ENTOMOLOGIST'S MONTHLY
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43
TAXONOMY, DISTRIBUTION AND NESTING BIOLOGY OF
WSPA BINGHAMI, U BASALIS, V UARIABILIS, U FERWDA,
U. LUCTUOSA, U MULTIMACULATA AND U BELLICOSA
(HYM., VESPINAE)
BY MICHAEL E. ARCHER
This paper completes the coverage of the remaining seven species of
in previous papers (Archer, 1991b, L992, 1993,
I994b, 1995,1996). The genus Vespa is considered to contain 23 species
(Archer, 1991a). Keys to the species are given in Archer (1989) and a
cladistic analysis to show possible phylogenetic relationships in Archer
Vespa not considered
Q99aa). Van der Vecht (1957,1959) provides important taxonomic papers
onVespa species giving much geographical distribution information.
Vespa binghami du Buysson, 1905
This species consists of two colour subspecies: the western nominate
subspecies with a largely dark brown unicolorous gaster and the eastern
V
binghami suprunenkoi Birula, 1925 with light and dark brown bands on
the gaster. Intermediately coloured females with less extensive light
brown gastral bands than those of V binghami suprunenkoi have been
found in Sichuan, China. Males may show a greater range in colour
variation than the females. A male V binghami suprunenlcoi has been
found in northern Thailand where the females are the nominate
subspecies.
The geographical distribution of V binghami (fig. 1) is: India (northem
West Bengal, Meghalaya), northern Burma, Bhutan, northern Thailand,
northern Laos, China (Tibet, Sichuan, Yunnan), Russia (Sakhalin
Primor'ye ["Maritime Kray']) and Korea.
Very little is known about the nesting biology
of V
bingharni.
Information on labels attached to specimens (van der Vecht,I959;Archer,
unpublished) indicate that this species is found between the altitudes of
20Om and 273Om, usually over L000m, and is active from March until
early November. The life cycle is annual and colonies are synchronised in
their development. The queens are larger than the workers and readily
separated by size.
Vespa basalis Smith, L852
V basalis is widespread in continental Asia extending onto the islands
of Sumatra and Taiwan. Its detailed geographical distribution (fig. 2) is:
Pakistan (Islamabad), India (Uttar Pradesh, Himachal Pradesh,
Meghalaya, Assam, northern West Bengal), Nepal, Burma (Kachin, Shan,
Karen, Tenasserim), northern Thailand, Laos, northern Vietnam, China
(Sichuan, Yunnan, Fujian), Korea, Sumatra and Taiwan. Further
distribution information is given for Nepal by Yamane (L974), Taiwan by
Starr (1992), and Korea by Kim et al. (1994).
29th March, 1999 Vol. 135 (1999)
44
ENTOMOLOGIST'S MONTHLY MAGAZINE
Fig. 1.
-o>=-2?
Fig.2.-The
world distribution of Vespa basalis.
45
MAGAZINE
The nesting biology of V basalis is known to some extent from
ENTOMOLOGIST'S MONTHLY
Williams (1919), van der Vecht (1957,1959), Yamane, 56. (1977, 1992),
Matsuura (1973), Yamane, Sk. (1974), Kuo & Yeh (1985), and Muzaffar
& Ahmad (1986). Information on labels attached to specimens has also
proved useful (Archer, unpublished). I am grateful to Dr S. Martin for a
translation of Kuo & Yeh (1985).
Typically, nests are found in mountain forests attached to the branches
of trees, sometimes in shrubs and on buildings, in rock crevices and in the
ground. Aerial nests in trees are usually at least four metres above the
ground. Underground nests are shallow so that the nest is visible from the
entrance. Nests which are started in holes are relocated during May or
June to aerial sites. In Taiwan, nest densities of ten nests per 150ha have
been found.
In Nepal, nests are found between 90Om and 1500m although adults
have been found between 700m and 1500m. In Taiwan, nests are found
from 100m up to 1500m altitude but mainly between 200m and 800m.
Generally, throughout the range of V basalis, adults have been found
between 1,0m and 2830m, but mainly between 500m and 190Om.
Queens overwinter in holes and crevices in walls, trees, and even in old
nests. The overwintering period is usually from late December until early
March, although adult activity has been recorded during January and
February. In Taiwan the queens establish nests during March and April
with the first workers appearing during April and May. Sexuals emerge
from October until December. The life cycle is very long: about nine
months or 270 days.
Queens carry out inter-specific usurpations on Vespa tropica (L.) at the
end of April (Kuo & Yeh, 1985), and on V velutina l-epeletier (Yamane,
reez).
At first, nests in bushes are egg-shaped with a short (2-3cm) ventral
tube. After relocation, the nest shape is at first spherical, becoming pearshaped with a conical roof. In one nest the conical roof occupied about
I2Vo of. the volume of the nest. The entrance to the nest is at first circular,
but later becomes a vertical slit (about Lcm wide and 17cm long). Mature
nests usually have three slit-like entrances on thc sides of the envelope.
Mature nests vary in height (22-90cm) and width (LG70cm), and in
the number of combs (5-16 or more). One large nest was estimated to
contain 40,100 cells, of which 27,600 were small cells used to rear the
workers and some males, and I2,5O0 large cells used to rear the queens
and most of the males. The small cells (diameter 7.1mm) were distinctly
smaller than the large cells (diameter 9.4mm). The upper older combs
consisted of small cells and the lower combs of large cells. Central combs
consisted of small and large cells with the proportion of large cells
increasing in the lower combs.
In Taiwan, the maximum number of immature brood is reached during
October. In a large colony the number of immature brood was estimated
to be in excess of 25,000 including sealed brood of 1,848 workers, 2,187
46
ENTOMOLOGIST'S MONTHLY MAGAZINE
queens, and2,74}males (about T40insmall cells and 2,000 in lar_g_e,celll)In ttris large colony, meconial counts showed that about 30,000 cells
(76.8Vo of Cells) had produced at least one adult: 44.9Vo of cel.ls producing
only one adult;'28.5% two adults; 3.3Vo three adults, and 0.1Vo four adults.
'itre developmental period of the brood varied from 4143 days (egg
stage 6-7 day3, larva L5 days, sealed brood 2V2I days). It is not clear
whlther this developmental period refers to the workers, queens or males.
Normally a colony reaches a maximum Population of about 6,000
workers, alihough a lirge colony may have L0,000. In one lq_gt colony, it
it would pioduce 37,600 workers, 4,000 males and
3,400 queens. Workers and queens differ in size.
was estimated ihat
Vespa variabilis du Buysson, L905
'itris sp""ies consist-s of two colour subspecies: the eastern nominate
,
subspeciei with the first to the fifth gastral terga having alternating black
or dark brown and light brown bands, and the western V variabilis fumida
van der Vecht, 1959 with the first to the fifth gastral terga black.
The geographical distribution of V variabilis (fig. 3) is: India (northern
West nEngit, Arunachal Pradesh, Sikkim), Nepal, Burma (Sagaing) and
China (Sichuan, Yunnan, Hubei, Fujian).
Fig. 3.
-
The world distribution of Vespa variabilis.
Very little is known about the nesting biology
of U
variabilis.
Information on labels attached to specimens (van der Vecht,l959;Archer,
unpublished) indicates that this species is found at altitudes between 670m
and 3000m but usually over 1000m, and therefore is a montane species.
Adults were collected from the end of February until the end of
November. The life cycle is annual with the queens overwintering for two
to three months. Colonies are synchronous in their development.
V fervida Smith, 1859
U fervida and U luctuosa are a
species
pair united by
the
synapomorphy of the apical margin of the seventh gastral sternum of the
male with two emarginations (Archer, I994a).
ENTOMOLOGIST'S MONTHLY
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47
V fervida is restricted to Sulawesi (Indonesia) but is found throughout
this island including its southern islands of Salayat and ButangVery little is known about the nesting biology of V fervida except that
it is found in forested areas (personal communication, G.R. Else, 1986).
Information on labels attached to specimens (van der Vecht, t957, 1959;
Archer, unpublished) indicate that this species is found at altitudes from
100m to 1080m so that probably it is restricted to forested areas rather
than a particular altitudinal range.
The- queens and workers clearly differ in size. Queens are found
throughout the year so it is possible that colonies are asynchronous in their
development.
V luctuosa de Saussure, 1854
The nomenclatural history of V luctuosa is complex. Bequaert (193a)
recognised five colour varieties of V luctuosa: 1, The nominate colour
variety; 2, semperi du Buysson, L905; 3, malayana, nom. nov- for
annulata Smith, 1858; 4, bellicosa de Saussure, 1854; and 5, luzonensis,
nova var.
Van der Vecht (1957) raised bellicosa to species rank and recognised
malayana as a synonym of the subspecies V multimaculata multimaculata
Perez,19L0. The other three colour forms of V luctuosa were considered
to be subspecies.
Kojima & Tano (1985) described V luctuosa negrosensls based on two
queens, which could not be distinguished from V luctuosa luctuosa, and
fbur workers, with very minor colour differences from V. luctuosa
luctuosa. With further study of a larger number of specimens this new
colour variety might be considered a microgeographic variation but not a
macrogeographic variation or subspecies. As such V luctuosa negrosensis
should be treated as a syn. n. of V luctuosa luctuosa.
V luctuosa is restricted to the Philippine Islands. V luctuosa semperi
(southern colour form) has been found on the island of Mindanao and is
distinguished from the other colour forms by its yellow sixth gastral
tergum. The other two colour forms have a black sixth gastral tergum.
V luctuosa luctuosa (northern-central colour form) has been found on
the islands of Luzon, Mindoro, Panay, Negros and Cebu. It is recognised
by the narrow yellow apical band on the first to the fourth or fifth gastral
terga. The width of the yellow band on the second gastral tergum extends
about one sixth of the way across the tergum.
V. luctuosa luzonensis (eastern colour form) has been found on the
islands of Leyte, Biliron, Panaon and Samar. It is recognised by the wide
yellow apical band on the first to the fourth or fifth gastral terga. These
yellow
bands extend at least one quarter of the way across the tergg.
Kojima (1982) and Kojima & Reyes (1936) give keys and distribution
notes on the colour forms of V luctuosa.
Alittle information is available on the nesting biology of V luctuoss in
Williams (1919), Kojima & Yamane (1980), Kojima (1982) and Starr &
48
ENTOMOLOGIST,S MONTHLY MAGAZINE
Jacobson (1990). Information on labels attached to specimens. and the
specimens'themselves (van der Vecht, 1957 ;Archer, unpublished) provide
information on female caste differences, altitude distribution, and months
of activity.
Nests are usually found in forested areas attached to the branches of
trees from Lm to 6m, but sometimes up to 2Om, above the ground. Nests
also have been found in bushes, and one was attached to a ceiling of an
abandoned house.
queen nest of V luctuosa luctuosa consisted of a single comb
completely covered by an envelope with a ventral entrance extended into
a ,rrty short (about 3mm long) cylindrical tube. Two queen nests- of V
luctuosa luzonensis had bell-shaped envelopes so that from a ventral view
A
the single comb was clearly visible. Later nests are . spherical and
completely covered by envelopes except fo1 a ventral circular or oval
entrince situated to one side. Mature nests elongate vertically but lack
a
roof cone.
Probably up to six combs are present in mature nests with about'l',250
cells, alttrougtr larger nests have been seen. Small and large cells are
s"putit"d. ftre oldest comb, at the top of the nest, consists of small
readily-occisionally
with a few peripheral large cells, while the other combs
cells,
consist of large c-ells, sometimes with small cells in the second comb.
Cells in the oldest comb have up to five meconia per cell, indicating
that a cell has been used to rear five adults. The second comb may have up
to four meconia per cell, while younger combs have progressively fewer
meconia per cell.
Adulti have been found from 2OOm to 150Om, and to be active in all
months except February. Males have been found during January, March,
May, September, and October. Queen, small and large nests all have been
found during March. Thus it is highly likely that colonies
are
asynchrononJitr their development. Queens and workers clearly differ in
size.
V multimaculata Perez, 1910
U multimaculata and U bellicosa are
synapomorphy just
a species pair although a
for these two species has not been found (Archer,
1994a).
Van der Vecht (L957) described the western colour form,
V multi-
maculsta pendleburyi, with wider yellow gastral telgal bands, e.g. on the
second teigum, at short distance from the middle of the tergum, the
yellow band extends about one quarter across the tergum. In the eastem
colour form, the nominate subspecies, it only extends about one sixth
across the tergum. I have seen specimens of V. multimaculata pendleburyi
and V multiiaculata multimacilata together at the same site both in the
range of the westem colour form (in Peninsular Malay-sia) Td the eastern
i
colSur form
maintained.
(in
Sabah).
Th" two
subspecies therefore should not be
ENTOMOLOGIST'S MONTHLY
The geographical distribution
of U
MAGAZINE
multimaculata
is:
49
Singapore,
Malaysia (Peninsular, Sarawak, Sabah), Brunei, and Indonesia (Sumatra,
Kalimantan).
Very little is known about the nesting biology of V multimaculata.
Martin (1995) found that it nested in the ground, especially under the roots
of trees with nests from 20 to 30cm in diameter. Nests are associated with
both lowland and montane forests (van der Vecht, 1957; Martin, 1995).
Information on labels attached to specimens (van der Vecht, 1957,
I959;Archer, unpublished) indicate that this species is found at altitudes
between 50m and 21,40m. Adults have been found throughout the year
except January. Queens have been found active away from the nest during
February, April, July and October, and males have been found during
February, March, April and July. Thus it is highly possible that colonies
are asynchronous in their development. Queens and workers clearly differ
in size.
V bellicosa de Saussure, 1853
The geographical distribution of V bellicosa is: Indonesia (Sumatra,
Kalimantan) and Malaysia (Sarawak).
Very little is known about the nesting biology of V. bellicosa. It is
with lowland forests (van der Vecht, t957; Martin, 1995).
Information on labels attached to specimens (van der Vecht, 1957,
I959;Archer, unpublished) indicate that this species is found at altitudes
between 75m and I25m. Adults have been found throughout the year
except for March and December so that it is highly likely that colonies are
asynchronous in their development. Queens and workers probably differ
associated
in size.
REFERENCES
Archer, M.8., 1989, A key to the world species of the Vespinae (Hymenoptera). Part 1,
Key, Checklist and Distribution: 147 Part 2, (72 figs): t1-34), Research Monograph of the
College of Ripon and York St John, No. 2; l99la, The number of species that can be
recognised within the genus Vespa (Hy-., Vespinae), Entomologistb mon. Mag., 127;
16l-164;1991b, Taxonomy and bionomics of the Vespa tropica group (Hym., Vespinae),
ibid., 127: 2?5-232; 1992, T\e taxonomy of Vespa crabro L. and V. dybowskii Andre
(Hym., Vespinae), ibid.,l28 157-163;1993, The life history and colonial characteristics of
the Hornet, Vespa crabro L. (Hym., Vespinae), ibid., 129 1,51-163; 1994a, A phylogenetic
study of the species of the genus Vespa (Hymenoptera: Vespinae), Ent. scand.,24:469478;
1994b, Thxonomy, distribution and nesting biology of the Vespa bicolor group (Hym.,
Vespinae), Entomologist's mon. Mog., 130: 14L158; 1995, Thxonomy, _distribution and
nesting biology of the Vespa mandarinia group (Hym., Vespinae), ibi.d., l3l: 47-53;1996,
Taxonomy, distribution and nesting biology of Vespa affinis and Vespa mocsaryann du
Buysson (Hym., Vespinae), ibid., 133l. 27-38. Bequaert, J., 1930, On the generic and
subgeneric divisions of the Vespinae (Hymenoptera), Bull. Brooklyn ent. Soc.,25: 59-70;
L934, I-es races de coloration de Vespa luctuosa de Saussure et de Polistes tenebricosus
r
Lrpeletier, Bull. Mus. Hist. nat. Belg.,10: 1-11. Kim, J.K., Moon, T.-Y. & Yoon, I.8.,
1994, Systematics of vespine wasps from Korea, I. Genus Vespa Linnaeus (Vespidae,
Hymenoptera), KoreanJ. Entom.,24:1'07-11'5. Kojima, J.,1,992? The genusVespa in the
ftritipplnes (Hymenoptera, Vespidae), KontyA, 50: 4344M. Kojimar_J. & Reye^s, S.G.,
t986,?tritippine Vespidae, I, Genus Vespa, Philipp. Ent.,6:445454. Kojima, J. & Thno'
50
ENToMoLocrsls MoNTHLy MAGAZINE
1985, A new subspecies of Vespa luctuosa from Negros Island, ibid.,6: 43H37.
Kojima, J. & Yamane, Slc, 1980, Biological notes on Vespa luctuosa luzonensis from
lryte Island, The Philippines, with descriptions of adults and larvae (Hymenoptera:
Vespidae), Insecta matsum.,19:79-87. Kuo, M.C. & Yeh, W.H., 1985, Studies of Vespidae
T.,
in Taiwan. II. Ecological studies of Vespa basalis Smith, Vespa velutina flavitarsis Sonan
and Vespa tropica pseudosoror Vecht, J. Natn. Chinyi lrut. Agric., 11: 95-106. Martin,
S.J., 1995. Hornets (Hymenoptera: Vespinae) of Malaysia. Malayan Nature J., 49:71-82.
Matsuura, M., 1973. Nesting habits of several species of the genus Vespa in Formosa,
Kontyfi,4l:28G293. Matsuura, M. & Yamane, Slc, 1990, Biology of the vespine wasps,
Springer-Verlag, Berlin. Muzaffa4 N. & Ahmad, R., 1986, Studies on hornets attacking
honeybees in Pakistan, Pak J. agric. Res., 7: 5943. Staq C.K., 1992, The social wasps
(Hymenoptera: Vespidae) of Thiwan, Bull. Nat. Mus. Nat..Scr.,3:9!138. Star, C.K. &
Jacobson, R.S., 1990, Nest structure in Philippine Hornets (Hymenoptera, Vespidae,Vespa
spp.),Jap.J. Ent.,58:125-143. VechtJ. van der,l95T,TheVespinaeof the Indo-Malayan
and Papuan areas (Hymenoptera, Vespidae), Zool. Verh. Leiden,34: 1-83; 1959, Notes on
Oriental Vespidae, including some species from China and Japan (Hymenoptera, Vespidae),
Zool. Meded.,36:205-232. Williams, EX., 1919, Philippine wasp studies, 2, Descriptions
of new species and life history studies, Bull. Hawaiian Sug. Pltrs' Ass. Srn. (Ent. Ser.), 14:
119-186. Yamane, Sk, 1974, On the genus Vespa (Hymenoptera, Vespidae) from Nepal,
KontyL, 42: 29-39. Yamane, 56., 1977, On the collecting of vespine nests, based on
practices through a survey in Thiwan from 1972to 1974 (Hymenoptera: Vespidae), Seibutsu
Ky6zai 12: 42-59; 1992, A huge nest of Vespa basalis collected in Taiwan (Hymenoptera:
Vespidae), Chinese J. Entom.,12: 1-11.
The University College of Ripon and York St John, l,ord Mayor's Walk, York, YO3 7EX,
England, U.K.
November 13th,1996.
Platyspathius sp. (Hym., Braconidae, Doryctinae) in S.W France, new to the
Palaearclic Region
A single male specimen of a species of Platyspathius Yiereck, 191 I
was collected in a Malaise Trap operated at Bernac, l-ot-et-Garonne, France by R.R. Askew
between 29 June and 31 July 1995. The trap was situated at the edge of ancient mature
woodland (Quercus, Juniperus, Cornus, Ace4 Prunas, etc.) and collected several rare
insects characteristic of such sites (e.g. Shaw, 1996, Entomologist\ mon. Mag.,132:204).
Platyspathius is a relatively small genus, previously known only from the Afrotropical and
Indo-Australian regions. As far as is known they are parasitoids of insects living in dead
wood. The French specimen runs smoothly to the type species of the genus, P pictipennis
Viereck, in Nixon's key (1943, Trans. R. ent. Soc. Lond,93: 173456) by virtue of the
distinctive structure of its hind leg and (while being near the edge of the range of variation
in its relatively light reddish coloration, rather weak sculpture, and slender build) it agrees
well with the seven male specimens of this species from Uganda and Sudan preserved in
the BMNH. However, more specimens, and especially females, would be needed to decide
its specific identity. The record is brought forward now because the Askews have moved on
from their former house at Bernac, substantially reducing the hope that more specimens can
be obtained, not only to demonstrate presence of a breeding population but also to enable
identification to species. I am grateful to Dick Askew for giving me the ichneumonoid
portions of Malaise Trap catches, to Sergei Belokobylskij for his opinion on the specimen,
and to Mike Fitton for access to the BMNH collection. The French specimen is deposited
in the National Museums of Scotland.
M.R. SHAW National Museums of Scotland,
Chambers Street, Edinburgh, EHl lJF: January 28th,1998.
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