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 vanewrighti Lees , sp. nov. LSID: urn:lsid:zoobank.org:act:6CE17307-FD07-47EB-A07A-70C0C94D3FB8 Prior references : sp. 63 ( Lees 1997 ). Type material., Deposition BMNH : Holotype : ♂ ( Fig. 9 A), Madagascar NW, Bekolosy, RS Manongarivo, 1280 m , low stature forest, 14.0295o S , 48.3095o E +/- 1 km , 1280 +/- 50 m , 18/12/1994 : 10:30, D.C. Lees: DLBEK 94_173, NHMUK 010289124 [ QTR barcode]; Paratypes : Deposition BMNH : ♀ ( Fig. 9 B), Madagascar NW, Bekolosy Mt., RS Manongarivo, 14.0427o S , 48.3028o E +/- 0.25 km , 1250 +/- 50 m , 23/12/1994 , D.C. Lees: DLBEK 94_175, NHMUK 010289125 [ QTR barcode]; ♂, NW, Bekolosy Mt., RS Manongarivo, 14.045o S , 48.306o E , 1162 m , at fruit, 22/12/1994 , D.C. Lees: DLBEK 94_130 [barcoded specimen: CCDB-02225-F04; BMAD 064-09; HM404241], IA219 [isotope voucher], BMNH (E) #697875, KA579 [=KA-P579, DNA extract voucher]; ♂, NW, Bekolosy, RS Manongarivo, Bekolosy, 1162 m , at fruit, 19/12/1994 : 10:45, D.C. Lees: DLBEK 94_163, IA328 [isotope voucher], NHMUK 010289126 [ QTR barcode]; ♂, NW, Bekolosy, RS Manongarivo, 1200–1230 m , 14.034o S , 48.31o E +/- 1 km , 1215 +/- 50 m , 20/12/1994 , 09:40–10:18, D.C. Lees: DLBEK 94_182, KA2055 [=KA-P2055; DNA extract number], NHMUK 010289127 [ QTR barcode]; ♂, NW, Bekolosy, RS Manongarivo, 1215 m , 14.034o S , 48.31o E +/- 1 km , 1215 +/- 50 m , 16/12/1994 , 07:00–07:52, D.C. Lees: DLBEK 94_162; IA329 [isotope voucher], NHMUK 010289128 [ QTR barcode]; ♀, NW, Bekolosy, RS Manongarivo, ridge ~ 1250 m medium stature forest, 14.0295o S , 48.3095o E +/- 1 km , 1295 +/- 50 m , 18/12/1994 , D.C. Lees: DLBEK 94_167, 1 greenish egg expressed 18/12: E53, NHMUK 010289129 [ QTR barcode]; ♂, NW, Bekolosy Mt., Manongarivo RS, 1200–1300 m , 14.0295o S , 48.3095o E +/- 1 km , 1250 +/- 50 m , D.C. Lees: 233 DL [genitalia], DL 9703 [ DNA voucher], IA466 [isotope voucher], NHMUK 010289181 [ QTR barcode]; Deposition MNHN: ♂, Madagascar NW, Bekolosy, RS Manongarivo, slope above camp, 1215 m , 14.034o S , 48.31o E +/- 1.5 km , 1215 m , 16/12/1994 , 07:34, D.C. Lees: DLBEK94_171; ♂, NW, Bekolosy, RS Manongarivo, near camp, 1163 m , 14.04575o S , 48.29623o E +/- 0.5 km , 1130 +/- 35 m , 12/12/1994 : 08:00–09:00, D.C. Lees: DLBEK94_176; ♂, data as above but: DLBEK94_172; ♂, NW, Bekolosy, RS Manongarivo, 1250 m , 14.0427o S , 48.3028o E +/- 0.25 km , 1250 +/- 50 m , 18/12/1994 : 09:20–10:30, D.C. Lees: DLBEK94_161; Deposition ABRI: ♂, Madagascar NW, Bekolosy, RS Manongarivo, 1250 m , 14.0427o S , 48.3028o E +/- 0.25 km , 1250 +/- 50 m , 12/12/1994 , 10:16–12:08, D.C. Lees: DLBEK94_168; Deposition PBZT: ♂, Madagascar NW, Bekolosy Mt., summit, RS Manongarivo, Madagascar NW, 1487 m , 14.0275o S , 48.3048o E +/- 0.5 km , 1482 +/- 25 m , 19/12/1994 , D.C. Lees: DLBEK94_118, IA602 [isotope voucher]; ♂, NW, Bekolosy, RS Manongarivo, 1350–1400 m , 14.0304o S , 48.3049o E +/- 0.16 km , 1375 +/- 25 m , 19/12/1994 , D.C. Lees: DLBEK94_111. Deposition summary : BMNH (HT, 5 PT ♂♂, 2 PT ♀♀), MNHN (3 PT ♂♂), ABRI (PT ♂), PBZT (2 PT ♂♂). Type locality. Madagascar NW, Bekolosy, RS Manongarivo, 14.0295o S , 48.3095o E +/- 1 km , 1280 +/- 50 m . Diagnosis. Especially in ♂♂, Ht. vanewrighti is the smallest species of Malagasy Heteropsis so far known (only in Africa are found smaller mycalesines such as Bicyclus nobilis (Aurivillius, 1893) : Condamin, 1973 ). It can normally be distinguished from Ht. harveyi by the lack of an androconial patch on the FWV, and by its restriction, as presently known, to the extreme northwestern part of the rainforest biome. In Marojejy, northeastern Madagascar , I found a population of a putatively undescribed species (equivalent to “ Hen. sp. 77” of Torres et al., 2001 ), which could be confused with Ht. vanewrighti , which sometimes exhibits extremely reduced FWV patch. However, counting from the start of the standard 358 bp COI-5P barcode, a barcoded individual of Ht. vanewrighti from Bekolosy (CCDB-02225-F04, BMAD064-09) has the following five potentially autapomorphic nucleotide states relative to other members (in parentheses) of the Ht. harveyi group: 268, 292, 547: T(C); 352: A(G); 451: C(T). FIGURE 9. Adults in DV (left), VV (right) A : HT ♂ of Heteropsis vanewrighti Lees , sp. nov. (Mt. Bekolosy; NHMUK010289124). B : PT ♀ of Ht. vanewrighti (Mt. Bekolosy; NHMUK010289125). C : HT ♂ of Ht. imerina Lees , sp. nov. (Mantadia; NHMUK010289132). D : PT ♀ of Ht. imerina (Ankasoka, route Lakato; DCL-DB-2906, MNHN). Scale bar: 30 mm. Description. Wings : Upperside dark brown, darkening towards FW costa and apex. The only ocellus strongly expressed is that of FWD CuA1, whose round black iris spans vein CuA1-CuA2 and has a very dull reddish-orange and fairly narrow concentric ring. HWD ocellus space-CuA1 faintly indicated as a black point, white pupil barely visible. A dark brown line flanked by slightly paler mid brown follows the strongly crenate HW border and the FW margin is also quite crenate. Underside mid brown with ocellus space-CuA1 a similar size to FWD with black iris also spanning veins CuA1-CuA2, more or less circular with a fairly narrow dull orange concentric ring. FWV space-M1 ocellus quite small, with black iris and faint very narrow orange ring. HWV space-CuA1 ocellus about half size of FWV one and elliptic with narrow dull orange ring. Rs-M3 and CuA2-1A ocelli expressed as minute white dots. Area between PMb and Mb dark brown-edged outside and inside respectively. HWV Mb irregular, a similar shape to Ht. andravahana as revised here, with a near right angle inflexion around vein CuA1, curving convexly around vein CuA2, and sharply convex above vein M2 to mid-costa. Distad of Mb there is mottling of mid and darker brown grey, with irrorated strigulae pronounced as on FWV. Four brown Cbs in FWV cell area do not stand out as a particularly distinct pattern from the general brown irroration, but tend to be parallel to one another and perpendicular to costa. A wavy Sml is indistinct but flanked distad with paler mid brown, while the distalmost Sml is dark brown and more distinct, following the crenate margin. Five short and convex-subtriangular dark brown tails at vein ends are interspersed with a few paler brown scales elsewhere in the fringe. Tails at M3- CuA2 are slightly more strongly produced than the others. Variation : ♂♂ similar, but on ventral surface, ocelli are sometimes expressed by stronger white dots (notably those of space-Rs-M3), and FWV space-CuA2 ocellus often includes a small black iris with faint orange ring, as in the ♀. ♀♀ tend to be larger with a paler mid-brown upperside and space-CuA1 ocellus tends to be slightly elliptic on HWD and strongly expressed in both wings, while FWD space-M1 ocellus is small and has a narrow dull orange ring. FWV space-M1 ocellus relatively large and circular to elliptic with a pale orange ring. HWV space-CuA1 Orng wider, dull red (in at least one ♀). The ventral surface of the ♀ has an especially mottled, ‘bark-cryptic’ pattern, consisting of intense russet brown irroration with ochreous on the HWV that is more strongly contrasting than in the ♂. Wingspan/fwl : range 30.2–33.7/ 15.8–17.4 mm (n=8, ♂♂), mean 32.05 +/- 1.4 SD/16.76 +/- 0.56 SD mm (n= 7 ♂♂), including HT ♂= 30.2/ 16.26 mm . Range 30.6–37.1/17.0– 18.7 mm (n=3, ♀♀); mean = 33.78 +/- 3.23 SD/ 17.99 +/- 0.86 SD mm (n=3 ♀♀). Androconia : discocellular brush ochreous-grey overlying a lenticular orange-brown patch. There is no androconial patch in FWV near 1A+2A as in related species. Variation not quantified. Palps : penultimate segment on outside face with narrow grey stripe, flanked by greyish cream scales, bordered by narrow dark brown scales; inside face greyish-ochreous, flanked by dark brown-black scales; ♀ similar. Male genitalia ( Fig. 8 B, 233DL, PT): from LV: tegumen shallowly domed (with small concave proximad notch from DV), with fairly distinct brow down to scythe-hooked uncus. Gnathos fairly straight, thin and tapered, in line with up-hook of uncus in specimen examined (slightly incurved and not sinuate from DV bending round from stout oblong base). Tegumen-vinculum division slightly more than maximum dimension of tegumen. Valve base fits in rather oblong frame with rather angular ‘shoulder’ at base of dorsal edge, leading abruptly to gently elbowed rather parallel-sided valve arm which has a very slender ‘head’ for club with stout small spine at tip, uprecurved and significantly proud of uncus tip (strongly incurved at tip from DV and with a few short distad teeth). Aedeagus very long and thin, dorso-ventrally flattened, more than one and a half times length of valve, gently and smoothly uprecurved distad from ostium, which is itself long and cupped proximad, winged into two lobes proximad from DV. Saccus relatively long, and juxta featuring a prominent down-lip, which also has small side lobes. Etymology. After R.I. Vane-Wright, who published the serious SEM study of mycalesine androconia (Vane- Wright, 1971), revised the subtribe in Sulawesi ( Vane-Wright & Fermon (2003)) , and who inspired the modern study of the subtribe in Madagascar . Discussion. The species was first recognized in the field by DCL as sp. 63 in December 1994 at Bekolosy mountain in the Réserve Spéciale de Manongarivo ( Lees: 1997 : 65), and subsequently found at the adjacent Mt. Antsatrotro in November 2011 . Although of a similar size, the HT of Henotesia wardii Butler, 1879 (= Heteropsis viettei Lees, 2003 ) has whitish shading distal of the HWV Mb. The only similar specimen actually described (but as detailed above not nomenclaturally available) is the HT of He. andravahana ab. marmorata Aurivillius, 1911 . This and a large series of specimens referred to Heteropsis harveyi in the Oberthür collection are clearly distinct from Ht. vanewrighti in being generally larger in size, possessing a FWV androconial patch, and occurring much further south in the central to southeastern parts of the rainforest biome. Additional information. DNA divergences : CO1-5P cluster number BOLD:AAE4111, with two haplotypes (exemplars BMAD064-09 (Bekolosy), BMAD212-15 and BMAD216-15 from Antsatrotro) is about 1.52% divergent to ‘ H . sp. 20’ (“sp. 77” of Torres et al., 2001 ; exemplars of cluster number BOLD:ACW5777 including HM 240620 ); for more details see under Ht. harveyi . Phylogeny/sister species : without a detailed phylogeographic analysis of the Ht. harveyi -complex, the sister taxon of Ht. vanewrighti is not yet certain, but the closest currently described relative is Ht. harveyi . Aduse-Poku et al., (2016 , in press) find it sister to Ht. sp. 20. Ecology and distribution. Habitat : montane rainforest with fine leaved climbing bamboos. Behaviour : at rest and in rainy weather, both sexes rest on tree trunks. Hostplant : unknown, assumed to be fine-leaved climbing bamboos among which the adults fly. Early stages : unknown. Distribution : Bekolosy and Antsatrotro mountains (where found in 2011), Réserve Spéciale de Manongarivo, to where, as far as is known, Ht. vanewrighti i s endemic ( Fig. 30 A, purple dots). Elevational range : 1045–1487 m . (n=73 including referred specimens and observational data). Referred specimens. ♂, Madagascar NW, Bekolosy Mt., RS Manongarivo, 14.045o S , 48.306o E +/- 3.5 km , 1150 +/- 350 m , 23/12/1994 , D.C. Lees: DLBEK 94_144A; ♂, NW, Bekolosy Mt., RS Manongarivo, 1370 m , 14.0263o S , 48.3075o E +/- 0.16 km , 1370 +/- 50 m , 16/12/1994 , 11:20, D.C. Lees: DLBEK 94_99; ♂, data as above but: DLBEK 94_100; ♂, NW, Bekolosy Mt., RS Manongarivo, 1163 m , 14.0458o S , 48.2962o E +/- 0.5 km , 1130 +/- 35 m , 12/12/1994 : 08:00–09:00, 11:20, D.C. Lees: DLBEK 94_169; ♀, NW, Bekolosy Mt., RS Manongarivo, 1225 m , 14.0335o S , 48.3098o E +/- 1 km , 1250 +/- 50 m , over river, 18/12/1994 : 09:30–10:20, 11:20, D.C. Lees: DLBEK 94_164; ♂, NW, Antsatrotro, c. 1250 m , 3/12/2011 : 13:52, D.C.Lees, IA610 [isotope voucher]; ♂, NW, Antsatrotro, c. 1260 m , 3/12/2011 , D.C.Lees, IA611 [isotope voucher]; ♀, NW Antsatrotro, c. 1250 m , 3/12/2011 , D.C.Lees, IA612 [isotope voucher]; ♂, NW, Antsatrotro, c. 1250 m , 3/12/2011 : 10:24, D.C.Lees, IA613 [isotope voucher]; ♂, NW, Antsatrotro, Manongarivo, 3/12/2011 : 09:25, WG46 [wing image], IA230 [isotope voucher]; ♂, NW, Antsatrotro, Manongarivo, 12/2011, DL 0033, KA753 [=KA-P753; DNA extract voucher].