(lepidoptera, saturniidae)
TRANSCRIPT
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ectdiit Trans, tepid. Soc, .1apan 56<3):184-192,June 2005
Life history ofActias ignescens (Lepidoptera, Saturniidae) reared on its natural
host plant, crimpteronia paniculata, on South Andaman Island
K. VEENAKuMARIi)*, PRAsHANTH MoHANRAJi'* and Richard S. PE[Gi.ER!''' Centra] Agricultural Research Institute, Port Blair, Andaman & Nicobar Islands. India
" Department of Biology, University of the Incarnate Word, 4301 Broadway, San Antonio, Texas, 78209-6397 U.S.A.
Abstract The immature stages and previously unknown female ofActias ignescens, an endcmic of
thc Andaman Islands, are dcscribed and figured in colour. The lurval host plunt in nature isCopterenia panicutata (Crypteroniaceae). There are two generations annually, with adults on the
wing in January and again in August and September. The immature stages are compared with
those of the nearest related species ol' Actias.
Key words Actias. Andamans, Ct ypreronia, India, meon muths, Myrtales, saturniid.
Introduction
Actias, a monophyletic genus (Shimada et at., 1995; Regier et at., 2002), is represented bytwo endernic species. A. catlandra Jordan and A. ignescens Moore, in the Andaman Islands,located in the Bay of Bengal. These along with Antheraea andamana Moore, which is an-other endemic species, were the first three saturniids to be reported from the AndamanIslands (Moore, 1877), Actias ignescens is a sexual]y dimorphic species that has beenfound only on South Andaman, one of the three 1arge islandg in the archipelago, which to-
gether with North and Middle Andamans constitute the Great Andaman. The Andaman and
Nicobar Islands comprise a political subdivision of India. Various authors cited in the fo1-
lowing paragraph classified Actias ignescens and A. maenas Doubleday in the generaArgema or Sonthonnaxia. However, Actias differs from the African genus Argema by the
fact that the tails of the hind wings are never crossed when the moths are at rest in Actias,but are in Argema, and the cocoons of Argema are dense and have pre-fbrmed exit valves,
whilst those of Actias are papery and lack the valves. The name Sonthonnexia is ajuniorsubjective synonym ofActias,
Acti.as ignescens was first described by Moore (1877) from the island of South Andaman.Later, Cotes (1891-1893) concluded that it was merely a form ofActias ntaenas Deubleday,but referred to maenas by the synonym leto, This he surmised solely on the basis of
Moore's scanty description, as no specimens ofA. ignescens existed in the Indian Museumand so could not be examined by him. Based mainly on the distribution of the dark areason the wings, Hampson (1892) distinguished it from A. maenas and retained it as a separate
species, Sonthonnax (]899) in his study of the silk-producing Lepidoptera synonymised itwith Atgema maenas Doubleday, He also figured a drawing of a male based on a specimen
in The Natural History Museum (London), labelling it Argema maenas variety ignescens.This was the first illustration of the species. Seitz (1928), who was a strong proponent ofthe subspecies concept, predictably arrttnged ignescens under maenas in his widely avail-
able standard reference. Arora & Gupta <1979), in their monograph on the non-mulberry
*Prescnt addrcss: Pro.jcct Directerate of Biologica] Control, P. B. No. 2491, H. A. Farm Post,
Hebbal, Bangalore-560024, India.
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Life History oi' A('tias i,gneseen{' 1S5
silk moths of lndia, synonymised A. ignescens with Actias ntaenas. once again without cx-
amining specimcns from the Andaman Islands. Thc 53 specimens ei' A, maenas in the
Indian Museun]. Calcutta. which Arora and Gupta examincd camc from Sikkim. Assam.Meghalaya, Shillong. and Dar.jeeling. Both Peigler (1989) and Nassig et al. (1996a) ho"s-cver. were ot' the opinion that the insu(ar A. igne,scens is significantly different fronM'e]atedforms in the genus to merit treatmcnt as a distinct species. D'Abrera (1998) prcscnted a
colour photograph that was life-size of a malc in exccllent condition from Moore's original
type serics, and designated that specimen as lectotype. In this report we describc the fe-male for the first time, and figure it in coJour. No fema]c specimens QfA. ignesc'ens are inThe Natural History Museum, which is why D'Abrera did not figure it. The naine ig-nescens refcrs to fire, so perhaps Moore chose thc name because of the burnt, brownish or-angc colour of the male,
Methods
The methods that we fo11owed in collecting and rearing A. ignescens were idcntical to thosewe adopted in our earlier studies on the Saturniidae of these islands (Veenakumari et ctt.,
1995, 1996: Prashanth Mohanraj et al.. 1996, 1998; Prashanth Mohanraj & Veenakumari.2002). One significant departure ",as the manner in which we confincd the adults, which
wc detail below.
Field collection
Two ]ate instar larvae collectcd on Cr}vJtetvnia pani('utata Blume (Crypteroniaceae) inSeptembcr 1998 from Mount Harriet National Park, South Andaman Island, pupated in thelaboratory. The resulting adults propitiously turned out to be a male and a female. Sincethe male emerged eartier we p]aced it in the refrigerator till the fernale a]so emerged when
we released the pair into a large cylindrical "'ire mesh cage that we improvised. Cloth wasscwn to thc two open ends of the cage and hunLg from the ceiling by a thread (Figs 1-2- ),
ws
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Figs 1--2, XNiire mesh cagc for mating and egg laying ofA('tias i,gnes('ens. 1. View oi' (hc "'holc
cage suspcndcd from thc cei]ing. 2. C]ose-up oi' confincd adults of .tL. igites('ens.
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Table I. Durations and dimensions of the immature stages ofActia.y ignescens on South Andaman,
Larval instarsEgg
I II III IV vPupa
Durations(days)
Ma]e
Fema]e
*
*
S.54± O.88 4.9Sti).7S 4.95 ±O.59 n=13 n=22 n=21
5.29± 1.86 5.05±O.69 4.86ti).57
n=17 n=20 n=21
6.0e±O.97 9.30±1.38 30.43±S.51
n=20 n=20 n=23
623±O.8] 11,4±1.70 35.53± 19.67
n=22 n=20 n=15
Dimensions(mm)
2.6] × 2.95 × 1.94 ±
Allindividuals O.03xO.05xO.08 9.6±O.7
n=14 n=1316.5
± 1.0 27.4± 1.6 43.0± 1.9 65.6±4.8 59.0× 23.0**
rt=43 n=29 n=19 n=22 n=2
' lncubation period for a]1 the eggs=9 days (n=47).
'" Male cocoons measured 67.9 × 31.3± 3.9 × I.6 mm (n=9). Female cocoons measured 8l,O× 37.3±4.1 × 2.4
mm (n=6).
We also released a second pair of freshly emerged adults which were caught resting on veg-
etation, about 2 metres above the ground, in Mount Harriet National Park about 2 km away
from the spot where we collected the earlier pair of larvae. We did not observe mating inthe cage, but both females began laying eggs. It is possible that the adult female col]ectedfrom the fOrest had mated before being captured by us. A total of 113 eggs were laid indi-vidually or in small clusters and were stuck to the surface ef the cloth or onto the metal sur-face of the wire mesh,
Results
Description of immature stages
We examined al] the immature stages of the moth under a microscope and described andmeasured them in the living condition (Table 1 & Figs 3-15),
Egg (Fig. 6), The eggs are spheroid; unifbrmly ash-grey to pale blue, with the usual shal-
low polygonal depressions giving a honeycomb appearance to the chorion, The majority of
the eggs however were dult, pale brown in colour as they had been smeared by the deepbrown viscous adhesive fiuid used by the females to glue their eggs onto the substrate. Themicropyle is situated at the centre of a dark grey circle,
First instar (Fig. 7). Head glossy black; bearing short transparent setae; antennae basallywhite and terminally brown; labrum and mandibles brown; clypeus white; thorax and pro-thoracic shield yellow-orange; a narrow transparent area along posterior margin of protho-racic shield through which internal contents of body are visible. On hatching these Iarvaewalked around the container, drank water and then sett]ed down in the characteristic sphinxposition on the undersurfaces of leaves. Abdornina] segments I to IV with black dorsal sad-dle extending up to the lateral scoli. Below the scoli these segments are yellow-orange, likeall the other segments. Anal claspers and margins of anal plate black. Scoli are terminally
transparent and bear transparent setae. Dorsal scoli on eighth abdominal segment fused,Ventral surface of larva whitish, Proximal two-thirds of legs black while the distal one-third is whitish, bearing brown claws. Prolegs, white with brown crochets,
Second Instar (Fig, 8). Head glossy and light brown except along the edges of the fronswhere it is a deeper shade of brown; these 2 bands converge towards, but do not meet at theapex of the frons. There are two prominent black spots behind the pale orange dorsal scoli
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on the anterior leading edge of the prothoracic shield. These black spots are separated by a
green band of the same shade as the rest of the body, Dorsal and latero-ventral scoli have
bright orange heads with black bristleslsetae on the thQracic sco]i while some are brown on
the abdominal scoli. Dorsal scolus on abdorninal segment 8 is fused. The subdorsal scoli
are the smallest and are further distinct in being golden honey co]oured and bearing brown
setae. Distributed a]1 over the green body surface are small white cupolae, each of which
bears a short, black terminal seta. The yentral surface is also green with brown legs and
claws. Prolegs bear a terminal brown patch and brown crotchets. The spiracles are whitish
and connecting the spiracles runs a pale yellow line or band.
Third instar (Fig, 9), Head glossy light brown and mottled; base of antennae cream;
clypeus and frons whitish; labial palps white at the base. No black spots on prothoracicshield, only a faint pale green line runs along the middle. Be]ow the orange head the dorsal
scoli on the mesothorax and metathorax sport a brown patch. BaTTing the head, the whele
larva is uniformly green. The dersal and ventral scoli are tipped with orange and bear black
and brown setae. Subdorsal scoli are small and yellowish. Dorsal scolus on abdominal
segment 8 is fused. There is apale green subspiracular stripe running along the length of
the larva and connecting the latero-ventral scoli. The whole surface of the body bears small
whitish cupolae tipped with short, brown setae. The anal plate is yellow with a deep brownarea along the anal edge and it also has brown spots from each of which arises a single
brown seta. The anal prolegs are brown distally and proximally cream.
Fourth instar (Fig. 10). Head glossy brown and sparsely setose. The setae are of variable
length with the longer ones being brown and the shorter ones being white. The labrum is a
deeper brown than the rest of the head and the mandibles are the deepest brown with the
serrated edges being black in colour. The firons is pale brown to dirty white. The larva is
green with distinct narrow yellow circum-annular bands along posterior margins of abdom-
inal segments 2 to 5. The subspiracular row of scoli divides the larva longitudinally into 2
shades of green lighter above and deep green below it. There is no subspiracular yellow
stripe, Spiracles are pale brown with a yellow halo and thin yellow longitudinal stripe.
Prolegs are terminally brownish-black with brown crochets. The dorsal and subspiracular
scoli (Fig. 15) are orange tipped with yellow-green dots from each of which arises a brown
spine. The setae progressively reduce in size from the anterior to the posterior segments as
does the intensity of the orange colour, such that the posterior most scoli become totally
green in colour. The dorsal scoli of the eighth abdominal segment are fused. The anal plate
(Fig, 12) is cream with small, brown warts each of which bears a short, brown, terminal
seta. The anal proleg (Fig. 11) is brown-black distally with glossy warts of the same
colour; each of the warts bears a whitish or a brown seta.
Fifth instar (Fig, 13). This is also a green larva with white dots resembling the fourth instar
but the cream, circum-annular bands are wider along posterior margins of abdominal seg-
ments I to VII and are more prominent. The rear margin of the prothoracic shield bears an
irregular patch whitish pale blue or pale grey. Both instars rest in the position with the head
raised above the substrate as is characteristic of a]l species ofActias. Ventrally, the regions
between the thoracic prolegs as well as that between the anal prolegs are very pale, almost
white in colour contrasting with the colour of the rest of the venrer. The subpiracular, yel-
!ow stripe is absent as in the previous instar. The late instar larva makes a scratching noise
by scraping its black mandibles against the base of the fOrelegs when disturbed. The shape
and general appearance of the mature larva closely resembles that ofActias maenas (Ntissig
& Peigler, 1984; Nassig et al., 1996b), but the whitish setae are almost all gone in the final
stage of A. ignescens, as is the case with the eastern Indonesian A. grvenendaeti Roepke
(Paukstadt & Paukstadt, 1993), The larva of A. ignescens is easily distinguished from that
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Life History ofActias ignescens 1g9
of the centraHndenesian Actias isis (Sonthonnax,) (Naumann, 1995: colour pl, 2): the latter
has a darker head, blackish scoli, prominent white setae, and much less yellow on the anal
prolegs.
Cocoon (Fig, 14). Papery white with perforations, constructed along a twig.
Female (Fig, 3). Forewing length 80 mm; hind wing length 137 mm inclusive of the tai];
n:=1. Ground colour solidly yellow; body pale yellow; thoracic collar, costal margin, and
outer margins purplish brown, dusted with gray; forewing ocellus (eyespot) crescentic, at-
tached to costal band, black, surrounded by dark yellow; antemedian line purplish, thin,
slightly angled; postmedian line brownish, broken into series of dashes; tails broader thanin male (Fig. 4), dusted with purple along centres. twisted into broader yellow fiaps termi-
nally, The female ofActia,s ignescens actually resembles most closely that ofA. isis frem
central Indonesia with their shared du]1 yellow colouration, and weak brown markings, The
females ofA. maenas are decidedly green in their ground colour, with fewer markings. The
female ofA, isis was figured in coleur by Naumann (1995) and D'Abrera ( 1998),
Specimen examined: 1 4 from Mt Haniet, South Andaman Island. 24. xii. 2001, leg. K.Veenakumari and Prashanth Mohanraj; in coll, Project Directorate of Bio]ogical Control,
Bangalore, India.
Field observations
The larvae of A, ignescens feed on leaves of Ci77)teivnia paiticttlata, a deciduous tree of
moderate size that attains a height of 12-18 metres. Although it grows in the lowland
forests of both the Middle and South Andamans, it is only on the tatter that we have found
this moth. The larvae fed on the new flush of tender green leaves ringed with purple, which
the tree puts forth after it sheds its old leaves, Ctyptervnia panicttlata is also now known to
be one of the larval food plants ofAttac'us mcmulleni Watson, but we crroneously reported
this plant as l/itex glabrata R. Brown in Veenakumari et at, (1995).
The larvae and cocoons were found in the trees at heights ranging from 6-10 metres. The
branches on which the larvae were present were extensively defoliated. The complete lami-
na of many leaves had been eaten with the bare petioles visible starkly against the blue of
the sky. Freshly hatched larvae when released on trees of C. panicttlata dispersed and dis-
tributed themselves with one larva only per ]eaf. Many of these completed their life cycles
on the trees on which they were released. The females that emerged rested on their respec-
tive cocoons for 3-4 days after which they fiew away possibly due to human disturbance.During this period we did not notice mating by any of the females. On the other hand the
males that emerged from the cocoons flew away, They were not seen to mate with the pre-viously emerged females.
Regarding the phenology, the larvae. cocoons and freshly emerged adults of both sexes
have been collected by us in the months of January, August and September. During this pe-riod we also found larvae of Samia .ftztva Jordan on Heteropanax ,f7agrans (Roxb.) Seem
(Araliaceae) (see Prashanth Mohanraj et al.. 1998: Peigler & Naumann, 2003), as well as
larvae of Actias catlandra on a tree known locally as pyinma (Lagerstroemia h.}・poteuc'a
Figs 3-IS. A('tias ignescens. 3. Femalc. 4. Male, 5. A malc-female pair. 6, Egg. 7. 1 instar, 8. II
instar. 9. M instar. 10. IV instar. 11. Claspers of anal prolcgs (e]ose-up). 12. Anal plale
(close-up). 13, V instar. 14. Cocoon, 15. Dorsal scoli.
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190 K. VEENAKuMARI, PRAsHANTH MoHANRAJ and Richard S. PEIGLER
Kurz, Lythraceae) in the same habitat in Mount Harriet National Park,
Discussion
The British occupied the Andaman lslands in 1858. By the end of the nineteenth century
three species of the Saturniidae, Antheraea andamana, Actias cattandtn and A. ig. nescens,
were known from these islands, In the next fburteen years three more species, Cricuta an-damanica, Samia julva, and Attacus mcmuUeni, were added to this list. No further addi-tions were made to the saturniid fauna of these islands ti11 2001 and 2002 when threespecies of Antheraea, viz., A, cernyi, A, meisteri and A, rudtcuffi were described from theAndamans (Brechlin, 2002), andA, insutaris Watson was eievated to fu11 species rank fromits forrner status as a subspecies ofA.frithi (Brechlin & Kitching, 2001), With the excep-
tion of these recent findings we have worked out the life cycles and described and illustrat-ed the immature stages of all the saturniids known frem the Andaman Islands. The six
species studied by us have all been reared on plants on which the larvae feed in nature. A.andamana is the enly one among the species which is known to occur on Little Andaman inaddition to the island of South Andaman. No species have so far been found by anyone on
the Nicobar Islands, Since the neighbouring areas of SoutheastAsia harbour a rich saturni-id fauna and the Nicobars have strong biotic arnnities with these areas it is very likely thatspecies of native Saturniidae await discovery on these islands. The recent discovery of
three new species ofAntheraea is a clear indication that knowledge of the saturniid fauna isfar from complete. Attacus mcmulleni and Cricuta anclamanica are the only ones among
the six species studied by us that have six larval instars, All the other species, including A.ignescens, haye only five larval instars. The larvae of Ac'tias spp. are the largest of all theSatumiini known from these islands and are equal in size to the sixth larval instar of A.mcmulteni.
This is the first time that a plant belonging to the family Crypteroniaceae is reported as the
host plant of larval Saturniidae from anywhere in the world. It is likely that this family en-tered Asia on India when it detached from eastern Africa and drifted northward to Asia
(Conti et at., 2002), pushing up the Himalayas. However, we doubt that the moth groupsare so ancient, and believe that the current distributions of Actias in eastern Asia and
Argema in Madagascar and mainland Africa are better explained by dispersal events across
northeastern Africa and southwestern Asia. Regarding the host plant relationships,
Crypteroniaceae belongs to the order Myrtales, which contains other plant families such as
Combretaceae, fythraceae, and Myrtaceae, all containing species used as food by Actias
and Argema. The fact that the family Rhizophoraceae also belongs to the Myrtales, rein-forces the hypothesis by us based on a cocoon discovery (Prashanth Mohanraj et al,, 1996)
that Rhizophora was indeed a food used by Actias caltandra.
Acknowledgements
The first two authors thank Dr A. K. Bandyopadhyay, fomier Director, Central AgriculturalResearch Institute, Port Blair for unfiinching support to all our research endeavours. We are
very grateful to Ulrich Brosch, J. D. Holloway, W. A. Ntissig and Stefan Naumann for gra-cious assistance whenever we contacted them. We thank R XL Sreekumar, Regional Station,Botanical Survey of India, Port Blair, for identifying the plants and B. Pardhasardhi for the
photographs. For assistance in the conduct of the work we acknowledge the assistance of
Bikas Mondal, Kinnuram, and Basanth. We also thank the Chief Wildlife Warden,Andaman and Nicobar Islands, for permission to collect and study the insect fauna in the
national parks and wildlife sanctuaries of these islands.
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References
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摘 要
ア ン ダ マ ン 諸 島の Ac.’tias ignescens Moore の 生活史 と ♀ の 発見 (K . VEENAKuMARI , PRAsEIANTHMoHANRAJ and Richard S. PEIGLER )
Ae.tiαs ignescens Moore は,ア ン ダマ ン 諸島に 固有の オオ ミズ ア オ属 の 1種 で,東南 ア ジ ア の 広域種, A.
tnaenas Doubleday の 代置種と考え られ て い る、1998年 9 月、南 ア ン ダマ ン 島の Harriet山国立公 園内の
Cr}/pteronia paniculata (フ トモ モ 目クリプ テ ロ ニ ア 科)か ら得られ た 2頭 の 老齢幼虫 か ら本種 の 9 ♀ が
羽化 し,こ れ ら を 飼育ケー
ジに 人れ て,交配 させ た.また,2001年 t2月 に は,同地 で 新鮮 な 〒個体が 得ら れ,こ れ も飼育 ケ
ージ に 入れて ,採卵 した.こ れ ら2頭 の ♀か らは合計 113卵が得 られ.幼生 期に つ
い て 詳細 な観察 を行 っ た,本報で は,卵か ら繭まで の 幼生期 を図 示 ・記載 し,従来未知 で あ っ た ti’に
つ い て も図示 ・記載 した.な お ,ヤ マ マ ユ ガ科 の 食樹 として の ク リ プ テ ロ ニ ア 科は 今回 が 初 め て の 記
録で あ る が,私達 (Vecnakumarl et al,.1995)が Atta(:us ’mctnulleni Watsonの 食樹.の
・つ として 記録 した
Vitex gtabrataは Cryptet’onia panicutataの 誤 りで あ っ た.
「.文責 :岸 田 泰則1
(Accepted January l 3,2005)
Published by the Lepidopterological Socicty of 亅apan ,5−20,Motoyokoyama 2, Hachi〔’ji. Toky 〔〕,]92−OO63 Japan
N 工工一Eleotronio Library