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PhytoKeys 4: 5–51 (2011)
New pteridophyte species and combinations from the Marquesas Islands...
doi: 10.3897/phytokeys.4.1602
reseArCh ArtICle
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Launched to accelerate biodiversity research
New pteridophyte species and combinations from the
Marquesas Islands, French Polynesia
David H. Lorence1,Warren L. Wagner2, Kenneth R. Wood1, Alan R. Smith3
1 National Tropical Botanical Garden, 3530 Papalina Road, Kalaheo, HI 96741–9599 2 Department of
Botany, MRC–166, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC 20013–7012 3 1001 Valley Life Sciences Building, no. 2465, University Herbarium, University
of California, Berkeley, CA 94720–2465
Corresponding author: David H. Lorence (lorence@ntbg.org)
Academic editor: Vicki Funk | Received 26 May 2011 | Accepted 15 June 2010 | Published 12 July 2011
Citation: Lorence DH, Wagner WL, Wood KR, Smith AR (2011) New pteridophyte species and combinations from
the Marquesas Islands, French Polynesia. In: Lorence DH, Wagner WL (Eds) Botany of the Marquesas Islands: new taxa,
combinations, and revisions. PhytoKeys 4: 5–51. doi: 10.3897/phytokeys.4.1602
Abstract
Intensive botanical exploration of the Marquesas Islands (French Polynesia) for the Vascular Flora of the
Marquesas Islands and Flore de la Polynésie française projects has resulted in numerous additional new
collections from these islands. Study of these collections has brought to light 11 new species of pteridophytes (ferns and lycophytes) which are described herein: Blechnum pacificum Lorence & A. R. Sm., sp.
nov., Cyclosorus castaneus A. R. Sm. & Lorence, sp. nov., Cyclosorus florencei A. R. Sm. & Lorence, sp.
nov., Dryopteris macropholis Lorence & W. L. Wagner, sp. nov., Dryopteris sweetorum Lorence & W. L.
Wagner, sp. nov., Polystichum kenwoodii Lorence & W. L. Wagner, sp. nov., Polystichum uahukaense Lorence & W. L. Wagner, sp. nov., Pteris hivaoaensis Lorence & K. R. Wood, sp. nov., Pteris marquesensis
Lorence & K. R. Wood, sp. nov., Pteris tahuataensis Lorence & K. R. Wood, sp. nov., and Thelypteris
marquesensis Lorence & K. R. Wood, sp. nov. One new combination is made: Cyclosorus marquesicus
(Holttum) Lorence & A. R. Sm., comb. nov. (based on Plesioneuron marquesicum Holttum). An analysis
of the conservation status of these new Marquesas Islands taxa reveals they are in need of inclusion in the
IUCN Red List with conservation status ranging from vulnerable (one species), and endangered (four
species), to critically endangered (five species).
Keywords
Blechnum, Cyclosorus, Dryopteris, French Polynesia, Marquesas Islands, Polystichum, pteridophytes, Pteris,
Thelypteris
Copyright David H. Lorence et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Introduction
The only recent treatment of the Marquesan pteridophytes (=ferns and lycophytes,
used interchangeably herein) is Forrest and Elizabeth Brown’s Flora of Southeastern Polynesia (Brown & Brown 1931). This is essentially a report on the plants collected by the Browns on the Bayard Dominick Expedition (1921–1922), by E. H.
Quayle, B. W. Jones, and R. Beck on the Whitney Expedition (1921–1922), and
by the Pacific Entomological Survey (1929–1932) (Lorence and Wagner 1997).
The Browns’ flora did not include earlier collections not then represented in the
Bishop Museum, except as they may have been recorded in the French Polynesian
flora of Drake del Castillo (1892). In his Flore de la Polynésie Française Drake
del Castillo recognized 142 fern and lycophyte species from the region, three of
which were described as new. Brown and Brown (1931) recorded 103 species from
southeastern Polynesia, or more specifically 122 taxa including varieties and forms
of which 59 (48%) were described as new. Although the Browns’ work suffers from
a number of shortcomings such as non-parallel descriptions, atypical typifications,
and errors in identification, it has remained the only modern pteridophytes flora
available for the Marquesas.
Copeland (1932) subsequently published a floristic treatment of the Society Islands
pteridophytes that included 164 species from this archipelago, but did not include the
Marquesas Islands. Most recently, Kato et al. (2008) published an illustrated flora of
ferns and lycophytes of the South Pacific islands, encompassing Samoa, Fiji, Vanuatu,
and New Caledonia. Revisions of only a few taxonomic groups occurring in French
Polynesia and the Marquesas have been published to date, e.g. Elaphoglossum (Rouhan
et al. 2008) and Hymenophyllaceae (Ebihara et al. 2006). Useful identification guides
include an online key and pictorial checklist of the pteridophytes of Moorea, available
at http://ucjeps.berkeley.edu/moorea/pteridophytes.html. A comprehensive published
treatment of Marquesan pteridophytes is clearly needed.
This paper increases our knowledge of the Marquesan pteridophyte flora by describing 11 new species and proposing one new combination. The present contribution forms part of a series of precursor publications with the goal of producing the first
complete Vascular Flora of the Marquesas Islands (Lorence and Wagner 1997). Currently available online as a Web-based flora with a searchable database of descriptions,
photos, literature and specimens, it may be accessed at http://botany.si.edu/pacificislandbiodiversity/marquesasflora/index.htm. This project will provide a comprehensive
treatment of all Marquesan pteridophytes, for which the essentially completed treatment is now available on the website. A primary goal is to publish the Vascular Flora of
the Marquesas Islands as a two volume printed work. A second goal involves collaboration on the Flora of French Polynesia project headed by Jacques Florence under the
auspices of the French Institute pour la Recherche et Developpement (IRD, formerly
ORSTOM). To date only two volumes of the modern Flore de la Polynésie Française
have been published (Florence 1997, 2004), although Dr. Florence is currently finalizing a field guide to the French Polynesian pteridophytes.
New pteridophyte species and combinations from the Marquesas Islands...
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Island Names
Orthographic variation exists for certain of the Marquesas Islands. For the sake of
consistency we herein utilize the names accepted by the French Polynesian Government (see website at: www.presidence.pf ) for the islands. In the following list accepted
names are in boldface and alternative spellings are listed in parentheses: Fatu Hiva
(Fatuhiva, Fatu Iva), Hiva Oa (Hivaoa), Mohotani (Motane), Nuku Hiva (Nukuhiva), Tahuata, Ua Huka (Uahuka), and Ua Pou (Uapou).
Conservation status
As the Marquesan environment is under serious threat from human impacts, feral
animals, and weeds (Florence and Lorence 1997) the conservation status was estimated for each new species. When evaluated using the IUCN criteria for endangerment
(IUCN 2001, see also www.iucnredlist.org/info/categories_criteria2001), all but one
of the new Marquesan pteridophyte species fall into the Endangered (EN) or Critically Endangered (CR) categories, which designates species facing the highest risk of
extinction in the wild. One species, Blechnum pacificum which occurs on five Marquesan islands, three Society Islands, and Rapa Iti in the Austral Islands, is considered
Vulnerable (VU). The IUCN Endangered (EN) criteria include: B1having known
ranges less than 5000 km2; B2 an area of occupancy of less than 500 km2; a, b severely
fragmented or known to exist at no more than five locations; c continuing decline in
the quality of habitat; or D, a populations size less than 250 mature individuals. The
IUCN Critically Endangered (CR) criteria include: B1 having known ranges of less
than 100 km2 and/or B2, an area of occupancy of less than 10 km2; D, population size
of less than 50 mature individuals; and estimates including at least two of the following: a, severely fragmented or known to exist at only a single location; b, continuing
decline in the extent and/or quality of habitat, extent of occurrence or occupancy, or
number of mature individuals.
Methodology
All measurements given herein are taken from dried herbarium specimens, although
certain features such as shapes were supplemented with information from field notes
and photos. Measurements are presented in the descriptions as follows: length × width,
followed by units of measurement (mm or cm). All specimens cited in this paper have
been seen by the authors. Specimens from the following herbaria were studied: AD,
BISH, BR, K, MO, NY, P, PAP, PTBG, and US. The area of occupancy (distribution)
was calculated using herbarium collection data and field observations, and the conservation status is proposed following the IUCN Red List Category criteria (IUCN 2001;
www.iucnredlist.org/info/categories_criteria2001).
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
systematics
BLECHNACEAE
Blechnum
Blechnum L. is a cosmopolitan genus of about 180 species especially well represented in
the southern hemisphere (Mabberley 2008). Brown and Brown (1931) recorded three
species from the Marquesas Islands: the endemic B. nukuhivense E. D. Br., the rather
widespread Pacific species B. vulcanicum (Bl.) Kuhn, and B. capense (L.) Schltdl. However, it is apparent that the latter name has been misapplied to Polynesian plants, which
we describe below as a new species. The three Marquesan Blechnum can be separated
by the following key.
Key to Blechnum in the Marquesas Islands
1a
1b
2a
2b
Sterile fronds 1-pinnate, nearly all the pinnules contracted at their bases and
free from rachis, the apex with a conform terminal pinna similar to lateral
pinnae.......................................................................................B. pacificum
Sterile fronds entire, pinnatifid, or pinnatisect with the segments adnate and
not contracted at their bases, or rarely the basal segments pinnate and the
basiscopic base contracted and not adnate, the apex lacking conform terminal
pinna ..........................................................................................................2
Sterile fronds entire or sometimes irregularly pinnatifid toward middle and
base...................................................................................... B. nukuhivense
Sterile fronds pinnatisect or rarely the basal segments pinnate ..B. vulcanicum
1. Blechnum pacificum Lorence & A. R. Sm., sp. nov.
urn:lsid:ipni.org:names:77112673-1
http://species-id.net/wiki/Blechnum_pacificum
Figs 1, 2, 14A
Species Blechno venoso Copel. affinis, sed stipitis squamis non tan numerosis, deciduis,
leviter castaneis usque ad castaneis, sterilibus pinnis cum axialibus glabris et paulatim
prominentibus venis, fertilibus pinnis cum viridi expansa textura prope basin differt.
Type. Marquesas Islands: Ua Pou: Poumaka Summit Trail, 690 m elevation,
9°23'33"S, 140°04'59"W, 19 June 2004, L. M. Dunn and D. H. Lorence 481 (Holotype: PTBG-041866!, PTBG-041867! [2 sheets]; Isotypes BISH!, P!, PAP!, US!).
Blechnum capense sensu E. D. Br. & F. B. Br., non Burm. f., non (L.) Schltdl.
Description. Large terrestrial ferns; rhizomes erect or suberect, rarely decumbent
and dorsiventral, short, stout, (10–)15–20 mm in diameter (excluding scales), apex
covered with scales; scales of rhizome and bases of stipes linear-subulate to oblongovate, 20–30 ×1–3(–5) mm, thin, light brown to brown, concolorous but somewhat
New pteridophyte species and combinations from the Marquesas Islands...
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Figure 1. Blechnum pacificum Lorence & A. R. Sm. A sterile frond, blade B sterile frond, stipe C sterile
frond, pinna base. Drawn from the type collection (Dunn and Lorence 481).
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Figure 2. Blechnum pacificum Lorence & A. R. Sm. A fertile frond, blade B, fertile frond, pinna base
showing expanded sterile portion C scale from base of stipe D part of scale from base of stipe, detail of
cells. Drawn from the type collection (Dunn & Lorence 481).
New pteridophyte species and combinations from the Marquesas Islands...
11
thicker and darker basally and in center, basifixed, base rounded or truncate, apex
sinuate, margins entire or subentire, cells linear, arranged in vertical rows. Fronds clustered near apices of rhizomes, usually dimorphic but both fertile and sterile pinnae
occasionally occurring on the same blade; stipes 41–102 cm long, 4–10 mm in diameter, stout, light to dark brown when fresh, drying stramineous, densely scaly toward the bases, sparsely so distally, eventually glabrescent, smooth, grooved adaxially.
Sterile blades 1-pinnate, oblong-ovate to oblong-elliptic, (35–)50–100 × (20–)30–60
cm, coriaceous to subcoriaceous, stiff, rachises stramineous or sometimes dark brown,
proximal pinnae only slightly or not at all reduced, apices acute, each blade with a conform terminal pinna, when young the rachises, costae, costules, and veins scaly with
thin, scurfy, brown to pale brown, oblong-ovate to subulate, contorted-sinuate scales
to 10 mm long with margins subentire or sometimes lacerate basally; pinnae 19–30
on a side, with a slightly swollen, mammiform aerophore at the base each pinna, basal
pinnae opposite and shortly stipitate (to 2 mm), the distal ones becoming subopposite
to alternate, sessile with basiscopic base often adnate to rachis, terminal pinna free,
conform, medial pinnae 11–30 × 1.5–2.5 cm, linear, margins serrulate, often decidedly undulate, apices acute or acuminate, serrulate, acroscopic bases oblique-cuneate,
basiscopic base truncate or rounded, the veins prominulous, simple or 1-forked, free,
each ending in a marginal tooth; fertile fronds subequal to or slightly smaller than sterile, with up to 40 pinnae pairs (sometimes the proximal pinnae sterile and distal pinnae fertile on the same frond), fertile pinnae approximately the same length as sterile
pinnae but narrower, 2.5–5 mm wide, the margins strongly revolute, the sori covering
most of the abaxial surface but usually with some expanded green tissue at the base,
adaxially glabrate. Sori linear, abaxial surface of pinnae bordered or partly covered by
the reflexed, scarious, erose blade margins except on expanded green bases. Spores subellipsoidal, 67 × 47 µm including a perispore 8–12 µm wide.
Distribution. Known from the Marquesas Islands (Nuku Hiva, Ua Pou, Hiva Oa,
Tahuata, and Fatu Hiva), Society Islands (Moorea, Raiatea, and Tahiti), and Austral
Islands (Rapa Iti).
Ecology. This new terrestrial species occurs in clearings or shade from 380 to 1500 m
elevation in lowland to montane mesic and wet forests, shrubland or fernland, in valleys,
on slopes or ridge crests and rocky banks. In the Marquesas associated species include
Crossostylis biflora J. R. Forst. & G. Forst., Dicranopteris linearis (Burm. f.) Underw., Freycinetia impavida (Gaudich. ex Hombr.) B. C. Stone, Metrosideros collina (J. R. Forst. &
G. Forst.) A. Gray, Pandanus tectorius Parkinson, Santalum insulare Bertero ex A. DC.,
and other native tree, shrub, and pteridophyte species. Threats include competition from
alien plant species and damage from feral ungulates. Although this is the most widely
distributed of the new species, it is clearly at risk due to habitat loss and degradation.
Etymology. The epithet refers to the Pacific distribution of this new species.
Conservation status. IUCN Red List Category: Vulnerable (VU): B1: total area
of occupancy less than 20,000 km2 (ca. 750 km2); b (i–iii), habitat continuing decline
inferred; B2: total area of occupancy less than 2000 km2 (ca. 750 km2); B2b (i–iii),
habitat continuing decline inferred. The suitable habitat for Blechnum pacificum on
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
most islands of occurrence is indicated as a declining or endangered environment,
threatened by human activity (deforestation, fire), feral animals, and invasive plants,
reducing the extent of the forest.
Specimens examined. Austral Islands: Rapa Iti. Perau–Mamuere summit, eastern peaks, 27°S, 144°W, 10–100 ft (3–30 m), Wood & Faraire 9777 (NY, PTBG [4
sheets]), Fosberg 11567 (UC), Fosberg 11601 (UC), Fosberg 11605 (UC); St. John &
Fosberg 15293 (UC). Society Islands: Moorea: Mt. Rotui, 149°50'18W, 17°30'38S,
916 m, Nitta & Vinette 212 (PAP, UC); Mt. Tohiea,149.819°, 17.553°, 994 m, Nitta
& Vinette 320 (PAP, UC). Tahiti. Pirae–Maoua Aorai trail, Quayle 105A (BISH,
UC); Orofena, east side of south ridge, ravine in rain forest, St. John & Fosberg 17114
(BISH); Mt. Marau road, crest between Tapaerui and Punaruu valley, 1250 m, Fosberg
62647 (PTBG [2 sheets], US); Orofena, south ridge, moist thicket on exposed ridge,
1600 m, St. John & Fosberg 17082 (UC); Mahina, Ahonu-tuauru, 2875 ft (876 m),
Grant 4394 (UC); Aorai, in summit shrubs, 6700 ft (2042 m), M. L. Grant 3791
(UC); Faaa, Mt. Marau, 3 km below TV tower, 1300 m, Hodel 1375 (UC); Fautaua,
below Diadem, 2830 ft (863 m), Grant 3546 (UC). Raiatea: Temehani Plain, Moore
175 (BISH [2 sheets]). Marquesas Islands: Nuku Hiva: Toovii, 1000 m, Brown &
Brown 528 (BISH); Toovii Plateau, spur of Mt. Ooumu, 790 m, Gagné 1039 (US);
Tauamaka, Toovii plateau, 1000 m, Mumford & Adamson 576 (BISH, UC); without precise locality, 1000 m, Quayle 1284 (BISH), Quayle 1298 (BISH). Ua Pou:
Mt. Tekahoipu, 800 m, Quayle 1138 (BISH [2]); Tekohepo, summit, 2500–3000 ft
(762–914 ft), 09°24'31"S, 140°04'21"W, Wood & Perlman 6455 (PTBG [2 sheets],
US); Drainage northwest of Teavahaakiti, 700 m, Wood 10458 (P, PAP, PTBG, US).
Hiva Oa: road from Atuona to Puamau, just below Ootua, 625–700 m, Sachet &
Decker 1904 (PTBG, US); Puamau, along Puamau–Atuona trail, 500–650 m, Decker
1190 (PTBG, US); Feani, 800 m, Brown 877 (BISH [3]); Atuona–Feani Trail, ridge
crest, 1200–1300 m, 24–26 Sep. 1963, Sachet & Decker 1160 (US [2]); Montagnes
NW du Temetiu, entre la haute vallée de Hanamenu et la crête de Temetiu–Feani, 850
m, Schäfer 5932 (US); Vaiata, NW slopes of Mt. Ootua, 800 m, Mumford & Adamson 355 (BISH); Feani ridge to upper slopes of dry side of island, 1150 m, Oliver &
Schäfer 3130 (US [2]); Temetiu, 1100 m, Pacific Entomol. Surv. 156 (BISH); above
Atuona, 800 m, Pacific Entomol. Surv. Ex 355 (BISH); Hanaiapa, 700 m, Jones 1613
(BISH [2 sheets]); without precise locality, 800 m, Brown 16 (BISH). Tahuata: de
Hamatea [Amatea] à la crête centrale de l’île, 750–850 m, Thibault 64 (US); Mt. Amatea, 1000 m, Jones 1796 (BISH). Fatu Hiva: ‘Omo’a–Ouia–Mounanui Trail, 690 m,
Gagné & Montgomery 2323 (BISH); trail from ‘Omo’a along Punaitai ridge crest to
base of Tekou peak, 550–840 m, Lorence et al. 6171 (BISH, PAP, PTBG); Hanavave,
600 m, Jones 1826 (BISH); Sentier d’Ouia, W du col, lieu-dit Tahuna, 620 m, Schäfer
5803 (US).
Discussion. The name Blechum capense Burm. f. has been erroneously applied to
this new Polynesian species (Brown and Brown 1931; Copeland 1932). The true Blechnum capense is confined to southern Africa and some nearby islands (Burrows 1990;
Roux 2001). This southern African species has also been treated under the name B. syl-
New pteridophyte species and combinations from the Marquesas Islands...
13
vaticum Schelpe (Schelpe 1979; Jacobson 1983), on the false assumption that the type
of B. capense Burm. f. was a mixed collection of two species (Jacobson 1983). However,
there seem to be good reasons for placing B. sylvaticum in synonymy under B. capense
(Roux 1982; Schelpe and Anthony 1986; Burrows 1990; Roux 2001).
The name Blechnum procerum G. Forst. has also been incorrectly applied to the
Polynesian plants. Quoting Nicholson and Fosberg (2004: 125–126), who stated that
the type of Blechnum procerum is from New Zealand: “Tindale (1960b: 254) published
a photo (t. 7) of the Goettingen material [Nova Zeelandia, Forster 295, GOET] as
‘Type specimen’ and referred to BM and K specimens as ‘Forster material.’ Chambers and Farrant (1998a: 4), without discussion, said ‘T. Noua Zeelandia, Forster;
lecto (here chosen): K; isolecto: BM, GOET (photo seen).’” Nicolson and Fosberg
(2004: 126) further stated “There is considerable confusion about the names applied to
this taxon and the following quotation from Brownsey and Smith-Dodsworth (2001)
summarized it well: ‘The name Blechnum minus was used incorrectly by Allan (1961)
and Crookes (1963) for the plant we call B. procerum. The true B. minus, or swamp
kiokio, is only doubtfully distinct from the common kiokio (previously known as B.
capense).’” Blechnum procerum has lower pinnae only slightly reduced (not less than
half the length of the median pinnae), fertile fronds are up to 50% longer than sterile
fronds, and the fertile pinnae show no expanded green tissue at the base in the type as
in B. pacificum. Clearly, the name B. procerum does not apply to the Polynesian plants.
In New Zealand, closest relatives appear to be B. novae-zelandiae T. C. Chambers
& P. A. Farrant and B. minus (R. Br.) Ettingsh., neither of which is conspecific with our
Polynesian plants. Blechnum novae-zelandiae is superficially similar but differs in having reduced proximal pinnae and distinctive “black-spot” rhizome and stipe scales with
dark brown or black centers and pale margins (Chambers and Farrant 1998). From
evidence presented in a recent paper on the phylogeny of New Zealand Blechnaceae
by Shepherd et al. (2007, Fig. 2), it seems likely that if Blechnum pacificum were sampled, it would fall somewhere in the clade containing B. wattsii Tindale and B. novaezelandiae.
Several species from New Caledonia, all considered endemic, form a confusing array of species somewhat similar to B. pacificum. These include B. confusum (E. Fourn.)
Brownlie, B. chauliodontum Copel., and B. subcordatum (E. Fourn.) Brownlie. Brownlie’s (1969) illustration and characterization of B. subcordatum suggests that it differs in
having smaller fronds with less scaly stipes and rachises, fewer pinnae pairs (5–15), and
sterile pinnae not so undulate at the margins. Blechnum confusum differs in its strongly
ascending, more sharply serrulate and less scaly pinnae (sterile blades are nearly glabrous). The closest species in Malesia (excluding Papua New Guinea) appears to be
B. vestitum (Blume) Kuhn, nom. cons., non B. vestitum T. Moore (see Chambers and
Farrant 2001; Chambers 2004; McNeill et al. 2006: 438). The Papua New Guinea
species B. dilatatum (Brause) T. C. Chambers & P. A. Farrant is similar to B. pacificum
in having fertile pinnae with an expanded basal sterile region, but in the latter the
margins are never revolute and the rhizome scales are thin, concolorous, and pale to
medium brown. No names of taxa with types from Fiji apply to the new species, the
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
closest species there being B. milnei (Carruthers) C. Chr. (historically also called B. procerum), which differs in having very large fronds with generally broader, less coriaceous
pinnae and a less scaly rachis and costae on the sterile blades, and fertile pinnae lacking
expanded green tissue at the base.
Among the Polynesian species of Blechnum, B. pacificum seems most closely related
to B. venosum Copel. from Rapa Iti in the Austral Islands. In addition to having copious, shiny, dark brown, almost blackish scales on the stipes as noted in the diagnosis,
the veins of B. venosum are very prominent and strongly raised above the surface on
the abaxial side of blades, whereas the veins in B. pacificum are visible abaxially but
scarcely, if at all, raised. This gives B. venosum a much harsher, more cartilaginous
appearance. Also, in B. venosum, there are very short hairs on the veins abaxially and
some hairs are even present between the veins on laminar tissue, but B. pacificum lacks
such hairs. In B. venosum, some of these hairs on the veins and laminar tissue appear
multicellular (septate, but uniseriate), and glandular or gland-tipped. Also, pinna margins in B. pacificum are more crenulate (scalloped) than in B. venosum which has entire
margins.
DRYOPTERIDACEAE
Dryopteris
Dryopteris Adanson is a large, essentially cosmopolitan genus of around 225 species
with its greatest diversity in north temperate regions (Fraser-Jenkins 1986a, 1989;
Mickel and Smith 2004). Previously only a single species, Dryopteris fatuhivensis E.
D. Br., was recorded from the Marquesas (Brown and Brown 1931), where it occurs
on Nuku Hiva, Fatu Hiva, and Ua Huka. Dryopteris fatuhivensis was placed in subg.
Dryopteris, sect. Hirtipedes by Fraser-Jenkins (1986b) along with related species from
Asia, including the widespread D. hirtipes (Blume) Kuntze which includes two subspecies, subsp. hirtipes and subsp. atrata (Kunze) Fraser-Jenk., both in southeast Asia. We
follow Fraser-Jenkens (1989) in recognizing D. fatuhivensis as a Marquesan endemic
species distinct from D. hirtipes.
Recent collections from the Marquesas have revealed the presence of two additional, distinctive endemic Dryopteris species distinguished by their large, 3-pinnate
to 3-pinnate-pinnatifid fronds. Both species have normally developed spores and lack
any morphological features suggesting hybrid origin. Morphologically they are quite
different than D. fatuhivensis, from which they may be separated by the characters in
the following key.
Key to Dryopteris in the Marquesas Islands
1a
1b
Blades 1-pinnate to 1-pinnate-pinnatifid ..............................D. fatuhivensis
Blades 3-pinnate to 3-pinnate-pinnatifid ....................................................2
New pteridophyte species and combinations from the Marquesas Islands...
2a
2b
15
Ultimate pinnule segments spaced 5–10 mm distant with margins serrate,
segments acute at apex ............................................................D. sweetorum
Ultimate pinnule segments spaced 3–5 mm distant, margins crenate or lobed
¼ –2/3 toward costule, segments truncate or obtuse at apex ...D. macropholis
2. Dryopteris macropholis Lorence & W. L. Wagner, sp. nov.
urn:lsid:ipni.org:names:77112674-1
http://species-id.net/wiki/Dryopteris_macropholis
Figs 3, 4, 14B, C, D
A ceteris marchinonicis speciebus integra margine maximis usque ad 80 × 14 mm squamis
(vel paleis) vestitis rhizomate atque stipitis base, ampla tripinnati-pinnatifida lamina, ultimis usque ad 12–19 × 4–8 mm inferne glabris pinnulis, truncatis vel crenatis, 1/4–2/3
costa dissectis lobis, sparsis parvis castaneis paleis vestita rhachidi, quaque pinnula 1–4
indusiatorum sororum paribus munita, glabro indusio, praecipue differt.
Type. Marquesas Islands: Ua Huka: Hitikau region, ascended via Matukuoha ridge
overlooking Hane, constitutes the summit of the single crater of Ua Huka, 700
m, UTM 0661697–9015668, 5 Dec 2003, K. R. Wood 10489 (Holotype PTBG041629!, PTBG-041630! [2 sheets]; Isotypes P!, PAP!, US!).
Description. Terrestrial ferns; rhizomes suberect, 20–25 cm long, 5–7 cm in diameter
(to 15 cm including scales), densely clothed with pale brown to reddish brown or dark
brown scales; scales of rhizome and base of stipe (10–)20–80 × (1–)2–5–14 mm, thin,
narrowly oblong-elliptic to linear-lanceolate, falcate, usually twisted distally, concolorous, lustrous, medium to dark brown or reddish brown, margins entire, cells narrowly
rectangular to linear-fusiform. Fronds clustered, 5–7 per rhizome, erect-arching; stipes
(35)49–75 cm long, 4–6 mm in diameter medially, about as long as the blades, adaxially grooved, reddish brown to stramineous, entire length clothed in dense, persistent,
spreading, lustrous, light to dark brown or reddish brown, linear-oblong to linear-lanceolate twisted scales to ca. 20 × 3 mm, margins entire or subentire and fringed with shortstipitate glands, bases darkened at point of attachment, mixed with smaller bristlelike
and hairlike scales, surfaces bearing short, gland-tipped hairs, scales progressively smaller
and finer distally and on rachis, stipes of older fronds punctate with dark scale bases;
blades thickly chartaceous, dark above green when fresh, paler beneath, 50–100 × 32–66
cm, ovate-deltate, 3-pinnate to 3-pinnate-pinnatifid at least in lower half, distally mostly
2-pinnate-pinnatifid; rachises stramineous to light brown, densely scaly with persistent
medium to dark brown, spreading bristlelike scales to 9 × 1 mm, margins entire or with
sparse sessile glands, mixed with short glandular hairs, rachises of older fronds punctuate with dark brown scale bases; pinnae opposite to subopposite, (11–)13–20 on a side,
spreading, ovate-oblong to linear-oblong, apex acuminate, lowermost pinnae the largest,
20–33 × 11–19 cm, with 11–16 pairs of pinnules, slightly inequilateral, basiscopic basal
pinnules 7–10.5 cm long, acroscopic basal pinnules shorter, 4–8.5 cm long, lowermost
pinnules usually the largest, distal pinnae stalked 3–6 mm becoming sessile, apices pin-
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Figure 3. Dryopteris macropholis Lorence & W. L. Wagner. Frond with A blade and B stipe. Drawn from
the type collection (Wood 10489) and field images.
New pteridophyte species and combinations from the Marquesas Islands...
17
Figure 4. Dryopteris macropholis Lorence & W. L. Wagner. A pinna from near base of lamina, lower
surface B lower surface of fertile pinnule showing sori C pinnule with sorus, sporangia, and scales D scale
from stipe e part of stipe scale showing detail of cells. Drawn from the type collection (Wood 10489)
and field images.
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
natifid; largest ultimate pinnules 12–19 × 4–8 mm, spaced 5–10 mm distant, obtuse to
truncate at apices, margins crenate or cut ca. ¼ –2/3 toward costule, adaxially glabrous,
abaxially glabrous except for scattered small, spreading, brown linear scales on rachises;
veins forking 1–2 times, scarcely visible to visible on both surfaces, depressed adaxially
and prominulous abaxially; each fertile pinnule usually with 1–4 pairs of inframedial sori.
Sori with indusia 0.4–0.6 mm in diameter, brown, thick, glabrous. Spores dark brown.
Distribution. Marquesas Islands, known from Nuku Hiva, Ua Huka, Ua Pou,
Hiva Oa, and Tahuata.
Ecology. This new species is rare and localized from ca. 700 to 1150 m elevation.
It occurs in transitional mesic to wet forests with Alsophila tahitensis Brack., Hernandia
nukuhivensis F. Br., and Sphaeropteris spp.; in wet forests dominated by Crossostylis biflora, Freycinetia spp., Hibiscus tiliaceus L., Metrosideros collina, Pandanus tectorius, and
with associates including Fagraea berteroana A. Gray ex Benth., Ficus prolixa G. Forst.
var. prolixa; Glochidion (Phyllanthus) marchionicum F. Br., Weinmannia marquesana F.
Br. var. marquesana, and Xylosma suaveolens (J. R. Forst. & G. Forst.) G. Forst. subsp.
pubigerum Sleumer; in montane wet forests of Metrosideros collina and Weinmannia
marquesana var. marquesana; in montane shrublands; and in and summit cloud forests
and shrublands with Alsophila tahitensis, Cyrtandra spp., Dicranopteris linearis, Freycinetia spp., Metrosideros collina, Psychotria spp., Sphaeropteris spp., and Vaccinium cereum (L. f.) G. Forst. var. adenandrum (Decne.) F. Br.and diverse pteridophytes. Threats
in most areas include human disturbance, feral pigs, and invasive weeds.
Conservation status. Proposed IUCN Red List Category Endangered (EN): B2a,
B2b (i–iii): B2: total area of occupancy less than 5000 km2 (ca. 904 km2). B1a, severely
fragmented; B1b (i–iii), habitat continuing decline inferred. The suitable habitat for
Dryopteris macropholis on Nuku Hiva (ca. 340 km2), Ua Huka (ca. 83 km2), Ua Pou
(ca. 105 km2), Hiva Oa (ca. 315 km2), and Tahuata (ca. 61 km2) is restricted to mountain slopes and summits, indicated as an endangered environment that is threatened
by human activity (deforestation and fire), feral animals, and invasive plants, reducing
the extent of the forest.
Specimens examined. Marquesas Islands: Nuku Hiva. Toovii, Ooumu area, top
of Tapueahu Valley off new road, 8°51'S, 140°19'W, 3500–3700 ft (1067–1128 m),
Wood & Perlman 4606 (P, PTBG [2 sheets], US). Route Taiohae–Toovii, branche
droite de la haute Taipivai, 8°53'S, 140°8'W, 750 m, Florence 8423 (P [6 sheets]);
Piste Nord de Terre Déserte, haute vallée de Tapueahu, 1150 m, 140°11'W, 8°52'S,
Florence et al. 9447 (P [3 sheets]). Ua Huka: Hitikau and the Vaikivi summit region,
8°54'S, 139°31'W, 800 m, Wood & Perlman 10761 (PAP, PTBG, US). Ua Pou: forested ridges and slopes to the N and W of Pouakei, 9°23'S, 140 °5'W, 2300 ft (701
m), Wood & Perlman 10840 (PAP, PTBG, US); Tekohepu, 2500–3000 ft (762–914
m), 9°24.31'S, 140°4.21'W, Wood 6499 (P [2 sheets], PTBG [2 sheets], US). Hiva
Oa: Temetiu, 9°49'S, 139°4'W, 3700 ft (1128 m), Wood 4381 (P, PTBG [2 sheets]);
Temetiu region, drainages to southeast of Vaimete et Vaiumioi (source), headwaters of
Hanamenu, UTM 0710665–8916125, 3500 ft (1067 m), Wood 10045 (PTBG [5
sheets]); piste de Hanamenu, NW du Mt. Temetiu, 1150 m, 9°48'S, 139 °5'W, Flor-
New pteridophyte species and combinations from the Marquesas Islands...
19
ence & Perlman 9665 (P [4 sheets]). Tahuata: summit ridge near Haaiputeomo, NE
of Vaitahu, 9°57.19'S, 139 °5.74'W, 2500–2700 ft (762–823 m), Wood 6572 (BISH,
P, PAP, PTBG [5 sheets], US).
Discussion. This remarkable new species resembles Dryopteris macrolepidota Copel.
(type from Tahiti), a species distinguished by it massive suberect rhizome with russet scales
to 30 × 3 mm, fronds to 160 cm long, stipes to 50–70 cm long, stipes and rachises paleaceous with long, thin russet scales with dark, thickened bases, 3-pinnate subcoriaceous
blades to 80 × 40 cm, the abaxial surfaces with scattered thin, linear scales, and supramedial sori. Society Islands collections in the Bishop Museum herbarium identified as Dryopteris dicksonioides (Mett. ex Kuhn) Copel. have massive prostrate to erect rhizomes to
50 cm tall, fronds 80–140 cm long with stipes 60–100 cm long scaly only near the base,
stipe scales dark brown and opaque to 20 mm long, rachises with small brown, scattered
scales, and 3-pinnate-pinnatifid blades to 80 × 40 cm, the segments thinly chartaceous
with linear brown scales on the lower surfaces. Scales of the rhizomes and stipe bases in
D. macropholis are much larger than in the former two species. Palmer (2003) considered
Dryopteris dicksonioides synonymous with D. glabra (Brack.) Kuntze, an exindusiate species endemic to the Hawaiian Islands. However, the type of D. dicksonioides is from Tahiti
and clearly does not represent D. glabra (Copeland 1932). This new species seems most
closely related to D. sweetorum, known only from Fatu Hiva, which differs by its rhizome
and stipe base scales smaller, lustrous, dark brown, 9–25 × 1–2 mm and its broadly ovatedeltate, 3-pinnate-pinnatifid blades (42–)62–93 × (38–)64–80 cm, the ultimate segments
to 15 × 5 mm, spaced 5–10 mm distant, oblique with tips acute, cut about halfway to
the costule, with margins acutely serrate, and the abaxial surfaces and rachises glabrous.
3. Dryopteris sweetorum Lorence & W. L. Wagner, sp. nov.
urn:lsid:ipni.org:names:77112675-1
http://species-id.net/wiki/Dryopteris_sweetorum
Fig. 5
A D. macropholi minoribus, atro-castaneis, ovato-lanceolatis vel lineari-oblongis squamis
12-25 × 1.5-2 mm supra rhizomates et stipitum bases, largis tripinnato-pinnatifidibus
frondibus, ultimis segmentibus usque ad 15 × 5 mm, intervallibus 5-10 mm distantibus,
obliquis, acutis culminibus, sectis ca. in medio ad costulam cum marginibus acute serratis,
abaxialibus superficiebus et glabris rachidibus differt.
Type. Marquesas Islands: Fatu Hiva: Teavapuhiau, ridge to Touaouoho, 2000 ft
(607 m), 8 September 1995, K. R. Wood 4493 (Holotype: PTBG-038471!, PTBG038472!, PTBG-038473!, PTBG-0384741! [4 sheets]; Isotypes BISH! [2 sheets], P!,
PAP! [3 sheets], NY!, US! [3 sheets]).
Description. Terrestrial ferns; rhizomes short creeping to suberect, 3–15 cm in
diameter including stipe bases, densely clothed with scales; scales of rhizomes and
bases of stipes lustrous, dark brown, 9–25 × 1–2 mm, ovate-lanceolate to linearoblong, concolorous, margins entire to subentire distally with short, irregular teeth,
20
David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Figure 5. Dryopteris sweetorum Lorence & W. L. Wagner. A stipe and part of rhizome B blade C detail
of stipe with scales D lower surface of fertile pinnule showing sori e rhizome scale F part of rhizome scale,
detail of cells. Drawn from the type collection (Wood 4493) and field images.
New pteridophyte species and combinations from the Marquesas Islands...
21
cells linear. Fronds densely clustered, 5–7 per rhizome; stipes (45–)60–100 cm long,
7–10 mm in diameter medially, adaxially grooved, stramineous to medium brown,
densely scaly basally, distally with progressively smaller scales and appressed hairlike scales, mostly glabrescent, surfaces brown-punctuate; blades firmly chartaceous,
broadly ovate-deltate, (42–)62–93 × (38–)64–80 cm, 3-pinnate-pinnatifid at base,
otherwise mostly 2-pinnate-pinnatifid, with pinnae opposite to subopposite, spreading, 8–13 on a side; rachises stramineous, with scattered narrow scales to 2–12 ×
0.2–0.6 mm; lowermost pinnae the largest, 25–38 × 19–25 cm, slightly inequilateral, basiscopic basal pinnules 10–15 cm long, acroscopic basal pinnules slightly
shorter, 7–12 cm long, distal pinnae stalked to 5 mm or sessile, apex pinnatifid; ultimate pinnules to 15 × 5 mm, spaced 5–10 mm distant, oblique, tips acute, cut ca. ½
toward costule, with margins acutely serrate, adaxially glabrous, abaxially glabrous or
rachises with scattered hairlike scales; veins forking, scarcely visible adaxially, prominulous abaxially. Sori (1–)3–6 pairs per segment, supramedial; indusia 0.9–1.3 mm
in diameter, brown, thick, glabrous except with a few short-stipitate glands in center.
Spores dark brown.
Distribution. Known only from Fatu Hiva, Marquesas Islands, in montane wet
forest at 600–640 m elevation.
Ecology. This new endemic species is apparently rare and localized, known only
from the region from Teavapuhiau ridge to Mt. Touaouoho where it grows scattered
among other ferns on hillsides in understory of open forest with Alsophila tahitensis,
Crossostylis biflora, Cyclophyllum barbatum (G. Forst.) N. Hallé & J. Florence, Freycinetia impavida, Hibiscus tiliaceus, Metrosideros collina, Pandanus sp., Weinmannia
marquesana var. marquesana, and Vaccinium cereum var. adenandrum, and Wikstroemia
coriacea Seem.
Etymology. We take great pleasure in naming this magnificent new species in
honor of Barbara K. and Cyrus B. Sweet, III, who have generously supported scientific
research at the National Tropical Botanical Garden and particularly the Vascular Flora
of the Marquesas Islands project.
Discussion. This new species differs from D. macropholis by the characters noted
in the diagnosis above, notably in the ultimate pinnules segments spaced 5–10 mm
distant with serrate margins and acute apices.
Conservation status. Proposed IUCN Red List Category: Critically Endangered
(CR): B1ab, B2a,b (i–iii). B1, total extent of occurrence less than 100 km2 (ca. 85
km2), a,b, known from a single location; B2a, estimated area of occupancy estimated
to be less than 10 km2 [three collections likely representing a single population are
known]; B2b (i–iii), habitat continuing decline inferred. The estimated area of occupancy for D. sweetorum on Fatu Hiva (less than 10 km2) is indicated as an endangered
environment, threatened by human activity (deforestation and fire), feral animals, and
invasive plant species, reducing the extent of the forest.
Specimens examined. Marquesas Islands: Fatu Hiva: Teavapuhiau, leeward side
of windswept ridge, UTM 758470–8840707, 2100 ft (640 m), Wood 10091 (PTBG
[3 sheets], PAP [2 sheets], US [3 sheets]); Teavapuhiau Pass (above Ouia Valley), 700
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
m, B. Gagné 1233 (US); Epaulement SW de Mt. Touaouoho, 770 m, 138°38'W,
10°29'W, Florence et al. 9536 (P [4 sheets]).
Polystichum
Polystichum Roth is a cosmopolitan genus of about 300 species (Mabberley 2008).
Brown and Brown (1931) recognized and described a single species from the Marquesas, P. marquesense E. D. Br. Recent collections from Hiva Oa, Tahuata, and Ua Huka
have revealed the presence of two new species which are described below. Polystichum
marquesense differs from both new species in having exindusiate sori and stipes clothed
with large, dull brown, ovate, overlapping scales.
Key to Polystichum in the Marquesas Islands
1a
1b
2a
2b
Blades uniformly bipinnate .................................................... P. uahukaense
Blades tripinnate, at least in lower portion ..................................................2
Largest stipe scales linear-oblong, 25–30 × 1–3.5 mm, reddish brown with
margins bearing short acicular teeth, concentrated base of stipe, mid to upper
part of stipe with smaller, spreading linear-oblong scales and hairlike scales;
largest pinnules and lobes of largest pinnae usually with 1 apical and occasionally 1–6 crenate marginal teeth, arista of apical tooth 0.5–0.7 mm long;
indusia present ......................................................................... P. kenwoodii
Largest stipe scales ovate to narrowly ovate, 10–20(–30) × 4–10 mm, dull, pale
brown with strongly erose-ciliate margins, overlapping and evenly distributed
along stipe, mixed with smaller ovate-ciliate scales and cobwebby scurf; largest pinnules and pinnule lobes of largest pinnae usually with 3–8(–11) teeth,
arista of apical tooth 0.5–2 mm long; indusia absent ............. P. marquesense
4. Polystichum kenwoodii Lorence & W. L. Wagner, sp. nov.
urn:lsid:ipni.org:names:77112677-1
http://species-id.net/wiki/Polystichum_kenwoodii
Figs 6, 14E
A Polytsicho marquisensi E. D. Br. longioribus et angustis rhizomatibus, stipitis squamis
20-35 mm, squamis prope stipitis basin congregates, soris indusiatis differt.
Type. Marquesas Islands: Ua Huka: Vaikivi summit region and drainage, 8°54'S,
139°31'W, 600 m, 16 June 2004, K. R. Wood 10759 (Holotype PTBG-044161!,
PTBG-044162!, PTBG-044163! [3 sheets]; Isotypes BISH!, P!, PAP!, US!).
Description. Terrestrial ferns; rhizomes erect or suberect, 7–20 cm long, 5–15 cm
in diameter including bases of stipes (from collector’s notes); rhizome scales 20–35 ×
2–2.5 mm, linear-oblong, concolorous, medium brown, margins entire, cells narrow
New pteridophyte species and combinations from the Marquesas Islands...
23
Figure 6. Polystichum kenwoodii Lorence & W. L. Wagner. A blade B stipe and part of rhizome C lower
surface of fertile pinnule showing sori and scales D sorus with indusium and scales e rhizome scale F part
of rhizome scale, detail of cells G lower surface of sterile pinnule showing scales. Drawn from type collection (Wood 10759) and field images.
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
linear, in sinuous files, mixed with slender hairlike scales. Fronds clustered; stipes 55–90
cm long, 5–6 mm in diameter medially, 2/5 as long as or equaling length of the blade,
adaxially grooved, pale brown, basally densely clothed with reddish brown, linear to
linear-oblong scales 10–30 × 1.5–3 mm, similar to rhizome scales, stipes distally with
progressively smaller and finer scales mixed with matted hairlike scales or glabrescent;
blades narrowly ovate to oblong-ovate, 50–83 × 30–32 cm, tripinnate at least in lower
pinnae, with 20–28 pinnae on a side; rachises stramineous, moderately to densely scaly
with reddish brown hairlike scales to 8 mm long; pinnae linear-oblong, apices longacuminate, largest pinnae 8–26 × 4.5–7 cm, lowermost 2(–3) pinnae pairs slightly
reduced, costules adaxially grooved; largest pinnae with up to 32 pairs of pinnules, pinnules deeply incised acro- and basiscopically forming 1–2 free, obovate lobes or blades
tripinnate in large fronds, pinnule apices obtuse, tips abruptly acuminate-aristate, both
margins serrate-crenate, blades stiffly chartaceous to subcoriaceous, adaxially glabrous
except for a few hairlike scales basally, abaxially with whitish or tan hairlike scales with
expanded lacerate-ciliate bases on costules and veins; veins forking, scarcely visible
adaxially, slightly prominulous abaxially. Sori with a peltate indusium 0.5–0.7 mm in
diameter, tan, thin and fugacious, with scattered marginal projections, not confluent
at maturity. Spores dark brown.
Distribution. Marquesas Islands, known from Ua Huka, Hiva Oa, and Tahuata.
Habitat. This new species occurs at 600–884 m elevation in diverse montane wet
forest and shrubland dominated by Freycinetia sp., Hibiscus tiliaceus, and Weinmannia
marquesana var. marquesana with other associates including Alstonia marquisensis M.
L.Grant, Boehmeria virgata (G. Forst.) Guill., Cheirondendron bastardianum (Decne.)
Frodin, Crossostylis biflora, Cyclophyllum barbatum, Ficus prolixa var. prolixa, Metrosideros collina, Myrsine grantii Fosberg & Sachet, Reynoldsia marchionensis F. Br., Xyolosma suaveolens subsp. pubigera, and often growing with numerous other pteridophytes.
It also occurs in transitional mesic to wet forest with Alsophila tahitensis, Crossostylis
biflora, Freycinetia sp., Metrosideros collina, Pandanus tectorius, and Weinmannia marquesana var. marquesana. Polystichum kenwoodii grows terrestrially in forest understory
or sometimes along boulder-strewn stream beds.
Etymology. We take pleasure in naming this new species for its collector Kenneth
R. Wood (1953–), whose excellent collections have contributed greatly to our knowledge of the Marquesas flora.
Conservation status. Proposed IUCN Red List Category: Endangered (EN):
B2a, B2b (i–iii): B2: total area of occupancy less than 500 km2 (ca. 460 km2); B2a,
severely fragmented with only three populations known; b (i–iii), habitat continuing
decline inferred. The suitable habitat for Polystichum kenwoodii on Ua Huka (ca. 83
km2), Hiva Oa (ca. 315 km2), and Tahuata (ca. 61 km2) is indicated as an endangered
environment, threatened by human activity (deforestation and fire), feral animals, and
invasive plants, reducing the extent of the forest.
Specimens examined. Marquesas Islands: Hiva Oa: Hanamenu region, up Hanamenu valley to the drainages below and west of Temetiu, 9°76'S, 139°0'W, 884 m,
K. R. Wood 10232 (BISH, P, PAP, PTBG [3 sheets], US). Tahuata: Amatea region,
New pteridophyte species and combinations from the Marquesas Islands...
25
around Haaoiputeomo satellite dish, 9°92'S, 139°8'W, 884 m, K. R. Wood 10257
(BISH, P, PAP, PTBG [3 sheets], US).
Discussion. Polystichum rapense E. D. Br. from Rapa Iti (Austral Islands) resembles
this new species but differs in having very dark brown, almost black scales 3–4 mm wide
on the rhizome and base of stipe and much smaller, bipinnate fronds (stipes 6–10 cm
long, blades 11–14 × 8–10 cm). Polystichum stokesii, also from Rapa Iti, has similar dark
brown to blackish, lustrous scales on the rhizome and base of stipe and shorter fronds
with stipes to 20 cm long and blades to about 50 × 40 cm compared with P. kenwoodii.
5. Polystichum uahukaense Lorence & W. L. Wagner, sp. nov.
urn:lsid:ipni.org:names:77112678-1
http://species-id.net/wiki/Polystichum_uahukaense
Figs 7, 14F, 15A
A Polytsicho marquisensi E. D. Br. minoribus rhizomatibus 3-4 cm. longis × 1.5-3 cm diametro, frondibus minoribus bipinnatibus cum laminis 26-58 × 12-27 cm, soris indusiatis differt.
Type. Marquesas Islands: Ua Huka: Hane/Hokatu cliff zone, 520 m, 11 December
2003, K. R. Wood & J.-Y. Meyer 10518 (Holotype PTBG-042930!; Isotypes AD!,
BISH!, K!, MO!, NY!, P!, PAP!, UC!, US!).
Description. Terrestrial ferns; rhizomes short, erect or suberect, 3–4 cm long, 1.5–3
cm in diameter; rhizome scales 20–28 × 2–2.5 mm, linear-oblong, concolorous, medium to dark brown, margins entire or with occasional short teeth, cells linear, in
sinuous files. Fronds 5–7 per rhizome; stipes 16–43 cm long, ½–2/3 length of the
blades, adaxially grooved, base with dense, linear to linear-oblong scales 10–20 × 1–2
mm, brown and similar to rhizome scales but with thinner, lacerate-dentate margins,
mixed with smaller hairlike scales with lacerate bases, mid- to upper part of stipe with
hairlike scales or glabrescent; blades ovate-oblong to narrowly oblong, 26–58 × 12-27
cm, bipinnate; rachises densely scaly with pale brown, hairlike scales 3-6 mm long
with expanded lacerate base; pinnae 20–30 on a side, narrowly oblong, apex narrowly
acute, largest pinnae 6–15 × 1.5–3 cm, lowermost 1(–2) pairs slightly reduced, costules
adaxially grooved, pinnules 8–16 pairs per pinna, each incised acroscopically with a
small auricle or basal pinnules of larger pinnae sometimes deeply incised forming a
nearly free obovate lobe, pinnule apices acute to obtuse, both margins crenate, apex
and lobes spinulose at tip; blades chartaceous to subcoriaceous, adaxially sparsely capitate-glandular when young, glabrate, abaxially with pale tan hairlike scales on costae,
veins, and margins, veins forking, scarcely visible adaxially, slightly prominulous abaxially. Sori each with a peltate indusium 0.5–0.6 mm in diameter, tan, with numerous
marginal projections, thin and often fugacious, not confluent at maturity. Spores black.
Distribution. Marquesas Islands, known only from the type locality on Ua Huka
in the Hane/Hokatu cliff zone at the head of the valley above Hane village.
Ecology. Polystichum uahukaense is extremely localized and known from a single
population on moist, mossy cliff faces in shade of wet forest with Hibiscus tiliaceus and
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Figure 7. Polystichum uahukaense Lorence & W. L. Wagner. A rhizome and stipe B blade C detail of
stipe with scales D lower surface of sterile pinnule e lower surface of fertile pinnule showing sori and scales
F rhizome scale, outline G part of rhizome scale, detail of cells. Drawn from the type collection (Wood
10518) and field images.
New pteridophyte species and combinations from the Marquesas Islands...
27
Pandanus tectorius dominant, at 520–525 m. Associated vegetation on cliffs includes
Bidens polycephala Sch. Bip., Boehmeria virgata, Macropiper latifolium (L. f.) Miq., Peperomia pallida (G. Forst.) A. Dietr., Phyllanthus pacificus Müll. Arg., and pteridophyes
including Dryopters macropholis, Microsorum grossum (Langsd. & Fisch.) Brownlie,
Nephrolepis sp., Pteris comans G. Forst., Selaginella arbuscula (Kaulf.) Spring., and Tectaria jardinii (Mett. ex Kuhn) E. D. Br. The main threats to this ecosystem include
rockslides and competition with naturalized plant species including Ageratum conyzoides
L., Coffea arabica L., Kyllinga brevifolia Rottb., Oplismenus compositus (L.) P. Beauv.,
Paspalum conjugatum P. J. Bergius, Psidium guajava L., and Zingiber zerumbet (L.) Sm.
Etymology. This new species is named for its only known island of occurrence.
Conservation status. Proposed IUCN Red List Category Critically Endangered
(CR): B2a, B2b (i–iii); D): B2: total area of occupancy less than 10 km2 (ca. 5 km2).
B2a, a single population known; b (i–iii), habitat continuing decline inferred. The
suitable habitat for Polystichum uahukaense on Ua Huka (ca. 83 km2) is indicated as
an endangered environment, threatened by feral animals and invasive plants, reducing
the extent of the forest. Estimated population size is 300 individuals (Wood 10735).
Specimens examined. Marquesas Islands: Ua Huka: Hane/Hokatu cliff zone,
520 m, 14 December 2003, K. R. Wood & J.-Y. Meyer 10552 (P, PAP, PTBG, US),
480 m, S 8° 54'37”, W 139° 31'32”, 12 June 2004, Lorence et al. 9303 (BISH, NY,
P, PAP, PTBG, US).
Discussion. Polystichum uahukaense differs from its Marquesas Islands congeners
in having uniformly bipinnate blades, a feature it shares with P. rapense from Rapa Iti
which differs in having very dark brown, almost black scales 3–4 mm wide on the rhizome and base of stipe and much smaller, bipinnate fronds (stipes 6–1 cm long, blades
11–14 × 8–10 cm). However, based on overall morphology these two species seem
closely allied to each other.
PTERIDACEAE
Pteris
Pteris L. is a large, pantropical genus comprising about 250 species (Mickel and Smith
2004). In their treatment of southeastern Polynesian pteridophytes, Brown and Brown
(1931) recognized two indigenous species of Pteris L. from the Marquesas: P. comans G.
Forst. and P. tripartita Sw. Recent collections made by Kenneth R. Wood in conjunction
with this project have yielded three additional, very distinctive endemic Pteris species described below.
Key to Pteris in the Marquesas Islands
1a
1b
Blade tripartite ...........................................................................P. tripartita
Blade 1- to 2-pinnate ..................................................................................2
28
2a
2b
3a
3b
4a
4b
David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Blade 2-pinnate ............................................................................ P. comans
Blade 1-pinnate to 1-pinnate-pinnatifid ......................................................3
Stipes thickly clothed in persistent, stiff, spreading, red-brown bristlelike or
hairlike scales .......................................................................P. marquesensis
Stipes glabrous or soon glabrate ..................................................................4
Apex pinnatifid, 6–7-lobed .....................................................P. hivaoaensis
Apex entire, the single terminal pinna free or sometimes adnate basally to the
distal pinnae pair ..................................................................P. tahuataensis
6. Pteris hivaoaensis Lorence & K. R. Wood, sp. nov.
urn:lsid:ipni.org:names:77112679-1
http://species-id.net/wiki/Pteris_hivaoaensis
Figs 8, 15B
Ab aliis Marquesas speciebus laminis 1-pinnatis usque ad 1-pinnato-pinnatifidis, apice
cum 6-7 loborum paribus distalibus falcatis, glabris vel glabrescentibus stipitibus differt.
Type. Marquesas Islands: Hiva Oa: Temetiu, windswept ridges and drainages, 9°49'S,
139°4'W, 930 m (3050 ft), 24 August 1995, K. R. Wood 4374 (Holotype PTBG038496!; Isotypes BISH!, P!, PAP!, US!).
Description. Lithophytic ferns; rhizomes erect, clumping together, 3–7 cm long,15–
22 mm in diameter, clothed in very narrow, golden-brown acicular hairlike scales 1–2
mm long. Fronds erect or arching-pendent, to 30 cm long, clustered at rhizome apex;
stipes ca. ½ length of frond, up to 1 mm in diameter medially, atrocastaneous, grooved
adaxially, sparsely pustulate-tuberculate, glabrous except for a few linear to linearlanceolate, falcate scales 2–3 × 0.1–0.5 mm at bases, tan-brown, margins subentire;
blades chartaceous, glabrous, narrowly ovate to lanceolate, 10–16 × 5–9 cm; proximal
2–3 pinnae consisting of subopposite, subequal pairs, 1-pinnate-pinnatifid for most of
length, sessile or stalked up to 2 mm, non-articulate, bearing 2–3 pair of falcate lobes
of increasing size proximally with maximum size at basiscopic lobe of lowest pinnae,
lobes linear-oblong or linear-triangular 0.3–2.5 × 0.3–11 mm, apices rounded to acute,
crenate to dentate, basal pinnae slightly reduced, blade distally pinnatifid, lanceolate,
the apex pinnatifid, with 6–7 pairs of falcate lobes 5–8 mm wide above their bases,
dilated basiscopically, margins mostly entire, tapering gradually to acute apices with
crenate-dentate margins, single terminal lobe falcate or tapering to narrowly linear crenate margin; margins of sterile pinnae finely dentate with one tooth per vein ending;
costae and costules grooved adaxially, rounded abaxially, similar to stipe in color; veins
netted with 1–2 rows of areoles. Sori occasionally interrupted at sinuses and absent at
apices of segment; indusia tan-brown with entire margins. Spores tan-brown.
Distribution. Known only from Hiva Oa, Marquesas Islands.
Ecology. This rare species is known only from the type location at 930 m on wet
stream walls adjacent to wet forest dominated by Cheirodendron bastardianum, Cros-
New pteridophyte species and combinations from the Marquesas Islands...
29
Figure 8. Pteris hivaoaensis Lorence & K. R. Wood. A habit B lower surface of fertile pinna C sterile
pinnule, detail of venation D lower surface of fertile pinnule showing sori e rhizome scale. Drawn from
the type collection (Wood 4374) and field images.
30
David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
sostylis biflora, Metrosideros collina, Weinmannia marquesana var. marquesana and other
associates including Alsophila tahitensis, Cyrtandra spp., Freycinetia sp., Leptochloa marquisensis (F. Br.) P. M. Peterson & Judz., Melicope spp., and Psychotria spp., interspersed
with a rich pteridophyte understory. The main threats to this habitat are feral pigs and
invasives plants including Elephantopus mollis Kunth, Psidium guajava, and Syzygium
cumini (L.) Skeels.
Conservation status. Proposed IUCN Red List Category Critically Endangered
(CR): B2a, B2b (i–iii), D); B2: total area of occupancy less than 10 km2 (ca. 5 km2).
B2a, a single population known; b (i–iii), habitat continuing decline inferred; D, population estimated to number fewer than 250 mature individuals. The suitable habitat
for Pteris hivaoaensis on Hiva Oa (ca. 315 km2) is indicated as an endangered environment, threatened by human activity (deforestation, fire), feral animals, and invasive
plants, reducing the extent of the forest. D, the rarity of this species is supported by
the lack of collections and the small extant area within a commonly collected island,
i.e. with a single collection.
Etymology. This new species is named for its only known island of occurrence.
Discussion. Pteris hivaoaensis is quite distinct from the other Marquesan Pteris
species, differing by its pinnate to pinnate-pinnatifid blades with glabrous or glabrescent stipes and the apex pinnatifid with 6–7 pairs of falcate lobes.
7. Pteris marquesensis Lorence & K. R. Wood, sp. nov.
urn:lsid:ipni.org:names:77112680-1
http://species-id.net/wiki/Pteris_marquesensis
Figs 9, 15C, D.
Ab aliis Marquesas speciebus laminis 1-pinnatis usque ad 1-pinnato-pinnatifidis, stipitibus
dense vestitis cum squamis persistentibus, rigidibus, setiformibus rubro-castaneis differt.
Type. Marquesas Islands: Tahuata, summit ridge near Haaiputeomo, satellite dish
region NE of Vaitahu, 2500–2700 ft, 9°57.19'S, 139°5.74'W, 17–19 July 1997, K.
R. Wood 6565 (Holotype PTBG 038520!, 038521!, 038522!, 038523!, 038524! [5
sheets]; Isotypes BISH!, NY!, P!, PAP!, UC!, US!).
Description. Terrestrial ferns; rhizomes dorsiventral, short creeping to suberect,
up to 2 cm in diameter (to 15 × 15 cm including stipe bases), scaly at apex, scales of
rhizome and base of stipe concolorous, reddish brown, lustrous, linear, sinuate distally,
15–30 × 0.5–0.7 mm, margins subentire or rarely with scattered short teeth, intermixed with smaller narrower scales. Fronds 4–7 per rhizome, clustered at rhizome apex;
stipes 60–134 cm long, 4–6 mm in diameter medially, adaxially grooved, stramineous,
densely scaly especially in lower ca. 1/3, above with scales spreading and less dense, acicular, bases dark brown, persistent, thickened, distally sinuate, apex filiform, margins
subentire; lamina chartaceous, 1-pinnate-pinnatifid or in smaller fronds 1-pinnate,
66–90 × 35–64 cm, ovate-triangular, base obtuse or truncate, apex acute, pinnatifid
and not conform; pinnae 6–10 pairs, basal 1–5 pinnae pairs rarely entire or more com-
New pteridophyte species and combinations from the Marquesas Islands...
31
Figure 9. Pteris marquesensis Lorence & K. R. Wood. A rhizome and stipe bases B fertile blade C lower
sterile pinna D lower surface of fertile pinnule showing marginal sori e rhizome scale F part of rhizome
scale, detail of cells G detail of stipe scales. Drawn from the type collection (Wood 6565) and field images.
32
David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
monly irregularly pinnatifid or lobed at least toward base, or occasionally to apex in
sterile fronds, basal pair of pinnae on stalks 2–6 mm long, largest basal pinnae 22–36 ×
11–18 cm, basiscopic lobes to 13 × 2.5 cm, linear-oblong, slightly falcate, nearly twice
as long as acroscopic lobes, acroscopic base often auriculate, apex acute to acuminate,
occasionally obtuse in shorter lobes, middle and upper pinnae sessile, basiscopic base
decurrent on rachis, acroscopic base free and nearly parallel to rachis, margins entire or
finely serrate towards lobe apices, rachis stramineous, grooved adaxially, with scattered
dark brown, acicular scales to 2 mm long, glabrescent or scales sometimes persisting in
pinnae axes; veins netted with 2–5 rows of areoles in larger pinnules. Sori with indusia
1 mm wide, entire, continuous along margins except at serrate apices; sporangia mixed
with paraphyses. Spores medium brown.
Distribution. Known from Tahuata and Hiva Oa, Marquesas Islands.
Ecology. Pteris marquesensis occurs in wet forest and shrubland with Crossostylis
biflora, Freycinetia sp., Metrosideros collina, and Weinmannia marquesana var. marquesana, dominant. Other associates include Cyrtandra spp., Morinda citrifolia, Psychotria
spp., Vaccinium cereum var. adenandrum, numerous pteridophytes including Asplenium spp., Doodia marquesensis E. D. Br., and Tmesipteris gracilis Chinnock, and diverse
bryophytes.
Etymology. This new species is named for the Marquesas Islands, where it is
known from three islands.
Conservation status. Proposed IUCN Red List Category Endangered (EN): B2a,
B2b (i–iii), D): B2: total area of occupancy less than 500 km2 (ca. 10 km2); b (i–iii),
habitat continuing decline inferred; D, population size estimated to number fewer
than 250 mature individuals. The suitable habitat for Pteris marquesensis on Hiva Oa
(ca. 315 km2) and Tahuata (ca. 61 km2) is indicatedas an endangered environment,
threatened by human activity (deforestation, fire), feral animals, and invasive plants,
reducing the extent of the forest.
Discussion. This striking, large new species is distinctive among all other Polynesian and Micronesian members of the genus. It resembles Pteris warburgii Christ
from Papua New Guinea which also has reticulate veins and pinnatifid blades, but that
species differs in having only a single pair of lateral pinnae and a terminal pinna, and
very few, inconspicuous rhizome and stipe scales. Pteris marquesensis also somewhat
resembles P. umbrosa R. Br. from Australia, but that species has fronds with free dichotomous veins and narrower pinnae adnate to the winged rachis, with the lowermost
pair of pinnae pinnate-pinnatifid.
Specimens examined. Marquesas Islands: Hiva Oa: Temetiu region, drainages to the SE of Vaimete and Vaiumioi, headwaters of Hanamenu, UTM 0710665–
8916125, 1067 m, Wood 10046 (P, PAP, PTBG, US); Ootua, W side of summit area
above road, 9°46'S, 138°58'W, 777 m, Wood 10871 (PTBG, 3 sheets), Wood 10072
(P, PAP, PTBG, US). Tahuata: Haaoiputeomo near satellite dish, NE from Vaitahu to
summit ridge, 2000–2500ft (610–762 m), Wood 4461 (BISH, NY, P, PAP, PTBG,
UC, US); satellite dish region NE of Vaitahu, 2500–2700 ft (762–823 m), 9°57.19'S,
139°5.74'W, Wood 6556 (BISH, NY, P, PAP, PTBG, US).
New pteridophyte species and combinations from the Marquesas Islands...
33
8. Pteris tahuataensis Lorence & K. R. Wood, sp. nov.
urn:lsid:ipni.org:names:77112681-1
http://species-id.net/wiki/Pteris_tahuataensis
Figs 10, 15E
Ab aliis Marquesas speciebus laminis 1-pinnatis usque ad 1-pinnato-pinnatifidis, distalibus pinnis non alatis, et glabris ver glabrescentibus stipitibus differt.
Type. Marquesas Islands: Tahuata: Hanatetena, main valley, first deep gulch to the
north, general UTM 0710938 – 8899189, 518 m, 2 Feb 2003, K. R. Wood 10083
(Holotype: PTBG-04297!; Isotypes P!, PAP!, UC!, US!).
Description. Terrestrial or lithophytic ferns; rhizomes creeping to suberect, 2–5 cm
long, 1–3 cm in diameter, clothed in fine, golden-brown acicular hairs 1–2.5 mm long;
fronds 15-20 per rhizome, pendent, 35–125 cm long; stipes ca. ½ length of frond,
up to 4 mm in diameter, atrocastaneous to stramineous, grooved adaxially, sparsely
pustulate, glabrous except for a sparse cover of linear to linear-lanceoate, tan-brown
scales 3–5 × 0.3–0.5 mm at base of stipes; blades chartaceous, glabrous, ovate, 18–65 ×
12–35 cm, ovate. with 5–7 pairs of pinnae; proximal 1–3 pinnae pairs 1–(2–) pinnatepinnatifid, up to 34 cm long, sessile or stalked up to 9 mm, base obtuse to truncate,
uniauriculate or bearing 2–3 pair of falcate lobes or with proximal pair occasionally
dividing into falcate pinnules 5–13 cm × 1.4–1.7 mm, reaching maximum length
basiscopically on lowest pinnae, apices acute to cuneate, or attenuate, margins crenate
to dentate; distal part of blade composed of 3–5 pinnae pairs, these simple, falcate,
subopposite, up to 23 cm long, the bases sessile, obtuse to truncate, sometimes uniauriculate acroscopically, tapering gradually to cuneate or attenuate apices with crenatedentate margins, the single terminal pinna free or sometimes adnate basally to the
distal pinnae pair, 6.5–23 × 1.0–2.5 cm, apex attenuate, crenate; costae and costules
grooved adaxially, rounded abaxially, similar to stipe in color; veins netted with 2–3
rows of areoles. Sori with indusia 0.5–0.9 mm wide, olivaceous; sori usually absent at
apices of pinnae, Spores castaneous.
Distribution. Known only from Tahuata, Marquesas Islands.
Ecology. This new terrestrial or lithophytic species occurs in wet forests and
shrublands from about 418 to 914 m elevation, usually on windswept vertical cliffs
in water seepage. A population of approximately 50–70 plants was observed at 914
m on windswept vertical cliffs around a natural spring on saturated basalt walls with
Leptochloa marquisensis (F. Br.) P. M. Peterson & Judz. and Selliguea feeioides Copel.,
mosses, and lichens in wet forest of Crossostylis biflora, Freycinetia sp., Metrosideros
collina, Reynoldsia marchionensis, Weinmannia marquesana var. marquesana, Myrsine
grantii, and Pandanus tectorius. A second population occurs at 518 m in mesic forest with Cerbera manghas L., Cyclophyllum barbatum, Hibiscus tiliaceus, Pandanus
tectorius, Sapindus saponaria L. and Xylosma suaveolens subsp. pubigerum. A third
population is from lowland degraded mesic forest at 418 m elevation with Cerbera
manghas, Cyclophyllum barbatum, Ficus prolixa var. prolixa, Hibiscus tiliaceus, Pan-
34
David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Figure 10. Pteris tahuataensis Lorence & K. R. Wood. A habit B blade C fertile pinna distal part D lower
surface of fertile pinnule showing marginal sori e rhizome scale. Drawn from the type collection (Wood
10083) and field images.
New pteridophyte species and combinations from the Marquesas Islands...
35
danus tectorius, and Xylosma suaveolens subsp. pubigerum, Sapindus saponaria. The
main threats to this species include competition from invasive alien plant species and
habitat destruction by feral goats and fire at lower elevations. Known only from three
collections, the one population at 418 m having an estimated 50–70 plants (Wood
10085), the others were isolated plants.
Conservation status. Proposed IUCN Red List Category Critically Endangered
(CR): B2a, B2b i–iii; D): B2: total area of occupancy less than 10 km2 (ca. 5 km2); B2a,
three populations known; b (i–iii), habitat continuing decline inferred; D, population
estimated to number fewer than 250 mature individuals. The suitable habitat for Pteris tahuataensis on Tahuata (ca. 61 km2) is indicated as an endangered environment,
threatened by human activity (deforestation, fire), feral animals, and invasive plants,
reducing the extent of the forest. D, the rarity of this species is supported by the lack
of collections and the small extant area, i.e. with only three known collections and an
estimated total population size of fewer than 250 plants.
Etymology. This new species is named for its only known island of occurrence.
Specimens examined. Marquesas Islands: Tahuata: Hapatoni, village to the
south of Vaitahu, ridge to summit above Patikoea point, UTM 0706136–8897872,
418 m, Wood 10085 (BISH, K, MO, NY, P, PAP, PTBG, UC, US); Amatea region,
locations around Haaoiputeomo satellite dish, 9°92'S, 139°8'W, 914 m, Wood 10250
(PTBG, US).
THELYPTERIDACEAE
With approximately 950 species (Smith et al. 2006) the Thelypteridaceae are one of
the largest and most diverse fern families in the Marquesas and the tropics in general. Holttum recognized 15 genera in the Pacific and Australasia (Holttum 1976,
1977). However, delimitation of Holttum’s genera is often not clear-cut, and some
of the most important characters he used for circumscribing them involve chromosome numbers and spore characters visible only at high magnification (e.g., under
the scanning electron microscope). Furthermore, although many of Holttum’s genera
seem monophyletic, a combination of characters is usually needed to circumscribe
them. For these reasons, many authors now recognize only about five genera of Thelypteridaceae (Kramer and Green 1990; Smith and Cranfill 2002; Smith et al. 2006).
For the purposes of the Marquesas Vascular Flora project we prefer to adopt this
broader generic circumscription of Thelypteris Schmidel. and Cyclosorus Link, but follow Kramer and Green (1990) in recognizing the subgenera as natural groups. Two
new species of Cyclosorus (subg. Plesioneuron) and one new species of Thelypteris (subg.
Coryphopteris) have come to light among specimens collected in the Marquesas and
are described below. One new combination is required, bringing the total number of
Thelypteridaceae species in the Marquesas to nine. The genera may be separated using
the characters in the following key.
36
David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Key to the genera of Thelypteridaceae in the Marquesas Islands
1a
1b.
Veins to 14 pairs per segment; segments oblique, falcate; indusia absent or
very small; sporangia generally setose ........................................... Cyclosorus
Veins to 7 pairs per segment; segments spreading or slightly oblique, not falcate; indusia relatively large, persistent; sporangia lacking setae .....................
............................................................... Thelypteris (subg. Coryphopteris)
Cyclosorus
Viewed in the broad sense, Cyclosorus is the largest genus of Thelypteridaceae, comprising
about 600 species in the tropics of both hemispheres. The seven indigenous or endemic
and one naturalized species in the Marquesas may be separated by the following key.
Key to the species of Cyclosorus in the Marquesas Islands
1a
1b
2a
2b
3a
3b
4a
Lower pinnae not reduced, lower 2–3 pairs usually not deflexed; veins all free;
sinus with thickened membrane ± obliquely decurrent as a ridge almost to
costa on lower surface ........................C. (subg. Plesioneuron) marquesicus
Lower pinnae not or only slightly reduced, lower 2 (−3) pairs deflexed; veins
free or uniting, but sinus lacking thickened ridge decurrent toward costa on
lower surface ...............................................................................................2
Blade with basal pinnae much narrowed on both sides near base, rudimentary
pinnae below them 0−2 pairs, inconstant and irregular; blades with sessile
spherical yellow glands present abaxially, especially along veins near segment
tips; scales absent along costae abaxially, costal hairs ca. 0.1 mm long ...........
................................ C. (subg. Amphineuron) opulentus (Kaulf.) Nakaike
Blade with basal pinnae narrowed or not on both sides, but rudimentary basal
pinnae lacking or several pairs regularly present; blades lacking sessile spherical yellow glands abaxially, or these spread ± evenly on laminar tissue; scales
present or absent along costae abaxially, costal hairs 0.1−1 mm long, or hairs
absent .........................................................................................................3
Fronds 1.5−3 m long; narrow, hairlike scales to 2 cm long present throughout
stipe and on basal part of rachis, often breaking and leaving spine-like bases;
lower surface of pinnae lacking sessile spherical yellow glands; sori exindusiate......................................C. (subg. Chingia) longissimus (Brack.) Ching
Fronds to 1.2 m long, usually smaller; stipe scales less than 2 cm long, these
usually confined to base of stipe, not hairlike or leaving spine-like bases; sessile spherical yellow glands scattered all over lower surface of pinnae (except
C. invisus); sori indusiate.............................................................................4
Rhizomes creeping; laminar tissue between veins lacking sessile glands
abaxially ................................................................................................ 5
New pteridophyte species and combinations from the Marquesas Islands...
4b
5a
5b
6a
6b
7a
7b
37
Rhizomes suberect or erect; laminar tissue between veins bearing numerous
sessile, round, yellowish to orange-red glands abaxially ...............................7
Blades glabrous or very sparsely hairy abaxially, hairs if present less than 0.1
mm long on rachises and costae abaxially; costae glabrous adaxially; indusia
absent; sporangia lacking hairs (Hiva Oa) .....................................................
........................................................... C. (subg. Pneumatopteris) florencei
Blades abundantly hairy abaxially, on costae, costules, veins, and between
veins, hairs to 1 mm long; costae hairy adaxially; indusia present, hairy ......6
Rhizomes long-creeping; pinnae lobed 1/3−2/5 toward costae; veins and costules glabrous adaxially; sporangia bearing hairs (Fatu Hiva) .........................
...............................C. (subg. Sphaerstephanos) invisus (G. Forst.) Copel.
Rhizomes short-creeping; pinnae lobed 1/2−2/3 toward costae; veins and costules, sometimes also laminar tissue between veins, with acicular hairs adaxially; sporangia lacking hairs (Ua Huka) .........................................................
...........................................C. (subg. Christella) dentatus (Forssk.) Ching
Abaxial costal hairs red-brown (castaneous), at least in part, curved toward
pinna tips, some hairs > 0.5 mm long (Nuku Hiva) ......................................
.......................................................C. (subg. Sphaerostephanos) castaneus
Abaxial costal hairs hyaline, spreading or ascending, straight or falcate, uniformly very short and < 0.1 mm long, or costae sometimes with scattered to
rather dense longer hairs to ca. 1 mm; Society and Marquesas Islands (Fatu
Hiva, Hiva Oa, Tahuata, Ua Huka) ..............................................................
...................... C. (subg. Sphaerostephanos) subpectinatus (Copel.) Ching
Discussion. Cyclosorus subpectinatus (Copel.) Ching, from the Marquesas and
Society Islands, is extremely variable and possibly represents more than a single species, but the material at hand seems to intergrade. Specimens from different islands
are all slightly different in various indument, venation, indusial, and size characteristics. Specimens with the largest indusia (to nearly 1 mm in diameter) and longest
costal hairs are from Ua Huka (e.g., Dunn 340, BISH; Lorence 9307, PTBG, UC).
The type of C. subpectinatus, from Tahiti, also has rather long, dense costal hairs,
but has smaller indusia only ca. 0.3 mm in diameter The sole collection seen from
Tahuata (Wood 10270, P, PAP, PTBG, US) has exceptionally large fronds, with pinnae to 28 × 3.5 cm. Most specimens seen from both the Society Islands (Huahine,
Moorea, Raiatea, and Tahiti), and the Marquesas Islands have relatively short, uniform costal hairs ca. 0.1 mm long and small indusia ca. 0.3 mm in diameter. Two
specimens from Fatu Hiva (Chapin 789, BISH, cited by Holttum, 1977, as the sole
specimen of the species from the Marquesas; and Florence 9509, BISH, P, US) seem
to be nearly or quite exindusiate. All specimens seen of C. subpectinatus have rather
numerous to moderate, tan, adpressed, lanceolate costal scales abaxially, as well as
dense, spherical, yellowish to orange-red sessile glands between the veins abaxially,
but lack laminar glands adaxially.
38
David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Figure 11. Cyclosorus castaneus A. R. Sm. & Lorence. A frond B upper surface of pinna C lower surface
of fertile pinnules showing sori, hairs, and glands D detail of lower surface of fertile pinnule with sorus,
hairs, and glands. Drawn from the type collection (Lorence et al. 6115).
New pteridophyte species and combinations from the Marquesas Islands...
39
9. Cyclosorus (subg. Sphaerostephanos) castaneus A. R. Sm. & Lorence, sp. nov.
urn:lsid:ipni.org:names:77112682-1
http://species-id.net/wiki/Cyclosorus_(Sphaerostephanos)_castaneus
Fig. 11
A Cyclosoro heterocarpo (Blume) Ching numerosis, nitidis, castaneis, falcatis pilis 0.3−1.5
mm longis super rachides, costis et costulis abaxialibus, parva indusia 0.4−0.6 mm diametro, magis marcatis aerophoribus, fortasseque longioribus, magisque bene-formatis caudicibus differt.
Type. Marquesas Islands: Nuku Hiva: Toovii region, trail along ridge from near
l’Economie Rurale complex to Ooumu peak, 860–1080 m, 17 July 1988, D. Lorence (with W.L. Wagner, J. Florence and S. Perlman) 6115 (holotype PTBG!; isotypes
BISH!, PAP!, US!).
Description. Terrestrial ferns; rhizomes erect, caudices 15–60 cm long; fronds clustered, 7–10 per rhizome; base of stipes to first large pinnae 40–50 cm, distal part of
stipes bearing 7–10 pairs of reduced pinnae 3–5 cm apart, 1–7(–35) mm long, the
largest ones sometimes trifid; rachises reddish brown, bearing numerous reddish, often
curved hairs; proximal large pinnae narrowed at their bases; blades, excluding reduced
proximal pinnae, 50–70 cm long; largest pinnae 11–16 × 1.5–2 cm, at their bases each
with a swollen, tuberculiform aerophore (most developed and scalelike, to 4 mm long
in young fronds), pinnae bases not auricled, apices caudate-acuminate, lobed 2/3–3/4
toward the costae (2–3 mm from costae), lobes slightly falcate, rounded at tips; costules ca. 3–5 mm apart; veins to ca. 8–9 pairs per segment, the basal pair from adjacent
segments anastomosing and producing an excurrent vein 1–1.5 mm long to the sinus,
the next acroscopic vein to sinus-membrane; abaxial surface of costae with very short,
hyaline, spreading to distally curved hairs 0.1 mm long and much longer and stouter
falcate hairs to 0.3–1.5 mm on costae, scales lacking or costae with a few adpressed,
tan, amorphous scales to 2 × 0.1 mm; yellowish sessile glands borne on abaxial laminar
surfaces, often dense, absent or very sparse on adaxial laminar surfaces; stout reddish,
falcate hairs adaxially on costae and costules to 1 mm long, with short, falcate hairs ca.
0.2–0.3 mm long sometimes on veins and between veins, especially below sinuses. Sori
medial; indusia reddish brown to tan, ca. 0.4–0.6 mm in diameter, glabrous or with a
few short hairs 0.1–0.2 mm long; sporangia with sessile, yellow glands on the capsules.
Distribution. Known only from Nuku Hiva, Marquesas Islands.
Ecology. This new species occurs in montane rain forest with fern understory. All
three specimens seen were collected on Nuku Hiva on or near Ooumu Peak in the
Toovii area and probably represent a single population.
Conservation status. Proposed IUCN Red List Category Endangered (EN): B2a,
B2b i–iii): B2: total area of occupancy less than 500 km2 (ca. 50 km2). B2a, a single
population known; b (i–iii), habitat continuing decline inferred. The suitable habitat
for Cyclosorus castaneus on Nuku Huka (ca. 340 km2) is indicated as an endangered
environment, threatened by human activity (deforestation), feral animals, and invasive
plants, reducing the extent of the forest.
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Etymology. Named for the castaneous, curved hairs that are borne on the costae
abaxially.
Specimens examined. Marquesas Islands: Nuku Hiva: Toovii Plateau, trail behind L’Economie Rurale toward Ooumu peak; 3100 ft (945 m), S. Perlman 10125
(BISH, PTBG); Toovii, Ooumu area, top of Tapueahu Valley off new Hwy, 0851 S
14019 W, 3500–3700 ft (1067—1128 m), K. Wood 4578 (BISH, PAP, PTBG, US).
Discussion. The three known specimens were previously determined as Sphaerostephanos (Cyclosorus)heterocarpus (Blume) Holttum, a related species known from
Southeast Asia, Malesia, Australia (Queensland), Melanesia (Fiji, Vanuatu), and
Samoa. From that species C. castaneus differs in the numerous shiny, castaneous,
falcate hairs 0.3–1.5 mm long, on the rachises, costae, and costules abaxially. These
hairs vary in length, but are frequently longer than 0.5 mm on the costae. Cyclosorus
heterocarpus shows considerable variation in indument (both length and disposition
of hairs); however, the hairs in C. heterocarpus are always hyaline, not so decidedly
curved, and are generally much shorter than those in C. castaneus. Holttum (1982)
recognized many unnamed forms of C. heterocarpus and produced a key to them
based primarily on the depth of pinna lobing, presence/absence of sessile glands on
the laminae adaxially, width of fertile pinnae, and presence/absence of hairs on the
laminae adaxially. Cyclosorus heterocarpus also has larger indusia than C. castaneus, often 1 mm in diameter or more, and the indusia envelop the sporangia when young.
Some variants of C. heterocarpus have sessile glands adaxially (rare in C. castaneous)
and less deeply incised pinnae (Holttum 1982). Aerophores in C. heterocarpus appear to be much less developed, only a lunate, slightly raised area, or mammiform
hump. In C. castaneus, the aerophores are tuberculiform, peglike, or even scalelike,
to 4 mm long. A single specimen from Ua Pou (Anakooma river valley just ESE of
Oave peak, 470 m, Lorence 9117, PTBG) resembles C. castaneus in having reddish
hairs on the costae abaxially, but the hairs are less deeply colored and spreading, not
falcate. Most sporangia seem empty, and what spores are formed are irregular, so it
is possible this is a hybrid involving C. castaneus and C. florencei. However, neither
parental species has been found on Ua Pou. Further study is clearly needed.
10. Cyclosorus (subg. Pneumatopteris) florencei A. R. Sm. & Lorence, sp. nov.
urn:lsid:ipni.org:names:77112683-1
http://species-id.net/wiki/Cyclosorus_(Pneumatopteris)_florencei
Fig. 12
A C. glandulifero (Brack.) Copel. absentibus falcatis, acicularibus pilis adaxialliter supra
axiales costas et presentibus reductarum proximalium pinnarum paribus non tan numerosis
(ca. 10 pro ca. 20), absentibus pilis supra sporangiales stipites et et receptacula differt.
Type. Marquesas Islands: Hiva Oa: Atuona, trail to Hanamenu, 9°48'S, 139°04'W,
wet forest, 29 Jul 1988, J. Florence (with D. Lorence, S. Perlman) 9598 (holotype
BISH!; isotypes P, PAP, neither seen).
New pteridophyte species and combinations from the Marquesas Islands...
41
Figure 12. Cyclosorus florencei A. R. Sm. & Lorence Holotype collection with frond and part of rhizome,
Florence et al. 9598 (BISH). Inset shows detail lower surface of fertile pinna with sori.
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Description. Terrestrial ferns; rhizomes creeping, to 10 mm in diameter; fronds
spaced, base of stipes sparsely scaly, scales brown, lanceolate; stipes to ca. 70 cm long
below first large pinnae, distal part of stipes to 9 mm in diameter at base, bearing ca. 10
pairs of greatly reduced pinnae 2.5–4 cm apart, these 1–10 mm long; proximal large
pinnae narrowed at their bases; blades subcoriaceous, excluding reduced, glanduliform
proximal pinnae, to ca. 60 cm long, gradually reduced distally to a pinnatifid apex;
rachises tan to stramineous, glabrous; large developed pinnae to ca. 30 lateral pairs, to
ca. 12 × 1.7 cm, at their bases each with a swollen, conical aerophore to ca. 0.5 mm
long, pinnae bases not auricled, apices caudate-acuminate, lobed 2/5–1/2 toward the
costae (3–4 mm from costae), lobes oblique and slightly falcate, subacute to rounded at
tip; costules ca. 4 mm apart; veins to ca. 8–9 pairs per segment, prominent (especially
abaxially) on both sides of laminae, the basal pair from adjacent segments generally
obtusely united and producing an excurrent vein 2.5–3 mm long to the sinus, the next
1–1 1/2 pairs merging with this excurrent vein or running to a cartilaginous, raised, sinus membrane; abaxial surface of rachis, costae, costules, and veins nearly lacking hairs
or with scattered, minute hairs less than 0.1 mm, costae bearing adpressed to slightly
spreading, tan, amorphous (cell walls not readily discernible at 30×) scales to 2 × 0.2
mm; yellowish sessile glands absent on both laminar surfaces, pustules also lacking;
hairs absent adaxially on costae, costules, and veins. Sori medial to supramedial; indusia absent; sporangia bearing yellowish capsular glands ca. 0.1 mm, lacking acicular
hairs on sporangia and from receptacles.
Distribution. Hiva Oa, Marquesas Islands, known only from the type collection
made along the trail from Atuona to Hanamenu.
Ecology. Occurs in montane wet forest with Crossostylis biflora, Freycinetia sp.,
Metrosideros collina, Weinmannia marquesana var. marquesana and other species characteristic of this habitat.
Etymology. We take pleasure in naming this new species for Jacques Florence
(1951–) who has done so much to advance our knowledge of the flora of the Marquesas Islands and that of French Polynesia in general.
Conservation status. Proposed IUCN Red List Category Critically Endangered
(CR); B2a, B2b i–iii; D): B1, extent of occurrence estimated to be less than 100 km2;
B2, area of occupancy estimated to be less than 10 km2 (ca. 9 km2), and B2a, a single population known; b (i–iii), habitat continuing decline inferred; D, population
estimated to number fewer than 250 mature individuals. The suitable habitat for Cyclosorus florencei on Hiva Oa (ca. 315 km2) is indicated as an endangered environment, threatened by human activity (deforestation and fire), feral animals, and invasive
plants, reducing the extent of the forest; and D, the rarity of this species is supported
by the lack of collections and the small extant area within a commonly collected island,
i.e. with a single collection.
Discussion. This new species is perhaps most closely related to Cyclosorus glanduliferus (Brack.) Copel., from Rarotonga (Cook Islands), Samoa, Solomon Islands, and
New Hebrides, and to C. stokesii (E. D. Br.) Ching, known only from Rapa Iti in the
Austral Islands (Holttum 1977). The three species are similar in rhizome habit, blade
New pteridophyte species and combinations from the Marquesas Islands...
43
dissection, and blade size. However, C. glanduliferus differs from C. stokesii in having
more numerous glanduliform pinnae along the stipes (to about 20 pairs), longer aerophores at bases of costae, presence of small aerophores at costule bases, fewer and shorter costal scales, presence of numerous acicular, falcate hairs on the costae adaxially, and
hairs borne on the sporangial stalks and receptacles. From C. stokesii, C. florencei differs
in having less scaly stipe bases (scales numerous and ovate in C. stokesii), fewer pairs
of glanduliform proximal pinnae (to 20 pairs of glanduliform pinnae in C. stokesii),
lacking aerophores at bases of costules, more numerous, longer costal scales, 4–5 pairs
of veins uniting below sinus or connivent at the sinuses, and lacking hairs from the
sporangial stalks. Spore differences mentioned by Holttum (1977) between C. glanduliferus and C. stokesii cannot be evaluated for C. florencei because the sporangia in
the type of C. florencei are immature. The label with the type indicates the presence
of small pustules abaxially, but we fail to see any pustules of the kind often found in
species of subg. Pneumatopteris; we do see numerous, slightly more reflective stomates
(guard cells) at 30 times magnification. Until now no species of subg. Pneumatopteris
has been recorded from the Marquesas Islands.
The following new combination in Thelypteridaceae is also required.
11. Cyclosorus marquesicum (Holttum) Lorence & A. R. Sm., comb. nov.
urn:lsid:ipni.org:names:77112686-1
http://species-id.net/wiki/Cyclosorus_marquesicum
Basionym: Plesioneuron marquesicum Holttum, Allertonia 1: 192. 1977.
Type. Marquesas Islands: Hivoa: Hana lafa, 700 m, October 1922, W. B. Jones 1618
(Holotype BISH!).
Distribution. Marquesas Islands (known from Hiva Oa, Tahuata, Ua Huka, and
Ua Pou) and the Society Islands (Moorea). A list of exsiccatae is available on the Flora
of the Marquesas website (http://botany.si.edu/pacificislandbiodiversity/marquesasflora/index.htm).
Thelypteris (subgen. Coryphopteris)
Holttum (1976, 1977) circumscribed Coryphopteris Holttum as a genus of about 50
species with its greatest diversity in New Guinea and the Malesian region. However,
we prefer to consider Coryphopteris as a subgeneric segregate of a more broadly circumscribed Thelypteris following the concepts of Kramer and Green (1990), Smith and Cranfill (2002), and Smith et al. (2006). For the purposes of the Marquesas Vascular Flora
project we prefer to adopt two genera, Thelypteris and Cyclosorus in their broader sense,
but follow Kramer and Green (1990) in recognizing the subgenera as natural groups.
Species belonging to Thelypteris subgenus Coryphopteris are small to medium-sized
terrestrial ferns (rarely epiphytic) resembling miniature tree ferns with an erect or
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
sometimes decumbent rhizome topped by a cluster of 1-pinnate fronds with the basal pinnae pairs deflexed in most species. Additional distinguishing characters include
relatively large, sessile glands especially dense on the lower surface of the lamina, often
abundant scales on the lower surface, and septate acicular hairs on the upper surface of
the rachis and costa of some species. Most species belonging to subgenus Coryphopteris
are restricted to low wet forest or mossy cloud forest habitat on high mountain ridge
slopes and crests, often above 1,000 m elevation, where they grow in leached, nutrient
poor soils (Holttum 1976).
Two species of Thelypteris subgenus Coryphopteris are known from the Marquesas,
T. quaylei (E. D. Br.) Ching [Coryphopteris quaylei (E. D. Br.) Holttum], and a diminutive new species described below, T. marquesensis. Thelypteris quaylei is a larger, more
robust species known from Nuku Hiva, Ua Pou, Ua Huka, Hiva Oa, and Fatu Hiva,
generally at lower elevations (580–884 m). Certain collections of T. quaylei from the
summit region of Ua Huka (700–884 m) tend to be relatively small, precociously fertile plants. However, the appearance and habitats of the two species are quite different
and they can be separated morphologically by the characters in the following key.
Key to Thelypteris subg. Corphyopteris in the Marquesas Islands
1a
Plants with stipes 1–5(–8.5) cm long, with scattered scales when young, with
hairs to 0.5–0.7 mm long; scales near base of stipe 1-2 mm long; blades
2.5–14.5 cm long; pinnae 5–14 pairs; pinnae lobed 1/3–4/5 toward to costa,
sessile glands lacking on surface, sori 1(–2) per segment; pinnule veins 1–5
pairs .................................................................................... T. marquesensis
1b
Plants with stipes 15–25(–33) cm long, densely scaly when young, glabrescent, with hairs to 0.2 mm long; scales near base of stipe 5–7 mm long; blades
20–30 cm long; pinnae (12–) 18–25 pairs; pinnae lobed 4/5–7/8 toward to
costa, sessile glands present on surface, sori (1–) 3–4 per segment; pinnule
veins 5–9 pairs ..............................................................................T. quaylei
12. Thelypteris marquesensis Lorence & K. R. Wood, sp. nov.
urn:lsid:ipni.org:names:77112689-1
http://species-id.net/wiki/Thelypteris_marquesensis
Figs 13, 15F
Species Thelypteri quayleii affinis, sed minori habitu, laminis cum 1-3 paribus inferis deminutisque pinnarum, et pinnarum abaxiali superficie sine sessilibus glandis differt.
Type. Marquesas Islands: Hiva Oa; Temetiu, windswept ridges and drainages, 3280
ft [999 m], 9°48'S, 139 °4'W, 26 August 1995, K. R. Wood 4408 (holotype PTBG038499!; isotypes, P!, PAP!, US!).
Small ferns, usually lithophytic, forming colonies over wet basalt rock faces (rarely
mossy tree trunks); rhizomes slender, decumbent to suberect, unbranched, radial, 1.5–7
cm long, 0.4–1.2 cm in diameter, densely clothed by stipe bases, sparsely covered in red-
New pteridophyte species and combinations from the Marquesas Islands...
45
Figure 13. Thelypteris marquesensis Lorence & K. R. Wood. A habit B lower surface fertile pinnae showing sori C lower surface of fertile pinna showing sori and hairs D rhizome scale. Drawn from the type
collection (Wood 4408) and field images.
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
Figure 14. A Blechnum pacificum, habit, habit, fertile and sterile fronds (Fatu Hiva, Lorence 6171)
B–D Dryopteris macropholis B frond (Ua Huka, Wood 10489, type) C habit, D rhizome and stipe bases
(C, D Hiva Oa, Wood 10045) e Polystichum kenwoodii, rhizome and stipe bases (Hiva Oa, Wood 10232)
F Polystichum uahukaense, habit (Ua Huka, Wood 10552).
New pteridophyte species and combinations from the Marquesas Islands...
47
Figure 15. A Polystichum uahukaense, frond (Ua Huka, Wood 10518) B Pteris hivaoaensis (Hiva Oa,
Wood 4374) C, D Pteris marquesensis C frond D rhizome and stipe bases (both Hiva Oa, Wood 10871)
e Pteris tahuataensis (Tahuata, Wood 10083) F Thelypteris marquesensis, habit, colony on wet rock face
(Hiva Oa, Wood 4408).
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David H. Lorence et al. / PhytoKeys 4: 5–51 (2011)
brown, unicellular acicular hairs; fronds 7–9 per rhizome; stipes 1–5(–8.5) cm long, medium brown, moderately covered in unicellular acicular, red-brown hairs 0.5–0.7 mm
long and scales, the scales sparsely scattered, those near base ovate-falcate, red-brown
to brownish black, 1–2 × 0.5–0.7 mm, margins entire or with sparse short projections;
blades 1-pinnate-pinnatifid, 2.5–8.5 × 1.2–2.8 cm, oblong-elliptic, rachises medium
to red-brown, moderately covered with light brown, curved acicular hairs 0.3–0.7 mm
long, apex pinnatifid, pinnae 5–12 pairs, largest pinnae 0.6–1.4 × 0.3–0.5 cm, apex
obtuse, lobed (1/3–)½ toward costa, lobes crenate, margins with acicular hairs 0.3–0.6
mm long, costules to 2 mm apart, lower pinnae slightly reduced and lower 2(–3) pairs
deflexed, basal acroscopic pinnule 1 mm longer than next, veins 2–3 pairs in basal lobe,
1–2 pairs in middle lobes, abaxial surface of rachises and costae hirtellous with brown
acicular hairs 0.2–0.7 mm long, adaxial surfaces glabrous or with occasional short acicular hairs 0.2–0.5 mm, sessile glands absent. Sori medial, 1(–2) per segment; indusia
reniform to subcircular, margins with sessile glands, otherwise glabrous.
Distribution. Known only on Hiva Oa, Marquesas Islands, from a single population in the vicinity of Mt. Temetiu, on the island’s central summit crest.
Ecology. Thelypteris marquesensis occurs at 1000–1200 m elevation in low, windswept, montane wet forests and shrublands with Alsophila tahitensis, Cheirodendron
bastardianum, Crossostylis biflora, Cyrtandra spp., Freycinetia spp., Leptochloa marquisensis, Melicope spp., Metrosideros collina, Psychotria spp., Weinmannia marquesana var.
marquesana, and abundant pteridophytes in the understory. This new species is rare
and localized, although in one area it occurs in dense colonies of hundreds of plants
that completely cover wet banks and rock faces in and around shallow depressions or
grottos, with fronds often appressed to the rock face (K. Wood, pers. obs.). Threats to
this species include rooting by feral pigs and invasion by aggressive alien plant species,
notably Elephantopus mollis, Psidium guajava, and Syzygium cumini.
Etymology. This new species is named for the Marquesas Islands, where it is
known currently known only from Hiva Oa.
Conservation status. Proposed IUCN Red List Category Critically Endangered
(CR): B2a, B2b i–iii): B1, extent of occurrence estimated to be less than 100 km2;
B2, area of occupancy estimated to be less than 10 km2 (ca. 9 km2), and B2a, a single
population known; b (i–iii), habitat continuing decline inferred. The suitable habitat
for Thelypteris marquesensis on Hiva Oa (ca. 315 km2), confined to Mt. Temetiu and
vicinity, is indicated as an endangered environment, threatened by human activity
(deforestation and fire), feral animals (pigs), and invasive plants, reducing the extent
of the forest.
Discussion. This new species differs from Thelypteris quaylei in its much smaller
habit (although some specimens of T. quaylei from the Ua Huka summit are atypically
small), fronds with 1–3, gradually reduced basal pairs of pinnae, and lack of sessile
glands on the lamina surfaces (but present on indusial margins), a feature it shares with
T. fasciculata Ching from New Caledonia, New Guinea, and the Celebes (Holttum
1977). Although T. marquesensis may occur in the vicinity of T. quaylei (Lorence et al.
8942, PTBG), the latter species tends to grow in wet forest understory either terres-
New pteridophyte species and combinations from the Marquesas Islands...
49
trially or on mossy boulders, although it is sometimes found in shady depressions on
steep ridge slopes either as individuals or forming small colonies.
Specimens examined. Marquesas Islands: Hiva Oa: Temetiu, windswept ridges and drainages, 3900 ft, 9°48'S, 139°4'W, Wood 4392 (P, PAP, PTBG, UC, US);
chemin d’Atuona a Hanamenu par Feani, pente vers Hanamenui, Schäfer 5195 (US);
Atuona–Feani trail, crest of Feani ridge, Sachet & Decker 1127 (US), 1192 (US).
Acknowledgments
The extremely useful discussions, field observations, photos, and specimens provided by
Jacques Florence of IRD [ex ORSTOM], Paris in conjunction with the ongoing Flore
de la Polynésie française project, Jean-Yves Meyer (Delegation à la Recherche, Polynésie
française), and Steve Perlman and Ken Wood (National Tropical Botanical Garden) all
have contributed significantly to this study. Important observations and new collections resulted from fieldwork carried out during the 1988 Fatu Hiva Expedition and
subsequent trips in 1995, 1997, 2003, 2004, and 2005 as part of the Vascular Flora of
the Marquesas Islands project, which was supported by a generous project donation from
private sponsors. We are grateful to the Delegation à la Recherche (Papeete, Tahiti),
the Musée de Tahiti et des Iles, and the Service du Développement Rural for logistic
support and permission to collect in the Marquesas and to the Delegation and the
Haut-Commissariat de la République en Polynésie française for permission to conduct
research there. We thank the curators of the following herbaria for loans or permission
to study specimens: BISH, K, MO, NY, P, PAP, PTBG, UC, and US. The magnificent
illustrations were skillfully drawn by Alice Tangerini (Smithsonian Institution), often
from large or otherwise challenging specimens. Val Romano (Smithsonian Institution)
kindly assisted with locating, managing, and updating of specimens. We thank Alain
Touwaide (Smithsonian Institution) for translating the Latin diagnoses. The contribution by WLW to this study was partially supported by the Smithsonian Research Opportunities Fund and while appointed as McBryde Chair at NTBG. Support for ARS
to study specimens at NTBG was provided by the McBryde Senior Research Fellow
program. We wish to thank the reviewers (Thomas Ranker and Kunio Iwatsuki) for
valuable suggestions and comments that improved the final version of this paper.
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