Research Article |
Corresponding author: Chris J. Müller ( chrismuller999@gmail.com ) Academic editor: Matthias Nuss
© 2016 Chris J. Müller.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Müller CJ (2016) A stunning new species of Jamides Hübner, 1819 (Lepidoptera, Lycaenidae), with notes on sympatric congeners from the Bismarck Archipelago, Papua New Guinea. ZooKeys 571: 113-131. https://doi.org/10.3897/zookeys.571.7356
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Jamides vasilia sp. n., from montane West New Britain Province, Papua New Guinea, is described and illustrated. The new species is strongly divergent from other known Jamides Hübner, 1819 in possessing a high antenna-forewing length ratio, long androconia on the hindwing upperside and a strongly convex forewing inner margin in the male. It is compared by external structures, male genitalia and mtDNA sequence data to putative related species in the cyta group of Jamides. Notes on various Jamides taxa from the Bismarck Archipelago are also provided, with J. pseudosias (Rothschild, 1915) and J. reverdini (Fruhstorfer, 1915) recorded from New Britain for the first time.
Taxonomy, Lepidoptera , Lycaenidae , Polyommatinae , new species, Bismarck Archipelago, androconia
Jamides Hübner, 1819, butterflies, commonly known as Caeruleans, belong to the subfamily Polyommatinae. The genus is distributed throughout much of the Oriental, Australian and Pacific region tropics (
The new species introduced herein is exceptionally distinct from all other Jamides species. A number of other phenotypically distinct butterflies have recently been recorded from West New Britain Province (
Examination of type and other relevant material was carried out in various institutions (as listed below). Adult specimens were photographed using a Nikon D300s Digital SLR Camera with a Nikon AF-S VR Micro-Nikkor 105mm f/2.8G IFED Macro lens and Nikon R1C1 Close-up Kit Flashes Speedlights. RAW images were edited using Adobe Photoshop CS6. Editing included alignment, auto contrasting and removal of background. A standardised procedure was followed with photography and image editing to ensure consistency of image output. Genitalia were extracted following maceration of abdomens in 10% KOH at room temperature for 36 hours. Genitalia were photographed in glycerol using the fore-mentioned camera body adapted to a Meiji Techno EMZ-5TR-P-FOI Trinocular Stereozoom Microscope, with OPTEK FL95E Fibreoptic Illuminator and twin arm optical fibre. Individual images were taken with the remote acquisition software DIYPhotoBits Camera Control 5.2. Sliced genitalia photographs were stacked and concatenated using the software Helicon Focus 6.0 and edited in Adobe Photoshop CS6. The plates were assembled in Adobe InDesign CS6 and the phylogeny tree in CorelDRAW X6. Genitalia were stored in small glycerol-filled vials pinned beneath the specimen.
Descriptions of external facies follow that of the numerical vein system of
Tissue material (two legs) was collected from representatives of all of the eight Jamides subgroups of
Individual sequence properties were assessed using MEGA, version 4.1 (
Sequences were uploaded to GenBank (http://www.ncbi.nlm.nih.gov/genbank/) and are listed for each individual in the phylogenetic tree presented in Fig.
CJMC
MNHU
NHM
SMT Staatliches Museum für Tierkunde, Dresden, Germany
Holotype ♂ (Figs
Jamides adults (left side upperside and right side underside, where halved) and label data. 1 Jamides vasilia holotype ♂ upperside 2 J. vasilia holotype ♂ underside 3 J. vasilia holotype ♂ label data 4 J. vasilia paratype ♂ (halved) 5 J. vasilia paratype ♂ (halved) (note rubbed hindwing) 6 J. vasilia paratype ♀ (halved) (note rubbed forewing) 7 J. vasilia paratype ♀ upperside 8 J. vasilia paratype ♀ underside 9 J. vasilia paratype ♀ (halved) 10 J. pseudosias holotype ♂ upperside 11 J. pseudosias holotype ♂ underside 12 J. pseudosias holotype ♂ label data 13 J. pseudosias ♂ (halved) (New Ireland) 14 J. pseudosias ♀ (halved) (New Ireland) 15 J. pseudosias ♀ (halved) (New Britain). Scale bar = 10 mm.
Both sexes of Jamides vasilia are highly distinctive and cannot be confused with any other known species. The uppersides of both sexes are brighter and more reflective than those of other species in the genus, the ground colour of the male in particular rivalling Morpho butterflies in intensity and radiance. The male of J. vasilia is unusual from a structural perspective, having long antenna (versus forewing length) that extend well beyond the cell and reach the postmedian area. The antenna/forewing length ratio is ~0.6, whereas in all other known Jamides species the antenna of the male is approximately half the length of the forewing. The shape of the male forewing is unique in that the tornus and inner margin are rounded, the latter being convex while in all other Jamides species the inner margin is straight. Also peculiar in the male is the large purple-brown patch occupying the costal one-third of the hindwing upperside. This patch is adorned with long androconial hairs (up to 5 mm in length). No other known Jamides bears such prominent structures. The male forewing upperside bears a thick terminal black border, tapering towards the tornus, not present in other species. On the underside, both sexes are easily recognised by the curved striae that are well displaced between veins on both wings and the two apical black spots on the hindwing are more rounded and pronounced than in other taxa. The area of orange bordered by metallic blue on the hindwing underside is extensive in both sexes, particularly the female, and extends from the inner margin all the way to space 5. The male forewing underside has the entire median area grey-white between the inner margin and vein 2.
The male genitalia of J. vasilia is unusual, bearing long teeth-like processes on the inner margin of the valva, with a spine-like process at the apex of the valva.
♂ Forewing length 15.8mm (Holotype), Antenna length 9.5mm (Holotype). Head grey; antenna long and extending well beyond end of cell, black ringed with white; thorax blue scaled on upperside, grey beneath; abdomen dark grey with blue scales near base on upperside; legs black with white between segments.
Forewing with tornus rounded (in congeners squared), and inner margin convex (in congeners straight).
Forewing upperside brilliant metallic sky blue, darkening slightly towards termen and apex, a prominent black termen border widening to ~1.5mm at apex; cilia black. Forewing underside deep grey-brown; inner margin broadly pale grey-white (in spaces 1a and 1b), with some metallic sky blue scales along vein 1b; termen narrowly white; a narrow grey-brown subterminal band, narrowly edged with white that forms triangular marks on basal edge; a postmedian band of similar colour to ground colour, about 2mm wide, narrowly edged with curved white on outside margin and with corresponding dark brown on inside margin, band is strongly displaced at veins by approximately 1mm, towards base in spaces 6 and 7 and incrementally towards base towards inner margin in spaces 2, 3 and 4; a median band at end of cell, approximately 1.5mm wide, curved and edged with white and dark brown as in postmedian band; cilia dark brown.
Hindwing rounded, with 2.5mm long, black tail at vein 2, tipped with white.
Hindwing upperside brilliant metallic sky blue, darkening slightly towards termen; a large purple-brown patch, clothed with long androconia along costa and in median area, occupying much of spaces 6 and 7 as well as part of cell and space 5; termen narrowly black; a series of diffuse black-dusted and indented subterminal spots, as well as two small black spots at tornus near intersection of vein 1b. Hindwing underside deep grey-brown; inner margin narrowly white, interspersed with black at vein ends; a large black apical spot (1.5mm diameter) in space 6 and another apical spot of similar colour and dimensions in space 7, both spots rimmed narrowly with white; two large subtornal black spots in spaces 2 (approx. 1mm across) and 3 (approx. 0.4mm across) and smaller, less regular black spots in spaces 1b and 4, each of these spots broadly edged along veins with metallic sky blue and basally with bright orange, and then fine white arcuate lines; an additional arcuate white subterminal line in space 5 linking the subapical and subtornal black spots; a postmedian band similar to that on the forewing underside, curved and strongly displaced at veins; a median band edged with white and dark brown at end of cell; a basal band approximately 1mm wide, edged narrowly with white, displaced at either side of cell; cilia dark brown.
♀: Forewing length 20.6mm, antenna length 10.8mm, antenna, thorax, abdomen and legs similar to male.
Forewing with inner margin straight.
Forewing upperside bright lustrous sky blue, darkening towards termen; apex of termen and apex broadly and sharply edged black (2.5mm and 8mm wide, respectively). Forewing underside similar to male, ground colour slightly paler and inner margin only narrowly white-grey and without blue scales, postmedian band extending to vein 1b, white edging to bands slightly wider and more diffuse.
Hindwing rounded, with black, white-tipped tail at vein 2 (approx. 4mm long).
Hindwing upperside bright lustrous sky blue, darkening towards termen; inner margin broadly black, basally transitional to brown, with narrow blue scaling along edge of space 6 and 7 adjacent to cell; a row of black subterminal spots (each averaging 0.6mm diameter) in spaces 2, 3, 4 and 5; two irregular black subtornal lines in spaces 1a and 1b, subparallel to termen; inner margin narrowly white. Hindwing underside similar to male, ground colour slightly paler, orange bordering subterminal spots wider and very extensive, reaching from the inner margin to space 5.
Male genitalia. Vinculum and tegumen ring broadly oval; sociuncus divergent; socii with lateral margin rounded, socii distinctly separated by straight, perpendicular sinus; saccus of even thickness, brachium widely bowed dorsally, yet sharply bent laterally, tapered dorsally; valva elaborate, hollowed, with serrated margin and teeth-like processes on inner margin, long narrow process stemming from near base of valva, weakly clubbed with pointed ventral surface on club; phallus with prezonal section approximately one tenth the length of postzonal section, slender, with apical rounded pencil-like process.
This enigmatic and exquisite new butterfly is named in honour of the author’s wife, Vasilia (Vicki) Savvas (Muller). Vicki has always supported the author’s obsession in butterfly research, despite the many sacrifices both on and off the field.
New Britain Island, Papua New Guinea.
Adults of J. vasilia inhabit moss forest and appear to have a more rapid, erratic flight than other members of the genus. Two females were initially observed flying around the base of a Syzygium R.Br. ex Gaertn. (Myrtaceae) sapling and resembled those of the lycaenid Arhopala thamyras (Linnaeus, 1758). The particular Syzygium plant had numerous, highly active, medium-sized brown ants present on the lichen-covered trunk but no early stages of Jamides could be located either on the foliage, trunk or in leaf litter surrounding the base of the plant. In the upper parts of the Whiteman Range (Figs
The phylogeny of Jamides, presented in Fig.
There is no significant variation in the type series of J. vasilia, with all specimens similar in size and shape. The male exhibits very slight variation in the width of the terminal black border.
The extent of distribution of J. vasilia on New Britain Island is not known. Based on the distribution of other endemic butterfly taxa on the island, it is unlikely to be restricted to the Whiteman Range, although no specimens have been observed during surveying, at a range of altitudes, of the Nakanai and Bainings Mountains, in central and east New Britain, respectively. J. vasilia appears to be a rare species.
Catochrysops cyta:
The type (?types) of Jamides cyta (Boisduval, 1832) were taken in New Ireland, during the voyage of the Astrolabe through the Indo-Pacific during the period 1826–1829. The Astrolabe, captained by Dumont d’Urville, visited at least three coastal sites in New Ireland; Port Praslin, Hèvre Cartret (Carteret Bay) and Likiliki in the Bay of Frondeurs (=Slinger’s Bay) (
“Ailes d'un bleu-argenté luisant; les inférieures avec une petite queue; dessous des quatre avec plusieurs raies blanches interrompues; les inférieures ayant en outré une rangée marginale de taches noires, don't les trois voisines du bord abdominal marquées de fauve et de vert doré.
Il a le port et la taille d’Elpis, auquel il resemble beaucoup.
Nouvelle-Irlande.”
Translated, this states that the insect (presumably a male), has “Wings a shiny silvery blue; the hindwings with a small tail; underside of the four [wings] with several broken white stripes; the hindwings additionally having a marginal band of black spots, of which the three adjacent [closest] to the abdominal [inner] margin are marked with fawn and golden green. It has the appearance and size of Elpis [Jamides elpis (Godart,[1824])], which it closely resembles. New Ireland.”
The description above is pertinent only to the male of J. cyta. The author has surveyed several sites throughout New Ireland, close to the localities visited by the Astrolabe, and found J. cyta to be particularly common at all of the lowland sites.
Jamides cyta adults (left side upperside and right side underside, where halved) and label data. 16Lampides amphissina var. malaguna lectotype ♂ upperside 17 As Fig.
Nominate cyta from the Bismarck Archipelago is highly distinct from the New Guinea mainland subspecies amphissina Grose Smith, 1894 (not to be confused with amphissa from Maluku), in that the Bismarck specimens are larger and brighter, with much more pronounced orange on the hindwing underside. Bismarck cyta taxa may be further separated in that females from New Britain possess more extensive and brighter blue on the upperside than those from New Ireland and may warrant subspecific status. The types of J. cyta amphissina are illustrated in Figs
J. cyta is an easily recognised species, the male bearing a pale blue-white upperside and both sexes bear a row of triangular subterminal black spots on the hindwing underside. Note that the illustration of the male J. ‘cyta’ (as Jamides ‘cytus’) in
Lampides pseudosias:
In the Bismarcks J. pseudosias was known only from one pair taken at an unspecified locality in New Ireland by A. F. Eichhorn in November 1923, which are held in the NHM (Figs
J. pseudosias is readily distinguished from its congeners by its long hindwing tail, which is at least 1.5 times that of related species in New Guinea and by the rich blue colour of the male upperside. The forewings of the male are narrower and the apex more rounded than in related species. The forewing underside median bar and postmedian band are continuous across vein 3, though slightly oblique, giving the band a curved appearance. Note that
Lampides elpis reverdini:
Jamides reverdini was previously known from the Bismarcks from just a single specimen from New Ireland, which
This insect is unique in its comparatively large size, bright silvery blue upperside and boldly patterned underside. On the underside, the ground colour is a distinct deep grey and, like J. pseudosias, the forewing median bar and postmedian band are connected at vein 3. J. reverdini is easily separated from J. pseudosias by, among other fore-mentioned features, the shape of the subterminal band on the hindwing underside. In J. reverdini, this band constitutes a series of rectangular markings bordered heavily with white, whereas in J. pseudosias these markings are distinctly triangular.
Jamides vasilia exhibits significant disparity from other members of Jamides. The male wing shape, with rounded tornus and convex inner margin, is unique, as is the black, tapering terminal border. On the forewing underside of the male the inner margin is broadly white-grey, relative to the remainder of the wing, a feature apparently unique in the genus. Also in the male, the hindwing upperside bears conspicuous androconia (Fig.
The male genitalia of J. vasilia (Fig.
Jamides adults (left side upperside and right side underside, where halved) and label data. 31 Jamides areas holotype ♂ upperside 32 J. areas holotype ♂ underside 33 J. areas holotype ♂ label data 34 J. areas allotype ♀ upperside 35 J. areas allotype ♀ underside 36 J. areas allotype ♀ label data 37 J. areas ♀ (halved) (Guadalcanal) 38 J. nitens holotype ♂ upperside 39 J. nitens holotype ♂ underside 40 J. nitens holotype ♂ label data 41 J. nitens ♂ (halved) (Telefomin) 42 J. nitens ♂ (halved) (Mianmin Range) 43 J. nitens ♀ (halved) (Telefomin) 44 J. reverdini ♂ (halved) (New Britain) 45 J. reverdini ♀ (halved) (New Britain). Scale bar = 10 mm.
Jamides male genitalia. (a genitalia in dorsal view with aedeagus removed, b genitalia in ventral view with aedeagus removed, c genitalia in lateral view with aedeagus removed, d aedeagus in dorsal view, e aedeagus in lateral view, f valva in lateral external view, g valva in lateral interior view, h valva in dorsal view, i valva in ventral view. 46 Jamides vasilia holotype ♂ 47 J. cyta cyta (New Britain) 48 J. cyta amphissina (Sepik, mainland NG). Scale bar = approx. 1 mm.
Jamides male genitalia. (a genitalia in dorsal view with aedeagus removed, b genitalia in ventral view with aedeagus removed, c genitalia in lateral view with aedeagus removed, d aedeagus in dorsal view, e aedeagus in lateral view. 49 J. pseudosias (Mianmin Range) 50 J. areas (New Georgia, Solomons) 51 J. nitens (Mianmin Range) 52 J. reverdini (New Britain). Scale bar = approx. 1 mm.
Jamides male hindwing apex, showing development (or absence) of androconia. 53 Jamides vasilia paratype ♂ 54 J. pseudosias (Mianmin Range) 55 J. cyta cyta (New Britain) 56 J. areas (New Georgia, Solomons) 57 J. nitens (Mianmin Range) 58 J. reverdini (New Britain). Scale bar = approx. 1 mm.
That J. vasilia was phylogenetically recovered, as a result of Bayesian Inference, within a clade of Jamides also comprising J. cyta and J. nitens is perhaps unexpected, considering that J. vasilia is phenotypically very different from either of these taxa. Indeed, there is significant genetic divergence (in COI barcode) between J. vasilia and J. cyta or J. nitens. However, a potential similarity between J. vasilia and J. cyta is that the latter has been recorded feeding on Syzigium in Australia (
J. vasilia is a montane species, having been recorded from 850 to 1050m. An additional female was observed at 700m in the Whiteman Range. Of the related Jamides taxa, J. nitens is also a montane insect,
Various phylogenetic analyses produced a range of trees for the Jamides dataset. Indeed, in several, including Neighbour-Joining, Maximum Likelihood and Maximum Parsimony, the resultant tree suggested that J. vasilia is sister to all assessed Jamides species groups and that it may represent a separate genus. Given the variable results from the different methods, COI alone is clearly not an adequate tool and additional sequencing of nuclear gene fragments would undoubtedly assist in better resolving the phylogenetic position of the new taxon.
Fifty percent majority rule consensus phylogram for selected Jamides, from a Bayesian analysis of 650 base pairs of COI gene fragment. Numbers at the nodes are the posterior probabilities of those nodes. The scale bar represents 2% genetic distance. Each terminal indicates the taxon name, locality and Genbank sequence accession number. Note the following abbreviations in parentheses for Province (Prov.), Papua New Guinea (PNG) and Australia (Aus).
A number of friends and colleagues assisted the author, who is indebted to all of them. John Tennent photographed relevant specimens in institutional collections and facilitated several of the author’s trips to the UK. Geoff Martin and Blanca Huertas (NHM) allowed the study of material in their care. Ullasa Kodandaramaiah, Kerala (India), sequenced certain material. Yusuke Takanami, Japan, provided photos of lectotypes held in MNHU and SMT. Jean Weiner,