( M i l l ipore membrane filter , 

0.45 ,pm pore) or thimerosal added as a bacteriocidal agent, 

necrotic lesion formation was markedly suppressed. In the few 

instances where lesions did form in supposedly "sterile" condi- 

tions, chitinoclastic bacteria were subsequently found. It 

appears that residual fecal pellets were a source of the bacteria 

which contaminated the system and overwhelmed the suppressive 

capability of the thimerosal. 

To show that epicuticular damage was indeed necessary for 

the initiation of lesions, as suggested by Rosen (1970), abraded 

test animals were compared to similarly treated specimens in 

which the abrasions were sealed water tight with clear nail 

polish. The results in Table 5 show a striking difference. 

Damaged cuticle when exposed to raw stream water containing 

chitinoclastic bacteria invariably developed necrotic lesions. 

Those with sealed wounds never developed lesions unless the seals 

were faulty and leaking. 

In all attempted isolations, the artificially induced 

lesions yielded chitinoclastic bacteria--almost exclusively of 

the Beneckea type. Scanning electron microscopy of the naturally 

occurring lesion and artificially induced lesion showed a remark- 

able similarity. In both cases, extensive erosion of the cuticle 

is evident and large aggregations of bacteria are seen. By con- 

trast, areas free of abrasive damage or erosion are essentially

free of bacteria. The activity of these bacteria on the chi- 

tinous substrate very closely resembles that described by Akin 

and Amos (1975) for cellulolytic bacteria and their degradation 

of cellulosic plant material. In both cases, the production of 

exoenzymes is likely responsible for hydrolytic degradation of 

the substrate. 

This study has shown that necrotic exoskeletal lesions are 

common to the endemic Hawaiian freshwater shrimp, A. bisulcata. 

The causal agent of these lesions have been shown to-be chitino- 

clastic bacteria which are ubiquitous in the stream environment 

and probably a component of the shrimp's microflora. Stream con- 

ditions which lead to damage of the cuticle--primarily the 

epicuticle layer--exposed the cuticle to bacterial invasion. 

Establishment of chitinoclastic bacteria on the cuticle and sub- 

sequent degradation of the chitinous substrate result in forma- 

tion of a necrotic lesion. The dark coloration characteristic of 

these necrotic areas is due to melanization of hemocytes which 

aggregate underneath the developing lesion. 

ACKNOWLEDGEMENTS 

The author wishes to acknowledge the support provided by the 

National Institute of Health Minorities Biomedical Support Grant 

No. 5 SO6 RR 08073-06 GRS and an additional grant from the 

Environmental Center, University of Hawaii. I am grateful to 

Messrs. Alfred Menino, Jr., Jason Moniz, Mark H. K. Greer, Rory 

Murata, and Miles Nagata for their assistance in various phases 

of this study. Mr. Ken Lesch, San Francisco State University, 

provided invaluable assistance in scanning electron microscopy.


Akin, D. E., and H. E. Amos. 1975. Rumen bacterial degradation 

of forage cell wall investigated by electron microscopy. 

Appl. Microbiol. 29: 692-701. 

Baumann, P., L. Baumann, and M. Mandel. 1971. Taxonomy of 

marine bacteria: The Genus Beneckea. J. Bacteriol. 107: 

268-294. 

Bright, D. B., F. E. Durham, and J. W. Knudsen. 1960. King crab 

investigation of Cook Inlet, Alaska. Bur. Commercial Fish. 

Lab., Auke Bay, Alaska. Mimeo Contr. Rep. 180 pp. 

Cook, D. W., and S. R. Lofton. 1973. Chitinoclastic bacteria 

associated with shell disease in Penaeus shrimp and the blue 

crab, Callinectes sapidus. J. Wildlife Diseases 9: 154-158. 

Hayat, M. A. 1978. Introduction to Eiological Scanning Electron 

Microscopy. Univ. Park Press, Baltimore. 323 pp. 

Hess, E. 1937. A shell disease in lobster (Homarus americanus) 

caused by chitinovorous bacteria. J. Biol. Bd. Canada. 

3: 358-362. 

Iversen, E. S., and G. L. Beardsley. 1976. Shell disease in 

crustaceans indigenous to South Florida. Progr. Fish. Cult. 

38: 195-196. 

Kubota, W. T. 1972. The biology of an introduced 

Macrobrachium - lar (Fabricius) in Kahana Stream. 

prawn 

M. S. 

Thesis, University of Hawaii, Honolulu. 185 pp. 

More, W. R. 1969. A contribution to the biology of the blue 

crab, Callinectes sapidus, Rathbun in Texas, with a 

descri~tion of the fisherv. 

Tech. 5eries No. 1. 31 pp. 

Texas Parks and Wildlife Dept., - 

Overstreet, R. M. 1973. Parasites of some penaeid shrimps with 

emphasis on reared hosts. Aquaculture 2: 105-140. 

Rosen, B. 1967. Shell disease of the blue crab, Callinectes 

sapidus. J. Invert. Pathol. 9: 348-353. 

. 1970. Shell disease of aquatic crustaceans. Pages 

409-415 S. F. Snieszko, ed. A symposium on disease of 

fishes and shellfishes. Am. Fish. Soc., Spec. Public. 

NO. 5. 

Sindermann, C. J. 1974. Diagnosis and control of mariculture 

diseases in the United States. National Marine Fisheries 

Service. Tech. Ser. No. 2. December 1974. 306 pp. 

Skerman, V. B. D. 1959. A guide to the identification of the 

genera of bacteria. Williams and Wilkins Co., Baltimore. 

217 pp.

TABLE 1. Occurrence of exdskeletal lesions in Atya bisulcata 

from selected Hawai'i Island streams (% in paren- 

theses). 

Number of Specimens Specimens with Lesions 

Sampling Site Total Female Male Total Female Male 

Wailuku River 242 30 212 163 27 136 

(67.4) (90.0) (64.2) 

Ka'ie'ie Stream 243 45 198 17 9 8 

(7.0) (20.0) (4.0) 

Pukihae Stream 363 146 216 135 6 4 7 1 

(37.3) (43.8) (32.9) 

TABLE 2. The location of exoskeletal lesions in Atya bi- 

sulcata from Wailuku River, Hawai'i. 

Body Part 

Cephalothorax 

Abdomen Segments 

Telson & U-ropod 

Walking Legs 

Swimmerets 

Total. 8 0 

Number* Percent of 

of Lesions Total 

* 36 lesions on left side, 30 lesions on right side, and 

14 lesions located medially

TABLE 3. Isolation of chitinoclastic bacteria from exoskeletal 

lesions. 

Number of Lesions Yielding Chitinoclasts 

Lesions 

Source Sampled Number Percent 

bisulcata 

lesion 

- 

Macrobrachium lar 

lesion 

A. - bisulcata 

normal cuticle 

TABLE 4. Induced exoskeletal lesions in Atya bisulcata. 

Total Percent 

Test Conditions Specimens With Lesion Without Lesion 

Chitinoclastic 

Bacteria in sterile 

stream water 3 3 

Raw stream water 16 

Sterile stream water 

with Thimerosal, ~ o - ~ M 20 

TABLE 5. The development of exoskeletal lesions in Atya bisul- 

cata with abrasions. (Specimens were held in raw 

=am water at 25OC for a minimum of 8 days.) 

Total Number Number of Specimens 

Treatment of Specimen Developing Lesions Without Lesions 

Abraded 

only 

Abraded and 

sealed 15

THE DISTRIBUTION OF MYRICA FAYA AND OTHER SELECTED PROBLEM 

EXOTICS WITHIN HAWAI 

NATIONAL PARK* 

Garvin Clarke 



Myrica faya was introduced to the Hawaiian Islands sometime 

in the late 1800's. In the mid-1940's the Territorial government 

had recognized the plant as an aggressive weed and a threat to 

pasture land. Intense eradication measures and bio-control 

efforts were instigated by the Territorial and State governments 

throughout the years, but by 1962 infestation had affected over 

21,000 acres with over 80% of the acreage situated on Hawai'i 

Island. Recently, M. faya has come of interest to Hawaii 

Volcanoes National ~arF as a possible threat to native forest 

systems, and is now being evaluated for various control methods. 

A survey of - M. faya distribution within the Park was undertaken 

1978. 

during a seven-week period from December 1977 to January 

Data collection was accomplished through direct field 

observation, helicopter surveillance, and to a lesser degree 

aerial photographs. 

Findings revealed that M. faya distribution constitutes a 

horizontal band between 2100 feet and 4000 feet elevation from 

Namakanipaio southeast to Panau. Infested areas were mapped and 

estimated densities established. Information concerning dis- 

persal mechanism, germination rates, control measures, and hab- 

itat diversity are noted. Other aggressive exotics were recorded 

as observed during g. faya mapping. 

M. faya was detected to be a very formidable foe to the 

native ecosystems within the Park. The ability of this tree to 

cover large areas within a matter of years is recorded. A real- 

istic approach coupled with applicable research is needed in an 

effort to understand and control this problem. 

* Abstract

FOREST BIRD POPULATIONS ON O'AHU 

Mark S. Collins1 and Robert J. Shallenberger2 

For the past three years Ahuimanu Productions has conducted 

various environmental assessments that have led to a greater 

understanding of the abundance and distribution of O'ahu's forest 

birds. Beginning in June of 1976 with a brief survey of South 

Halawa Valley, a series of four avifaunal surveys was conducted 

in consideration of the alternate routes for the proposed H-3 

trans-Ko'olau highway (1, 2, 3, 5). The most intensive as well 

as extensive survey of the series was initiated on December 19, 

1977, and was completed on March 9, 1978. This most recent sur- 

vey involved 200 man-days in the field and was conducted in eight 

central Ko'olau valleys and ridges extending from Moanalua Valley 

in the south to Poamoho Trail in the north. In addition a bird 

and mammal survey of Army lands on O'ahu and Hawai'i was con- 

ducted during 1976 to 1977 under contract to the Army Engineers 

Division, Pacific Ocean (5). As a part of this contract, a 

significant portion of the Northern Ko'olau Range and a less 

extensive area in the Wai'anae Range were surveyed for forest 

birds. The compilation of these avifaunal surveys represents the 

most intensive study of O'ahu's forest birds ever conducted. 

In addition to extensive data on abundance and distribution 

of O'ahu's common forest birds, significant information gathered 

from these avifaunal studies include: 

Observations that confirm the substantial range exten- 

sion of three exotic species (Vanikoro Swiftlet, 

Red-vented Bulbul, Yellow-faced Grassquit) (3, 4, 5); 

Three separate observations of the endangered O'ahu 

Creeper in the central Ko'olau Range (5); 

The first nesting record for the Vanikoro Swiftlet in 

Hawai'i (5); 

The probable sighting of a female O'ahu 'Akepa at the 

summit of Schofield-Waikane Trail (4) ; 

U. S. Forest Service, Hawaii Volcanoes National Park, Hawaii 

96718. 

2 President, Ahuimanu Productions, Kailua, Hawaii 96734.

5) The observation of the "Mystery Garrulax" on Poamoho 

Trail, a bird that had not been seen in nearly two 

decades (5); 

6) The discovery of a substantial population of I i w i in 

the Wai'anae Range below Ka'ala (4). 

Copies of these contracted survey reports are not currently 

available for mass distribution. However, the compiled data are 

now being prepared for future publication. 

Shallenberger, R. J. 1976. Avifaunal survey of South 

Halawa Valley. Prepared under contract to Parsons, 

Brinkerhoff-Hirota Assoc. (June 1976). Unpublished. 

Shallenberger, R. J. 1976. Avifaunal survey of North 

Halawa Valley. Prepared under contract to Parsons, 

Brinkerhoff-Hirota Assoc. (Sept. 1976). Unpublished. 

Shallenberger, R. J. 1977. Avifaunal survey of North 

Halawa Valley, Oahu. Prepared under contract to 

Parsons, Brinkerhoff-Hirota Assoc. (August 1977). 

Unpublished. Appears in reference no. 6 (Vol. 5). 

Shallenberger, R. J. 1977. Bird and Mammal study of 

Army lands in Hawaii. Prepared on contract to the U. S. 

Army Corps of Engineers. Ahuimanu Productions, Kailua. 

598 pp. (3 Vols.). 

Shallenberger, R. J. 1978. Avifaunal survey of the 

central Koolau Range, Oahu. Prepared under contract to 

Parsons, Brinkerhoff-Hirota Assoc. (April 1978). 

106 pp. Unpublished.

LEK BEHAVIOR AND ECOLOGY OF TWO 

HOMOSEQUENTIAL SYMPATRIC HAWAIIAN DROSOPHILA : 

DROSOPHILA HETERONEURA AND DROSOPHILA SILVESTRIS 

Patrick Conant 




According to Loiselle and Barlow (1978), "Lekking is the 

temporary synchronus aggregation of sexually active males for 

reproduction." It is a type of communal display. Theoretically, 

males displaying communally are more conspicuous and have a 

better chance of encountering females. Until recently, most of 

the research on lek behavior has been done on birds, especially 

grouse (Tetraonidae) and mannikins (Pipridae). Lekking in fish 

has only begun to be investigated (Loiselle & Barlow 1978) and 

reports of work on insect lek behavior are not numerous (Wilson 

1975). 

Spieth (1968) was the first to describe lek behavior in 

Hawaiian Droso hila Two species of Hawaiian 

Spieth foun &. to engage in . lek behavior are 

and - D. silvestris. Carson and Stalker (1968) found these two 

species to be homosequential, that is, the banding patterns on 

the polytene chromosomes are identical, indicating they are very 

closely related. They also found that heteroneura and silvestris 

share a polymorphic inversion, although silvestris has six other 

paracentric inversions. 

Ahearn et al. (1974) found that in the laboratory, etho- 

logical isolation between heteroneura and silvestris was strong 

althouqh incomplete. Unfortunately, quantitative comparisons of 

the action patterns observed in the courtships of these two 

species have not been made. Spieth (pers. comm.) has, however, 

found some qualitative differences. 

Hybrid progeny of successful pairings in the laboratory are 

vigorous and fertile (Carson & Kaneshiro 1976). Recently, 

Kaneshiro has even collected hybrids in the wild at Kahuku Ranch 

on the island of Hawai'i (Kaneshiro & Val 1977). 

The two flies belong to the Planitibia complex of picture 

wing Drosophila and are widely sympatric on the island of 

Hawai'i. The other two species that belong to this complex are 

differens found on Moloka'i and planitibia on Maui. Morpho- 

logical, ethological, and chromosomal data indicate that these 

two flies are ancestral to heteroneura and silvestris (Kaneshiro 

1976). It is most likely that planitibia is ancestral to 

silvestris. D. heteroneura may be derived from planitibia but 

differens is the more probable ancestor.

My objectives in this Study were to observe and describe the 

lek behavior and ecology of these two species, and to determine 

if there were differences in these aspects of their systematics 

that would explain their reproductive isolation in the wild. 

Two primary study areas were established on Kealakekua Ranch 

in Kona to study differences in habitat preference. The lower of 

the two sites (1067 m/3500 ft) , was called Ha~u'u. It is in an 

~ ~ . 

~~~ ~~ 

'ohi'a (~etrosideros sp.) forest with a den& tree fern (Cibotium 

sp.) understory. The higher study site (1304 m/4280 ft), called 

Oiki, was i n a remnant 'ohi'a forest with scattered Acacia koa 

and diverse tree strata interspersed with open pasture. 

- 

Two other sites where observations were made were on Keauhou 

Ranch at 1432 m (4700 ft) near Hawaii Volcanoes National Park 

and Kahuku Ranch near South Point. The habitats whete the flies 

occur on these two ranches are similar. Both sites are open pas- 

ture with small clumps of dense remnant 'ohi'a-tree fern forest. 

Observations were made at two separate elevations on Kahuku 

Ranch, 1058 m (3800 ft) and 1235 m (4050 ft). 

Observations were also made in 'Ola'a forest at 1237 m (4060 

ft) adjacent to the University of Hawaii Volcano Experiment 

Station. This forest is similar to the hapu'u study area; it is 

an 'ohi'a forest with a dense understory of tree ferns. 

My observations of the behavior of heteroneura on food 

supported Spieth's contention that predation by birds acts as a 

selection pressure favoring the evolution of lek behavior in 

Hawaiian Drosophila. That is, courtship is not on the food as 

with most Drosophila, but on branch-territories nearby. 

Mark-release experiments showed that marked flies often 

occupied the same branch when consecutive sightings of them we re 

made. At morning and afternoon sightings, and at afternoon and 

the following morning sightings, marked flies had moved only 

small distances (about 0.5 m). 

Field observations suggested that the morning temperature 

threshold for feeding behavior for both species is about 10°C 

(50°F). Males of both species alight on their branch- 

territories presumably after feeding. Males of both species were 

active at the lek if weather conditions remained optimal. Condi- 

tions outside these limits caused flies to remain inactive or to 

leave their branch-territories. Males of both species leave 

their territories near sunset. Temperature decrease below a 

threshold probably stimulates this behavior if decreasing sun- 

light does not do so first. No temporal differences in atten- 

dencies of the two species at their respective leks were evident. 

The initial response of a heteroneura male to the approach 

of another arthropod moving on its territory is usually an 

approach, and is often followed by patrolling behavior, the 

repeated walking of a distance of 10 to 80 cm over a section of 

vegetation. Patrolling behavior is most often observed (in

either species) after an encounter with another arthropod; how- 

ever, the frequency and duration of walks appears to decay with 

time. While on their territories male heteroneura frequently 

make short flights in an erratic path away from and back to their 

territory. 

Patrolling behavior may perform three different functions. 

The visual stimuli of a patrolling male may serve to keep the 

males together, interacting and competing for territories. It 

may also attract females. Finally, patrolling may be a means by 

which the most fit males may defend territories and insure that 

they inseminate the females that alight there. 

Female heteroneura (and presumably silvestris) are usually 

found at about the same height as the males (about 2 m) resting 

on the undersides of branches; Many hours of observations of 

male heteroneura on their territories revealed that females 

alighted on the territories infrequently. 

Premating isolation can be maintained through two different 

means. Behavioral isolation between two species prevents them 

from exchanging genetic material. Through ecological isolation 

species are kept separate by the influence of environmental 

factors. 

Differences in body color between the two species suggested 

they might be adapted to different levels of incident light under 

the forest canopy. However, the analysis of incident light level 

data measured on flies suggested that flies do not seek different 

light levels in their habitat. Instead the results lend support 

to the hypothesis that differences in body color (particularly 

abdomen color) may be most important as stimuli in mate 

recognition. 

Observations on Kealakekua Ranch, Kahuku Ranch, and 'Ola'a 

forest did not reveal any differences between leks of the two 

species. Heights of male silvestris in vegetation at the 

different sites were all found to be about 2 m off the ground. 

Observations at "shared lek sites" (leks under the same 

continuous canopy in close enough proximity that interspecific 

interactions were possible) did not reveal any differences in the 

leks of the two species at these shared sites. There was no 

species-specific preference for plants on which territories were 

defended. In fact, where branches or fern stipes were long 

enough, I occasionally saw males of both species "on station" 

(Spieth 1966) on the same branch or stipe. At certain of these 

shared lek sites some spatial separation of the two leks was evi- 

dent. One possible explanation at certain sites might be that 

behavioral interactions might cause the flies to congregate where 

they only infrequently encounter the other species on the terri- 

tories. Species-specific pheromones released into the air might 

also serve to keep both sexes of both species together at the lek 

but no evidence of this has been reported. Another resource, 

besides leks, these species shared was food. Often, both species 

were seen interacting on the same food.

Further evidence for resource sharina was found when both 

< 

species were reared from the same decaying Clermontia branches 

collected on Kahuku Ranch. 

The only ecological variables to which the two flies 

appeared to respond to differently were changes in moisture and 

pressure (that is, lapse rate) associated with elevation. 

Results of past collections of these species from different 

elevations on Kahuku and Keauhou ranches, Kilauea Forest Reserve, 

'Ola'a forest, and kipukas on the Saddle Road, suggest that 

silvestris is more common at higher elevations. It is possible, 

however, that silvestris may just have a wider range of eleva- 

tional distribution since it is always found where heteroneura 

occurs. 


Ahearn, J.N., H.L. Carson, T.H. Dobzhansky, and K.Y. Kaneshiro. 

1974. Ethological isolation among three species of the 

planitibia subgroup of Hawaiian Drosophila. Proc. Nat. 

Acad. Sci. 8. S. A. 71: 901-903. 

Carson, H.L., and K.Y. Kaneshiro. 1976. Drosophila of Hawaii. 

Systematics and ecological genetics. Ann. Rev. Ecol. Syst. 

7: 437-543. 

Carsan, H.L., and H.D. Stalker. 1968. Polytene chromosome 

relationships in Hawaiian species of Drosophila. 11. The 

- D. planitibia subgroup. Univ. Texas Publ. 6818: 355-366. 

Kaneshiro, K.Y. 1976. Ethological isolation and phylogeny in 

the planitibia subgroup of Hawaiian Drosophila. Evolution 

30: 740-745. 

Kaneshiro, K.Y., and F.C. Val. 1977. Natural hybridization 

between a sympatric pair of Hawaiian ~roso~hila. Am. 

111: 897-902. 

Nat. 

Loiselle, P.V., and G.W. Barlow. 1978. Do fishes lek like 

birds? - In E.S. Reese, and F.J. Lighter, eds. Contrasts in 

behavior. Wiley-Interscience, New York. 

Spieth, H.T. 1966. Courtship behavior of endemic Hawaiian 

Drosophila. Univ. Texas Publ. 6615: 245-313. 

. 1968. Evolutionary implications of sexual behavior in 

Drosophila. Pages 157-193 in T.H. Dobzhansky, M.K. Hecht, 

and W.M.C. Steere, eds. ~vzutionary biology. Vol. 2. 

Appleton-Century-Crofts, New York. 

Wilson, E.O. 1975. Sociobiology, the new synthesis. Belknap 

Press, Cambridge, Massachusetts. IX + 653 pp.


3. THE KILAUEA RAIN FOREST ECOSYSTEM* 

Sheila Conant 

Department of General Science 



The analysis of this rain forest was focussed on its struc- 

ture and dynamics with regard to both plant and animal life. 

Species were assessed quantitatively according to their life 

forms and ecological roles which they have assumed in the various 

general niches of this forest. 

The structure and behavior of the forest plant community 

w i l l be briefly characterized and the activity patterns of the 

birds, tree arthropods, and introduced mammals w i l l be high- 

lighted. A few conclusions w i l l be drawn as to the maintenance 

trends of this ecosystem under natural conditions. 

* Abstract

BIRDS OF THE KALAPANA EXTENSION 

Sheila Conant 




INTRODUCTION 

In recent years the National Park Service has been working 

to develop a feasible and sound management program for lands 

included in the approximately 49,000 acres of the Kalapana 

Extension (Fig. l), which was acquired by Hawaii Volcanoes 

National Park between 1938 and 1960 (National Park Service 1974). 

The first step in planning analysis has consisted largely of 

inventory-type research aimed at identifying and locating the 

resources of the Kalapana Extension. An up-to-date inventory of 

the avifauna was called for as part of this research. 

Most of the information on birds presented in the Draft 

Planning Analysis (National Park Service 1974: Map D, henceforth 

Fig. 2 of this report) was based on data gathered by Baldwin 

(1953) more than 25 years ago. Surveys by Dunmire (1962) and 

Berger (1972), as well as verbal communications from other 

researchers, provided some additional data. It should be noted 

that the map shows only what were called "unimpaired" bird hab- 

itats. Presumably, some forest birds had and still have today 

more extensive distributions than is suggested by this map. 

MATERIALS AND METHODS 

In 1976 I began field surveys under contract with the Coop- 

erative National Park Resources Studies Unit (CPSU). This report 

summarizes data collected through December of 1977, including 

observations reported to me by other researchers. 

Censuses of birds were conducted using either Emlen's (1971) 

transect count method or Reynolds et al. (in press) variable 

circular plot method. For the rarer species not of ten detected 

during censuses, such as the '10 (Buteo solitarius) , sighting 

locations were recorded. Although I was unable to exhaustively 

survey all parts of the Kalapana Extension, I attempted to inven- 

tory the avifauna of the major habitats, concentrating my field 

work in the closed Metrosideros rain forests near Napau Crater 

where Baldwin (1953) had reported several endangered species. I 

was interested to know if these species still occurred in the 

areas listed as "unimpaired habitats" as of 1974 (Fig. 2).


Fifteen exotic (63%), two indigenous (a%), and seven endemic 

(29%) bird species were observed during this study or by other 

researchers during the study period (Table 1). 

Four native forest birds previously reported in or near the 

Kalapana Extension were not recorded there during this study. 

They are the 'O'u (Psittirostra sittacea), the 'Akiapola'au 

(Hemignathus wilsoni), the H a w a i h a (Loxops coccineus 

coccineus), and the 'I'iwi (Vestiaria coccinea). The first three 

are endanqered species, and all belonq to the Hawaiian Honey- 

creeper Family - ( ~re~inididae). 1; seems 

'Akiapola'au or the Hawai'i 'Akepa may still 

recent sightings of the 'O'u (W. E. Banko, 

'Ola'a Tract and near Park Headquarters and 

that this species could still be found, 

northernmost parts of the Kalapana Ex tension. 

person survey such as mine, the species could 

doubtful that the 

occur there, but 

pers. comm.) in the 

residences suggest 

very rarely, in the 

In a brief, one- 

easily be missed. 

Because the native forest bird distribution plotted in 

Figure 2 included only "unimpaired habitats," it is impossible to 

determine whether or not the extent of native bird distribution 

has really changed. I suspect that present-day native forest 

bird distribution (Fig. 3), regardless of habitat quality, is 

essentially the same as it was in 1974. No doubt volcanic 

activity has reduced bird distribution since Baldwin's (1953) and 

Dunmire's (1962) studies due to habitat destruction. This 

change, as well as extensive alteration in forest habitats 

wrought by feral pig and goat activity and exotic plant invasion, 

may account, at least in part, for the disappearance of several 

forest bird species from the Kalapana Extension. There are vir- 

tually no "unimpaired" habitats remaining. It should be noted 

that the National Park Service (1974) report must have inadver- 

tently omitted Hawai'i 'Oma'o (Phaeornis obscurus obscurus) from 

the forest birds listed in Map D. 

Whether or not absolute densities of native birds have 

decreased or increased is not known because this information has 

not been available previously. If the indigenous Kolea (%vialis 

dominica) and Noio (Anous tenuirostr is melanogenys) are 

included, native birds have been observed in virtually - every type 

of habitat in the Kalapana Extension (Table 2). 

Although rain forests of the Extension appear to be suitable 

habitats for most of the Hawai'i Island forest birds, these areas 

were unusually low in avian diversity, with 'Apapane (Himatione 

sanguinea sanguinea) and Hawai'i 'Oma'o dominating the avifauna, 

although Hawai'i 'Elepaio (Chasiempis sandwichensis sandwich- 

- ensis) and '10 also occurred there. In fact, the Kalapana Exten- 

slon harbors the most extensive population of 'Oma'o to be found 

within Hawaii Volcanoes National Park. It is the only form of 

Phaeornis (Hawaiian thrushes) not yet considered endangered.

Notably and inexplicably absent from rain forests were the 

i w i and the Pueo (w flammeus sandwichensis). I feel these 

species - mav - occur there. but because of their verv low numbers. 

were not observed. Hawai'i 'Amakihi (Loxo s virens virens) were 

present in mesic and dryland forests above -% ~ m t m a t i o n , 

and Pueo and '10 were recorded in lowland scrub. Kolea are 

likely to be found in any open grassy area between approximately 

September and April, and Noio can be seen regularly along the 

coast1 ine . 

The endemic passerines reached relatively low elevations, 

some of which were recorded by ob'servers other than the author. 

Low elevation records for native passerines were: 'Elepaio 400 

feet; 'Oma'o 1600 feet; 'Amakihi 50 feet; and 'Apapane 400 feet. 

East of Mauna Ulu flows, 'Amakihi were observed no higher than 

1600 feet, nor lower than 400 feet, except occasionally 

(J. Jacobi & F. R. Warshauer, pers. comm.). However, west of 

Mauna Ulu flows they were consistently observed in suitable 

habitat, regardless of elevation. While the 'Amakihi does not 

regularly inhabit homogeneous closed rain forest in substantial 

numbers, it can usually be found in openings and edges. This was 

not the case in the forests east of Mauna Ulu flows. Perhaps the 

lack of understory diversity characteristic of much of this feral 

pig-disturbed area reduces available resources for birds, giving 

the much more abundant 'Apapane a competitive edge. This ques- 

tion is in need of further study. 

No specific searches were made for the nocturnal 'Ua'u 

(Pterodroma phaeopygia sandwichensis), which was previously 

reported to nest in Makaopuhi Crater, on the northern border of 

the Extension (U. S. Fish & Wildlife Service 1974). 

The Nene (Branta sandvicensis) was recorded in western por- 

tions of the Extension durinq this studv and by P. Banko (pers. 

comm.) and J. and Z. Jacobi (pers: comm.): P. ~anko (in piep.) 

is presently working with the National Park Service on a new 

recovery program aimed at reintroducing a breeding population of 

Nene to lowland habitats in Hawaii Volcanoes National Park. 

The only widespread endangered species in the Extension was 

the '10, which has been observed in or over most habitats, except 

for the very dry scrub and grassland communities, east of Mauna 

Ulu flows either during this study or by other workers. 

The Japanese White-eye (Zostero s japonicus) was by far the 

most abundant and widely ;------E lstrlbuted bird species (Table 3). 

Only the Cardinal (Cardinalis cardinal is) was as widely distr ibuted 

as the JaDaneSe - White-eve, and it was much less abundant 

A . 

(Table 3). The House Finch (Car odacus mexicanus) was the only 

other exotic bird that was elt --I?-- er very common or widely distributed. 

Several species of game birds (Tables 1 & 3) were observed 

during this study and by J. and Z. Jacobi (pers. comm.). With 

the exception of the Ring-necked Pheasant (Phasianus colchicus), 

recorded from lowland scrub, game birds seem to have entered 

Kalapana Extension habitats from the 'Ainahou Ranch area. These 

species were probably introduced to the Ranch before it became 

part of the National Park.

One exotic bird not found but previously recorded in the 

Kalapana Extension was the Red-billed Leiothrix (Leiothrix 

lutea). This species has undergone a rapid decline, for unknown 

reasons, in its Hawaiian Islands range during the last decade 

(Hawaii Audubon Society 1978). 

CONCLUS ION 

Four species ( including three endangered forms) of native 

birds apparently no longer occur in the Kalapana Extension where 

they were reported about 25 years ago (Baldwin 1953). However, 

all the habitat types in the Extension are still occupied by some 

native bird species, with closed Metrosideros rain forests, open 

Metrosideros forests, including scrub forests, harboring the 

greatest diversities of native birds. The closed Metrosideros 

rain forests provide the best and most extensive habitats avail- 

able in Hawaii Volcanoes National Park for the Hawai'i 'Oma'o, 

the only unendangered form of Phaeornis (Hawaiian thrushes). 

Furthermore, should populations of endangered Hawai'i Island 

forest birds (e.g., 'O'u, 'Akiapola'au, 'Akepa, Hawai'i Creeper) 

stage a comeback in the Park, it would most likely occur in 

extensive tracts of rain forest, such as those found in the 

'Ola'a Tract and the Kalapana Extension. For these reasons 

forest habitats should be managed to protect their integrity. Of 

extreme importance in this regard is the control of feral pigs 

and exotic plants. 

When the natural features of the Kalapana Extension, espe- 

cially those geological and botanical, but also ornithological 

and archaeological, are measured against its questionable value 

as homestead land (e.g., high volcanic risk, unsuitability for 

agriculture and ranching), it seems clear that management of 

these lands should have conservation of natural resources as its 

highest priority. 


I thank June Saito for typing the manuscript. Terry Parman, 

Maile Stemmermann, and Rick Warshauer assisted with field work. 

The National Park Service has been particularly helpful with 

logistics. I appreciate Dr. Cliff Smith's assistance with 

several aspects of the project.


Baldwin, P. H. 1953. Annual cycle, environment and evolution in 

the Hawaiian honeycreepers (Aves: Drepaniidae). Univ. 

Calif. Publ. 2001. 54: 285-398. 

Banko, P. 1978. Nene reintroduction program and research in 

Hawaiian National Parks. In C. W. Smith, ed. Proceedings, 

Second Conf. in Natural Sciences, Hawaii Volcanoes National 

Park. CPSU/UH (Univ. of Hawaii, Botany Dept.). 

Berger, A. J. 1972. Birds of Hawaii Volcanoes National Park. 

Island Ecosystems IRP, US/IBP Tech. Rep. 8. 49 pp. 

Dunmire, W. W. 1962. Bird populations in Hawaii Volcanoes 

National Park. Elepaio 22: 65-70. 

Emlen, J. T. 1971. Population densities of birds derived from 

transect counts. Auk 88: 323-341. 

Hawaii Audubon Society. 1978. Hawaii's birds. Second Edition. 

Hawaii Audubon Society, Honolulu. 96 pp. 

Mueller-Dombois, D., and F. R. Fosberg. 1974. Vegetation map of 

Hawaii Volcanoes National Park. CPSU/UH Tech. Rep. 4. 

(Univ. of Hawaii, Botany Dept.). 44 pp. 

National Park Service. 1974. Draft Planning Analysis, Kalapana 

Extension Homesites, Hawaii Volcanoes National Park. 

Western Region, National Park Service, Dept. of the 

Interior. 43 pp. Appendix of 6 maps. 

Reynolds, R. T., J. M. Scott, and R. A. Nussbaum. A variable 

circular plot method for censusing birds. Condor. (In 

press). 

U. S. Fish and Wildl 

forest birds. 

Division of Fish 

ife Service. 1974. 

U. S. Fish and Wildl 

and Game, Honolulu. 

Hawaii's endangered 

ife Service and Hawaii

TABLE 1. A list of the birds in the Kalapana Extension of Hawaii Volcanoes 

National Park (* = Endangered, E = Endemic, I = Indigenous, X = 

Exotic). 

Scientific Name Vernacular Name Hawaiian Status 

* Branta sandvicensis 

- 

* Buteo solitarius 

Imphortyx californicus 

Francolinus erckel ii 

Francolinus adspersus 

Phasianus colchicus 

Phasianus versicolor 

Pluvialis dminica 

- 

Anous tenuirostris 

melanogenys 

Streptopelia chinensis 

- 

Geopl ia str iata 

- 

Asio flmus 

sanawichensis 

Alauda arvensis 

Garrulax canorus 

Phaeornis obscurus 

obscurus 

Hawaiian Goose Nene 

Hawaiian Hawk '10 

California mail 

Erckel Francolin 

Closebarred Francolin 

Ring-nec ked Pheasant 

Green Pheasant 

Golden Plover Kolea 

Hawaiian Noddy No io 

Spotted D3ve 

Barred Dove 

Hawaiian Ckrl Pueo 

Skylark 

Melodious Laughing-thrush 

Hawai' i %rush ' h a' o

TABLE l--Continued. 

Scientific Name Vernacular Name Hawaiian Status 

Chasienpis sandwichensis 

sandwichensis 

Zosterops japonicus 

Acridotheres tristis 

~oxops virens virens 

Himatione sang 

P uinea 

sangulnea 

Lonchur a punctulata 

Passer dmesticus 

Cardinalis cardinalis 

Carpodacus mexicanus 

Hawai' i ' ~lepaio 

Japanese mite-eye 

-on Ypna 

Hawai' i 'Amakihi 

' Apapane 

Spotted Munia 

House Sparrow 

Cardinal 

House Finch 

'Elepaio E 

X 

X 

'Amakihi E 

' Apapane E

TABLE 2. Densities (birds/4O ha) of native bird species in the different 

vegetation types of the Kalapana Extension (P = T 1 bird/4O ha; 

+ = irregularly present). Vegetation types are based on Mueller- 

Dgnbois and Fosberg (1974). 

Species cM cM(ns) OM oM(C) MD olf s HEAn r Corranents 

'I0 P P P 

Kolea 

Noio 

Pueo 

'Qna' o 23 18 4 

' Elepaio 4 2 2 

' Amakihi + + 5 

'Apapane 100 102 106 

P P P 

P P P 

P P 

P (shoreline) 

above 1600 ft 

above 400 ft 

P 400-1600 ft E of 

Mama Ulu flows 

cM = closed Metrosideros forests, various understory types 

above 400 ft 

cM(ns) = closed Metrosideros forests with native shrub understory 

OM 

OM( C) 

MD 

olf 

s 

HEAn 

r 

= open ktrosideros forests, various understory types 

(includes scrub Metrosideros canmunities) 

= open Metrosideros-Cibotimn forests 

= Metrosideros-Diospyros forests, various understory types 

= open mixed lowland forests 

= mixed lowlard scrub canmunities 

= lowland Eleteropgon-Eragrostis-Andropogon grasslards, sometimes 

with mixed shrubs 

= rocklard canmunities with scattered grasses ard shrubs, includes 

salt spray communities

WLE 3. Densities (birds/40 ha) of introduced bird species in different 

vegetation types of the Kalapana Extension (P = < 1 bird/40 ha; 

+ = irregularly present). Vegetation types are based on Mueller- 

Dombois and Fbsberg (1974). 

Species CM cM(ns) OM oM(C) MD olf s HEAn 

California Quail 1 P 

Erckel Francolin' P 

Closebarred Francolin' P 

Ring-necked Pheasant 

Green Pheasant] 

Spotted dove 

Barred mve 

Skylark 

Melodious Laughing-thrush 

Japanese White-eye 

Cmon Myna 

Spotted Wnia 

Cardinal 

House Finch 

These species probably limited to plant canmunities in the northwestern 

corner of the Kalapana Extension, adjacent to upper 

'Ai~hou Ranch. 

portions of the 

CM = closed Metrosideros forests, various understory types 

cM(ns) = closed Metrosideros forests with native shrub understory 

OM = own Metrosideros forests, various understory types 

( incl-trosideros canmunities) 

oM(C) = open Metrosideros-Cibotiun forests 

MD = Metrosideros-Diospyros forests, various understory types 

olf = open mixed lowland forests 

s = mixed lowland scrub canmunities 

HEAn = lowland Heteropogon-Eragrostis-Andropogon grasslands, sanetimes 

with mixed shrubs

FIGURE 1. A map showing the locations of the main Hawaiian 

Islands, the Island of Hawai'i, Hawaii Volcanoes' 

National Park, and the Kalapana Extension.

01 

FIGURE 2. A map of the known (as of April 1974) "unimpaired habitats" of native forest w 

birds in the Kalapana Extension, according to National Park Service (1974: 

Map D).

FIGURE 3. A map showinq native forest bird distribution in the Kalapana Extension 

acco;ding to-this study. Includes many habitats considered to be impaired 

by feral animals and exotic plants.



BIRDS OF THE CRATER DISTRICT 

Sheila Conant 


and 

Maile A. Stemmermann 

Department' of ' Zoology 



INTRODUCTION 

Field stu, dies of avian distribution and abundance were un dertaken 

in Haleakala National Park between June 1976 and April 

1978. We conducted most of the field work during the summer 

months, but also took several trips into the study area at other 

times of the year to evaluate seasonal changes in bird activity. 

Most species densities for different habitat types have been 

derived from censuses using the transect count method described 

by Emlen (1971 ) , or the circular plot method of Reynolds et al. 

(in press). Some species densities could not be calculated by 

these methods (e.g., game birds, House Finch) : in such cases, we 

estimated dens :ities by averaging census totals per unit area 

covered. 

Although this survey encompassed the entire Crater District, 

certain regions received particular attention due to high density 

or diversity of birds. These areas included scrub habitats in 

the eastern end of the Crater, the Paliku area, the eastern 

boundary of Kaupo Gap, and Pu' u Mamane. 


Twenty-two species of birds representing 14 families were 

found in the Crater District during this survey. Thirteen of 

these species (amrox. . -- 60%) were exotic. Of the native species 

about 30% are endemic to.~aui and two species (the Nene, Branta 

sandvicensis, and the 'U'au, Pterodroma phaeopygia sandwichensis) 

are considered endanqered. Table 1 shows the approximate densities 

of bird species - in five qeneral - habitat types - within the 

Crater District.

Several patterns in avian distribution and diversity are 

apparent from our data. Species diversity in the Crater is 

strongly affected by vegetation patterns. Bird diversity gen- 

erally increases with increasing plant cover, with the lowest 

number of species occurring in the arid western region of the 

Crater, and the highest number occurring in mesic forest, partic- 

ularly those along the eastern boundary of the Crater District. 

In a similar fashion, species diversity on the outer Crater 

slopes increases with decreasing altitude, reflecting a corres- 

ponding increase in vegetative cover and plant species diversity 

down the altitudinal gradient. 

Specific distribution patterns are highly reflective of the 

niche components (especially the feeding niche) of the birds in 

question. Distributions of native and non-native species are 

broadly separable on this basis. Exotic species generally have 

wide distributions; each species may have a distribution encom- 

passing several different habitat types and may be common 

throughout most of its range. This tendency towards wide distri- 

butions shown by exotics is in many cases reflective of their 

generalized ecologies. As Ralph (1978) has found, the broad 

feeding niches of many of these birds enable them to utilize 

diverse habitat types. 

The distributions of two common exotics, the Japanese 

White-eye ( Zosterops j aponicus) and the House Finch (Carpodacus 

mexicanus) are good examples of these patterns. Both species 

occur in habitat types between the extremes of arid scrub and 

grasslands, and wet forests. As shown in Table 1, both species 

occur in fairly high densities even outside their optimal hab- 

itats. The House Finch seems to have a greater ability to use 

marginal habitats than does the White-eye, possibly because of 

its greater flocking tendency and mobility. 

Other exotic species have broad ranges similar to those of 

the _ White-eye and House Fi~n-ch,. but occur i n lower densities. 

These species typically occur in fewer habitat types than their 

more abundant counterparts, and may have more specialized feeding 

habits. Amona the s~ecies showina . such distributions are the 

Ring-necked *pheasant (~hasianus colchicus) , the Mockingbird 

(Mimus polyglottos) , and the Skylark (Alauda arvensis) . Each of 

these birds occurs over a large area of the Crater, but as indicated 

in Table 1, none occurs in very large numbers in any one 

habitat type. 

Some exotic species are rare in the Crater District, either 

because their ranges are expanding into the area, or because they 

are poorly adapted to Crater habitats. The Grey Francolin 

(Francolinus pondicerianus) is an example of the first category: 

it is uncommon in the Park and has localized distribution in the 

west side of Kaupo Gap. The bird is common in Kaupo ranchlands, 

but was not recorded in the Park prior to this study. Other 

- 

exotics, mostly non-game species such as the Melodious Laughingthrush 

(~arrulax canorus) and the Cardinal ( Cardinalis cardinalis) 

have been sighted on a sporadic basis, generally on the 

periphery of the Crater District. These birds are unable to

persist in Crater habitats as yet, possibly due to feeding 

limitations or an inability to cope with the rigorous climatic 

conditions. 

Native species shou distribution trends similar to the exo- 

tics, although they show more restricted distributions and are 

often uncommon outside limited areas. The dietary specificity of 

the native forest birds limits them to small areas of . suitable 

habitat. Table 1 illustrates the restricted ranges of these 

birds. The generalized native forest birds (e.g., the 'Amakihi, 

[Loxops virens wilsoni] and the 'Apapane [Himatione sanguinea 

sanguinea]) have larger distributions in the Crater District and 

are in less danger of extirpation from the area than the more 

specialized Maui Creeper (Loxops maculatus newtoni) and 'I'iwi 

(Vestiar ia coccinea) . The ranges of the latter-species within 

- - 

Haleakala are limited and are hiqhlv sensitive to seasonal shifts 

in resource abundance, much more so than the more generalized 

Drepanids . 

Native non-passerines tend to have broader ranges than do the 

honeycreepers, but no natives are as abundant as the broad-ranged 

exotics such as the Chukar (Alectoris chukar) or Pheasant. As 

illustrated in Table 1, the ranges of the Nene and the Pueo (a 

flammeus sandwichensis) are similar in many respects to those of 

the broad-ranged exotics, except as regards density values. The 

low densities of native non-passerines may be attributed to 

several factors, among them, competition between native and 

exotic species, and the resulting exclusion of natives from sub- 

optimal habitats, habitat destruction, and predation by exotic 

mammals . 

Management recommendations for the two endangered species in 

the Park center on the last two problems mentioned above. We 

feel strongly that control and elimination programs for nest 

predators should continue, and be expanded during the breeding 

seasons of both the 'Ua'u and Nene. Predation, especially by 

rats (Kjargaard 1978), is a serious threat to the nesting success 

and continued survival of both species in the Crater. In addi- 

tion, populations of both the 'Ua'u and Nene should be carefully 

studied in order to define not only the sizes of breeding 

populations, but also the nesting success of both species. 

In keeping with Park goals, ecosystem management and maintenance 

of native habitats of these species should be of high 

priority. ~limination of exotic organisms in the Park (particularly 

feral mammals and the more aggressive exotic plants) will 

significantly contribute to the enhancement of native bird 

habitats in Haleakala.


Eden, J. T. 1971. Population densities of birds derived from 

transect counts. Auk 88: 323-341. 

Kjargaard, J. I. 1978. The status of the Hawaiian Dark-rumped 

Petrel at Haleakala, In C. W. Smith, ed. Proceedings, 

Second Conf. in Natural Scznces, Hawaii Volcanoes National 

Park. CPSU/UH (Univ. of Hawaii, Botany Dept.) . 

Ralph, C. J. 1978. Habitat utilization and nichd components 

in some endangered Hawaiian forest birds. In C. W. Smith, 

ed. Proceedings, Second Conf. in Natural scZrl6es, Hawaii 

Volcanoes National Park. CPSU/UH (Univ. of Hawaii, Botany 

Dept.). 

Reynolds, R. T., J. M. Scott, and R. A. Nussbaum. A variable 


press) .

TABLE 1. Density values (birds/40 ha) in five Crater District habitats. (P = present at a 

density less than 1 bird/40 ha). 

Scientific Name 

Pterodrma phaeopygia sandwichensis 'Ua'u P 

Habitat* 

Hawaiian or Density (bird/40 ha) 

Vernacular Name 1 2 3 4 5 

Phaethon lepturus dorotheae Koa' e-kea P P P 

Branta sdvicensis Nene 2 5-10 1-5 1 

Alectoris chukar Chukar 1-20 5 1-10 2-4 

Francolinus pondicerianus Gray Francolin P 

Phasianus colchicus Ring-nec ked 3-4 5 1 1-25 

Pheasant 

Plwialis dminica Kolea 5 5 

- Asio flamneus sdwichensis 

Eueo 2 P 

Tyto - alba 

Barn Cw1 2 P 

Colunba - livia 

mck mve P 

Alauda arvensis Skylark 2 2 1 

Garrulax canorus ~ldious 

Laughing- thrush 

P 

Leiothrix - lutea 

Red-billed Leiothrix 1-18 3-20 

Mimus polyglottos Wckingbird P 1 1-4 1-2

TABLE 1--Continued . 

Scientific Name 

Zosterops j aponicus 

Acridotheres tristis 

Mxops virens wilsoni 

Mxops maculata newtoni 

~imatione sanguinea sanguinea 

Vestiaria coccinea 

Mnchura punctulata 

Cardinal is cardinalis 

Carpodacus mexicanus 

* Habitat types: 

Habitat* 

Hawaiian or Density (bird/40 ha) 

Vernacular Name 1 2 3 4 5 

Japanese P 7 1-8 

White-eye 

Cornon m a P 

Maui 'Amakihi 1 

' Alauwahio 

' Apapane 6 

'I'iwi 

Spotted Mia P 

Cardinal 

House Finch P 1-40 P-35 

1) Rock, cinder, open native scrub communities; crater floor; crater slopes 

2) Grasslands 

3) Savannah 

4) Closed native scrub 

5) Native rain forest

EXPERIMENTAL HYBRIDIZATIONS IN HAWAIIAN METROSIDEROS 

Carolyn A. Corn 

Hawaii State Division of Forestry 

Department of Land & Natural Resources 


The genus Metrosideros of the family Myrtaceae is native, 

but not endemic to Hawai'i. The genus occurs scattered from 

eastern Australia through the high islands of the Pacific Ocean, 

with the largest number of species located in New Zealand. If 

New Zealand is considered to be the probable center of origin for 

the genus because it has the largest number of extant species, 

then the high islands of the Pacific to the east and north could 

have served as "stepping stones" in the migration of the genus to 

Hawai'i by long distance dispersal (Corn 1972b). The most common 

species occurring on oceanic islands of the ~scific is M. collina 

(J. R. & G. Forst.) Gray, whose distribution ranges from the New 

Hebrides on the west to Pitcairn Island on the east to the 

Hawaiian Islands on the north. The species is fairly uniform in 

the western part of its range (i.e., Tahiti and Marquesas) and 

reaches its greatest - variability - in the Hawaiian Islands (Smith 

1973). This species in Hawai'i is presently called - M. polymorpha 

Gaud. (St. John 1979). 

The genus is abundant in the relatively undisturbed portions 

of the six largest Hawaiian Islands. It occurs in diverse 

edaphic, topographic, and climatic habitats as either a tree or 

shrub. The plants are variable in height, shape, vegetative and 

floral characteristics. - Taxonom-ic treatments of the genus 

(Hillebrand 1888; Rock 1917; Skottsberg 1944; Porter 1972; 

St. John 1979) are difficult since the taxon may be best 

described as a polymorphic group of plants. Sastrapradja and 

Lamoureux (1969) found no distinct patterns of variation in wooa 

variation. Anatomical and morphological evidence of leaf varia- 

tion in mature plants along an altitudinal and rainfall transect 

on Mauna Loa, Hawai' i, often suggests clinal patterns of varia- 

tion commonly found among outbreeding forest trees (Corn 1979). 

This paper includes information from observations and field 

hybridizations of seven varieties of - M. polymorpha present'on 

Mauna Loa, Hawai'i. A basic number of n = 11 chromosomes was 

described by Niimoto (1950), Skottsberg (1955), and Carr (1978), 

which corresponds to the basic count for New Zealand material by 

Mouse1 (1965). However, Skottsberg also noted counts of n = 12 

and n = 13 in material from the Island of ~awai'i' for varieties 

incana and glabrifolia, respectively. Carpenter (1976) in a 

paper on plant-pollinator interactions described two yellowflowered 

plants as self-compatible, but the red-flowered plants 

as being partially self-incompatible.

The objective of this paper is to ascertain if hybridization 

can occur between plants that are morphologically different. 

METHODS 

Pollen samples were obtained by two methods. The first 

method involved coating glass slides with either vasoline or 

Scotch tape and placing them around and under blooming Metro- 

sideros trees for four days and nights to see if Metrosideros 

pollen was wind-dispersed. The second method was devised to see 

if birds were transmitting Metrosideros pollen between trees at 

1220 to 2012 m elevation in Hawaii Volcanoes National Park and 

Kilauea Forest Reserve. Birds were caught in mist nets, their 

head feathers around their beaks were sampled with scotch tape, 

and the scotch tape was fastened to microscope slides and viewed 

for Metrosideros pollen. 

Experimental field hybridizations were prepared by removing 

all but five to seven buds in an inflorescence and emasculating 

the remaining buds. A wire frame was erected around the prepared 

inflorescence which was covered by an organdy bag, so that the 

bag did not touch the elongating styles during windy weather. 

The bag was secured by cotton and string at its base to prevent 

ants and other insects from crawling into the bag. A small plas- 

tic umbrella was erected above the inflorescences used in nectar 

production studies to prevent rain from diluting the nectar. 

Some of the prepared bags with emasculated flowers served as con- 

trols which were not crossed. Other bagged inflorescences were 

crossed with pollen from select plants about 10 days after emas- 

culation. Pollination bags were again secured after the flowers 

were crossed and labelled. Approximately four to seven months 

later when the capsules were mature, the labelled bags were 

cli-pped off the female parent plant and brought into the 

laboratory for analysis. 

Each bag containing mature capsules was air-dried until the 

capsules opened. One hundred seeds from each cross were placed 

into petri dishes, given light and water for germination trials. 

After one month the percent seed germination for each cross was 

tallied. The seedlings were then planted into soil and grown. 

RESULTS 

Metrosideros grows as a tree or shrub which bears conspic- 

uous flower clusters at the terminal portions of its branches. 

Although it flowers most commonly during the spring and summer, 

sporadic blooms may be seen on a few trees throughout the year. 

The inflorescence is composed of a flower cluster that may vary 

in number from several to about 30, but normally between 18 and 

24 flowers.

Flowers normally have no scent and are red in color. How- 

ever, their color on different trees may vary from deep red to 

various shades of red, salmon, orange, yellow, and very rarely 

white. The flowers are perfect with five small petals and numer- 

ous protruding stamens which are the showiest portion of the 

flower. The flowers open with the modified floral cup facing 

upward which allows the secreted nectar to be retained within the 

cup. The height of the stamens in relation to the style varies, 

as does the space between the row of stamens and the central 

style. 

Nectar production begins as the petals and stamens unfold, 

and is greatest several days later when the anthers begin to 

dehisce. It ceases three to five days later when the anthers and 

petals abscise. The receptive period of the stigma may vary from 

one to two days after the stamens begin to exert (Carpenter 1976) 

to several days after the anthers begin to dehisce (Corn 1972~). 

Within an inflorescence it is common to have flowers in all 

stages of the blooming cycle. 

Nectar is 10 to 15% solid (by refractometer measurements) 

when flowers are enclosed within a plastic bag. However, these 

same flowers when exposed to wind, low humidity, and no nectar- 

gathering animals, may have nectar concentrations of >60% (Corn 

1979). Analysis of nectar yields low protein (or histidine) 

content. 

The stamens produce abundant sticky pollen which attract 

various hymenopterans, including native and introduced bees and 

wasps. These hymenopterans may also obtain nectar on sunny, hot 

days. Other insects seen on the flowers are nocturnal cater- 

pillars that live in the flower buds during the day and emerge at 

night. They feed on the anthers and young succulent portions of 

the flower buds. Moths, crickets, ants, and even centipedes have 

been seen on the blossoms. No wind-dispersed Metrosideros pollen 

was obtained using sticky slides placed under and around blooming 

trees. 

Although the flowers are open and relatively unspecialized, 

their dimensions and position on the branch are probably best 

suited to bird pollination. Various native and introduced birds 

visit the flowers for nectar and/or insects. The pollen adheres 

to the feathers and beaks of birds visiting the flowers. Twentythree 

of 27 sampled birds had Metrosideros pollen (Table 1). 

Species carrying pollen include: 'Amakihi (Loxops virens) , 

'Apapane (Himatione sariguinea), Hawaiian Creeper (Loxops 

maculatus m) 

, Japanese White-eye (Zosterops japonica) , House 

-rpodacus mexicanus frontalis), and House Sparrow (Passer 

domesticus). Since the birds commonly flit from tree to tree 

visiting the flowers, they can serve as active and efficient 

cross-pollinators. Carpenter (1976) found more capsules produced 

on trees that birds were visiting the blossoms, than on trees 

where insects but no birds were visiting the blossoms.

Selective field hybridizations yielded mixed results. None 

of the uncrossed emasculated flowers produced seed or mature cap- 

sules. Since apomixes is not usually associated with diploidy, 

it is probably safe to asspme that apomixes plays no role in seed 

production. 

Hybridizations were made between trees within one site and 

between varieties at various sites. In some cases successful 

crosses were made in one direction between two plants, but the 

reciprocal cross was not successful. Carpenter (1976) suggested 

a partial self-incompatibility system was present in red- 

flowering plants. Of nine self-pollinated red-flowering plants, 

four of these (36%) set no seed. When these same plants were 

outcrossed, five of 36 crosses (or 14%) produced no seed. Seed 

germination from selfed individuals commonly yielded few 

seedlings. 

Crosses were attempted among the seven varieties found on 

Mauna Loa (Fig. 1). All but one possible combination was at- 

tempted. Of the 20 combinations tried, seven of them did not 

produce capsules and seed. Many factors could have contributed 

to these failures. For example, some bags were broken off the 

trees between the time of pollination and capsule maturity; some- 

times the style was injured during emasculation; the timing of 

the cross was poor; the pollen too old; or climatic factors 

unfavorable. Before self-incompatibilities are attributed to 

these failures, additional trials and cytological studies need to 

be made. 

Not all successful crosses diagramed in Figure 1 had their 

capsules collected at the proper time. Some capsules had de- 

hisced and the seeds exposed to rain. When this happened the 

seeds became darker in color and did not germinate. 

A subset of data shown in Figure 1 (Fig. 2) illustrates 

those instances where seed germination could be tabulated. Some 

of the 55 crosses that were tried on 16 trees failed to produce 

capsules. Where capsules were produced, germination varied from 

Therefore, the occurrence of these varieties in certain hab- 

itats may limit their chances of crossing with other varieties 

that are not present in the immediate area. However, these vari- 

eties may still be able to hybridize if given the opportunity. A 

good example of this can be seen in a cross I made two years ago 

between a plant from the Marquesas Islands and a plant from 

O'ahu. The Marquesas Island plant which is growing at Lyon 

Arboretum, is distinctly different in appearance from the O'ahu 

plant. Although no seed was produced using the O'ahu tree as the 

female parent, 22% seed germination was obtained using the 

Marquesas Islands tree as the female parent. 

In summary, various crosses have been attempted among the 

seven recognized varieties found on Mauna Loa, Hawai'i. Some of 

these crosses produce viable seeds with the hybrids now being 

grown for future analysis and crosses. Other crosses did not 

result in seed set. The reasons for these failures are not 

known. Additional work needs to be done to verify if these 

crosses are genetically incompatible. 

ACKNOWLEDGMENTS 

Support for research of the paper was made possible by 

research grants from Island Ecosystems IRP, US/IBP Hawaii (NSF GB 

23230) and Pacific Tropical Botanical Garden. Travel to the 

conference was furnished by Hawaii Division of Forestry. 


Carpenter, F. L. 1976. Plant-pollinator interactions in Hawaii: 

Pollination energetics of Metrosideros collina (Myrtaceae). 

Eco~. 57(6): 1125-1144. 

Carr, G. D. 1978. Chromosome numbers of Hawaiian flowering 

plants and the significance of cytology in selected taxa. 

Amer. J. Bot. 65(2): 236-242. 

Corn, C. A. 1972a. Genecological studies of Metrosideros. 

Pages 88-95 in D. Mueller-Dombois, ed. Island ecosystems 

stability and evolution subprogram: Second progress report 

& third-year budget. Island Ecosystems IRP, US/IBP Tech. 

Rep. 2. 

. 1972b. Seed dispersal methods in Hawaiian Metrosideros. 

Paqes 422-435 in J. A. Behnke, ed. Challenqing biological 

problems: ~Eections toward their solution. -oxford ~niv. 

Press, New York.

Corn, C. A. 1979. Variation in Hawaiian Metrosideros. Ph.D. 

Dissertation in Botany, University of Hawaii, Honolulu. 

295 pp. (Unpublished). 

Hillebrand, W. F. 1888. Flora of the Hawaiian Islands. Fac- 

simile ed., 1965. Hafner Publ. Co., New York. 673 pp. 

Mousel, B. 1965. Contribution a l'etude cytotaxinomique des 

Myrtacees. Mem. Mus. Nat. Hist. Nat. Ser. B 16: 91-125. 

Niimoto, D. 1950. Chromosome counts in some members of 

Myr taceae : Stain experiments with the plants studied. 

Univ. Haw. Dean Prize for Undergraduate Research, 

University of Hawaii, Honolulu. 12 pp. (Unpublished). 

Porter, J. R. 1972. The growth and phenology of Metrosideros in 

Hawaii. Ph.D. Dissertation in Botany, University of Hawaii, 

Honolulu. 291 pp. (Unpublished) . 

Rock, J. F. 1917. The ohia lehua trees of Hawaii. A revision 

of the Hawaiian species of the genus Metrosideros Banks, 

with special reference to the varieties and forms of Metro- 

sideros collina (Forster) A. Gray subsp. polymorpha (Gaud.) 

Rock. Bot. Bull. Hawaii Board Agr. For. 4: 1-76. 

St. John, H. 1979. Metrosideros polymorpha (Myrtaceae) and its 

variations. Hawaiian Plant Studies 88. Phytologia 42(3): 

Sastrapradja, D. S. 

wood anatomy 

Bogoriensis 5: 

, and C. H. Lamoureux. 1969. Variations in 

of Hawaiian Metrosideros (Myr taceae) . Ann. 

1-83. 

Skottsberg, C. 1944. Vascular plants from the Hawaiian Islands. 

IV. Acta Horti. Gotoburg. 15: 402-410. 

. 1955. Chromosome numbers in Hawaiian flowering plants. 

A r k i v fur Botanik 3: 63-70. 

Smith, A. C. 1973. Studies of Pacific Island plants. XXVI. 

Metrosideros collina (Myrtaceae) and its relatives in the 

southern Pacific. Amer. J. Bot. 60(5): 479-490.

. 

rl 

4 

... 

N O N 

rlrlrl 

m 

s 

rl

POLY MORPHA 

---- NO SEED 

- SEED PRODUCED 

Figure 1. Field hybridizations attempted between 

Metrosideros polymorpha varieties.

POLY MORPHA 

- 0.1 TO 15% SEED GERMINATION 

- 16 TO 30% SEED GERMINATION 

31 TO 42% SEED GERMINATION 

Figure 2. Seed germination for a subset 3f field 

hybridizations shown in Figure 1 of 

Metrosideros polymorpha varieties.

THE RARE AND THREATENED PLANTS IN THE AHUPUA'A OF MANUKA, 

KAULANAMAUNA, AND KAPU'A, SOUTH KONA, HAWAI'I* 

Lisa K. Croft, and Paul K. Higashino 




There is no simple description for a typical community with- 

in the ahupua'a of Manuka, Kaulanamauna, and Kapu'a, as a conse- 

quence of climatic patterns in South Kona. Within this location 

there is a mosaic of communities. 

Subalpine, pioneer communities on lava, kipukas, "fog inver- 

sion layer" rain forests, mixed mesophytic forests, dry transi- 

tional forests, and coastal strand communities are all found 

within this one area. 

This paper focusses on the rare' and threatened plants that 

are located within each of these zones, as identified by the 

project systematists. For species that have been identified on 

certain lists as rare and endangered, preliminary information-- 

habitat, condition (vigor, regeneration), and general obser- 

vations--has been compiled. This research was sponsored by the 

National Science Foundation under the Student Originated Studies 

Program to conduct a baseline survey of the archaeological and 

natural resources in the ahupua'a of Manuka, Kaulanamauna, and 

Kapu'a, during the summer of 1977. 

* Abstract

HUMAN SETTLEMENT AND ENVIRONMENTAL CHANGE 

AT BARBERS POINT, OIAHU 

Bertell D. Davis 

Archaeological Research Center Hawaii, Inc. 


and 

Department of Anthropology 



ABSTRACT 

Recent studies at Barbers Point, O'ahu, have demon- 

strated an unparalleled potential there for coordinated 

research on human settlement and environmental change 

in leeward "marginal" regions of the Hawaiian Islands. 

The following paper presents some of the more signif- 

icant results: the available data are summarized, and 

several interpretive models from different sources are 

offered. 

The current level of knowledge is tantalizing to say the 

least. Clearly our principal limitation is one of sampling. It 

is expected, however, that with the definition of specific 

research goals this limitation can be overcome. 

Preliminary environmental impact studies at Barbers Point, 

O1ahu, have demonstrated the unique potential there for signif- 

icant contributions to the cultural and natural history of the 

Hawaiian Islands. Several areas of investigation are of partic- 

ular interest: the survival of rare and endangered species, the 

extinction of endemic Hawaiian avifauna, the structure of the 

leeward lowland forest prior to man's arrival in the islands, and 

the native Hawaiian settlement of a presumed "marginal" environ- 

ment. To better accomodate such diverse but parallel concerns, 

continuing research is coordinated in phases, beginning with 

intensive survey of the terrestrial biology and archaeological 

resources. Fieldwork for this Phase I study was completed over 

the past 12 months, and a detailed report is now in preparation 

(Davis & Griffin 1978). 

The following paper summarizes the findings of the current 

study. Using data collected from the cultural survey, together 

with that recovered from excavation by the Bishop Museum (Sinoto 

1976, 1978), I suggest a tentative model for interpreting the 

archaeological remains at Barbers Point. The model focuses on 

patterns of settlement and subsistence, and several alternatives 

are considered as testable hypotheses. This paper is of course 

necessarily quite preliminary. My purpose here is primarily to

open ideas for consideration and to stimulate discussion for 

developing and refining the direction of future research. 

TO begin with, the study area is located along the south- 

western coast of O'ahu on a broad plain of emergent fossiliferous 

coral-algal reef (Fig. 1). Because the seaward portions of the 

region are relatively isolated from alluvial encroachment, the 

exposed limestone has weathered to form a shallow karst land- 

scape. The area is characterized by numerous solution sinkholes, 

irregular rock masses, and poor soil development. Local climate 

is generally arid with intense sunshine, warm dry winds, and low 

annual rainfall. However, the coral plain is a major aquafer, 

and many of the sinkholes in the study area penetrate to the 

water table. Thus despite superficially arid conditions, the 

availability of fresh water alone is not the principal constraint 

on human settlement and subsistence. Indeed, as is typical of 

atoll environments in the tropical Pacific, the more significant 

factors are those resulting from the alkaline substrate and the 

general paucity of inorganic sediments. It is these conditions 

which placed particular requirements upon the adaptive strategies 

employed by the former inhabitants of Barbers Point. 

The intensive botanical survey described and mapped the 

modern vegetation zones of the study area, and complementary 

wildlife studies defined primary habitats. The presence of a 

second threatened plant species, Achyranthes splendens var. 

rotundata Hbd., was confirmed in addition to the originally iden- 

tified Euphorbia skottsbergii var. kalaeloana Sherff. The popu- 

lation W r i b u t i o n of 50th species have been determined and 

pertinent recommendations presented (Miura & Sato 1978). An 

uncommon terrestrial shrimp, Halocaridina rubra, was also found 

in a large, flooded cave, and is briefly reported on by Miura and 

Sato (1978). 

From this baseline study we can now begin to consider the 

nature of the former environment at Barbers Point. Excavations 

by the Bishop Museum yielded extensive remains of terrestrial 

mollusca and extinct avifauna as well as cultural material. In 

addition to known coastal, forest, and predatory species, the 

avian assemblage includes several new taxa, two of which are 

flightless forms (Sinoto 1976; Ziegler 1978). Continuing 

analysis of the avifauna should yield substantial information 

regarding the birds themselves and their habitats. The potential 

for paleoenvironmental reconstruction has been especially en- 

hanced with the recovery of terrestrial rnollusca. Because they 

are habitat specific, these snails are highly responsive to local 

conditions. Although sampling for land snails has been somewhat 

limited, the available collection indicates a varied population 

in which eleven species have been identified to date (Kirch 

1978). Morgenstein (1978) observes that the rnollusca occur in a 

complete and continuous biostratigraphic profile. Preliminary 

sediment analyses--including pollen, spores, and phytoliths-- 

correlate with the inferred deposition of avian and molluscan 

remains, and attest to significant changes in the vegetation of 

the study area. It is suggested that an initial reduction in the 

resident aviary probably occurred prior to a major transition

from small-shrubbery to heavy-forest vegetation, and before human 

settlement of the area (Morgenstein 1978). What factors precip- 

itated these events, however, are still uncertain. Clearly more 

extensive sampling will be needed before further assessments are 

possible. 

Discussion of the archaeological remains may be initiated 

with three simple observations. (1) The settlement at Barbers 

Point was one of functionally integrated, multi-household resi- 

dence groups. (2) The settlement was minimally long-term, recur- 

rent occupation of the same habitation areas. And (3) the local 

subsistence pattern focused on the exploitation of marine-strand 

resources integrated with limited horticulture involving tree 

and/or root crops. Let us now consider these propositions in 

detail. 

What is the evidence for functionally integrated, multi- 

household residence groups? 

I have defined the archaeological evidence for such a group 

as the occurrence of functionally different, but contemporaneous 

features clustered in close spatial association. Here the 

assumption is that at the time of occupation, the various fea- 

tures of the cluster served a range of uses which, when combined, 

encompassed a set of activities that defined the residence group. 

I now suggest that the minimal group was an extended family 

incorporating several households similar to that outlined by 

Handy and Pukui (1972). The similarity is not complete, however. 

This is, first, because Handy and Pukui's description of the 

Hawaiian family is based on ethnohistoric material gathered 

during the 1930's in Ka'u, Hawai'i; and secondly, because many of 

the structural features listed for their residence group are 

apparently missing from the Barbers Point settlement. Specific 

features of this model include separated cooking and eating 

houses for men and women, sleeping houses, storage facilities, 

work areas, and a menstrual house or other place of seclusion for 

women (Handy & Pukui 1972: 7-11). This model clearly reflects 

the segregation of sexes according to the kapu system. Such 

proscriptions, however, may not necessarily have operated uni- 

formally throughout the islands, among all levels of society, or 

during all periods of cultural development. Indeed, while Malo 

(1951: 118-126) describes a similar residential organization for 

"respectable' people, he nevertheless observes that "no accounts" 

often followed no such proprieties. 

The habitation features recorded at Barbers Point were sub- 

jected to metric analysis using total area, interior floor area, 

and maximum wall width as the principal discriminating variables. 

Frequency distribution curves of the combined variables distin- 

guished three classes of structures ultimately based on overall 

size (Fig. 2). Except for three anomolous enclosures, Class I 

features are C-shaped structures less than 10 m2 in total area. 

Class I1 features include both C-shapes and walled enclosures of 

1735 m2. And finally, Class I11 features are rectangular 

enclosures 24 m2 or more in area.

Presence-absence trait analysis further defined these 

classes in functional terms. Because of their small size, shape, 

and virtual absence of other distinctive attributes, Class I 

structures are inferred to be for the storage of tools and 

materials, or possibly for such produce as yams and sweet 

potatoes. Class I1 features are the ordinary dwellings of indi- 

vidual households. Class 111 enclosures also appear to be ordi- 

nary dwellings. Although these enclosures are rather large when 

compared to other habitatiqn features in the study area, none 

approach the usual size range or structural complexity to 

indicate more specialized features, such as men's hoyses. 

A fourth class of habitation feature is defined from the 

trait analysis. These are distinguished by the presence of scat- 

tered midden and large mounds of burned coral associated with 

open, elevated floors. Class IV features are therefore inferred 

to bs cooking areas using surface imu, or ovens, although oven 

pits may be found under the raised floors. 

Extensive disturbances in the study area hive left few prob- 

able clusters intact. Of the remaining examples, one cluster 

especially, includes seven Class I storage features, six Class I1 

dwellings, and five Class IV cooking features in two separate 

areas within the cluster (Fig. 3). The habitation features are 

associated with structurally modified sinkholes and clearings of 

soil-humus deposits--both inferred to be garden areas, and with 

other unmodified sinkholes used as refuse dumps. The entire 

cluster is situated on a slight rise of limestone outcropping 

immediately adjacent to an area of surface drainage. Despite 

considerable evidence for surface runoff, the relatively minor 

difference in elevation does not seem sufficient to explain this 

location as a direct response to flooding alone. What is of 

interest is that shallow, but numerous pockets of silt are found 

in these drainages. The proximity of the habitation area to the 

accumulated sediments suggests that the drainages were also uti- 

lized. This may have been for additional gardening, or perhaps 

as a source of alluvial materials for use in mulched garden pits 

(sinkholes) . 

The one feature conspicuously missing from this cluster is 

the Class 111 enclosure. Although a Class 11 or several Class I 

features may be located nearby, the larger enclosures are in fact 

quite dispersed throughout the study area without any apparent 

pattern to their location. It is possible that these enclosures 

are the residences for individual households, and that the 

"clusters" may have functioned as communal foci for a larger 

settlement group. If so, then the whole nature of the residence 

group in terms of spatial distribution may require rethinking. 

Alternatively, the Class I11 enclosures may not be contem- 

poraneous with the feature clusters, and their distribution may 

reflect changes occurring in the settlement of the area. Trait 

analysis suggests that Class I11 features are either very late 

prehistoric, or wholly historic phenomena. Although precontact 

dates of A.D. 1666k41 and 1743+41 were obtained from one such 

enclosure tested by the Bishop Museum (Sinoto 1976: 87), the

artifact assemblage was clearly historic. This was true also of 

surface remains recorded in an adjacent feature during the 

present study. 

Whether these or other changes, and the conditions which may 

have induced these changes, were occurring at Barbers Point 

during the period of settlement are intriguing and important 

questions for investigation. 

What is the evidence that the settlement was minimally long- 

term and recurrent occupation of the same habitation areas? 

Adequate temporal controls have yet to be established. 

Nevertheless, that feature clusters initially conforming to my 

definition of a residence group are present in the study area 

indicates that the resident population must have included whole 

families. This would not likely have been the case if Barbers 

Point were merely the temporary campsite of itinerant spe- 

cialists. In the one instance where a range of dates are avail- 

able from a single feature (a Class 11 enclosure), the span of 

occupation is over 250 years (Sinoto 1976: 87). At least two 

separate events altered the nature of this site: the filling Of 

a sinkhole adjacent to a living floor followed by the building of 

the present enclosure wall. Whether or not this represents con- 

tinuous occupation has not been resolved. But it does suggest 

that use of the site may have become more formalized through 

time. Again, this does not seem to be consistent with short- 

term, or transient habitation. 

It is evident that an argument can initially be made for 

extended, recurrent residence in the study area. The possibility 

of permanent habitation, however, remains an open question. 

Tight stratigraphic control within and temporal control between 

habitation features are minimally essential for resolving these 

and other crucial questions raised by the available data. 

What is the evidence for local subsistence based on the 

exploitation of marine-strand resources integrated with limited 

horticulture? 

The artifact and midden assemblage clearly shows the use of 

marine and strand resources. Shellfish were collected from the 

shoreline and the reef. Octopus was taken from the reef on hook 

and line using cowrie-shell lures. And in-shore fishing was done 

with lines using small rotating and jabbing fishhooks. Off-shore 

line fishing is also indicated by large, one-piece rotating fish- 

hooks (Sinoto 1976), by fragments of large points probably for 

two-piece trolling lures (Lewis 1970; Sinoto 1978), and by the 

remains of tuna in the midden (Sinoto 1978). 

Although the availability of fresh water is not considered a 

major obstacle to the growth of cultivated plants, the present 

evidence for horticulture is largely circumstantial. The numer- 

ous walled sinkholes, enclosed humus-filled depressions, and

small clearings of soil-humus deposits are inferred to be garden 

areas. Ethnographic correlates for the walled sinkholes are 

found elsewhere across the tropical Pacific. On the low coral- 

line atolls tree and root crops are frequently grown in mulched 

garden pits to utilize the limited ground water, and to overcome 

the excessive alkalinity of the carbonate substrate (Barrau 

1961). Similar strategies for conserving moisture, such as low- 

walled windbreaks and intensive mulching, have been documented 

from arid localities elsewhere in Hawai'i. Then there is the 

continued survival of several native economic plant species, 

particularly noni and ti, found thriving in the study area. 

Depending upon the plants involved, gardening at Barbers 

Point would potentially have been a labor-intensive activity. 

Tree crops, for instance, probably required only occasional 

tending once they were well started. The persistence of noni, 

ti, and others in the deeper sinkholes indicates that these 

plants are capable of supplying sufficient natural mulch in the 

form of leaf litter to ensure continued growth. On the other 

hand, the level of mulching required to reduce excessive alka- 

linity and provide a suitable medium for productive growth of 

root crops would have been a demanding activity. If indeed root 

crops were grown in the study area, then this point can be taken 

further to suggest that at least a part of the population at 

Barbers Point was permanently resident there. 

To obtain direct evidence for cultivation, or indirect evi- 

dence of the potential for cultivation, w i l l require largely a 

comparative approach. Both the inferred horticultural features 

and the "natural ," or unmodified, features must be sampled to 

meet two criteria. It must be demonstrated (1) that the deposits 

in the culturally modified features are consistently similar from 

one feature to the next, and (2) that the deposits in the mod- 

ified features are significantly different from those in the 

unmodified features. Once these conditions are fulfilled, it 

must be further established that plant- cultivation is the most 

reasonable explanation accounting for the differences between the 

two sample groups. Direct evidence may include macrofossils like 

yam or sweet potato tubers, or more likely, microfossils such as 

the pollen, spores, or phytoliths from the cultivated plants. 

Although less satisfying, indirect evidence would largely be 

based on first determining the proper growth medium required by 

the presumed cultivated plants. Secondly, it must be shown that 

the proper conditions are, or were present in the modified 

features, and not in the unmodified features. 

The most intriguing question, and the one for which the 

least empirical data is currently available, is the relationship 

between the remains of the extinct avifauna and that of the human 

settlement. It has already been suggested that the major reduc- 

tion in the Barbers Point aviary apparently occurred prior to 

human settlement. However, this is a very preliminary assessment 

requiring further confirmation. Direct affects of human preda- 

tion, or significant disruption of habitat resulting from other 

unassociated events, cannot yet be discounted as possible 

contributive factors in the extinction of selected bird species.

SUMMARY 

Barbers Point is a geologically unique area for the high 

volcanic islands of Hawai'i. Located on the coast of an exten- 

sive raised coral-reef plain and isolated from alluvial encroach- 

ment, the exposed limestone has weathered to form a shallow karst 

landscape. Karst environments are alkaline, ideal for the 

preservation of bone and vegetal debris. Yet the area had 

received little serious attention and was generally viewed as too 

marginal to have supported a significant precontact population. 

Routine environmental impact studies in conjunction with a 

proposed harbor development have since reversed this opinion. 

Intensive cultural surveys indicate that Barbers Point was 

more extensively settled than had previously been considered. 

Pending detailed excavations, the initial data suggest a settle- 

ment of multi-household residence groups over a time span of 

c. A.D. 1600 to 1870. Local subsistence was based in part on 

marine-strand resources, and apparently integrated with limited 

cultivation adapted to make use of the many sinkholes that 

characterize the area. Other relationships have yet to be 

determined. 

For the natural sciences, biological surveys have identified 

an uncommon endemic terrestrial shrimp and rare and endangered 

plant species. Of greater significance is the potential for 

paleontological and paleoenvironmental studies, particularly 

regarding the bird life and forest structure of the leeward low- 

lands prior to man's arrival in the islands. Limited test exca- 

vations have recovered the remains of several new taxa of extinct 

birds, including flightless forms. Varieties of land snails have 

also been identified from the excavations. Highly responsive to 

local conditions, these snails attest to significant changes in 

the vegetation at Barbers Point.


Barrau, J. 1961. Subsistence agriculture in Polynesia and 

Micronesia. B. P. Bishop Museum Bull. 223. Honolulu. 

Davis, B. D., and P. B. Griffin (Eds.). 1978. Studies in natu- 

ral history and human settlement at Barbers Point, O'ahu. 

Interim Report I: Present environment and archaeological 

survey of the Proposed Deep-Draft Harbor Area, Barberg 

Point, 'Ewa, O'ahu, Hawai'i. Archaeological Research Cente~ 

Hawaii, Inc., Manuscript Report No. 14-1151. Lawai, Kauai. 

Handy, E. S. C., and M. K. Pukui. 1972. The Polynegian family 

system in Ka-'u, Hawai'i (Original 1958). Charles E. 

Tuttle, Rutland. 

Kirch, P. V. 1978. Report on recent subfossil land Mollusca 

from Barbers Point, Oahu. B. P. Bishop Museum Manuscript 

No. 120777. Honolulu. 

Lewis, E. 1970. The Campbell project: A preliminary report. 

Manuscript prepared for Graduate Seminar in Anthropology, 

University of Hawaii, Honolulu. 

Malo, D. 1951. Hawaiian antiquities. B. P. Bishop Museum Spe- 

cial Publication 2 (Second edition). Bishop Museum Press, 

Honolulu. 

Miura, M. T., and G. Sato. 1978. Botanical and faunal survey of 

the Proposed Deep-Draft Harbor Area, Barbers Point, O'ahu. 

- In B. D. Davis and P. B. Griffin, eds. Studies in natural 

history and human settlement at Barbers Point, O'ahu. 

Interim Report I: Present environment and archaeological 

survey of the Proposed Deep-Draft Harbor Area, Barbers 

Point, 'Ewa, O'ahu, Hawai'i. Archaeological Research Center 

Hawaii, Inc., Manuscript Report No. 14-1151. Lawai, Kauai. 

Morgenstein, M. E. 1978. Geoarchaeological reconnaissance of 

Barbers Point. Hawaii Marine ~ese'arch, Inc., Kailua, Oahu, 

Hawaii. (Manuscript). 

Sinoto, A. 1976. A report on cultural resources survey at 

Barbers Point, Island of Oahu. B. P. Bishop Museum Nanu- 

script No. 122476. Honolulu. 

. 1978. Archaeological and paleontological salvage at 

Barbers Point, Oahu. B. P. Bishop Museum Manuscript No. 

030178 (Draft). Honolulu. 

Ziegler, A. C. 1978. Prehistoric Hawaiian birds. In C. W. 

Smith, ed. Proceedings, Second Conf. in ~aturarscience, 

Hawaii Volcanoes National Park. CPSU/UH (University of 

Hawaii, Botany Dept.). (In preparation).

0 4 8 mi. 

LANA,," 'L) 

KAHO'OLAWE * 

1;> 

HAWAI'I 

FIGURE 1. Map of O'ahu and the Hawaiian Islands showing the 

location of the Barbers Point Study Area.

WIDTH OF LARGEST WALL (cm) 

TOTAL AREA (m2) 

FLOOR AREA (m2) 

FIGURE 2. Frequency distribution graphs for total area, interior 

floor area, and width of largest wall for habitation 

features from the Barbers Point Study Area.

FIGURE 3. Topographic map of a portion of the Barbers Point Study 

Area showing the distribution of habitation features 

and the location of surface drainages. Stippled area 

indicates feature cluster discussed in text.

ESTABLISHMENT OF SOME RECENT IMMIGRANT INSECTS 

IN HAWAII VOLCANOES NATIONAL PARK 

C. J. Davis 

Hawaii Field Research Center 



Between 1961 and 1978 over 300* new immigrant arthropods 

were recorded in the State of Hawaii. Most of these were dis- 

covered on O'ahu but a few were first recorded from the neighbor 

islands such as the false dandelion gall wasp which was dis- 

covered on the Mauna Loa Strip Road in June 1966 and the bristly 

rose "slug" which was discovered in Volcano on 6 October 1973. 

While most of these newly reported organisms were insects, 

some were mites and miscellaneous arthropods. 

Between 1966 and 1978, 11 of these immigrant insects were 

recorded in Hawaii Volcanoes National Park. Undoubtedly there 

are others which have not been detected in the Park to date or 

reported by other sources not available to the writer. 

With the exception of a sphingid, Theretra nesus (Drury), 

most of the Park immigrants are firmly established. 

A summary of exotic insects which reached Hawai'i Island 

from O'ahu is presented in Table 1 and their relationship to the 

Park flora and other organisms is discussed. 

. . .. . . . - -. . . .~ . . - . ~. 

* Proc. Haw. Ent. Soc.

1) Xylosandrug compactus (Eichhoff). (Black twig borer) 

This is a tiny black beetle that bores into the twigs, 

branches, and boles of livinq trees . The male is brownish in 

color, smaller than the female, and about 1 mm or less in length. 

The female excavates a small chamber in the pithy portion of the 

twig or stem and deposits 30 or more white eggs in this niche. 

Upon hatching, the legless larvae feed on ambrosia fungus which 

is stored in dorsal pouches of the female and liberated for the 

developing brood. 

Together with this fungus the black twig borer has become an 

important enemy of fruit trees, ornamentals, orchids, and native 

and exotic forest trees in Hawai'i. Over 100 hosts have been 

recorded. 

This beetle has strong host preferences and w i l l attack vig- 

orous seedlings as well as mature trees. Often young trees are 

top killed and gradually succumb. Trees that have been weakened 

by drought or other factors are readily susceptible to borer and 

associated fungus organisms. 

The first host record in the Park was at Waha'ula in October 

1975, where it was found infesting twigs and branches of the 

native lama, Diospyros ferrea. 

The altitudinal range of X. compactus is sea level to 914 m 

elevation and of the 11 new-park insects listed in this paper, 

this beetle has the greatest plant pest potential. 

2) Coptosoma xanthogramma (White). (Black stink bug) 

The black stink bug was collected at Kukalau'ula anti-goat 

enclosure, 244 m elevation on Canavalia kauensis, 18 March 1977 

--a new host and Park record. It is the first known represen- 

tative of the family Plataspidae to become established in the 

Hawaiian Islands (Beardsley 1967). 

The adults are black in color, broadly oval in shape, about 

2 mm long, and odiferous when handled. 

The nymphal stages vary in color and, like the adults, pre- 

fer to feed on the succulent growth of host plants. Both adults 

and nymphs have sucking mouth parts. 

With one or two exceptions, legumes are the principal hosts 

and both exotic and native species are represented in the coastal 

areas of Hawaii Volcanoes National Park. 

The buqs are qreqarious with as many as 1000 adults and 

- - 

nymphs being observed on a four foot branch of a Sesbania tree on 

windward O'ahu.

An egg parasite*, Trissolcus sp., was reared from para- 

sitized eggs collected on O'ahu in January 1968, and at Hilo, 

Hawai'i, in February 1968. It is very likely established in the 

Park and hopefully keeping the black stink bug below pest levels. 

According to P. M. Marsh, United States Department of Agri- 

culture Taxonomist, this parasite probably came in with its host 

from the Philippine Islands. 

3) Psylla uncatoides (Ferris & Klyver). (Acacia psyllid) 

- 

In July 1970 this important pest of koa (Acacia koa) was 

found at high population levels on terminal foliar grow- Mauna 

Loa Strip Road, 1645 m elevation. This was also the first record 

of this forest pest on Hawai'i Island. 

It is a native of Australia and occurs in New Zealand and 

California. The major hosts are Acacias and Albizzias. In addi- 

tion to koa, another species (A. koaia) occurring at Kawaihae 

Uka, Mt. Kohala, was heaviiy attacked in the early 1970's by 

P. uncatoides. 

The adults are small, about 1 mm or less in length and 

resemble a tiny cicada. There are five nymphal instars and both 

nymphs and adults are gregarious and have sucking mouth parts. 

Leeper and Beardsley (1973, 1976) studied the Acacia psyllid 

in Hawaii Volcanoes National Park and at the A. koaia sanctuary, 

Kawaihae Uka, Mt. Kohala, and concluded in-their initial study 

that imported natural enemies were needed for the control of this 

new immigrant pest. 

This was subsequently accomplished with the introduction of 

two species of lady bird beetles, Harmonia conformis and Diomus 

pumilio. To date, Diomus has not been recovered -and since the 

aforementioned authors' last publication, Harrnonia was found on 

koa at Hilina Pali and clustered in a weathermter off the 

Mauna Loa Strip Road, 1646 m elevation on 25 October 1977--an 

indication that this beneficial lady bird beetle is well estab- 

lished in the Park. A number of these beetles were also found on 

the summit of Mauna Kea, 4205 m elevation on 21 August 1976. 

These were most likely wind borne. 

4 ) Gilletea taraxaci Ashrnead. (Dandelion gall wasp) 

The dandelion gall wasp was collected from false dandelion 

(Hypochaeris radicata) on the Mauna Loa Strip Road, 1818 m ele- 

vation in June 1966, a new State record. It was subsequently 

found on Mt. ~aleakala, Maui, at 3030 m elevation on the same 

host, 22 May 1969. 

* Proc. Haw. Ent. Soc. 20(2): 261.

Since the false dandelion is an exotic weed, the gall wasp 

can be regarded as a beneficial immigrant. 

5) Pollenia rudis (Fabricius). (Cluster fly) 

This calliphorid parasite of earthworms was first observed 

at Kamuela, Hawai'i, in April 1968 and by 1969 had spread rapidly 

around the Island becoming very abundant in Hawaii Volcanoes 

National Park and vicinity. 

The adults are nuisance pests of buildings, usually entering 

in late afternoons. Between 1969 and 1971, thousands of these 

flies were observed in the tack room of the stables located near 

the Tree Molds and in various homes. 

In recent years cluster fly populations have been at low 

population levels except for brief upsurges. The reasons for 

this are not fully understood. Lowering of earthworm populations 

and adult predation by plovers, spiders, skinks, and other 

organisms may have been responsible. 

6) Antianthe expansa (Germar). (Solanaceous treehopper) 

The plant hosts of the solanaceous treehopper include 

Cestrum, Solanum, and Acnistus. 

On 31 August 1977, nymphs of this insect were found on 

potted Nothocestrum by tree nursery personnel at Ainahou Nursery, 

914 m elevation. This was the first record of the solanaceous 

treehopper in the Park as well as a new host record. 

These are small bizarre insects having the head vertlcal and 

the nymphs are queerly ornamented with spines. The adults w l l l 

sometimes move beh-ind a leaf or around a branch to escape 

capture. 

Both adults and nymphs have sucking mouth parts, occur in 

large numbers, and may have pest potential. 

7) Papilio xuthus Linnaeus. (Citrus swallowtail) 

According to the literature, the caterpillars of the citrus 

swallowtail butterfly feed on various kinds of citrus trees, 

lime berry, Tr iphasia tr ifolia, prickly ash, Zanthoxylum 

americana, and Fagara spp., all members of the family Rutaceae. 

The attractive butterflies were first sighted in Volcano 

residential area in January 1974, and were officially recorded in 

Kailua, Kona, in June 1974. 

In October 1977, National Park tree nursery personnel found 

citrus swallowtail caterpillars feeding on kawa'u'kua-kulu-kapa, 

Fagara (Zanthoxylum dipetalum), a new Hawai'i host record. The

caterpillars were found on young nursery stock in the old tree 

nursery. 

Eggs were found on plants that were transferred to the new 

tree nursery but were not viable. 

Three mature larvae pupated and two normal adults were 

placed in the Park collection. 

8) Cladius difformis Panzer. (Bristly rose slug) 

The bristly rose slug was found severely damaging rose 

foliage on 6 October 1973, in the Volcano residential area. The 

adults are small black wasps and are known as sawflies. They are 

parthenogenic and oviposit in the petioles and midribs of the 

leaves. The caterpillars are slug-like in appearance and have 

chewing mouth parts. 

This was the first record of the Hymenopterous family 

Tenthredinidae in the State of Hawai'i and it was observed at 

Kilauea Military Camp in January 1977. The bristly rose slug is 

restricted to roses. 

A similar sawfly is found on wild blackberry and is well 

established in Volcano, Kipuka Ki, Kipuka Puaulu, and in other 

Park localities. Blackberry is the preferred host but it will 

feed on 'akala, Rubus hawaiiensis. It was purposely introduced 

for biological control of wild blackberry. 

9) Anua indiscriminata (Hampson). (Myrtaceous moth) 

10) Theretra nessus (Drury). (Yam sphingid) 

11) Macroglossum pyrrhostictum Butler. (Maile pilau hornworm) 

The last three immigrants are among the most recent arrivals 

in the Park and their relationship to the Park flora has not been 

determined. The myr taceous moth is a noctuid whose caterpillars 

feed on guava, eucalyptus, and possibly 'ohi'a foliage. The yam 

sphingid is doubtfully established and the maile pilau hornworm 

has not been found feeding on native Rubiaceae. Under laboratory 

conditions, however, the caterpillars have been reared to matu- 

rity on pilo, Coprosma sp. The adults are attracted to light and 

are frequently observed in the Park as well as Volcano District 

feeding on honeysuckle and impatiens flowers.


Beardsley, J. W. 1962. On accidental immigration and establishment 

of terrestrial arthropods in Hawaii during recent 

years. Proc. Haw. Ent. Soc. 18(1): 99-109. 

-- - . 1965. Insect invaders, our unwelcome immigrants. Haw. 

Farm Science 14(3): 1-4. 

. 1967. Coptosoma xanthogramma (White) (Hemiptera: 

Plataspidae) a new pest of lequmes in Hawaii. Proc. Haw. 

Ent. Sic. 19(3): 367-372. 

Davis, C. J. 1970. Black twig borer threatens native trees. 

Newsletter, Haw. Bot. Soc. 9(5): 38-39. 

Leeper, J. R., and Beardsley, J. W. 1973. The bioecology of 

Psylla uncatoides in the Hawaii Volcanoes National' Park and 

the Acacia koaia sanctuary. Island Ecosystems IRP, US/IBP 

Tech. Rep. 23: 1-3. 

. 1976. The biological control of Psylla uncatoides 

(Ferris & Klyver) (Homoptera: Psyllidae) on Hawaii. Proc. 

Haw. Ent. Soc. 22(2): 307-321. 

-

'IRBLE 1. Sunnnq of sm recent imniqrant insects rum established in Hawaii Volcames National Park. 

Insect 

1) Xylosandrus conpactus (Eichhof f) 1961 

(black twig b3rer) 

2) Coptosom ~anthcgram~ (White) 1965 1967 1965 

(black stink bug) O'ahu (Oct) (Sept) 

3) Psylla uncatoides (Ferris & Klyver) 1966 1967 1966 

(Acacia psyllid) I O'Au 1 (JW I (Mar) 

4) Gilletea taraxaci A sbad 

(dandelion gall wasp) 

6) Antianthe expansa ( G e m ) 

(solanamus treehopper) 

7) Papilio xuthus Limaeus 

(citrus dlckutail) 

Cladius dif fonnis (Panzer) 

(bristly rose slug) 

-1 197471974 11974 

0' ahu (Nov) (Aug) 

11) Macr lossum rrhostictm (Butler) 

*a- 1 1976 1 1976 11976 

O'ahu . (Nov) (Jun) 

Hawaii Volcanoes 

National Park 

1975 

(Oct) 

1977 

(Jan) 

1978 

(Jan)

A MATHEMATICAL MODEL OF 'OHI'A DIEBACK 

AS A NATURAL PHENOMENON 

William E. Evenson* 



Honolulu. Hawaii 96822 

INTRODUCTION 

By means of a very general model system, the possibility of 

interdependent dying, as contrasted to individual, random dying, 

in large areas of forest is investigated. In particular, insight 

is provided into the possible roles of natural mechanisms in pro- 

ducing such interdependent collapse behavior and what the proper- 

ties of such mechanisms must be. In this study, mechanisms of 

change in the forest are characterized as "natural" if they nave 

been part of the forest environment over an evolutionary time 

scale. Otherwise, they are characterized as "introduced." 

This model is applied to the ~roblem of dieback in the 

native 'ohira (~etrosideros collina kubsp. polymorpha) forests of 

Hawai'i. The focus of the model is on the "trigger" of the dieback. 

That is, we look only at the transition of trees from a 

healthy to a declining state (or vice versa). Mechanisms at work 

in thesubsequent death or reinvigoration of trees are beyond the 

scope of this model. 

The model allows separate examination of the effects of 

interaction between trees, external factors affecting growth, and 

physiological factors. Thus, it gives insight into the relative 

importance of various features of the 'ohi'a dieback phenomenon. 

It is of especial interest for 'ohi'a dieback to know 

whether natural factors could produce interdependent decline or 

whether an "introduced" epidemic (e.g . , disease or insects or 

some combination of introduced factors) is necessary to explain 

the field observations. The model presented here, while very 

general, deals with the plausibility of these various types of 

mechanisms for interdependent dieback. 

Before discussing the model considered in this paper and its 

application to 'ohi'a dieback, the value of such models from 

physias in biological problems and the kind of information they 

* Permanent address: Department of Physics and Astronomy, 

Brigham Young University, 

Provo, Utah 84602

can be expected to provide will be considered (Weidlich 1971; 

Callen & Shapero 1974). 

Problems of the sort considered here concern the behavior of 

"incompletely-specified systems." That is, they deal with sys- 

tems (e.g., a forest) for which there is either insufficient data 

to predict the behavior of each individual or incomplete knowl- 

edge of the laws which govern individual and system behavior, or 

both. In fact, these systems are intrinsically incompletely 

specified since we are not really interested in predicting the 

detailed behavior of each individual and since the data necessary 

for that task is impossibly extensive. Incompletely specified 

systems are probabilistic systems by their nature. The sub- 

discipline of statistical mechanics in physics is directed at 

such problems in Hamiltopian systems (Hobson 19711, and both 

models and methods originally developed in physics may be appli- 

cable to problems of incompletely-specified systems arising in 

other contexts. 

In particular, while the causes of interdependent behavior 

among atomic spins in magnets and interdependent behavior among 

'ohi'a trees in a dying forest are obviously rather different, 

the statistical behavior of these two systems does show 

interesting similarities. 

The most serious objection to the approach taken in this 

paper is that models from physics, like the one discussed below, 

are such an oversimplification of reality that their results can- 

not be relied upon. While this is always a danger, such diffi- 

culties can be minimized by focusing attention on features which 

are insensitive to specific details of the model. It is argued 

that the collapse behavior in the present study is such a fea- 

ture. In addition, if detailed simulation is desired, it is 

often possible to add specifics to such models and successively 

improve the simulation of reality. 

DESCRIPTION OF THE MODEL 

Imagine a forest of - N uniform-age, essentially identical 

trees. Let each tree be in one of two states: healthy or 

declining. (The effect of modifying this model to include individual 

differences in trees and continuous variation in vigor 

w i l l be discussed below.) This simple, very general model shows 

the important features of transition from healthy to declining in 

a manner which is qualitatively the 

model. 

same as a more realistic 

The forest of identical trees in two possible states can be 

modelled as a spin-+ classical magnet in two dimensions--the 

Ising model in statistical mechanics (Weidlich 1971; Callen & 

Shapero 1974). We use this analogy to analyze the behavior of 

our model forest. There are three important classes of variables 

in this problem:

(1) Tree interaction parameters which represent the effects 

of the trees on each other. We label these parameters Ji , for 

the interaction between trees i and i. Since we consider oily a 

single species in this model, the i represent only intra- 

specific competition. Interspecific compejition is treated as an 

individual (negative) growth factor in its effects on the trees. 

These parameters are analogous to spin-spin interactions in 

magnets. 

(2) Individual growth factors which influence the tree toward 

the healthy or declining state. These factors represent the 

net effect on each tree of such influences as limiting resources, 

disease or insect attacks, moisture relations, and interspecific 

competition. They may be favorable or unfavorable. We label 

these factors gi for tree i, 1 , . . , These factors are 

analogous to local magnetic fields in the magnetic systems. 

(3) A stand condition parameter which represents the rela- 

tive susceptibility of trees to stresses which might induce die- 

back. We label this parameter S. Large S characterizes stands 

which are resilient in meeting stress aFd resist the transition 

from healthy to declining. The magnitude of S w i l l depend on 

tree physiology, including maturity and oyher factors which 

affect resistance to stress. Such factors may include environ- 

mental restrictions to a tree's ability to respond to stress; for 

example, substrate limitation of root development. This para- 

meter plays a roll in the model analogous to inverse temperature 

in the magnetic system. 

We study the simplest case: i.e., we assume that the individual 

growth factors are the same for all N trees in the stand; 

and we assume that there is no tree interaction except for pairs 

of trees which overlap in canopy, rhizosphere, etc. (i.e., close 

neiqhbors). Hence, - G; = G, independent of which tree is con- 

& - 

sidered, and 

-ij 

I. = - I if trees - i and i overlap, while Iij = 0 

otherwise. 

So the picture is of N trees interacting with close neigh- 

bors and under the influence of a growth factor, which may either 

induce or retard growth. Some of these trees are healthy, and 

some are declining. The susceptibility of trees to transition 

from healthy to declining is governed by a stand condition 

parameter. 

MODIFICATIONS AND RELEVANCE TO 'OHI'A DIEBACK 

The two basic assumptions of the model are that the trees 

are identical and that there are two discrete states for the 

trees: healthy or declining. It is, however, possible to relax 

these assumptions. Since 'ohi'a is a pioneer species on new lava 

flows in Hawai'i, often with a readily available seed source from 

an adjacent, older lava flow, and since it does not regenerate in 

its own shade, it tends to grow in uniform-age stands (Mueller-

Dombois 1977). Of course the trees are still not identical: 

there are genetic and micro-environmental differences which produce 

differences in trees. However, since these variations are 

limited by the nearly uniform seed source, age, and macroenvironment 

of a stand, they can be accounted for by allowing our 

parameters, Li., Gi, and S, to vary from tree to tree (i.e., to 

vary with - i and 17 with a-limited distribution. 

The assumption of two discrete states for the trees can be 

replaced by a continuous spectrum of tree vigor or by the possi- 

bility of a tree's state being described as a mixture of the 

healthy and declining states. The latter suggestion actually 

seems most reasonable since trees do die in stages and sections 

--e.g., the crown may die but leave vigorous trunk sprouts 

(Mueller-Dombois 1977). 

It is clear that the model, even with relaxed assumptions, 

oversimplifies the description of a stand of 'ohi'a trees in 

nature. For this reaaon, we only address questions of general 

behavior which are not likely to depend critically on the details 

of the model. In particular, the presence or absence of widespread 

decline (i.e., of interdependent collapse behavior like a 

phase transition) is insensitive to the modifications described 

above, with the possible exception of allowing the stand condition 

parameter, S, to vary from tree to tree. (Allowing S to 

vary removes theconcept of an equilibrium "temperature" and-suggests 

that the forest is not in internal equilibrium.) However, 

- 

S should only vary over a narrow range through a given even-aged 

stand and hence can be treated as approximately constant in the 

systems of interest to this study. The effect of variations of S 

with a restricted distribution will be to broaden the "phase 

transition" so that it does not occur so sharply in time and 

space. But interdependent collapse behavior will still be 

evident in such a system. 

In sum, then, the modifications needed to make the propo-sed 

model more realistic will not affect the qualitative nature of 

any interdependent collapse which occurs. Hence, one can with 

some confidence draw conclusions from this model relative to the 

plausibility of various possible trigger mechanisms for 'ohi'a 

dieback. 

BEHAVIOR OF THE MODEL 

In the steady state, we can express the behavior of the 

model described above in terms of the steady-state probability, 

- P(h), that the forest stand consists of h percent healthy trees. 

In detailed analysis of the model, we yind that - P(h) - depends on 

the three parameters, I, 5, 2, only in the two combinations 

- G times S and I times S. So we introduce two new parameters: 

q = s, ana - k = - 13. hat-the state of the forest depends on the 

parameters of the model in these combinations accords well with 

our expectations for biological systems: the effect of a given

environmental stress (negative G) depends not only on the magnitude 

of the stress but also on -the resilience of the forest 

i.., on stand condition, - S). And similarly for the effects of 

competition. 

Figure 1 shows a sequence of - P(h) - for N = 100 with relatively 

large k and three values of 1. ~Fese curves illustrate 

the possibility-of interdependent change of a stand from healthy 

to declining condition.* Such a situation could occur if the 

effect of the trees on each other (through nutrient depletion for 

example) were quite strong, while the conditions for growth 

measured by - G fluctuated from favorable to unfavorable in an 

extreme year. 

When the individual growth factors or the stand condition 

parameter are neutral, so 5 = 0 (Fig. l[b]), the forest stand is 

in a critical state. The probability that the stand will be pre- 

dominantly declining is the same as that it will be predominantly 

healthy. So small fluctuations could cause a collapse from 

healthy to declining. The converse is also possible: the use of 

fertilizers, thinning, or similar techniques to make growth more 

favorable could revive a stand which has begun to die back. An 

alternative outcome might be the growth of separate healthy and 

declining regions within the larger stand. For the case where 

g = 0, one could have about equally large clumps of dying trees 

and of healthy trees. The establishment of such clumps will be a 

function of microenvironment. This may be what happens in a 

"hot spota'--the small patches of dieback in dryland areas. 

For contrast, Figure 2 shows a similar sequence of P(h) but 

with & = 0. There is no bimodality here because there is no 

interdependent response. Rather, examination of the model shows 

that the change from a healthy to a declining stand takes place 

randomly by individual trees in this case, resulting in random 

thinning. To get as strong a dieback condition (or as strong a 

healthy condi-tion) as in Figures l(a) and (c) requires much 

larger values of 9 (3 = 1.0 when - k = 0 gives about the same condition 

as 9 = 0.02 when k = 2.5). So the swings in growth factors 

have to be much greater to produce a recognizable dieback 

when interaction between trees does not play a significant role. 

An introduced disease could produce such a large change in 3. 

However, relatively light environmental stress may be sufficient 

to produce collapse when it is associated with interactions 

between the trees. Such light stress could easily be produced by 

changes in nutritional status, moisture, or 

constraints. 

other environmental 

* The mathematical expression of the model and its analysis to 

produce the curves discussed here are being prepared for sepa- 

rate publication. The analysis and results are very similar to 

Weidlich (1971).

We see from the two figures that interactions between trees 

introduce an interesting kind of stability, with an associated 

fragility, into the forest system. The stability comes from the 

tendency of the forest to maintain its state even for relatively 

small 9. However, once 9 becomes negative, there is a tendency 

for the whole system to "collapse." Hence a fragility, when 

close to critical values of the growth factors and stand condi- 

tion parameter, is closely connected with the stability of the 

forest. 

DISCUSSION AND CONCLUSIONS 

The motivation for this work has been to examine the plausi- 

bility of the hypothesis that natural mechanisms within the 

'ohi'a forest ecosystem can produce an interdependent collapse of 

a forest stand. 

As illustrated in the previous section, the simplified model 

discussed here does show such behavior. It has been noted that 

the modifications required to make the model realistic do not 

change the qualitative behavior of the collapse, which is the 

focus of our interest. We see that collapse due to a natural 

mechanism is a possible outcome which is an alternative to random 

thinning, or, in a more extreme case, to introduced epidemic. 

The reason that collapse occurs rather than random thinning is 

related to the way in which 'ohi'a stands are established as 

pioneer, uniform-age stands on new lava flows with no regenera- 

tion in their own shade. 

The fact that dieback in 'ohi'a forests seems to occur only 

in fairly mature forests, perhaps at intervals of several hundred 

years, suggests that the magnitude of environmental stress (i-e., 

the reduction in q) required to produce collapse is quite large 

and hence unusual. This fact also suggests that the interaction 

parameter, which increases as the trees get larger and more able 

to affect one another, must be fairly large. In addition, S will 

be larger for mature trees than, for example, for seedlings: So 

the requirements for collapse (i.e., that k be large and g go 

from a significant positive value to a significant negaeive 

value) are associated with the long time scale of several hundred 

years from establishment of the stand to dieback. This long time 

scale has made the collapse more difficult to recognize as a natural 

phenomenon in the sense addressed here. Similar phenomena 

on a shorter time scale have been much easier to recognize.* For 

example, one could apply the same type of model to the yearly 

dieback of annual plants. Annual life-forms provide a mechanism 

to respond to environmental stress (sharp decrease in - G). This 

* The example given here was originally suggested to me by 

N. Balakrishnan.

decrease in G (perhaps accompanied in some cases by a decrease 

in S) leads t o very rapid transition of a stand from healthy to 

deciining. The response pattern which has been illustrated in 

Figure 2 would serve for this example as well. 

This study also suggests that it would be profitable to seek 

out ways of measuring the interaction parameter, I, in the field 

to verify the interaction mechanism suggested-here for 'ohi'a 

dieback. This parameter will include all factors which cause 

of environment. For example, nutrient competition will bring 

about similar deficiencies in neighboring trees when nutrients 

are depleted below a critical level. Since this effect is due to 

interaction between trees, it is one of the components of the 

parameter - I. A l l such components must be considered. 

SUMMARY 

The possibility of collapse of the 'ohi'a forest from 

healthy to dying condition due to natural factors is investigated 

by means of a very general mathematical model. The model is 

closely related to the Ising model of magnetism. The role of 

interactions between the trees, i.e., intraspecific competition, 

is contrasted with the roles of environmental stress, disease, or 

insect epidemic in producing a collapse of the model forest. It 

is argued that the collapse behavior persists through modifica- 

tions of the model which bring it into closer correspondence with 

reality. Study of the model leads to the following conclusions: 

(1) Canopy collapse can plausibly be triggered by natural mech- 

anisms including competition, as well as by introduced epidemic 

factors . (2) Collapse is an alternative to random thinning or 

external epidemic, any of which can occur in the model under 

appropriate conditions. (3) Investigation of interaction mech- 

anisms between trees in the 'ohi'a forest is an especially 

important and potentially rewarding avenue for further research. 


Callen, E., and D. Shapero. 1974. A theory of social imitat 

Physics Today 27 (July) : 23-28. 

Hobson, A. 1971. Concepts in statistical mechanics. Gordon 

Breach Science Publishers, New York. 172 pp. 

Mueller-Dombois, D. 1977. Ohia rain forest studv: ecoloq i cal 

investigations of the ohia dieback problem in-~awaii. Final 

Report. CPSU/UH Tech. Rep. 20 (University of Hawaii, Botany 

Dept.) 117 pp. 

Weidlich, W. 1971. The statistical description of polarization 

phenomena in society. Br. 3. Math. Statist. Psychol. 

24: 251-266. 

on. 

and

Ole HEALTHY, h 

FIGURE 1. Probability distribution for healthiness 

of forest stands with strong interactions 

between trees.

Ole HEALTHY, h 

FIGURE 2. Probabilitv distribution for healthiness 

of forest stands with no interactions 

between trees. 

0

EVALUATION OF A NEW TECHNIQUE FOR HERBICIDAL 

TREATMENT OF MYRICA FAYA TREES 

- 

Donald E. Gardner 



Among the many exotic plant species occurring within Hawaii 

Volcanoes National Park, the firetree (Myrica faya Aiton) is well 

recognized as being among those which pose the greatest threat to 

the composition of the Park's native ecosystems. Because of its 

demonstrated ability to quickly and aggressively become estab- 

lished in many Park habitats occurring roughly between 2000 and 

4000 feet elevation, and since its current abundance is such that 

the species threatens to exceed the l i m i t s of practical control, 

high priority has been placed by resource management personnel 

upon developing efficient eradication methods. The present 

exotic plant control program, which is directed largely at the 

firetree, involves direct uprooting of smaller individuals and 

spraying the stems or trunks of larger plants with a diesel-Kuron 

(silvex) mixture, often facilitated by cutting into the bark. 

Large trees are routinely cut down and their stumps treated with 

the herbicide mixture to prevent resprouting. While herbicide 

treatment in this manner has proven to be effective in killing 

trees, certain disadvantages are also inherent in this method: 

Diesel as a solvent for Kuron is in general more diffi- 

cult to obtain and to work with in comparison with 

water. Application equipment is difficult to clean fol- 

lowing use as is the protective clothing of the workers 

them se 1 v5 s-. -Accldenrally - spil-led material presents 

greater cleanup problems. 

Storage of large quantities of diesel presents a poten- 

tial fire hazard. 

The herbicide must be sprayed entirely around the lower 

tree stem for best results, often in combination with 

stem scoring. Access to the complete circumference of 

the stem is often limited by bushy lower growth, heavy 

surrounding vegetation, or rough terrain. 

The spraying process inevitably results in accidental 

spray contact with native vegetation types growing close 

to target trees. Care that must be taken to avoid such 

contact lessens the efficiency of control work. Fire- 

trees in some habitats are often associated. so closely 

with 'ohi'a trees that effective treatment of the exotic 

without affecting the native tree is extremely 

difficult.

5) Although spraying is generally limited to the lower stem 

as compared to the entire crown, rather large quantities 

of herbicide solution are required per tree. Trans- 

porting such quantities to remote areas of infestation 

is often costly. 

To define and evaluate possible alternate methods of chem- 

ical control of M. faya, the herbicide Roundup (a product of the 

Monsanto company)-was selected for testing. Although at the time 

experimentation was begun Roundup had not been approved for 

routine application in a park exotic plant control program, the 

lack of approval was due to the relatively recent entry of the 

product on the market rather than to any demonstrated undesirable 

qualities. Roundup has since received this approval and is in 

use in the Park's efforts to control infestations of the exotic 

fountain grass (Pennisetum setaceum). 

Qualities of Roundup which led to its consideration were its 

demonstrated systemic activity in target plants other than 

- M. faya and the relative ease with which it is handled due to its 

water solubility. The current higher cost of the product compared 

to diesel-Kuron on an equal volume basis presents a recognized 

disadvantage to its use. This factor is the basis for the 

non-use of Roundup by the State of Hawaii Exotic Plant Control 

Division in M. faya control. Another product, Tordon 22K, which 

is available for use by the State gives satisfactory control 

(Walters & Null 1970; Robert Kami, Noxious Weed Specialist, 

Hawaii State Department of Agriculture, pers. comm.). A mixture 

of Tordon 212 and 2,4,5-T amine has also proven effective in 

killing firetree in Hawaiian state forest reserves (Kim 1969). 

The above-mentioned ease with which Roundup is taken up and 

translocated offers potential for use through methods other than 

those conventionally utilized . The advantages of such techniques 

may include reduction in time and effort required per treated 

tree, ability to transport equipment- and material to treat trees 

in difficult to reach areas, and avoidance of damage to other 

vegetation in the vicinity of the target tree. 


The Byron Ledge area of Hawaii Volcanoes National Park was 

selected as a suitable study area due to its accessibility and 

the large number of firetrees growing in a relatively open area 

unencumbered by heavy undergrowth. 

Trees for the preliminary tests fell within a size range 

from approximately 1.5 m high and 2 cm basal diameter to 2.5 m 

and 4.5 cm. One branch per tree of approximately 0.75 to 1 cm 

diameter, depending upon tree size, was clipped off and a small 

plastic vial containing 20 ml of either 0.5, 1, 2, 5, or 10% 

aqueous Roundup solution was attached to the cut branch end of 

each tree such that the cut surface extended through a hole cut

in the vial cap and was immersed in herbicide solution. Actual 

amounts of solution absorbed by each tree were not determined 

since initial attempts at sealing the vials upon the branches 

often allowed leakage to occur. Results of these tests indi- 

cated, however, the feasibility of killing firetrees with Roundup 

through a single application point. In subsequent tests, treat- 

ment variables including dosage in relation to tree size, growth 

form of the tree, and effect of position of application on the 

tree were considered. Other factors such as influence of pheno- 

logical or seasonal states upon treatment time were considered, 

although the observed lack of distinct dormant and growing 

seasons among trees in the study area resulted in less emphasis 

being placed upon the latter. 

Thirty-two test trees were selected and separated into two 

size classes: the smaller ranged from 16.3 cm basal circum- 

ference to 25 cm and from 2.55 m tall to 5 m; the larger included 

basitone growth forms (several equally large branches at the 

base), and heights up to 6.5 m. Crown cover (bushiness) was also 

visually evaluated as a size class factor. 

Four treatment locations upon trees were selected and test 

trees were further placed into categories accordingly: the upper 

portion of the crown; the lowest branch of suitable diameter on 

the mainstem (or of a major branch in a basitone tree); distal 

from the mainstem (the treatment branch was clipped off no less 

than 90 cm from the mainstem); proximal (the branch was clipped 

as near the mainstem as possible). Two herbicide concentrations, 

2 and 5%, were superimposed upon the size and treatment location 

categories. These concentrations were selected on the basis of 

preliminary test results as reasonable experimental dosages. 

Fifty-ml syringe barrels from which the plungers had been 

removed were fitted to lengths of bicycle inner tubing such that 

the rubber was stretched over the larger barrel opening. The cut 

branch end of each treated tree was inserted into the syringe 

barrel through the tube section immediately following clipping 

and the rubber was tightened securely around the branch with a 

screw-clamp. Petroleum jelly was also applied around the clamped 

area to insure the seal. The combined volume of the syringe 

barrel and rubber tubing was approximately 60 ml after displace- 

ment by the approximately 1 cm diameter cut branch. 

The syringe barrel attached to each tree was filled with the 

appropriate Roundup concentration immediately upon the attachment 

of the latter to the cut branch. The herbicide was introduced 

with a 50-ml syringe provided with an 18-gauge needle. All trees 

of this experiment were treated in February 1978. 

A test to determine the effectiveness of undiluted Roundup 

relative to that of 2 or 5% aqueous dilutions was conducted 

simultaneously with the latter. Smaller (12-ml) syringe barrels 

were sealed to freshly cut branch ends of trees with waterproof 

silicone rubber caulking compound. Trees of similar size range 

to those described above were selected. Proximal or distal

treatment locations were again chosen for each tree and the 

position of application along the mainstem axis was again 

considered. 

Trees were treated with either 1.2 or 3 ml of undiluted 

herbicide, the equivalent amount of 60 ml of 2 and 5% dilutions, 

respectively. Periodic observations were conducted and treatment 

results were recorded in May 1978. 


Reactions to treatment of firetrees with Roundup by the 

above-described methods varied from slight effects to complete 

death of trees. The results of the 2 and 5% dilution treatments 

are summarized in Table 1. 

Although variation existed among individual trees, these 

results indicate a trend toward increased effectiveness among 

those trees treated through a lower branch cut near the mainstem. 

The latter treatment resulted in general death of the tree with 

most branches being affected, whereas treatment in an upper 

distal location usually resulted only in localized death. Move- 

ment of the herbicide throughout the tree was evident only 

through limited terminal dieback of other branches. Basitone 

trees were more difficult to treat effectively through single- 

site herbicide introduction than were trees with dominant main- 

stem growth forms since the former required substantial downward 

movement of the herbicide through the treated branch to reach the 

vascular system of the other major branches. The apparent 

greater efficiency of herbicide distribution in an upward direc- 

tion indicated a close correlation with xylem transport of water 

and in some instances a high degree of early selection for 

individual branches along the mainstem. 

Variation in rates of diluted herbicide absorption was noted 

among individual trees. Complete absorption of the entire 60 ml 

quantity occurred among some trees within 2 days, although most 

required somewhat longer for complete uptake. A few trees 

(Table 1) never completely absorbed the available amount during 

the test period. Dosage comparisons involving the latter trees 

are therefore difficult to make, although effects of the solution 

amounts actually taken up were noted. 

Trees treated with small quantities of undiluted Roundup 

reacted in a manner comparable to that described for Roundup 

dilution treatments. Again, herbicide introductions proximal to 

the mainstem and in the lowermost possible position tended to be 

most effective, although variation in percentage of death among 

individual trees was again noted, presumably resulting largely 

from inconsistencies in effectiveness of internal vascular 

connections. Complete absorption of undiluted Roundup quantities 

into fresh cut branch ends was markedly and uniformly rapid,

usually requiring only a few minutes. In contrast to tests 

utilizing Roundup dilutions, in no case did undiluted Roundup 

remain unabsorbed. 

To discover the extent to which firetrees were able to 

absorb undiluted Roundup, 12-ml syringe barrels or sections of 

half-inch inside diameter rigid-walled plastic tubing were sealed 

to cut branch ends of smaller and larger trees, respectively, 

with caulking compound. These containers were regularly replen- 

ished with quantities of herbicide such that during a 45-day 

period in excess of 100 ml of Roundup were absorbed by each 

smaller tree and in excess of 300 ml by each larger tree over a 

32-day period. Rapid uptake continued even after portions of 

trees near treatment areas had developed severe visual signs of 

poisoning. These results dispel former concerns that attempts to 

introduce concentrated Roundup through a single cut branch may 

cause immediate death of the branch with subsequent inhibition of 

further uptake, resulting in ineffectiveness of the treatment 

method. 

Treatment with undiluted Roundup therefore offers no 

apparent disadvantage compared to the use of various dilutions, 

whereas the relatively small quantities required per tree and 

rapid absorption of the former through a single treatment site 

present obvious advantages. In no case was a tree killed or 

severely affected by either a diluted or an undiluted Roundup 

treatment observed to show any regenerative activity through root 

or lower stem sprouting. 

Further experimentation emphasizes determination of effec- 

tive treatment methods and herbicide quantities for trees larger 

than those here described and for those growing in other Park 

elevations and habitats. Also, alternate methods for effective 

treatment of trees unsuitable for the cut branch method are under 

investigation. 


Kim, J. Y. 1969. Myrica faya control in Hawaii. Down to Earth 

25(3) : 23-25. 

Walters, G. A., and W. S. Null. 1970. Controlling firetree in 

Hawaii by injection of Tordon 22K. USDA Forest Service Res. 

Note PSW-217.

TABLE 1. Effects of Roundup absorbed through cut branch ends of firetrees. 

Tree size 

class & 

herbicide 

concentration 

Large 

2% 

Small 

2 % 

Lower 

Proximal 

Lower 

Distal 

upper 

Proximal 

upper 

Distal 

a a number scale from 0 (unaffected) to 10 (completely dead) indicates the approximate 

degree to which each tree was affected. 

parentheses indicate a lack of complete absorption of herbicide solution from the 

container. In all other cases the solution (approx. 60 ml) was completely taken in 

by the tree.

FACTORS CONTROLLING THE DISTRIBUTION OF EXOTIC 

PLANTS IN THE KO'OLAU MOUNTAINS, O'AHU 

Grant Gerrish 




A study of native and exotic plants was conducted in two 

rain forest communities in the Ko'olau Mountains, O'ahu. One 

study area is on Mt. Tantalus in the southern Ko'olau's; the 

other, 24 miles to the north, is at Pupukea. The study areas are 

between approximately 1500 and 2000 feet in elevation and are 

within the 'Ohi'a Vegetation Zone as described by Egler (1939). 

Eighteen 400 mz plots were sampled in the more natural vegetation 

of Tantalus, 16 such plots were placed at Pupukea. All species 

of vascular plants growing in the sample plots were recorded and 

their abundance estimated. 

Of the 110 species of vascular plants found in the sample 

plots of either study area, 38 are exotic. Of these 38 exotic 

species, only a few are important in determining the structure 

and appearance of the vegetation. If frequent is taken to mean 

"occurring in at least half of the sample plots of one or the 

other study area," and abundant to mean "having cover greater 

than twenty percent in at least one sample plot," then only seven 

species of exotics are both frequent and abundant in these study 

areas. These seven are the trees: Psidium guajava, P. cattleianum, 

and Citharexylem spinosum; the smaller woody -: 

Cordyline terminalis and 

- 

Clidemia hirta; and the two grasses: 

Andro o on vir inicus and setaria palmaefolia. These species 

i r respective vegetation layers and locally 

give the vegetation the appearance of being dominated 

plants. 

by exotic 

Concern about the impact of exotic plants on Hawaiian eco- 

systems and the detrimental effects that these introductions may 

have on the endemic flora necessitates the examination of the 

ecology and behavior of exotic species and to ask the question 

"Why of the more than 4000 species of exotic plants found in 

Hawaii (St. John 1973) have these seven species become frequent 

and abundant in these rain forest communities?" Svecies which 

are obviously "trailside weeds," such as Erechtites hieracifolia, 

w i l l not be discussed in detail, since any damage to the native 

vegetation associated with the presence of these plants can be 

more directly attributed to the disturbance that allowed them to 

become established. At the same time, it is not realistic to 

discuss these ecosystems under conditions of no human-induced 

disturbance. The presence of pigs and goats in the southern 

Ko'olau's, and at least occasional human presence throughout the

ange insure that the vegetation will always be subjected to some 

disturbance. Exotic plants which can exploit this minimal level 

of disturbance are of prime interest. 

The occurrence of 11 exotic species at Tantalus which do not 

occur at Pupukea supports the hypothesis that more exotics are 

found in the southern Ko'olau's because the effects of man are 

greater at that end of the range, especially in the Honolulu 

area, to which Mt. Tantalus is adjacent. This hypothesis pro- 

poses that the distribution of exotic plants is largely a func- 

tion of seed availability and disturbance of the vegetation by 

man or recently introduced animals. Obsetvations and research 

indicate that now, as well as in the past, the Tantalus study 

area is subjected to more disturbances of these kinds than is the 

Pupukea study area, and that Tantalus has available to it a 

greater source of potential exotic invaders in the form of orna- 

mentals and other introduced plants grown in botanic and private 

gardens in the Honolulu area. 

The presence of escaped cultivars on Tantalus, such as 

Cinnamomum zeylanicum and Ilex paraguariensis, and of the ornamentals 

Ardisia crispa and Tro aeolum majus, ,clearly indicates 

that proximity to Honolu u contro111ng factor of these 

species distributions. None of these exotic species are reported 

elsewhere in the Ko'olau's. Citharex lem caudatum was introduced 

into Lyon Arboretum in Manoa -T---% 

Va ley at t e foot of Mt. Tantalus. 

This t;ee now forms dense stands on Tantalus and elsewhere on the 

periphery of Manoa Valley and has been reported as "occasional" 

in the Schofied Barracks area (USACH, unpublished). 

However, there are species of exotic plants which have dis- 

tribution patterns that do not fit the hypothesis that the spread 

of exotics is mediated by man alone. There are exotics which are 

found in the Tantalus study area and near, but not in, the 

Pupukea study area; and exotics that occur at Pupukea and in the 

vicinity of, but not in, the Tantalus study area. These distri- 

bution patterns would not be found if seed availability were the 

only factor governing the distribution of exotlc species. 

The common guava, Psidium guajava, is the most abundant 

exotic tree on Tantalus, but only a few scattered individuals of 

this species occur in the Pupukea study area. Guava occurs on 

all sides of the Pupukea study area, indicating that unavail- 

ability of seeds is not the reason this species is not abundant 

there. The distribution of Schinus terebinthifolius is similar 

to that of the guava. 

On the other hand, the noxious shrub - Clidemia hirta, another 

melastomaceous weed, Pterolepis lomerata, and the grass Andro- 

pogon virginicus, are very common =-K in t e Pupukea study area and 

elsewhere in the Ko'olau's. These three exotics are found near 

Tantalus, but not in the Tantalus study area. Considering the 

great dispersability of the seeds of all three of these species, 

it is not possible that unavailability of seed is responsible for 

the absence of these three species on Tantalus.

With respect to the distribution of some exotic plants, it 

would appear that each study area is an island surrounded by 

these species, but free of them. Distribution patterns of some 

native plants coincide with these exclusive distributions of 

exotics. On Tantalus and at lower elevations in the Pupukea 

area, Acacia koa is a co-dominant in the 'ohi'a forest. Koa is 

lackinginthe Gukea study area. Metrosideros tremuloides is 

the most common species of the genus at Tantalus; only one individual 

of this species was found-in the Pupukea study area. Many 

native species which are common at Pu~ukea are absent or of low 

These differences in the native and exotic floras of the two 

areas suggest that important environmental differences exist. 

According to published meteorological data (Voorhees 1929?; 

Taliaferro 1959), the annual amount and the monthly distribution 

of rainfall is quite similar in both areas. No major climatic 

differences exist. 

Analyses of the soils of the sample plots of the study areas 

were conducted. The soil at Pupukea is mapped as the Kapaa 

Series (USDA 1972). These highly weathered clays are classified 

as Oxisols. They are derived from Ko'olau basalt and have a high 

gibbsite content. Analyses showed the mean pH to be 4.6. Avail- 

able phosphorous was undetectable in most samples. Bases, espe- 

cially calcium and magnesium, were present only in very low 

concentrations. All analyses indicated that the soils of the 

Pupukea study area are extremely infertile. 

In the Tantalus study area, two very distinct soils were 

found . A majority of the study area is on cinder-derived soils 

of the Tantalus Series. This inceptisol is derived from ash and 

cinder of the Tantalus eruption (USDA 1972) which is dated to 

less than 100,000 years age (MacDonald & Abbott 1970). These 

soils are youthful and were found to retain higher concentrations 

of calcium and magnesium than the soils of Pupukea. Mean A 

horizon pH is 5.6 and available phosphorous was detected in all 

samples. 

Several sample plots in the Tantalus study area were found 

to be on soils derived from the ancient Ko'olau basalt rather 

than Tantalus cinder. These highly weathered clays have a mean 

pH of 4.6 with no detectable available phosphorous or calcium in 

the A horizon. These lava-der ived soils, like the lava-der ived 

soils of Pupukea, are extremely infertile. 

The influence of soil fertility on the vegetation structure 

and quantity of biomass supported at each study area can be seen. 

At Pupukea, the canopy above 2 m high is rarely more than 50% 

closed and above 5 m is approximately 70% closed. Thus, the 

forest at Pupukea can be called "open," while the canopy at

Tantalus is "closed" and of higher stature. While no measurement 

of biomass was made, it is evident that a larger standing crop is 

maintained on the more fertile cinder-derived soils of Tantalus 

than at Pupukea. 

In some cases, the distribution of exotic species can be 

directlv correlated with the canovv characteristics of the community 

.* For example, shade-loving Setaria palmaefolia and 

Commelina diffusa are found on the forest floor of Tantalus. 

Neither species occurs in the open forest of the Pupukea study 

area. At Pupukea, Andropogon virginicus locally dominates the 

qround cover under the open tree canow. *- - but this srass is not 

found on Tantalus. t he-distributions of these and-other species 

appear to be controlled by biotic characteristics of the ecosystems, 

and only secondarily by the environmental factors which 

determine the general structure of the community. 

In other cases, the effect of environmental factors is 

directly expressed. The near absence of the common guava, 

Psidium guajava, from the Pupukea study area is not a result of 

seed unavailability, lack of vegetation disturbance, or an 

unsuitable vegetation structure. The most reasonable explanation 

is that this tree can not tolerate the stress of the acid, infer- 

tile soil of Pupukea. Similarly, Clidemia hirta is in the 

vicinity of Tantalus and has been shown to be relatively shade 

tolerant (Wester & Wood 1977) but it is not found in the Tantalus 

study area. This species is very tolerant of infertile soil but 

lacks the genetic attributes needed to compete with faster grow- 

ing plants on a less stressful site. This tree appears to be 

both tolerant of soil infertility and capable of competing on 

more fertile sites. 

That soil fertility is a controlling factor in the distri- 

bution of both native and exotic plants is supported by the find- 

ing that the vegetation of the infertile lava-derived soils of 

Tantalus is more likethe vegetation-of Pupukeathan like that-of 

the more fertile soils of Tantalus. The vegetation of these 

lava-derived soils of Tantalus exhibits an open canopy; lacks a 

number of species common in other Tantalus communities, such as 

and possesses several species common at Pupukea 

on Tantalus, such as Dicranopteris linearis. 

In conclusion, it has been found that a number of exotic 

species have escaped cultivation into the native vegetation 

around Honolulu. Some of these do not appear to be aggressive; 

others, such as Citharex lem caudatum, do. Some exotic species, 

especially g r a s s e m 

have distribution patterns deter- 

mined by community biotic factors, such as degree of canopy 

cover. The distribution of other species is controlled by soil 

fertility. 

It is suggested that the difference in soil fertility be- 

tween the Tantalus and Pupukea study areas is one of several or 

many environmental barriers that can be found within a climat- 

ically similar zone of the Ko'olau Mountains. These barriers may 

effectively prevent the spread of an exotic species through the

native rain forest. However, the several exotic species that are 

capable of crossing all or most of these barriers are the plants 

that must be considered a threat to the integrity of native vege- 

tation and to the existence of local endemics. From this study 

it has been found that the most threatening exotic plants in the 

rain forests of the Ko'olau Mountains are Psidium cattleianum, 

Citharexylem caudatum, and Clidemia hirta. 


Egler, F. E. 1939. Vegetation zones of Oahu, Hawaii. Empire 

Forestry J ournal 18(1): 1-14. 

MacDonald, G. A., and A. T. Abbott. 1970. Volcanoes in the sea. 

Univ. Hawaii Press, Honolulu. 441 pp. 

St. John, H. 1973. List and summary of the flowering plants in 

the Hawaiian Islands. Pac. Trop. Bot. Gdn. Mem. 1. 519 pp. 

Taliaferro, W. J. 1959. Rainfall of the Hawaiian Islands. 

Hawaii Water Authority, Honolulu. 

USACH. Installation environmental impact statements. Botanical 

Survey. United States Army Support Command, Hawaii. Held 

in Dept. of Botany, Univ. Hawaii. (Unpublished). 

USDA Soil Conservation Service. 1972. Soil survey of the 

islands of Kauai, Oahu, Maui, Molokai, and Lanai, State of 

Hawaii. U. S. Government Printing Office, Washington, D. C. 

754 pp. 

Voorhees, J. F. 1929? A quantitative study of the rainfall of 

the island of Oahu. Paradise of the Pacific, Honolulu. 

Wester, L. L., and H. B. Wood. 1977. Koster's curse (Clidemia 

hirta), a weed pest in Hawaiian forests. Environmental 

Conservation 4(1): 35-42.

RESOURCE TRACKING PATTERNS IN ACARI ASSOCIATED WITH BIRDS 

IN HAWAII VOLCANOES NATIONAL PARK: A PRELIMINARY REPORT* 

M. Lee Goff 


Bernice Pauahi Bishop Museum 


For some time now, host-parasite relationships have been an 

area of important and frequently controversial inquiry among sys- 

tematists. Studies of Mallophaga infesting birds by Clay (1949, 

1950, 1957); the streblid batflies by Wenzel et al. (1966); and 

the Macronyssidae and Laelapidae of bats by Radovsky (1967, 1969) 

have provided examples which have led to the somewhat optimistic 

statement: "parasite phylogeny paralleles host phylogeny" 

(Kethley & Johnston 1975). In more rigorous terms this may also 

be expressed : "parasite inter-relationships are congruent with 

host inter-relationships" (Kethley & Johnston 1975). Recently 

Acari infesting birds have been studied with some emphasis given 

to host-parasite co-evolution (Kethley 1971). These Acari may be 

divided into three groups based on their interaction with the 

bird host (Fig. 1). Group I contains the host-dwelling mites. 

Here, most or all of the life cycle of the mite is spent on the 

host. Group I1 contains the nest-dwelling ectoparasites. These 

mites visit the host only to feed and spend the remainder of 

their life cycle in the nest. Group I11 mites are the field 

parasites. These mites, most notably the chiggers and ticks, are 

associated with the host only for feeding. Wide host ranges are 

typically associated with Group I11 mites. 

Group I parasites have been the primary source of data for 

the construction of patterns of radiation, as shown in Figure 2 

for the Acaridei, and for tentative phylogenies, as shown in 

Figure 3 for the parasitic Gamasina (after Radovsky 1969). It is 

of interest that in the Acaridei (Fig. 2) there is an actual dif- 

ference in the life cycle corresponding to the difference in 

habitat, as the deutonymphal stage, or hypopus, is absent from 

parasitic forms. A tendency toward reduction in stages in the 

life cycle, or tachygenesis, is common among parasitic forms. 

The house dust mites (family Pyroglyphidae) are placed in an 

intermediate position between the nest habitat and the parasitic 

habitat in Figure 2. This results from the belief that these 

* Studies upon which this report is based were supported in part 

by Cooperative National Park Resources Studies Unit Contract 

CX 8000-7-0009 to the University of Hawaii and in part by 

National Institutes of Health Grant A1 13893 to Bishop Museum.

mites are parasitic forms which have secondarily reverted to the 

nest habitat (Wharton 1976). In both Figures 2 and 3, the nest 

habitat is shown to be an intermediate step toward parasitism. 

The nest serves as a stable food source for nidicolous mites as 

well as a concentrating mechanism for their mating. In many 

instances, this association has been shown to be a forerunner of 

parasitism, for example, in the parasitic Gamasina (Radovsky 

1969). It is also of some interest to note that the bird- 

infesting Rhinonyssidae are believed derived from the bat- 

infesting Macronyssidae (Fig. 3). In this instance, strict 

adherence to the previously stated concept of host-parasite con- 

gruence would require that one derive the birds from the bats. 

With relative safety, I feel that I can say this is slightly 

unreasonable. In the past such non-congruencies have been 

explained by invoking either the "accidental transfer" or 

"historical accident," which imply some form of selection error 

on the part of the parasite. 

The possibility that something other than an accidental 

transfer was operating was suggested by Kethley and Johnston 

(1975), based on studies of quill mites of the family Syringo- 

philidae (Kethley 1971). They observed that the mites had not 

tracked their hosts as a unit through their evolution, but 

instead, topographic sub-units of the host. In the case of the 

Syringophilidae the parameters of quill diameter and wall thick- 

ness were the major determining factors in the mite population. 

Thus distantly related, or in some instances apparently non- 

related, hosts which had structurally similar feathers were 

observed to support closely related mite populations. 

With this background, it might be expected that similar patterns 

would be found in mites infesting the external portions of 

feathers. The feather mites of the superfamily Analgoidea comprise 

a complex of over 50 families. These are highly derived 

mites with varying degrees of host specificity (Krantz 1971). 

Feather mites are primarily grazers on the surface of the feather 

and do not normally appear to cause any injury to the host, thus 

large numbers of mites per host are common. Members of the 

family Analgidae are frequently noted from a wide range of hosts 

(Krantz l97l), but generally from hosts with similar feathers. 

Among species of Proctophyllodidae, most notably the genus - Proc- 

tophyllodes, a high degree of host specificity is noted, with 38% 

of the species reported from a single host species (Atyeo & 

Braasch 1966). Thus in the feather mites, there are indications 

that both co-evolution and resource tracking are present. Due to 

the relative isolation of the Hawaiian Islands, and the number of 

endemic birds present, Hawaii Volcanoes National Park presents an 

ideal situation for the study of these patterns in the birds. At 

present both endemic and introduced birds are being collected and 

processed for ectoparasites in conjunction with a study of avian 

malaria, sponsored by the Cooperative National Park Resources 

Studies Unit (CPSU).

Identifications completed to date are presented in Tables 1 

and 2. While conclusions cannot be drawn at this time, several 

undescribed taxa have been encountered and new records for hosts 

and localities are present in the data. Prior to this study only 

the families Analgesidae and Proctophyllodidae had been reported 

from the Hawaiian Islands (Garrett & Haramoto 1967). In the 

Proctophyllodidae, only the genus Proctophyllodes was reported. 

All other feather mite records are new. The recovery of spec- 

imens of cytoditids from the Red-billed Leiothrix constitutes a 

new host record. The recovery of Neharpyrhynchus sp. from an 

'Amakihi constitutes both a new host record and the first record 

of this genus from Hawai'i. 

Following completion of taxonomic studies, host-parasite 

relationships will be studied to determine which patterns are 

present in the feather mites associated with endemic birds and 

these results compared to currently available taxonomic 

structures for the species involved. 


I am indebted to Dr. Warren T. Atyeo, University of Georgia, 

for providing identifications of feather mites. Harpyrhynchidae 

were identified by Dr. Wayne W. Moss, Philadelphia Academy of 

Sciences. Birds were collected and processed under the direction 

of Dr. Charles van Riper 111.


Atyeo, W. T., and N. L. Braasch. 1966. The feather mite genus 

Proctophyllodes (Sarcoptiformes: Proctophyllodidae) . Bull. 

Univ. Nebraska State Mus. 5: 1-354. 

Clay, T. A. 1949. Some problems in the evolution of a group of 

ectoparasites. Evolution 3: 279-299. 

. 1950. A preliminary survey of the distribution of the 

Mallophaga (feather lice) on the class Aves (birds). Bombay 

Nat. Hist. Soc. 49: 429-443. 

. 1957. The Mallophaga of birds. Pages 120-156 - in 1st 

Int. Symp. on host specificity among vertebrates. 

site de Neuchatel. 

Univer- 

Garrett, L. E., and F. H. Haramoto. 1967. A catalog of Hawaiian 

Acarina. Proc. Hawaii. Entomol. Soc. 19: 381-414. 

Kethley, J. B. 1971. Population regulation in quill mites 

(Acarina: Syringophilidae) . Ecology 52: 1113-1118. 

Kethley, J. B., and D. E. Johnston. 1975. Resource tracking 

patterns in bird and mammal ectoparasites. Misc. Publ. 

Entomol. Soc. Am. 9: 231-236. 

Krantz, G. W. 1971. A manual of acarology. Oregon State Univ. 

Book Stores, Inc., Corvalis, Oregon. Pp. 1-335. 

Radovsky, F. J. 1967. The Macronyssidae and Laelapidae 

(Acarina: Mesostigmata) parasitic on bats. Univ. Calif. 

Publ. Entomol. 46: 1-288. 

. 1969. Adaptive radiation in the parasitic Mesostigmata. 

Acarologia 11: 450-483. 

Wenzel, R. L., V. J. Tipton, and A. Kiewlicz. 1966. The 

streblid batflies of Panama (Diptera: Calypterae: 

Streblidae). Pages 405-676 in Ectoparasites of Panama. 

Field Museum of Natural ~istorc Chicago, Illinois. 

Wharton, G. W. 1976. House dust mites. J. Med. Entomol. 

12: 577-621.

TABLE 1. Mites recovered from body washes of birds. 

Bird 

' Amakihi 

(Loxops virens) 

Apapane Laelapidae 

(Himatione sanguinea) Rhinonyssidae 

Mites Recovered 

Family Genus & Species 

Rhinonyssidae Ptilonyssus sp. 

Harpyrhynchidae Neharpyrhynchus sp. 

'&alo Rhinonyssidae Ptilonyssus sp. 

(Phaeornis obscurus) 

Red-billed Leiothr ix Cytoditidae 

(Leiothrix lutea) 

Cytodites sp. 

Japanese White-eye Cheyletidae Neocheyletiella sp. 

(Zosterops japonicus)

TABLE 2. Feather mites recovered £ran endemic and introduced Hawaiian birds. 

Bird Mite Family Mite &nus & Species 

' Apapane 

(Himatione sanguinea) 

House Finch 

(Carpodacus mexicanus) 

House Sparrow 

(Passer danesticus) 

'I'iwi 

(Vestiar ia coccinea) 

Laysan Finch 

(Psittirostra cantans) 

Red-billed Leiothr ix 

(Leiothrix - lutea) 

'6na'o 

(Phaeornis obscurus) 

Rice Bird 

(Lonchura punctulata) 

Japanese White-eye 

( Zosterops japnicus) 

Analg idae 

Proctophyllcd idae 

Trouessar tiidae 

Xolalgidae 

Analgidae 

Proctophyllcdidae 

Analges sp. 

Proctophyllcdes sp. 

Ptercdectes sp. 

Proctophyllodidae Proctophylldes pinnatus 

Proc tophyllcdidae Proctophyllodes troncatus 

Analgidae 

Proctophyllcdidae 

Analges sp. 

Proctophylldes sp. 

Analgidae Analges sp. 

Analg idae 

Proctophyllodidae 

~rouessar tiidae 

n. gen. & n. sp. 

Analges sp. 

Proctophylldes sp. 

Trouessartia sp. 

Xolalgidae n. gen. & n. sp. 

Analgidae Cnychalges sp. 

Analgidae Anhernial es sp. 

d u s sp. 

Pteronyssidae buchetla dofichosikya 

Trouessartidae 

Calcealges yunkeri 

Trouessartia sp.

NASAL MITES 1 

RHINONYSSIDAE 1 FIGURE 1. MITES ON BIRDS 

ASCI DAE (PHORETI C) 

EREYNETIDAE 1 

TROMBICULIDAE 111 

CYTODITIDAE 1 

TURBINOPTI DAE 1 

CHEYLETIDAE 1 

HARPYRHYNCHIDAE 1 

EPIDERMOPTIDAE 1 

KNEMIDOCOPTI DAE 1 

SUBCUTANEOUS ~UBCUTANEOUS MITES 

LAMINOSIOPTIDAE 1 

HYPODERIDAE 11 

OTHER MITES 

DERMANYSSIDAE 11 

MACRONYSSI DAE 11 

ARGASIDAE 11 

IXODIDAE 11 & 111 

TROMBICULIDAE 11 & 111 

1 HOST DWELLING 

1 1 NEST DWELL1 NG 

111 FlFln PARASITES 

ANALGOIDEA (50+ FAMILIES) 1 

QUILL MITES 

SYRINGOPHILIDAE 1 

DERMOGLYPHIDAE 1 

SYRINGOBI IDAE 1 

CHEY LETIDAE (PREDATORY) 

DOWN MITES 

ANALGIDAE 1

Hirstionyssidae Rhinonyssidae 

(Rodents & insectivores) (Bird respiratory 

Alphalaelapinae 

(Mountain beavers) I / parasites) 

\ 

/ 

Macronyssidae 

(Microchiroptera, 

/ 

secondarily mammals, 

Omentolaelapidae 

birds & reptiles) 

(1 sp. under scales 

Of African snake) / Spelaeorhynchidae 

/ Entonyssidae 

(Snake respira- 

(Ears of New World 

bats) 

Laelapinae Spinturnicidae 

(on mammals) (Winqs - of bats) 

tory 

Dermanyssidae 

. I \ I 

~Godorh~nchidae Dasyponyssidae 

(Colubrid snakes) (New World armadillos) 

(1 genus mammals;, , 

1 genus birds) . 

\ / 

\ 

\ 

\ 

/ 

/ 

/ 

. 

\ \ 

/ Myonyssinae 

/A genus on rodents, 

Hystrichonyssidae - - - - - Androlaelaps lagomorphs & insectivores) 

. I 

(Old World porcupines) 

. 

# 

0 

0 

Raillietidae ' 

(Ears of ungulates) 

,(Nest 

0 

associate) 

Hypoaspis / 

(Free-living predator) 

Haemogamassinae 

(Free-living nest 

/ associate) 

FIGURE 3, Adaptive radiation in the parasitic Gamasina 

(after Radovsky 1969) .

ALTITUDINAL EFFECTS ON THE GENERAL DIVERSITY 

OF ENDEMIC INSECT COMMUNITIES IN A LEEWARD HAWAIIAN FOREST SYSTEM 

MANUKA FOREST RESERVE, SOUTH KONA, HAWAI'I 

Samuel M. Gon I11 

Department of Zoology 



INTRODUCTION 

The amazing diversity of form and habitat in the Hawaiian 

native insect fauna is currently an object of review and study. 

Perhaps 4000 species of endemic insects have been described, and 

new species are often encountered in general collections of 

remoter areas. Thus, a pioneering stage still prevails in 

Hawaiian forest entomology. 

Many species of introduced insects (especially those of 

economic importance) have been well studied, and aspects of their 

taxonomy, physiology, behavior, and ecology are relatively well 

known. In contrast, many of the Hawaiian insects are so poorly 

understood that all the information available may be a type 

specimen and general collection information. Some species holo- 

types are defined from as little as wing fragments from a single 

specimen, never again re-encountered (Zimmerman 1948). 

One of the recent trends in Hawaiian entomology is the utilization 

of transect techniques along altitudinal gradients. 

These studies have demonstrated that species are often restricted 

to well-defined altitudinal ranges (~agn6 1976). There are usually 

habitat limitations related to altitude which define optimal 

ranges and distribution boundaries. 

This study examines the distribution of endemic insect 

families along an altitudinal gradient between 1670 and 580 rn, 

and investigates differences in the diversity of the families at 

various elevations. The trends uncovered would provide insight 

to environmental and biotic factors related to altitude, affect- 

ing the distribution of the Hawaiian insect fauna.

STUDY SITE 

The ahupua'a of Manuka, Kaulanamauna, and Kapu'a lie upon 

the southwestern flank of the Mauna Loa shield volcano in the 

South Kona district of the island of Hawai'i. These triangular 

sections of land extend downslope from their common apex at Pu'u 

Ohohia (1690 m) along the Southwest Rift Zone of Mauna Loa, fan- 

ning outward to include a 12 km expanse of shoreline at their 

bases (USGS topographic quadrangles 1967). 

Transects and sample stations 

METHODS 

Field study was conducted from June 15 to July 31, 1977. 

The survey of insect fauna was one aspect of a holistic ecosystem 

baseline survey of the Manuka area conducted under the auspices 

of the National Science Foundation. 

Twelve sampling stations were established along two parallel 

transect access lines in altitudinal increments of 290 m running 

from Pu'u Ohohia downslope. The arrangement of sampling stations 

is illustrated in Figure 1. This placement established two rep- 

licate stations at the 1440, 1150, and 875 m elevations, and one 

replicate at the 585 m elevation. The study area boundaries 

precluded establishing replicates at the 1690 m level, while 

commercial agriculture at one of the 585 m plotsites made consid- 

erations of sampling at that location moot. 

Whenever possible, the entomology sampling station plotsites 

coincided with that of the Manuka Research Project vegetation 

ecology survey sites, in order to have available detail-ed vege- 

tation data. 

Sampling techniques 

In this study, only the night fauna was considered. Night 

collection commenced at 7:00 PM and ran continuously to 10:OO PM. 

Each night collection utilized a simple sheet light trap. A 

Coleman-type lantern provided attractive illumination. A1 1 

insects that came to rest upon the sheet were collected during 

the 3-hour period, and kept for identification. Large insects 

were collected with sweep nets, while smaller insects were 

aspirated into vials. 

Data analysis 

The number of families encountered at each plot were cumu- 

lated, and mean values for isoaltitudinal plots were computed. 

The trends in the diversity of insect communities at the family

level were analysed against altitude distributions of the sample 

station plotsites via correlation coefficients generated by: 

= S x ~ where 2xy (covariance). = 1 (Cxiyi - 1 XxiCyi) 

SxS y n-l n 

and 2, = hi2 - (CX?) '/n j .-I 

- sy = jiyi2 - (~yi)~/n 

n - 1 

where 2, and sy are standard deviations, x and y being values of 

two discrete and changing factors: alfitude and the number of 

insect families encountered. 

RESULTS 

There was a general linear increase in the number of 

families collected with decreasing altitude (Table 1). Correlation 

coefficients calculated using the mean number 

collected at isoaltitudinal plots substantiated 

of families 

this linear 

relationship (r 

n = 5). 

- = -0.72, - 

With decreasing altitude, greater numbers of families appeared 

during the first half of the collection periods. At the 

1690 m station, 50% of families encountered were collected by the 

midpoint of the collection period. At 1150 m, this figure had 

increased to 70%. while at 875 m it was 79% and at 585 m, the 

insects collected by the first half of the collecting period 

amounted to 84% of the total yield. The rise in this temporal 

packing correlated with altitude (r - = -0.74, - n = 12). 

An interesting exception to the linear relationship of both 

family diversity and temporal packing trends ocurred at the 

1440 m elevation, where both diversity and packing exceeded figures 

characteristic of the 1150 m plots, and were only slightly 

lower than values at the 875 m level. This "hump" is easily seen 

in Figure 2. If the data from the 1440 m plots are deleted, and 

correlation coefficients are again calculated, the linear fit 

against altitude is far more precise (family diversity/altitude 

- r = -0.82, - n = 9; temporal packing/altitude - r = -0.91, - n = 9). 

The distribution of entomology sites, vegetation ecology 

relevi stations, habitat descriptions, and climatic observations 

along the altitudinal gradient are presented in Figure 3. The 

number of vascular plant species and insect families encountered 

at identical sites are compared in Figure 4. The data is uti- 

lized in the discussion to assess vegetational correlations and 

to incorporate several situations to interpret both general 

trends and the "hump" phenomenon.

DISCUSSION 

The trend of increasing diversity in insect communities with 

lower altitude that was uncovered in the study area was expected 

by this author. Major dimensions critical to the survival and 

fitness of insect species, such as temperature and moisture, ap- 

proached more favorable conditions with decreasing altitude. In 

addition to these environmental aspects, the effect of vegeta- 

tional diversity can be considered. 

Environmental Aspects 

Temperature 

On the even, lee slope of the Mauna Loa shield volcano, mean 

temperature decreases with altitude. It is not unusual to flnd 

morning frost at the 2000 m level, while at sea level, nights are 

often uncomfortably warm. There is some evidence that temper- 

ature effects contributed to limiting the upper range of many 

insect species in this study. The total number of specimens per 

collection increased with decreasing altitude, and the numbers of 

insect families which appeared by the first half of the collec- 

tion period increased from five families (50% of the total fam- 

ilies collected) at 1690 m, to 18 families (84% of the total 

families collected) at 585 m. Clearly, not only were there more 

insect families encountered, but more total specimens collected, 

and a much greater relative activity at lower elevations. The 

difference in activity was especially notable. At 1690 m, less 

than 100 specimens were collected, and long periods of inactivity 

were prevalent by the end of the collection period, when night 

temperature was estimated at 5°C. In contrast, at the 585 m 

plotsites, several hundred specimens were collected, and near the 

end of the collecting period, the collectors were hard-pressed to 

keep up with the insects clustered on the sheet and swarming 

around the light. The estimated temperature at the end of the 

night collection there was 15°C. 

Beetles in the family Scolytidae were not encountered except 

below 875 m, where they were quite common. Scolytid beetles are 

known in Metrosideros wood, and Metrosideros sp. exists as the 

dominant macrophanerophytic species in the entire study area. 

Thus, food limitations cannot be imposed. Increasing moisture 

conditions toward 1150 m is not a limiting factor. Scolytids are 

known from rain forests as well as mesophytic forests. Predation 

is a factor to consider; however, bird species such as Loxo s and 

Himatione exist at both low and higher altitudes, and d n c i- 

dence of predacious and parasitic insects increases with lower 

altitude. It seems likely that temperature is the limiting 

factor. Little is known of the temperature tolerances of the 

Hawaiian insects, and because no systematic temperature readings 

were taken in this study, no conclusions can yet be made.

Moisture 

The climatic pattern along the Kona Coast of the island of 

Hawai'i is clearly defined from the predominant tradewind situa- 

tion in the Hawaiian Islands. Due to the massive obstacle of the 

Mauna Loa shield volcano which negates tradewind influences, the 

Kona (leeward) flank climate is determined largely by a diurnal 

convective cycle. 

In the day, land heat-induced winds pull moisture-laden 

ocean air up the slope of Mauna Loa. An inversion layer near 

1100 m induces cloud formation in a band above this level, where 

highest precipitation occurs. Above and below this level, aver- 

age rainfall decreases along gradients (Blumenstock & Price 

1967). From the barren cinder above. 1700 m, moisture levels 

increase downslope toward fog forest at 1440 m, and mixed meso- 

phytic forest below this to approximately 300 m, where near-xeric 

conditions prevail. Within the altitude range of the study, how- 

ever, moisture conditions are seen to increase downslope. If 

moisture is considered as a limiting factor, then the good agree- 

ment between moisture gradients and insect family diversity leads 

to the conclusion that the increase in moisture and the increase 

in insect diversity are related. 

In a study of the distribution of canopy-associated arthro; 

pods along a transect on the windward slope of Mauna Loa, Gagne 

(1976) suggested that factors contributing to the distribution of 

the more restricted arthropods would appear to be related to cli- 

mate. For example, in his study, the exotic detritivorous roach, 

- Allacta similis, was apparently excluded from higher montane 

environments, a phenomenon which ~agn; attributed to cooler 

temperature and greater moisture in the lower rain forest (below 

1800 m). In the same manner, he noted that the detritivorous 

tree cricket, Paratrigonidium spp: , predominated in moister, 

warmer sites at the lower portlons of the transect (below 

1500 m). .In a moregeneral 

-- 

trend, Gagne found , that arthropod ~~. 

species diversity in Acacia koa tree canopies was relatively high 

at low and mid-elevations, but decreased markedly with altitude, 

which he attributed to climatic causes. 

In the Manuka study, moisture conditions were considerably 

different from the Mauna Loa findings, where tradewind influences 

push the inversion layer higher, and mesic conditions prevail to 

about 200 m, with rain forest environments restricted to below 

1400 m. In the Manuka study area, rain forest environment does 

not exist, and mesic conditions exist in a band of precipitation 

largely below 1400 m and above 400 m. We would expect that the 

upper altitudinal limit, if it is determined by relative humid- 

ity, would be lower than those observed along the Mauna Loa 

transect. 

In the day collections at Manuka, Allacta similis was not 

found above 1150 m plotsites. In nocturnal collections, the 

crickets, ~aratr igonidium spp., likewise were limited to below 

1150 m. A rarer, brachypterous ground cricket, Leptogryllus sp., 

was found only at 860 m, despite concerted searches of leaf

litter habitats at lower and higher altitudes. Although both 

Paratrigonidium and Le to r llus 

scleroterized, 

Le to r llus is an especla ly soft bodled insect. The tendency 

xeric conditions at lower elevations, and the 

tendency toward colder, xeric conditions at higher elevations 

seem to have restricted the range of this fragile, wingless, 

detritivorous cricket. 

q_pyy__ zre , 

Desiccation pressure and the alleviation of this factor at 

lower altitudes seems likely to play a role in determining the 

increase of diversity seen in the study area. However, until 

determinations can be made about the optimum moisture conditions 

for any Hawaiian insect species, we have no quantitative indica- 

tion of moisture-related limitations of insect distribution in 

the Hawaiian systems. 

Biotic Aspects 

Vegetational diversity 

Temperature and moisture regimes determine the distribution 

of endemic plant species. Krajina (1963) described 14 biogeo- 

climatic zones, elaborating on the works of Rock (1913), 

Ripperton and Hosaka (1942), Fosberg (1961), and others, com- 

piling ecological observations and studies of topographic, 

geological, climatic, and biotic factors. Five of these zones 

exist in the Manuka study area, reflecting a leeward forest 

pattern. At lower altitudes (300-470 m) open mixed xerophytic 

and mesophytic forest grades into closed mixed mesophytic and 

xerophytic forest (470-850 m) which in turn gradually passes into 

mesophytic marine tropical and subtropical forest (850-1470 m) 

and rather suddenly passes into open mixed mesophytic and xero- 

phytic scrub forest (1470-1690 m) and finally to open xerophytic 

scrub (1690 m to subalpine and alpine elevations). The area has 

been largely characterized as a "dry transitional forest," based 

upon the vegetational community gradients, but the diversity of 

the communities is affected by differences in substrate, making 

the area far more complex. 

Aerial photographs of the study area taken in 1962 show that 

the substrate varies in both age and basic composition. Unweath- 

ered 'a'a and cinderfalls may be found alongside sections of 

older, weathered substrates with good soil developement. Dif- 

ferential vegetation type is seen to correspond to differing 

substrata. 

Results of a vegetation survey and ecological study con- 

ducted concurrently with the entomology collections of this study 

show that vegetational diversity changed with altitude. For 

example, at the 1690 m site, a plant species count tallied a 

maximum of 15 vascular plant species. At 1440 m, 52 vascular 

plant species were collected, and at 1150 m, 35 vascular plant 

species were encountered. At 875 m, 37 vascular plant species 

were tallied, and at 585 m, 39 vascular plant species were

counted at the main transect relev6s. When the correlation coefficient 

r was computed for insect family diversity and plant 

species count, a strong correlation was seen (r = 0.811, - n = 10). 

The relationship between the number of plant species and the 

number of insect families at identical plotsites reflects the 

correlation (Fig. 4). 

Two major factors may be responsible for the relationship: 

trophic relationships between insects and plants, and the effect 

of spatial heterogeneity on species diversity. 

The major relationship between insects and plants is trophic, 

and thus the ecology of the vegetation has a direct bearing 

on insect ecology. This would be most evident in stenophagous 

insects (e., insects with narrowly limited diets). Swezey 

(1954) compiled an annotated checklist of the insect faunas of 

Hawaiian forest plants, making note of stenophagous species. The 

percentage of stenophagous insects ranged from an 11% incidence 

of stenophagy (on Acacia koa), to a 72% incidence of stenophagy 

(on Pelea spp.). ~ o r m * r i t ~ of endemic plant species, however, 

the incidence of stenophagy ranged between 30% and 45%, and 

the mean incidence of stenophagy in the 

insects in 36.55%, a sizeable percentage. 

Hawaiian herbivorous 

The various Hawaiian plant species are host to a diverse 

number of insect species, ranging from eight known associated 

species on Osteomeles sp. to more than 128 known to feed on 

Acacia e. Each plant species in the vegetational community 

could "contribute" its complex of associated insects to the total 

insect fauna. In the same manner, the absence of a plant species 

in a given area would mean that its stenophagous insect comple- 

ment would be missing from the fauna. In our study area, the 

coleopteran families Bostr ichidae, Cerambyciidae, and Carabidae 

were restricted to the 875 m plotsites and lower. The homopteran 

families Cicadellidae and Cixiidae likewise become extremely com- 

mon below 875 m. Both families' predominance seem to be related 

to a change in vegetation below 940 m. Psychotria, a tree 

species in the family Rubiaceae becomes common in the canopy. 

Various dryland tree species such as Antidesma, Drypetes, and 

Diospyros make their appearance below 800 m and occur in their 

highest frequency just below 600 m. The increase in the number 

of tree species may explain the appearance of the coleopteran 

families, notable wood borers of various Hawaiian trees. The 

prevalence of the homopteran families may be related to the prev- 

alence of Psychotria, from which both cixiids and cicadellids 

were collected in large numbers in day collections. 

The predatory families Chrysopidae, Braconidae, and Bethyl- 

idae become prominent in the lower altitude plotsites, although 

the chrysopid lacewings and the braconid wasps were present at 

much lower numbers at higher plotsites. It is likely that either 

prey species became more numerous at lower altitudes, or that 

biomass of the prey population became greater. Both situations 

were seen to occur. The increase in predatory and parasitic 

families can be considered an indirect result of the increase in

plant species diversity. The more diverse the vegetational com- 

munity, the greater is the potential for species packing in the 

consumer community and likewise in the entire trophic network. 

In addition to trophic relationships, increased vegetational 

diversity results in an increased spatial heterogeneity. There 

is an increase in the complexity of the physical environment. 

For example, the higher abundance of Psychotria sp. in lower 

altitude plotsites means that there are differences in the trophic 

niche, habitat changes in the litter layer, unique resiance 

opportunities for day-inactive insect species, a new bark habitat 

for the psocopteran and orthopteran families, etc. Ip short, the 

more complex the physical environment becomes, the more complex 

the plant and animal communities supported, and the higher the 

Species diversity. MacArthur (1965) suggested that betweenr 

habitat diversity is a major scheme in determining tropical 

species packing. For example, MacArthur and MacArthur (1961) 

determined that the extent of foliage stratificatio~ in a forest 

community was more important than the species diversity of the 

vegetational community alone in affecting the faunal diversity. 

In the Manuka study, foliage stratification increased with lower 

altitude, as conditions improved for tree species. A t the 1150 m 

plotsites, the canopy of Metrosideros was non-interlocking and 

rose to about 6 m. Forest conditions developed by 875 m, however, 

with densely interlocking canopies of Metrosideros and 

Psychotria, reaching crowns at 25 m. The substantially tall 

canopy created adequate room for a well-developed middle-story of 

Cibotium tree ferns, M rsine lessertiana, and Vaccinium calycinum, 

and an understory ---T o ferns and small vascular plants. In 

the585 m plotsites, species diversity was even higher, and the 

complexity~of foliage stratification was very well developed. It 

is not surprising that the increase in species diversity and 

foliage stratification downslope in the study area is paralleled 

by a correspondant 

community. 

increase in the diversity of the insect 

- . - 

Combination of factors 

All of the factors discussed this far cannot be realis- 

tically considered independently. The combination of environ- 

mental aspects and biotic factors is a dynamic process, the 

synthesis of which is the final characteristics of insect distri- 

bution we have observed. For example, moisture characteristics 

probably determined the basic vegetational diversity at 875 m; 

however, the presence of vegetation can create microclimates in 

which moisture and temperature conditions are quite different 

from adjacent, barren areas at the same altitude. As a result, 

the increase in the complexity of the environment created by 

microclimates would allow for a higher potential in species 

diversity. What will be discussed next is an example of the 

resultant effect of a combination of environmental and biotic 

factors: the "hump" phenomenon at 1440 m.

The "hump" phenomenon 

The exceptionally high diversity of the 1440 m plotsite col- 

lection can be considered in terms of the interaction of several 

factors which exist at that elevation. It can be seen that per- 

haps three aspects unique to the 1440 m plotsites could contri- 

bute to the "hump" phenomenon: moisture conditions, vegetational 

diversity, and kipuka effects. 

An inversion layer fog belt which exists from 1080 to 1860 m 

creates a high humidity situation without heavy precipitation. 

It has been suggested that for some species of Hawaiian insects, 

the physical effect of rain showerg may restrict their presence 

in zones of precipitation (Gagne 1976). In addition, high air 

moisture creates favorable conditions for the growth of fungus, 

and results in suitable habitats for detritivorous and fungivorous 

insects, which constitute a considerable percentage of 

Hawaiian forest insect communities (~agn6 1976). In our study, 

the dipteran families Dolichopodidae 

restricted between 1160 and 1500 m. 

and Cecidomyiidae were 

The importance of fog drip as a major mode of water uptake 

in the Hawaiian forest has been documented (Juvick & Perreira 

1973). In the Manuka study, we found that not only is there a 

more luxuriant vegetational situation created, but vegetational 

diversity is highest in the fog zone. 

In addition, the 1440 m plotsite sampling was conducted in a 

kipuka of moderate size. Generally, a kipuka, or regional uncon- 

formity (Pukui & Elbert 1971) is a section of vegetation sur- 

rounded by the relative infertility of fresher lava substrate. 

In many cases, the change in species diversity and community 

Structure of vegetation moving from barren lava to within the 

lush kipuka environment promises to show consistent patterns, 

allowing the kipuka concept to be defined ecologically. Yoshi- 

naga and- Anderson -(1977; unpub; ms .)-studied the kipuka systems 

of the Manuka-Kapu'a area and found that because of the insular 

character of kipukas, their habitat differs from continuous 

stands of old substrata as well as from that of the surrounding 

fresh lava. Compared to the open lava, the more weathered kipuka 

substrate has better moisture-holding capacity, more available 

nutrients, and better rooting opportunities. The more closed 

canopy offers shelter for species unable to tolerate the hot, dry 

open environment of the surrounding undecomposed lava. Litter 

may collect more effectively than in the open. Along the perim- 

eter of a kipuka exists a strip of edge habitat, which can 

support species which might be unable to grow in closed forest. 

Just outside the kipuka is a boundary habitat, where the harsh 

environment of the lava flow may be somewhat ameliorated by the 

effects of the kipuka such as shade, fog drip, and leaf litter. 

The plant species list for soil kipukas in our study area was 

similiar to a combined species list for open lava and continuous 

stand vegetation on soil. Thus, in the kipuka situation, a 

combination of high plant species complement, higher potential

vegetational situation, and habitats unavailable to both sur- 

rounding lava and continuous stand forest, can be considered as 

factors contributing to an increased insect diversity. 

In summary, Figure 3 illustrates the diversity of habitat 

a ~ d climatic conditions which contribute to the "hump" phenomenon 

at 1440 m: fog conditions, general vegetational diversity, and 

kipuka effects. In addition, Figure 3 demonstrates gradients in 

moisture and vegetation that are pertinent to the general 

relationship of altitude and insect diversity uncovered in this 

study. 

CONCLUSION 

In the Manuka-Kapu'a study, nocturnal insect diversity at 

the family level increased as altitude decreased, with the 

exception of a disproportionately high diversity at the 1440 m 

elevation. The general altitude effect was attributed to the 

interrelation of three major factors: temperature, moisture, and 

vegetational diversity. Although it is clear that temperature 

and moisture conditions contributed indirectly to insect distri- 

bution by determining vegetational diversity and structure, it is 

not known what direct limiting effect thay have on the Hawaiian 

insect species. It is recommended that physiological studies be 

conducted, focusing on the tolerances of Hawaiian species to a 

wide range of temperature and moisture conditions, determining 

survivorship thresholds and optimum ranges. The information from 

such studies could be used to test whether temperature and 

moisture limitation is a mechanism directly determining the 

distribution of Hawaiian insects in field studies. 

The exceptionally high diversity at 1440 m was attributed to 

three major factors not present at other-plotsite locations: fog 

belt conditions, unique vegetational species complement, and the 

effects of kipukas.


Blumenstock, D. I. ,and S. Price. 1967. Climates of the states: 

Hawaii. US Government Printing Office, Washington, DC. 

Fosberg. 1961. 

~agn6, W. C. 1976. Canopy associated arthropods of Metrosideros 

and Acacia koa on the Mauna Loa Transect, 

Ecosystems m, US/IBP Tech. Rep. No. 77. 

Hawaii. 

32 pp. 

Island 

Juvik, J. O., and D. J. Perreira. 1973. The interception of fog 

and cloud water on windward Mauna Loa, Hawaii. Island 

Ecosystems IRP, US/IBP Tech. Rep. No. 32. 11 pp. 

Krajina, V. J. 1963. Biogeoclimatic zones on the Hawaii 

Islands. Haw. Bot. Soc. Newsletter 2(7): 93-98. 

MacArthur, R. H. 1965. Patterns of species diversity. Biol. 

Rev. 40: 510-533. 

MacArthur, R. H., and J. W. MacArthur. 1961. On bird species 

diversity. Ecology 42: 594-598. 

Pukui, M. W., and S. H. Elbert. 1971. Hawaiian dictional: 

Hawaiian-English, English-Hawaiian. Univ. of Hawaii Press, 

Honolulu. 188 pp. 

Ripperton , and Hosaka. 1942. 

Rock, J. F. 1913. The indigenous trees of the Hawaiian Islands. 

Private publication, under patronage. Honolulu. 

Swezey. 1954. 

U.S. Dept. of the Interior, Geological Survey. 1967. 

Topographic Quadrangles. 

Zimmerman, E. C. 1948. Insects of Hawaii Vol. 1-9. Univ. Press 

of Hawaii, Honolulu. 

References to Yoshinaga and Anderson 1977 are in preparation: 

NSF-SOS Program, Technical Report for Grant No. NSF SOS SMI 

76-07608.

TABLE 1. mmber of families encountered in night collec- 

tions of isoaltitudinal plotsites, Manuka Forest 

Reserve, South Kona, Hawai'i (A = main transect; 

B = auxilliary transect; K = Kapu'a transect). 

- - 

Transect Total x by Total by X by 

Plotsite Families Altitude 8:30 P.M. Altitude

FIGURE 1. 

Arrangerent of tratlsect lines and platsites, Mm& Fbrest -, 

South Xona, Hawai'i. 

zone in the ahupua'a of Kapu'a. 

-- ----- transect access trail

KEY: 

Nmhr of families 

colleded fran: 

@ Kapu'a transect 

8 Auxilliary (B) transect 

0 Main transect 

Mrmber of families Vertical solid 

collected fran all and hatched lines 

transects appearing denote range of 

by 8: 30 PM (midpint values. 

of collecting period) 

I 

FIGURE 2. The n&r of insect families encountered in night collections oj 

isoaltitudinal plotsites, plan& Forest Reserve, South &ma, Hawaili. Imu 

diately notable is a general decrease of family diversity with increase in 

altitude. Exever, data collected from the 4600 ft (1400 m) plotsites do 

not fit vd.1 in the txend, creating a "hq" phen-n. This "hmp" is 

mst pronounced in data frcnn the rain transect collections, but is present 

in all transect collections. Ihe solid lines connect rrean values f m all 

plotsites and derronstrate both the general trend and the deviance m the 

4600 ft (1400 m) plotsites. The 

blackened data points damnstrat 

that the percentage of £&lies 

appearing by the midpint of tk 

collection period decreases witk 

altitude, and is similar1 

diswrdant at the 4600 ft 

(1400 m) level. 

I T, I I 1 I I I I , I 

2000 3000 4000 5000 

: ELFVATIa (feet abve sea level)

(smwu) ze3 uo~wat3-s~a~a~ ~ = G I u o w w h 

(0691) OOPS (00~~)' 009P (0SIT) I OOLE (sL~) ~OEZ (58s) 'SL~T 

(sxrw~) 7333 uo?wam--savs ~ T - W 

I 1 I I 

(0691) OOPS (OOP'C) 009P (OS'CT) OOLE (SL.8) 0082 (585) ISLET 

I 1 I I 1 I I I I I I 

009 

ooLr OOST OOET (sm=) ooor 008 

'JJFlMaT3 

000s OOOP (w33) OOOE oosz oooz 

I I I I I I I 1 I I I I " """'1"""'"I"""'



MOSSES OF THE CRATER DISTRICT 

William J. Hoe 




The biological distinctiveness and the lack of serious bryo- 

logical collecting in Hawaiian alpine areas was recognized as 

long ago as 1930. At that time Edwin B. Bartram, who would 

later publish the Manual of Hawaiian Mosses (1933), wrote to Otto 

Degener (dated September 30) and expressed the opinion that "The 

most likely places for new and interesting additions will be 

around the rim of Haleakala and above 6 or 7000 ft on Mauna Loa 

and Mauna Kea..." Since then, collections by Degener and others, 

primarily in conjunction with more generalized surveys, have 

tended to bear out these predictions. 

The Resources Basic Inventory (RBI) surveys, conducted 

during the summers of 1975 through 1977, have provided an oppor- 

tunity to study the moss flora of upper Haleakala as well as the 

distribution of the taxa. The intensive collecting in 55 repre- 

sentative sites has yielded what is probably a complete picture 

of the moss flora. With the taxonomic basis now understood, 

future research could include investigations of the moss commu- 

nities present and their relationships to the general vegetation 

as well as physiological adaptations to the rigors of Hawaiian 

alpine conditions. 

Identification of the mosses collected during the survey is 

virtually completed, and will result in a technical report summa- 

rizing the taxa present, their general distribution and phyto- 

geographic relationships. It would perhaps be most appropriate 

at this time to summarize general relationships of the mosses of 

upper Haleakala and to discuss a few of the phytogeographically 

significant species. 

Bartram, in introductory remarks to his Manual, concluded 

that the affinities of the Hawaiian moss flora lay almost exclu- 

sively with the region to the southwest, i.e., to the Indo- 

Pacific region. Gemmell (1955), in further analysis of Bartram's 

data, came to the same conclusion. Based upon the incomplete 

data then available, these conclusions were certainly correct. 

However, availability of more recent collections, particularly 

from the poorly-known Hawaiian alpine areas, has shown that the 

desert-like areas above the tree line contain a surprisingly 

large and diverse flora. The sometimes abundant representation 

of genera such as Andreaea, Encalypta, Grimmia, Ptychomitrium,

Racomitrium, and Tortula are not at a11 reminiscent of Indo- 

Malesian or even of tropical floras but of the northern and 

southern hemisphere temperate regions instead. The non-endemic 

species and close relationship of many of the endemics further 

suggest Boreal relationships. The purely Austral elements are, 

in fact, represented by only a few taxa. Table 1 summarizes the 

phytogeographic relationships of Haleakala's alpine species. Of 

the 36 taxa which can be identified with confidence, 15 (42%) are 

represented in both the North and South temperate areas, 13 (36%) 

are Boreal, with only 3 (8%) Austral. With the possible excep- 

tion of three endemic species in the "unknown" category, there is 

no relationship with the Indo-Pacific floras. This suggestion 

should not be as surprising as it may first seem. In terms of 

dispersal distance the Hawaiian Islands are considerably closer 

to Boreal than to possible Indo-Pacific or Austral sources of 

diaspores for its alpine flora. Both the jet stream and the 

trade winds originate in the Boreal regions. 

One may reasonably ask, then, about the origins of the three 

austral Haleakala taxa. Amphidium tortuosum, although widely 

distributed in the temperate Southern Hemisphere, seems to have 

migrated northward along-the American cordilleras, reaching into 

Central America. The local uo~ulations, therefore, may well be 

descendent from American rathe; than ~ust;al sources. -~ortella 

fra ilis var. tor telloides, originally described from Antarctica, 

h r e 

and in Hawai'i' may simply represent stress forms 

resulting in similar morphological responses rather than one 

being derived from the other. That is, the two populations are 

probably not the result of long distance dispersal. Andreaea 

acutifolia, known from such Austral areas as the Falkland 

Islands, Auckland Islands, Campbell Island, New Zealand, 

Kerguelen, and southern South America, is generally considered to 

be variable and very close to the extremely variable and cosmo- 

politan Andreaea rupestris: Whether it deserves specific or even 

varietal recognition is eing investigated. 

The Boreal representatives clearly outnumber the Austral 

(>4:1) in the Hawaiian alpine. The odds, then, would seem to 

favor Boreal origins for the majority of the mosses of cosmo- 

politan-temperate affinities. If this hypothesis is correct, 

then Haleakala's alpine flora is basically of Boreal origin and 

has little relationships with the downslope, primarily Indo- 

Pacific rain forest species. 

The Crater District of Haleakala National Park is inter- 

esting to a bryologist not only because of its alpine flora but 

because it contains upper rain forest representatives in areas 

such as Paliku. Areas such as Paliku are, in many ways, clearly 

transitional. They serve as the upper boundary for the many taxa 

of the lowland forest which are only sparingly represented and as 

the lower l i m i t for alpine taxa which are present only in exposed 

sites. There is, however, a surprisingly large number of Boreal 

forest elements present. New Maui or Hawaiian Islands records of 

this type discovered during the past three summers include

Isopterygium Plagiothecium cavifolium, Tr ichostomum 

tenuirostre, Le flexifolium ana Orthodontium pellucens, 

among others. The flrst three are new state records, and may 

eventually be found on other islands as well. 

The lower and middle Hawaiian rain forests are undoubtedly 

the best known regions bryologically. The mosses of such areas 

are often obvious and abundant. Historically, these forests have 

been the easiest of access. The phytogeograhpic affinities of 

these species generally lie with Oceania and SE Asia. In very 

simple terms, these species require all but brief periods of con- 

stant moisture and high humidity. They are probably also frost 

intolerant. Although the Paliku area is probably sufficiently 

wet for at least most of these rain forest taxa, the cold air 

draining from the Crater and the surrounding slopes may well 

represent the single most important factor in limiting their 

presence. The rain forest taxa w i l l be discussed in the tech- 

nical report: I would like to emphasize that they are only spar- 

ingly present and are never as abundant as they would be further 

downslope in rain forest areas. 

The remnant dryland forest of eastern Kaupo Gap has a number 

of structural similarities with the dryland forest of the 

Wai'anae Mountains of O'ahu. Both have a well-developed canopy 

cover, with a nearly absent understory and herbaceous ground 

cover. Several moss species, common in the Wai'anae Mountains, 

in the Park are confined to the Kaupo Gap dry forests. These 

include Entodon solanderi and ~issidens intermidius. 

In addition to the Boreal forest elements discussed and the 

locally attenuated lower and middle rain forest taxa, the Paliku 

area contains an upper rain forest montane element. This general 

habitat and associated species assemblage is found on all of the 

Hawaiian Islands between 4000 to 6000 feet. In the absence of 

quantitative data, about all that one can say is that there is a 

clear change in the genera and species which predominate when 

compared with the lowland forest. This difference is suffi- 

ciently marked to be noticed in the field by a person familiar 

with the lower forests. Unlike the alpine, in which Boreal 

representatives predominate, and the lowland-middle elevation 

forests in which the Indo-Malesian elements are the most impor- 

tant, the upper rain forest is comprised of species from 

American, continental Asian as well as Indo-Pacific sources. 

These remarks on the mosses of Haleakala are clearly only 

introductory. They are intended primarily to point out the 

uniqueness and the value of the Crater District from a scientific 

as well as a resource management point of view and to encourage 

future studies. At this time, it is impossible to provide 

definitive answers to the questions this brief review must have 

raised.


Bartram, E. B. 1933. Manual of Hawaiian mosses. 8. P. Bishop 

Museum Bull. 101, Honolulu. 275 pp. 

Gemmell, A. R. 1955. A preliminary study of the Hawaiian moss 

flora. M i t t . Thur. Bot. Ges. 1: 71-86.

TABLE 1. Phytogeographic relationships of Haleakala's alpine 

moss flora. Taxa preceeded by an asterisk are con- 

sidered endemic. 

A. Cosmopolitan B. Boreal 

C. Austral D. Unknown or Other 

1. Amphidim tortuosum * 1. Orthotrichum hawaiicum 

2. Andreaea acutifolia * 2. Pohlia baldwinii 

3. Tortella fraqilis * 3. Pofilia mauiensis 

var . tortelloides * 4. -mitrium mauiense 

(Central American) 

5. Bryum ceramiocarpum 

(Andean Venezuela)


4. THE HAWAIIAN LAVA TUBE ECOSYSTEM 

F. G. Howarth 




Lava Tubes as Life-Support Systems 

How and why an animal would lose its eyes, color pattern, 

and other characters, and restrict itself perpetually to the rig- 

orous environment of caves has long intrigued biologists. Much 

is known of the ecology of limestone caves in temperate regions 

(Vandel 1965; Barr 1968). The realization that lava tubes also 

harbor an analogous specialized fauna is more recent (Torii 

1960). In addition to Hawai'i, significant lava tube faunas are 

now known from the Galapagos (Leleup 1967, 1968), Japan (Ueno 

19711, and North America (Peck 1973). In Hawai'i the fact that 

at least seven native groups have independently evolved troglo- 

bitic species on at least two islands indicates that adaptation 

to lava caves is a general process. 

Troglobites (obligatory cavernicoles) are restricted to the 

true dark zone of caves. The dark zone environment, as outlined 

by Howarth (1973) is similar to that described by Poulson and 

White (1969) for temperate limestone caves. It is a rigorous one 

defined by the absence of light; the relative constancy of tem- 

perature at or near the average annual temperature of the region 

and of relative humidity above the physiological limits of most 

terrestrial animals; and the absence of many environmental cues. 

There is generally a rocky substrate and often an illusion that 

food is scarce. 

Lava tubes are destroyed by erosion in a relatively brief 

geologic time. However, they are continually being created 

during volcanic eruptions since oceanic volcanoes are charac- 

teristically built with vesicular basalts which often flow as 

pahoehoe, and such flows almost always build lava tubes (Peterson 

& Swanson 1974). Lava tubes are a common land form on younger 

oceanic islands. Further , since the voids in basaltic lavas 

offer some avenues for subterranean dispersal from older caves to 

younger caves, we can expect that a specialized cave fauna will 

develop wherever new basaltic lava flows continually cross older 

flows over a long enough period of time, and the climate allows 

the continuous colonization of caves.

In Hawai'i rain water percolates rapidly into the young 

porous basalt. Only where the water table is near the surface is 

significant water found in the cave. This occurs near the coast 

and a remarkable aquatic fauna, including many troglobites, in- 

habits isolated coastal pools of brackish water (anchialine hab- 

itat) in young lava flows (Holthuis 1973; Maciolek & Brock 1974). 

In younger lava tubes the substrate is usually barren lava 

rock. However, this can vary considerably in texture, e.g., a 

polished glazed surface, irregular pile of breakdown blocks, 

highly vesicular porous lava, and even ashy rubble. The absence 

of organic detritus is often surprising. Lava tube slimes which 

represent insipient soil formation often cover large areas. In 

the oldest caves clay and soil have filled most smaller voids and 

cover the floor in many areas. 

Limestone caves also have a rocky substrate, often as irreg- 

ular blocks, but there is a greater variety of minerals including 

crystals, biogenic minerals, and alluvial sediments. A fine 

residual silt from solution of limestone is characteristic, and 

is closely associated with terrestrial troglobites (Barr & Kuehne 

1971). 

- - 

The main enerav sources in Hawaiian lava tubes are plant 

roots, especially 'ohi'a (Metrosideros collina var. 01 mor ha), 

slimes deposited by organically rich percolating groun -r, 

and accidentals which are those animals that blunder in. In contrast, 

the main energy sources in continental limestone caves are 

trogloxenes, especially bats and crickets, and debris washed in 

with sinking streams, especially during floods. Additional 

energy is supplied by accidentals, percolating ground water, 

autotrophic bacteria, and aerial plankton. 

The greater overburden of limestone caves often precludes 

the importance of tree roots and, .except for Paulian and Grjebine 

(1953) who discovered an intermediate cave-adapted cixiid in 

Madagascar (Synave 1954), most biospeleogists have disregarded 

roots in their surveys. However, the discovery in Hawai'i of a 

cave cixiid (Howarth 1972) stimulated other researchers to check 

roots in caves. Recently Fennah (1973b) described three species 

of troglobitic fulgoroids, two from Mexico and one from Aus- 

tralia, and Peck (1975) listed an undescribed species from 

Jamaica caves. 

The absence of native trogloxenes in Hawai'i may be related 

to the absence of winter and of a need to hibernate, and also to 

the fact that the continental trogloxenic groups did not colonize 

Hawai'i. Hawai'i's onlv bat and only native land mammal 

(Lasiurus cinereus semot;s) is a forest species and is not known 

to enter caves. 

Sinking streams are not important energy sources in Hawaiian 

lava tubes. It is unusual for a stream to enlarge a lava tube; 

rather, it speeds the siltation and erosion processes. The few

lava tubes visited that had captured a temporary stream were 

shortly blocked with silt, had signs of periodic flooding, and 

had a poor fauna. 

Cave Fauna 

Native cavernicolous animals are predominantly arthropods. 

In Kazumura Lava Tube, 10 troglobitic and seven native troqlo- 

philic (facultative cavernicoles) species occupy nine general 

niches: three primary consumer niches, feeding on living and 

dead roots; one omnivore; two predatory niches differing in 

strategies e . , with and without snares); one sarcophagous 

niche; and two saprophagous niches, one feeding on fungus and one 

generalist. There is broad niche overlap as underscored by 

omnivore niche occupied by the troglobitic cricket. 

Only one species of root penetrates into Kazumura Lava Tube 

to any great extent, and five native and four exotic arthropods 

feed on it. The troglobitic cixiid which is probably the most 

abundant arthropod in the cave is a sapsucker. The living root 

chewers are represented by three species of moths of the genus 

Schrankia. One of these is a weak flier and appears to be trog- 

lobitic. An undescribed troglobitic millepede occupies the 

living and dead root feeding niche. 

At the top of the food chain is a large striking troglobitic 

wolf spider which stalks its prey and does not build a web. The 

other five native predators are less well known and probably live 

in cracks and rarely enter larger passages. A troqlobitic ter- 

restrial water treader scavenges on dead arthropods. The two 

saprophagous niches are occupied by four troglophilic taxa. 

To date no native organisms have been found boring into or 

specifically feeding on large diameter roots, and this is pos- 

sibly an empty niche. However, in arthropod surveys it is diffi- 

cult to generalize on negative evidence, i.e., I can only say I 

have not found it, not that it does not exist. As explained by 

Janzen (1977) natural functioning ecosystems have few if any 

empty niches unless there is a major new disturbance, such as 

that created by an exotic species. 

Even though 15 exotic species have colonized Kazumura Lava 

Tube, with one exception (snare building predators) they do not 

appear to have invaded the niches to a great extent. 

On the younger flows on Hawai'i Island the troglobites have 

a wide distribution, but many lowland troglobites have not been 

found above 1000 to 1500 m where other species often occur. 

Roots are more important in higher caves. The few lava tubes on 

Maui are significant because their fauna provides a control group 

for Hawai'i Island studies. At least six native groups have 

independently evolved troglobitic species on the two islands.

Most continental troglobites are considered relicts of past 

climatic changes, especially glaciation and changes in sea level 

(Vandel 1965; Barr 1968; Mitchell 1969). However, many of the 

Hawaiian troglobites have close surface relatives still extant. 

Three such species pairs, Caconemobius varius--C. fori; Oliarius 

golyp- hemus --0. . inaequalis; and ~ e s i d i o- m aLa-rselium have 

een recocinizgd ((Fennah 1973a: ~aan6 & Howarth 1975: Gurnev & 

Rentz 1978; ~oharth 1979):' ~iiis strongly suggests that tioglobites 

are relicts only if the surface species have become 

extinct, not that they become caveadapted after extinction of the 

surface population. There are many examples of adaptive radiation 

among continental cave animals, but-most apparently are true 

relicts due to the more complex geological history of the 

continents. 

Cave Perturbations and Maintenance Trends 

Continental caves are often viewed as islands and their ecosystems 

share an apparent fragility in response to perturbations. 

Cave ecosystems on islands may be in double jeopardy and several 

of the newly discovered arthropods are candidates for endangered 

species status. What then is the future of this unique ecosystem, 

not even recognized before 1971? If perturbations had 

caused its demise sometime during the last 200 years, biologists 

would have continued to believe no such fauna had ever existed in 

Hawai ' i . 

On Hawai'i Island there are still many avenues of dispersal 

between lava tubes and continual new flows can be expected; 

therefore, one can expect the survival of the cave fauna, barring 

any major catastrophies. On the older islands of Maui and Kaua'i 

the caves are eroded remnants, many of the avenues of dispersal 

are closed through erosion, and the cave animals lead a tenuous, 

threatened, or endangered existence. 

The major perturbations facing the Hawaiian cave ecosystems 

are as follows: (1) destruction of the forest by grazing ani- 

mals, fire, exotic plants, agriculture, and urbanization; (2) 

creation of new entrances and increased siltation and filling of 

caves by the erosion resulting from the above activities; (3) 

colonization by exotic animals; (4) use of caves for refuse 

disposal; and (5) direct disturbance by human visitation. 

(1) Since the main energy source is plant roots, the de- 

struction of the overlying forest removes this energy source. 

The obligatory primary consumers die off rapidly and the food web 

shrinks to a few scavengers and predators feeding on accidentals. 

A few primary consumers, notably Schrankia and the millepede, can 

switch to some exotic root species. 

(2) Troglobites are restricted to the true dark zone where 

the relatively constant environmental conditions are maintained. 

New entrances introduce surface climatic influences and the cave

fauna is destroyed. Further, the erosion of the surface causes 

rapid filling of the voids in the lava and eventually the cave 

itself. 

(3) The introduction of exotic species, or biological pollu- 

tion, is perhaps the most insidious perturbation because it is 

normally irreversible and it often pervades the ecosystem in 

unforeseen ways. The impact of exotic species will be discussed 

below. 

(4) Using caves for refuse dumps hastens the filling of the 

cave and introduces large amounts of food that alters the cave 

food web in several ways. (a) In relatively closed caves it can 

foul the air and kill the inhabitants. (b) By composting it can 

raise the cave temperature and dry the cave. (c) Most impor- 

tantly, it often allows the colonization of the cave by oppor- 

tunistic scavengers and predators. The high populations of these 

being supported on refuse can swamp the endemic biota. 

Kaua'i has few extant lava tubes and the two known troglo- 

bites are among the most bizarre discoveries to date. Regret- 

tably, the fields with the largest caves known on Kaua'i were 

covered by 5 m of sugarcane bagasse shortly before I visited the 

area. The caves are now gone, the fauna extinct, and no one will 

ever guess what that fauna might have been! 

The entrance to Offal Cave, a relatively large lava tube on 

Haleakala Volcano, Maui, was used as an offal pit by the local 

slaughterhouse, and the tallow, rotting bones, and other garbage 

are piled high near the entrance and scattered throughout the 

downslope portion of the cave. The cave is no longer used for 

this purpose, but even now the cave ecosystem approaches that 

reported for large bat caves in tropical continental areas, where 

a huge amount of animal matter is introduced into the cave and 

supports a large population of many troglophiles. In Offal Cave 

these troglophiles are almost all exotic carrion feeders, scav- 

engers, and predators, including such groups as cockroaches, ear- 

wigs, ants, moths, spiders, millepedes, and isopods. This is the 

only cave where many of these organisms have invaded any signif- 

icant distance from the entrance, and it is assumed that it is 

the rich, novel food supply that allows them to colonize the zave 

environment. Only one endemic troglobite, the omnivorous cricket 

Caconemobius howarthi, occurs in this section of the cave. 

Unfortunately, grazing has destroyed all native trees over 

Offal Cave and no roots now penetrate into the deep cave, so that 

it remains hypothetical whether other troglobites may have sur- 

vived the influx of offal and associated biota if the rest of the 

ecosystem were intact. I believe most would not have survived, 

as such a large amount of organic matter would have heated the 

cave and dried or otherwise affected the environment. Such a 

large influx of exotic predators, sustained by the high popula- 

tion of exotic scavengers, also would have preyed on any cave 

species so that few endemics would have survived.

(5) Caves are fragile ecosystems and, like other discrete 

geologically defined habitats such as montane bogs and sand 

dunes, are easily disturbed by human visitation. Normal weath- 

ering processes are so changed and attenuated that even foot- 

prints can remain for centuries. In Hawai'i careless or 

destructive visitors kill or break tree roots, mark walls, litter 

the cave, and trample animals and their habitats. Tobacco smoke 

is a strong insecticide and smoking in the enclosed cave environ- 

ment may be lethal to the fauna. The heat from both the body and 

a torch, if used, can dry the cave. Any smoke also introduces a 

large number of condensation nuclei to the saturated cave atmos- 

phere. Littering is related to the using of caves as refuse 

dumps as discussed above. Cave visitation by the public should 

be discouraged until adequate protection of sample caves and 

ecosystems is assured. 

Impact of Exotics 

Many of the arthropods recently introduced by man, espe- 

cially household pests and soil forms coming in with plant 

materials, have successfully colonized certain Hawaiian lava 

tubes. Such animals as the cockroaches, centipedes, millepedes, 

isopods, spiders, and other groups have been successful in low- 

land caves. Some of these exotics have surely altered the 

ecology of the caves, but it is unknown whether any replaced 

native species in the cave ecosystem. This is the region most 

disturbed by man. Many exotic species of roots penetrate these 

caves and mostly exotic species occupy these exotic niches. 

These are also the caves littered by visitors and used for dumps, 

as explained above. 

Exotic scavengers also exploit the dead accidental exotic 

mammals (roof rat and mongoose) in Kazumura Cave. Barr and 

.. Kuehne. . (1971 ) reported . a . slmil~ar ph.er?om.enon . . in . . Mammoth. Cave, 

Kentucky. They felt that litter from human activities has 

allowed the colonization of the cave by troglophiles 

not otherwise be able to do so. 

that would 

Of the nine niches found in Kazumura Lava Tube the two pred- 

atory niches showed the greatest intrusion by exotics, followed 

by the saprophagous niches. In this cave only one species of 

root is present and so restricts potential phytophagous invaders. 

With an increase in species diversity there is a greater chance 

of an exotic or native species finding a suitable niche. 

Thus those secondary and tertiary consumers that are gener- 

alists have the highest diversity of prey or food to choose from, 

and it follows that these niches would have the most species, 

both native and exotic, in most habitats. The invasion by an 

exotic does not imply that there had been an empty niche but that 

the invader was able to create one. 

One exotic spider, Nesticus mogera from Japan, is common in 

mid- to high elevation caves on Hawai'i Island where its sloppy 

inverted webs are found between adjacent protuberances and in

cracks in the walls in nearly the same situation as one sometimes 

finds the rare troglobite Erigone stygius. Both species are 

about the same size and both build similar sized webs. Although 

their webs are quite different, they probably capture similar 

prey. Even though N. mogera may not be as well adapted to the 

cave environment as E. styqius its cave population is constantly 

being augmented by- individuals from surface habitats. As the 

prey is depleted the native spider loses. This circumstantial 

evidence implies that - N. mogera is replacing the endemic species. 

Other exotic troglophiles have not colonized Hawai'i's 

caves. For example, the predatory snail Euglandina rosea is a 

common troglophile in its home region in Florida. In Hawai'i its 

shells are common in low to mid-elevation caves but apparently 

the absence of both calcium for its shells and suitable prey have 

prevented this species from colonizing island caves. Other 

examples from Offal Cave are discussed above. 

With one exception exotic trogloxenes have not yet become 

established. Attempts to introduce cave bats to Hawai'i in the 

1920's were unsuccessful (Tomich 1969). The Edible Nest Swiftlet 

was released in 1962 and is now established in a small area on 

O'ahu (Shallenberger 1976). 

Five main groups of trogloxenes exist in the world. Bats 

and rhaphidophorine crickets are nearly worldwide in caves except 

at high latitudes and on a few oceanic islands. Cave rats 

(~eotoma spp.) are widespread in North America. The Oilbird 

(Steatornis caripensis) inhabits some Neotropical caves, and cer- 

- - 

tain Swifts (familv A~odidae) nest in caves of the Old World 

tropics. These groups ca;ry in organic matter as food, excre- 

ment, nesting material, and dead bodies. 

The advent of trogloxenes in Hawai'i will result in a major 

new energy source in the caves and the establishment of a new 

food web, drawing almost entirely on exotic organisms for its 

cycle. 

Had a trogloxene colonized Hawai'i naturally before man, the 

cave ecosystem would certainly have been greatly altered, but 

native species would have eventually adapted to exploit the new 

niches. The present situation is quite different, as a great 

many potential exotic troglophiles that live in guano caves else- 

where are already established as inquilines of man or his domes- 

tic animals or as soil animals. These species w i l l be able to 

invade and flourish in Hawai'i's caves when a food source is 

there, and few native animals will have a chance to adapt to the 

new conditions. A preview of this phenomenon was described above 

for a cave that was used for an offal dump, and the ecosystem was 

drastically a1 tered with only one native species surviving. 

I suggest that the apparent fragility and instability of 

island ecosystems when compared to continental ecosystems is more 

related to the degree, type, or harshness of the perturbation, 

than to some inherant weakness in the workings of the system. 

Evolution dictated that island ecosystems functioned well before

the current onslaught. It is true that given the disharmonic 

nature of island biota, those missing groups that exploit their 

environment in an innovative way w i l l drastically alter the 

island ecosystem when they are introduced. Examples are troglo- 

xenes in Hawai'i's caves and grazing mammals which convert forest 

to grassland. But these groups were missing once on the conti- 

nents, too. Did not the forests of the eastern great plains of 

North America give way to the bison and man, as did the eastern 

Mediterranean forests to the goat? Most examples on the conti- 

nents have been obscured because of the complex geological and 

evolutionary history there. 

One of the reasons islands are so interesting biologically 

is that they have not had as complex a biological and geological 

history as the continents. Islands can be studied as experi- 

mental controls for the evolutionary and ecological processes 

which are occurring but are obscured on the continents. 


This work was supported by NSF Grant No. GB23075 ISLAND ECO- 

SYSTEMS IRP/IBP HAWAII and by separate grant to the author, NSF 

Grant No. DEB75-23106. I thank Dr. D. Mueller-Dombois, Depart- 

ment of Botany, University of Hawaii at Manoa, for assistance in 

analyzing the data.


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lobites. Evol. Biol. 2: 35-102. 

Barr, T. C., and R. A. Kuehne. 1971. Ecological studies in 

the Mammoth Cave system of Kentucky. 11. The ecosystem. 

Annales de Speleologie 26(1): 47-96. 

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tubes, 4. Go new blind Oliarus (Fulgoroidea: Cixiidae) . 

Pacif. Ins. 15: 181-184. 

. 1973b. Three new cavernicolous species of Fulgoroidea 

omop opt era) from Mexico and western Australia. Proc. Biol. 

Soc. Wash. 86: 439-446. 

Gagnd, W. C., and F. G. Howarth. 1975. The cavernicolous fauna 

of Hawaiian lava tubes, 7. Emesinae or thread-legged bugs 

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the description of one new genus and four new species. 

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of Hawaii. Science 175: 325-326. 

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Hawaii Island: an ecosystem supported by wind borne debris. 

Pacif. Ins. 20: 133-144. 

Janzen, D. H. 1977. Why are there so many species of insects? 

Proc. XV Int. Cong. Entomol. 1976: 84-94. 

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aux iles Galapagos. Noticias de Galapagos, nos. 5-6, 1965: 

4-16. 

+ 

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Galapagos et an Ecuador (N. et J. Leleup, 1964-65) Rgsultats 

scientifiques. Koninklijk Museum voor Midden-Africa=Mus&e 

Royal de L'Afrique Centrale. Premiere Partie: 9-34.

Maciolek, J. A., and R. E. Brock. 1974. Aquatic survey of the 

Kona Coast ponds, Hawaii Island. Sea Grant Advisory Report 

UNIHI-SEAGRANT-AR-74-04: 73 pp. 

Mitchell, R. W. 1969. A comparison of temperate and tropical 

cave communities. Southwestern Naturalist 14: 73-88. 

Paulian R., and A. Grjebine. 1953. Une campagne speleologie 

dans la Reserve naturelle de Namoroka. Naturaliste Malgache 

V: 19-28. 

Peck, S. B. 1973. A review of the invertebrate fauna of vol- 

canic caves in the western United States. Bull. Natl. 

Speleol. Soc. 35: 99-107. 

. 1975. The invertebrate fauna of tropical American 

caves, Part 111: Jamaica, an introduction. Int. J. 

Speleol. 7: 303-326. 

Peterson, D. W., and D. A. Swanson. 1974. Observed formation of 

lava tubes during 1970-71 at Kilauea Volcano, Hawaii. 

Studies in Speleology 2: 209-222. 

Poulson, T. L., and W. B. White. 1969. The cave environment. 

Science 165: 971-981. 

Shallenberger, R. J. 1976. Avifaunal survey of North Halawa 

Valley. 'Elepaio 37: 40-41. 

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galeries souterraines du systeme de Namoroka (Hemiptera- 

Homoptera). Le Naturaliste Malgache z(l953): 175-179. 

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Special Publication 57. Bishop Museum Press, Honolulu. 

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troglobionts of Japanese caves. (I). Jap. J. Zool. 12: 

555-584. 

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san . I. Introductory and historical notes. Bull. Natl. 

Sci. Mus. Tokyo 14: 201-218, pls. 1-4. 

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Oxford. 524 pp.

DESCRIPTION OF A NEW LARGE-SCALE 

VEGETATION MAPPING PROJECT IN HAWAI'I 

James D. Jacobi 

U. S. Fish and Wildlife Service 

Off ice of Endangered Species 


and 




INTRODUCTION 

Vegetation mapping is a method which is often used to dis- 

play the distribution of plant communities or vegetation types in 

a two-dimensional format. It entails the delineation and 

description of more-or-less homogeneous patterns of the vegeta- 

tion as interpreted usually either from aerial photographs or on 

the ground. The degree of homogeneity in the map units depends 

on what types of vegetation characteristics are being viewed, and 

what the overall purpose of the map is. 

Two research projects are currently being conducted in the 

native forests of Hawai'i, for which it was considered essential 

to have an adequate map of the vegetation patterns in the dif- 

ferent study areas. One of these projects, the Hawai'i Forest 

Bird Survey conducted by the U. S. Fish and Wildlife Service 

(USFWS), is attempting to determine the status and distribution 

of the native forest birds, with emphasis on the listed rare and 

threatened species on all of the islands. Field work for this 

survey is presently being conducted on the island of Hawai'i. 

In the second project, the 'Ohi'a Forest Study, aspects of 

the dynamics of the native montane rain forests are being exam- 

ined in detail, with particular emphasis on the phenomenon known 

as the 'ohi'a dieback. This project, directed by Dr. Dieter 

Mueller-Dombois of the University of Hawaii, was funded from 

1975-1977 by a grant from the National Park Service. 

An attempt was first made to utilize existing vegetation 

maps for these two projects. However, none of the available naps 

were found to be suitable for this purpose. Therefore, a new 

vegetation mapping project was initiated which will eventually 

cover the native forest areas on all of the major islands. This 

mapping project is supported primarily by the USFWS; however, 

additional support has come from the 'Ohi'a Forest Study for work 

on the windward side of the island of Hawai'i where the study 

areas over lapped.

In the fallowing paper, a quick summary is made of the types 

Of vegetation maps which currently exist, particularly for the 

island of Hawai'i, and the new mapping project is describe.j in 

detail. 

Discussion of Map Scale 

A major factor to be considered when working with vegetation 

maps is the scale at which they were produced. A small-scale map 

Shows units which are rather generalized and includes a consid- 

erable amount of variation. A large-scale map, on the other 

hand. shows map units which are quite detailed and includes con- 

siderably less variability. Table 1 gives a general summary of 

the different ranges of map scales and indicates the kinds of 

information which each can display. 

Review of Some of the Vegetation Maps Which Have been Published 

'for Hawal'l 

There have been numerous different vegetation maps pub1i:;hed 

for Hawai'i, some depicting the general vegetation on all of the 

islands, while many others deal with small specific areas in 

greater detail. 

Probably the most familiar map is one published by Ripperton 

and Hosaka (1942) entitled Vegetation Zones of Hawai'i. This map 

fits into the intermediate-scale range of maps with all of the 

islands except Hawai'i mapped at approximately 1:500,000. The 

island of Hawai'i was mapped at the scale of 1:1.5 million, so 

all of the islands could be included on a single small map sheet. 

Ten different vegetation zones are distinguished on this map 

which depict a combination of both actual and potential vegeta- 

tion coverage as determined mainly byclimatic aria eaaphic coiidi- 

tions. This map is useful for getting a general overview of the 

vegetation; however, it is difficult to work with in any detail 

on the ground. 

Several other maps are available at this general scale which 

are very similar to Ripperton and Hosaka's, most notable being 

maps published by Knapp (1965) and Lamoureux (1973). 

At the large map scale range, all areas on all of the major 

Hawaiian Islands were mapped at 1:62,500 by Honda and Klingen- 

smith (19631, as part of the Hawaii Forest Type-Map series pro- 

duced by the U. S. Forest Service and the Hawaii Division of 

Forestry. The map units in this case describe (a) land use 

class, (b) forest type (i.e., tree species composition), (c) den- 

sity of tree cover, and (d) tree stand size class in terms of 

sawtimber classes. These vegetation types were interpreted' from 

aerial photographs taken in 1954, but were compiled with only a 

minimal amount of ground verification.

Another large-scale map was published by Mueller-Dombois and 

Fosberg (1974) which describes the vegetation types of Hawaii 

Volcanoes National Park (HVNP). The vegetation patterns in this 

case were also interpreted from the 1954 photographs, and were 

ground checked in detail in the more accessible area. The map 

units were based primarily on dominant species and structural 

criteria (such as plant spacing and height) of the vegetation. 

Both the Hawaii Forest Type-Maps and the HVNP map are at the 

map scale at which actual patterns of the vegetation are dis- 

played with sufficient detail to serve as base maps for the Fish 

and Wildlife Service Forest Bird Survey and the study of the 

dynamics of the 'ohi'a rain forest. Unfortunately, the Hawaii 

Forest Type-Maps were not ground checked adequately and are, 

therefore, too inaccurate for practical use. The HVNP map, on 

the other hand, is much more accurate for what is displayed. 

However, it covers only a small portion of the forests in which 

we were interested. Additionally, since it was based on photo- 

graphs taken nearly 25 years ago, it is unusable in areas in 

which the vegetation has changed considerably, particularly as 

the result of land clearing, and of 'ohi'a dieback in the 'Ola'a 

Tract forest section of the Park. 

We, therefore, decided that the best way to approach the 

problem at hand was to produce a new vegetation map series, also 

at the large-scale level. 

The Current Mapping Project 

In the current project the vegetation types in most habitats 

dominated by native species on all of the major Hawaiian Islands 

will be mapped. The Fish and Wildlife survey was initiated in 

the summer of 1976 on the island of Hawai'i, and we expect to 

finish field work on Kaua'i in the summer of 1981. An attempt is 

being made to keep the vegetation mapping running concurrently 

with the field survey work for each of the different study areas. 

Figure 1 shows the areas which w i l l be mapped for the island 

of Hawai'i. To date, the map for one area, the Ka'u Forest, has 

been finished and is currently being published (Jacobi, in 

press). The preliminary mapping has been completed for the 

Hamakua, Waiakea, 'Ola'a, and Mauna Kea sections, and the final 

versions for each of these areas will be completed in the very 

near future. Currently, field work is being concentrated in the 

Hualalai and Kona regions. 

Description of the Map Units 

In this new map series, the vegetation is displayed at two 

levels of resolution. The first level shows the distribution of 

the general plant associations which in this case are defined by 

the predominant species composition of the dominant vegetation 

layer. So far at least 10 different general plant associations

have been identified which include, for example, alpine and sub- 

alpine scrub, grassland, 'ohi'a forest, 'ohi'a-koa forest, and 

tree fern dominated communities. For this purpose, the tree 

layer was considered to be dominant in open and closed forest 

stands. 

The second level of resolution describes the vegetation in 

much greater detail. In this case, four major components of the 

vegetation are taken into account in determining the map units: 

(1) tree canopy crown cover, (2) tree canopy height, (3) dominant 

species composition of the tree layer, and (4) understory or 

groundcover composition. Tables 2 and 3 show the possible attri- 

butes fo each of the vegetation components. An example of a 

vegetation type symbol is shown in Table 4. 

Mapping Procedure 

Preliminary map units are first delineated on aerial photo- 

graphs with the aid of a mirror stereoscope. Several types of 

aerial photos have been used for the different areas mapped so 

far. For the Ka'u Forest and Mauna Kea maps, the 1965 black and 

white EKL series photos (Soil Conservation Service) at the 

approximate scale of 1:24,000 were used. For the Hamakua, 

'Ola'a, and Waiakea maps, two sets of photographs were used, one 

being a set of true color photos at the approximate scale of 

1:12,000 taken in 1972 for the State Division of Forestry, and 

the other a set of color infrared photos, roughly at the scale of 

1:50,000, taken by NASA in 1974-1975. Recently the U. S. Geolog- 

ical Survey has released an excellent set of air photos covering 

all of the island of Hawai'i and most of the other islands. 

These black and white photos were taken in 1976-1977, and are at 

the approximate scale of 1:40,000. We plan to use this series of 

photographs for most of the future mapping work on this project. 

Once the preliminary mapping for an area has been completed, 

the boundaries on the photographs are compiled into an undis- 

torted map overlay at the scale of 1:24,000. This involves 

optically transferring the vegetation boundaries onto a rectified 

(i.e., corrected for photo distortion) base map. 

Field Verification of the Map Units 

One of the most important steps in preparing any type of map 

is verification of the map units in the field. For this project, 

the preliminary map units are checked in two ways: from the air 

in a small airplane or helicopter, and on the ground. The air 

reconnaissance has proved to be extremely valuable for getting a 

tree-top view of the different vegetation types. Many of the 

problem areas identified in the preliminary air photo mapping can 

be resolved in this way.

Verification on the ground is carried out along transects 

running mauka-makai at 2-mile intervals through the study area 

(Fig. 2). These are the same transects on which the bird census 

is conducted by the Fish and Wildlife survey teams (Scott 1979). 

The advantage to working along these transects, besides increased 

access into the forest, is that sampling points called "stations" 

have been regularly and accurately located along each line 

(12 stations/mile) . 

Once the preliminary maps have been corrected, new overlay 

maps are drawn which w i l l be overlain onto USGS topographic quad- 

rangle maps for final publication. For ease of use, the scale of 

the final printed maps will be reduced to 1:48,000. 

A#plications of the Vegetation Maps to Other Studies in the 

ative Forests 

The major objective in producing this new series of vegeta- 

tion maps is to relate forest bird distribution to vegetation 

types, and to provide a framework on which to study the dynamics 

of the 'ohi'a rain forest. I expect, however, that the map 

series w i l l be useful to other persons and agencies whose work 

involves the native forests, particularly in such areas as eco- 

logical and geological research, and land use planning, most 

notably connected with preservation of our natural areas. 

The major reason for this paper, therefore, is to make more 

people aware of this project, and to solicit additional sugges- 

tions which may increase the overall applicability of the maps to 

other types of studies.

LLTERATURE CITED 

Htmda, N., and J. Klingensmith. 1963. Hawaii Forest Type-Maps 

(1:62,500). USDA, Pacific Southwest Forest and Range Exper- 

iment Station, and State of Hawaii, Division of Forestry. 

Jacobi, J. D. 1978. Vegetation map of the Ka'u Forest and 

adjacent lands, island of Hawai'i. 

Knapp, R. 1965. Die Vegetation von Nord-und Mittelamerika und 

der Hawaii-Inseln. Gustav Fischer Verlag, Stuttgart. 

373 pp. 

by A. 

Translation of Vegetation of the Hawaiian Islands 

Y. Yoshinaga and H. H. Iltis, Hawaiian Botan. Soc. 

Newsletter 14(5): 95-121. 

Lamoureux, C. H. 1973. Plants. Pages 63-66 - in R. W. Armstrong, 

ed. Atlas of Hawai'i. The University Press of Hawaii, 

Honolulu. 

Mueller-Dombois, D., and H. Ellenberg. 1974. Aims and methods 

of vegetation ecology. John Wiley and Sons, New York. 

547 pp. 


Hawaii Volcanoes National Park (at 1:52,000). CPSU/UH Tech. 

Rep. 4. (Univ. of Hawaii, Botany Dept.) . 44 pp. 

Ripperton, J. C., and E. Y. Hosaka. 1942. Vegetation zones of 

Hawaii. Hawaii Agric. Exp. Sta. Bull. 89. 

Scott, J. M. 1979. Forest birds survey of the Hawaiian Islands. 

- In C. W. Smith, ed. Proceedings, Second Conf. in Natural 

Science, Hawaii Volcanoes National Park. CPSU/UH (University 

of Hawaii, Botany Dept.). (In preparation).

TlBLE 1. Sumiuy of ranges of map scales and the types of vegetation information which they 

can display. (Pdapted £ran Wller-Dcanbois and Ellenberg 1974). 

W SCALE TYPE SCALE RANGE 

( approx. ) 

TYPES OF INFOFMATION DISPLAYED MINIMUM UNIT SIZE 

1. Small scale 1: >million* Generalized potential vegetation ' 2500 ha 

(*1 an = >lo Ian) 

2. Intermediate 1:l mill to 1:100,000* Regional maps, potential 2500-25 ha 

(*1 an = 1 Ian) vegetation associations 

3. Iarge 1:100,000 to 1:10,000* Generalized actual plant 25-.25 ha 

(*1 an = 100 m) associations 

4. Very large 1:10,000 to 1:100* Detailed plant associations, 2500 mz-1 m2 

(*1 an = 1 m) Individual trees 

5. Chartmaps 1:

TABLE 2. Cmpnents for the tree layer which are used in the vegetation map symbols. 

TREE CANOPY CRW COVER 

d = Dense; '85% cover 

c = Closed; >60-85% cover 

o = Open;>20-60% cover 

s = Scattered trees; C20% cover 

TREE SPECIES CCMRXITION FORMAT 

A-B 

A-B, C 

Other possible 

forms 

Only sp. A present; canprises 

all of the crown cover indicated 

for this layer 

Species A dominant comprising 

560% of indicated cover; species 

B present but comprising 20-40% 

of indicated cover 

Species A and B codaninant, each 

comprising 40-60% of indicated 

cover 

Species A and B codaninant, with 

species C present, comprising 

20-40% of indicated cover. 

A-B-C; A, B, C 

TREE CANOPY HEIW 

1 = Short-stature trees; 3-5 m tall 

2 = Moderate-stature trees; 5-10 m tall 

3 = Tall-stature trees: > 10 m tall 

TREF. SPECIES NAME ABBREVIATICNS 

- 

Ac = Acacia koa 

Ch = Cheircdendron trigynun 

ELI = Euphorbia sp. 

Is = Introduced species 

Me = Metrosideros collina 

MK = Myrsine lessertiana 

My = Myoprun sandwicense 

So = Sophora chrysophylla

TABLE 3. Canpnents for the ground cover which are used in the vegetation map. 

SPECIES mOUPS 

The format for listing ground cover is the tq = sedges, rushes, grasses, and herbs 

sane as for listing tree species composition in boggy situations 

unless otherwise noted. the around < cover is ds = native alpine or subalpine shrubs. 

&suned to cover >60% ' 

(styphelia , Vacciniun ,- Dodonaea, . 

Geranim, mbautia, etc.) 

If the ground cover is

TABLE 4. Example of a vegetation type symbol showing the interpretation of the different 

symbol canponents. 

1. Tree Crown Cover 3. minant Tree Species Composition 

l 0 3 C 

2. Tree Canopy Height 5. Other Information 

1. nee crown cover > 20-60%. 

2, Canopy height >10 m. 

4. i round Cover 

3. Tree canopy dominated by Wtrosideros collina. 

4. Ground cover codaninated by native rain forest shrub species and herbaceous plants growing 

in boggy situation with some tree ferns. 

5. Canopy trees in "dieback" condition at time of mawing (i.e., with many starding dead or 

defoliated trees).

* 3 a: 

4: 0 0 4 

W I * f n z - 0 

Y a 4: 4 1 

4: W I I d 3 

1 4 Y Y C a Y 

a z 4 : 4 : 4 4 : < a 

T 3 T - < Z 3 Z Z a I 

0 4 : 4 : < J ~ 4 : 0 0 3 0 

Y T ~ E O ~ Y Y Y I ~

ETGLTRE 2. Location of the U. S. Fiah and Wildlife Service's Forest Bird Survey 

transects on the island of Hawai'i.

THE IMPACT OF THE SWEET POTATO 

ON PREHISTORIC HAWAIIAN CULTURAL DEVELOPMENT 

Michael W. Kaschko and Melinda S. Allen 

Department of Anthropology 



ABSTRACT 

Recent research in Hawaiian archaeology suggests 

that commencing about 1000-1100 A.D. substantial change 

and development occurred in various aspects of prehis- 

toric Hawaiian culture, and this trend continued in at 

least some respects to European contact. It is clear 

that the sweet potato was of increasing importance to 

Hawaiian agriculture during this time. The New World 

origin of the sweet potato, as opposed to the Asiatic 

nature of the other Oceanic cultigens, further presents 

the possibility of a separate and post-initial settle- 

ment introduction of this crucial crop into the prehis- 

toric Hawaiian agricultural complex. 

The possible temporal alternatives for the arrival 

of the sweet potato in Hawai'i are considered. The pre- 

dictable effects of sweet potato introduction at various 

points in the prehistoric sequence are examined in 

relation to the actual evidence for major cultural 

developments: agricultural expansion-intensification, 

population growth, increasing social complexity, etc. 

Specific methods are suggested for incorporation into 

future archaeological research to provide data relevant 

to the role of the sweet potato in the prehistoric 

Hawaiian adaptation. 

Early Western explorers, beginning with Captain James Cook 

in 1778, commented repeatedly on the extent, intensity, and quality 

of native Hawaiian sweet potato (I omoea batatas [L.] Lam.) 

production, particularly in contrast ---ii to ot er areas of Polynesia 

(Yen 1974: 311-317). As late as 1823 the Reverend William Ellis 

described extensive agricultural field systems on the island of 

Hawai'i where sweet potato was the primary crop (Newman 1970: 

112-120). These early accounts clearly attest to the importance 

of sweet potato in the Hawaiian economy at European contact. 

Late prehistoric Hawaiian culture was characterized by a complex 

ranked society, high population density, and an economy primarily 

reliant upon well-developed agricultural systems. The possible 

role of the sweet potato in the development of these cultural 

features is our concern.

The sweet potato is unique in that it is the only Polynesian 

cultigen of American origin (Brand 1971; Yen 1971, 1974). All 

other Polynesian cul tigens are of Asian derivation. Although 

taro is characterized as the preferred food of the Hawaiians, as 

a crop sweet potato has several advantages (Handy & Handy 1972: 

124-128) : 

1) It can be grown in less favorable locations with respect 

to sunlight and soil. 

2) The tubers mature in three to six months as opposed to 

the nine to 18 months required for taro. 

3) It requires less labor in planting and less care in 

cultivation. 

4) The tubers w i l l keep in the ground without rotting for 

several months after maturing. 

5) The sweet potato is not seasonally restricted. 

Present evidence indicates that the sweet potato was intro- 

duced to Eastern Polynesia from South America sometime after 

initial settlement, about the first to third century A.D., but 

prior to the colonization of New Zealand which occurred before 

1000 A.D. (Green 1975: 604-624). Carbonized sweet potato frag- 

ments have been recovered from archaeological sites in New Zea- 

land (Leach 1976: 145), Easter Island (Rosendahl & Yen 1971: 

379), and Hawai'i (Rosendahl & Yen 1971: 383) with dates of 1650- 

1850 A.D.; 1526+100 A.D.; and 1425-1725 A.D., respectively. In 

these areas sweet potato was of major agronomic importance. In 

contrast, sweet potato was a minor crop and dietary element in 

central East Polynesia at contact where it was used primarily for 

pig feed (Yen 1974). 

Studies at Palliser Bay provide the earliest indirect evi- 

dence for sweet potato cultivation in New Zealand (Leach 1976). 

Intensive examination of environmental conditions past and pre- 

sent, in conjunction with archaeological excavations, showed 

sweet potato to be the only possible crop for that particular 

area. These field systems date to the 12th century A.D., and it 

is assumed that the arrival and dispersal of the sweet potato 

predates the systems by a hundred years or more. For Easter 

Island it has been suggested that the sweet potato arrived with 

the initial colonizers (Yen 1974: 294) in the fifth century A.D. 

(McCoy 1976: 10). In Hawai'i evidence for swidden agriculture 

has been substantiated early in the temporal sequence (Kirch 

1975; Green, in press). However, in these Hawaiian sites the 

specific cultigen has not been identified. Until more direct 

evidence is available, the possibility of sweet potato intro- 

duction subsequent to initial settlement but prior to the estab- 

lishment of the major dryland field systems remains viable. The 

necessary agronomic techniques were already an integral part of 

the Hawaiian's cultural knowledge, but if a more adaptable plant 

was introduced it could have allowed for intensive exploitation 

of previously unused land.

In 1823 Ellis described five dryland agricul tural field 

systems on Hawai'i where sweet potato was the primary cultigen 

(Newman 1970: 114-115). These were located in leeward North 

Kohala, in Kona between Kailua and Ka'awaloa, at Wai'ohinu and 

Kapapala in Ka'u, and near Kamaili in Puna. Two of these areas 

have been investigated archaeologically: the North Kohala system 

at Lapakahi (Newman 1970; Rosendahl 1972; Tuggle & Griffin 1973) 

and the Kona system at Kealakekua (Soehren & Newman 1968; Newman 

1970: 123-137). Research in Makaha Valley, O'ahu (Green, in 

press) and Halawa Valley, Moloka'i (Kirch 1975) provide addi- 

tional data on dryland cultivation in prehistoric Hawai'i. 

The earliest indirect evidence in Hawai' i for sweet potato 

cultivation is from Makaha Valley. Field shelters dated at 1100 

to 1300 A.D. were found in stratigraphic association with evi- 

dence for swiddening (slash and burn agriculture) (Green 1970: 

101). It is assumed that these fields were used for either sweet 

potato or dry taro cultivation. This initial swidden agriculture 

was followed by more permanent inland pondfield systems with 

associated dryland farming (Green, in press). In Halawa Valley, 

geological and malacological evidence suggest a sequence of 

forest burning resulting in slope instability and erosion by 

1200 A.D. Based on this data, in conjunction with the cultural 

remains, Kirch (1975: 175-176) suggests that swiddening was prob- 

ably a major agricultural activity in the valley from the date of 

initial settlement (ca. 650 A.D.) onward. It was concluded that 

a quantitative shift in relative emphasis from swidden to pond- 

field cultivation gradually took place. The extensive field 

system at Lapakahi dates from the late 1400's to historic con- 

tact. It is in this area that the first direct evidence of sweet 

potato was found. Carbonized sweet potato tuber fragments were 

excavated from a field shelter in upland Lapakahi with associated 

dates ranging from 1425 to 1725 A.D. (Rosendahl & Yen 1971; 

Rosendahl 1972). Less directly, some coastal Lapakahi sites date 

to about 1300 A.D. (Tuggle & Griffin 1973: 58) and - may indicate 

the initial stages of sweet potato cultivation. Sweet potato 

tuber skin has also been tentatively identified from one of the 

Mauna Kea Adze Quarry rockshelters (B.P. Bishop Museum, current 

analysis). Only one radiocarbon date has been obtained for the 

rockshelter, 1492 A.D. (corrected), and the identified material 

is from a layer that is stratigraphically more recent, but 

definitely prehistoric. 

Recent research has led some archaeologists to hypothesize a 

prehistoric expansion of permanent settlement into the dry lee- 

ward areas of Hawai'i beginning as early as 1000 to 1100 A.D. 

(Cordy 1978). It is suggested that this settlement was accom- 

panied by inland agricultural expansion and substantial popula- 

tion increase, as exhibited by the extensive remains of dryland 

field systems and numerous coastal habitation sites. There were 

also certain social developments that took place which resulted 

in the formation of complex ranked societies during the late 

1400's (Hommon 1976). This is in part documented by the appear- 

ance of large house sites and heiau (temples) which represent 

social stratification. The areas affected are notably those in 

which the sweet potato would be the most productive crop. It is

tempting to suggest that the introduction of this adaptable cul- 

tigen was an important factor in these developments. However, 

evidence to support such an inference has been elusive thus far. 

Three alternative time periods are suggested for the intro- 

duction of sweet potato to Hawai' i: 

1) With initial settlement ca. 600 to 750 A.D. (Yen 1973: 

81), or possibly as early as 300 A.D. (Cordy 1978: 25). 

This would mean that the sweet potato was involved with 

but not the catalyst for the previously mentioned 

cul tur a1 developments. 

2) At a later date, about 1000 A.D. or somewhat later. The 

sweet potato could have precipitated or been directly 

requisite for certain cultural developments (Hommon 

1976: 258-269). 

3) By a historically unrecorded Spanish vessel in the 

1500's (Dixon 1932; Stokes 1932). In this case the 

sweet potato would have been introduced after major 

cultural developments had taken place. 

There is a paucity of both direct and indirect data relevant 

to the prehistoric presence or absence of sweet potato in 

Hawai ' i. Only the third possibility noted above is seriously 

questioned by present archaeological evidence. We suggest the 

following research methods may assist in defining the chronology 

of sweet potato introduction and establishment in the Hawaiian 

horticultural complex, as well as identifyinq the effects this 

cultigen 

Hawaiian 

1) 

2) 

3) 

may have had upon the development of prehistoric 

culture: 

The controlled archaeological sampling of prehistoric 

dryland agr icul tural field systems can be improved. 

This would be accomplished in part by obtaining con- 

sistent samples along two lines of variation in an indi- 

vidual field system, that is, along the longitudinal 

axis and the inland-seaward axis. Such a sampling 

procedure, combined with comprehensive absolute dating, 

should result in a complete view of the development of 

an agricultural system from its initial stages through 

European contact. 

Direct evidence is obtainable in the form of sweet 

potato macrofossils. Archaeological excavations could 

be concentrated on those sites, structures, and features 

which will most likely contain preserved sweet potato 

remains. In addition, specific field recovery tech- 

niques can be developed. 

The techniques of macrofossil identification need 

refinement, and comparative collections and keys should 

be developed.

4) Microfossil identification and analysis is not fully 

developed in Hawai'i and previous attempts at isolation 

of archaeological pollen have been relatively unsuc- 

cessful for a variety of reasons. However, pollen 

analysis may still prove productive, particularly in dry 

areas. It should be noted that for sweet potato pollen 

low percentages would be expected as the pollen is not 

windborn and the plant usually was harvested before 

flowering occurred. 

5) Climatic-edaphic conditions of a specific micro-environ- 

ment can be compared to the physical requirements of a 

particular cultigen as was done by Leach (1976). 

Ultimately all of these methods depend on the accurate and 

reliable dating of archaeological sites through basaltic glass 

hydration rind measurement and radiocarbon dating techniques. 

This very brief discussion of the possible role of the sweet 

potato in prehistoric Hawai'i is clearly quite preliminary and 

incomplete. All of the data relevant to this topic have not been 

recounted nor have all the possible interpretations been noted. 

However, recent archaeological research and dating in Hawai'i, 

combined with minimal direct evidence presently available on the 

sweet potato, is rather enticing. It is possible that a focus on 

the role of this adaptable food plant could have major explan- 

atory potential in future interpretations of the prehistoric 

development of Hawaiian culture. Until new data more clearly 

defines the prehistoric position of the sweet potato, in terms of 

the time of introduction and the effects upon Hawaiian culture, 

the concerns raised here remain a legitimate area of inquiry for 

archaeological research in Hawai'i.


Brand, D. D. 1971. The sweet potato: an exercise in method- 

ology. Pages 343-365 in C. L. Riley and Others, eds. Man 

across the sea: problems of pre-Columbian contacts. 

University of Texas Press, Austin. 

Cordy, R. H. 1978. A study of prehistoric social change: the 

development of complex societies in the Hawaiian Islands. 

Ph.D. Dissertation in Anthropology, University of Hawaii, 

Honolulu . 

Dixon, R. B. 1932. The problem of the sweet potato in 

Polynesia. American Anthropologist 34(1): 40-66. 

Green, R. C. (ed.). 1970. Makaha Valley Historical Project, 

Interim Report No. 2. Pacific Anthropological Records 

No. 10, Anthropology Dept., B. P. Bishop Museum, Honolulu. 

. 1975. Adaptation and change in Maori culture. Pages 

591-641 - in G. Kuschel. Biogeography and ecology in New 

Zealand. The Hague, Netherlands. 

. Makaha Valley Historical Project Report No. 5: Makaha 

y e f o r e 1880 A.D. Pacific Anthropological Records, Anthropology 

Dept., B. P. Bishop Museum, Honolulu. (In press). 

Handy, E. S. C., and E. G. Handy. 1972. Native planters in old 

Hawaii: their life, lore and environment. B. P. Bishop 

Museum, Bull. 233. Honolulu. 

Hommon, R. J. 1976. The formation of primitive states in pre- 

contact Hawai' i. Ph.D. Dissertation in Anthropology, 

University of Arizona, Tuscon. 

Kirch, P. V. 1975. Halawa Valley in Hawaiian prehistory: dis- 

cussion and conclusions. Pages 167-184 in P. V. Kirch and 

M. Kelly, eds. Prehistory and ecology in a windward 

Hawaiian valley: Halawa Valley, Molokai. Pacific 

Anthropological Records No. 24, Anthropology Dept., B. P. 

Bishop Museum, Honolulu. 

Leach, H. M. 1976. Horticulture in prehistoric New Zealand: 

an investigation of the function of the stone walls of 

Palliser Bay. Ph.D. Dissertation in Anthropology, Univer- 

sity of Otago at Dunedin, New Zealand. 

McCoy, P. C. 1976. Easter Island settlement patterns in the 

late prehistoric and prohistoric periods. Bulletin Five of 

the Easter Island Committee of the International Fund of 

Monuments Inc. New York.

Newman, T. S. 1970. Hawaiian fishing and farming on the island 

of Hawaii in A.D. 1778. Division of State Parks, Dept. of 

Land and Natural Resources. Honolulu. 

Rosendahl , P. H. 1972. Aboriginal agriculture and residence 

patterns in upland Lapakahi, island of Hawaii. Ph.D. 

Dissertation in Anthropology, University of Hawaii, 

Honolulu. 

Rosendahl, P. H., and D. E. Yen. 1971. Fossil sweet potato 

remains from Hawaii. Journal of the Polynesian Society 

EO(3): 379-385. 

Soehren, L., and T. S. Newman. 1968. The archaeology of 

Kealakekua Bay. B. P. Bishop Museum and University of 

Hawaii Special Report. Honolulu. 

Stokes, J. F. G. 1932. Spaniards and the sweet potato in Hawaii 

and Hawaiian-American contacts. American Anthropologist 

34(4): 594-600. 

Tuggle, H. D., and P. B. Griffin. 1973. Lapakahi, Hawaii: 

archaeological studies. Asian and Pacific Archaeology 

Series No. 5. Social Science Research Institute, University 

of Hawaii, Honolulu. 

Yen, D. E. 1971. Construction of the hypothesis for distribu- 

tion of the sweet potato. Pages 328-342 in C. L. Riley and 

Others, eds. Man across the sea. Unicrsity of Texas 

Press, Austin. 

. 1973. The origins of oceanic agriculture. Archaeology 

and physical anthropology in Oceania 8(1): 68-85. 

. 1974. The sweet potato and Oceania: an essay in 

ethnobotany. B. P. Bishop Museum, Bull. 236. Honolulu.

DEDICATION ADDRESS FOR HAWAII FIELD RESEARCH CENTER* 

1 June 1978 

Bruce M. Kilgore 

Associate Regional Director 

Resource Management and Planning 

Western Region, National Park Service 

San Francisco, California 94102 

When Bob Barbee asked me to offer some comments this morn- 

ing, I thought about my past role as a park scientist and my 

present role as a manager of professional programs. I felt I 

should not miss this opportunity to comment on some issues that 

concern me very much dealing with the National Park Service (NPS) 

science programs and scientists and their relationship to 

resource management programs and managers. 

During our first century of managing national parks, we took 

it upon ourselves to "play God" because we decided which natural 

processes were "good" and which were "bad," But how did we 

assign such moral qualities to fire in the forest or to predators 

among species of wildlife? 

In 1963 we were reminded by the Leopold Report that "playing 

God" was not what our mission was all about. And as scientists 

and managers, I find it useful from time to time to look at some 

of its major points again. You remember the catch phrases: 

"National Parks should be a vignette of primitive America," and 

"A reasonable illusion of primitive America can be recreated 

. . . using the utmost in skill, judgment, . . . and ecologic 

sensitivity." 

But there were other important ideas, too: 

1. It pointed out the folly of tinkering with natural processes 

without understanding these processes. 

2. It said that the NPS must recognize the enormous complexity 

of ecologic communities and the diversity of management pro- 

cedures required to perpetuate them. 

3. It said that management without knowledge would be a dan- 

gerous policy. 

* Portions of this paper were adapted from the Keynote Address 

at the NPS Pacific Northwest Region's Science/Resource Manage- 

ment Workshop, April 18, 1978.

When I began my present assignment in the Western Region, I 

wrote a memo to my boss, Howard Chapman, in which I raised sev- 

eral basic questions about science and scientists and attitudes 

of managers toward them. I said that perhaps the first question 

we must ask ourselves and answer honestly is: "Do we really want 

professionals and scientists in the NPS?" If we do, we must pay 

for this service, both through adequate funding and through 

strong commitment to the highest standards of professional activ- 

ity. Such activity must include: (1) high-quality, in-house 

research to provide essential facts to quide management programs; 

and (2) publication of these results in professional journals. 

Our past performance, while it has been improving recently, 

still has a long way to go, as both the Robbins and Leopold 

Reports in 1963 pointed out. In summary, these reports said four 

things: 

1. We need a permanent, independent, identifiable research unit 

within the National Park Service. 

2. Most of the research by the Park Service should be mission- 

oriented. 

3. The NPS should itself plan and administer its own mission- 

oriented research program. 

4. The results of research undertaken by the Park Service should 

be publishable and should be published. 

Such concepts form the basis for my personal philosophy of 

what our objectives and goals ought to be for a natural science 

research organization in the Park Service. But I think there are 

differences in approaches between some managers and researchers 

on these points. 

The Manager Needs the Sound, Scientific 

Support of the Scientist 

While many managers may sense they need information upon 

which to base their management of forest resources or wildlife 

resources or fisheries resources, they do not always think they 

need a real scientist. 

"Just get me the data," some say. "Give it to me in a 

report with management recommendations I can understand. 

But don't bother to write it up for those ivory-tower sci- 

entific journals. That's just the scientist doing his thing 

with his scientific peers. That's for his own personal 

benefit. It doesn't help me."

I want to say that I strongly disagree with this philosophy. And 

I want to tell you why. There is no way that a manager can be 

assured his scientist's information is solid unless he operates 

like a scientist and is recognized by his peers and the scien- 

tific community as a scientist. And for this to happen, there 

are few viable shortcuts to the process of careful design of a 

research project, careful review of that design by the most 

knowledgeable professional peers, careful gathering of data 

(often by research technicians, not the scientist himself), and 

professional analysis of the results and drawing of conclusions 

which are then subjected to a number of review processes, with 

publication as the final product. 

This is a point I have trouble with in discussions with many 

managers and some researchers. Yet, I feel strongly that if a 

field research scientist does not publish, the research mission 

of the Park Service will certainly perish in the sense that it 

will come to have zero influence in or out of the Service. 

So, what I am saying is the National Park Service must in- 

creasingly learn to support your local scientist and your local 

Cooperative National Park Resources Studies Unit (CPSU) when they 

seek to establish a reputation for solid scientific achievement. 

We must learn to support the process of presenting papers at sci- 

entific meetings and preparing the results for publication in the 

best possible scientific journals. 

Attitude. of the NPS Scientist 

On the other hand, let me warn the National Park Service 

field-area scientist and the CPSU scientist that a part of the 

reason we lack management support for science stems from atti- 

tudes of some Service scientists and research biologists. There 

are those scientists--few, I hope--who are inclined to use fancy 

equipment and procedures to do a job that less sophisticated pro- 

cedures could do equally well and with better management support 

and understanding. If you need computers and sophisticated 

equipment, use them. But do not play science games. And do not 

try snow jobs on managers. 

The National Park Service scientist, who does not fully 

understand that the primary function of Service scientists is to 

produce mission-oriented results for those problems identified by 

management as being top priority problems, has done great damage 

to the image of science in the NPS. Such an individual may feel 

he is free to study whatever strikes his fancy, because anything 

he learns w i l l benefit society and hence, the NPS. While most 

basic research has some interpretive value, there is no quicker 

way to lose support of the hard-pressed manager with a tight 

budget and an early deadline than to operate this way.

But we can make it over this hump if we have two things: (1) 

greater understanding on the part of the manager that good solid, 

science is costly and takes some optimum minimum time, and that 

following through to publication is a worthwhile investment both 

for the scientist and the manager; (2) greater commitment on the 

part of the NPS scientist to working with the manager at the out- 

set to select his highest priority projects to study, and then a 

continuing effort to gain a mutual understanding of what both 

hope to achieve by the research. This should sometimes include 

how data gathering--whatever is decided upon--will help the 

manager make a decision. In other words, we need desperately to 

better understand one another. We need better bridging of the 

communications gap that exists between manager and scientist. 

Bridging the Gap Between Scientist and Manager: 

The Role of the Resource Management Specialist 

I feel that bridging the communications gap between the 

scientist and the Superintendent or manager is a key role that 

resources management specialists can and must play. This role is 

vitally important, and they need background experience and pro- 

fessional training as nearly equivalent to that of the scientist 

as possible. As I would see it, researchers and resources 

management specialists relate to each other in the following way: 

1. The scientist develops the basic strategy--a sound rationale 

for ecological action programs of prescribed burning or goat 

and pig reduction or reintroduction of extirpated species. 

2. Then the resources management specialist--the second half of 

an essential team--deals with the tactical operations of ac- 

tually doing controlled burning in a regular way or guiding 

rangers in reducing exotic animal herds. 

An extremely important need in the Service now is to develop 

a solid, professional resources management program. We need a 

career ladder for resources management specialists, an effective 

training program for such specialists, and a separate grade eval- 

uation system to encourage them to become highly skilled spe- 

cialists and not have to transfer to line management or to 

research in order to advance professionally. We should be able 

to recruit prospective resource management specialists directly 

from universities or from other assignments where their back- 

ground experience qualifies them well. 

I would see scientists and resources management specialists 

forming essential teams in larger parks, splitting the strategy 

and tactics of resources management, while in smaller parks, the 

scientist part of the team would be provided by scientists 

stationed at CPSU's.

We will see how far these ideas get in the next few years in 

the NPS. But some effective system for bridging the research- 

management gap must be found because managers need mission- 

oriented research. But not just the short-term brush fire 

efforts. Once you have identified a major isue, you need to go 

into in-depth studies of the various aspects of the ecosystem 

that are related to that particular problem. In no way can we be 

superf icial in our approach. 

Where the Researcher Fails the Manager.. . 

And the Manager Fails the Researcher 

All too often, researchers fail in their job to assist 

managers, and managers fail in their job to support researchers. 

The researcher most often fails the manager when he: 

--carries out overly-sophisticated studies that are 

unrelated to management; 

--makes little effort to communicate the results of his 

research to the manager (including recommendations for 

action) ; 

--does not set up mutually agreed upon objectives at the 

beginning of the project and then follow through with 

reports and publications that are of value to the manager. 

The researcher owes a manager at least two thinqs: a solid study 

that leads to publication; and recommendations on how his 

research relates to management. 

The manager may fail the researcher when he: 

--undercuts the researcher's efforts to work steadily 

on primary projects, often by involving him in 

"brush fire" projects; 

--does not communicate management problems he needs 

research answers for in a timely way, or does not 

seek a researcher's input on whether a given re- 

sources problem should have priority consideration 

for limited research funding; 

--puts research at the bottom of the priority list 

for funding (maybe cutting it first in order to 

fill chuckholes in his road); 

--discourages a researcher's papers at professional 

meetings or discourages him from finishing publica- 

tions.

Hawaii Volcanoes: A Success Story 

But with all its problems and controversies, resource man- 

agement and research at Hawaii Volcanoes National Park has been a 

real success story, and the research center we are dedicating 

here today is a concrete example of the tremendous progress being 

made. Looking back briefly at where we have been in research in 

the National Park Service during the past 50 years, gives us some 

perspective. Lack of knowledge about natural resources and 

natural processes in parks has been a serious threat to the eco- 

logical health of many parks. Such was the case here in Hawai'i 

in the early 1960's with the two large park areas of Hawaii 

Volcanoes and Haleakala. 

It is well documented that more species of native Hawaiian 

plants and animals have become extinct in these islands--and more 

are threatened with extinction--than in any other biological pro- 

vince on earth. We in the National Park Service are especially 

concerned with the problems in Hawai'i since the Service is the 

largest Federal land agency in the State, and because the Service 

is charged with a Congressional mandate to conserve the scenery, 

natural objects, and wildlife on all national park lands. 

Research programs in the Service really began in the late 

1920's when an advisory committee on problems in the national 

parks recommended a research program to fill some of the gaps in 

scientific information needed to administer and interpret the 

nation's national parks. In response, a Branch of Research and 

Education was created in 1930, headed by Dr. Harold Bryant, a 

student of Joseph Grinnell. Two years later, the Wildlife 

Division of the NPS was established as the first organization 

created solely for the purpose of ecological research and manage- 

ment of biological resources. It was led by George Wright, 

another Grinnell student. 

At about this time, the Civilian Conservation Corps (CCC) 

presented the Park, Service with a unique opportunity for expand- 

ing its conservation role on the national scene. CCC Camps were 

established all over the country; many were placed in national 

and State parks. The National Park Service administered portions 

of the CCC Program, and was able to acquire a large sum of 

Federal funds for research and management activities in national 

parks. (The site of the present Hawaii Field Research Center was 

at one time a CCC Camp.) 

Unfortunately, George Wright was killed in 1936 and the CCC 

Program was abolished in the early 1940's. This led to a def- 

inite drop in NPS research efforts. It was not until nearly 25 

years later, in the early 1960ts, that the biological problems of 

the parks were again recognized as needing extensive NPS research 

commitment. In 1958, the Service obtained its first official 

budget solely for research--a meager sum of $28,000 for the 

entire NPS.

While the $28,000 would not even build a single comfort sta- 

tion for one national park, the money had some real psychological 

and fiscal pump-pr iming effects. Several Regional Offices and a 

few parks added their own funds to materially augment this ini- 

tial sum. As such, this stimulated research institutions to 

produce several dozen reports by 1962 on critical ecological 

situations in a number of parks. 

In the early 19601s, the Secretary of the Interior requested 

two surveys. The first was the "Leopold Report" I mentioned 

earlier. The second survey was one by the National Academy of 

Sciences on Research needs in national parks. It outlined the 

steps necessary to set up an effective research organization to 

handle park management problems. 

Part of the Academy of Sciences Report stated that research 

centers should be established in national parks when justified by 

the nature of the park, and that such research centers should not 

only serve the staff of a national park but should be used 

jointly by personnel from universities, other organizations, and 

other Government agencies. 

The late 1960's signaled the end of the 25-year period of 

frustration for biological research and management in the 

national parks, and the beginning of a new period of opportunity 

and hope for a better future. 

Today, 10 years later, the annual research budget for Hawaii 

Volcanoes alone totals about $250,000. Considering our sister 

agencies, in the latter 1960's the Fish and Wildlife Service 

assigned a biologist to Hawai'i to research the probable causes 

for the decline of Hawaiian birdlife, and in 1969 the two agen- 

cies jointly established the Mauna Loa Field Station in Hawaii 

Volcanoes, manned by one research biologist from each agency. 

Then in 1970, ecosystem research in Hawai'i received a tre- 

mendous "shot-in-the-arm" with a several-million-dollar grant 

from the International Biological Program (IBP). Many of the 

5-year studies undertaken wholly or in part in the Hawaiian 

national parks required adding space for offices and labora- 

tories. 

These present facilities of the Hawaii Field Research Center 

were used at that time by the Kilauea Job Corp Camp, but when the 

camp was vacated in 1973, plans were immediately formulated to 

use the buildings (1) for needs of the IBP Program; (2) for other 

scientists including personnel from the Cooperative National Park 

Resources Studies Unit at the University of Hawaii; and (3) to 

expand the operations of the Mauna Loa Field Station into a 

several-person facility for research on endangered Hawaiian eco- 

systems. Since 1973, there has been increasing momentum in the 

development of the Center as a major facility for research in 

Hawaii and in the national parks.

In 1977, the U. S. Forest Service joined the ra,nks of the 

National Park Service and the U. S. Fish and Wildlife Service as 

the third Federal agency in Hawai'i to be concerned with research 

and management of endangered Hawaiian biotas. Total funding for 

the three agencies approaches about $750,000 a year. 

The combined research efforts of the three agencies will 

make this one of the really significant interagency efforts in 

the country. The combined total of more than 20 permanent and 

seasonal employees from the three agencies should enable us to 

carry out a far more effective program of research on the decline 

and present status of endangered flora and fauna than would be 

possible by any single agency effort. We hope there w i l l be the 

synergistic interactions which will help us all, and that some 

critical mass has been achieved that will ensure that this 

research facility, with these biennial conferences, with monthly 

seminars attended by leading scientists in Hawai'i, and with on- 

going research carried out by staffs of three Federal agencies 

will become the leading research facility of its kind in the 

State. 

It is appropriate, therefore, at this stage of development 

that we officially recognize the potential for the Hawaii Field 

Research Center to become Hawai'i's leading facility for research 

and management of natural resources, and as one of the best in- 

stitutions of its kind in interagency cooperation anywhere in the 

nation. 

In doing so, I want to express my personal commendation, and 

--I think I can safely say--that of the Western Region and the 

Washington Office of the National Park Service, for the heroic 

research and resource management efforts which have been made at 

Hawaii Volcanoes and in Hawaiian parks generally. 

of: 

This is a tribute to the dedicated and long-standing efforts 

--Bob Barrel, Hawaii State Director of the National Park Service; 

--Bryan Harry, Past Superintendent, and Bob Barbee, Present 

Superintendent of Hawaii Volcanoes; 

--The researchers, past and present, ... who have contributed 

immensely to knowledge needed for active management programs-- 

Ken Baker, Garrett Smathers, Dieter Mueller-Dombois, Cliff 

Smith, their graduate students, and many others from the 

University of Hawaii; 

--and perhaps one of those who has contributed most has been 

Don Reeser, Resource Management Ecologist at Hawaii Volcanoes, 

who with his dedicated staff has contributed immensely to the 

positive values of the resources management program at Hawaii 

Volcanoes which is recognized as one of the finest active 

resource management programs in the Western Region and the 

National Park Service as a whole.

By no means, however, do I imply that our work is done! 

There is much left to do, including several items about which 

there is much controversy, and where we will need productive 

interchange between scientists and resource managers to resolve 

the issues. But I feel we all have two common objectives, as 

stated in the Master Plan which was recently approved. Namely: 

(1) Protect the park's remnant Hawaiian ecosystems-- 

including endangered species--from further depredation 

and competition by those exotic animals and plants 

introduced by modern man. 

(2) Reestablish the park's endemic species into their 

former ranges, concentrating efforts on those species 

which are in danger of extinction, and those that are 

key components of major native ecosystems. 

It is my honor on behalf of the National Park Service to 

declare the Hawaii Field Research Center as an official function 

of the National Park Service research and management effort, and 

to acknowledge that the Center is a facility for use and coopera- 

tion by other Federal agencies and educational institutions con- 

cerned with the conservation of Hawai'i's natural flora and 

fauna.

THE STATUS OF THE 

HAWAIIAN DARK-RUMPED PETREL AT HALEAKALA 

John I. Kjargaard 

Haleakala National Park 

Maui, Hawaii 96768 

The Hawaiian Dark-rumped Petrel, or 'Ua'u (Pterodroma 

phaeopygia sandwichensis), a rare and endangered oceanic seabird, 

has probablv been nestinq at Haleakala continuouslv for manv cen- 

turies. he eggs and yo;ng were considered a deiicacy by the 

Hawaiians who learned to excavate a hole in the burrow through 

which they could "harvest" the birds every year. Dogs were also 

sometimes used to locate the burrows and dig out their occupants. 

Although the species was rediscovered to science at Hale- 

akala by Richardson and Woodside in 1954, it was heard by Ted 

Rodrigues and other CCC personnel in the mid-1930's and observed 

as well as heard by Clifford McCall and other Park Rangers in the 

late 1940's. 

A history of the Hawaiian Dark-rumped Petrel compiled by 

Winston E. Banko in 1971 is the most complete study of the popu- 

lation status and distribution of the species in Hawai'i. Be- 

tween 1966 and 1971 James Larson, Warren King, Jitsumi Kunioki, 

and others initiated work to locate and monitor the Haleakala 

population (Appendix A). From 15 known burrows in 1966 the total 

has now risen to 437; of the original 15 known burrows only 47% 

were active, and of that number active one-third failed to pro- 

duce fledglings. Predation by rats and cats were considered the 

major problem. 

Monitoring records prior to 1970 are limited; however, since 

that time fairly complete reports have been made on population 

studies, banding predator control, and bird mortality. Since 

1968 the percent of activity in known burrows has ranged from 39% 

in 1969 to 95% in 1970, with recent years averaging about 69%. 

With the beginning of a predator trapping program in 1968 the 

nesting success rose to 93% and went as high as 99.9% in 1973. 

The average has been 96.5%. 

Since 1969, 22 dead adults and six juveniles have been 

recovered. Most adult deaths have been attributed to collisions 

with rocks and, in a couple of instances, with automobiles whose 

lights appear to attract and blind the birds. Juvenile mortality 

has been attributed primarily to predators although other factors 

such as parent death, adverse weather, and burrow collapse 

undoubtedly contribute.

Due primarily to lack of personnel Haleakala National Park 

has done very little work on the breeding biology of the species 

and has instead concentrated on population studies. Access to 

the primary nesting areas on White H i l l and Kilohana Pali is dif- 

ficult and hazardous due to the steep unstable terrain which has 

helped to l i m i t work on the species. The other major problem in 

conducting research on the Petrel at Haleakala has be,en the dif- 

ficulty in locating burrows, even known ones. Each located 

burrow is numbered with white spray painted numerals and one or 

more white spots which serve both to catch the eye and to 

indicate the entrance. 

Magnetic disturbances are evident in many places on the 

"Petrel slopes" and even with the use of an artificial north, 

conventional mapping techniques cannot be applied to determining 

exact locations of individual burrows. Prior to the development 

of the Haleakala Petrel Burrow Location System in 1977 burrows 

were frequently plotted by "guestimate" and although they were 

occasionally close to their actual location, more often they were 

many yards off. 

To rectify this situation a photographic based burrow plot- 

ting system was produced with the aid of a grant from the Hawaii 

Natural History Association. The tactic was to photograph all 

the "Petrel slopes" from the air from three different angles. 

Four series of pictures totaling 67 photographs were produced 

with each burrow plotted on at least two photos. The system is 

cross referenced by area and burrow number to increase versa- 

tility and although it takes about a day's practice to become 

familiar with the methods involved, the accuracy is so high that 

the location time has been cut significantly. 

Last year 290 successful breeding pairs were recorded in the 

primary nesting area and it is safe to estimate that there are at 

least another 20 pairs in more remote areas of Haleakala. It has 

been estimated that the population that nests at Haleakala, in- 

cluding all those birds that have not yet reached breeding matu- 

rity, is somewhere in the vicinity of 1600t500 individuals. 

Although the breeding population has remained more or less stable 

during the last 10 years there is little cause for optimism. 

Feral animal predation remains the largest single problem. 

Recently feral pigs moving up from Ko'olau Gap have dug up bur- 

rows in the Holua area, Dogs have never been observed but cats, 

mongoose, rats, and mice have all been trapped in the primary 

nesting areas. Perhaps one of the greatest problems is the 

almost total lack of breeding biology data which makes it diffi- 

cult to accurately assess all the factors that may be involved in 

the survival of this species.


Banko, W. E. 1971. Hawaii race of Dark-rumped Petrel (draft). 

Buxbaum, K., and J. Kunioki. Report on Dark-rumped Petrel 

observations, Haleakala National Park, Summer 1972. 

Gill, D. E., and L. N. Huber. Breeding status of Dark-rumped 

Petrel in Haleakala Crater Maui. 

Harris, M. P. 1970. The biology of an endangered species, The 

Dark-rumped Petrel (~terodioma phaeop~gia) , in the Galapagos 

Islands. Condor 72: 76-84. 

Huber, L. N. Survey of a breeding colony of Dark-rumped Petrels 

in Haleakala Crater, Maui, Hawaii. 

King, W. Haleakala Crater Maui. 

King, W. B. May 13, 1971. Report on research conducted June- 

August 1970 on the status of the Dark-rumped Petrel in 

Haleakala National Park, Hawaii. 

Kjargaard, J. I. September 1975. The Petrel Papers. 

. 1977. The Haleakala Petrel Burrow Location System. 

Kunioki, J. 1970. Dark-rumped Petrel observations, Summer 1970, 

Haleakala National Park. 

. 1971. Dar k-rumped Petrel observations, Summer 1971, 


. 1973. Dark-rumped Petrel observations, Summer 1973, 





~ a l e a k a l National a Park. 


~ a l e a k a l National a Park. 



Larson, J. W. 1967. The Dark-rumped Petrel in Haleakala Crater, 

Maui, Hawaii. 

National Park Service. 1968. Dar k-rumped Petrel observations, 

Summer 1968.

National Park Service. 1969. Dark-rumped Petrel observations, 

Summer 1969. 

Richardson, F., and D. H. Woodside. 1954.. Rediscovery of the 

nesting of the Dark-rumped Petrel in the Hawaiian Islands. 

Condor 56: 323-327.

APPENDIX A 

Summation of Haleakala Petrel data canpiled by King, Qth, Larson, Kunioki, Kjargaard, Others 

Total Known Wlrrows 15 15 15 36 226 ? 344 344 344 362 428 

Total Burrows Checked 15 0 15 36 210 113 322 250 275 315 334 

Percent Active 47 - 60 39 95 71 82 76 62 65 67 

Percent Failed 33 - 7 8 5 1 3 0.1 4 3 2 

Total Fats Trapped - -- - -- 15 4 24 ? ? 12 6 

Rattus norvegicus - -- -- - 2 4 ? - -- 

A Rattus rattus -- 11 0 ? - - 11 

- - -- - -- -- 

0 0 

-- -- 

1 

Rattus exulans 2 0 ? 1 5 

Total Mice Trapped -- -- - - 0 0 3 ? ? 17 0 

Total Cats Trapped - -- - -- 6 0 0* 0 * 0 0 0 

Total bngoose Trapped -- -- -- -- 0 0 0 1 0 1 0 

Dead Pdults Recovered - -- -- 3 1 1 1 3 3 2 6 

Dead Juveniles Recovered -- - - 0 ? 2 3 1 0 0 0 

* observed but not trapped


7. IMPACT OF EXOTIC PLANTS AND ANIMALS IN HAWAI'I* 

Charles H. Lamoureux 




,I thouah biologists have long assumed that the influx of 

exotic plants and animals reaching Hawai'i in the past 200 years 

has caused severe disturbances in natural ecosystems and to 

native organisms, few reliable data have been obtained. Some 

such data have been obtained in recent years, mostly in connec- 

tion with Hawaii IBP and with the National Park Service. This 

paper will discuss a number of different effects of introduced 

species, including their roles in destruction or displacement of 

native species, spreading fire, biological control, changes in 

moisture regimes, and disease introduction and spread. The 

literature will be summarized and gaps in our knowledge requiring 

further work will be indicated. 

It is postulated that the rate of introduction of exotic 

species, and hence the rate at which perturbations have been 

imposed on Hawaiian ecosystems, has been an important factor in 

accounting for the magnitude of change which has occurred in 

these ecosystems. 

* Abstract

POHAKULOA PROPAGATION PROJECT: 

A CONTINUING SUCCESS STORY 

Ah Fat Lee 

Wildlife Branch 

Hawaii State Division of Fish & Game 

Department of Land & Natural Resources 


INTRODUCTION 

In the latter part of the 18th century, the population of 

Nene, (Branta 

Henshaw -tea 

sandvicensis) was estimated at 25,000. 

that "the time w i l l inevitably come. 

In 1902, 

and that 

soon, when this goose will need protection £;om sportsmen (and 

introduced predators) to save it from its otherwise inevitable 

fate of extermination." The Board of Agriculture and Forestry of 

the Territory of Hawaii operated a restoration project on O'ahu 

from 1927 to 1935. Nene were distributed to various interested 

people at the close of the project. By 1950, only one of these 

birds, a gander, was still alive and accounted for. The present 

Nene Restoration Project was started in Pohakuloa, Hawai' i, in 

1949, with wildlife staff personnel in charge of the flock. At 

that time the world population was estimated to be approximately 

50 Nene. Thirty of the 50 were estimated to be free flying in 

the fields; 20 were in captive flocks in three different locations. 

The captive flocks carried the same blood lines. The 

original two pairs at Pohakuloa were obtained on a breeding loan 

from Mr. Herbert Shipman, a cattle rancher from 'Ola'a, Hawai'i. 

~ecause the Shipman flock was the only one available, it was very 

likely that they were of an inbred line. 

Captive Rearing 

The inbreeding continued in the Project with poor results. 

Our records show that a high percentage of the inbred ganders 

were sterile. Embryos were of weak vitality and a high per- 

centage died in the shell. Between 1953 and 1962 eight Nene were 

obtained from the fields. They included captured adults, young 

goslings, and a stray egg. Since they were not all taken from 

the same geographical area it is assumed that they were not too 

closely related. With this infusion of wild stock from the 

fields, the fertility and hatchability greatly improved in the 

Project (Table 1). The percent fertility of all eggs laid by 

Shipman strain geese was 54.5%; fertility of eggs laid by geese 

of different ages ranged from a low of 7.4% to a high of 75%. 

The percent fertility of all eggs laid by Wild strain geese was 

76.7%; fertility of eggs laid by geese of different ages ranged 

from a low of 41.4% to a high of 100.0% (Table 2).

Management Techniques 

The first clutch of eggs were removed from each goose at the 

completion of the clutch. This encouraged the goose to produce a 

second clutch of eggs during the breeding season. 

Several methods of incubation have been tried in the Pro- 

ject. The use of setting Muscovy ducks, Silky bantam chickens, 

and mechanical incubators were tried. Ducks and bantams proved 

satisfactory as incubators; however, since the Nene laying season 

is between November and March, we had difficulty finding enough 

available setting hens and ducks. Fall and winter are, of 

course, the normal times for domestic fowl to go into molt. We 

have not had a high degree of success with mechanical incubators 

(Table 3). 

Our best results have been to permit the goose to incubate 

her eggs. When the goslings of the first clutch begin to pip, 

the eggs are taken away from the goose and placed in an incu- 

bator-hatcher. The nest of the goose is broken up, and we have 

been successful in having Nene lay second clutches of eggs during 

the season. Three to five eggs are laid with an average clutch 

size of four. We have had a few six-egg clutches, and one suc- 

cessful hatching of six goslings out of six eggs. The incubation 

period for Nene eggs is 30 to 32 days. 

The goslings of the first clutches are hand-reared. The 

parent pair are permitted to raise the goslings of the second 

clutch. 

The diet of the Nene goslings consists primarily of greens, 

commercial poultry feeds, and vitamin/mineral supplements. The 

favorite choice of greens is Sow thistle (Sonchus oleraceus), and 

we go out of our way to collect it in the lower elevations for 

the flock. In Pohakuloa a rye grain-barley cross, Zetra petra, 

is planted during the fall and winter months when f r o m l l s the 

local vegetation. This has proven to be very acceptable to the 

Nene; however, they - w. i l l also qraze on any qreen qrass, including 

kikuyu (~ennisetum clandestinum) , chickweed (stellaria media) ; 

and varieties of clover. 

We keep accurate pedigree records of all Nene bred, raised, 

or kept at Pohakuloa. Pairs of Nene are mated as long as they 

are productive and compatible. We have pairs of producing Nene 

in the breeding flock that are 14 years of age. Most Nene breed 

in their second year, but we had one'goose that produced young 

her first year. Mated pairs have been successfully broken up, 

when the need arises, and mated to other mates. 

The strain of "hairy down" goslings has been successfully 

bred out of the flock during the past three years. 

At six weeks of age, the goslings are cloacally sexed and 

banded with numbered aluminum U. S. Fish and Wildlife Service 

bands. Goslings are capable of flight at 10 weeks, so their pri- 

maries are clipped while they are in the pens. Prior to release,

each Nene is banded wi.th colored plastic leg bands. The order of 

the color bands is a "color band combination" for a specific 

individual, and no two Nene are banded alike. As many as three 

color bands have been placed on each leg of an individual Nene. 

The aluminum U. S. Fish and Wildlife Service bands are removed 

before release, and our records contain the given serial number 

as well as the color band combination. Biological studies in the 

field are facilitated by this method of record keeping. 

Goslings are taken out to the gentle-release pen in the 

fields between two and eight months of age. In years when we 

produce a large number of goslings, the young are sent out in 

groups of about the same age. In years when production is 

smaller, the goslings are held until the last broods are ready 

for flight, for a single yearly release. 

The development of techniques for captive breeding and 

rearing Nene for successful release has been a challenge to all 

of us associated with the Project. Between 1949 and 1978, we 

have been successful in raising 1699 Nene at Pohakuloa (Table 4). 

1225 have been released on the Island of Hawai'i; 268 have been 

released on the Island of Maui in Haleakala Crater (198 Nene were 

shipped to Hawai'i from the Wildlife Trust in England, and those 

birds, as well as seven Nene from a private breeder in Connec- 

ticut, have been released on Maui). 

Now, as the Endanqered Species Propaqation - - Project, we continue 

our work with-~ene, Koloa (Anas - wyvillianaj , Laysan teal 

(+ laysanensis) , and the 'Alala (Corvus tropicus). 

LITERATURE 

CITED 

Anonymous. 1972. A report of the Nene restoration program. 

State of Hawaii, Div. of Fish and Game, Honolulu. 

. 1976. Survival and recruitment of wild and released 

pen-reared Nene on Hawaii. Project W-18-R-1, Job No. R-1-0, 

July 1, 1975 to June 30, 1976. State of Hawaii, Div. of 

Fish and Game, Honolulu. 

Henshaw, H. W. 1902. Birds of the Hawaiian Islands. Thos. G. 

Thr um, - Honolulu. 

Schwartz, C. W., and E. R. Schwartz. 1949. The game birds in 

Hawaii. Board of Agriculture and Forestry, Terr. of Hawaii, 

Honolulu. 

Smith, J. D. 1952. The Hawaiian goose (Nene) restoration 

program. Wildlife Mgt. 16: 1. 

Walker, R. L. 1977. How to save an endangered species...the 

Hawaiian goose (Nene). Address given before the American 

Assoc. of Zoo Veterinarians, Honolulu, Hawaii, Oct. 31, 

1977. (Unpublished).

!IWLE 1. Summary of production at 6hakuloa £ran 1953-1954 through the 1971-1972 

breeding season. 

1953- 1954- 1955- 1956- 1957- 1958- 1959- 1960- 1961- 1962- 

Season 1954 1955 1956 1957 1958 1959 1960 1961 1962 1963 

No. Breediq Pairs 6 4 5 6 6 15 14 15 16 16 

No. Producing 

First Clutch 

Second Clutch 

Third Clutch 

Fourth Clutch 

4 

4 

0 

0 

0 

4 

4 

1 

0 

0 

5 

5 

1 

1 

0 

6 

6 

2 

2 

1 

6 

6 

5 

4 

2 

15 

15 

12 

6 

3 

14 

14 

13 

8 

0 

15 

15 

14 

10 

0 

16 

16 

14 

9 

2 

16 

16 

16 

13 

1 

Total Eggs 

Eggs per Clutch 

14 

3.5 

16 

3.2 

27 

3.8 

46 

4.1 

76 

4.4 

118 

3.3 

149 

4.3 

174 

4.4 

181 

4.4 

204 

4.4 

~ggs per Goose 3.5 4.0 5.4 7.7 12.6 7.9 10.6 11.6 11.3 12.7 

No. Fertile, K* 6 4 18 22 15 28 56 97 87 130 

No. Fertile, E** 0 1 0 0 1 4 5 18 18 17 

No. Infertile 8 10 8 16 47 76 86 49 68 50 

NO. Damaged 0 1 1 8 13 10 2 10 8 7 

Percent Eggs with 

Fertile, LG 42.8 35.0 66.7 47.8 19.7 23.7 37.5 55.7 48.1 63.7 

Percent Qgs Fertile 42.8 31.2 61.7 47.8 21.1 27.1 40.9 66.2 58.1 72.7 

No. Hatched 4 4 8 14 8 11 27 40 52 65 

Percent Hatchability2 66.8 100.0 44.5 63.7 53.3 39.2 48.2 41.2 59.8 50.0 

Mortality3 0 0 0 2 4 1 8 7 6 5 

Percent Mortality 0 0 0 14.3 50.0 9.1 29.6 17.5 11.5 7.8 

Goslings per Gooseb 1.0 1.0 1.6 2.3 0.7 0.7 1.9 2.7 3.2 4.1

TABLE 1--Continued. 

1963- 1964- 1965- 1966- 1967- 1968- 1969- 1970- 1971- 

Season 1964 1965 1966 1967 1968 1969 1970 1971 1972 

No. Breediq Pairs 17 17 20 24 30 40 30 29 26 

No. Produciq 16 16 20 23 27 38 30 29 21 

First Clutch 16 16 20 23 27 38 30 29 21 

Secord Clutch 15 15 16 19 18 26 13 25 18 

Third Clutch 13 8 11 7 2 0 1 5 2 

Fourth Clutch 1 0 0 0 0 0 0 0 0 

Total Qgs 202 176 197 208 196 259 180 260 185 

Eggs per Clutch 4.5 4.5 4.2 4.2 4.2 4.0 4.1 4.5 4.5 

qgs per Goose 12.6 11.0 9.9 9.1 7.3 6.8 6.0 8.6 8.9 

No. Fertile, LG* 106 121 138 143 151 200 145 191 139 

No. Fertile, DG** 28 22 18 10 6 7 12 28 4 

No. Infertile 56 26 32 43 26 36 19 29 29 

NO. Damaged1 11 7 9 12 13 16 10 12 13 

Percent Qgs with 

Fertile, I& 52.7 68.7 70.1 68.7 77.0 77.2 80.6 73.5 75.0 

Percent Qqs Fertile 66.3 81.3 79.2 73.7 80.2 80.0 88.3 84.2 77.0 

No. Hatched 47 50 81 93 121 176 122 145 111 

Percent Hatchability2 44.3 41.3 58.7 65.0 80.1 88.0 78.7 66.2 77.6 

~ortality' 5 6 11 1 5 13 8 13 7 

Percent brtality 10.6 12.0 13.6 1.1 4.1 7.4 6.5 8.9 5.0 

Goslings per ~oose' 2.9 3.1 4.1 4.0 4.5 4.6 3.8 5.0 4.0 

1 These are eggs broken in the nest; soft-shelled; abnormally mall, etc. 

Fertility undetermined. 

2 

Percent of eggs with fertile, live germs that were successfully hatched. 

3 Only post-hatch mo&ality (occurring within the first two weeks) 

included here. 

is 

* This represents production per goose of goslings successfully hatched. 

**LG means live germ at 10 days of incubation. 

**DG means dead germ at 10 days of incubation.

PART A: Shipnan Strain Geese (14 Geese) 

TABLE 2. Fertility of eggs in relation to the age of geese. 

No. of Geese 8 12 10 10 9 9 8 6 6 6 5 

No. of Eggs 68 111 119 119 118 110 94 70 60 59 46 

No. Fertile 5 30 65 62 79 66 56 56 43 43 29 

Percent Fertile 7.4 27.0 54.7 52.1 67.0 60.9 59.6 71.4 71.7 72.8 63.1 

No. of Geese 27 15 9 7 4 3 1 

No. of Eggs 212 143 83 66 29 17 4 

No. Fertile 164 121 66 43 12 15 4 

PercentFertile 72.4 84.7 79.5 65.2 41.4 88.3 100.0

Table 3. Hatching success as related to method of incubation. 

Method of Incubation 

IWscovy Silky Bantam 

Ducks1 Hens Mechanical ' Neneb 

Number with Fertile, Live Germs 43 156 294 335 

Number with Fertile, mad Cgrms 1 16 43 10 

Number Infer tile 11 22 55 49 

Number Damaged 6 7 1 22 

Number Incubated Full-Term 43 156 294 335 

Number Hatched 17 78 88 294 

Percent Hatchability + 39.6% 50.0% 29.9% 88.0% 

* All eggs are frcm first and second clutches. 

' ~ucks were used from 1957-58 through 1959-60. 

Hens were used from 1961-62 through 1964-65. 

Mechanical incubation was used from 1960-61 through 1966-67. 

' Data for incubation by ~ene is from 1958-59 through 1968-69. 

Percent hatchability in terms of nmber hatched divided by the number incubated 

full term. 

Note: Ducks were used to supplement Nene for incubating eggs during the early 

stages of the propagation program. The hatchability was comparatively poor 

because the geese were predominantly Shipnan strain geese. These blood lines 

had been found to have low fertility and hatchability.

WLE 4. Gn5 restoration project record (July i, 1949 throqh June 30, 1977). 

~ene kared 

at Ehakuloa 

Year Mnnber 

1949-50 2 

1950-51 3 

1951-52 2 

1952-53 1 

1953-54 4 

1954-55 4 

1955-56 8 

1956-57 12 

1957-58 3 

1958-59 15 

1959-60 17 

1960-61 32 

1961-62 45 

1962-63 54 

1963-64 38 

1964-65 41 

1965-66 69 

1966-67 84 

1967-68 123 

1968-69 156 

1969-70 114 

1970-71 131 

1971-72 104 

1972-73 109 

1973-74 134 

1974-75 141 

1975-76 160 

1976-77 47 

Totals 1653 

Year 

&leased 

1960 

1961 

1962 

1963 

1964 

1965 

1966 

1967 

1968 

1969 

1970 

1971 

1972 

1973 

1974 

1975 

1976 

1977 

En6 @leased Island of Hawai'i* 

K$uka 

Keauhou Keauhou 2 Kahuku 'Ainahou 

Sanctuary Sanctuary Sanctuary Sanctuary 

20 - 

- 

11 20 

- 35 - 

- 42 - 

- - - 

30 19 - 

- - 

- - 75 

- - 85 

- 33 122 

106 - - 

94 - - 

2 35 - 

13 - - 

61 

- - - 123 

- - - 135 

- 164 - - 

- - - - 

276 348 282 319 

All of the NGn5 released on Hawai'i wre reared at Ehakuloa. 

Total 

20 

31 

35 

42 

- 

49 

- 

75 

85 

155 

106 

94 

37 

74 

123 

135 

164 

- 

1225 

NEnE Released Island of 

FTOm 

Ran Wha- Comec- 

Englani kuloa ticut 

- - - 

- - - 

30 5 

19 

20 

5 

8 

5 

- 

24 8 2 

- 25 - 

- - - 

- - 

20 

50 22 - 

55 - - 

- - - 

- 44 - 

- 50 - 

- - - 

- - - 

- 34 - 

- 48 - 

198 269 7 

Maui 

Total 

- 

35 

29 

28 

34 

25 

- 

20 

72 

55 

- 

44 

50 

- 

34 

48 

Total 

NEnE 

&leased 

20 

31 

70 

71 

28 

83 

25 

75 

105 

227 

161 

94 

81 

124 

123 

135 

198 

48 

474 1699

STUDIES IN THE LIFE HISTORY OF THE 'ALALA IN CAPTIVITY* 

Barbara Lee 

Division of Fish and Game 

State Department of Land & Natural Resources 

Honolulu, Hawaii 

This illustrated report describes observations on behavior 

of hand-reared 'Alala (Corvus tropicus) maintained as captive 

breeding stock at the State of Hawaii's Endangered Species 

Project, Pohakuloa, Hawai' i. The development of husbandry tech- 

niques and the problems and experiences related to establishing 

and maintaining a breeding population of 'Alala in captivity will 

be briefly discussed as well as the objective study of social, 

instinctive, and reproductive activities of the only six 'Alala 

in captivity. 

* Abstract

HUMAN PERCEPTION OF THE HAWAIIAN ENDANGERED SPECIES: 

A PRELIMINARY REPORT ON A THREE-YEAR RANDOM SURVEY 

Mark David Merlin 




INTRODUCTION 

The rare and endangered species of Hawai'i represent one of 

the major problems facing those interested in preserving the 

exceptional natural heritage of the Hawaiian archipelago. It is 

well known that the Hawaiian Islands have a diverse and unique 

native biota. Most regretfully, a relatively large number of 

endemic species have already become extinct within historic times 

and several more are on the verge of disappearing forever. 

There are a number of reasons for the extraordinary demise 

of so many of the native Hawaiian species. Exploitative land 

use, the impact of large feral herbivore populations, the intro- 

duction of aggressive weed species, fires, predatory and patho- 

genic organisms, and general habitat destruction have all taken 

their toll on the unique and vulnerable endemic species of 

Hawai'i. The impact of these problems has become increasingly 

acute in recent years. 

OBJECTIVES 

The basic assumption underlying the research discussed below 

is that a general consensus regarding the importance of the rare 

and endangered species is lacking. In order to determine citizen 

attitudes relevant to this issue, an ongoing research effort was 

initiated in 1976 to survey human perception of the problem. The 

goal of this project has been two-fold in nature: on one hand, 

there has been an attempt to quantify perception of the real or 

potential economic, scientific, aesthetic, ecological, and bio- 

logical value of the rare and endangered species; on the other 

hand, it has also been the aim of this research to stimulate more 

study into the problem so that an objective measurement of the 

public's attitudes can become known. In other words, this is 

basically a pioneering effort to monitor popular feelings about 

an issue of growing concern and urgency. Furthermore, it has 

been hoped that the survey process will in some way elevate 

public awareness of the problems so that educated decisions af- 

fecting the future of the native Hawaiian plants, animals, and 

habitats can be made.

METHODOLOGY 

In order to measure the public's attitudes regarding the 

real or potential value of the species in question, a series of 

random surveys was taken of citizens from various parts of the 

Island of O'ahu. Over a three-year period, some 15 undergraduate 

students administered random surveys throughout the island as 

partial fulfillment of a course requirement in the General 

Science Department of the University of Hawaii at Manoa. 

The sample survey size for each student was approximately 

200. In 1976, eight students (Arthur Horibe, Rose Souza, Cynthia 

Hara, Jean Higa, Ronna Hazel, Mary Sniffen, Lori Fowler, & Diane 

Rose) completed their field work. In 1977, three students (Alva 

Young, Bobbie Daniels, & Ann Kagawa) completed their field work. 

And in 1978, four more students (Harold Yap, Debra Yuen, Joni 

Tanonaka, & Terry Tamura) completed their field work. The com- 

bined effort so far has compiled a sample size of approximately 

3000. 

The majority of the individual surveys were taken at various 

shopping centers located on O'ahu. Randomness and general objec- 

tivity in survey procedure were stressed. However, although 

these aspects were crucial to the usefulness of the data and the 

validity of the interpretations, common problems facing the 

social scientist may not have been under satisfactory control. 

For example, many people refused to answer the survey; and as in 

many surveys of human perception, there is always the question as 

to whether or not the persons surveyed did respond to the ques- 

tions according to their true attitudes rather than socially 

acceptable ones. Moreover, one can argue that the structure of 

the questions themselves may have influenced the respondents to 

answer in a socially approved way. 

Indeed it is hard to study the attitudes in question without 

biasing the responses in favor of preservation. This is a par- 

ticularly difficult problem when there is no "price" involved in 

giving the "right" answer. In fact, this difficulty (i.e., per- 

sonal financial commitment) is generally problematic in studies 

of quality of life (Dr. Earl Babbie, pers. comm. 1976). With 

this basic problem in mind, we revised the original survey admin- 

istered in 1976 so that those taken in 1977 and 1978 had (what we 

considered to be) less ambiguity and better research design. 

Copies of the 1976 and revised 1977-1978 surveys are pre- 

sented in the Appendix. Note that data regarding age, length of 

residency in Hawai'i, educational background, ethnicity, and sex 

was also solicited from those answering the survey. Generally 

these showed relatively close correlations to these same charac- 

teristics manifested in the overall state population. Cross 

tabulation analysis of these characteristics (of the people sur- 

veyed) and the attitudes reflected in their answers to the first 

nine questions regarding their perceptions of the importance of 

the endangered species may reveal some interesting aspects of

public opinion concerning the issues at hand. This data is still 

in the process of being analysed. 

RESULTS 

Combined tabulations for the individual years 1976, 1977, 

and 1978 are presented in Tables 1 and 2. It should be noted 

that questions 4, 5, and 6 for 1976 have been shifted to ques- 

tions 5, 6, and 7, respectively, in the 1977 and 1978 surveys; 

and question "7" for 1976 has been shifted to "4" in the 1977 and 

1978 surveys. 

A cursory examination of the data reveals an apparently 

strong concern for the protection of Hawai'i's endangered plants 

and animals. The majority of those surveyed over the three-year 

period feel that these species have an important research poten- 

tial, have significant roles in the Hawaiian ecosystems, serve 

useful purposes, are important parts of Hawai'i's heritage, and 

have significant aesthetic value. However, non-native plants are 

also considered to be of equal value and the percentages of unde- 

cided responses to some questions tends to reduce the overall 

positive response of the public to the questions regarding the 

protection of the endangered species of Hawai' i. 

The general difficulties of social survey research notwith- 

standing, it is hoped that this preliminary effort will stimulate 

other students, scientists, and concerned citizens to improve on 

the research design and possibly produce a more complete descrip- 

tion and explanation of the public's attitudes pertinent to this 

problem.

(Question) 

Variables Strongly Agree 

TABLJI 1. Survey of perception of edangered species (1976-1978). 

- 1976 

Agree St rongly Disagree Disagree 

31( 2%) 

593(39%) 

99( 7%) 

517(34%) 

38( 3%) 

188(13%) 

34( 2%) 

612(41%) 

19( 1%) 

4i 1%) 

165(34%) 

25( 5%) 

33( 7%) 

204(42%) 

22( 5%) 

50(10%) 

157(32%) 

8( 2%) 

5( 1%) 

268(42%) 

17( 3%) 

23( 4%) 

226(35%) 

13( 2%) 

70(11%) 

189(30%) 

8( 1%) 

Uncertain 

52( 3%) 

207 (14%) 

271 (18%) 

91( 6%) 

58( 4%) 

266(15%) 

94( 6%) 

237(16%) 

37( 2%) 

4( 1%) 

113(23%) 

79(16%) 

20( 4%) 

48(10%) 

17( 3%) 

74(15%) 

143(29%) 

14( 3%) 

6( 1%) 

85(13%) 

56( 9%) 

36( 6%) 

40( 6%) 

36( 6%) 

32(13%) 

137(21%) 

29( 5%)

(Question) 

Variables No Bsponse 

TABIE 2. Survey of perception of endangered species (1976-1978). 

NO Response 

113 

7.5% 

NO Response 

NO Response 

Under 15 

Less %an Yr 

Inter. Sch 

Japanese 

High Sch 

564 

37.5% 

Caucasian 

452 

30.0% 

- 21-30 

379 

25.2% 

- 5-10 

160 

10.6% 

College 

557 

37.0% 


109 

7.2% 

- 31-40 

202 

13.4% 

- 11-20 

321 

21.4% 

- Other 

87 

5.8% 

Filipino 

65 

4.3% 

41 or Cwer 

289 

19.2% 

- More 

443 

29.5% 

- Other 

342 

22.8%

TABLE 2-Continued. 

(Question) 

Variables ~o ~espnse Under 15 15-20 21-25 26-30 31-40 41-50 Over50 

----- 

9 24 135 78 64 80 59 38 

2% 5% 28% 16% 13% 16% 12% 8% 

- 1 

- 2-4 

5-1 0 

- 11-20 - Over 20 

94 34 42 151 166 

19% 7 % 9% 31% 34% 

Intermediate Sch High Sch College - Other 

24 233 222 5 

5% 48% 46% 1 % 

Japanese Caucasian Chinese Hawaiian Filipino Korean Samoan Other No Response 

97 

20% 

79 

16% 

138 

28% 

20 

4 % 

37 

8% 

27 

6% 

11 

2% 

57 

12% 

21 

4 % 

- Male Female ~o Wsponse 

197 184 7 

40% 38% 2%

(Question) 

Variables 

10 

11 

12 

13 

14 

tie Response 

11 

2% 

NO Response 

19 

3% 

NO Response 

1 

0% 

No Response 

16 

3% 

NO Response 

8 

1% 

Under 15 

16 

3% 

Less Than 1 Yr 

52 

8% 

Inter. Sch 

244 

38% 

Japanese 

186 

29% 

- Male 


309 

48% 

- 1978 

- 15-2 0 

318 

50% 

- 1-4 

61 

10% 

High Sch 

233 

35% 

Caucasian 

163 

25% 

Female 

323 

50% 

- 21-30 

123 

19% 

- 5-1 0 

67 

10% 

College 

160 

25% 


31 

5% 

- 31-40 

74 

2% 

- 11-20 

281 

44% 

- Other 

3 

0% 

Filipino 

44 

7% 

Over 40 

98 

15% 

Over 20 

160 

25% 

- Other 

200 

31%

APPENDIX 1. Questionnaires used during the perception of endangered species 

study (1976 & 1977-78). 

This is a survey to find out how the people of Hawaii feel about the rare 

- - - 

native plants and animals of Hawaii. All questions are optional; but please 

answer as best you can. Mahalo. 

Can you guess how many native Hawaiian plants and animals are in danger of 

disappearing completely from::the Hawaiian environments? (How many?) 

(Please check the appropriate box for each of the following questions.) 

SA = strongly agree 

A = agree 

D = disagree 

SD = strongly disagree 

U = undecided (no opinion) 

1. The endangered species, like all species, have a right to live. 

SAD A n D n sDu '-'I 

2. Economic progress is more important than the native plants and animals. 

SAU A D SDU u n 

3. The species provide important research potential. 

SAD A l l D O SDU U O 

~o 

4. Native plants and animals serve no useful purpose. 

SACI A D DU s ~ a 

5. The native plants and animals are an important part of Hawaii's 

natural heritage. 

SAfl A n D n snn u n 

6. Non-native plants and animals are equally desirable. 

SA17 A D n u SDI7 u n 

7. The ecological functions of the native plants and animals should be 

protected. 

SAO A n D U SDU u u 

8. The protection the native plants and animals would limit recreational 

9. The native plants and animals add to the environmental beauty of Hawaii. 

S A D A a D D S D D u n 

What should be done about the endanger plant and animal situation? 

Suggestions? Use the back of this paper. 

10. Your age: under 15 15-20 21-25 26-30 31-40 41-50 over 50 

11. How long have you lived in Hawaii? 

12. Educational Background: Intermediate School High School - 

College 

13. What ethnic background best describes you?

This is a survey to find out how the people of Hawaii feel about the 

endangered native plants and animals f Hawaii. All questions are optional; 

but please answer as best you can. Mahalo. 

(Please check the appropriate box for each of the following questions.) 

SA = strongly agree 

A = agree 

D = disagree 

SD = strongly disagree 

U = undecided (no opinion) 

The endangered plants and animals, like all 

species, should be protected by man. 0 O n 1 0 

Economic progress is more important than the 

preservation of the endangered native plants u R D U D 

and animals. 

The endangered plants and animals may have. an 

important research potential. n n u n 1 

The endangered plants and animals may have an 

important role in Hawaii's ECO~O~Y. fl O U I I R 

The endangered plants and animals serve no 

useful purpose. 0 I l U U O 

The endangered plants and animals are an 

important part of Hawaii's natural heritage. 

(3 n u n n 

Non-native plants and animals are just as u ~ u I n 

valuable as the endangered ones. 

The protection of the endangered plants and animals (1 0 [1 

would limit recreational use of their habitats. 

The endangered plants and animals of Hawaii add 

to her environmental beauty. a n u u 1 

Your age: under 15 15-20 21-25 26-30 31-40 41-50 over 50 

How long have you lived in Hawaii? 

Educational Background: Intermediate School High School 

(Highest level completed) College 

Of which ethnic background do you consider yourself? 

Japanese Filipino 

Chinese Korean 

Caucasian Samoan 

Hawaiian Other 

Your sex: Male Female

ACID RAIN IN HAWAI 'I* 

John M. Miller, and Alan M. Yoshinaga 

Mauna Loa Observatory 

National'Oceanic and Atmospheric Administration 

Hilo, Hawaii 

Precipitation samples have been collected for chemical anal- 

ysis on the island of Hawai'i since June 1975. Though the number 

of collection sites varied from 5 to 10, the permanent sites 

included Kapoho (sea level), Hilo (60 m), Kulani Mauka (2500 m), 

and Mauna Loa Observatory (3400 m). Samples were collected 

either on an event basis or every three days depending on the 

accessibility of the site. The pH and conductivity tests were 

performed immediately after collection of all samples. Anion 

analysis was made on selected samples using an ion chromatograph. 

The results show that pH values in precipitation ranged from 3.7 

to 5.5 with an all-island average of 4.5. The acidity of the 

precipitation increased with elevation from sea level to 3400 m. 

Analysis in terms of air trajectories and surface winds at the 

Mauna Loa site indicates the more acidic rains come from the 

northern quadrant. The effects of the volcano on the acidity 

will also be discussd. 

* Abstract

THE SEPTEMBER 1977 ERUPTION OF KILAUEA VOLCANO, HAWAI'I' 

Richard B. Moore, Daniel Dzurisin, 

Gordon P. Eaton, Robert Y. Koyanagi, Peter W. Lipman, 

John P. Lockwood, Gary S. Puniwai, and Rosalind Tuthill Helz2 

The latest eruption of Kilauea Volcano began on 13 September 

1977, after a 21-month period of quiescence. Harmonic tremor in 

the central east rift zone and rapid deflation of the summit 

occurred for 22 hours prior to the outbreak of surface activity. 

The first spatter cones formed along a discontinuous, en 

echelon, 7-km-long fissure system trending N 70°E between two 

prehistoric vents, Kalalua and Pu'u Kauka. During the first 

week, eruptive activity was concentrated at two spatter cones, 

one near the center and one at the west end of the new fissure. 

The most voluminous phase of the eruption began late on 

September 25. An irregular spatter rampart formed along a 500-m 

segment near the center of the new fissure, but within 24 hours 

activity became concentrated at the east end of this segment. 

One flow from the new, breached, 40-m-high cone at this site 

moved rapidly (up to 300 m/hr) southeast, eventually reaching a 

point 700 m from the nearest house in the evacuated village of 

Kalapana. The total volume of material produced during this 

19-day eruption is estimated to be 25-50 x 10 m. 

Samples from active flows and vents indicate that a differ- 

entiated tholeiitic basalt was erupted. Plagioclase is the only 

significant phenocryst, and augite and minor olivine accompany it 

as microphenocrysts. This mineralogy, although uncommon in 

Kilauea lavas, is similar to that of the 1955 basalt. Some vari- 

ation in bulk composition occurred throughout the eruption, but 

the last basalt produced also appears to be differentiated, 

suggesting that the magma involved in summit deflation has not 

erupted. 

Abstract 

National Center, U. S. Geological Survey, Reston, Virginia.


1. BRIEF INTRODUCTORY SURVEY 

Dieter Mueller-Dombois 




IBP, the International Biological Program was the first 

internationally coordinated, mu1 ti-disciplinary ecological 

research program which focussed on the ECOSYSTEM as the study 

object. Officially, the research objective was to study the biological 

basis of or anic roduction in the world's major ecosystems. 

About -bg,.\. na Ions par lclpated in the Program. Rather 

large research teams were organized in Russia, ~apan, East and 

West Germany, England, France, Canada, Australia, and the U. S. 

The operational phase lasted a decade, from 1965 through 1975. 

In the U. S., two major component programs were developed, a 

Human Ada tabilit component, concerned-with the study of human 

-a-g in extreme environments, and an Environmental 

component, concerned with the natural science aspects, partic- 

ularly with the structure and function of major ecosystems. 

Teams of each 80-150 natural scientists were organized to study 

five mainland biomes in the U. S.: Tundra, Grassland, Desert, 

Eastern Deciduous Forest, and Western Coniferous Forest. The 

research emphasis of the Biome studies was on ecosystem metab- 

olism, i.e., Photosynthesis, Respiration, Decomposition, Consumer 

Relations, and Mineral Cycling. In addition, three smaller 

research projects, with teams of 20-50 scientists were organized 

on the theme of Ecosystem Structure and Evolutionary Biology. 

Two of these worked on comparisons of similar ecosystems in geo- 

graphically disjunct places: one was the Mediterranean Scrub 

ecosystem comparison between California and Chile; the other, the 

Desert Scrub and mesquite ecosystem comparison between Arizona 

and Venezuela. Ours was the third, focussing on Island Ecosystem 

Stability and Evolution. 

By legislative mandate, the National Science Foundation 

received an annual allotment of about $6 to 10 million. Of this, 

the mainland biome studies received $1 to 2 million per year. 

The three Ecosystem Structure and Evolution Study programs 

received from $200,000 to $500,000 per year. 

The funds were highly competitive. We got the first slice 

in 1970 after three times rewriting our proposal, and then 

managed to maintain approximately a $300,000 per year budget for 

our suggested five-year program from 1970-75.

General Conclusions 

1. Zonal (or community) boundaries are evident from our data on 

species distributions, but boundary criteria need to be 

defined. 

2. Different organism-groups show different gradient sensitiv- 

ities which are peculiar to the organism group. ... 

3. Spatially associated species groups occur in nearly all 

organism groups. 

4. Native species show a higher degree of spatial integration 

than introduced species among plants, birds, and Droso- 

philids. However, the origin (whether a species is native or 

non-native) is not a general predictor for species distri- 

bution patterns. These depend on the species ecological 

properties and the degree of naturalization (i.e., habitat 

saturation) among the exotics. 

5. There is not only one generalized pattern of species distri- 

bution as Whittaker has claimed for temperate mountain gra- 

dients. Instead we can recognize at least three dominant 

patterns: 

1) Individually distributed species 

2) Spatially associated, but overlapping species groups 

3) Various forms of widely distributed species. The domi- 

nance of the latter groups are probably a characteristic 

related to biota distribution on young island mountains.

FIGURE 2. Distribution of the three rodent species 

sampled along the Mauna Loa transect 

(Oct. 71 through Sept. 73). Abundance 

scale in equal units of 5 animals trapped 

per year: Maximum is 9 units = 41-45 

animals (M. musculus at site 4), 8 units = 

36-40 animals, 7 units = 31-35, 6 = 26-30, 

etc. , horizontal line = 1 animal (e.g. , 

R. rattus at site 12). K = closed kipuka 

forest in savannah zone IV. Origin: TeC 

= Temperate-continental Asia, TeL = Temperate- 

littoral Asia, TrP = Tropical-Pacific Asia.

- 

)RIGIN 

- 

TeC 

- 

TeL 

- 

TrP 

- 

11.000 

9.000 

7.000 

5.m 

3.m 

11-11 

'RANSECT ZONES I 1 1 1 1 m ll PI? 

9 

IBP SITES 14 13 12 11 IO/B 7 6 5 4 K 3 2 1 

I I I I I I I 

ALTITUDE DISTANCE I kml

FIGURE 3. Distribution diagram of populations of selected 

fungal species from soils along the Mauna Loa 

transect. Curve heights are based on relative 

population levels rated from very low to high. 

Dashed lines imply presence at very low levels. 

Species group numbers along the right-hand column 

refer to spatially associated groups in the two- 

way table (50/10 rule). Ungrouped species are 

at the bottom of the diagram. FI = Fungi Imperfecti, 

P = Phycomycete, S = Sterile Isolate.

A 

SOIL FUNGUS SET I B C 

ZONES SET2 1 m 

Fusorlum Iolerltlum 

Mortlarsllo hyqrophllo 

UYCW froqllls 

I 

Cyllndrocorpon mqnyrlonum 

I 

Mortlarello ramanniono 

1 A 

I 

Trlchodcrmo vlrlds I 

ALTITUDE DISTANCE (km)


8. ISLAND ECOSYSTEMS: WHAT IS UNIQUE ABOUT THEIR ECOLOGY? 

Dieter Mueller-Dombois 




We may now ask the question whether we have found anything 

unique or different in the ecology of island ecosystems from our 

studies. This is not an easy question. 

It is clear that the biological evolution of our island ecosystems 

has been rather unique. Four factors stand out which 

contribute to the unique biological evolution of ecosystems here. 

These are the extreme isolation of the island group, the small 

- size of the island habitats, the recenc of the oceanic islands 

as a group, and their perturbation dy in connection with 

volcanism. 

The extreme isolation had a significant "screening effect" 

on what organism groups could get here and establish themselves 

successfully. This screening effect excluded any plants with 

large seeds or small seeds of short longevity. It also excluded 

among animals, all terrestrial mammals (except the hoary bat), 

large reptiles, and primates, except man. 

The small size of the island habitats is the result of small 

isiand land masses jutting h-i-gh ou~f-o~f the-ocean. Thus, we have 

distinct altitudinal segregations of habitats with their own 

temperature regimes. Furthermore, these small land masses are 

segregated into windward (pluvio tropical) and leeward (xerotropical) 

habitats with their own rainfall regimes. The habitats 

are further fractioned by great variations in substrate, ranging 

from recent volcanic flows to old, sceletized, and nutrientdepauperated 

latosols. This island habitat mosaic brings about 

another factor of important ecological consequences, and that is 

the very limited recurrence of similar habitats across the island 

chain. These narrow habitat dimensions strongly l i m i t the 

population sizes of the island biota. 

The recency of oceanic islands as a group is undoubtedly of 

evolutionary significance also. They originated in the Tertiary, 

when the modern angiosperm flora had already evolved. In con- 

trast, some of the continental tropical ecosystems evolved with- 

out major geological disturbances forming a primary succession 

from seed fern forests to primitive gymnosperm and angiosperm 

forests to modern angiosperm forests. These continental eco- 

systems developed during a much greater evolutionary time span.

The high tree species diversity of some continental tropical 

forests may be largely attributable to this. 

Volcanism causes major geological disturbances. These per- 

turbations are a significant part of island ecosystem development 

from the highest mountain top to sea level. Volcanic pertur- 

bations are of many kinds and differing degrees and are erratic 

or unpredictable. They were once effective on each island and 

left their traces long after individual volcanoes became extinct. 

As such they had and still have a great effect on the evolution 

of the island biota. 

There is little doubt then, that the island biota evolved 

under unique environmental conditions. Much has been written 

also about their adaptive characteristics, which sometimes 

resulted in the development of rather bizarre island life forms. 

But has this also made the ecology of island ecosystems 

different? 

The answer, as revealed from our studies, appears to be that 

ecological principles do not differ for island ecosystems. How- 

ever, our studies have brought out some new dimensions to island 

ecosystem ecology, which should add to both their scientific 

understanding and appropriate management. 

Distributional Characteristics of Island Biota 

The spatial distribution analysis along the Mauna Loa moun- 

tain gradient confirmed Whittaker's individualistic species 

distribution model established for temperate mountains. It also 

confirmed the spatial association model of species distribution 

which is, in part, an affirmative answer to MacArthur's question 

on species and community patterns in the tropics. However, spa- 

tially associated species groups along environmental gradients 

are not to be considered unique for tropical areas, since such 

patterns have been demonstrated many times also in temperate 

environments. They are, like Whittaker's individualistic species 

distribution patterns, a universal phenomenon applicable to 

islands and continents, temperate and tropical environments 

alike. 

However, we found a number of other distribution trends, all 

of them more or less wide-ranging (e.g., bimobal, multi-modal, 

and broadly overlapping) . These all reflect generalistic ten- 

dencies of species behavior. The high proportion of these 

generalists in our biota groups are perhaps characteristic for 

geologically young areas or those relatively poor in species. 

This tendency may not be found, in geologically older areas, and 

thus also not so much on oldeg volcanic islands. 

An island characteristic, which may have interesting appli- 

cations, is that soil fungi, soil algae, and soil arthropods are 

probably mostly indigenous. This would imply as a hypothesis 

that they may form a community-similarity link with continental

mountain habitats in similar climatic and soil regimes. Con- 

versely, the other island community members, the higher plants, 

birds, canopy arthropods, tree borers,and Diptera flies all form 

much more unique species compositions. 

Community Structure and Niche Differentiation 

Island communities have the same gross-structural charac- 

teristics as found in continental communities. For example, 

montane tropical rain forests and lava tube ecosystems are also 

found on continents. At the species level, our island commu- 

nities are almost totally unique. But they are not so unique at 

the higher taxon lev'el. At this higher level one can find 

interesting similarities and departures from continental 

ecosys tems. 

In our community structure analysis we focussed on the 

general niche level, a functional ecological unit concept, inter- 

mediate between the individual species and the total ecosystem. 

We identified general niches by species of closely similar func- 

tion and structure, i.e., by life-form types (i.e., synusiae in 

plants, guilds in animals). 

We did not really find "empty niches" in the sense of 

absence of important life forms among the native species. The 

life-form spectra appeared complete in all organism groups ana- 

lyzed, i.e., plants, birds, canopy arthropods, and cave animals. 

This does not mean that "empty niches" may not be found in other 

island ecosystems, but it may imply that the empty niche phenom- 

enon is probably an exception rather than a rule in developed 

island ecosystems. What appeared to be a departure from conti- 

nental ecosystems of similar kind was that several important 

life-form groups had only one or a few native species, often with 

high quantitative importance. These appeared to occupy the more 

stable positions (or general niches) in the ecosystems in the 

sense that few exotics had invaded them. Conversely, exotic 

species invasion appeared to occur more readily in those general 

niches or life-form groups in which several native species occur 

with relatively small populations. However, this is a new 

hypothesis, which needs further testing. We have not yet given 

special attention to the ecology and relative stability of rare 

and endangered native species, which as a rule are probably more 

specialized. 

Instead our studies brought out the ecological versatility 

of some of the dominant native island biota. For example, in the 

Kilauea rain forest, all native tree species can grow on mineral 

soil and as epiphytes. The same applies to most of the herba- 

ceous native ferns. This phenomenon argues for the stability of 

native species composition under temporarily adverse forest floor 

conditions (e.g . , pig disturbance, flooding, ash blanket 

deposits) .

It is probable that ecological generalists among the native 

species prevail in the island ecosystems, which we analyzed. 

Both the Kilauea rain forest and the lava tubes are relatively 

young ecosystems, which support biota which are able to survive 

perturbation effects associated with volcanism. Frank Howarth 

mentioned the underground dispersal modes for the Hawaiian cave 

fauna. Thus, here we are dealing largely with biota, which are 

still displaying pioneering traits, a charactistic which all 

island biota must have had for becoming successfully established 

in the new island environment.

VARIABILITY IN DORSAL PATTERNING AMONG POPULATIONS OF 

HAWAIIAN "HAPPY-FACE" SPIDERS (THERIDION SP. OR SPP.) 

ON THE BIG ISLAND* 

William P. Mull 




Since the December 1972 discovery on O'ahu of an apparently 

undescribed comb-footed spider (Theridiidae) with a striking 

"happy-face" design on its back, close relatives have been found 

on Maui and the Big Island. Preliminary surveys of Big Island 

forests have produced over 100 specimens and revealed a remark- 

able spectrum of dorsal patterning variation among populations of 

these tiny (3-4 mm body length) Hawaiian leaf-dwelling spiders. 

Pending study by a specialist, it is unclear whether these popu- 

lations represent a single very variable species or several 

species. They are tentatively assignable to the widespread genus 

Theridion, which includes 10 described species (plus one sub- 

species) endemic to Hawai'i, only one of which bears resemblance 

to the "happy-face" group. These previously unknown Hawaiian 

spiders seem to be another example of the extremes of "genetic 

plasticity" and "evolutionary flux" expressed among other 

closely-related groups of Hawaiian organisms (e.g., Anomalochrysa 

lacewings, Achatinella tree snails, Metrosideros trees) in 

response to qenerous ecological opportunities and minimized 

inhibiting pressures presented to thbse groups by the unique 

Hawaiian environment. 

* Abstract

THE ROLE OF THE HAWAIIAN TWO-LINED 'OHI'A BORER, PLAGITHMYSUS 

BILINEATUS SHARP, IN THE DECLINE OF 'OHI'A-LEHUA FORESTS 

ON THE ISLAND OF HAWAI 'I* 

Richard P. Papp 




P1,agithmysus bilineatus Sharp (Coleoptera: Cerambycidae) , 

the Hawaiian two-lined 'ohi'a borer, is an integral part of the 

'ohi'a forest decline. This insect has been closely associated 

with the onset of decline symptoms (crown chlorosis and crown 

loss) in 'ohi'a trees at eight widely differing sites on the 

island of Hawai'i. Experimental implantation of - P. bilineatus 

larvae has also produced severe crown symptoms in otherwise 

healthy trees. Our data analyses indicate that P. bilineatus is 

a secondary invader of physiologically weakened tfees, but it is 

of primary importance as the only organism yet to be both consistently 

associated with decline and experimentally proven to be 

capable of producing decline symptoms in healthy trees. Although 

abiotic factors continue to be suspect in the initiation of 

'ohi'a forest decline, the role of this organism is now clear as 

an accelerating factor. Furthermore, since this species appears 

to be symptomatic rather than incitative in the demise of the 

'ohi'a forest overstory, it can be regarded as a beneficial 

organism. P. bilineatus hastens the destruction of the declining 

forest canopy, opens the forest floor to light, and promotes the 

rapid resurgence of the shade-intolerant native understory, thus 

helping to preserve the integrity of the rain forest ecosystem. 

* Abstract

ACARI ON MURINE RODENTS ON MAUNA LOA, HAWAI'IL 

Frank J. Radovsky, JoAnn M. Tenorio, 

P. Quentin Tomicha, and James D. JacobiJ 



As part of the U. S. International Biological Program, 

rodents were trapped seasonally during a two-year period at 

14 primary sites from 840 to 2440 m on a transect along the 

southeastern slope of Mauna Loa; rodents were also intensively 

collected in the Kilauea Forest near the transect. Three of the 

four murine species present in the Hawaiian Archipelago were 

taken: - Mus musculus, Rattus rattus, and R. exulans. Ectoparasites 

were recovered from rodents by a standardized washing 

technique. Mammalogical and parasitological data were analyzed 

by computer. The occurrence, host associations, and spatial distribution 

of some Acari are treated here. The occurrence of some 

parasitic mites was found to be partially independent of 

factors and associated with local differences in climate. 

host 

Abstract 

Research Unit, State of Hawaii, Department of Health. 

J Department of Botany, University of Hawaii at Manoa, Honolulu, 

Hawaii 96822.

HABITAT UTILIZATION AND NICHE COMPONENTS 

IN SOME HAWAIIAN ENDANGERED FOREST BIRDS* 

C. John Ralph 

Institute of Pacific Islands Forestry 

U. S. Forest Service 


In the native forests on the island of Hawai'i there is a 

species assemblage of 10 common passerine birds. Two of them are 

introduced, and six of the remainder are honeycreepers (three of 

these are endangered). A l l species except one are primarily in- 

sectivores. Four species utilize nectar sources to some extent 

and four utilize fruit. 

Detailed observations of habitat use and foraging behaviors 

coupled with indices of abundance of flowering and fruiting, as 

well as population estimates of the birds, provide a preliminary 

estimate of a wide variety of niche components. Introduced 

species appear to have a somewhat broader foraging niche than 

native species in the same guild. Competitive interactions seem 

to play a role in limiting populations of one of the endangered 

species, while another appears to be limited by its rather narrow 

niche. 

Interrelationships between phenology, abundance, and spatial 

distribution of food plants are all critical factors in deter- 

mining habitat utilization by the birds. Interspecific inter- 

actions appear to play a secondary role at most times. 

* Abstract

PLANTING, A TOOL FOR NATIVE ECOSYSTEM RESTORATION 

Don Reeser 



Natural Resources Management objectives for Hawaii Volcanoes 

National Park are to protect and restore native Hawaiian eco- 

systems. Making progress towards these objectives is a difficult 

task primarily because of introduced non-native agents competing 

with or directly destroying native conditions. We are making 

some headway, however. 

The primary tool at the command of the resource manager is 

"extraction," i.e., the removal or control of non-native ele- 

ments. Priority is given to those species which are considered 

to be the most destructive and competitive and for which tech- 

niques for control or elimination are feasible. Freeing Park 

ecosystems of goats, pigs, mongooses, feral cats, and certain 

exotic plants is the logical course of action where management 

emphasis has been placed in recent years and where it w i l l prob- 

ably remain in the future. Unfortunately extraction alone, 

regardless of how effective, will not counteract over 150 years 

of disturbance many areas of the Park have sustained. Large 

areas will continue to be dominated by exotic flora and fauna, 

and many rare or endangered species will continue to decline. 

Besides extraction, the only other significant tool the 

resource manager has left to pursue established objectives is the 

reintroduction of appropriate native species, primarily plants, 

where and when conditions are suitable. Propagating and reintro- 

ducing native plants is an integral and vital component of Hawaii 

Volcanoes National Park's Natural Resources Management Plan and 

is consistent with National Park Service Management policy which 

is: 

The reintroduction of native species into parks is encour- 

aged, provided that: 

--the species being reintroduced most nearly approximates 

the extirpated subspecies or race; 

--the species disappeared, or was substantially diminished, 

because of human-induced change--either directly or 

indirectly--to the ecosystem. 

The planting program is not new at Hawaii Volcanoes National 

Park. It has been a sporadic activity since the 1920's. Only in 

recent years, however, has it become a full time operation. 

Nearly a year ago a modern greenhouse was built here at the

Hawaii Field Research Center. The planting program is in its 

infancy, relative to its potential to contribute to the restora- 

tion and maintenance of native Hawaiian ecosystems. Some of the 

ways this program is or will be contributing are as follows: 

1. Restoration of man-made scars 

Following a disturbance such as rerouting the Crater Rim 

Road at Waldron Ledge an ugly scar has been created. If 

left alone, natural succession will produce a swath of 

exotic grasses through which it is unlikely 'ohi'a or 

other native trees can become established. We are 

planting this area to advance succession so that the 

scar will blend into the adjacent terrain. 

An ugly scar near the research center was created during 

the 1930's by bulldozing probably for the purpose of a 

recr,eation site for Civilian Conservation Corp. We are 

now raising 'ohi'a trees which will be transplanted here 

in an attempt to return this area to a more nearly 

native condition. 

2. Restoration of selected lowland sites following goat 

removal 

Significant changes are taking place in once goat- 

infested areas. Many areas of the lowland are dominated 

by exotic shrubs and grasses. Historical accounts and 

examination of remnant vegetation and natives which have 

reappeared provide clues to the orginal flora of the 

area. We are making some experimental plantings of 

selected areas to see if natives can be reestablished 

and to what extent succession following goat removal can 

be influenced to produce a more native flora. 

On the slopes of ~akahanu Pali several wiliwili trees 

survived 150 years of goat browsing. As a result of 

this goat browsing and a 7.2 earthquake in 1975, only 

one is left representing the last gene pool from this 

site. We hope to reestablish a population in this area. 

Canavalia first appeared inside a goat enclosure at 

Kukalau'ula and has also come up at Kaone several miles 

away. We planted Canavalia on the top of Pu'u Kapukapu 

on a site composeil entireTy of exotic grasses to see if 

it w i l l produce a native cover and give competition to 

the exotic grass. 

3. Exotic Plant Control 

Much effort and money is spent on trying to control 

certain aggressive plants. For some, mechanical cutting 

and/or herbicidal treatment is possible, but for others

this technique is futile. Native plantings could be 

used to compete with exotic species now that goats are 

removed and the thorniness or natural herbivorous 

defenses of many exotic plants do not necessarily give 

them the advantage. 

For example, Lantana is a noxious weed for which the 

State of Hawaii has introduced many insects for biolog- 

ical control. One technique which we are exploring is 

the feasibility of removing the problem exotic and 

immediately replacing it with a native before exotics 

can reinvade. 

4. Preservation and protection of rare and endangered 

species 

Possibly several dozen plants considered to be rare and 

endangered are not reproducing in the wild. We are 

hopeful that with the control of goats and pigs they 

will begin to reproduce. It appears there are other 

regeneration problems such as exotic grass cover, de- 

struction of seeds by rats, and exotic insects. There- 

fore, until solutions to these problems are found it is 

imperative that we continue to propagate and learn as 

much about these plants as possible so that their 

survival is ensured. 

5. Historical restoration and interpretation 

Greenhouse plant propagation can assist reestablishment 

of the historical scene and reestablish native or Poly- 

nesian plants used by early Hawaiians. These are 

located at archeological sites primarily along the Kala- 

pana coast and are planted out in consultation with the 

Pacific archeologist and historian. 

Plantings are recorded in books maintained at Park Headquarters 

and at the greenhouse. All the vital information on 

each planting is recorded and each site is pinpointed on maps. 

These records are always available for use by other researchers, 

or other interested persons. Monitoring of plantings is done on 

a sporadic basis whenever greenhouse personnel have an opportunity 

to get to the site. Size, condition, and mortality are 

recorded of a random sample of a given planting. More precise 

monitoring is planned to be able to follow the effectiveness of 

the program. As a side benefit of the planting program, an enor- 

~,ous amount of information is being collected and recorded such 

as location of rare and endangered species, flowering and fruiting 

times, etc. Insect collections are continuously made which 

are being identified and mounted by Mr. Cliff Gavis. In the 

greenhouse, germination techniques and a host of other information 

are being recorded which w i l l contribute to the overall 

program.

It is understandable that there is concern that the program 

does, in fact, truly enhance native ecosystems as intended. We 

fully recognize that there have been mistakes made in the past 

sueh as bringing in species which were never suspected of occur- 

ring in the Park or planting. species outside of their natural 

range. However with the evolving comprehensive ecosystem res- 

toration plan, rather than just a part time-greenhouse operation 

as it was a few years ago, errors of this kind should be avoided. 

A primary safeguard, however, is that the program be open to 

scientific scrutiny. The present resources management plan list 

of species to be propagated was compiled through consul tation 

with ecologists, botanists, taxonomists, and other interested 

persons so species, propagating material source, and planting 

locations can be evaluated, and input on the desirability of same 

can be received. It has been suggested that we go even. further 

than this in encouraging scientific input and expertise on the 

program by developing a discussion group composed of scientists 

from many disciplines who would meet regularly with Park per- 

sonnel. This would be a more fornal method of receiving advice 

and suggestions, and we are interested in discussing further the 

practicability of forming such a group. Another safeguard is to 

continually keep sight of the goal and to evaluate the impact of 

any resource management action on the native flora and fauna. It 

is native ecosystems which continue to decline islandwide. 

Hawaii Volcanoes National Park is one of the few places in 

liawai'i where objectives for ecosystem preservation and restora- 

tion are clear and unencumbered by conflicting land use policy. 

In this Park we have the opportunity to make some lasting headway 

in native ecosystem preservation and it w i l l be done through eco- 

logically sensitive resource manipulation. The reintroduction of 

native species is an important tool needed for doing the job.

FOREST BIRD SURVEY OF THE HAWAIIAN ISLANDS 

J. Michael Scott 

U. S. Fish and Wildlife Service 

Patuxent Wildlife Research Center 

P. 0. Box 44 

Hawaii National Park 


Despite Hawai'i's relatively small size, much of its flora 

and fauna may still be undescribed or undiscovered. A new genus 

of bird was discovered as recently as 1973 (Casey & Jacobi 1974) 

and many plants and insects remain to be described. Basic infor- 

mation on distribution, abundance, and biology is lacking even 

for many of the most common birds. This lack of information, 

combined with the limited area of natural vegetation, mu1 tiple 

and often conflicting demands for the land, and the vulnerability 

of island ecosystems in general, makes it imperative that we 

learn as much as possible about the native birds of Hawai'i now. 

A survey of the distribution and abundance of birds and 

their habitats on all the major islands was begun in 1976 by the 

U. S. Fish and Wildlife Service in cooperation with the Hawaii 

Division of Fish and Game, Hawaii Division of Forestry, U. S. 

Forest Service, National Park Service, and Private landowners. 

The objectives of the survey are to determine: 

1) the distributional areas for all forest birds in the 

study area; - 

2) the density (birds/km2) by vegetation type and eleva- 

tional strata for all birds within the study area; 

3) the population size for all forest birds for each 

vegetation type, elevational strata, and study area; 

4) habitat preferences for all forest birds in study area; 

5) occurrence of major vegetation types relative to the 

distributional patterns of birds; 

6) land use patterns and stability of habitats within each 

distributional area; and 

7) areas in which more detailed studies can be undertaken 

to clarify distributional anomalies and to identify 

limiting factors for endangered species.

To accomplish the objectives of the survey, transects are 

laid at intervals of 3.2 km (2 miles) on the Island of Hawai'i 

(Fig. I), and at 1.6 km (1 mile) intervals on the other major 

islands. Stations are placed every 134 m along these transects. 

The vegetation at each station is characterized according to tree 

height, canopy cover, species composition of the canopy and 

understory, and the presence of major habitat modifiers such as 

'ohi'a dieback, pig damage, banana p6ka (Passiflora mollissima), 

browsinq, - qrazina, - and loaqina. In addition, the fruitinq and/or 

flowering of thedblapa (~heirodendron spp. ) and 'ohi 'a - ( ~etiosideros 

collina) is recorded tor 10 plants at each stationxe 

occurrence of birds at each station is determined during eight 

minute count periods conducted during the first 4 hours after 

first light. By recording all birds heard and seen at each 

station and their initial detection distance it is possible to 

determine the density for each species encountered at a station. 

The area surveyed is a circle whose diameter varies with the 

species being censused as well as with the observer and vegetation 

type (Ramsey & Scott 1979; Reynolds et al., in press). The 

information on the occurrence of birds is related to the vegetation; 

multivariate statistics are used to determine habitat 

correlates. 

Additional information is obtained on the occurrence of rare 

plants and birds as a result of incidental observations made 

after the regular census period. 

The forest bird survey is conducted by a team of six trail 

cutters, 11 avian biologists, four botanists, and one statis- 

tician. On site observations as well as aerial photographs and 

direct aerial observations are used to conduct the various 

studies. To date, 815 km of trail have been laid, censuses have 

been conducted at 6247 stations, the vegetation characterized at 

3100 stations, and over 3000 man-days of work expended in our 

efforts to quantify the distribution and abundance of Hawai'i's 

birds and their habitats. Transects 1 t o 75 (Fig. 1) have been 

censused and it is anticipated that all the islands w i l l be 

surveyed by September 1981. 

Preliminary analyses of the data have greatly changed our 

understanding of the distribution and abundance of the rare and 

endangered birds on Hawai'i. Revised priorities for research and 

management have been adopted as the result of these findings. 

Threats to native birds and their habitats have been identified 

as well as possible ways to reduce or eliminate their impact. 

Distributional anomalies for endangered birds have been iden- 

tified which, when studied in detail, should assist us in deter- 

mining the limiting factors for these species. New study areas 

are being located and important information on breeding biology 

and habitat preferences is being obtained. Additionally, it 

should be possible to stratify sampling effort in future studies 

of Hawai'i's plants and animals, thus significantly reducing the 

time and money necessary to complete future studies.

When analysis of the data is completed it will be possible 

to state, for each species, its habitat preferences, area occu- 

pied (ha), the acreage of each habitat type within its range, 

relative stability of habitat types within its range as well as 

the density and population size for any subdivision of elevation, 

vegetation, or geography that is of interest. It is hoped that 

all this information will be used by land managers as well as 

researchers in their efforts to better understand and protect the 

plants and animals of Hawai' i. 


Casey, T. L. C., and J. D. Jacobi. 1974. A new genus and 

species of bird from the island of Maui, Hawaii (Passeri- 

formes: Drepanididae). Occ. Pap. B. P. Bishop Museum 

24 (12): 216-225. 

Ramsey, F. L., and J. M. Scott. 1979. Estimating population 

densities from variable circular plot surveys. In N. N. 

Cormack, L. P. Patil, and D. S. Robson, eds. 1nterGtional 

Ecological Congress Satellite in Statistical Ecology. 

Vol . 5. Sampling Biological Populations. Pennsylvania 

State Univ. Press. 

Reynolds, R. T., J. M. Scott, and R. N. Nussbaum. A variable 


press).

FIGURE 1. Map of the island of Hawai'i showing locations of 

transects used during Forest Bird Survey.

DIRECT SOWING OF TREATED MAMANE SEEDS: 

AN INEFFECTIVE REGENERATION TECHNIQUE 

Paul G. Scowcroft 





Most of you are familiar with the mamane (Sophora chrysoand 

mamane-naio (Myoporum sandwicense) forest ecosystems 

W'Mauna Kea Forest Reserve. I need not recount the history 

of this reserve--you know it as well as I. Suffice it to say 

that feral herbivores have been major contributors to the degradation 

of these ecosystems. 

Since 1970, the number of feral - sheep (Ovis aries) has averaged 

1500 animals. Mouflon sheep (Ovis musimon) havetotaled 200 

to 300 head. And feral goats (Capra hircus) have numbered about 

150 to 200 animals. Today, the number of browsing animals in the 

mamane and mamane-naio ecosystems is about 1/20th of what it was 

in the mid-1930's. 

Despite these relatively small populations, regeneration of 

mamane has not occurred in some areas while in others it has 

occurred at a slower rate than I would expect. This situation 

would probably persist even if all browsing pressure were elimi- 

nated. Such has been the case within several sheep exclosures, 

two of which are 15 years old. Elimination of browsing pressure 

within the exclosures has not been followed by an increase in 

mamane seedlings. Regenerating such areas with mamane would, 

therefore, involve the planting of seedlings and/or direct sowing 

of seeds (i.e., artificial regeneration). In my opinion, these 

efforts would fail if sheep still roamed the area. 

If browsing pressure is eliminated from the ecosystems, thus 

making artificial regeneration of mamane a practical management 

option, information about successful regeneration methods will be 

needed. The study reported here deals with artificial regenera- 

tion by direct seeding. The objective was to determine the ef- 

fect of seed coat treatment and sowing depth on mamane seedling 

emergence, survival, and growth under field conditions. 

The 1-ha Wailuku River sheep exclosure located at 2750 m 

elevation on the east flank of Mauna Kea (Fig. 1) was selected 

for the experiment because mamane regeneration was lacking in a 

nearby exclosure built in the early 1960's.

METHODS 

Old mamane seed pods were collected from Hale Pohaku, 2750 in 

elevation. Seeds were extracted by hand and sorted to remove 

damaged ones. Intact seeds were kept in sealed plastic bags at 

room temperature until sowing. 

The two factors examined in this study were seed coat treat- 

ment and sowing depth. Four seed coat treatments were tested: 

1. Acid soak: 

2. Sanded: 

3. Hot water soak: 

Seed immersed in concentrated 

sulfuric acid for 60 minutes. 

Seed abraided between two sand- 

ing blocks for about 1 minute. 

Seed immersed in water at 100°C 

and allowed to soak until the 

water reached room temperature 

(overnight). 

4. Control: Seed not treated. 

Seeds were sown at six depths: broadcast over the surface, 

spot sown at 1.3 cm, and at every 2.5 cm thereafter to a depth of 

about 11 cm. 

Three blocks, each divided into four plots, were laid out 

within the Wailuku exclosure. Seed coat treatments were randomly 

assigned to the plots in each block. Plots were further divided 

into six subplots each and sowing depths were randomly assigned 

to them (split-plot design.). Movement of surface sown seeds was 

restricted by means of buried hardware cloth around the perimeter 

of the appropriate subplot. - - 

Eighty treated seeds were sown in each subplot during the 

first week of March 1974 when the upper 13 cm of soil were moist 

to sight and touch. Seed spots were made by inserting a rein- 

forcing rod to the desired depth. Spacing was 10 by 10 cm. Only 

one seed was sown in each spot after which soil was poured into 

the hole and lightly tamped. 

As seedlings emerged, numbered plastic markers were stuck 

nearby in the ground. The condition and height of each seedling 

were recorded. 

Weekly measurements were made for the spring test to the 

15th week after sowing. Thereafter, measurements were made every 

4 weeks through the 54th week when the test was terminated. 

Differences in seedling emergence, survival, and height due 

to seed coat treatment and sowing depth were examined by standard 

ANOVA techniques and multiple range tests using appropriate 

transformations. In addition, height-age regressions were fitted 

to the data and the coefficients were compared.

RESULTS 

None of the seed sown on the soil surface germinated during 

the test. Therefore, I did not include this sowing depth in the 

analyses. 

Seedling emergence 

Analysis revealed that both seed coat treatment and sowing 

depth significantly (p

Seedling survival 

Seedling survival was low. Of the 602 seedlings that 

emerged, only 99 (16%) were still alive 54 weeks after sowing. 

Was survival affected by sowing depth? By type of seed coat 

treatment? After looking at my data, I decided I could not 

answer these questions for the control and hot water treatments 

because so few seedlings emerged from these. Therefore, I only 

compared seedling survival between acid and sanded treatments 

(Table 2). 

Depth at which seed were sown did not significantly affect 

end-of-test seedling survival. However, seedling survival was 

affected by type of seed coat treatment; significantly greater 

survival was observed for seedlings originating from sanded seed. 

I examined the relationship between percent survival and 

week of emergence and found that survival was not dependent on 

week of emergence. Seedlings that emerged relatively early had 

just as good a chance of surviving as those that emerged later. 

Another expression of survival is seedling age--that inter- 

val between the time a seedling was first seen and the time it 

was declared dead or the test ended. Analysis showed that seed- 

ling age was not significantly affected by week of emergence, 

seed coat treatment, or sowing depth. 

One unexpected discovery was that some seedlings tallied as 

dead suddenly reemerged, one as much as 24 weeks later. Of the 

602 seedlings that emerged, 57 exhibited this behavior. Thirteen 

of these were still alive at the end of the test, thus accounting 

for about 13% of the surviving seedlings. 

Seedling height 

Height-age regression curves were fitted to my data (Fig. 3). 

Comparisons of the regression coefficients for the acid seed coat 

treatment showed that growth was significantly greater for seed- 

lings from the 11.4 cm depth than for those from other depths. 

The same was true for the sanded treatment. Regression curves 

for the control treatment--3.6, 6.4, and 8.9 cm depths only--were 

significantly different from each other with seedlings from the 

6.4 cm depth growing tallest. 

The effect of seed coat treatment on seedling height was not 

clear from my data. 

The coefficients of determination (r2) for the regressions 

were low. Obviously, factors other than seedling age, seed coat 

treatment, and sowing depth were affecting height growth.

About 70% of the seedlings died back either prior to their 

death or prior to the end of the study. The greatest proportion 

of seedlings exhibiting dieback (77%) came from the 3.8 cm depth 

followed in order by the 6.4 cm (71%), 8.9 cm (70%), 11.5 cm 

(64%), and 1.3 cm depth (47%). 

DISCUSSION 

The management implications of these results are clear; 

direct sowing is not viable regeneration technique on sites simi- 

lar to the one I used. Maximum emergence was only 54%. Survival 

was low and height growth slow. 

Compared to planting of nursery-grown seedlings, direct 

sowing is not effective for regenerating mamane. About the time 

the test began, 36 containerized seedlings with an average height 

of 24 cm were planted in the Wailuku exclosure. Survival was 47% 

in April 1978 compared to 3% for seedlings from this study. 

Average height of survivors was 52 cm for the planted seedlings 

and 31 cm for the others. Planting was clearly the superior 

regeneration method.

TABLE 1. Comparisons of percent seedling emergence means 

for each combination of seed coat treatment and 

sowing depth. 

Seed Coat Sowing Depth 

Treatment 1.3 3.8 6.4 8.9 11.4 

Percent 

Acid 6dL 54a 34b 15c 15c 

Sanded 3def 32b 34b 22bc 15c 

Control og 6d 4de 3def < lefg 

Hot Water c lfg ldefg ldefg < lfg < lfg 

1 Means followed by the same letter are not significantly 

different (Duncan's Multiple Range Test). 

TABLE 2. Average percent survival of mamane seedlings 

by sowing depth for acid and sanded seed coat 

treatments. 

Seed Coat Sowing Depth (cm) 

Treatment 1.3 3.8 6.4 8.9 11.4 Average 

Percent Survival 

Acid 0 9 12 9 11 10 

Sanded 3 3 25 2 4 21 20 2 4 

Average 14 15 18 16 17 --

Figure l--Map of Mauna Kea, island of Hawaii, showing the location of the Wailuku 

exclosure study site.

254 

Figure 2--Cumulative number of emerged mamane seedlings over time for each sowing 

depth and seed coat treatment during the test period, March 7, 1974 to 

March 18, 1975. 

100 - 

80 - 

60 - 

40 - 

..*. 

/ ................................................................................... 

Acid 

I.... Sanded 

I ; ... 

1: 

Control 

Hot water 

li 

80 , /;.Y*\ _ - _ - - 

60 . 

40 . 

20 - 

0. 

.................................. 11.4cm depth 

I 

March 

-.- 

, 

--, 

____________- 

fl ____---_--_-----I__________ 

9 

t.*t*.*t.,t.t.*.*. 6-4.m depth *... 

: 

1 /i Acid and sanded curves 

coincide 

... 

.a 

/- ~ 

I 

June 

/,' 

'~..'*.*.*..*.....'*.**+. 

________.__--__-------____--------------. 

*.*.**.*., .*.,*,.* .*,* e.gcm ".*~."*' ."*t.*.,. 

............. ......... ^ ................................ 

60 1 

...... 

...... 

~ . 

~- 

............................. 

I I 

September 

Weeks Since Sowing 

" ...... 

I 

oecember March

FIGWIE 3--SEEDLING HEIGHl OVER TIME FOR EACH SOWING DEPTH WITHIN A GIVEN SLED COAT 

TREATllENT DURING THE SPRING TEST. 

SO~IING DEPTH ~ r n 

I?? EQUATION SOWING DEPTH icn, R' EQUATION 

6 3.8 0.205 7.959 + 5.970LNCXI a 3.8 0.147 7.285 + L.B~OL~~ 

b.Y 0.n7 3.243+9.139LNiX> s - r b.LI 0.00L 10.838 + O.LLILLN 

+ 8.9 0.2(13 LO.1123 + 2.847LNCX> 9, 

'I

COMMUNICATIONS TECHNIQUES AND THE SCIENTIST 

Jerry Y. Shimoda 

City of Refuge National Historical Park 

Kona, Hawaii 

(Write word SEX on the Board) 

Now that I have your attention, let me tell you why I think 

I am here. Perhaps the reason for having me present my paper 

first has something to do with Superintendent Barbee's notice of 

April 10 to all participants of this Science Conference. In it, 

he appealed to us to transmit our research into easily understood 

language so that our papers can be enjoyed by the highly mixed 

group of persons here today. 

I consider myself a layman in the midst of scientists, so 

let me express some of my concerns. Most of you will be using 

what you think is the easiest form of public communications--the 

lecture. Actually, although it is the fastest and easiest form 

to use, it is the most difficult one to transmit messages with. 

anyl lay people shudder at the thought of attending a Science 

Conference because they will not understand what is going on. 

But if the scientist needs public support for his research proj- 

ects, he must somehow be able to arouse the interest of these 

people and help them to understand. A good communicator always 

looks at his presentation with this key thought: "How can I help 

my listenegs to better understand what I am saying?" 

he -scientist of ten 'uses scientific names of plants; birds, 

and animals without giving the common name. I suggest writing it 

out: Colocasia esculenta--Taro! (Write scientific name on flip 

chart). Then, use the word "taro" for the rest of the - presentation. 

One of the dangers that any group of people can fall 

into is the "in-house language." Anyone who communicates-with 

the public needs to be aware of this. For example, (write on 

board) I work for the NPS, - DI, our central office is - WASO, and it 

establishes policies tE are interpreted and comes to CIRE 

through - WRO and - HISD. Permit me to explain those terms. N P ~ 

National Park Service, DI is Department of the Interior, WASO is 

the Washington, D. C., Office, CIRE is City of Refuge National 

Historical Park, WRO is Western Regional Office, HISD is Hawaii 

State Director's office. 

Permit me to present another example of "in-house talk." 

Some years ago, I worked at Saratoga National Historical Park, a 

Revolutionary War Battlefield, in upstate New York. We sold a 

booklet about the battle there. One day, I noticed some visitors 

looking at the booklet and chuckling among themselves. I found 

out why! For those of you who are not familiar with the Battle

of Saratoga, it occurred in the fall of 1777. The American army 

was commanded by General Horatio Gates and the British army by 

General John Burgoyne. The passage in question in the booklet 

was describing the advance of Burgoyne's army toward the American 

position. It said, "Burgoyne's right and left flanks lay in the 

woods, but his front was open." We quickly revised that passage, 

of course! 

In good communications we must SIMPLIFY (write on board)! 

But this does not mean reducing your presentation to a childish 

one. Perhaps the scientist needs to look at things in the fol- 

lowing ways. He needs to be bilingual and at the same time have 

a good understanding of objectives, human relations, and the 

techniques of presentations--both verbal and non-verbal. 

Let us look at each of these items separately. Be BILINGUAL 

(write on board)--like knowing a second language. Learn the 

language of the communicator, e.g., use the active voice--"I went 

to the woods" instead of "I had gone to the woods," "We decided" 

instead of "It had been decided by us." Concentrate on short 

sentences instead of those long ones with 40, 50, or more words. 

Your listener w i l l find it easier to follow your presentation, 

instead of having to follow you through all of the semicolons, 

commas, adjectives, and adverbs, until you finally get to your 

point. 

Use words that touch your listener's emotions so that he 

w i l l become involved in what you are saying. For example, it may 

be better to say "stink" instead of "odoriferous," or "home" 

instead of "house," and "killed" instead of "dispatched ." 

Graphic words are better in public communications, but - not to the 

point of nausea. 

One may say, "But there is no simple word for that flower or 

that animal." My reacton to that is, "Use the word, but explain 

it. Write it on the board!" (Point to Colocasia esculenta writ- 

ten on flip chart). Sueakinq of words, a speaker should avoid 

profanity -or delGing Into the gory as-a shock treatment for they 

tend to turn off a good part of his audience, and cause them to 

become negative toward him and what he is saying. 

In speaking, leave out as many "1's" as possible, like "I 

did this" and "I did that." When you do even little things like 

this, you are "removing the static so that your listeners can 

hear the music." 

In preparing a speech, decide your objective for giving it, 

even before you start your outline. For example, in preparing 

for this paper, "Communications Techniques and the Scientist," 

the first thing I did was to decide that my objective would be 

change. I would try to change your thinking so that you would 

think of your listeners, and try to help them to better under- 

stand what you are saying, instead of telling your listeners of 

the great things you have accomplished, in language they cannot 

understand.

In a ten-minute paper like today's, a clear objective is 

needed because you do not have much time to reach one. Choice of 

words and thoughts also become important. As the presentations 

are made today and in the following two days, you will notice 

that some of the ten-minute papers will seem short and others 

will seem long. That will be because of the way they are written 

and the way they are presented. Remember that there is not a 

boring subject in the world. It is all in how you tell it! 

To make a good speech, it is necessary to understand human 

relations. Sensitivity to the listeners' feelings is important. 

Remember that without the listeners, there is no need for a 

speaker! Try to relate what you are saying to your listeners' 

experience as much as possible. -- Do not present the same paper 

you presented last night at the dinner meeting of botanists, to a 

luncheon meeting of the local Lions Club today. Too many times 

this happens because we are not interested enough in having the 

listeners understand what we are saying, but are interested only 

in having our names in the newspaper that says we made a speech 

somewhere. 

Now, let us review a few of the things any speaker should be 

aware of, and look at some new things. A person who is going to 

present a paper or give a talk on a particular subject must be 

willing to pay the price of preparation. He must have an objec- 

tive in mind and must use an outline. The beginner should write 

his talk out, re-write until he is satisfied, then practice out 

loud on lay persons (with graphics if he is going to use any). 

Then, re-write again based on their feedback. 

Arrive early at the meeting place to set up your equipment, 

and to look at the layout of the room. It is always better to 

provide your own equipment, e.g., movie projector, slide pro- 

jector, or tape recorder, because you are more familiar with it 

than with one that the facility provides. 

When you come up to the stand, be ready to speak. Do not 

play with your papers, your glasses, etc. Take ten deep breaths 

to help pull yourself together, then begin. 

In making your presentation, relax, be natural, and throw 

your voice out to the people in the last row. Eye contact with 

the audience is also very important. Movements help, but not 

distracting movements, like playing with your glasses, shifting 

your weight from one foot to the other, or playing with the coins 

in your pocket. Use gestures, but they must look natural. 

People say "But I can't do gestures because when I try them they 

feel forced." Then practice! I am doing gestures deliberately 

right now to show you that deliberate gestures need not appear 

forced. 

In communicating with the public, the burden is on you, the 

speaker. Keep in mind that the hardest parts of a day to give a 

speech are: bright and early in the morning, right after lunch, 

and right after a dinner. You need to use all the tricks in the 

book to keep the audience's attention.

under grass than in roots exposed to direct sunlight. Most roots 

with suckers show no obvious scars of damage. 

Rooting of cuttings under mist 

Mist rooting of cuttings provided a higher percentage of 

rooting than did air layering, but the rooted cuttings had a much 

lower percentage of survival after transplanting than did the air 

layers. Therefore, it is difficult to determine which of these 

tdo successful methods of propagation is the better. 

Most of the successes occurred when perlite was used as the 

rooting medium and 3000 mg/kg IBA in talc as the rooting sub- 

stance (Table 4). Rooting was obtained only once on a phyllo- 

dinous cutting. One experiment (Nos. 7, 8, 9) indicated that 

rooting could be improved by supplying nutrients to the cuttings 

while they were under mist. In the potassium nitrate (KN03) 

solution portion of this experiment, one phyllodinous cutting in 

addition to half of the true-leaf cuttings rooted. Unfortu- 

nately, these results could not be duplicated when the experiment 

was repeated on two occasions. The difference may have been that 

in the first test the cuttings were placed under mist the same 

day that they were obtained. The other experiments were started 

with 1- to 2-day-old material. 

Although an average of 20% of cuttings rooted when IBA and 

perlite were used, only 14 propagules survived transplanting. 

The low percentage resulted because koa cuttings were very slow 

to root and produced very few roots per cutting. Rooting usually 

took 2 months or longer. By the time the cuttings rooted, almost 

all the leaflets had dropped and when they were transplanted, the 

rest of the leaflets were usually lost. 

The methods we found most successful were to collect root 

suckers that h a d reached t h e round rather than. .sqware.stem. stage. 

These were placed under mist the same day they were collected. 

We rubbed the freshly cut end in 3000 mg/kg IBA in talc and put 

the cuttings in individual styrofoam cups of perlite to reduce 

fungus contamination. Cuttings that rooted were transplanted as 

soon as discovered to black plastic bags containing a mixture of 

mica-peat and perlite. The bag was then closed and sealed around 

the stem to reduce moisture entry. The packaged transplant was 

then held under mist for about 3 weeks before transfer to a shade 

house. 

Recently, we have obtained an excellent source of stool 

shoot and root sucker material from one superior tree that was 

mistakenly cut by loggers and using the methods described we have 

twice achieved the usual 20% rooting, and have increased the 

ultimate survival rate to 12% of the original cuttings started. 

Seven rooted cuttings have been planted at Laupahoehoe and 

five at Keauhou. The seven at Laupahoehoe have grown in a very 

similar fashion to the air layers. One died a few months after 

planting. Three others have become infested with Uromyces koJe,

ut are now growing in a normal upright fashion. Two others show 

plagiotropism. All have formed phyllodes, but all are so far 

much less vigorous than seedling trees growing in the same area. 

Clonal variation 

The percentage of rooting of cuttings and air layers indi- 

cates a possible pronounced clonal variation in rootability 

(Table 5). These data are only for rooting, not for ultimate 

survival of the propagules and are, of course, strongly weighted 

by the variation in numbers of cuttings worked. Only those trees 

that produced propagative material were listed. Six trees-- 

numbers 1, 10, 18, 19, 20, and 22--provided a fair size sample of 

material which did not root. They can be compared with the quite 

good rooting performance of material from trees 2, 8, 9, 15, and 

26. These results are only indicative of possible differences. 

They cannot be validly compared statistically because of the 

differences in conditions at each attempt in rooting. 

Since these data were obtained, we have started more than 

100 cuttings from tree number 2 of which 10 to 12 will survive. 

The key to success in all this propagation work is to use large 

amounts of material, and this has proved to be impossible with 

the superior trees up to now. 

CONCLUSION 

Acacia koa can be propagated by air layering or rooting of 

cuttings under mist. However, neither procedure has yet been 

developed sufficiently so that it can be considered a practical 

method of propagating forest-grown superior trees. So far only a 

few propagules of such trees have been produced, and many of them 

are showing slow growth, disease susceptibility, plagiotropism, 

or poor form. 

The only practical way of propagating the species is from 

seed. This has several obvious disadvantages for tree improve- 

ment. Few seeds are produced by the tall, well-formed superior 

trees which have crowns above the general forest canopy where 

they are subjected to strong winds and, probably, low populations 

of insect pollinators. The seeds are open pollinated, so progeny 

expresses only a portion of the genotype. 

Most of our tree improvement work with koa will be concen- 

trated on seedlings. We are gradually building up a supply of 

sufficient superior tree seed to do a progeny test which w i l l 

become a future seed orchard. We can also collect seedlings from 

beneath the trees to transplant to progeny tests.

We are continuing to explore ways of improving vegetative 

propagation of the species. The latest technique we are trying 

is to air-layer root suckers in the field and then bring them in 

as cuttings to root under mist once swelling has occurred above 

the girdle. This works well with southern hardwoods, but our 

trials of the method are not yet far enough along to report on. 


Hartmann, H. T., and D. E. Kester. 1975. Plant propagation 

principles and practices. Third Ed. Prentice-Hall, N. J. 

662 pp. 

Kormanik, P. P., and C. L. Brown. 1974. Vegetative propagation 

of some selected hardwood forest species in the Southeastern 

United States. N. Z. J. Forestry Sci. 4(2): 228-236. 

Murashige, T., and F. Skoog. 1962. A revised medium for rapid 

growth and bio assays with tobacco tissue cultures. Physio. 

Plant. 15: 473-497.

Table 1--Grafting trials of Acacia k s 

Reference 

Stocks 

Type 1/ 2/ 

number Grafts of graft- Scions- and age Remarks 

Side veneer 

Side veneer 

t-bud 

Side veneer 

t-buds 

Side veneer 

Top cleft 

Patch buds 

Side veneer 

Side veneer 

Top cleft 

Approach 

Side tongue 

Whip 

Side veneer 

Shoot tip 

(juv. & mature) 

Root sucker tip 

(juv. ) 

Root sucker lateral 

(juv. 

Stem sprouts 

(juv. ) 


(juv. ) 


(juv.) 


(juv.) 


( juv. 


(juv. ) 

Seedling branch 

(10 mo. old) 

Seedling tip 

(juv. 

Root sucker 

propagde 

Seedling tip 

(juv.) 

Seedling tip 

(mature) 


(juv.) 

v~escriptive terms follow those by Hartmann and Kester (1975). 

Seedling 

4 mo. 

Seedling 

5 mo. 

Seedling 

5 mo. 

Seedling 

6 mo. 

Seedling 

7 mo. 

Seedling 

7 mo. 

Seedling 

8 mo. 

Seedling 

9 mo. 

Seedling 

9 mo. 

Seedling 

10 mo. 

Forest 

trees to 

1 yr. 

Seedling 

5 mo. 

Seedling 

7 mo. 

Seedlina 

10 mo. 

Seedling 

10 mo. 

- 

~/JUV. indicates juvenile or true-leaf stage; mature indicates phyllode stage. 

18 (9 of each scion) 

kept under mist 

One-half kept under mist 

One-half kept under mist 

Sprouts girdled, graft 

at swell above girdle 

Raffia and grafting wax, 

expert grafter 

Raffia and grafting wax, 

expert grafter 

Scion and stock same plant 

Seedling tips to forest- 

grown tree stumps 

Scions were potted 

cuttings and air layers

Table 2--Rooting and survival of air layered root suckers and 

stem sprouts of superior trees 

Tree 

number Air layers Xooted Surviving 

Total 159

Table 3--Treatments applied to Acacia koa roots intended to 

induce suckering 

rea at men@ Trees Treatments 

Knife wounding (deep) 

Knife wounding (shallow) 

Expose root to sun 

"Chew" with pliers 

Pound with hammer 

Heat with torch 

Girdle root 

Raise root on rock 

Bury exposed root 

Wound - kinetin (100 rng/L) 

Wound - NAA (500 rng/ll) 

Wound - Ethrel (100 mg/L) 

Wound - B (500 rng/i) 

Wound - IAA (200 rng/ll) 

Wound - GA (500 rng/ll) 

Air layer (untreated) 

Air layer B (500 rng/L) 

L/IAA = indoleacetic acid; NAA = napthaleneacetic acid; 

B = benzyladenine; GA = gibberellic acid.

N m b x Appl ica- 

Ref. Cut- Ontogenetic Aw in t ion 

No. tiqs Stage source Auxin' Strength bktixd mting Wiun Mnnber Fematk 

True-leaf 

True-leaf 

True-leaf 

True-leaf 

True-leaf 

True-leaf 

True-leaf 

Rue-leaf 

hue- leaf 

True-leaf 

Rue-leaf 

True-leaf 

Fmtsuckers IEA 

Fmtsuckers IBA 

Stem sprouts IBA 

Ste'Tl sprouts IBA 

Stem sprouts IBA 

mtsucker IBA 

mt sucker IBA 

mot sucker IBA 

mt sucker IBA 

Stem sprouts None 

sten sprouts None 

stm sprouts None 

--- - 

' IBA = Indolebutyric acid; NAA = Napthaleneacetic acid. 

3000 q/kg Talc 



3000 -/kg Talc 


3000 ml/kg Talc 




- - 

Perlite 

Perlite 

Funning water 

Runniq water 

Running water 

Perlite 

Perlite after 10 min 

NaOH 

Perlite after 10 sec 

H2SO4 

Perlite 

Perlite after 10 min 

NaOH 

Perlite after 10 sec 

H2SO4 

Perlite 

7 11% rooted 

10 27% rooted 

0 

0 

0 

- 

- 

- 

19 18% rooted 

3 - 

1 - 

1 - 

- 

0 

0 

0 

- 

- 

.

Table 5--Clonal variation in rootinq of root suckers and stem sprouts 

of superior trees 

Percent 

Tree both rooting 

number Air layers Rooted Cuttings Rooted methods

STAND ANALYSIS OF AN INVADING FIRETREE 

(MYRICA FAYA AITON) POPULATION, HAWAI'I 

- 

Garrett A. Smathers 

CPSU Western Carolina University 

Cullowhee, North Carolina 28723 

and 

Donald E. Gardner 



INTRODUCTION 

Fi retree (Myrica fa=), an aggressive, noncomm ~ercial, exotic 

species that is native to the Azores, Madeira, an d Canary Islands. 

has been s~readina ra~idlv in Hawai'i for approximately 

80 years. This treeLwas introd;ced- in Hawai' i for reforestation 

in the late 1800's, but by 1944 it had become so aggressive in 

colonizing agricultural and forested land that the Board of Agriculture 

and Forestry was pursuing a control program to eradicate 

it (Neal 1965). 

Distribution and Controls 

Firetree concentration is dense on the islands of Maui and 

Hawai'i. However, the major efforts of control have been on the 

island of Hawai'i. In 1961 Kawasaki (unpublished) reported that 

the major concentrations on the island of Hawai'i were along the 

Hamakua Coast from Laupahoehoe to Honoka'a, then mauka (toward 

the mountain) to the Parker and Kukaiau Ranches. A smaller popu- 

lation covering 300-400 acres in the 'Ola'a Forest near Hawaii 

Volcanoes National Park (HVNP) was also reported in the 1961 

survey. 

In a 1966 survey Kawasaki (unpublished) estimated that the 

'Ola'a Forest infestation had increased to 4500 acres, including 

1500 acres on Forest Reserve Land and 25 acres in HVNP. Two ad- 

ditional infestations were observed in the National Park: (1) a 

50-acre site on the northeast rim of Kilauea Crater; and (2) a 

150-acre site at the intersection of the Chain of Craters Road 

and the 'Ainahou Escape Road. 

Invasion of firetree on the island of Hawai'i has increased 

exponentially. A 1970 survey revealed that more than 40,000 

acres of the island were infested with firetree (Walters & Null 

1970). The 225 acres infestion in HVNP in 1966 had increased to 

approximately 9000 acres in 1977. Infested acreage varied from 

light (1 tree/acre) to heavy (1000 trees/acre) concentrations,

with the major distribution being in the seasonal dry forest sec- 

tion of the Park. 62,776 firetrees were removed from the Park 

from 1967 to 1974, and from 1975 to 1978 an additional 30,884 

were destroyed, making a total of approximately 100,000 trees 

removed over a 10-year period (Donald Reeser, Resources Manage- 

ment Biologist, pers. comm.), and yet the plant continues to 

spread (Smathers 1976). The National Park Service considers the 

firetree invasion an unnatural phenomenon that threatens and im- 

pairs the natural and historic quality of the Park's vegetation. 

The United States Forest Service and the Hawaii State Board 

of Forestry consider firetree to be an aggressive exotic with no 

commercial value, occupying land which should be utilized for 

agriculture and commercial forestry purposes. The State of 

Hawaii has conducted a control program for nearly 20 years, but 

funding and manpower have caused considerable fluctuations in 

this effort. Herbicides are the primary means of control. Of 

the various herbicides, Tordon 22k has proved the most successful 

in giving complete canopy kill and 99% control of sprouting 

(Walters & Null 1970; Walters 1973). 

Controls by the National Park consist of uprooting small 

trees and using Kuron herbicide (2,4,5-TP) on medium and large 

trees. In addition to chemicals, several species of insects have 

been tested for control of firetree, but none have been success- 

ful (Krauss 1964). 

Ecological Evaluation 

As yet no comprehensive ecological evaluation has been made 

of the long-term impact of firetree upon the native vegetation. 

It seems reasonable that such a study should be conducted, con- 

sidering the long period firetree has been colonizing the island 

ecosystems, and since its complete eradication is not likely. 

The latter is especially true in wildlands where agriculture and 

forestry are not practiced. Such a study would reveal the eco- 

logical role that firetree has with native species as well as 

with other exotic species now naturalized to Hawai' i. Informa- 

tion of this type would provide resource managers a better knowl- 

edge of how to evaluate the firetree's presence in light of their 

agency's mission and policy. 

There is now an excellent opportunity to study firetree as 

it invades a series of ecosystems in Hawai'i. Since 1971, fire- 

tree has been invading the Devastation Area of the 1959 Kilauea 

Iki Crater eruption site in HVNP (Fig. 1) where a similar compre- 

hensive ecological study has been underway for nearly 20 years. 

A main objective of the Devastation Area study is to determine 

the competitive relationship between native and exotic plants as 

they colonize recent volcanic substrates. Results of this study 

have shown that exotic and native plants have both a competitive 

and complementary relationship. In all habitats, native woody 

plants were eventually capable of replacing or holding their own 

with exotics. The Park manager has now set aside a part of the

Devastation Area for a concentrated study of the invading 

firetree population. 

STUDY AREA 

In December 1959 Kilauea Iki, a pit crater on the summit of 

Kilauea Volcano, erupted and deposited a blanket of pumice over 

an area of 500 hectares. Later the entire area, which is approx- 

imately 1200 m in elevation, became known as the Devastation 

Area . The latter name was given to the area because of wide- 

spread destruction of both a montane rain forest and a seasonal 

dry forest. With its variety of climates, substrates, and con- 

tiguous populations of native and exotic plants, the area pro- 

vided a unique opportunity to study the formation of new plant 

communities. 

Immediately after the eruption, a study was begun of plant 

invasion and recovery within the six habitats. These habitats 

were recognized by kinds of substrate and remains of the former 

vegetation. A series of permanent photo stations, belt transects 

(Fig. 2), and quadrats were established to record the chrono- 

logical sequence of plant succession and recovery. The results 

of this study have provided information heretofore unknown on the 

phytosociological relationships of native and exotic plants 

(Smathers and Mueller-Dombois 1974; Smathers 1976). 

During the 15-year observation period (in 1974) a small 

population of firetree seedlings was recorded in the western part 

of Habitat 5 near Byron Ledge. 

invasion started about 1971. 

It is estimated that the initial 

The habitat is characterized by the large number 

+ 

of surviving 

native 'ohi'a (Metrosideros collina subsp. 01 mor ha) 

trees with a pumice layer that varies from approximate y 30 cm to 

3 m in depth (~abitat 5 in Fig. 3). It is in-the lee o? the cinder 

cone that formed during the 1959 eruption and slopes gently 

in a southwesterly direction. This habitat is somewhat protected 

from the prevailing northeasterly trade winds. However, it 

receives greater insolation in the lower sectors because of 

decreased cloud cover. The approximate mean annual air temperatur 

e is 17°C and the mean annual rainfall approximates 

2700 mm. Mean evaporative rate from a Livingston atmometer was 

6 ml/day/week with a standard deviation of 2.7. The climate is 

characterized by humid mild winters and warm dry summers 

(Smathers & Mueller-Dombois 1974). 

METHODS 

A survey was made of the firetree infestation pattern in 

Habitat 5 to determine its boundary, homogeneity, and the direc- 

tion of invasion before placement of transects for stand anal- 

ysis. Two permanent belt transects at right angles to one 

another were established in the infested area to meet the above 

criteria. It was not possible to determine direction of invasion

as size classes (height and diameter) were found evenly distri- 

buted throughout the populated area. One transect, C-C', which 

was 180 m long and consisted of contiguous 10 x 10 m plots, was 

originally established in 1960 for the Devastation Area study. 

The other transect W-W' , was 70 m long and consisted of 10 x 20 m 

contiguous plots (Fig. 2). 

Ninety-six firetrees were sampled within the transects for 

height, basal diameter, vigor, and phenological characteristics. 

A vigor rating of good to average included specimens with dark 

green to green foliage, mature fruits, and strong to medium ter- 

minal growth. A rating of poor included specimens with numerous 

pale green to chlorotic leaves, defoliated branches, fruit fall- 

ing before maturity, and terminal die back. Associated plant 

species were recorded as to their physical position in relation 

to each firetree. Parameters of density, frequency, and percent 

cover were determined for the various height and diameter ranges. 

Unusual growth forms were also recorded. 


Stand Structure and Vigor Characteristics 

The structure and vigor of the firetree population are shown 

in Table 1. By totaling both transects (C-C' and W-W' ) the 

highest number of trees (61) is in the 2-5 cm range; the second 

highest (32) is in the < 2 cm seedling range; and the next lowest 

number (2) is in the 6-7 cm range. The number trend in diameter 

sizes indicates that firetree is making a steady-progressive 

invasion of the area, but the small number of trees with mature 

fruits (Table 2) could not likely be the parent stock of the 

large number (32) of seedlings (< 2 cm). 

Of the 96 firetrees examined, 89 were directly associated 

with the native 'ohi'a trees by being rooted beneath their 

crowns. In the < 2 cm diameter range, 27 seedlings, having a 

height range of 0.20-2.00 meters, grew beneath 'ohi'a trees, and 

10 of these over 1 m tall were beginning to interlock with the 

lower 'ohi'a branches. In the 2-3 cm range, 34 shrubby firetrees 

were growing beneath 'ohi'a trees, and 25 of these were inter- 

locking with 'ohi'a branches. In the 4-5 cm range and up to the 

4 m height range, 25 shrubs grew beneath 'ohi'a with 20 exhib- 

iting strong interlocking of branches with 'ohi'a. In the 6-7 cm 

small tree range, 100% had interlocking of most branches with 

both 'ohi'a and firetree seedlings become established beneath 

'ohi'a trees and then grow upwards and into the 'ohi'a crown with 

interlocking branches. This direct aggressive behavior of flre- 

tree toward 'ohi'a would appear to end in competitive replacement 

of the latter. However, in all but one firetree'ohi'a interlock 

situation, 'ohi'a was exhibiting average to good vigor. The only 

interlocking 'ohi'a with low vigor was recorded in plot 8 of 

transect C-C' . On the other hand, flretree was not faring as 

well as shown in Table 1. In Transect C-C' its vigor was from 

average to good in the < 2 cm range. However, this condition

shifted in the 2-3 cm range where 82.4% of the firetrees exhib- 

ited average vigor, but on reaching the 4-5 cm range, average 

vigor had decreased to 54.5%, and 27.3% of the trees exhibited 

poor vigor. In transect W-W' a similar relationship existed with 

poor vigor continuing to increase with diameter range. At range 

6-7 cm, 50% of the trees showed poor vigor, and at 10-11 cm the 

single tree recorded in this range had poor vigor. 

The foregoing data show that within the area considered in 

this study firetree tends to lose vigor as it increases in size. 

Cause of the vigor loss may be a lack of available soil water. 

The recent pumice soil is exceedingly dry regardless of the high 

rainfall (2700 mm) for Habitat 5. Available water for plants 

ranges only from 2% to 3%. Thus, there is less water available 

in this new volcanic material than in most sands, and plants in 

open areas will have water for growth only for a short period 

after showers (Smathers & Mueller-Dombois 1974). 

The surviving 'ohi'a trees have created a mesic microhabitat 

beneath their canopy, in comparison to the xeric soil environment 

outside the canopy. Beneath each tree an accumulating litter 

layer has formed that is periodically moistened by through- 

falling rain, and protected from desiccation by the crown cover. 

Thus the microhabitat conditions beneath 'ohi'a favors firetree 

seed germination and seedling development up to a diameter range 

of 2-3 cm. Further growth causes a drain on the soil water, and 

this condition is reflected by loss of vigor. When the firetree 

reaches the 6-11 cm range, soil water is practically unavailable 

for growth, and thus vigor becomes poor. The fact that firetree 

has never invaded the dry, barren soil of Habitat 4 tends to 

support the foregoing hypothesis. 

It is not known whether there is competition between the 

'ohi'a and firetree root systems. Surviving 'ohi'a were rooted 

in the old soil layer prior to the 1959 ash fallout layer. In 

localities where the ash fallout was over 0.5 m, a secondary root 

system developed on the buried 'ohi'a trees, thus there could be 

competition between the two trees for water and nutrients. In 

any case, 'ohi'a would still have the advantage by having a 

rooting system in the old soil, which still receives water 

filtering through the new soil. 

Quantitative Characteristics 

The basal diameter, height, density, and frequency of fire- 

trees are shown in Table 3. In transect C-C', the 2-3 cm range 

had the highest density, frequency, and second highest percentage 

cover, thus showing its dominance in the community structure 

(stratification). In transect W-W', the 2 cm class had the 

highest density (1.64/100 m2) and frequency, but the 2-3 cm and 

4-5 cm ranges had the same frequency and also the highest per- 

centage cover (3.88 and 5.72, respectively). The higher total 

density, frequency, and percent cover of the W-W' transect than 

the C-C' transect is because 'ohi'a trees are spaced farther 

apart in the C-C' transect than in the W-W' transect. Therefore,

there were more available microhabitats.('ohi'a crowns) in tran- 

sect W-W' for firetree to colonize. 

Phenological Character istics 

Phenological characteristics of this firetree population are 

shown in Table 2. The following information on flowering and 

fruiting was obtained in August 1977. Most noticeable was the 

senescence of staminate flowers and green fruits developing on 

several plants. Also, considerable defoliation was occurring on 

the branches of three plants that bore both staminate flowers and 

fruits. Approximately one-fourth of the fruits observed were 

purple, which indicated they were ripe. With the exception of 

defoliation, the flowering and fruiting cycle of firetree in 

Hawai'i probably approximates that in the plant's native habitat. 

In Madeira and the Canary Islands, Krauss (1964) observed fire- 

trees in June with an abundance of male flowers and green fruits. 

However, he noted that the staminate flowers were drying up. 

From July to September he reported that most of the green fruits 

had turned purple, and by November there were many ripe fruits 

with some on the ground. 

Flowering and fruiting starts with the Devastation Area fire- 

trees in the 2-3 cm range. Of the trees in transect C-C' 17.6% 

had staminate flowers or fruits, while in transect W-W' 16.7% had 

staminate flowers or fruits. The percentage of trees with fruits 

increased with height and diameter size. At the 4-5 cm range, 

53.8% of the trees had staminate flowers or fruits (both C-C' and 

W-W' combined). At the 6-7 cm range 100% of the trees had stami- 

nate flowers or fruits. There were no seedlings beneath those 

trees that bore fruit, even though the ground beneath some trees 

was covered with fruits. Overall, 24.0% of the trees had fruits: 

4.2% with immature fruits, 6.3% with mature fruits, and 13.5% 

with both mature and immature fruits. 

Thus, firetrees begin to produce seed at an early age, and 

the seed crop continues to increase as the stand gets older. 

Therefore, numerous seeds are available for stand regeneration 

and dispersal. This type of accelerated productivity, associated 

with horde invasion, tends to characterize some pioneering 

species, more so than a long-term colonizer. It is not known why 

defoliation of terminal branches of three trees occurred after 

producing flowers and fruits. These trees were relatively large 

ranging from 3.95-5.67 cm in diameter and 3.1-3.9 m in height. 

Two exhibited poor vigor and one average vigor. Their condition 

may have resulted from stress brought on by lack of available 

soil water. It could also indicate a response of conserving 

energy for fruit development. 

The means of dispersing firetree throughout Habitat 5 is 

still unknown. As previously pointed out, it is not likely that 

the large number of seedlings were offspring from the small num- 

ber of trees capable of producing mature fruit. Also, there was

no invasion pattern characterized by a sequential trend of diam- 

eter ranges along a directional gradient. The random distribu- 

tion of diameter sizes suggests a dispersal agent that would be 

following a similar pattern. In addition, the fact that 93% of 

the firetrees were established underneath 'ohi'a crowns, suggests 

a strong correlation with the distributional pattern other than 

just microhabitat conditions for seed germination. 

There is good reason to believe that birds are involved in 

the dispersal of firetree seed. While collecting field data, the 

investigators observed numerous Japanese White-eye (Zosterops 

'aponica) foraging in the 'ohi'a trees. The White-eye is an 

2xotic bird in Hawai'i. It is native to Japan, and it has been 

observed to feed on insects, nectar, and fruits in Hawai'i (Guest 

1973). In Australia, Gannon (1936) reported that White-eye 

spread blackberry, lantana, and several other species of plants. 

It seems logical that as Japanese White-eye forage among the 

'ohi'a flowers for nectar or insects, they could be depositing 

firetree seeds obtained from trees outside the Devastation Area. 

The presence of numerous firetree seedling beneath 'ohi'a crowns 

and oftentimes close to the trunk, tends to support the assump- 

tion that seeds are being deposited by birds. Another exotic 

bird that is common to the area, and may also be capable of 

spreading firetree is the Red-billed Leiothrix (Leiothrix lutea) . 

Growth Form Characteristics 

The growth form characteristics are shown in Table 1. Fire- 

trees in the 2-3 cm and 4-5 cm ranges exhibited a high degree of 

basitonic branching (multiple branching near base of main stem). 

It was not determined what produces this type of adventitious 

budding. It seems likely that the basitonic branching is in 

response to a stressful condition. For example, one specimen 

found in the 1974 survey of the Devastation Area (Smathers 1976) 

is believed to have survived the 1959 ash fallout. This tree had 

a stem 10 cm in diameter that appeared to have been burned off by 

the hot falling ash. It was approximately 10 cm below the 1959 

ash level, underneath and close to the trunk of a surviving 

'ohi'a tree, and with numerous branches sprouted from the burned 

stump. These branches had been unable to penetrate the dense- 

basal branches of the surviving 'ohi'a, and thus they had grown 

outward, prostrate on the ground, beyond the periphery of the 

crown and then upward. This growth response could indicate fire- 

tree's low shade tolerance under high crown densities. A similar 

condition was also observed in transect C-C' and W-W' where a 

majority of firetrees grew into 'ohi'a with open crowns; however, 

where the basal canopy was dense, they grew outward, prostrate on 

the ground.

CONCLUSIONS 

The Devastation Study Area provides a unique opportunity to 

study firetree ecology. Here, a self-contained population can be 

eventually studied under six different habitat conditions that 

range from rain to seasonal dry forests types. 

The initial phase of the present study has provided here- 

tofore unknown ecological information on firetree in Hawai'i. 

Although several factors have been evaluated, the results are 

preliminary. Additional investigation is needed before defin- 

itive conclusions can be made. Notwithstanding, the present 

results show that the firetree population is not competitively 

replacing 'ohi'a trees nor any other native vascular plant. On 

the contrary, firetree shows a decided loss in vigor as it 

develops into tree size, apparently a function of low avail- 

ability of soil water for an increasing biomass. However, the 

close interlocking of 'ohi'a and firetree crowns has a threat- 

ening characteristic which must be further evaluated. 

To evaluate the apparent close, physical, competitive rela- 

tionship between 'ohi'a and firetree will require long-term 

observations on a permanent site. Data derived over an extended 

period of time will reveal whether firetree can competitively 

replace 'ohi'a, and in addition whether firetree can regenerate 

itself on the same site. 

Kawasaki's (unpublished) observation that firetree forms a 

dense-closed canopy forest with nothing growing beneath, seems to 

indicate that it is a shade intolerant species. In addition, it 

must be determined whether firetree can recolonize where it was 

previously eradicated by herbicides or natural succession. 

Although the present location of firetree correlates with 

the foraging pattern of fruit-eating birds, primarily the 

Japanese White-eye, it cannot be definitely stated that this is 

the dispersal agency. Considerable observations, seed viability, 

and germination study will be needed to test this hypothesis. 

It is imperative that firetree be observed in its native 

habitats (Azores, Madeira, and Canary Islands). This would pro- 

vide a better understanding of its potential ecological role in 

the various ecosystems of Hawai'i. Now that firetree has become 

naturalized in Hawai'i, as have hundreds of other exotics, it 

seems that the prudent course of action would be to learn as much 

as possible about how it fits into the new vegetation. Knowledge 

of this type w i l l provide a better understanding of what con- 

trols, if any can be effective in eliminating or stabilizing 

firetree populations. This viewpoint is shared by some of the 

foremost ecologists who have studied the nature of exotic 

invasions (Elton 1977).


Elton, C. S. 1977. The ecology of invasions by animals and 

plants. Chapman and Hall, London. 181 pp. 

Gannon, G. R. 1936. Plants spread by the silvereye. Emu 35: 

314-316. 

Guest, S. J. 1973. A reproductive biology and natural history 

of the Japanese white-eye (Zostero s a onica a onica) in 

urban Oahu. Island Ecosystems *,%h* IRP 

29. 

95 PP. 

Krauss, N. L. H. 1964. Insects associated with firebush (Myrica 

Aiton). Proc. Haw. Entomol. Soc. 18(3): 405-411. 

Neal, M. C. 1965. In gardens of Hawaii. B. P. Bishop Museum 

Special Publication 50, Bishop Museum Press, Honolulu. 

924 pp. 

Smathers, G. A. 1976. Fifteen years of vegetation invasion and 

recovery after a volcanic eruption in Hawaii. Pages 207-211 

- 

in C. W. Smith, ed. Proceedings, First Conf. in Natural 

Science, Hawaii Volcanoes National Park. CPSU/UH (Univ. of 

Hawaii, Botany Dept.) 

Smathers, G. A., and D. Mueller-Dombois. 1974. Invasion and 

recovery of vegetation after a volcanic eruption in Hawaii. 

National Park Service Science Monogr. Ser. 5. 129 pp. 

Walters, G. A. 1973. Tordon 212 ineffective in killing firetree 

in Hawaii. USDA Forest Service Res. Note PSW-284. 

Walters, G. A., and W. S. Null. 1970. controlling firetree in 

Hawaii by injection of Tordon 22k. USDA Forest Service Res. 

- ~ 

Note PSW-217.

TABLE 1. Habitat 5. Structure and vigor of firetree pplation. 

- ~ 

Transect: CX', 18 plots, 10 x 10 meters. mtal cover 1800 m2 

Vigor Class 

~unter of Treed N h r 

Basal 

Diameter W r 

mnge of 

(a Trees 

mterwith 

Basitonic 

Branchilr~ 

8 of Diameter Range 

o + ++ 

(por) (average) (gocd) 

N h r 

Interlockiq 

with 'ohi'a 

Nlrmber growing 

Werneath 

'ohi'a 

Associated 

with Species 

other than 

'ohi'a 

mnter 

Growing 

in @en 

' 2 9 0 O/O 6/66.7 W33.3 2 7 0 0 

- 17 5 1b.9 2/1.2 11 6 0 

0 

4-5 11 2 3/27.3 6/54.5 2/18.2 7 3 0 1 

Subtotal 37 7 4fl0.8 26n0.3 7/18.9 20 16 0 1 

Transect: WU', 7 plots, 10 x 20 meters. mtal rover 1400 m2 

2 23 0 1/4.3 7/30.4 1V65.2 8 10 2 3 

2-3 18 4 3A6.7 10/55.6 5/27.8 14 3 1 0 

4-5 15 4 2/13.) 2/13.) 13 2 0 0 

6-7 2 0 1L50 1/50 2 0 - - . 

8-9 - - --- -- - -- - - - 

mth Transects Canbind (Sllbtotals) 25 plots, 3200 d. Total Cover 

Totals 96 15 21 46 29 58 31 3 4

mLC 2. Habitat 5. Phenolcqical characteristics of firetree population. 

Ransect: C-C', 18 plots, 10 x 10 meters. lbtal cover 1800 I$ 

Basal mature Mature Both Mature 

Diameter N&K Ruits Ruits 6 mature 

mge of Ruits aees with 

(an) Trees (8/8/77 ) Ruit (%) 

2 9 

2-3 17 

4-5 11 

Subtotal 37 

Transect: W-W', 7 plots, 10 x 20 meters. lbtal cover 1400 m2 

Both hansects Ocmbined (Subtotals) 25 plots, 3200 4. lbtal Cover 

Totals 96 4 6 13 24.0

VBLC 3. Witat 5. Wantitative characteristics of firetree ppllation. 

Ransect: C-C', 18 plots, 10 x 10 meters. Total mer 1800 & 

Subtotal 37 2.05/100m2 72.2 6.49 

subtotal 

Transect: W-W', 7 plots, 10 x 20 meters. mtal cover 1400 m2 

Both ltansects Canbind (Subtotals) 25 plots, 3200 m2. lbtal Cover

KILAUEA CALDERA 

0 

HALEMAUMAU 

- 

METERS 

RAIN FOREST-WC) 

-------_____ 

SEASONAL FOREST-cM(nr1 

1974 LAVA FLOW 

HABITAT NO. 1 

MASSIVE LAVA WlTH JOINT CRACKS 

HABITAT NO. 2 

WlTH TREE SNAGS 

HABITAT NO. 4 

PUMICE AREA WlTH TREE SNAGS 

X-WEATHER STATIONS 

RVlVlNG TREES 

FIGURE 1. Map showing location of ~ilauea 

Iki 

crater in reference to Hawaii Volcanoes 

National Park.

HABITAT PROFILE - NOS. 1.2,4,5,6 OF TRANSECT A-A' 

18.S0 C 

APPROXIMATE MEAN ANNUAL AIR TEMPERATURES AND MEAN ANNUAL RAINFALL FOR 1967 AND 1968 

16.3OC 

2203 mrn ) 3280 rnm 

KAU DESERT 

DEAD TREES (METROSIDEROS) 

SURVIVAL TREES OF 

DEEPEST BURIAL (METROSIDEROS) 

TREES WlTH 

LITTLE INJURY (METRDSIDERDS) 

SURVIVING SHRUBS WlTH 

SECONDARY ROOTS 

(VACCINIUM. ETC.) 

WEATHERED REGOSOL 

WlTH HIGH ORGANIC MATTER 

SCRUB TREES OF KAU DESERT 

(METROSIDEROS) 

ClBOTlUM TREE FERN 

SADLERIA FERN 

AERIAL ROOTS(METR051DEROS) 

HERBACEOUS VEGETATION 

(NEPHROLEPIS, ANDROPOGON. ETC.) 

i loo 

150 { 

4 200 

1 250 

METERS 

\ 

\ 

1 

KILAUEA IKI 

FIGURE 2. Habitat types and study transects of the 1959 

~ilauea Iki eruption site (crater floor and 

pyroclastic deposit). 

19.5 C 

THIS PROFILE SEGMENT APPROXIMATES 

TRANSECTS 0,c IN FIG. 7

FIGURE 3. Northeast-southwest profile of eruption 

site extending from ~Ilauea Iki crater 

to upper Ka'u Desert. 

I 

KAUAl 

STATE OF HAWAII 

160° W 

HAWAII VOLCANOES 

NATIONAL PARK 

KILAUEA IKI 

288

HALEAKALA NATIONAL PARKCRATER DISTRICT 


INTRODUCTION AND GENERAL OVERVIEW 

C. W. Smith 




The Crater District of Haleakala National Park lies between 

latitudes 20'41' and 20'46' N and longitudes 156O08' and 

156"15' W. The crater is in fact an erosional feature where two 

amphitheater-headed valleys, Kaupo and Ko'olau (Keanae), met in 

the Pu'u Mamane to Kapalaoa area. Later flows of the Hana 

volcanic series obscured all but the most obvious features of the 

erosion. 

The pre-contact Hawaiians used the crater area mainly as a 

transisland corridor later to be paved by Kihapiilani. Other 

uses, particularly religious and adze quarrying, were made of the 

area. The impact of this usage was minimal, involving minor 

disturbances of rocks and some fire-building. 

After 1788, the impact in the area remained minimal until 

the Wilkes Expedition's (1840-1841) report described the summit 

and surrounding areas in somewhat melodramatic terms. The moun- 

tain had in fact been climbed 12 years earlier in 1828 by three 

missionaries. The silverswords were the first to suffer, as 

tourists collected them to verify their ascent (Bryan 1915). The 

establishment of the Rest House and later the Silversword Inn 

resulted in an escalation of the visitor impact on the silver- 

swords. Perhaps the most unfortunate abuse was Maui's entry in 

Washington's Birthday Annual Floral Parade in 1911 in which a car 

was completely covered with large silverswords. 

Later, cattle grazed in and were driven through the crater. 

Pasture improvement was encouraged, at least in the Kaupo Gap, by 

burning the vegetation. Goats have been established in the area 

for a long time. However, before the Second World War the prob- 

lem was sufficiently acute that massive goat drives were orga- 

nized. The impact of goats in Haleakala has been well documented 

by Yocum (1967). The problem is still as acute today as it was 

in the recorded past. 

The lowest point of the Crater District is just below 4000 

feet, the highest just over 10,000 feet. The majority of the 

area is above the inversion layer and a significant percentage is 

above the diurnal frost line at 8000 feet. One might ascribe

much of the xeric scrub and open to almost absent plant commu- 

nities to extensive periods of drought. However, Whiteaker 

(pers. comm.) has evidence from climate diagrams which indicates 

that only at the Observatory is there any consistent drought 

period and then only for the month of June. The Paliku area has 

a climate diagram typical of a rain forest area. Thus other 

factors have to be identified to account for the paucity of vege- 

tation. The acute disturbances previously alluded to are 

undoubtedly part of the explanation. Alpine ecosystems are noto- 

riously slow to recuperate. The alpine edaphic factors and the 

paucity of soil in many areas are also contributing factors. It 

is my opinion that what we see today in Haleakala Crater is a 

meager remnant of the previous ecosystem. Even if the distur- 

bances are eliminated the recovery w i l l be an extremely slow 

process. 

The objective of the Resources Basic Inventory was to iden- 

tify all the plants and animals in the Crater District which 

would be presented as annotated lists. Distribution maps of 

individual species would also be produced. Resource management 

problems would also be identified and possible remedies dis- 

cussed. Finally, a detailed vegetation map and description of 

the vegetation units would be produced. 

The methodology of the Inventory was to use seven transects, 

five along a north-south axis and the other two along an approx- 

imately east-west axis. Study sites were established irregularly 

in areas along the transect which were obviously different from 

other areas. In all, 55 study sites were established. At each 

site, all species were recorded or collected, and their relative 

abundance using the Braun-Blanquet system was estimated. Obser- 

vations and collections were made in numerous other localities 

not formally sampled. 

The vegetation map and vegetation unit description program 

was conducted on a more formal basis. Potential vegetation units 

were later identifed, refined, and mapped from aerial photo- 

graphs. The unit boundaries and authenticity were verified by 

visiting each delineated area. Formal sampling areas were then 

established in each vegetation unit, the vegetation sampled and 

soil and other environmental parameters measured. 

Various aspects of these studies w i l l be presented in the 

following papers. Six areas are covered and I ask you to bear 

with the very limited scope of information presented in each 

which is necessitated by the severe time constraints of the 

Conference.


Bryan, W. A. 1915. Natural History of Hawaii, Book 1. The 

Hawaiian Gazette Co., Honolulu. 596 pp. 

Yocom, C. F. 1967. Ecology of feral goats in Haleakala 

National Park, Maui, Hawaii. Amer. Mid. Natur. 77: 418-451.



THE LICHEN FLORA 

C. W. Smith 




INTRODUCTION 

The lichens of Haleakala have received scant attention in 

the past. The only comprehensive collection prior to the 

Resources Basic Inventory was by Skottsberg during the Hawaiian 

Bog Survey in 1938. This collection was studied by A. H. 

Magnusson and forms a significant element of his Catalogue of the 

Hawaiian Lichens. A few other botanists have collected in the 

Crater including the Abbe Faurie, J. F. Rock, and 0. Degener. 

However, none of these collectors were specialists in lichens; 

their collections were incidental to their other interests, 

mostly flowering plants. 

This report is a preliminary investigation of the lichens of 

the Crater District of Haleakala National Park with notes on the 

principal lichen associations found in the area. Unfortunately, 

several groups are very inadequately understood from a taxonomic 

point of view and the omission of some of them seriously limits 

the reliability of a few of the assessments. This reservation is 

particularly true of the rock-inhabiting species at higher eleva- 

tions. A significant number of species, about 25%, still await 

determination by specialists. In the majority of instances, 

their lack of response is the result of their own current uncer- 

tainty on these Hawaiian specimens. As further informa tion 

becomes available the list w i l l be updated and the ecological 

assessments revised. 

WHAT ARE LICHENS? 

Lichens are an obligate symbiotic association between a 

fungus and an alga. The association produces a plant which is 

uniquely different from that of either the fungus or alga growing 

alone. Perhaps the single most significant ecological feature of 

lichens is that they must undergo periodic dessication. If they 

are not allowed to dry out within a three or four day period they 

become moldy and die. Thus these plants are ideally suited to 

areas where water is not continuously available. One may think 

of deserts in this respect but many situations in more equable

climates experience alternating periods of wet and dry, for exam- 

ple, rock surfaces, leaves, tree trunks, and branches and even 

the surface of soil. 

There are three basic growth forms in the lichens: 

crustose, foliose, and fruticose. Crustose species form a thin 

crust or film over the substratum. They are firmly attached to 

or embedded in the rock or bark. Foliose species lie flat on 

the substratum and are usually attached to it by hairs or 

rhizines. Foliose species can generally be separated from the 

substratum. Fruticose species are generally pendent or erect as 

in the familiar Usneas and British Soldier lichens. They are 

normally easily detached from the substratum. 

It is generally true to say that the drier an area the more 

likely you will find crustose species, the wetter an area foliose 

and fruticose species. This generalization is as true on the 

microscale as it is on the macroscale. If you look at the twigs 

on the edge of a tree or bush you w i l l normally find crustose 

species only. On the larger deeper shaded branches you w i l l 

probably find foliose or fruticose species. The complicating 

factor to this generalization in Haleakala National Park is that 

many areas are frequently inundated in clouds which encourages 

the growth of the foliose and fruticose species in situations 

that would normally only support crustose species. On the other 

hand, in areas where rainfall and fog interception result in 

infrequent dessication, mosses and liverworts replace the 

lichens. 

LICHEN ECOLOGY 

All lichens in the Hawaiian Islands are presumed to be 

native or endemic. No exotic species are known, a situation 

which is likely to change in the near future because of the 

introduction of large numbers of plants from various regions of 

the world, e.g., orchids introduced to the Foster Botanic Gardens 

frequently have live lichens associated with them; Christmas 

trees from the Pacific Northwest nearly always have lichens on 

their trunks, especially Hypo h sodes. On the other hand, 

endemic and native species p:E?A:shrted from Haleakala 

have not been located during this study. or example, the genus 

Umbilicaria is represented by three endemic species in the literature 

(Magnusson 1956). None of these species was found on the 

recent survey. Unfortunately, the type locality of one of these 

species, U. acifica, is "at the top of Halemau (sic) Trail." 

Since thg species --li as been collected from this area only, it may 

be assumed that it is now extinct or extremely rare in the area. 

The recent heavy pig impact w i l l have made the former alternative 

more probable. 

The lichen communities generally follow the flowering plant 

community distributions outlined in Whiteaker (1979) in his 

Vegetation Map of the Crater District of Haleakala National Park. 

However, two environmental variables modify the lichen community 

distributions so that they do not conform precisely to Whiteaker.

The diurnal frostline is of little significance in the dis- 

tribution of rock-inhabiting lichens though it does have an 

impact on bark-inhabiting species because of the reduced avail- 

ability of substrate. Lichens are capable of carrying out their 

life functions at much lower temperatures, even to freezing 

point, than flowering plants as long as there is sufficient mois- 

ture available. The other environmental variable, cloud inunda- 

tion, extends the distribution of foliose and fruticose lichens 

beyond their expected distribution in mesic plant communities. 

Lichens are extremely efficient at absorbing water from air and 

can become quite wet in a short period of time when submerged in 

clouds. As a consequence, their growth and abundance are in- 

creased so much that the relative cloud cover can be fairly 

accurately mapped from the abundance of the epiphytic foliose 

lichens. For example, the eastern side of the Ko'olau Gap up to 

and beyond Pu'u Mamane is more frequently immersed in cloud than 

the western or central portions of the Gap. 

Rock Communities 

The lichen communities on rock are ~robablv the least disturbed 

or altered in the study area. A few species have disappeared, 

for example, Umbilicaria pacifica, and the abundance of 

others may have been chanqed by habitat alteration. By and 

large, the-community structure is p;obably the same now as it was 

prior to the impact of western man. 

At the top of the mountain and in other areas which are pre- 

dominantly devoid of vegetation, the rocks are colonized by a 

community in which Acaros ora and Lecidea are dominant with occa- 

sional specimens o f d a , Candelar iella, and Rhizocarpon 

geographicum. Only the stable rocks and boulders are colonized; 

the loose cinder is too freauentlv disturbed bv wind and rain for 

lichens to become established. es en on the rocks the lichens are 

always in very protected situations where the microenvironmental 

conditions offer some relief from the rigorous climate of the 

area. With decreasing elevation, the lichens are found in more 

exposed situations with increasing frequency and other species, 

for example, Stereocaulon vulcani and Placopsis gelida begin to 

appear in the community. 

The lichens in this harsh environment do not grow very 

rapidly. Colony sizes are always small. The activity of lichens 

as primary colonizers in such situations is very low. Conse- 

quently, the rate at which they decompose the rock is low. Rain 

and other edaphic factors are probably more important in soil 

formation than are the lichens. At lower elevations or where 

moisture is more abundant, for example, the summit of Kuiki, the 

lichens probably play a significant role in soil formation. 

Where moisture from cloud or rainwater is more abundant, the 

lichen communities on rock are more luxuriant in terms of both 

biomass and species diversity. The species of the genus Stereo- 

caulon show an interesting series of communities which are

correlated with the amount and physical phase of the available 

water, as well as the age of the rock on which they are growing. 

Stereocaulon vulcani, the primary colonizer of most lava 

flows in Hawal'i, characteristicallv - qrows - where the annual rainfall 

is above 30 inches a year. Though one would expect to find 

it at the summit which supposedly receives this amount of rain 

each year, it is extremely rare and very poorly developed there. 

Its near absence is probably because most of the rain comes in 

two or three major kona storms each year. It is found throughout 

the rest of the Crater District but below 6000 feet its distribution 

is regulated by the growth of other organisms. The occurrence 

of this species in any appreciable quantity below 6000 feet 

is generally a good indication of the recent disturbance 

community. 

of the 

Stereocaulon octomerellum grows on large boulders in the high 

rainfall area of the eastern side of the Kaupo Gap. It is 

normally found only on well-weathered, exposed rocks. 

Stereocaulon ramulosum has an almost intermediate ecological 

position between the above two species. It favours environments 

in which cloud inundation is frequent. The height and fertility 

of the plants is indicative of the frequency of the cloud cover, 

the lower stature and infertile specimens indicating drier, 

harsher conditions. 

Litter Communities 

Where plant litter accumulates and areas where the humus 

content of the soil is high, the endemic Cladonia leiodea is dom- 

inant. The species is not tolerant of shading so it is charac- 

teristic of the open scrub communities. The luxuriance and 

colony size of the plants are an indication of the moisture 

regime of the area. The largest specimens are found in the 

wetter areas. The plant is particularly sensitive to mechanical 

disturbance and may serve as an indicator of past pig activity 

when absent from an area in which it should logically appear. 

A rather unusual litter community occurs under the dead but 

still standing leaves between Descham sia clumps. All of the 

species are very attenuated and none ___P_. 

are ertlle which is probably 

due to the suboptimal levels of light filtering down between 

the leaves. Pseudocyphellaria crocata, ~ticta weigelii, 

Peltigera golydactyla and Cladonia scabriuscula are the most 

common svecles in this situation. I know of no similar communitv a 

type adapted to such low light intensities. 

Leaf Communities 

In the gullies behind Paliku a fragmentary lichen community 

is found on the leaves of Pelea. Two species are present and 

represent the upper elevational limit of a specialized community 

normally found below 1000 feet. Their occurrence in this highly

protected environment illustrates the unusual nature of these 

gullies. 

Bark Communities 

The complex chemical nature of bark results in unique lichen 

communities on each tree or shrub species. Since chemical and 

physical surface characteristics change with the age of the bark, 

the associated lichen communities also change. Thus the lichen 

community on twigs will be different from that on the trunk. For 

example, the twigs and branches of mamane are colonized by 

Ochrolechia allescens H otrach na sinuosa, and species of 

- 

candelaria, Bue o d a n e r e a s the trunk 

has Parmelia dominicana, Pannaria rubiginosa, and Heterodermia 

speciosa. Every other tree and shrub has its own spectrum of 

species. ~onse&entlv, it is extremely difficult to describe the 

general distribition of bark-inhabiting species of lichens in any 

meaningful manner. Any attempt to do so is beyond the scope of 

this study which was not designed with this type of analysis in 

mind. 

The distribution of Usnea and Alectoria on pukiawe and 

'ohelo closelv .. ~arallels 

- 

the areas inundated bv cloud for sianificant 

periods. Usnea is found where clouds p;obably cover2 the 

area at least h a l f e days of the year whereas Alectoria smithii 

occurs in areas where the cloud cover is significantly less. 

RECOMMENDATIONS 

There are no formally designated threatened or endangered 

lichens. That does not mean that there are no rare lichens. 

Even if they were to be listed there would be very little that 

could be done to promote the species other than habitat protec- 

tion and conservation. As with many other groups studied during 

this survey, the removal of the feral herbivores would be a 

significant management action to preserve the lichen communities 

in the Park. 


Magnusson, A. H. 1956. A catalogue of the Hawaiian lichens. 

Ark. f.. Bot. 3: 223-402. 

Whiteaker, L. D. 1978. The vegetation and environment of the 

Crater District of Haleakala National Park. M.S. Thesis in 

Botany, University of Hawaii, Honolulu. 159 pp.



THE VASCULAR FLORA OF HALEAKALA 

Lani Stemmermann 




Haleakala includes one of the four alpine ecosystems in the 

Hawaiian archipelago (the other three being located on the island 

of Hawai'i). Its flora has been studied by numerous botanists 

since the first collections in this area were made by members of 

the United States Exploring Expedition in 1841. Hawaiian alpine 

ecosystems (only one of the Park's vegetation types) have been 

recognized in the past as having a high percentage of endemic 

species (Skottsberg 1931)--plants that grow nowhere else--and 

even now, despite the presence of numerous exotic species in the 

Park and the extinction of native taxa (both a result of past and 

present disturbances), the Park's flora exhibits a large number 

of endemic species (Table 1). 

The Haleakala National Park Crater District Resources Basic 

Inventory (RBI) integrated research program was undertaken to 

provide a biological inventory of the Park, and to identify 

resource management problems. When this report, currently in the 

final stages of preparation, is finished, a computer print-out 

will be available which will provide the following information. 

1) A general catalogue of the species with notes on general 

abundance and distribution within the Park, and their 

status (Indigenous, Endemic, Exotic) within the State: 

2) Inclusion of any of the species in any of the Rare and 

Endangered Species Lists (Fosberg & Herbst 1975: U. S. 

Fish & Wildlife Service 1976) or the State's noxious 

weed list (Office of Environmental Quality Control 

1972, revised 1976); 

3) Scientific, English, and Hawaiian names (when known). 

Use of the computer for storage of information facilitates up- 

dating the list as new information is gathered. 

In addition to an inventory of plants found in the Park, 

voucher specimens have been collected as reference material for 

Park personnel with some duplicate materials to be distributed 

to the Bishop Museum (BISH) and University of Hawaii (HAW) 

herbaria.

Thelast comprehensive review of the Park's flora was 

prepared by Mitchell (1945). Ruhle (1959 revised 1968, 1975) 

presented a good review of the natural history, but gathered no 

new information on the Park's flora or vegetation. Our studies 

during the years 1975-1978 have resulted in the listing of many 

more species for the Park than previously known, although certain 

genera, for which Mitchell listed numerous species and subspecific 

taxa, are not so finely circumscribed. For instance, 

17 taxa were recorded by Mitchell for the genus Railliardia, 

while far fewer are considered in our recent compilation. In 

such instances this is due to a more conservative interpretation 

of taxonomically difficult qroups, rather than extinction. 

Unfortunately certain taxa, such as-~lermontia haleakalensis, are 

probably extinct, while others, such as ~illebra-~anunculus. 

- - Previouslv reoorted from the Crater reqion. - 

confined to ~i~ahilu valley and Ko'olau Gap. 

a r e w 

Over the 30 years between Mitchell's study and the present 

one, there has been little change in the flora of the Park, but 

some of those changes are well worth mentioning. Several species 

which have the potential of being aggressive weeds were not noted 

by Mitchell (1945). while some of these may have been overlooked 

(a problem all too familiar to anyone who has attempted to com- 

pile species lists), others are no doubt relatively recent intro- 

ductions. These, and others which were previously reported from 

the Park and should be considered problematic, are listed in 

Table 2. Some of these species are officially considered 

"noxious weeds" (as indicated), but many are not, and only a few 

are presently under Park management. At present, a number of 

species with localized populations should be contained to prevent 

their spread throughout the Park. Table 2 does not include. all 

the weedy species in the Park, but only those which are thought 

to be most in need of control and not those already hopelessly 

out of control. 

A few rare or new species not previously noted from the Park 

In conclusion, certain taxa present in the Park are endemic 

not only to Hawaiian alpine and subalpine ecosystems, but to 

Haleakala proper such as Artemisia mauiense, Argyroxiphium macrocephalum, 

Stenogyne crenata (2 varieties), Geranium cuneatum, 

- G. arboreum, Santalum haleakalae, and others. The deleterious 

effects of feral qoats - on certain veaetation < tvDes - within the 

Park (especially in areas more-or-less inaccessible to hunters) 

cannot be overemphasized. Since many of the endemic species are 

of limited distribution, they should be considered threatened by 

the continued presence of goats. Delays in the implementation of 

an effective goat control program must be considered a serious 

threat to the native biological resources of Haleakala National 

Park .


Degener, 0. 1930. Fe rns and flowering plants of Hawaii National 

Park. Honolulu S tar-Bulletin Ltd., Honolulu. 

Fosberg, F. R., and D. Herbst. 1975. Rare and endangered 

species of Hawaiian vascular plants. Allertonia 1: 1-72. 

Mitchell, A. L. 1945. Checklist of higher flowering plants, 

grasses, sedges, rushes and ferns of the Haleakala Section, 

Hawaii National Park. In-house Document, Haleakala National 

Park . 

Office of Environmental Quality Control, Environmental Center, 

University of Hawaii. 1972 (revised 1976). Hawaii environ- 

mental laws and regulation. Vol. 11, A: PI: 33-37; A: PI: 

43-47. 

Ruhle, G. 1959 (revised 1968, 1975). A guide to the crater 

area of Haleakala National Park. Hawaii Natural History 

Association Publication, Hawaii. 

Skottsberg, C. 1931. Remarks on the flora of the high Hawaiian 

volcanoes. Goteborges Botaniska Tragard. 6: 47-65. 

U. S. Fish and Wildlife Service. 1976. Endangered and 

threatened species: Plants. Federal Register 41(117): 

24524-24572.

TABLE 1. Percentages of the indigenous, endemic, and exotic vas- 

cular plant flora of Haleakala National Park Crater 

District. 

Pteridophytes Monocots Dicots Total 

Indigenous 44 13 2 11 

Endemic 

Exotic

TABLE 2. Potentially aggressive weeds in Haleakala National Park Crater District. 

Species Status Distribution in Park 

Cirsiun vulgare (Savi) Tenore 1,2 Present especially in heavily goat infested areas. 

Eupatoriun denophorun Spreng. *,2,4 Maui pamakani is found in several large ppula- 

tions at mid-elevations in the Park such as at 

kipuka S of Laie cave, and S of Hanakauhi. It 

probably is no longer spreading but nevertheless 

should be controlled. 

Eupatoriun ripariun Spreng. *,2,4 This species is recorded £ran the Homer Grove 

area, and while ptentially dangerous in lower 

elevations it is probably not a threat in the Park 

but should be watched. 

Lantana Camara L. I~OLKJ~ probably not a problem or even a ptential 

problem above 5000 feet in the Park, the presence 

of Iantana at low elevations in Kaup Gap within 

the Park should be watched carefully and controlled 

when practical. 

Cpuntia megacantha Salm-Dyck l?, Panini is presently known from only t mall 

3,4 populations in mid-lower Kaupo Gap, and near hie 

Cave. It is probably under control' within the 

Park.


Species Status Distribution in Park 

Passiflora subpeltata Ortega 2,4 A few'plants were noted along Kaup Trail, and 

while the populations are currentlv not a problem 

any detect2 $ants of this genus should b;! con- 

trolled. 

~ennisetun clandestinun Hochst. 2,4 Kikuyugrass is found along trails and roads 

ex Chiov. throughout the Park, and at low elevations along 

Kaup Trail it is the dcminant cover species. 

- Pinus spp. 

Poa gratensis L. 

Ricinus communis L. 

- Rubus penetrans Bailey 

1,2, Certain pines and other gymnosperms have spread 

3? frcm their planting sites, notably near Homer 

Grove. Aggressive species should be controlled. 

2,4 The Kentucky bluegrass is common in danp areas 

throughout the Park, such as under trees. It is 

replacing the native Deschanpsia grassland in 

Kaluanui, and work should be done to f id a- means 

of controlling its spread. 

2,4 A large community of castor bean is found along 

and to the east of the Kaup Trail near the bound- 

ary of the Park. 

*,2,4 The prickly Florida blackberry has becane a nui- 

sance in the Paliku Horse Pasture, could spread 

elsewhere in the Park. It is high on the list of 

species needing inmediate control.

- Rubus rosaefolius Sn. 

2 !l%e thimbleberry is found occasionally in lower 

east Kaupo Gap in danp shaded areas; though perhaps 

not in danger of spreading its populations 

should be watched. 

Schinus terebinthifolius Raddi 2,4 A single specimen of Christmas berry was seen in 

the goat-ridden western part of Kaupo Gap; elevation 

ca. 4400 feet. It should be removed. 

- Ulex europaeus L. 

*,1,4 Cegener (1930) reprts that gorse was planted as a 

hedge in Olinda to contain sheep but within a 

decade of it being planted it had become a pest. 

~t one time territorial prisoners were employed to 

eradicate this plant. Within the Park it is only 

known from a mall patch below Park Headquarters. 

* Included in the State's noxious weed list 

1 Currently being controlled by Park personnel or volunteer groups 

2 Species not presently controlled which are in need of control by the Park 

3 Biocontrol agents available but apparently not presently effective 

4 Not included in Mitchell's species list 

Note: those species thought to be "hopelessly out of control" in the Park, or at least have 

probably reached their greatest distribution with little probability of control are 

not listed but include Anthoxanthm, Caryophyllaceae spp., Dactylis, Heterotheca, 

~olcus, Hypchaeris, Lapsana, - Emex acetosella, etc.

THE ACQUISITION OF NATURAL AREAS IN HAWAI'I 

Kim0 Tabor 

The Nature Conservancy 

Honolulu, Hawaii 

The Nature Conservancy's acquisition policy in Hawai'i has 

been shaped by several factors. 

Expert advice from knowledgeable individuals who know 

Hawai'i's uniqueness in specific areas of biological importance 

is critical to selection of natural areas, a process which is 

exceedingly intricate. It is desirable to acquire a piece of 

property of viable ecological significance after the finanacial 

and legal tangles are resolved. 

Acquisition of a general type of system rather than a spe- 

cific type of biota was the primary consideration in the acqui- 

sition of Maulua Nui. Once the preliminary selection was made 

from some 40 alternatives, a party of three people did a brief 

field reconnaissance of the property. Steven Montgomery, for 

many years scientific assistant to the Natural Area Reserve 

Systems Commission, and James Jacobi, a botanist and student of 

Dr. D. Mueller-Dombois with wide experience on the island of 

Hawai'i, provided a quick scientific assessment of the Maulua 

property during a three-day field trip in early January 1977 

(covering a distance of 11 miles, or 18.2 km). 

The location of Maulua Nui on the slope of Mauna Kea pro- 

vides a classic Hawaiian ahupua'a land form. It faces northeast 

against the tradewinds, from sea level to an elevation (at the 

boundary) 1 mile high on Hawai'i's highest mountain, on the 

State's largest land mass, the island of Hawai'i. This area is 

accessible to birds and seeds from the North American Continent 

given the wind patterns of perceived history. It is an area 

which has a high rainfall and sunshine ratio thus highly desir- 

able from the high growth rates possible; an attraction also to 

the forest industry. The mouth of the ahupua'a is a large fault 

valley 21 miles northeast of Hilo. The emerging valley stream 

provides an interesting, though disturbed, estuarine area. It is 

also an area of some local historic interest. 

Maulua Nui is a unique addition to the inventory of Nature 

Conservancy lands nationwide. It provided forest habitat for 

endangered forest birds of the native passerines and the Hawaiian 

Hawk, or '10.

Land availability is necessary to acquisition. An unwilling 

seller or an exorbitant price would prevent a transaction from 

occurring. An additional consideration born from the experience 

of the Kipahulu, Maui, acquisition of The Nature Conservancy is 

the desirability of getting firm, total, fee simple title for the 

monies expended. In this case the title was clearly established 

by one of the Kingdom of Hawai'i's most astute legal minds. The 

Nature Conservancy was fortunate in finding an extended family, 

rapidly changing its priorities, which was anxious to convert 

their real property into a liquid asset while maintaining the 

traditional land form of the ahupua'a. Integrating ownership of 

the ahupua'a with corollary emotional attachments to the family 

history was a factor also. From this point of view, The Nature 

Conservancy was the ideal vehicle as economic agricultural lands 

were not of interest. Continuing revenue and a family memorial 

Of sorts was created while providing the family with some liquid- 

ity to assist the changing, and sometimes conflicting, priorities 

within the extended family. 

Remoteness of the property and the conservation zoning made 

the property suitable for one other use--forestry. Forestry pro- 

vides a slow financial return, an eight-year harvest cycle in the 

most ambitious forecast, and is one which has extant political 

and biological problems given the intensive exotic cultivation 

intended. A general economic slowdown also encouraged a 

reasonable price. 

Maintenance of the ecosystem at low cost is a point of 

debate. The issues are feral pigs and people. Feral pigs are an 

enormous nuisance to the botanical integrity of an area as they 

are a biological plow complete with seeder. People from a 

consumer-oriented expendable economy and their transport systems 

resemble mechanical plows with not-so-degradable littering 

systems. While remoteness encourages the pigs, it discourages 

the people, in a qualified sense. Maintenance costs are there- 

fore subjective and are dependent upon the ultimate use or 

pressure on the land itself. 

Problems in acquisition were highly specific occurrences of 

some generally discussed situations. In the acquisition of 

Maulua Nui, orchestrating 15 different individuals in three 

different families through more-or-less uniform sets of legal 

documents including an Offer to Option, Option, Conservation 

Easement, Subdivision Applications, waivers for survey, access 

improvment, and water system; and deeds on the resulting three 

subdivided parcels (which are subject to reconsolidation) ; the 

swap of fractional interests between parcels and family members, 

plus the negotiation for, acquisition of, another parcel used in 

barter for a major percentage interest in two of the Maulua par- 

cels, directly involved at least 50 people, excluding the 12 

attorneys representing their clients. Needless to say scheduled 

closing was delayed.

For many reasons The Nature Conservancy needed fee simple 

title for expenditure of the donors' money. It was necessary 

therefore to subdivide the property to receive title to a spe- 

cific area representing the sum of that donation which inciden- 

tally carried with it a conservation easement on the boundary. 

The remainder of the ahupua'a was to be purchased on an "offer 

to-option" basis for eventual reintegration of the ahupua'a with- 

out unduly extending The Nature Conservancy. To accomplish this, 

it was necessary to subdivide the property in simultaneous 

proceedings at State and County level. Each proceeding was con- 

tingent upon the other. 

Sociological considerations were similarly important. The 

most immediate reason was to ensure the rapid success of sub- 

division necessary to the transaction. The other premise was 

that The Nature Conservancy was to be a long-term member of the 

Hawaiian community rather than a mainland mentor of local values. 

Consequently, members of the community closest to the property 

were informed of the transaction and questions poised were 

directly and sincerely answered. Community leaders and heads of 

pertinent special interest groups were similarly informed largely 

through mutual acquaintances. 

The Nature Conservancy is a private organization working 

with private landowners. Many other organizations also acquire 

Natural Areas with a view to preserving these. In a small geo- 

graphical area such as Hawai' i, where competing interests vie 

with intensity, the size and necessity of natural area acqui- 

sition will become an ever increasing question. The question of 

redundancy appears valid to the layman. The response to the 

criticism is complicated by the explanation of island by island 

variations of biota, varying State of Hawaii and Federal Govern- 

ment (Department of Interior, National Parks, Fish & Wildlife 

Service, U. S. Forest Service, Department of Agriculture) acqui- 

sition criteria and responsibilities. Currently these considera- 

tions are handled within the community on a consensus basis. The 

quality of information in each agency's decision-making process 

varies. The need for similarly evaluated information and stan- 

dard criteria is apparent. Some areas designated as "Natural 

Areas" may be too large, too small, or repetitive in maintaining 

specific ecosystems. Some land may have been designated a nat- 

ural area about which no one knows anything specific that w i l l 

prove useful to comparative analysis. 

The solution is for specific information of uniform criteria 

for the entire State of Hawaii to be housed in a single location 

by type of ecosystem, and by specific location. The Nature 

Conservancy pioneered the rationalization of such information and 

has continued to encourage State and Federal governments to 

introduce this type of information system so that planning 

departments of highways and other civil works projects can avoid 

impact on sensitive areas, avoiding the ' snail darter' syndrome 

and concurrent economic waste, while preserving the remnants of 

complex ecosystems. Such an information system builds by cata- 

loging the existing information, then begins to fill voids in 

geographical and biological information through field studies.

Conf irmatlon of existing information validates previously col- 

lected information, providing all users firm, dependable "intel- 

ligence" which, with one reference location, saves time and 

enormous frustration. 

Additional Natural Area acquisitions w i l l require this type 

of well-conceived identification and documentation process to 

enhance the case for acquisition. 

The Fish and Wildlife Service of the Department of Interior 

has already implemented a corollary program through its forest 

bird survey administered by Mike Scott, and the State of Hawaii 

has a similar program in its 'Alala (Hawaiian Crow) survey. The 

Maulua acquisition itself was influenced by information generated 

in a logical consistent manner by the forest bird survey, as w i l l 

additional Nature Conservancy acquisitions. A shift to pre- 

serving wildlife systems will become the responsibility of The 

Department of Interior if Senate Bill 1820 or the equivalent 

House Bill passes during 1978 or 1979. These bills provide for a 

National Heritage Program, identifying not only flora and fauna, 

but also within the same information matrix, cultural and prehis- 

toric sites of significance. Each State w i l l be responsible for 

adhering to the criteria to qualify for Federal funds. 

Locally, the State of Hawaii and the County of Hawaii recog- 

nize adaptation of their codes to reflect subdivision for non- 

economic uses may enhance the values which are so frequently sold 

as Hawai'i. Areas may be left without visible habitation.

STUDIES OF LEPTOSPIROSIS IN NATURAL HOST POPULATIONS: 

I. SMALL MAMMALS OF WAIPI'O VALLEY, ISLAND OF HAWAI'I* 

P. Quentin Tomich 

Research Unit, State of Hawaii 

Department of Health, Honokaa 

The small Indian mongoose (Herpestes auropunctatus), 

Carnivora: Viverridae, and the roof rat (Rattus rattus) and 

Polynesian rat (Rattus exulans), both Rodentia: Muridae, are 

abundant in Waipi'o Valley, island of Hawai'i. Two other murid 

- - 

rodents, the house mouse (Mus musculus) and the Norway rat 

(Rattus norvegicus), are 

carriers of serotypes of 

sporadic or rare in occurrence. As 

bacterial leptospires (Leptospira), 

which are transmissable to man, this assemblaqe of alien mammals 

is of public health significance and numerous- cases of leptospirosis 

have been traced to the valley. Population density of 

the mongoose was estimated at 2.3 per acre; for rats it fluctuated 

seasonally from 1 to 11 per acre. The serotypes 

- L. icterohemorrhaqiae and L. sejroe were found in the mongoose in 

a 40:60 ratio. Of 33 house mice tested, L. ballum was isolated 

from 21 and L. icterohemorrhaqiae from 2. One isolation of 

- 

L. icterohemorrhagiae was made from d Norway rats examined. For 

126 roof rats tested, 68% of adults and 26% of young were infected; 

and for 175 Polynesian rats, 34% of adults and 26% of 

young were infected. L. icterohemorrhagiae made up 95% and 

- 

L. ballum the remaining 5%-of infections in the roof rat. For 

the Polynesian rat the ratio was 75:25. Free-ranging rats under 

observation for as long as 8 months acquired or lost infections. 

The wet subtropical climate of Waipi'o Valley supports conditions 

for transmission of leptospirosis even in times of drought. No 

prominent differences were observed in the infection rates in the 

lower valley at 30 feet above sea level and 1.7 miles inland at 

120 feet. In the forested watershed of the valley rim at 3000 

feet, conditions of infection matched closely those on the valley 

floor. Tests of 152 water samples from streams, ponds, and taro 

paddies resulted in isolations only of saprophytic leptospires. 

* Abstract

A NECROPSY PROCEDURE FOR SAMPLING DISEASE 

IN WILD BIRD POPULATIONS* 

Charles van Riper 111, and Sandra G. van Riper 

Cooperative National Park Resources Studies Unit 



INTRODUCTION 

When the demography of wild birds is analyzed, disease is an 

important but often overlooked factor. Although disease can be a 

primary factor of population regulation, its overall importance 

is probably more closely related to increasing the susceptibility 

of the host to other mortality factors (Kennedy 1975; van Riper, 

in prep.). It is therefore important that researchers be able to 

determine levels of parasites and diseases, if they are to draw 

meaningful conclusions concerning demographic parameters of a 

host population. Our purpose is to outline a procedure which 

would enable an ornithologist, who does not have sophisticated 

laboratory facilities, to examine birds correctly and to find 

answers concerning diseases present within an avian population. 

For accurate disease diagnosis it is first necessary to 

establish a definite postmortem sequence, so that each animal is 

examined in a similar manner and data are organized and easily 

retrievable. Ornithologists often feel limited in their ability 

to understand and diagnose diseases, and in many instances pa- 

thologists are not readily available for consultation. Further- 

more, budgetary constraints frequently l i m i t the number of 

specimens that can be sent out for diagnosis; of those that are, 

the time lag before obtaining results is often considerable. It 

is therefore important that workers be able to perform their own 

diagnosis, and to do this the development of a necropsy form 

applicable to wild bird populations is essential. 

The majority of avian necropsy techniques available today 

have been developed for poultry (e.g., Hungerford 1969; Zander 

1975). Those designed specifically for other species usually 

place emphasis upon caged birds (Keymer 1961; Arnall & Keymer 

1975), in particular canaries (Serinus sp.) and the Budgerigar 

(Melopsittacus undulatus) (Stone 1969). Many necropsy procedures 

are geared for veterinarian use and consist of pages with only 

general headings after which findings are placed (Ensley et al. 

1976; Carpenter, pers. comm.). 

* This is a prepublication; anyone wishing to reference the mate- 

rial herein, should first contact the authors.

The necropsy technique discussed herein was developed for 

small passerine birds, but with slight modifications can be 

applied to most avian groups. It is based on described disorders 

present in poultry (Hofstad et al. l972), pet and caged birds 

(Petrak 1969; Arnall & Keymer 1975), and wild birds (Davis et al. 

1971), and should account for most diseases commonly encountered 

in birds from the field. Included is a checklist of potential 

symptoms interspersed with dissection directions (Part I), sup- 

plemented with detailed instructions on which parts of a bird to 

save when a symptom is encoutered (Part 11). Short sections are 

presented which (1) outline the materials and facilities neces- 

sary to carry out a postmortem analysis; (2) give general han- 

dling techniques which should be used during postmortem analysis; 

and (3) give detailed instructions on how to prepare different 

materials which are to be sent to laboratories for diagnosis. 

Diseases and the avian orders in which they have been reported 

are summarized in tables. Following the procedures outlined in 

this paper, most ornithologists should now be able to perform 

their own postmortem analyses. 

Required Materials 


The basic equipment required for this postmortem technique 

is minimal. It is, however, very important that the working area 

have limited access so as to reduce bio-hazards. Safety is an 

important consideration in doing postmortem analysis of any avian 

species, because many organisms that cause disease in birds are 

also pathogenic to man. Use standard procedures in handling 

diseased tissue and liberal amounts of a strong detergent (e.g., 

Tincture Green Soap) and disinfectant (e.g., Phenol or Pine Oil). 

Both a dissecting and compound microscope are necessary. 

Necropsy of small birds is tedious, and unless fine instruments 

are used much information can be lost. Opthalmic tools are 

ideal, and we have found iris microdissecting scissors, watch- 

makers and microdissection forceps, as well as microprobes 

invaluable. Other essential equipment should include a small 

piece of glass for examining the gastrointestinal tract, clean 

microscope slides and cover slips, sterile swabs and syringes, 

sterile petri dishes and vials for collecting tissue samples, 

sterile plastic bags for freezing tissue, and an alcohol lamp. 

Required chemicals and solutions for processing necropsy 

material include: 10% buffered formalin (add a pinch of CaC03 

per gallon); 70% alcohol glycerine-alcohol (90 parts 70% ethyl 

alcohol, 10% parts glycerine); F.A.A. (50 parts 95% ethyl 

alcohol, 10 parts commercial formalin, 2 parts glacial acetic 

acid, 40 parts distilled water); absolute methyl alcohol; sterile 

transport medium for fungi (e.g., Sabouraud's agar available from 

Difco Laboratories, Detroit, Michigan 48201; or Mycotic media 

available from Baltimore Biological Laboratory, Inc., BioQuest

Division, P. 0. Bos 243, Cockeysville, Maryland 20030); sterile 

transport medium for bacteria (e.g., Stuart's medium, a modified 

form packaged with a sterile swab available from Culturette, 

American Hospital Supply Corp., McGraw Park, Illinois 60085); and 

dry ice. Optional, but often extremely useful supplies include: 

filters, Lugol's solution (5 g iodine, 10 g potassium iodide, 

100 ml distilled water; dilute with 5 times the distilled water 

before use); Hoyer's mounting medium (30 g gum arabic, 50 ml dis- 

tilled water, 20 ml glycerol, 200 gm chloral hydrate; mix in 

order listed and filter through fine gauze); 10% solution of 

potassium hydroxide or 20% solution of sodium hydroxide; sterile 

transport medium for viruses (available from Colab Laboratories, 

Chicago Heights, Illinois 60412). 

Postmortem Methods 

General handling of the bird. A necropsy should be per- 

formed as soon as the bird is received because decomposition of 

internal organs is rapid and postmortem migration of parasites 

might occur. Take measurements immediately because weight, in 

particular, will change. Size measurements are important for 

aging purposes and may later prove useful as indicators of 

specific diseases within the population. Feather wear, cloaca1 

protuberance, and brood patch w i l l better define the breeding 

condition of the specimen. Tag and label the bird, and every 

sample taken from this animal should have the same necropsy 

number (recorded on the necropsy form); indicate if samples are 

"sterile" or "non-sterile" and the type of medium in which it is 

preserved. Obtain a detailed history of the specimen. 

Preparation and examination of smears. Several types of 

smears are useful in the diagnosis of disease. Direct micro- 

scopic examination (such as fecal material) is important because 

some organisms are much more readily detected when alive. By 

using Lugol ' s solution, fungal hyphae and protozoa become more 

visible. Impression smears of organs or exudate prepared for 

gram or Ziehl-Neelsen stain (fix by drying over heat) are impor- 

tant for laboratory analysis of bacteria. Blood smears and 

impression smears of organs stained with Glemsa are essential 

when searching for blood haematozoa; fix for 30 seconds in abso- 

lute methyl alcohol. 

Collection of blood serum. Serological tests w i l l require 

blood serum. Collect blood aseptically from the heart ,and let 

clot overnight. Centrifuge for 10 minutes and then transfer 

serum to sterile vials. Refrigerate or freeze for shipment. 

Preparation of tissue samples. There are a variety of fix- 

atives used in preparing tissue (e.g., Zenker's is useful for all 

except nervous system tissue), but a good general fixative for 

most histopathological work is 10% buffered formalin. Cut tissue 

samples in pieces no larger than 1 x 2 x 0.5 cm and place in

10 volume equivalents of formalin. After 24 hours the tissue can 

be packed with less formalin or left as is. When freezing tis- 

sue, use dry ice to rapidly lower the temperature below 60°C. 

Glass may shatter so plastic bags are best for samples. Ship in 

a styrofoam container with dry ice. 

Preparation of cultures. In general, collect samples to be 

cultured before the intestine is open. If an organ has been col- 

lected under nonsterile conditions, sear the surface with a hot 

spatula, incise tissue, and sample the cut surface. Sample moist 

membranes and soft organs with sterile swabs. Place the entire 

swab directly in medium for shipment. Collect joint or nasal 

exudate with a sterile hypodermic needle or swab. If solid agar 

is used, place the tissue firmly against agar or embed several 

small pieces in agar. 

Fixation of helminths. Nematodes can be fixed directly in 

glycerine-alcohol solution and shipped. Cestodes, trematodes, 

and acanthocephalans should be placed in F.A.A. for 24 hours and 

then transferid to 70% alcohol before shipment. 

Fixation of arthropods. Mites, lice, and small insects may 

be placed directly in 70% alcohol; larger organisms (such as 

fleas) may be killed first by placinq - in steamins - water and then 

transfering to 70% a1cohoi.- For permanent mounts of ectoparasites, 

drop the specimen directly into Hoyer's mounting 

medium, pass over a flame to relax specimen, and then cover with 

a cover slip. Fungal hyphae should be mounted in 10% potassium 

hydroxide or 20% sodium hydroxide and heated gently to clear the 

specimen. 

Sporulation of Coccidian Oocysts. A fecal suspension should 

be placed, with a thin layer of 1% formalin, in a petri dish for 

one to four days so that sporulation w i l l occur and specles can 

be identified. The fecal material may be shipped in this medium. 

Laboratory facilities. A problem often occurs in trying to 

find a laboratory which will process the tissue and identify 

pathogens. There are several aqencies that have well-established 

Taboratories which specialize in avian diseases (e.g., Fish & 

Wildlife Service), but they are usually hesitant to accept 

material from independent researchers due to lack of time or per- 

sonnel to process material. Better places to try include the 

Department of Agriculture, Department of Health, Veterinary 

Pathology laboratories, and university laboratories in Medical or 

Veterinary Sciences. Check with the particular laboratory for 

their preferences of tissue preservation.

Instructions 

RESULTS 

The following postmortem analysis (Part I) is organized to 

include initial cataloging of specimen, the history of the spec- 

imen, and necropsy and laboratory analysis. The actual examina- 

tion is outlined with dissection directions. The examiner need 

only follow the instructions in parentheses until a particular 

symptom occurs, check the space, circle the disorder (symptom) , 

and then using the number at the right of the line as a guide, 

turn to Part 11. In Part I1 are instructions for collection of 

relevant material necessary for laboratory analysis. The numbers 

on the postmortem form also are listed in Tables I and I1 under 

specific diseases; by referring to the tables the examiner can 

determine possible disorders to suggest to the laboratory for 

consideration. Underlined numbers in the tables refer to charac- 

teristic symptoms. However, disease symptoms were determined 

from poultry diseases to a large extent and they may differ in 

other species of birds. Furthermore, many diseases may share 

common symptoms, especially those that undergo a septicemic 

phase. Therefore, in most cases it is only by a thorough 

necropsy analysis supplemented with laboratory tests that a 

particular disease can be positively identified.

PART I 

SPECIES : FIELD I: NECROPSY # 

Area collected: Body measurements: 

Collector: Total length: mm 

Date collected: Wing length: mm 

Date examined: Tail length: mm 

Examiner: Beak length: mm 

Age : Weight: g Tarsus length: - m 

Sex: Gonad meas.: mm Condition of plumage: 

Fat: Skull : Head molt: Tail molt: 

P.M.State: Body molt: Wing molt: 

Preserved in: Worn plumage: Area: 

Brood patch: C1o.P.: 

History of bird: 

MATERIAL TO LABORATORY 

Smears: peripheral blood , heart , liver , spleen 

bone marrow , lungs , kidney , brain , fecal 

other 

Tissue: entire bird , heart , liver , spleen , lungs 

intestine , proventriculus , gizzard , esophagus 

crop , gall bladder , pancreas , kidney , eye 

brain , gonads , nerves , trachea 

bursa of fabricius , muscle , endocrine glands , legs 

feet , other 

Body Wash: Crop Contents: 

Parasites: helminths: 

Cultures : 

Other: 

arthropods: 

other: 

NECROPSY SUMMARY: 

LABORATORY RESULTS: 

DIAGNOSIS:

I. External Analysis 

(Examine bird externally for disorders.) 

a. HEAD AND BEAK 

F a c e : lesions; crusty or scaly scabs (1) 

F a c e 

or sinuses swollen (2) 

Ear disorder (3) 

(Cut eyelids back and expose eyeball.) 

E y e : inflamed; swollen; cloudy; exudate; helminths (4) 

(Cut off beak at nostrils; examine with dissecting microscope.) 

N a s a l chamber: lesions; nodules; exudate (5) 

N a s a l parasites (6) 

Other 

b. BODY AND WINGS 

K e e l prominent (7) 

Vent soiled; diarrhea (8) 

Feathers: dry, easily broken, or absent; follicles infected (9) 

- Skin: dermatitis, ulceration, swelling; uropygial infected (10) 

c. LEGS AND FEET 

Legs or feet: lesions; crusty or scaly scabs (1) 

Legs or feet: swollen; enlarged bones; inflamed joints (11) 

Missing appendages (12) 

Other 

d. BODY WASH FOR EXTERNAL PARASITES 

(Tape bill shut, shake in soapy water, let settle, decant to alcohol.) 

Comments : 

11. Internal Analysis 

a. BODY SURFACE 

(Skin body, neck, head; record fat and skull condition.) 

Muscles: lesions; discolored; hemorrhage (13) 

Feather, follicle, or skin parasites (14) 

0 ther : 

b. UPPER DIGESTIVE AND RESPIRATORY SYSTEMS, SKULL, AND BRAIN 

(Cut from mouth down neck exposing trachea and esophagus.) 

Mouth, pharynx, esophagus: lesions; nodules; cheesy masses (15) 

(Cut esophagus to proventriculus; preserve crop contents in alcohol.) 

Crop: lining thickened; contents sour (15) 

(Cut up length of trachea; examine with dissecting microscope.) 

Trachea: lesions; nodules; exudate (16) 

Tracheal parasites (16) 

(Remove skull; examine brain.) 

Brain: lesions; nodules; discolored; hemorrhage (17) 

Other: 

c. CELOMIC CAVITY AND AIR SACS 

(Cut out breast and slowly lift sternum. Examine air sacs.) 

Air sacs: lesions; nodules; exudate (18) 

(Examine internal organs and if any suggest infection, process immediately to 

avoid contamination.) 

Abdomen: lesions; nodules; exudate (19)

d. HEART AND PERICARDIUM 

(Examine pericardium and remove.) 

Pericardium: exudate; inflamed; discolored; hemorrhage (20) 

(Examine heart; prepare smear from heart blood and fix for Giemsa stain.) 

Heart: enlarged; lesions; nodules; hemorrhage (21) 

Other: 

e. LIVER, SPLEEN, GALL BLADDER 

(Examine liver, gall bladder; measure spleen: -- x nun. 

Liver disorder (22): 

S p l e e n disorder (23): 

Gall bladder disorder (24) : 

(Remove liver and spleen.) 

f. INTESTINAL TRACT AND PANCREAS 

(Examine external appearance of intestinal tract and membranes.) 

Peritoneum: nodules; discolored; inflamed (25) 

Intestine: external ballooning or hemorrhage (26) 

(Remove gastrointestinal tract and straighten on glass plate.) 

(Intestine length: - nun. Separate proventriculus and gizzard.) 

(Cut down length of intestine and lay open.) 

Intestine: lesions; nodules; hemorrhage: 

(Record location of parasites: 

helminths (27) 

- Ceca: lesions; nodules; exudate; hemorrhage; thickened; helminths 

Bursa of Fabricius abnormal (28) : 

(27) 

Other : 

(Take intestinal and cecal smears; check for protozoa parasites.) 

- 

Coccidiosis, Trichomoniasis, Histomoniasis, Other (27): 

(Cut open proventriculus and gizzard.) 

- 

Proventriculus: lesions; nodules; hemorrhage; erosion; helminths (30) 

Gizzard: lesions; nodules; hemorrhage; erosion; helminths (30) 

(Examine pancreas.) 

Pancreas: lesions; chalky; hemorrhage (31) 

- 

h. LUNGS 

(Examine and remove lungs.) 

Lungs: lesions; nodules; exudate; inflamed (32) 

Other: 

i. UROGENITAL SYSTEM AND ADRENAL GLANDS 

(Measure and remove gonads; record sex; examine adrenals.) 

- 

Gonads or assoaatedstructures abnormal (33): 

Adrenal glands abnormal (34) 

(Examine and remove kidneys.) 

Kidneys: lesions; nodules; discolored; enlarged (35) 

Other: 

j . NERVOUS SYSTEM 

(Examine nervous plexus.) 

Nerves: lesions; discolored; swollen (36) 

- 

Other : 

k. SKELETAL SYSTEM 

(Examine vertebrae; break leg bone and examine bone marrow.) 

Bone marrow abnormal (37): 

- Vertebrae or other bones infected (37): 

Other: 

1. COMMENTS:

PART I1 

(1) Face, legs, or feet: lesions; crusty or scaley scabs. 

Scrape part of a lesion onto a microscope slide (with 

water or mineral oil) and examine for mites; preserve in 70% 

alcohol. If exudate is present, prepare a smear for gram stain. 

Divide the remainder of infected tissue into three parts; pre- 

serve one in 10% formalin, freeze part on dry ice, and culture 

the rest in a mycotic medium. 

(2) Face or sinuses swollen. 

Smear a portion of the exudate on two clean slides and 

fix for gram stain. Either freeze an exudate sample or swab the 

sinus area and place in a virus transport medium. 

(3) Ear disorder. 

If the ear is crusty, examine a wet smear for mites. 

Preserve in 70% alcohol. 

(4) Eyes: inflammed; swollen; cloudy; exudate; helminths. 

Examine the eyes for helminths; fix and preserve. Smear 

exudate on a clean slide for gram stain. Fix the infected tissue 

in 10% formalin, cutting a window through the eyeball so that 

fixatives can reach the internal structures. Be aware that this 

is often a secondary symptom and primary disease disorders will 

probably occur elsewhere. However, if there were nervous symp- 

toms before the bird died, preserve the brain and nerve tissue at 

end of necropsy by freezing on dry ice. 

(5) Nasal chamber: lesions; nodules; exudate. 

From the exudate prepare a smear for gram stain and col- 

lect two swabs. Place one swab in a transport medium for virus 

and the other in bacteria transport medium. Preserve half of the 

infected tissue by freezing and the other half in 10% formalin. 

This may be a secondary symptom; therefore, examine the remaining 

respiratory system carefully. 

(6 ) Nasal parasites. 

Examine the nasal chamber with a dissecting microscope; 

preserve and fix parasites.

(7) Keel prominent. 

A prominent keel is often indicative of a chronic dis- 

ease; however, since it is also a secondary manifestation of many 

disorders, look for other symptoms. 

(8) Vent soiled; diarrhea. 

A soiled vent indicates diarrhea, a symptom of many dis-. 

eases. Look for other disorders and be especially careful to 

examine the digestive tract. 

(9) Feathers: dry, easily broken, or absent; follicles 

infected . 

Examine feathers under a microscope for ectoparasites; 

preserve in 70% alcohol. The entire feather may be placed in 

alcohol rather than removing the parasite. Follicles and inner 

shaft should be examined for mites. If the skin is dry, or 

scaley and powdery, preserve a section in 10% formalin. 

(10) Skin: dermatitis, ulceration, swelling; uropygial infec 

ted . 

Proceed as in #(1). If possible collect parasites and 

preserve in alcohol. 

(11) Legs or feet: swollen; enlarged bones; inflammed joints. 

Using a sterile syringe, collect fluid from joints (in- 

cluding wing joint), and smear for gram stain. Place a portion 

(or a swab) into a bacteria medium and either freeze (-60°C) the 

remaining exudate or place in viral transport medium. Preserve 

some infected tissue by freezing and the rest in 10% formalin. 

(12) Missing appendages. 

If appendages are missing consider the bird's history in 

terms of trauma (e.g., freezing) or past viral (e.g., Pox) infec- 

tions. Be sure to include this in the history of the bird. 

(13) Muscles: lesions; discolored; hemorrhage. 

If nodules are obvious on pectorals, open one and examine 

for nematode larvae or mites; preserve in 70% alcohol. Smear 

necrotic lesions and prepare for a gram stain. If exudate is 

present (e.g., blood) prepare a swab and place in a bacteria 

medium . Fix infected tissue in 10% formalin. This is often a 

secondary symptom so be careful to look for other indications of 

disease.

(14) Feather, follicle, or skin parasites. 

Examine the internal surface of the skin for mites; if 

present preserve in alcohol. Proceed as outlined in #(9). 

(15) Mouth, pharynx, esophagus: lesions; nodules; cheesy 

masses. Crop: lining thickened; contents sour. 

Lesions in the mouth should be smeared for gram stain 

or preserved in 10% formalin. If cheesy, culture on a mycotic 

medium for shipment. A wet smear may reveal protozoa (especially 

if crop lining is thickened); proceed as outlined in #(29). Pre- 

serve crop contents and tissue in formalin. Fix and preserve any 

helminths. 

(16) Trachea: lesions; nodules; exudate. 

Freeze or swab exudate and place in viral transport 

medium. Prepare an exudate for gram stain. Preserve lesions and 

tracheal tissue in 10% formalin and on dry ice. The entire 

length of the trachea should be examined for nematodes; fix and 

preserve. 

(17) Brain: lesions; nodules; discolored; hemorrhage. 

Swab tissue and place in bacteria medium. Make two 

impression smears (one fixed for gram stain and one for Giemsa). 

Divide the remaining brain tissue in half; freeze part and place 

the rest in 10% formalin. Collect a sample of body fat; freeze. 

(18) Air sacs: lesions; nodules; exudate. 

Make a tissue smear for gram stain and search tissue for 

mites. Prepare diseased tissue by freezing and in 10% formalin. 

Also freeze (even though they may appear normal) sinus tissue, 

trachea, lungs, and cloaca. 

(19) Abdomen: lesions; nodules; exudate. 

Lesions or exudate in the abdomen should be smeared for 

gram stain. If the lesions are nodular, examine for mites or 

nematode larvae; fix and preserve. The remaining diseased tissue 

should be place in 10% formalin. 

(20) Pericardium: exudate; inflammed; discolored; hemorrhage. 

Prepare two smears (for gram stain and Giemsa) from 

exudate. Preserve the remaining tissue in 10% formalin. Since 

this is often a secondary symptom, samples of tissue from the 

kidney, bone marrow, bursa of Fabricius, lungs, liver, spleen, 

and brain should be routinely collected.

(21) Heart: enlarged; lesions; nodules; hemorrhage. 

Prepare lesion, organ impression, and heart blood smears 

for gram stain and Giemsa stain. Swab tissue and place in a 

bacteria medium. Preserve what remains of the heart in 10% 

formal in. Tissue from other representative areas of the body 

(e.g., liver, spleen, kidney, bone marrow, lung, brain--freeze 

part--and bursa of Fabricius) should also be collected. 

(22) Liver disorder. 

Prepare two organ impression as well as lesion and exu- 

date smears for gram stain and Giemsa stain. If tubercles are 

present on the liver smash a small (2 nun) tubercle between two 

slides and prepare for Ziehl-Neelsen stain. Swab tissue and 

place in bacteria medium. Divide the remaining tissue, placing 

part in 10% formalin and freezing the rest on dry ice for ship- 

ment. A sample of heart blood should be prepared for Giemsa 

stain and blood serum collected. Collect spleen, heart tissue, 

lung, and bone. 

(23) Spleen disorder. 

Follow the procedure outlined in # (22) and be sure to 

collect a sample of the liver. 

(24) Gall bladder disorder. 

Using a sterile syringe collect some bile and place in 

bacteria medium. Examine the bile ducts for helminths and if 

present, preserve. 

(25) Peritoneum: nodules; discolored; inflammed 

Prepare a gram stain from lesion. If small tubercles are 

present smash some between two slides and fix for Ziehl-Neelsen 

stain. The remaining tissue should be placed in 10% formalin. 

(26) Intestine: external ballooning; hemorrhage. 

Examine carefully without opening the intestine. Prepare 

smears of any lesions for gram stain. 

(27) Intestine: lesions; nodules; hemorrhage; helminths. 

Ceca: lesions; nodules; exudate; hemorrhage; thickened; 

helminths. 

Before proceeding, prepare a swab from lesions and place 

in bacteria medium. Omit if intestinal contents have contam- 

inated the area such that lesions cannot be seared and incised.

Examine a wet smear of intestinal contents at several places for 

intestinal protozoa (see #I291 ). Smears of lesions should be 

prepared for gram stain. Fix the intestinal and cecal contents 

in 1% formalin and the tissue in 10% formalin. Freeze sections 

of the bursa of Fabricius, liver, spleen, bone marrow, and take 

blood serum. Sections of the heart, lung, kidney, liver, and 

spleen should be placed in 10% formalin. Examine the length of 

the intestine for helminths and if present, preserve. 

(28) Bursa of Fabricius abnormal. 

Divide the organ in half and preserve part in 10% 

formalin and freeze the rest on dry ice. 

(29) Intestinal Protozoa. 

To check for protozoa parasites, a wet smear w i l l usually 

suffice. However, Lugol's solution w i l l often facilitate obser- 

vation. 

(30) Proventriculus: lesions; nodules;' hemorrhage; erosion; 

helminths. 

Gizzard: lesions; nodules; hemorrhage; erosion; helminths. 

If lesions are present, swab and place in a bacteria 

medium. Prepare a smear from heart blood for gram stain. Any 

cheesy exudate should be transfered to a mycotic medium for 

culture and/or placed in 10% formalin. Be sure to remove the 

gizzard lining and examine for helminths; if present, preserve. 

(31) Pancreas: lesions; chalky: hemorrhage. 

Place the entire organ in 10% formalin. 

(32) Lungs: lesions; nodules; exudate; inflammed. 

From the exudate prepare two swabs; place one in a bac- 

teria medium and one in a transport medium for viruses. Fix an 

exudate smear for gram stain. Any cheesy exudate should be cul- 

tured on a mycotic medium. Preserve half the remaining tissue in 

10% formalin and freeze the rest. Fix a heart blood smear for 

Giemsa stain and separate serum into a sterile vial. 

(33) Gonads or associated structures abnormal. 

Prepare a swab for bacteria medium and make two impres- 

sion smears for gram stain and Giemsa stain. Freeze some of the 

remaining tissue and place the remainder in 10% formalin. 

Samples of the bone marrow, heart blood, liver, spleen should 

also be collected.

(34) Adrenal Glands abnormal. 

Look for other disorders, but note especially if shock 

might be suspected (e.g., hemorrhage of heart). 

(35) Kidney: lesions; nodules; discolored; enlarged. 

Prepare swabs from lesions for bacteria medium. Make 

impression smears for gram and Giemsa stain. Divide the remain- 

ing tissue in half and freeze part; examine the other half care- 

fully for tramemodes and Protozoa and if present, preserve. 

Collect blood serum in sterile vials and refrigerate. Also 

collect tissue from the heart, liver, and spleen. 

(36) Nerves: lesions; discolored; swollen. 

Preserve nerves that are diseased in 10% formalin. ~r e- 

pare a heart blood smear for Giemsa stain and see #(17) for 

possible treatment of the brain. Even if brain appears normal, 

preserve it in 10% formalin. 

(37) Bone marrow abnormal; vertebrae or other bones infected. 

Prepare two bone marrow smears, one for gram stain and 

one for Giemsa stain. Freeze part of the tissue and preserve the 

rest in 10% formalin. If vertebrae are implicated, check 

carefully for joint involvement and place exudate in bacteria 

med ium .

SUMMARY 

More work needs to be done on disease in wild birds, espe- 

cially studies which delimit the entire parasitic fauna present 

within a host population. Recording levels of a single pathogen, 

as most surveys to date have done, cannot possibly determine the 

impact disease is playing upon wild populations. A multi-disease 

approach is necessary, one which reveals the inter-relationship 

between all parasites and diseases within a host population. 

We have presented this postmortem technique in hopes that 

more ornithologists w i l l be inspired to attempt such multi- 

disease studies. These surveys will hopefully provide enough 

information so that disease interactions can be defined, possibly 

simulated in laboratory situations, and control measures can be 

found which would be applicable to the natural state.


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Stone, R. M. 1969. Clinical examination and methods of treatment. 

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Iowa.

TABLE 1. Summary of diseases in avian hosts '. 

Disease Pathogen Major ~ost ~ymptcans~ Hosts Wported Susceptible3 

Bacterial Diseases 

Arizonosis Septicemic; (4,17,18, 

19,21,22,27,32,33,36) 

Anthrax 

Botulisn 

Chlamyd iosis 

(Ornithosis) 

Cholera 

Clostridia 

Colibacillosis 

Erysipelas 

Infectious Coryza 

Bacillus anthracis 

Clostridiun botul inm 

Chlanydia psittaci 

Pasteurella multocida 

Clostridiun sp. 

Erysipelothr ix insidosa 

Hemophilus sp. 

Nervous 

Nervous; (u,36) 

Septicemic; (2,4,5, 

15,16,17,18,19,22,25, 

- 27,30,32,33,36) 

Wound infection; 

Digestive; (10,13, 

22,2Jr30,35,T) 

Str ,Ci ,A,F,C 

G (necrotic enteritis); 

probably all species suscep 

tible to wound infection or 

gangrene 

Str ,A,F,G,Gr,Ch,Ps,St,P

Disease Pathogen Major ~ost ~ymptcms ~osts &pr ted ~usce~tible' 

Bacterial Diseases (Con' t. ) 

Infectious Serositis Pasteurella anatipestifer 

(Duck Septicenia) - P. septicaeniae 

Listeriosis 

Mycoplasnosis 

Pseudo tuberculosis 

Salmonellosis 

Spirochaetosis 

Staphylococcus 

Streptococcus 

Listeria monocytogenes 

Mycoplasna sp. 

Pasteurella 

pseLdotubercul&is 

Salmonella sp.(over 

1000 pathogenic sp.) 

Borrelia anserina 

Staphylococcus aureus 

Streptococcus sp. 

septicmic; (4,17,18, A,F,G,Q,Ch,C,Ps,St,P 

19,2_0,2l,2,23r27, 

32,35,36) 

Respiratory; F,G,C,Ps,P 

Skeletal; (2,5,10, 

-I-,-, 11 16 18 19r20r22, 

Z,35) 

Septicemic; (13,18, 

19 22 23 27,321 

-I- l- I- 

Digestive; 

Se~ticenic: (4.11. 

A,F,G,Q,Ch,C,Ps,Cu,St,T, 

Cor ,Pic ,P 

Skeletal; (1,2,3,10, Str ,Ci,A,F,G,Q,Ch,C,Ps,Ap, 

- 11,22,37) 

C,P (mon on skin and 

mucous membrane)

TABIE l--Continued '. 

Disease Pathogen Major ~ost Syrnptuns2 Hosts &ported Susceptible 

Bacter id Diseases (Con' t .l 

Tuberculosis Mycobacter iun aviun 

Ulcerative Corynebacter iun sp. 

Enter itis 

Vibrio Infections Vibrio sp. 

Fungal Diseases 

Aspergillosis Aspersillus fmiqatus 

Candidiasis Candida albicans 

Cryptococcus Cryptococcus neoformans 

Favus 

Protozoan Diseases 

Microsprun sp. 

Tr ichophyton sp. 

Coccidiosis Isospora; Eimeria; 

and others. 

Haemoproteus Haemoproteus sp. 

Parahaemoproteus sp. 

Viscera; (1,11,18,19, Sp,Str ,Rh ,Ca ,Ti ,Ga ,Fel ,Ci ,A, 

- 22,23 -8- 27 I 32,33,37) 

F,G,Gr,Ch,C,Ps,Cu,St,Ap,Cor, 

Pic ,P 

Digestive; (22,23,27) F&,C 

Liver; Digestive; (20, Sp,Pod ,Pro ,Pel ,Ci ,&,F&,Gr, 

21,~,23,~,33,35) Ch,C,Ps,St,P 

I 

Respiratory; (l,2,4, Sp,Str ,Rh ,Ti ,Ga ,Fro ,El ,Ci , 

- 5,9,10,15,17,&3,22,32) A,F,G,Gr,Ch,C,Ps,St,T,Cor, 

P 

Digestive; (l5,27,30) Sp,Fh,Ci,A,G,Q,Ch,C,Ps,Cu, 

Ap,Pic,P 

Skin; (1,5,2,lJ, 

16,18,32) 

Digestive; (a,a, Pro,A,G,C,Ps,P others? 

29,35) 

- 

Blood; (17,18,22, Pod,Ci,A,F,G,Gr ,Ch,g,Ps,Cu, 

23,32) 

St,Cap,Ap,T,Cor ,Pic,P

TABLE 1--Contin& ' . 

W 

N 

Disease Pathogen Major Host Symptans2 Hosts Reported Susceptible OD 

Protozoan Diseases (Con't.) 

Histmoniasis 

Leucocytozoonosis 

Malaria 

Other blood 

Protozoa 

Sarcospx id iosis 

Tr ichamoniasis 

Trypanosomiasis 

Viral Diseases 

Arbovirus 

Bl uecomb 

Duck Virus Enteritis 

(Duck Plague) 

Histomonas meleagr id is 

teucocytozoon sp. 

Plasmod iun sp. 

Aegyptianella, 

Lankestrella, 

Toxoplasoa 

Sarcocystis rileyi 

nichomonas sp. 

Trypanosoma sp. 

Digestive; (22,.27,29) 

Blood; (17,21,2,23) 

Blood; (17,22,23) 

Blood; (11,17,18, 

20,21,22,23,27,32, 

35) 

Blood; (37) 

Se~ticenic: Diqestive: 

Duck Virus Hepatitis Liver; (17,2,23,35) 

- G 

Ci,A,F,G,Gr,Ch,C,Ps,Cu,St, 

T,Cor ,Pic ,P 

SprCitA,F,G,Gr rChrCrO~rStr 

Cor ,Pic,P 

Ci,A,F,G,Gr,Ch,C,Ps,Cu,St, 

Cap,Ap,Cor ,Pic,P 

Ci ,A,F,GtGr ,C,Cu,St,Ap, 

Pic ,P

TABLE 1--Continued '. 

Disease Pathogen Major Host ~ymptms~ Hosts kprted Susceptible 

Viral Diseases (Con' t . 

Encephalmyel itis 

Hemorrhagic Enter itis 

Infectious Bronchitis 

Infectious Bursal 

Disease 

Influenza 

(over 80 types) 

(Fowl Plague) 

Laryngotracheitis 

Monocytosis 

Newcastle Disease 

POX 

Puffinosis 

Quail Bronchitis 

Nervous; (4 ,lJ,30r 36) 

Respiratory; (2,4,5,l6, 

17,18,19,20,22,23,25,32, 

33,35) 

~espiratory; (2,4,5,15, 

16) 

Digestive; Viscera 

systemic; Nervous; Respi- 

ratory; (4,5,16,11,1_8,20, 

22,23,24,Zr3J,32,33,1_6) 

Skin; Mucous menbranes; 

(~r5tlOrllr12,~,16) 

Skin (feet); Nervous; (I, - 

17) 

- 

Respiratory; (2,4,5,16, 

18,321 

- 

probably all species of birds

TMIE 1--Continued ' . 

Disease Pathogen Major ~ost symptoms2 Hosts Reported Susceptible3 

Viral Diseases (Con' t.) 

Turkey Viral Hepatitis 

Viral Arthritis 

Neoplastic Diseases 

Erythroblastosis 

Hemang iana 

Leukosis complex 

Marek' s Disease 

Myelocytanatosis 

Nephroblastma 

Osteopetrosis 

Other Neoplasns 

Circulatory; (9,10,13,18, - G 

22,gr32,z,37) 

Skin; Viscera; (9,g) G 

viscera; (18,19,2lrZr1_3~ Ci,A,G,C,Ps,P 

28,30,32,33r35r37) 

- 

Nervous; Viscera; (1,4, A,F,G,C,Ps,St ,P 

13,~,19,21,22,23r25r27, 

28,30,31,33,35,36) 

- 

Bone marrow; (9,22,23,35, - G 

37) 

- 

Skeletal; (lJ ,13,2) G 

Kidney; (35) - 

- 

- G 

Skeletal; (E,37) G 

viscera; Various sites Str ,G,C,E,P - 

- 

-

' Information not our own is £ran Garnham (1966), Davis et al. (1971), Ebfstad et al. (1972), 

Arnall and Keymer (1975), Qeiner et al. (1975), McClure et al. (1978). 

Numbers refer to symptoms listed on postmortem form and outlined in Part 11; these nunbers 

indicate the symptans most likely to be found when that disease is present, and the under- 

lined nmbers are very characteristic symptoms. 

Letters refer to the orders of birds in which the diseases have been reported. Wild, 

domestic, cage, and laboratory groups that have shown susceptibility are incllded. The 

underlined orders indicate a host group in which that disease is particularly common. 

Key to orders: Sp = Sphenisciformes, Str = Struthioniformes, Rh = Rheiformes, Ca = Casuarii- 

formes, Apt = Apterygiformes, Ti = Tinamiformes, Ga = Gaviiformes, Pod = Podicipediformes, 

Pro = Procellariiformes, Pel = Pelecaniformes, Ci = Ciconiiformes, A = Anseriformes, 

F = Falconiformes, G = Galliformes, Gr = Gruiformes, Ch = Charadiiformes, C = Colunbiformes, 

Ps = Psittaciformes, Mu = Musophagiformes, Cu = Cuculiformes, St = Strigiformes, 

Cap = Caprimulgiformes, Ap = Apodiformes, Col = Coliiformes, T = Trogoniformes, Cor = Cora- 

ciiformes, Pic = Piciformes, P = Passeriformes.

BRINGING BACK THE MONARCH OF HAWAIIAN FORESTS--ACACIA - KOA 

Gerald A. Walters 


Forest Service, U. S. Department of Agriculture 

Berkeley, California, stationed in Honolulu, Hawaii 

Koa has been called the "Monarch of Hawaiian Forests." This 

is a fitting title for a species found on about 500,000 acres in 

the State. Trees may reach 120 feet tall, 10 feet in diameter, 

and more than 100 feet in crown spread. Koa is important as a 

component of the native forest to birds, insects, mollusks, and 

different plant species with which it grows in association. Its 

wood is valued highly for furniture, cabinets, veneer, and craft 

pieces. The technical properties of koa wood are very similar to 

those of black walnut (Juglans ni ra) . The koa industry, including 

harvesting, manufactur~ng , *sale of finished products, 

both here and elsewhere, generates about $7,000,000 annually. 

Koa forests are not as extensive as they once were. The 

principal reason is the effect of grazing anlmals that eat or 

otherwise damage young trees. Fire, insects, and diseases have 

destroyed many stands. Harvesting does not necessarily reduce 

the area of koa forest if natural regeneration that develops 

after site disturbance is allowed to grow and develop into a new 

forest. Too often, however, harvesting has been the first step 

in converting forest to pasture. An estimated 100,000 acres of 

koa forest have been converted to pasture in the past 50 years. 

Cattle, of course, are very effective in preventing reestab- 

lishment of a koa forest. 

In an effort to rehabilitate denuded watersheds, many seed- 

lings of many different species were planted on the Forest 

Reserves from 1900 to 1940. About 1.1 million koa seedlings were 

planted, making it the fourth most widely planted species. When 

the once barren watersheds were revegetated and when labor became 

scarce because of World War 11, interest in reforestation with 

koa or other species largely ceased. Little reforestation of any 

kind was done during the 1940's and the early 1950's. 

In the late 19501s, people began to realize the multiple 

values of the trees that had been planted in previous decades. 

Interest in reforestation was renewed, but not with koa. The 

Hawaii Division of Forestry built a bare-root nursery in 1961 

--principally for growing pine and eucalyptus seedlings. The 

bare-root system of production, transport, and planting requires 

a hardy species if it is to work satisfactorily. And koa is 

definitely not a hardy species. Although interest in forestation 

with koa has increased during the last 10 years, efforts to use

the bare-root system have failed. Koa seedlings had been suc- 

cessfully raised in and planted from flats and tin cans, but 

because of the high cost of labor, this method was not econom- 

ically feasible. 

About 5 years ago, in cooperation with the Division of 

Forestry, I began developing a new system for successfully rais- 

ing, transporting, and planting seedlings. The system is based 

on a small, specially designed container called the "Hawaii 

Dibbling Tube." The container is 5 inches deep and 1-1/8 inches 

inside top diameter. Four ridges that extend from top to bottom 

on the inside of the tube prevent root spiraling. The tubes are 

filled with rooting medium, then seeds are sown and covered. 

After about 4 months, seedlings are ready for outplanting. Seed- 

lings are removed from the tubes, packed in wax-lined boxes, and 

shipped to the planting site. Seedlings are planted using a 

dibble which, when driven into the ground makes a hole the same 

size and shape as the seedling root system. The tree planter 

makes the hole and drops in the seedling. The dibbling tube sys- 

tem is proving to be efficient in terms of seedling production, 

transport, and planting. Its real worth is best measured by the 

degree of seedling survival after planting. And the bottom line, 

of course, is that the trees generally survive after planting. 

The first planting of dibbling tube seedlings was made about 

4 years ago. About 100 koa seedlings were planted in the hapu'u 

harvest area in the Kilauea Forest Reserve. The seedlings, grown 

by the green-thumb method, were of reasonable quality. We did 

not know then, nor do we know now, what constitutes the best 

seedlings in terms of stem height and diameter, leaf number and 

area, shoot/root ratio, etc., for maximum survival and growth on 

a variety of sites under a variety of weather conditions. Nor do 

we know the cultural treatments, such as fertilizer formulation 

and concentration, light intensity, temperature, etc., to obtain 

the best seedlings. Even with these unknowns, this first plant- 

ing of koa was successful. About 95% survived, and they showed 

rapid initial growth. About a year later we made another 

planting with similar results. 

Results of these two plantings indicated that koa could be 

successfully planted in terms of survival, growth, and costs. 

Reforestation with koa again became feasible. 

These first efforts with koa and other species were on a 

research basis. In other words, we grew a few seedlings of dif- 

ferent species and planted them on different sites to test an 

idea. The idea worked so we expanded from a research basis to a 

pilot-scale production basis--expanding from a scale of hundreds 

to a scale of thousands of seedlings. The pilot-scale production 

nursery was constructed at the Division of Forestry bare-root 

nursery at Kamuela. 

The first crop of 40,000 koa seedlings from the pilot-scale 

production nursery was contracted for by the Bernice Pauahi 

Bishop Estate. In growing these trees, we tried to do everything 

to develop seedlings which would have high survival and growth

potential. For example, we collected nitrogen-fixing nodules 

from roots of koa seedlings growing in the area where the 

nursery-grown seedlings were to be planted and isolated the bac- 

terium responsible for nitrogen fixation. The bacterium was 

applied to all the koa seedlings in the nursery. Seedlings were 

watered, fertilized, exposed to full sunlight, etc., according to 

the green-thumb instincts of the nurseryman. When we thought the 

seedlings were ready for field planting, we packed them in wax- 

lined boxes and shipped them to the Keauhou-Kilauea Forestry 

Center for planting. 

The Keauhou-Kilauea Forestry Center is a project sponsored 

by the Bishop Estate. This project, on about 200 acres of cut- 

over and grazed-over koa-'ohi'a forest, is aimed at restoring koa 

for eventual sustained-yield management. Technical guidance for 

the project is being provided by State, private, and Federal 

organizations. 

The 200-acre area was fenced and divided into four 50-acre 

sections. It was decided to harvest merchantable koa trees, pre- 

pare the site, and plant where necessary on 50 acres at a time. 

That way, if mistakes were made or better methods were developed, 

other areas would benefit. The area was fenced, of course, to 

keep out cattle. 

Although natural koa reproduction generally develops in 

adequate numbers following site disturbance, their spatial dis- 

tribution is often irregular. Seedlings are most common where 

seed trees once were. Koa seedlings are planted to obtain 

uniform stocking within an area. 

A total of 36,000 seedlings were planted among the natural 

seedlings to obtain a 5- by 5-foot spacing between all seedlings. 

The first 18,000-seedling planting was done in August 1977 by 

Kamehameha School students and several adults. The second 

planting was done by welfare recipients in November. 

We did learn from the first planting as evidenced by the 

fact that only 56% survived compared to about 98% for the second 

planting. Seedlings- for the second planting were much hardier 

than those used for the first planting. Also, we were luckier 

the second time as rainfall was more than adequate. As of May 

1978, there were about 2600 natural and planted seedlings per 

acre. Natural seedlings averaged about 20 inches tall. Seed- 

lings planted in August averaged about 24 inches tall: those 

planted in November averaged about 16 inches tall. 

We had some mortality in both natural and planted seedlings 

due to frost. Apparently, the opening made in the forest by har- 

vesting and site preparation resulted in greater damage from cold 

air during periods of freezing temperatures in December and 

January. Less than 5% of the seedlings were affected.

Results of the efforts on the first 50-acre section were 

successful enough that work was started on the second 50 acres 

early in 1978. Seedlings were planted in May 1978. If we get 

sufficient rain, I feel certain that the survival rates will 

again be high. Also, because the seedlings were fertilized with 

1 ounce of 10-30-10 placed in a hole next to the seedling, ini- 

tial seedling growth should be rapid. This fertilizer treatment 

was based on a study we did which indicated that initial growth 

rate could be more than doubled with just 1 ounce of 10-30-10. 

On the basis of results of the first crop from the pilot- 

scale container nursery, the Division of Forestry plans to 

develop it into a full production nursery with a capacity of 

1 million seedlings per year. 

The planting of 36,000 seedlings at Keauhou-Kilauea was the 

first major koa reforestation project in about 35 years. Now we 

have just had the second. It is exciting to think that koa 

reforestation is now biologically and economically feasible. We 

not only have the potential to bring back the monarch of Hawaiian 

forests, we have the ability. Now we have to do it.



VEGETATION MAP OF THE CRATER DISTRICT 

Louis D. Whiteaker 




A vegetation map of the Crater District of Haleakala 

National Park was produced at a scale of 1:24,000 that can over- 

lay a composite of the USGS topographic quadrangle maps that 

cover the same area. After an initial field reconnaissance, the 

mapping was carried out using 1: 12,000 aerial photographs. The 

map units were field checked as to the accuracy of the boundary 

positions and the structural and floristic composition of the 

vegetation units. The boundaries were transferred to overlays on 

1:12,000 prints of NASA false infrared color aerial photographs. 

These maps served as a base for the final map which was produced 

with photographic methods. 

The mapped vegetation has been classified into 53 

structural-floristic communities that are grouped into four 

structural vegetation-types: forest communities, scrub commu- 

nities, grassland communities, and high altitude desert commu- 

nities. Forest communities were defined as areas with the 

tallest vegetation layer composed of woody vegetation greater 

than 5 m tall that had at least 30% crown cover. Scrub commu- 

nities were defined as areas in which the uppermost vegetation 

layer consisted of woody vegetation greater than 0.3 m but less 

than 5 m in height with crown cover greater than 30%. Grassland 

communities were defined as areas in which grass species had more 

than 30% cover while woody species had less than 30% cover. High 

altitude desert communities were defined as areas with less than 

30% total plant cover. 

Cover has been defined by Mueller-Dombois and Ellenberg 

(1974) as the vertical projection of the crown or shoot areas of 

a species to the ground and expressed as a percentage of the ref- 

erence area. In this study, closed cover was defined as greater 

than 60%, open cover as between 30%-60%, and sparse cover as less 

than 30%. 

The communities were labeled using a combination of symbols 

derived from generic names or other predominant surface cover 

that correspond as closely as possible to and are used in a sim- 

ilar manner as those used in Mueller-Dombois and Fosberg's (1974) 

vegetation map of Hawaii Volcanoes National Park. A total of 19 

symbols were used in combinations to classify the vegetation into 

the 53 structural-£lor istic communities that were mapped.

Areas for these vegetation communities were generated using 

an electronic planimeter. Scrub communities had the largest 

total area of 3691.7 hectares (9122 acres). High altitude desert 

communities were next with an area of 3119.7 hectares (7708 

acres) and grassland communities covered 568.6 hectares (1405 

acres). Forest communities had the smallest total area of 164.9 

hectares (407 acres). 

Using the map as a base, three topographic vegetation pro- 

files were constructed to aid in the interpretation of the map 

units. The courses of the profiles are shown in Figure 1. These 

courses were chosen so as to cross as much of the study area as 

possible while illustrating as much of the range in vege- 

tation-types and environmental factors as possible. Profile 1 

runs from Pu'u Nianiau at the Park boundary on the northwestern 

outer slope at 2087.5 m (6849 ft) elevation to Kilohana on the 

western rim of the crater at 2926.1 m (9600 ft). Profile 2 runs 

from Kilohana, across the crater floor, to the eastern rim above 

Paliku Cabin that separates Haleakala Crater from Kipahulu Valley 

at 2133.6 m (7000 ft). Profile 3 runs from the southern Park 

boundary in Kaupo Gap at 1158.2 m (3800 ft) up over Kalapawili 

Ridge at 2484 m (8150 ft) to the northern Park boundary on the 

northern outer slope at 2316.5 m (7600 ft). 

Profile 1 (Fig. 2) shows a decrease in both mean annual 

precipitation and mean annual temperature to be associated with 

the increase in elevation. An apparent effect of the temperature 

gradient on the vegetation can be seen at approximately 2590.8 n 

(8500 ft) where the vegetation becomes very sparse and is termed 

a high altitude desert. This change may be associated with the 

diurnal frost boundary, above which freezing temperatures occur 

at ground level every night of the year. Mueller-Dombois (1967) 

found the diurnal frost boundary to occur at about this elevation 

on Mauna Loa. 

Profile 2 (Fig. 3) shows an increase in mean annual temper- 

ature associated with the decrease in elevation, and an increase 

in mean annual precipitation associated with the west-east orien- 

tation. The increase in rainfall is related to a greater expo- 

sure to the effects of the predominant northeasterly trade winds. 

These factors result in a gradual increase in cover and stature 

of the vegetation from a low growing very sparse vegetation (high 

altitude desert), through several variations of scrub 

communities, to a low stature 'ohi'a rain forest. 

Profile 3 (Fig. 4) shows a decrease in mean annual temper- 

ature associated with the increase in elevation, and an increase 

in mean annual precipitation resulting from greater exposure to 

the effects of the northeasterly trade winds associated with the 

south to north orientation. Also, the lower end of this profile 

extends below the inversion layer which occurs between 1700 and 

2300 m (5000-7000 ft) elevation (Blumenstock & Price 1967). This 

complex of factors is associated with the occurrence of several 

forest communities between 1292 and 1890 m (4240-6200 ft) which 

are unique to this section in the study area.

This variation of the climatic regimes within the Crater 

District and along these profiles can be viewed graphically on 

climate diagrams constructed by the method of Walter (1963) 

(Fig. 5). Mean annual temperature ranges from 17.6OC (64°F) just 

below the southern Park boundary in Kaupo Gap at 1088 m (3570 ft) 

elevation to 8.0°C (46OF) at Haleakala Summit at 3055 m (10,025 

ft). Mean annual precipitation ranges from 1077 mm (42.4 in.) at 

the summit to 4508 mm (177.5 in.) at Paliku Cabin at 1945 m (6341 

ft) elevation. All five diagrams have a mean monthly temperature 

curve that shows little seasonal variation indicating a tropical 

climate at all elevations. Also, all five diagrams show a sim- 

ilar annual pattern of precipitation with the wettest months 

being January and December and the driest month being June. 

Drought conditions are indicated if the mean monthly precipi- 

tation curve crosses below the mean monthly temperature curve and 

is indicated on the diagrams by the stipled areas. This occurs 

at three stations: Haleakala Summit, Haleakala Ranger Station, 

and Holua Cabin, but only for a short period in June. The mean 

monthly precipitation curve for Paliku Cabin is above 100 mm for 

all months and thus indicates a rain forest climate. 


Blumenstock, D. I., and S. Price. 1967. Climates of the states: 

Hawaii. U.S. Department of Commerce, Environmental Science 

Services Administration, Washington, D.C. 

Mueller-Dombois, D. 1967. Ecological relations in the alpine 

and subalpine vegetation on Mauna Loa, Hawaii. The Journal 

of the Indian Botanical Society 46(4): 403-411. 

Mueller-Dombois, D., and H. Ellenberg. 1974. Aims and methods 

of vegetation ecology. John Wiley and Sons, Inc., New York. 


Hawaii Volcanoes National Park. CPSU/UH Tech. Rep. 4 (Univ. 

of Hawaii, Botany Dept.) ii+44 pp. 

Walter, H. 1963. Climatic diagrams as means to comprehend the 

various climatic types for ecological and agricultural 

purposes. Pages 3-9 in A. J. Rutter and F. H. Whitehead. 

The water relationsaf plants. John Wiley and Sons, Inc., 

New Yor k.

FIGURE 1. Courses of the topographic vegetation profiles 

of the Crater District of Haleakala National 

Park.

1342 rnrn t MEAN ANNUAL PRECIPITATION- 1118rnm- 1077 mm 

11.5.~ -MEAN ANNUAL TEMPERATUREt------ 8.6'~- 8.0'~ 

3 

- 

Y)TrTRI 1- U U AFlBm€SC€NT WVBS) a FLAT GRoWlNC SHRUBS ISTYRZLU.RAlLL)LRMA ETCJ 

3 Y~RL~~SISITT~ELU.VPLWIW.ETCJ MIXED W S E S ~)NTWXMTHUM.FESTW,U&CUS.ETCI 

4. W7W LDI Sim (CCS?W9U.-A.GER*HIUWETC 1 V.1 U101 ALTITUDE TUSSW GHASSES (DEXMMEU.TRIYlLM. ETCI 

FIGURE 2. Topographic vegetation profile 1 of the Crater District 

of Haleakala National Park.

2500 mm ---+ MEAN ANNUAL PRECIPITATION 4508 mm----+ (est. 3500 mm) 

17.2"~4- MEAN ANNUAL TEMPERATURE -12.3Oc --------, 10.3Oc 

9 MIRSINE LANAIENSIS 

R A N G E OF LNVERSION LATER 

I 

I mile 

:u 

I 

KALAPAW 

cnr-m. cO sns-mx 

I I 

Q GLOBOSE SnRUBS (STYPHELIA, VACCINIUM, ETC) 

NATIVE LOW SHRUBS (COPROSMA. RAILLARDIA. STYPHELIA, ETC) 

aru MIXED GRASSES (PENNISETUM, FESTUCA. HOLCUS ETCI 

W 

VVV HIGH ALTITUDE TUSSOCK GRASSES IDESCHAMPSIA, HOLCUS, ETC) .p 

W 

FIGURE 4. Topographic vegetation profile 3 of the Crater District of Haleakala 

National Park.

FIGURE 5. Climate diagrams and locations of stations for 

the Crater District of Haleakala National Park.

FOREST BIRD POPULATION VARIATION AS RELATED TO HABITAT TYPE* 

Claire M. Wolfe, C. John Ralph, and Paul K. Higashino 



A 16-ha site was studied on the Keauhou Ranch, Hawai'i, to 

determine the interrelationships between vegetation type and 

presence of native and exotic birds. The area was subdivided 

into 64 0.25-ha plots and the vegetation sampled on each plot. 

On a twice-monthly basis, birds were systematically mapped on the 

study site, and these data compared with the vegetation samples. 

Birds were identified as to species, and when possible as to sex 

and age. One year of data has been collected and preliminary 

analysis shows inter-specific differences in habitat preference. 

- 

* Abstract

VEGETATION OF THE HANA RAIN FOREST 

HALEAKALA NATIONAL PARK 

Alvin Y. Yoshinaga 




As part of the Hana Rain Forest Project in 1973, the vegeta- 

tion of the rain forest zone of the northeastwindward slope of 

Haleakala was studied. The study area included Kalapawili Ridge 

from below Wai'anapanapa to 1610 m (5400 ft), and part of the 

Ko'olau Forest Reserve northeast of Pu'u Alaea. The vegetation 

consisted of 'ohi'a (Metrosider0.s collina) montane rain forest, 

with occasional bogs and shrub stands. 

At bogs, frequency and cover were measured. A t forest 

sites, the vegetation was sampled separately in three different 

strata. For trees, densities and basal areas were measured; for 

saplings and arborescent shrubs, densities and frequencies; for 

understory, frequencies in 1 m2 quadrats. The data were pro- 

cessed by computer. Bray and Curtis ordination was used to 

interpret the relations between the sites, and a clustering 

program was used to group the sites into sites according to 

similarity. 

Data for the tree layer were used to divide the sites into 

four types, arbitrarily called Types 1-4. Understory data were 

similarly used to identify four site types, called Types a-d. 

.~.. . ~ . 

These were rela-ted to the-site typesfo-r trees; Forall-types, 

Metrosideros collina was the dominant tree; ~heirodendron 

trigynum was usually second. 

Type 1 included the lowest sites, from about 1670 m 

(5500 ft), near the Park boundary on Kalapawili Ridge. These 

were poorly drained, with scrubby forest alternating with open 

bogs. The tree layer consisted mainly of low (ca. 4 m) Metrosideros 

collina with a few Cheirodendron trigynum and - Ilex 

a n o m a 1 a . n tree species, though present, seldom reached full 

tree size. Tree density, basal area, and cover were low compared 

to the other types. The understory in some spots was transitional 

to bogs, at other spots it was of the type described below 

as Type a. 

Type 2 occurred on better drained uplands, mainly from 

1710 m to 2010 m (5600-6600 ft) on Kala~awili Ridse. The tree 

layer consisted mainiy of 5 m to 9 m tall '~etrosideios collina. 

Cheirodendron trigynum 

+- 

or, less frequently, Ilex anomala were 

second in density. M rsine lessertiana was lessxndant than at 

Type 3 sites. The ar orescent shrub layer consisted of Coprosma

SPP . r Pelea sp., and Vaccinium calycinum. Where the tree layer 

was open, a layer of large bushes consisting . of ~roussaisia 

arguta, clermontia sp., Labordia sp. , and, in places, Rubus 

hawallensis was often present. The understory was mainly Type a. 

At higher elevation sites the understory was transitional to Type 

Type 3 sites occurred on well drained uplands from 2010 rn to 

2070 m (6600-6800 ft) along Kalapawili Ridge, and in most of the 

Ko'olau sites. The tree layer was 5 m to 8 m tall along Kalapawili 

Ridge and 8 m to 13 m tall in the Ko'olau Forest Reserve 

(KFR). The basal areas were generally greater, and the canopies 

more closed, than in Type 2 sites. Metrosideros collina was the 

dominant tree, followed by ~heirodenaron trigynum or, less often, 

Myrsine lessertiana. - Ilex anomala was less abundant than at 

Type 2 s m o s m a spp., Pelea spp., and Vaccinium calycinum 

often reached t-. The =story was more open and poorer 

in species than at Type 2 sites. ~stelia sp., ~lermontia sp., 

Gouldia terminalis, Labordia sp., and Phyotegia sp. were 

uncommon or absent. Pteris excelsa, Carex alllgata, and Rubus 

hawaiiensis were more common than at Type 2 sites. At ~alapawili 

Ridge Type 3 sites, the understory was generally Type b. At 

lower KFR Type 3 sites, the understory was ~ y ~ e or~~ype 

- b d; at 

upper KFR Sites, Type c. 

Type 4 included the uppermost sites on Kalapawili Ridge, 

from 2060 m to 2110 m (6800-6900 ft). The canopy was low, ca. 

5 m, and dense. The tree layer consisted of Metrosideros collina 

with occasional Cheirodendron trigynum. Other tree species 

occur, but do not reach tree size. Small Metrosideros and 

Coprosma spp. were particularly abundant in the sapling and arbo- 

rescent shrub layer. The understory was an open version of 

Type b. 

Common to all four types of understory recognized were 

Athyrium spp. (incl. - A. microphyllum and A. sandwicianum), Dryopteris 

spp. (incl. - D. labra D. hawaiiensrs, and D. wallichiana), 

~ o g l o s s u m hirfum%'E~ wawrae, ~o!ypodiiini pellucidum, 

Sadleria sp., Unclnia uncinaFa, and Peperomia spp. 

Type a, usually associated with a Type 1 or Type 2 tree 

layer, was both the densest and richest in species. Typical of 

~ype a sites was high frequencies of Asplenium spp.: Astelia 

spp., Gouldia terminalis, and Myrsine lessertiana seedlings. 

Type b, generally associated with a Type 3 tree layer and a 

more closed canopy, was poorer in species and usually more open 

than Type 2. Carex alligata and Rubus hawaiiensis were present 

more often than in Type a. 

Type c, generally associated with Type 3 tree layer at 

higher elevation KFR sites, was more open and poorer in species 

than the other three types. The presence of Pteris excelsa, 

Carex alligata, and Rubus hawaiiensis was characteristic.

Type d was a catch-all category consisting mainly of lower 

elevation sites occurrina under various tvDes of tree lavers. 

Their main similarity is p;esence of Carex aliigata, Ilex anbmala 

seedlings, Nertera granadensis, Styphelia tameiamelae, and 

Vaccinium berber ifolium. 

In general, understories at lower elevation sites have more 

species than higher elevation sites. There are few species 

typical of higher elevation sites. Rather than turnover along 

the gradient, the trend is for species to drop out with in- 

creasing elevation, with few new species coming in. For tree 

species, the trend is different: More arborescent species reach 

tree size at the higher elevation sites than at the lower ones. 

At the time of the study in 1973, there were few if any 

exotic plants in the understory. Feral pigs were almost absent 

from Kalapawili Ridge from between 1710 m to 2200 m (5550- 

7200 ft), although present both above and below those elevations, 

and in KFR. Since 1973, pigs have become much more common along 

Kalapawili Ridge. In 1978, the 1973 sites were relocated, 

marked, and resampled in order to evaluate the pigs' effects on 

the vegetation. Effects seem small so far; the situation will be 

monitored to observe any changes as they develop.

PREHISTORIC HAWAIIAN BIRDS * 

Alan C. Ziegler 

Department of Vertebrate Zoology 



Until recently, the prehistoric Hawaiian avifauna was known 

only by the 1926 find of an extinct goose under 25 m of lava at 

Pahala on Hawai'i. Since 1971, however, remains of 20 or more 

previously unknown prehistoric Hawaiian bird taxa have been 

recovered from Moloka'i and Kaua'i (windblown sand dunes), Maui 

(lava tube), and O'ahu (solution pits in raised limestone reef). 

A radiocarbon age of 26,000 years for a Moloka'i goose skeleton 

is the only date presently available. 

A remarkable flightless component of this extinct avifauna 

comprises several geese and rails, and the first known flightless 

ibis. Yet-undescribed flighted birds include eagle, owl, and 

raven, as well as a variety of finch-like passerines. Hawaiian 

Hawk, Nene, Hawaiian Duck, and Chaetoptila (a meliphagid), or 

closely related forms, apparently occurred contemporaneously. 

Present-day native wading and marsh birds are relatively scarce 

or absent in the prehistoric deposits. Remains of modern Dre- 

panididae seem lacking in all sites except those of Kaua'i, pos- 

sibly because of a relatively more recent date for these latter 

deposits. 

Absence of terrestrial predators originally allowed survival 

of flightlessness in Hawai'i, and lowered metabolic requirements 

of the flightless individuals constituted a selective advantage. 

Evolution of flightlessness in the Islands probably represents 

neoteny, rather than the more common long-term incremental selec- 

tive process. Time and cause of extinction of this prehistoric 

avifauna is unknown but, although no evidence has thus far been 

found, it is quite possible that original Polynesian settlers or 

their associated animals were involved. 

* Abstract

LIST OF PARTICIPANTS 

Ken Adee, U. S. Forest Service 

Gregory A. Ahearn, Zoology Dept., Univ. of Hawaii at Manoa 

Jayne N. Ahearn, Genetics Dept., Univ. of Hawaii at Manoa 

Rothwell K. Ahulau, Univ. of Hawaii at Manoa, 

and Hawaii Institute of Marine Biology 

Barbara F. Allen 

Suzy Allen, Hawaii Volcanoes National Park 

Russell A. Apple, National Park Service, Honolulu 

Ken Baker, Hawaii Volcanoes National Park 

Winston E. Banko, Hawaii Volcanoes National Park 

Bob Barbee, Hawaii Volcanoes National Park 

Bob Barrel, National Park Service, Honolulu 

Carmen M. Baybayan, Univ. of Hawaii at Hilo 

John W. Beardsley, Entomology Dept., Univ. of Hawaii at Manoa 

Marc A. M. Bell, Biology Dept., Univ. Victoria, B.. C., Canada 

Tim J. Bertrand, U. S. Fish & Wildlife Service 

Theodore P. Bodner, U. S. Fish & Wildlife Sevice 

Dawn Breese, U. S. Forest Service 

Paul L. Breese, SRCF Kapa'au 

Kent W. Bridges, Univ. of Hawaii at Manoa 

Gerald D. Carr, Botany Dept., Univ. of Hawaii at Manoa 

Hampton L. Carson, Genetics Dept., Univ. of Hawaii at Manoa 

Meredith S. Carson 

John G. Chan, Biology Dept., Univ. of Hawaii at Hilo 

Gar Clarke, Hawaii Volcanoes National Park 

Mark S. Collins, U. S. Forest Service 

Patrick Conant, Entomology Dept., Univ. of Hawaii at Manoa 

Sheila Conant, General Science Dept., Univ. of Hawaii at Manoa 

Carolyn A. Corn, Univ. of Hawaii at Manoa, 

and Hawaii Division of Forestry 

Lisa K. Croft, Botany Dept., Univ. of Hawaii at Manoa, 

and U. S. Forest Service 

Linda W. Cuddihy, Botany Dept., Univ. of Hawaii at Manoa 

Gordon Y. Daida, Botany Dept., Univ. of Hawaii at Manoa 

Bertell D. Davis, Univ. of Hawaii at Manoa, 

and Archaeological Research Center Hawaii 

Clifton J. Davis, Hawaii Volcanoes National Park 

Joyce A. Davis, Botany Dept., B. P. Bishop Museum 

Isa Degener, New York Botanical Garden 

C. H. Diong, Zoology Dept., Univ. of Hawaii at Manoa 

Ann E. Dunmire, Cooperative Park Studies Unit, Univ. of Hawa 

Jon W. Erickson, Hawaii Volcanoes National Park 

William E. Evenson, Botany Dept., Univ. of Hawaii at Manoa, 

and Dept. of Physics & Astronomy, Brigham Young University 

-

List of Participants (Continued) 

Dennis B. Fenn, National Park Service, San Francisco 

Evangeline J. Funk, Botany Dept., Univ. of Hawaii at Manoa 

Peter C. Galloway, Univ. of Hawaii, Cons. Council for Hawaii 

Donald E. Gardner, Hawaii Volcanoes National Park 

Ruth A. Gay, Botany Dept., Univ. of Hawaii at Manoa 

Grant Gerrish, Botany Dept., Univ. of Hawaii at Manoa 

Jon G. Giffin, State Division of Fish & Game 

M. Lee Goff, Entomology Dept., B. P. Bishop Museum 

Samuel M. Gon, 111, Zoology Dept., Univ. of Hawaii at Manoa 

Arnold H. Hara, Entomology Dept., Univ. of Hawaii at Manoa 

Duane D. Harding, Mauna Loa Observatory 

D. Elmo Hardy, Entomology Dept., Univ. of Hawaii at Manoa 

Dennis Hashimoto, Hawaii Volcanoes National Park 

Mary M. Helfrich, Lyman House Museum 

Dorothy S. Henderson, Univ. of Hawaii at Manoa 

Derral R. Herbst, U. S. Fish & Wildlife Service 

Paul K. Higashino, U. S. Forest Service 

Stephen A. Holmes, Hamakua District Development Council 

Beverly M. Hookano, Univ. of Hawaii at Hilo 

Marcia E. Horner, Entomology Dept., Univ. of Hawaii at Manoa 

Frank G. Howarth, Entomology Dept., B. P. Bishop Museum 

James D. Jacobi, Botany Dept., Univ. of Hawaii at Manoa, 

and U. S. Fish & Wildlife Service 

Francis H. Jacot, National Park Service, San Francisco 

Terrell J. Jones, Cooperative Park Studies Unit, Univ. of Hawaii 

Dina Kageler, Hawaii Volcanoes National Park 

Seiso Kamimura, Dept. of Land & Natural Resources, 

Division of State Parks 

Michael W. Kaschko, Anthropology Dept., Univ. of Hawaii at Manoa 

Larry K. Katahira, Hawaii Volcanoes National Park 

Cameron B. Kepler, U. S. Fish & Wildlife Service 

Bruce M. Kilgore, National Park Service, San Francisco 

John I. Kjargaard, Haleakala National Park 

Charles H. Lamoureux, Botany Dept., Univ. of Hawaii at Manoa 

Ah Fat Lee, Wildlife Branch, Fish & Game, State of Hawaii 

Barbara Lee, Wildlife Branch, Fish & Game, State of Hawaii 

H. Franklin Little, Univ. of Hawaii at Hilo 

Marcia H. Little 

Tim K. Lowrey, Botany Dept., Univ. of California - Berkeley 

Anna Manis, The Nature Conservancy 

Doris Mann 

Herber J. Mann, National Oceanographic and Atmospheric Admin. 

Marcia May, Univ. of Hawaii at Hilo 

Phyllis H. McEldowney, U. S. Fish & Wildlife Service 

Hans Megens, B. P. Bishop Museum 

Christina B. Meller, Life of the Land 

Douglas Meller, Shoreline protection Alliance


Mark D. Merlin, General Science Dept., Univ. of Hawaii at Manoa 

John M. Miller, Mauna Loa Observatory 

Shirlene S-L. Miyashiro, Dept. of Land & Natural Resources, 


Geary S. Mizuno, Zoology Dept., Univ. of Hawaii at Manoa 

Steve L. Montgomery, Entomology Dept., univ. of Hawaii at Manoa 

Richard B. Moore, USGS--Hawaiian Volcano Observatory 

Christine E. Morgan, U. S. Forest Service 

Dieter Mueller-Dombois, Botany Dept., Univ. of Hawaii at Manoa 

Mae E. Mull, Hawaii Audubon Society 

William P. Mull, B. P. Bishop Museum 

Gail M. Murakami, Botany Dept., Univ. of Hawaii at Manoa 

Timothy G. Myles, Univ. of Hawaii at Manoa 

Jean I. Nishida, Dept. of Land & Natural Resources, 


Timmy J. Ohashi, U. S. Forest Service 

Gregory P. Owen, Hawaii Outward Bound School 

Richard P. Papp, B. P. Bishop Museum 

Carol Pearson, U. S. Forest Service 

Dana R. Peterson, Univ. of Hawaii at Manoa, 

and Haleakala National Park 

Lawrence J. Pinter, Navy Environment Office 

C. J. Ralph, U. S. Forest Service 

Don Reeser , Hawaii Volcanoes National Park 

Faith M. Roelofs, Botany Dept., Univ. of Hawaii at Manoa 

Christa A. Russell, Hawaii Volcanoes National Park 

Howard F. Sakai, U. S. Forest Service 

Simon C. Sanidad, Entomology Dept., Univ. of Hawaii at Manoa 

Mike Scott, U. S. Fish & Wildlife Service 

Paul G. Scowcroft, U. S. Forest Service 

Richard J. Scudder, Governor's Off ice of Environmental 

Quality Control 

Deborah A. Shaw, Field Studies Unit 

Clara H. Shimoda, City of Refuge National Historical Park 

Jerry Y. Shimoda, City of Refuge National Historical Park 

Roger G. Skolmen, U. S. Forest Service 

Garrett H. Smathers, National Park Service, Biology Dept., 

Western Carolina University 

Clifford W. Smith, Botany Dept., University of Hawaii at Manoa, 

and Cooperative National Park Resources Studies Unit, U. H. 

Linda L. Smith, Botany Dept., Univ. of Hawaii at Manoa 

Jim Sorenson, Botany Dept., Univ. of Hawaii at Manoa 

Wayne H. Souza, Dept. of Land & Natural Resources, 


John D. Stein, U. S. Forest Service 

Lani Stemmermann, Botany Dept., Univ. of Hawaii at Manoa, 

and B. P. Bishop Museum 

Maile A. Stemmermann, Zoology Dept., Univ. of Hawaii at Manoa


Carol S. Tabata, Coastal Zone Management, Univ. of Hawaii 

at Manoa 

Raymond S. Tabata, Sea Grant, Univ. of Hawaii at Manoa 

Kim0 Tabor, The Nature Conservancy 

Howard A. Takata, Sea Grant, Univ. of Hawaii at Manoa 

Avery L. Taylor, U. S. Forest Service 

JoAnn Tenorio, B. P. Bishop Museum 

Glenn I. Teves, Entomology Dept., Univ. of Hawaii at Manoa 

P. Quentin Tomich, State Health Department 

Charles van Riper 111, Univ. of Hawaii, and Cooperative Park 

Studies Unit, Univ. of Hawaii 

Sandra G. van Riper, Cooperative Park Studies Unit, Univ. 

of Hawaii 

Jerry A. Walters, U. S. Forest Service 

John F. Walters, Oceanography Dept., Univ. of Hawaii at Manoa 

Richard E. Warner, Field Studies Unit 

Rich Warshauer, U. S. Fish & wildlife Service 

Deborah A. Weiner, Cooperative Park Studies Unit, Univ. of Hawaii 

Art Whistler, Botany Dept., Univ. of Hawaii at Manoa 

Louis D. Whiteaker, Cooperative Park Studies Unit, 

Univ. of Hawaii 

Claire M. Wolfe, U. S. Forest Service 

David H. Woodside, Dept. of Land & Natural Resources 

Faye Yates 

Alvin Y. Yoshinaga, Botany Dept., Univ. of Hawaii at Manoa, 

and Cooperative Park Studies Unit, Univ. of Hawaii 

Ernest R. Yoshioka, State Dept. of Agriculture 

Chris J. Yuen, Youth Conservation Corps 

Alan, C. Ziegler, Division of Vert. Biology, B. P. Bishop Museum 

Nicholas Zimmer, Hawaii Volcanoes National Park

Acacia koa, 260, 333 

Acari, m, 237 

Acid Rain, 217 

'Alala, 207 

Animals, 198 

, 42 

SUBJECT INDEX 

Biological, 26 

Bird, 52, 199, 207, 309, 345 

Birds, 6, 17, 59, 71, 125, 193 

238, 243, 349 

Communications Techniques, 256 

Ecosystem, 58 

Ecosystem Restoration, 2 

Endangered, 17 

Endangered Forest Birds, 

Endangered Plants, 37 

Endangered Species, 208 

Exotic Plants, 120, 198 

Exotics, 51 

Geology, 1, 218 

Haleakala, 30, 71, 193, 289, 

292, 297, 337, 346 

Hana Rain Forest, 346 

Hawaii Volcanoes National Park, 

51, 98, 125 

Hawaiian Dark-rumped Petrel, 193 

Hawaiian Forests, 333 

Hibiscadelphus, 2 

Human Settlement, 87 

Hybridizations, 37, 77 

Insect, 134, 236 

Insects, 30, 41, 54, 98, 235, 237 

Island Ecosystems, 231 

Kalapana, 59 

Kilauea Rain Forest, 58 

Kilauea Volcano, 218 

- 

Lava Tube, 155 

Leptospirosis, 

Lichen, 292 

Limiting, 17 

Mamane, 247 

Mammals, 2,237,308 

Mauna Loa. 222.237 

Metrosideros, 77 

Mosses, 150 

Myrica faya, 51, 114, 274 

Natural Areas, 304 

Necropsy, 309 

Nene, 6, 199 

O'ahu, 52, 87, 120 

'Ohi'a, 105, 236 

Planting, 239 

Plants, 297 

Pohakuloa, 199 

Propagation, 260 

Rat, 2 

Regeneration Technique, 247 

Reintroduction, 6 

Rodents, 237 

Seismology, 1 

Shell Disease, 42 

Shrimp, 42 

Silversword, 37 

South Kona, 86, 134 

Spiders, 235 

Sweet Potato, 177 

Temporal Patterns, 34 

Threatened Plants, 86 

Variation, 41 

Vegetation Map, 165, 337 

Weed, 26 

Whitney, 1

June 1 - 3 , 1978 - University of Hawaii at Manoa

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