Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision Author C, Lees David text Zootaxa 2016 4118 1 1 97 journal article 38930 10.11646/zootaxa.4118.1.1 a4e408c4-2575-4b5c-878b-e1b9010ec619 1175-5326 264597 CFA586DA-10EE-468B-80EE-35351E3845FD Heteropsis barbarae Lees & Kremen , sp. nov. LSID: urn:lsid:zoobank.org:act:DC87986A-3822-43C9-B1B9-83A2714D088B Prior references : sp. 16 ( Lees 1997 ; Torres et al., 2001 : 462). Type material., Deposition MNHN : Holotype : ♂ ( Fig. 20 A), Madagascar NE, Maitsoarongana, Sikiory River, near camp, 15.1885o S , 49.5163o E +/- 0.02 km , 685 +/- 23 m , 12/12/2001 : 15:29, D.C. Lees: DL 01-227, BMNH (E) 2008-69 [accession number], BMNH (E) 1717096 [ QTR barcode]. Paratypes (accession BMNH (E) 2008-69 ) : ♂, Madagascar NE, Maitsoarongana, Sikiory R., 660–750 m , 15.1907o S , 49.5309o E +/- 2 km , 750 +/- 10 m 12/12/2001 , D.C. Lees: DL 01-241, BMNH (E) 1717097 [ QTR barcode]; ♂, NE, Maitsoarongana, c. 710 m , 15.1885o S , 49.5163o E +/- 0.02 km , 710 +/- 23m , 13/12/2001 : 06:00, D.C. Lees: DL 01-247, BMNH (E) 1717098 [ QTR barcode]; ♂, NE, Maitsoarongana, ridge just S. of camp, 15.1907o S , 49.5309o E +/- 2 km , 750 +/- 10 m 7/12/2001 : 16:39, D.C. Lees: DL 01-132, BMNH (E) 1717099 [ QTR barcode]; ♀ ( Fig. 20 B), Madagascar NE, Sikiory R., riparian, 15.1848o S , 49.4659o E , +/- 5.42 km , 710 +/- 5 m , 12/ 12/2001 : 17:12, D.C. Lees: DL 01-231, BMNH (E) 1717100 [ QTR barcode]; ♂, Madagascar NE, Maitsoarongana, near path, 15.1907 o S , 49.5309o E , +/- 2 km ., 750 +/- 100 m ., 13/12/2001 , D.C. Lees: DL 01-248, BMNH (E) 1717095 [ QTR barcode]. Deposition summary : BMNH (HT ♂, 4 PT ♂♂, 1 PT ♀). Type locality. Madagascar NE, Maitsoarongana, Sikiory River, 15.1885o S , 49.5163o E +/- 0.02 km , 685 +/- 23 m . Diagnosis. The ventral wings of Ht. barbarae are similar to some other members of the Ht. strigula group that exhibit a crenulate HW margin, notably Ht. menamenoides sp. nov. and Ht. strigula . Upperside dark brown, lacking light orange wash and smooth HW margin of Ht. menamenoides ( Fig. 21 A) and Ht. maeva ( Fig. 21 D) and the HWD orange crescent (Mf) typical of Ht. strigula . Fresh specimens of Ht. barbarae are distinctive in their strong slightly dingy orange ventral cast ( Fig. 20 A,E), and the ventral scales are easily displaced/rubbed. Worn ♀♀ can easily be confused with those of often sympatric Ht. pauper and Ht. undulans . Indeed the ♀ of Ht. undulans is sometimes almost indistinguishable ( Fig. 20 D) to that of Ht. barbarae ( Fig. 20 B), but Ht. undulans ♀♀ often tend to be more flushed with reddish scales on the upperside, while Ht. barbarae can be distinguished by the combination of rich and dense slightly dingy orange scales on the ventral surface which lack a yellow cast (such as in Ht. strigula ), combined with a less contrasting, pale orange band between Pml and Sml ( Fig. 20 C,D,F), which is pale ochreous in both sexes of some populations of Ht. undulans (however that is a very variable species), but there will always be a few especially more worn specimens of Ht. barbarae that are hard to place without dissection. The similarly lowland Ht. maeva has a brighter orange underside ( Fig. 21 D), denser black ventral brush in the ♂, and is distinctive in that its upperside is orange too, except for HWs of western specimens which are usually suffused with brown (but Ht. barbarae is anyway absent in the west of Madagascar ). Description. Wings : Upperside mid-brown with a slight russet brown cast. Space-CuA1 ocellus expressed strongly in FWD; orange ring sub-hexagonal., M1 ocellus very small black white pupilled spot surrounded by a narrow orange ring. HWD ocellus CuA1 fairly small and sub-elliptic, surrounded by a narrow orange ring. Underside has a lighter dingy orange cast, only slightly lighter distad of the Mb. Mb darker orange-brown, gently convex and slightly irregular (relatively straight in the HT HWV) with paler orange highlights distal to it in space- M2 to CuA1. FWV space-CuA1 ocellus similar size to FWD (in the HT), as is FWV M1 ocellus, which is reduced to a very small white point surrounded by a narrow black ring and orange ring that slightly contrasts with background. FWV space-CuA1 ocellus follows concave shape of the FWV darker orange-brown Mb before it bends back toward mid costa, with a ring that is orange near the black iris (which is significantly smaller than on FWD) and is yellowy-orange towards the greatest inflection of the bend in the Mb, forming a ‘spiral arm’ (Ore). HWV space-CuA1 ocellus similar size and shape to that on dorsal surface, surrounded by pale orange ring not much lighter than background; no other ocelli conspicuously expressed on the HWV nor HWD. The HWV basal area is sparsely irrorated with brown strigulae, the area between PMb and Mb more evenly and finely irrorated. A wavy-zigzag darker diffuse brownish line inset from margin follows a double narrow Sml divided by the ground colour, not very ochreous as usual in Ht. undulans ) that closely tracks the margin of both wings, the margin being relatively smooth in the FW and distinctly crenulate in the HW. The reddish/dark orange-brown HWV PMb is slightly more convex than the Mb and both lines terminate abruptly at 1A. In FWV in the cell area are four dark orange-brown Cbs meeting almost perpendicular at the costa, the outermost pair delineating relatively paler areas, the outermost of which is more convex than the others, with its borders diverging towards the costa (in the HT). Variation. ♀♀ similar but larger and slightly paler ( Fig. 20 B). Space-CuA1 ocellus FWD is a little wider proximad in some specimens, and FWD M1 ocellus sometimes not visible. HW space-CuA1 ocellus is more circular in some specimens. FIGURE 20. Adults in DV (left), VV (right) for mounted specimens and adults in natural posture. A : HT ♂ of Heteropsis barbarae Lees & Kremen , sp. nov. (Maitsoarongana, Makira; BMNH(E) 1717096). B : PT ♀ of Ht. barbarae (Sikiory R., Makira; BMNH(E) 1717100). C : LT ♂ of Culapa undulans Oberthür, 1916 (Fito; BMNH(E) #674752). D : ♀ of Ht. undulans (Antanandava, PN Andohahela). E : ♀ of Ht. barbarae (Makira; 27/01/2003). F : ♀ of Ht. undulans (Ankirindro, Makira; 17/01/ 2003) (photos: D. C. Lees). Wingspan/fwl : range 33.5–38.5/ 17.8–20.5 mm (n=14 ♂♂), including HT ♂, 36.3/ 20.7 mm ; mean 35.7 +/- 1.1 SD/19.2 +/- 0.97 SD mm (n=10 ♂♂, including HT). Range 36.7–40.4/ 20.1–21.5 mm ; mean 37.0+/- 2.6 SD/20+/- 1.5 SD mm (n=4 ♀♀; referred specimens). Androconia : (observations on ♂ specimen MAD182 from Marojejy, referred specimen). Discocellular brush HWD dark grey at base, rufous/reddish-brown in distal ½; underlying patch light brown to reddish-brown, long, very narrow lenticular, surrounded by silvery scales or yellowish ones distal to brush. HWD Cub (cubital brush) above 1A+2A base, mid brown, dispersed, juxtaposed potentially under abdominal sternal segments A2–A5, approximately where there is a conspicuous black androconial patch, which is not divided by lighter overlying scales as it is in Ht. undulans . Inflated vein 2A swollen within basal half. Palps : Ochreous, with intermingled brown scales on outside face away from compound eye, dark brown scales above where potentially brushing eye. Etymology. After Barbara, the mother of Claire Kremen. Discussion. This species was first recognized in the field in 1991–1993 by Claire Kremen in the survey of what is now Masoala National Park ( Lees 1997: 64 ). It had been collected once before on the expedition of M.I. Evans to RS Ambatovaky in 1988 (specimen in BMNH) but surprisingly there were no (other) historical specimens in BMNH nor MNHN. It is a quite striking and common lowland riparian species in large parts of the Northeast of Madagascar , so obviously completely overlooked in historic collecting in the region. All available types in the Ht. subsimilis and Ht. strigula groups (all in BMNH) were examined. In particular, among the ‘orange underside’ species, the following specimens are typified. These include two of up to three (although in his original description, Mabille does not state the number of specimens; but see also Mabille [1887]: 77) possible ♂♂ STs in BMNH of Mycalesis maeva Mabille, 1878 ; LT ♂, here designated ( Fig. 21 D): BMNH(E) 673788; Lectotype |MadagascarNo10 [?sic; or ‘MadagascarNW’?]|Ex Coll. CHRIS WARD|Ex Oberthür Coll. Brit. Mus. 1927-3.| Culapa [sic] maeva Mabille |P.E.L. Viette de. 1968 Mycalesis maeva ♂ LECTOTYPE . For Mycalesis maeva , the potential ST male BMNH(E) 673789, “NW Madgsr” from Joicey Bequest (presumably from the Grose- Smith collection) is here designated PLT. Regarding Culapa laetifica Oberthür, 1916 , a LT ♂ is here designated ( Fig. 21 B): the specimen bearing labels “BMNH(E) #674741; Lectotype |Nord-Madagascar (Antakares) Isokitra a Diego Suarez Mai a Octobre 1891 E.&B. Perrot; illustrated by Oberthür (1916: Pl. 368, f. 3072) ”. For the last species, the text by d’Abrera (1980 : 185) “♀ (? holotype ) as illustrated”, is here considered ambiguous, but the corresponding PLT specimen (also illustrated by Oberthür (1916: Pl. 368, f. 3073)) is now numbered BMNH(E) #674742, while four additional PLT ♂♂ of Culapa laetifica are numbered BMNH(E) #674743-674746. Among STs of Culapa laeta Oberthür, 1916 , a LT ♂ in BMNH is here designated ( Fig. 21 C), the specimen bearing labels “LT|Nord-Madagascar (Antakares) Isokitra à Diego-Suarez Mai à Octobre 1891 E.&B. Perrot| Culapa laeta Obthr ♂ type |P.E.L. Viette det. 1968 Culapa laeta Ch . Obthr ♂ LECTOTYPE |Supposed “ Type ” ( syntype ) of Culapa laeta Oberthür )|[Copy of f. 3075]|Ex Oberthür Coll. Brit. Mus. 1927-3.| Lectotype ♂ of Culapa laeta Oberthür D.C. Lees , det. Sept. 2000 |BMNH(E) 675416”; a surviving further 5 of 7 ♂ STs and all 4 ♀ PLTs of Culapa laeta become PLTs. The STs of Culapa undulans Oberthür, 1916 were also examined; the LT ♂ is here considered to have designated by d’Abrera, 1980 : 184 by the text “♂ ( holotype ) and ♀ as illustrated”, as illustrated on p. 185; the LT ( Fig. 20 C) bears the labels “ Madagascar Fito Mai a Aout 1897 Perrot Freres|BMNH(E) #674752| Culapa undulans ♂ Type Oberthür [handwriting]|[label bearing copy of f. 3062]” and is indeed the specimen previously illustrated by Oberthür (1916: Pl.368, f. 3062) ; the corresponding PLT ♂ of Culapa undulans from the same locality is numbered BMNN(E) #674753. The specimen illustrated with no abdomen by d’Abrera (1980 : 185; BMNH(E) #674891) represents a PLT of Culapa undulans as treated syntypically by Oberthür (1916: 220, pl. 367, f. 3063, from ‘Antsianaka’) but is misidentified and in fact represents a ♀ of Ht. pauper , to which “ Culapa (var. ou espèce séparée) pseudonarcissus ” Oberthür (1916: 223, fasc. 11: 41) (that d’Abrera, 1980 : 184 treated under Ht. undulans ) also would belong. The last form (represented by a ♀ without abdomen; Fig. 14 E) is here designated LT of Culapa pseudonarcissus , bearing labels “Nord Madagascar Antakares Isokitra a Diego Suarez Mai a Octobre 1891 | Culapa pauper var? pseudonarcissus (Bdv. in ms) Obthr. Type ♀)|BMNH(E) #674890” and this is presumably a dry season individual; although this has no consequence, Oberthür (1916, pl. 368, f. 3068) also captioned it “ Culapa pauper - pseudonarcissus ”. Additional information. ♂ genitalia : 141DL ( Fig. 18 C, referred specimen): the basic configuration is fairly typical for Ht. strigula group. From LV, the approximately parallel-sided uncus is very slightly longer than the tegumen, the dorsal margin approximately in line with that of the tegumen and slightly upraised after its brow, with a smoothly round ‘head’ before the ventral hook (inflated from DV before tip), which projects forward rather than down. The gnathos originates on an elliptic base and is gently tapered and straight to an approx. 35-degree angle to uncus in the specimen examined; from DV it features a typical ‘bull’s horn’ sinuate appearance with pointed tips inrecurved. Only a slight constriction is found at hinge with vinculum. The valve has a fairly parallel-sided, flattened arm which is bent over at tip (inwards from DV) exposing a dense coverage of spinoid setae and the uncus tip when not downflexed is about the same extension as, not proud of the valve tips. The saccus is of normal length for the Ht. strigula group and the aedeagus is about the same length as the valve and strongly recurved distad of the (also proximally uprecurved) ostium. The juxta is narrow and not sharply lipped proximad. DNA divergences: COI-5P cluster number BOLD:ACW4995 (exemplar BMAD248-15, DL14M-0033, Marojejy) is 4.86% divergent to Ht. tianae (BOLD:AAE4112), and 5.63% divergent to Ht. maeva (BOLD:AAE5488), its sister species in the study of Aduse-Poku et al., (2015). In the COII dataset of Torres et al., (2001) , Ht. barbarae (their “ Hen . sp. 16”; AY 040128 based on 1 ♀ from Ivontaka Sud) was 6.54% divergent to Ht. maeva ( AY 040132 based on 3 ♂♂, and 1 ♀ from Masoala; 390 bp comparable). Phylogeny/sister species : possibly Ht. menamenoides (combined tree in Aduse-Poku et al. , 2016 , in press). Torres et al., (2001 : 467) suggested a topological relationship with Ht. maeva in their cladistic analysis of COII sequences, but this was not supported ( Torres et al., (2001 : 466). Ecology and distribution. Habitat : riparian primary rainforest. Behaviour : flies low and both sexes come easily to fruit bait. Hostplant : unknown but almost certainly low grasses. Early stages : unknown but for expressed eggs, which were whitish. Distribution : endemic to the northeastern rainforests of Madagascar , from Marojejy and Masoala south to Ambatovaky ( Fig. 30 C, light green dots). Elevational range : 0– 890 m . (n= 153, including referred specimens and observations). Referred specimens. ♀, Madagascar NE, Marojejy PN, Camp II [above], 14.4344o S , 49.759o E +/- 0.25 km , 850 +/- 50 m , 12/11/2006 : 10:00, D.C. Lees: DL 06-298; ♂, NE, Marojejy PN, 14.4344o S , 49.7606o E +/- 0.5 km , 730 +/- 50 m , 11/11/2006 : 15:48, D.C. Lees: DL 06-269, DSC _0826 [photo]; ♂, data as above but: 3/12/2006 , D.C. Lees: DSC _0826 [photo]; ♀, NE, Manantenina camp 2, Marojejy PN, 14.4347o S , 49.7601o E +/- 0.5 km , 730 +/- 50 m , 10/11/2006 , D.C. Lees: DL 06-169; specimen, NE, Marojejy PN [between camp 1 and camp2], 14.4362o S , 49.7679o E +/- 0.85 km , 613 +/- 125 m , 10/11/2006 , 15:58, D.C. Lees: DL 06-170; ♀, NE, Marojejy PN, 14.4377o S , 49.7756o E +/- 0.25 km , 400 +/- 50 m , 20/11/2006 : 15:07, D.C. Lees: DL 06-456; ♂, NE, Marojejy PN, 14.4425o S , 49.7818o E +/- 0.5 km , 450 +/- 150 m , 10/11/2006 : 10:11, D.C. Lees: DL 06-205; ♂, NE, Marojejy PN, 1 km before [after?] entrée, 14.45015o S , 49.786o E +/- 1.78 km , 326 +/- 175 m , 10/11/2006 : 13:16, D.C. Lees: DL 06-204; ♂, NE, Marojejy PN [around entrée], 14.4626o S , 49.7964o E +/- 0.5 km , 156 +/- 100 m , 10/11/2006 : 12:48, D.C. Lees: DL 06-168; ♂, NE, Marojejy, 650 m , 30/1/2014 : 12:57, D.C. Lees: MAD 182 [pheromone voucher], IA387 [isotope voucher]; ♂ [ BMNH ], NE, Andranobe, Masoala Peninsula, 15.6816o S , 49.9573oE +/- 1.31 km , 0 m, 3-Dec-91 , C. Kremen: CK826, BMNH (E) 2008-69 ; ♂, NE, Masoala, Anaovandrano, 14/1/1994 , D.C. Lees: WG5 [wing image], IA38 [isotope voucher]; ♂, NE, Masoala, Ambavaony, 125 m , 27/11/1994 , D.C. Lees: DCL 47a, IA108 [isotope voucher]; ♂, NE, Masoala, Ambanivony, 25/11/1993 , C. Kremen: CK26, IA109 [pheromone voucher]; ♀, NE, Makira, Vohitaly, 15.4414o S , 49.5309o E +/- 0.15 km , 645 m , 30/12/2002 , D.C. Lees: DL-SF00, IA112 [isotope voucher]; ♀, NE, Makira, Ankirindro, 15.2931o S , 49.5472o E +/- 0.15 km , 675 +/- 25 m , 15/1/2003 , D.C. Lees: DL- SE00, IA114 [isotope voucher]; specimen, NE, Ankiatomboka forest, c. 710 m , 15.179o S , 49.412o E , 710 +/- 50 m , 9/12/2001 , D.C. Lees: DL 01-169; ♂, NE, Bivontro, Makira, 15.3984o S , 49.4477o E +/- 0.1 km , 762 +/- 25 m , 21/12/ 2002 , D.C. Lees: DL-SG77; ♂, NE, Vohitaly-Anjiahely, 15.3989o S , 49.5211o E +/- 1.82 km , 435 +/- 50 m , 31/12/ 2002 , BMNH (E) #672331 [ DNA voucher]; ♀, NE, Anjiahely, 0.37 km E., 15.4120o S , 49.5022o E +/- 0.25 km , 380 +/- 5 m 12/12/2002 , D.C. Lees: DL-SG1, BMNH (E) 2008-69 , BMNH (E) 1717101; ♂, NE, Sahantaha, near ‘fanihy’ roost area, 15.1963o S , 49.5564o E +/- 0.98 km , 517 +/-54 m , 13/12/2001 : 10:46, D.C. Lees: DL 01-257; ♂, NE, Anjanaharibe Mt., 15.188o S , 49.6139o E +/- 0.1 km , 475 +/- 5 m , 6/2/2003 , D.C. Lees; ♀, NE, Tampolo, W. Masoala, ca. 150 m , primary rainforest, 15.7301o S , 49.9604o E +/- 2 km , 6/11/2001 , D.C. Lees: DL 01-1; ♀, NE, Tampolo, W. Masoala, course of old railroad 1–2 km SE of camp, 15.7279o S , 49.9751o E +/- 1.8 km , 32 +/- 25 m , 9/ 11/2001 , D.C. Lees, DL 01-27, BMNH (E) #697188 [ DNA voucher], KA556 [=KA-P556; DNA extract voucher]; ♀, NE, Andranobe, W. Masoala, 240 m , 15.4o S , 49.58o E +/- 1.78 km , 240 m . 3/2/1991 , C. Kremen et al., Genitalia157 [genitalia voucher]; ♀, NE, Andranobe, W. Masoala, 90 m , 15.6816o S , 49.9573o E +/- 1.79 km , 90 m , 14/11/1991 , C. Kremen et al.: Biodiversity voucher; ♀, NE, W. Masoala, Andranobe field station, T2S2 [site], 90 m , 15.6816o S , 49.9573o E +/- 1.79 km , 90 m , 14/11/1991 , C. Kremen et al. , Biodiversity survey voucher|Gen158 [genitalia], IA594 [isotope voucher]; ♂, data as above but: 2/11/1991 , C. Kremen et al., Biodiversity voucher; ♂, NE, Andranobe, W. Masoala, 520 m , 15.65o S , 49.983o E +/- 1.79 km , 520 m , 14/11/1991 , C. Kremen et al., Biodiversity voucher; ♂, data as above but: 599 m , 20/10/1991 , C. Kremen et al. , Biodiversity voucher; ♂, data as above but: 8/11/1991 , C. Kremen et al. : Biodiversity voucher; ♀, data as above but: 100 m , 26/10/1991 , C. Kremen et al. : Bio|Gen156 [genitalia]; ♀, data as above but: 29/10/1991 , C. Kremen et al. : Bio|Gen159; ♀, NE, Andranobe, W. Masoala, 520 m , 15.65o S , 49.983o E +/- 1.79 km , 520 m , 8/11/1991 , C. Kremen et al. : Biodiversity voucher; ♂, NE, Be Dinta, upper Anaovandrano R., W. Masoala, 500 m , 15.6733o S , 49.9885o E +/- 0.15 km , 500 +/- 50 m , 18/2/ 1993 , D.C. Lees: DL 93-0012, IA593 [isotope voucher]; ♂, NE, Masoala E, Be Dinta, 550 m , 17/02/1993 , D.C. Lees: DL 93-0029, IA592 [isotope voucher]; ♂, NE, Be Dinta, upper Anaovandrano R., W. Masoala, 500 m , 15.6733o S , 49.9885o E +/- 0.15 km , 500 +/- 50 m , swamp forest, 18/2/1993 , D.C. Lees: DL 93-0030; ♂, NE, Be Dinta, upper Anaovandrano R., W. Masoala, 500 m , 15.6733o S , 49.9885o E +/- 0.15 km , 500 +/- 50 m , 17/2/1993 : 16:30, D.C. Lees: DL 93-0031; ♂ [ BMNH ], Madagascar NE, Masoala, Be Dinta, R. Anaovandrano, 500 m , 20/2/ 1993 , D.C. Lees; ♀, NE, Iketry [Iketra], lower Anaovandrano R., W7-L2, Masoala E., 15.7216o S , 50.1753o E +/- 0.15 km , 270 m , 18/1/1994 , D.C. Lees; ♀, NE, Manosona, Anaovandrano R., E Masoala, 15.784o S , 50.2164o E +/- 0.15 km , 45 m , 1/2/1994 , D.C. Lees; ♂, NE, Masoala E., Antsamanarana R., 275 m , 15.295o S , 50.227o E +/- 0.15 km , 275 +/- 50 m , site W2L2-275M 5 LO, 5/12/1993 , H. Raharitsimba: DL 93-0028; ♂ [developmental abnormality LHS CuA1 HWV], NE, E Masoala, Ambavony R. [Ambanivony], W2-L1-125M [site], disturbed forest, 15.2813o S , 50.2877o E +/- 0.72 km , 223 +/- 50 m , 29/11/1993 , C. Kremen: CK93-0009; ♀, NE, E Masoala, [Ambavaony R.], bifurcation to 370 m , riparian, W2L1 Saharand. [Sarahandrano] 15.28o S , 50.2860o E +/- 1.4 km , 240 +/- 190 m , 25/11/1993 , H. Raharitsimba: TR 46A; ♂, NE, Masoala E., Antsamanarana R., W2L2 [site], 275 m , 15.2813o S , 50.2877o E +/- 0.72 km , 5LO [fruit trap], riparian, 11/12/1993 , H. Raharitsimba: DL 93-0026; ♀, data as above but: 275 m , 5/12/1993 , H. Raharitsimba, GEN 117 DL [genitalia]; ♂ ( Fig. 18 C), NE, Antsamanarana R., Masoala E., [ 50–520 m ], 15.307o S , 50.233o E +/- 0.1 km , 100 m , 29/11/1993 , C. Kremen: CK30, GEN 141 DL [genitalia]; ♀, data as above but: 15.2968o S , 50.2271o E +/- 1.31 km , 294 +/- 225 m , 10/12/1993 , D.C. Lees: E64 (egg voucher); ♂, NE, Antsamanarana R., Masoala E., 250 m , 15.3o S , 50.23o E +/- 0.15 km , 269 m , 12/12/1993 , H. Raharitsimba, GEN 142 DL [genitalia]; ♀, NE, Ambavaony R., E Masoala, [ 50–380 m ], 15.279o S , 50.288o E +/- 0.12 km , 150 m , 1/12/1993 , C. Kremen: CK37, EGG [egg voucher]; ♀, data as above but: 223 m , 29/11/1993 , C. Kremen; ♀, NE, E Masoala, Ambavaony R., riparian, W2-L1-50M [ SITE ], 15.2813o S , 50.2877o E +/- 0.72 km , 50 m , 2LO [fruit trap] 28/11/1993 , D.C. Lees: E3 [egg voucher; “pearly white, smallish, like Ht. pauper ”]; ♂, NE, E Masoala, Ambavaony [Ambanivony], 380 m , W2L1 [ SITE ] 15.2813o S , 50.2877o E +/- 5 km , 30/11/1993 , H. Raharitsimba: TR 61; ♂, NE, Ambavaony R., E Masoala, 125 m , 15.279o S , 50.288o E +/- 0.12 km , 125 m , 27/11/ 1993 , C. Kremen, GEN 124 DL [genitalia]; ♂, NE, Masoala, 1993, C. Kremen: CK93-0005, IA595 [isotope voucher]; ♂ [ BMNH ], NE, Reserve Speciale d'Ambatovaky, 700 m [ 16.8o S , 49.1222o E , 580 m ], 11/2/1990 , M.I. Evans [#1.16], 254 DL [genitalia].