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Heteropsis barbarae Lees & Kremen, sp. nov.
No full textHeteropsis barbarae Lees & Kremen, sp. nov. LSID: urn:lsid:zoobank.org:act:DC 87986 A- 3822 - 43 C 9 -B 1 B 9-83 A 2714 D088B 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, 15.1885 o S, 49.5163 o 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, 15.1907 o S, 49.5309 o 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, 15.1885 o S, 49.5163 o 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, 15.1907 o S, 49.5309 o 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, 15.1848 o S, 49.4659 o 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, 49.5309 o 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, 15.1885 o S, 49.5163 o 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-CuA 1 ocellus expressed strongly in FWD; orange ring sub-hexagonal., M 1 ocellus very small black white pupilled spot surrounded by a narrow orange ring. HWD ocellus CuA 1 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- M 2 to CuA 1. FWV space-CuA 1 ocellus similar size to FWD (in the HT), as is FWV M 1 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-CuA 1 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-CuA 1 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 1 A. 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-CuA 1 ocellus FWD is a little wider proximad in some specimens, and FWD M 1 ocellus sometimes not visible. HW space-CuA 1 ocellus is more circular in some specimens. 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 MAD 182 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 1 A+ 2 A base, mid brown, dispersed, juxtaposed potentially under abdominal sternal segments A 2 –A 5, 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 2 A 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 |MadagascarNo 10 [?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: 141 DL (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- 5 P cluster number BOLD:ACW 4995 (exemplar BMAD 248 - 15, DL 14 M-0033, Marojejy) is 4.86 % divergent to Ht. tianae (BOLD:AAE 4112), and 5.63 % divergent to Ht. maeva (BOLD:AAE 5488), 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], 14.4344 o S, 49.759 o E +/- 0.25 km, 850 +/- 50 m, 12 / 11 / 2006: 10:00, D.C. Lees: DL 06- 298; ♂, NE, Marojejy PN, 14.4344 o S, 49.7606 o 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, 14.4347 o S, 49.7601 o E +/- 0.5 km, 730 +/- 50 m, 10 / 11 / 2006, D.C. Lees: DL 06- 169; specimen, NE, Marojejy PN [between camp 1 and camp 2], 14.4362 o S, 49.7679 o E +/- 0.85 km, 613 +/- 125 m, 10 / 11 /2006, 15: 58, D.C. Lees: DL 06- 170; ♀, NE, Marojejy PN, 14.4377 o S, 49.7756 o E +/- 0.25 km, 400 +/- 50 m, 20 / 11 / 2006: 15:07, D.C. Lees: DL 06- 456; ♂, NE, Marojejy PN, 14.4425 o S, 49.7818 o 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, 14.45015 o S, 49.786 o E +/- 1.78 km, 326 +/- 175 m, 10 / 11 / 2006: 13: 16, D.C. Lees: DL 06- 204; ♂, NE, Marojejy PN [around entrée], 14.4626 o S, 49.7964 o 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], IA 387 [isotope voucher]; ♂ [BMNH], NE, Andranobe, Masoala Peninsula, 15.6816 o S, 49.9573 oE +/- 1.31 km, 0 m, 3 -Dec- 91, C. Kremen: CK 826, BMNH (E) 2008 - 69; ♂, NE, Masoala, Anaovandrano, 14 / 1 / 1994, D.C. Lees: WG 5 [wing image], IA 38 [isotope voucher]; ♂, NE, Masoala, Ambavaony, 125 m, 27 / 11 / 1994, D.C. Lees: DCL 47 a, IA 108 [isotope voucher]; ♂, NE, Masoala, Ambanivony, 25 / 11 / 1993, C. Kremen: CK 26, IA 109 [pheromone voucher]; ♀, NE, Makira, Vohitaly, 15.4414 o S, 49.5309 o E +/- 0.15 km, 645 m, 30 / 12 / 2002, D.C. Lees: DL-SF00, IA 112 [isotope voucher]; ♀, NE, Makira, Ankirindro, 15.2931 o S, 49.5472 o E +/- 0.15 km, 675 +/- 25 m, 15 / 1 / 2003, D.C. Lees: DL- SE00, IA 114 [isotope voucher]; specimen, NE, Ankiatomboka forest, c. 710 m, 15.179 o S, 49.412 o E, 710 +/- 50 m, 9 / 12 / 2001, D.C. Lees: DL 01- 169; ♂, NE, Bivontro, Makira, 15.3984 o S, 49.4477 o E +/- 0.1 km, 762 +/- 25 m, 21 / 12 / 2002, D.C. Lees: DL-SG 77; ♂, NE, Vohitaly-Anjiahely, 15.3989 o S, 49.5211 o E +/- 1.82 km, 435 +/- 50 m, 31 / 12 / 2002, BMNH (E) # 672331 [DNA voucher]; ♀, NE, Anjiahely, 0.37 km E., 15.4120 o S, 49.5022 o E +/- 0.25 km, 380 +/- 5 m 12 / 12 / 2002, D.C. Lees: DL-SG 1, BMNH (E) 2008 - 69, BMNH (E) 1717101; ♂, NE, Sahantaha, near ‘fanihy’ roost area, 15.1963 o S, 49.5564 o E +/- 0.98 km, 517 +/-54 m, 13 / 12 / 2001: 10: 46, D.C. Lees: DL 01- 257; ♂, NE, Anjanaharibe Mt., 15.188 o S, 49.6139 o E +/- 0.1 km, 475 +/- 5 m, 6 / 2 / 2003, D.C. Lees; ♀, NE, Tampolo, W. Masoala, ca. 150 m, primary rainforest, 15.7301 o S, 49.9604 o 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, 15.7279 o S, 49.9751 o E +/- 1.8 km, 32 +/- 25 m, 9 / 11 / 2001, D.C. Lees, DL 01- 27, BMNH (E) # 697188 [DNA voucher], KA 556 [=KA-P 556; DNA extract voucher]; ♀, NE, Andranobe, W. Masoala, 240 m, 15.4 o S, 49.58 o E +/- 1.78 km, 240 m. 3 / 2 / 1991, C. Kremen et al., Genitalia 157 [genitalia voucher]; ♀, NE, Andranobe, W. Masoala, 90 m, 15.6816 o S, 49.9573 o E +/- 1.79 km, 90 m, 14 / 11 / 1991, C. Kremen et al.: Biodiversity voucher; ♀, NE, W. Masoala, Andranobe field station, T 2 S 2 [site], 90 m, 15.6816 o S, 49.9573 o E +/- 1.79 km, 90 m, 14 / 11 / 1991, C. Kremen et al., Biodiversity survey voucher|Gen 158 [genitalia], IA 594 [isotope voucher]; ♂, data as above but: 2 / 11 / 1991, C. Kremen et al., Biodiversity voucher; ♂, NE, Andranobe, W. Masoala, 520 m, 15.65 o S, 49.983 o 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|Gen 156 [genitalia]; ♀, data as above but: 29 / 10 / 1991, C. Kremen et al.: Bio|Gen 159; ♀, NE, Andranobe, W. Masoala, 520 m, 15.65 o S, 49.983 o 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, 15.6733 o S, 49.9885 o E +/- 0.15 km, 500 +/- 50 m, 18 / 2 / 1993, D.C. Lees: DL 93 -0012, IA 593 [isotope voucher]; ♂, NE, Masoala E, Be Dinta, 550 m, 17 /02/ 1993, D.C. Lees: DL 93 -0029, IA 592 [isotope voucher]; ♂, NE, Be Dinta, upper Anaovandrano R., W. Masoala, 500 m, 15.6733 o S, 49.9885 o 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, 15.6733 o S, 49.9885 o 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., W 7 -L 2, Masoala E., 15.7216 o S, 50.1753 o E +/- 0.15 km, 270 m, 18 / 1 / 1994, D.C. Lees; ♀, NE, Manosona, Anaovandrano R., E Masoala, 15.784 o S, 50.2164 o E +/- 0.15 km, 45 m, 1 / 2 / 1994, D.C. Lees; ♂, NE, Masoala E., Antsamanarana R., 275 m, 15.295 o S, 50.227 o E +/- 0.15 km, 275 +/- 50 m, site W 2 L 2-275 M 5 LO, 5 / 12 / 1993, H. Raharitsimba: DL 93 -0028; ♂ [developmental abnormality LHS CuA 1 HWV], NE, E Masoala, Ambavony R. [Ambanivony], W 2 -L 1-125 M [site], disturbed forest, 15.2813 o S, 50.2877 o E +/- 0.72 km, 223 +/- 50 m, 29 / 11 / 1993, C. Kremen: CK 93 -0009; ♀, NE, E Masoala, [Ambavaony R.], bifurcation to 370 m, riparian, W 2 L 1 Saharand. [Sarahandrano] 15.28 o S, 50.2860 o E +/- 1.4 km, 240 +/- 190 m, 25 / 11 / 1993, H. Raharitsimba: TR 46 A; ♂, NE, Masoala E., Antsamanarana R., W 2 L 2 [site], 275 m, 15.2813 o S, 50.2877 o E +/- 0.72 km, 5 LO [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], 15.307 o S, 50.233 o E +/- 0.1 km, 100 m, 29 / 11 / 1993, C. Kremen: CK 30, GEN 141 DL [genitalia]; ♀, data as above but: 15.2968 o S, 50.2271 o E +/- 1.31 km, 294 +/- 225 m, 10 / 12 / 1993, D.C. Lees: E 64 (egg voucher); ♂, NE, Antsamanarana R., Masoala E., 250 m, 15.3 o S, 50.23 o E +/- 0.15 km, 269 m, 12 / 12 / 1993, H. Raharitsimba, GEN 142 DL [genitalia]; ♀, NE, Ambavaony R., E Masoala, [50–380 m], 15.279 o S, 50.288 o E +/- 0.12 km, 150 m, 1 / 12 / 1993, C. Kremen: CK 37, EGG [egg voucher]; ♀, data as above but: 223 m, 29 / 11 / 1993, C. Kremen; ♀, NE, E Masoala, Ambavaony R., riparian, W 2 -L 1-50 M [SITE], 15.2813 o S, 50.2877 o E +/- 0.72 km, 50 m, 2 LO [fruit trap] 28 / 11 / 1993, D.C. Lees: E 3 [egg voucher; “pearly white, smallish, like Ht. pauper ”]; ♂, NE, E Masoala, Ambavaony [Ambanivony], 380 m, W 2 L 1 [SITE] 15.2813 o S, 50.2877 o E +/- 5 km, 30 / 11 / 1993, H. Raharitsimba: TR 61; ♂, NE, Ambavaony R., E Masoala, 125 m, 15.279 o S, 50.288 o E +/- 0.12 km, 125 m, 27 / 11 / 1993, C. Kremen, GEN 124 DL [genitalia]; ♂, NE, Masoala, 1993, C. Kremen: CK 93 -0005, IA 595 [isotope voucher]; ♂ [BMNH], NE, Reserve Speciale d'Ambatovaky, 700 m [16.8 o S, 49.1222 o E, 580 m], 11 / 2 / 1990, M.I. Evans [# 1.16], 254 DL [genitalia].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 67-70, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis tianae Lees & Kremen, sp. nov.
No full textHeteropsis tianae Lees & Kremen, sp. nov. LSID: urn:lsid:zoobank.org:act:E 6864 E 90 -B 8 CF- 4 E 5 F- 99 C 4 -AC 879 E 6 F 70 A Prior references: sp. 49, 69 (Lees, 1997: Torres et al., 2001: 462). Type material., Deposition BMNH: Holotype: ♂ (Fig. 24 A), Madagascar C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1375 +/- 75 m, 2 / 3 / 2004, D.C. Lees: DL- 4-182, NHMUK 010289158 [QTR barcode]. Paratypes: Deposition BMNH: ♂ ( Fig. 25 A), Madagascar C, Anjozorobe 1400 m, 12 / 11 / 1994, C. Kremen, 228 DL [genitalia], NHMUK 010289191 [QTR barcode]; ♀ (Fig. 24 B), Madagascar C, Amboasary-an-ala, Vanjamanitra-Vazimba forest, Anjozorobe; towards start of Vanjamanitra trail, 18.4368 o S, 47.9469 o E +/- 0.25 km, 1316 +/- 5 m 3 / 3 / 2004: later PM, 1 egg expressed 4 / 3 /04; D.C. Lees: DL- 4-360; CCDB-02225-F09, BMAD 069- 0 9, HM 404245 [DNA barcode voucher], BMNH (E) # 676763 [DNA voucher; cytochrome b], IA 85 [isotope voucher]; ♂, C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 12–14 m canopy forest, flat summit, 18.4497 o S, 47.9404 o E +/- 0.15 km, 1320 +/- 50 m, 2 / 3 / 2004: 09: 27, D.C. Lees: DL- 4-169, 561 [= DL 0561; DNA extract number], BMNH (E) # 671926, IA 122 [isotope voucher]; 2 ♀♀ [BMNH], “Andrangalooka Forest, Madagascar ” [=Andrangoloaka, E. Lac Mantasoa], BMNH (E) # 674766 (Godman-Salvin Coll.; Fig. 24 C) and BMNH (E) # 674767; Deposition MNHN: ♂ [MNHN], Madagascar Centre 8 km S.E. d’Anjozorobe forêt de Vanjamanitra 1380 m 20 / 23 -X- 1966 P. Griveaud, J. Vadon et P. Viette|DCL-DB- 4471; Deposition ABRI: ♂, Madagascar C, Anjozorobe, 18.413 o S, 47.946 o E +/- 0.02 km, 1310 +/- 50 m, 22 / 10 / 2014, D.C. Lees: DL 14 Z-066. Deposition summary: BMNH (HT ♂, 2 PT ♂♂, 3 PT ♀♀), MNHN (PT ♂), ABRI (PT ♂). Type locality. Madagascar C, Amboasary-an-ala, vic. Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1375 +/- 75 m. Diagnosis. Heteropsis oberthueri, described below, is hard to distinguish from Ht. tianae except by ♂ genitalia (the gnathos is more flattened at the base than in that species) and by mitochondrial DNA sequence (DNA barcode highly distinct from ‘sp. 28 ’ to be treated in a subsequent paper, and from Ht. turbans (Oberthür, 1916); see under DNA divergences). On average, the HWV Mb of Ht. tianae is slightly more outdented near space-M 2 (Fig. 24 A– C), and the HWV tends to have the space-CuA 2 ocellus expressed as small black-ringed ocellus. However, Ht. oberthueri (Fig. 24 D–E, Fig. 26 A) can be distinguished from the sympatric Ht. andasibe by the distinctly more outangled HWV Mb of that species (Fig. 22 C; Fig. 26 B). In the Angavo massif, I observed the last species to prefer habitat only a few metres from the forest margin. The ventrally very similar Ht. roussettae usually has a smoother outcurve to the HWV Mb at space-M 2 and strong tendency to expression of a small ocellus in space-M 3 on the HWD (Fig. 22 AB). Description. Wings: upperside uniform mid brown, FW space-CuA 1 ocellus with an orange ocellus ‘ring’ which is sometimes rather hexagonal in shape and eccentric, being wider at proximad edge. FW space-M 1 ocellus small with narrow orange ring. HW space-CuA 1 ocellus the only one expressed there and somewhat elliptic. Darkish brown diffuse, slightly wavy hair-brush emanating mainly from below vein 1 A+ 2 A as far as mid-vein. Underside greyish brown, rather more uniform and less strongly irrorated than in many Heteropsis, rather uniformly irrorated brown and ochreous brown. Space-CuA 1 ocellus FWV with orange ring yellowing proximad, this Ore following the darkish brown concave curve of the Mb before it bends back to mid-costa. Space-M 1 ocellus FWV reduced to white pupil with narrow black iris without trace of orange ring. On HWV, space-CuA 1 ocellus slightly elliptic with narrow black iris and small also, without orange ring trace. HWV space-M 1 -M 2 ocelli as white ‘pupil’ points. HWV Mb darkish brown and fairly straight, only gently curving. FWV cell area with four darker brown rather equally spaced transverse Cbs. Basal areas are not much darker than areas distad of the Mb, and a darker brown PMb is weakly represented. Variation. ♂♂ similar to each other, but seasonal variation has not been quantified. HWV Mb sometimes with a yellow Mf distad of it in space-M 2. HWV space-CuA 2 ocellus may be slightly expressed as small white pupil with elliptic black iris. ♀♀ similar in pattern, but larger and somewhat lighter dorsally with a more rounded HW shape. Wingspan/fwl: range 34.7–40.2 / 18.1–20.4 mm (n= 3 ♂♂), including HT ♂: 34.7 / 18.56 mm; mean 37.0 +/- 2.1 SD/ 19.4 +/- 1.0 SD mm (n= 5 ♂♂). Range 35.2 –41.0/ 18.3–22.1 mm; mean 38.2 +/- 3.3 SD/ 20.4 +/- 1.7 SD mm (n= 5 ♀♀). Androconia: both 1 A+ 2 A and 3 A have a tapered ‘balloon-like’ inflation, more swollen distad, from base to mid-vein, covered in thin grey-brown scales (Lees 1997: 96, Fig. 3 A: sp. 49). HW veins M 2, M 3, CuA 1 and CuA 2 are also narrowly inflated throughout their lengths and have specialized scales on the dorsal surface (Lees 1997: 96, Fig. 3 A). Abdominal black androconia indistinctly visible ventro-laterally around A 2. HWD discocellular brush fawn to yellowish/blonde at tip. Discocellular patch hwd (orange, small, lenticular, composed of narrow yellow scales) (observations on MAD 239–242, MAD 244; Anjozorobe, n= 5). Palps: penultimate segment with narrow brown medial strip, flanked by yellow and fringed by dark brown scales, with yellow scales mainly on mesad face. ♂ genitalia: 228 DL, PT (Fig. 25 A): from LV, uncus slightly proud of dorsal curve of tegumen and longer than it, fairly straight and parallel-sided to hook without a distinct ‘head’ although slightly broadened ventrally towards tip (evident from the SV), and tegumen fairly narrow at hinge with vinculum; tegumen viewed laterally less tapered ventrad than many species. Valve arm with fairly short neck (distinctly recurved and bowed inwards from the SV) and slight club at tip covered in spinoid setae, with distinct ‘beak’ oriented towards uncus and slightly mesad, uncus tip distinctly proud of valves. Gnathos from rather small base with distinct flattening, more so than in Ht. oberthueri (although not with ear-like structure) near base, recurved downwards at tapered tip (and inwards from the SV) with slight serration on dorsal surface. Saccus moderate length and parallel sided, aedeagus slightly shorter than valve and strongly recurved twice, towards and away from the also proximally uprecurved ostium. Etymology. Etymology. After ‘Tiana’ Raharitsimba (= Heritiana Rahagalala), who found some of the first modern exemplars of this species at Anjozorobe. Discussion. Historical specimens, one from Andrangoloaka forest in the Angavo massif, probably late 19 th Century, a bit south of the type locality near Lac. Mantasoa (♀) and two ♀ (“ Madagascar ”) were found in BMNH, but the species was subsequently found in the field in Anjozorobe forest from 1994 by C. Kremen and coworkers where it was called “sp. 69 ” (Lees, 1997: 65), and in MNHN from 1966 collection. STs of the similar species Culapa antsianakana Oberthür, 1916 and of C. anceps Oberthür, 1916 (LT ♂♂ designated above under Ht. roussettae; see Fig. 22 C–D) were examined and are clearly different, along with Ht. oberthueri, described below. Additional information. DNA divergences: COI- 5 P cluster number BOLD:AAE 4112 (exemplar BMAD 200 - 15, DL 14 Z-051) is about 2.58 % divergent to Ht. oberthueri (cluster number BOLD:ACW 4996, exemplar BMAD 242 - 15, DL 06- 11, RNI Zahamena) and about 4.17 % divergent from ‘sp. 28 ’ (BOLD:AAE 5458), which is not likely to be the sister species. In their dataset for COII (Torres et al., 2001), Ht. tianae (as their “ Hen. sp. 49 ”, AY 040160, based on a ♂ from Anjozorobe, 1400 m.; see correction below) is considerably less pairwise divergent (4.76 %), however, to Ht. turbans (AY 040130, based on 2 ♂♂ and 1 ♀ from Ranomafana National Park; which has COI- 5 P cluster number BOLD: ACD 8579). Phylogeny/sister species: probably Ht. oberthueri based on the close relationship of the COI- 5 P barcode (confirmed by Aduse-Poku et al., 2016, in press). In their cladistic analysis of COII sequences, Torres et al., (2001: 467) suggested a topological relationship of Ht. tianae (sp. 49) with Ht. ankova but that was not supported (Torres et al., 2001: 466). Ecology and distribution. Habitat: montane rainforest. Behaviour: both sexes come readily to fruit bait. Flies in substratum of forest. Hostplant: unknown, presumed to be grasses. Early stages: unknown. Distribution: endemic to the Angavo massif, including Anjozorobe and Andrangoloaka, as far as is known (Fig. 30 C, dark pink dots). Due to a formatting error, the locality was incorrectly given in Table 1 of Torres et al., 2001 as “Anjanaharibe Sud” instead of the line below, “Anjozorobe 1400 m.” Elevational range: 1320–1375 m. (n= 27 including referred specimens and observations). Referred specimens. ♂ [MNHN], Madagascar Centre, forêt a l’Est du lac de Mantasoa, Andrangoloaka, 27 -II/ 6 -III- 1970 1389 m, P. Griveaud|DCL-DB- 2240; 2 ♂♂ [MNHN], data as above but: DCL-DB- 4469, DCL-DB- 4470; ♂ [BMNH; probably referable to Ht. tianae], Madagascar, Crowley bequest 1901 - 78 Rcvd as Mycalesis iboina Ward F.A.H. | 246 DL [genitalia]; 4 ♂♂, 4 ♀♀, Anjozorobe, 1380 m. [18.4084 o S, 47.9377 o E +/- 1.5 km], 4 / 12 / 1994, C. Kremen; ♂, Anjozorobe, “volo” circuit, 18.413 o S, 47.946 o E +/- 0.02 km, 1310 +/- 50 m, 22 / 10 / 2014 13: 51; D.C. Lees: DL 14 Z-051, BMAD 200 - 15 [DNA barcode voucher]; ♀, C, Anjozorobe, 1400 m, 18.4084 o S, 47.9377 o E +/- 1 km, 1400 +/- 100 m, 11 / 12 / 1994; D.C. Lees: DL 94 -0003A, BMNH (E) # 672411 [DNA voucher]; ♀, same data but 14 / 12 / 1994; C. Kremen: CK 753, IA 211 [isotope voucher]; ♀, same data but 138 m 14 / 12 / 1994, C. Kremen: CK 754; ♂, C, Anjozorobe, 18.4498 o S, 47.939 o E +/- 0.2 km, 1323 +/- 25 m, 2 / 3 / 2004; D.C. Lees et al.: DL- 4 -166, 2576 [= DL 2276; DNA extract number], BMNH (E) # 676776, IA 9 [isotope voucher]; ♂, C, Amboasaryan-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1375 +/- 75 m, D.C. Lees: DL- 4- 165; ♂, same data but DL- 4-180; ♂, same data but DL- 4-181; ♀, same data but DL- 4-382; ♂, same data but DL- 4- 429; ♂, same data but DL- 4-395; ♂, same data but R. Ranaivosolo: DL- 4-405; ♂, same data but 2 / 3 / 2004: 09: 26, R. Ranaivosolo: DL- 4-198; ♀, same data but 09: 36, R. Ranaivosolo: DL- 4-199; ♂, same data but 13: 15, R. Ranaivosolo: DL- 4-212; ♂, same data but DL- 4-214; ♂, same data but 15: 15, R. Ranaivosolo: DL- 4-218; ♂, C, Amboasary-an-ala, Vanjamanitra-Vazimba forest, Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1375 +/- 75 m, 3 / 3 / 2004: 15: 41, D.C. Lees: DL- 4-366; ♂, data as above but: later PM, D.C. Lees: DL- 4-347; ♂, data as above but: D.C. Lees: DL- 4-356, 582 [= DL 0582; DNA extract number], IA 123 [isotope voucher]; ♂, C, Amboasary-an-ala, Vanjamanitra-Vazimba forest, Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1375 +/- 75 m, 3 / 3 / 2004: later PM, D.C. Lees: DL- 4 -361, 1051 [= DL 1051; DNA extract number], BMNH (E) # 672416, HDO, F-A, H-A, ABD, H-M [androconia sampled; K. Bubbinga study]; ♂, C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1375 +/- 75 m, 4 / 3 / 2004: 10: 52, D.C. Lees: DL- 4-417; ♂, data as above but: DL- 4- 418; ♂, C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, ‘just by S 3 ’, 18.4505 o S, 47.9399 o E +/- 0.15 km, 1323 m, 2 / 3 / 2004: 08: 56, D.C. Lees: DL- 4-167, IA 19 [isotope voucher]; ♂, C, Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1330 m, 4 / 2 / 2014: 15: 20, D.C. Lees: MAD 240 [pheromone voucher], IA 403 [isotope voucher]; ♂, Anjozorobe, Babakoto trail, c. 1350 m, K. Aduse-Poku, 04/02/ 2014, MAD 243 [pheromone voucher], KA 2059 [=KA-P 2059; DNA extract voucher]; specimen, Anjozorobe, KAP-MAD 1408, KA 1018 [=KA-P 1018; DNA extract number].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 79-82, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis oberthueri Lees, sp. nov.
No full textHeteropsis oberthueri Lees, sp. nov. LSID: urn:lsid:zoobank.org:act:FEE 8 CA 6 F-A 53 B- 43 A 5-8 C 9 F-E 5 D 69 AD0E 468 Prior references: sp. 89, sp. 87 (on some envelopes). “KA 518 _Heteopsis_ anceps ” [sic] in Aduse-Poku et al., (2015, Fig. 1). Type material. Deposition BMNH: Holotype: ♂ (Fig. 24 D), Madagascar E, Zahamena NW, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 5 / 11 / 2006, D.C. Lees: DL 06-064, G04 [leg for DNA], NHMUK 010289159 [QTR barcode]. Paratypes: Deposition BMHN: ♂, Madagascar E, Ambavala camp, Zahamena, 17.5451 o S, 48.7237 o E +/- 0.5 km, 1300 +/- 50 m, 6 / 11 /2006, 13: 10, D.C. Lees: DL 06- 130 A, NHMUK 010289160 [QTR barcode]; ♀ (Fig. 24 E), E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: DL 06- 130, IA 585 [isotope voucher], NHMUK 010289161 [QTR barcode]; ♂, E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: 371 DL, NHMUK 010289192 [QTR barcode]; ♂, E, Zahamena NW, 17.53 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 6 / 11 / 2006, D.C. Lees: DL 06-086, IA 586 [isotope voucher], NHMUK 010289162 [QTR barcode]; ♂, E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6- 7 / 11 / 2006, D.C. Lees: DL 06- 1001, IA 324 [isotope voucher], NHMUK 010289163 [QTR barcode]; ♀, E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 /2006, 11: 30, D.C. Lees: DL 06- 112, IA 587 [isotope voucher], NHMUK 010289164 [QTR barcode]; ♀, E, Towards Ambavala camp, Zahamena, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: DL 06- 141, IA 588 [isotope voucher], NHMUK 010289165 [QTR barcode]; ♀, E, returning to Bemoara camp, Zahamena, 17.5356 o S, 48.71875 o E +/- 0.70 km, 1180 +/- 100 m, 5 / 11 /2006, 14: 59, D.C. Lees: DL 06-059, IA 589 [isotope voucher], NHMUK 010289166 [QTR barcode]; Deposition MNHN: ♂, Madagascar E, Ambavala, Zahamena NW, 17.5451 o S, 48.7237 o E +/- 0.25 km, 1325 +/ - 50 m, 6 / 11 /2006, 14: 11, D.C. Lees: DL 06- 117, IA 584 [isotope voucher], DSC 03096.jpg [image]; Deposition ABRI: ♂, Madagascar E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: DL 06- 121. Deposition summary: BMNH (HT ♂, 4 PT ♂♂, 4 PT ♀♀), MNHN (PT ♂), ABRI (PT ♂). Type locality. NW Zahamena, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m. Diagnosis. The species is particularly similar to Ht. tianae described above and in fact few obvious distinguishing features are apparent that are likely to distinguish all examples, but that species is however allopatric, only known from the Angavo massif, and on average, the HWV Mb of Ht. oberthueri is relatively straight and there is usually no space-M 3 ocellus expressed on the HWD (Fig. 24 D–E). The wings have a slightly greyer cast. Ht. oberthueri also strongly resembles Ht. andasibe. It can be distinguished from that last species (which has slightly more conspicuous white spots on the ventral wing surfaces) by the less straight, usually more evenly outangled HWV Mb at space M 2 in Ht. andasibe (Fig. 22 C), and from more yellowish lowland Ht. strigula which have a conspicuous orange crescent on the HWD, and from Ht. ankova which is smaller and has ocellus space-M 3 as well as in space-CuA 1 expressed on the HWD (Fig. 19 C). The ventrally very similar Ht. roussettae has a strong tendency to expression of a small ocellus M 3 on the HWD (Fig. 22 A–B). The potentially sympatric (in ‘Antsianaka’ region) Ht. anceps has a relatively straight HWV Mb and a yellowish ventral cast with sparse light brown irroration and a distinct black ventral androconial patch on the ♂ sternal abdomen (Fig. 22 D). The ♂ genitalia are similar to those of Ht. tianae, differing by the more strongly inpointed subterminal spine on the valve and the terminal head covered in spinoid setae being enlarged. They also differ from the potentially sympatric Ht. anceps (Oberthür, 1916), being smaller and with a narrower distal valve tip (Lees, 1997: 107). DNA differences separating Ht. oberthueri from Ht. tianae are indicated by Aduse-Poku et al., (2016, in press). See below regarding male genitalia. Description. Wings: description is similar to that of Ht. tianae. Upperside uniform mid brown, with FW space-CuA 1 ocellus featuring an orange ocellus ‘ring’ which is sometimes rather ‘squashed’ along the diagonal from mid-costa to tergal angle. FWD space-M 1 ocellus small with narrow orange ring. HWD space-CuA 1 ocellus is the only one expressed there and is somewhat elliptic, with a narrow orange ring. Darkish brown, diffuse, slightly wavy and uprecurved hair-brush emanates mainly from below vein 1 A+ 2 A, as far as mid-vein. Underside greyish brown, similar to Ht. tianae, rather more uniform and less strongly irrorated than in many Heteropsis, rather uniformly irrorated brown and ochreous brown. Space-CuA 1 ocellus with orange ring yellowing proximad, following darkish brown concave curve of Mb before it bends back to mid-costa. Space-M 1 ocellus FWV a white pupil in HT. On HWV, space-CuA 1 ocellus small and slightly elliptic with narrow black iris, also without orange ring trace. HWV space-Rs-M 2 ocelli as white pupil-points and space-CuA 2 ocellus hardly expressed in HT. HWV Mb darkish brown and fairly straight but minutely irregular, only gently curving, with very small yellow Mf distad of it in space-M 2. FWV cell area with four darker brown rather equally spaced transverse Cbs. Basal areas not much darker than areas distad of Mb, and darker brown PMb weakly represented. Variation. ♂♂ similar to each other, but seasonal variation unknown as the entire type series was taken in a few days in early November. Space- M 1 ocellus FWV sometimes features a narrow black iris and space-M 2 sometimes just as a white pupil without narrow black iris nor trace of orange ring. Space-CuA 2 ocellus HWV slightly expressed in some specimens. ♀♀ similar in pattern, but larger and somewhat lighter dorsally with a more rounded HW shape. Wingspan/fwl: range 31.2–35.5 mm./17.0– 18.79 mm (n= 6 ♂♂), mean= 33.8 +/ 1.44 SD/ 17.87 +/- 0.54 SD mm (n= 6 ♂♂), including HT ♂ 32.4 / 18.79 mm. Range 34.0– 39.7 / 17.8 –20.0 mm (n= 4 ♀♀), mean = 36.2 +/- 3.03 SD/ 18.75 +/- 1 SD mm (n= 4 ♀♀). Androconia: HWD vein 1 A+ 2 A has a narrowly tapered inflation and 3 A is much more swollen distad, from base to mid-vein, covered in thin grey-brown scales. The length of HW veins M 2, M 3, CuA 1 and CuA 2 are also narrowly inflated, and have specialized scales on the dorsal surface. Abdominal black androconia are just visible ventro-laterally around A 4 –A 5. HWD discocellular brush dark brown. Sdp HWD grey, small, lenticular, composed of narrow grey scales). Palps: penultimate segment with narrow brown medial strip, flanked by yellow more towards where labial palp potentially wipes compound eye and fringed by dark brown scales, with yellow scales mainly on mesad face, brown inside towards tip. ♂ genitalia: 371 DL (Fig. 25 B, PT): description more or less as for Ht. tianae: from LV, uncus very slightly proud of dorsal curve of tegumen (which has a small proximad notch from DV) and slightly longer than it, fairly straight and parallel-sided to hook without a distinct ‘head’ towards tip (inflated from DV before tip), fairly narrow at hinge with vinculum. Valve without prominent dorsal ‘shoulder’. Valve arm with fairly short ‘neck’ and slight club at tip covered in spinoid setae (its crest particularly raised from DV), with distinct ‘beak’ oriented towards uncus and slightly mesad, with none of valve tip proud of uncus. Gnathos from rather small base with deepened (although not ear-like structure) near base, recurved downwards at tapered tip (appearing sinuate from DV and slightly inrecurved at pointed tip). Saccus not very long and parallel sided, aedeagus slightly shorter than valve and quite recurved, both towards and away from ostium, also proximally uprecurved. ♂ genitalia different from those of potentially sympatric (Ht. anceps (Oberthür, 1916)), which are larger and with broader distal valve tip (Lees, 1997: 107). Etymology. after Charles Oberthür, who in 1916 produced one of the first treatments, a seminal work on the Malagasy Mycalesina. Discussion. No historical material has been found in museums. The species was first found in the field during an expedition to northwest RNI Zahamena in November 2006, where it was hoped to rediscover Ht. anceps. The STs of the similar species Culapa antsianakana Oberthür, 1916 and of Culapa anceps Oberthür, 1916 (LT ♂ designated above under Ht. roussettae) were examined but the species strongly resembles Ht. tianae (see above). Additional information. DNA divergences: COI- 5 P cluster number BOLD:ACW 4996 (exemplar BMAD 242 - 15, DL 06- 11), about 2.58 % divergent to Ht. tianae (BOLD:AAE 4112, exemplar BMAD 200 - 15, DL 14 Z-051). Phylogeny/sister species: this species was not known at the time of Lees (1997). Closely related to Ht. tianae (sister in Aduse-Poku et al., 2016, in press). which it closely resembles, and based on the close relationship of the COI- 5 P barcode. Ecology and distribution. Habitat: primary rainforest. Behaviour: flies low. Hostplant: unknown, but likely to be low-growing grasses. Early stages: unknown. Distribution: as far as is known, endemic to RNI Zahamena (Fig. 30 C, brown dots). Elevational range: 1180–1310 m. (n= 27, including referred specimens). Referred specimens. ♀, Madagascar E, returning to Bemoara camp, Zahamena, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 5 / 11 / 2006, D.C. Lees: DL 06-060, G03 [leg for DNA]; specimen, data as above but: DL 06-066; specimen, data as above but: DL 06-067; ♀, E, Zahamena NW, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 5 / 11 / 2006, D.C. Lees: DL 06-061; ♂, E, Zahamena NW, 17.53 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 6 / 11 / 2006, D.C. Lees: DL 06-070; ♂, data as above but: DL 06- 118, BMAD 242 - 15 [DNA barcode]; ♀, E, just above cascade, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006: 15: 29, D.C. Lees: DL 06- 084; ♂, data as above but [without time]: DL 06- 132, DSC _0825 [photo]; ♀, data as above but: DL 06- 119, IA 432 [isotope voucher], KAP 518 [=KA-P 518; extract number, sequenced specimen in Aduse-Poku et al., 2015, mislabelled as Heteropsis anceps); ♀, data as above but: DL 06- 124; ♂, E, Ambavala camp, Zahamena, 17.5451 o S, 48.7237 o E +/- 0.5 km, 1300 +/- 50 m, 6 / 11 /2006, 13: 23, D.C. Lees: DL 06- 126; ♂, data as above but: 13: 16, D.C. Lees: DL 06- 127; ♂, E, Ankosy summit, Zahamena, 17.4946 o S, 48.733 o E +/- 1 km, 1321 m, 7 / 11 /2006, 10: 37, D.C. Lees: DL 06- 156; ♂, E, near Ankosy summit, Zahamena, 17.49405 o S, 48.73325 o E +/- 0.067 km, 1300 +/- 25 m, 7 / 11 / 2006, D.C. Lees: DL 06- 188; ♀, E, Zahamena, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 6 / 11 / 2006, D.C. Lees: DL 06- 67, IA 69 [isotope voucher]; ♂, E, Zahamena NW, 17.4946 o S, 48.733 o E +/- 1 km, 1321 +/- 25 m, 6 / 11 / 2006, D.C. Lees: 364 DL [genitalia voucher], IA 29 [isotope voucher], leg sample [KAP].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 83-85, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis kremenae Lees, sp. nov.
No full textHeteropsis kremenae Lees, sp. nov. LSID: urn:lsid:zoobank.org:act:D 5567 C 45 - 4 EB 9 - 4 E 62 - 8 F 67 -ABF 2 C 4008 F 4 E Prior references: sp. 14 A (Lees 1997, Torres et al., 2001: 462). Type material., Deposition BMNH: Holotype: ♂ (Fig. 13 A), Madagascar NE, Ambavaony R. [= Ambanivony R.], R. Onive (Ankavanana), E Masoala, 250 m, 15.282 o S, 50.288 o E +/- 0.1 km, 250 +/- 50 m, 28 / 11 / 1993, D.C. Lees: DL 93 -0021, NHMUK 010289135 [QTR barcode]. Paratypes: Deposition BMNH: ♂, Madagascar NE, Ambavaony R., E. Masoala, 125 m, 15.279 o S, 50.288 o E +/- 0.12 km, 125 m, 27 / 11 / 1993, H. Raharitsimba: DL 93 -0004, NHMUK 010289136 [QTR barcode]; ♀, data as above but: DL 93 -0005, fruit trap 14 LO, disturbed forest, NHMUK 010289137 [QTR barcode]; ♂, NE, Ambavaony R., E. Masoala, 280 m, 15.282 o S, 50.288 o E +/- 0.1 km, 280 +/- 50 m, 28 / 11 / 1993 13: 46, riparian streamside, shade rainforest, D. C. Lees: DL 93 -0022, NHMUK 010289138 [QTR barcode]; ♀ (Fig. 13 B), NE, Masoala Peninsula, Manosona, 45 m. slope, 15.784 o S, 50.2164 o E +/- 0.15 km, 45 m, 28 / 1 / 1994, D.C. Lees: DL 94 -0004, E 105 [egg voucher], W 7 L 1-45 M 1 LO [fruit trap], NHMUK 010289139 [QTR barcode]; ♀, NE, Ambavaony R., E Masoala, " 200 m. ", 15.282 o S, 50.288 o E +/- 0.1 km, 200 +/- 50 m, 28 / 11 / 1993; D.C. Lees: DL 93 -0009 [DNA voucher], E 4 [egg voucher: “ 3 greenish eggs expressed 29 / 11 ”], NHMUK 010289140 [QTR barcode]; ♂ (Fig. 15 A), NE, Masoala E, Ambanivony [=Ambavaony], 15.288 o S, 50.288 o E +/- 0.5 km, 50 +/- 25 m, site W 2 -L1, 50 m, riparian, fruit trap 2 lo, 26 / 11 / 1993, D.C. Lees: 19 DL [genitalia], NHMUK 010289183 [QTR barcode]; ♀, NE, Antsamanarana R., Masoala E., 15.295 o S, 50.227 o E +/- 0.15 km, 275 m, 11 / 12 / 1993, H. Raharitsimba: CK 93 -0020; NHMUK 010289184 [QTR barcode]. Deposition MNHN: ♂, NE, Ambavaony R., E. Masoala, ca, 100 m, disturbed streamside, 15.279 o S, 50.288 o E +/- 0.12 km, 180 +/- 75 m, 28 / 11 / 1993, C. Kremen: DL 93 -0007; ♂, NE, Antsamanarana R., Masoala E., "~ 50 m, Piste C", 15.307 o S, 50.233 o E +/- 0.1 km, 115 +/- 75 m, 11 / 12 / 1993; D.C. Lees: DL 93 -0008; ♀, NE, Antafononana lower Anaovandrano R., W 7 -L 2, Masoala E., 15.7458 o S, 50.1858 o E +/- 0.15 km, 40 m, H. Raharitsimba: DL 94 - 0 0 0 3, W 7 L 2-40 M 2 LO [fruit trap], IA 600 [isotope voucher]; Deposition ABRI: ♂, NE, Ambery R., Masoala E., 15.3716 o S, 50.4263 o E +/- 0.98 km, 26 m, 22 / 12 / 1994, D.C. Lees: DL 93 -0006; ♀, NE, Masoala PN, W 7 L 2, Antafonona, 40 m, fruit trap 1 Hi, riparian, 15.738 o S, 50.182 o E +/- 1.85 km, 185 +/- 145 m, 17 / 1 / 1994, H. Raharitsimba: DL 94 -0002. Deposition summary: BMNH (HT ♂, 3 PT ♂♂, 3 PT ♀♀); MNHN (2 PT ♂♂, PT ♀); ABRI (PT ♂, PT ♀). Type locality. Madagascar NE, Ambavaony R., R. Onive, E Masoala, 250 m, 15.282 o S, 50.288 o E +/- 0.1 km. Diagnosis. Apparently most similar to Ht. pauper, from which it differs by rounder wings especially in FW wingshape (as in Ht. subsimilis and Ht. avaratra sp. nov.), and more generally by a larger space-CuA 1 HWD ocellus and more contrasting orange and yellow banding/shading on the underside, as well as a more consistently orange FWV ‘shoulder’ at the base of the costa). ♂♂ can be distinguished by HWD Spdb tending to be blackish (whitish in Ht. pauper). Description. Wings: Dorsal wing surface uniform mid-brown, FWD space-CuA 1 ocellus with black iris at least spanning veins CuA 1 -CuA 2, light orange ring narrow to wide and sub-hexagonal, sometimes compressed along diagonal from tergal angle to mid costa. FWD space-M 1 ocellus very small, with narrow pale orange ring. HWD ocellus is the only one expressed there and fairly round with fairly narrow pale orange ring and spanning about 2 / 3 of vein CuA 1 -CuA 2, on FWV space-CuA 1 ocellus similarly expressed to FWD but with wider ring that is pale orange with yellow area proximad marking the tapering arm of a spiral that closely hugs the concave brown Mb before it bends back to mid costa. On HWV, space- 1 A ocellus much more strongly expressed than on HWD and sub-elliptic, black iris spanning most of vein CuA 1 -CuA 2 and with a concentric yellow ring that spans the whole space-CuA 1. Space-Rs ocellus is on HWV very reduced in HT. HWV Mb irregular but not jagged, russet and darker towards its distad margin, convex to intersection with vein M 2 and then fairly straight to mid space- CuA 2 and angled back to 1 A. Yellow-ochreous highlighting distad of Mb. Russet colouring shadows Mb proximad to PMb and within this area, the background grades from rich orange to ochreous (this variegation of orange and yellowish is a good field character for the species) towards base and there is quite strong irroration with russetbrown. A diffuse wavy grey-brown line follows margin in both wings and the Sml is distinct and dark brown, hugging closely the margin, which is particularly crenate in the HW (more so than any other of the Ht. subsimilis group). Distad of the yellowish highlighting beyond the Mb, yellowish background is interspersed with grey-brown irroration. In the FWV cell, the four russet brown, slightly convex arcs/lines delineate two more yellowish Cbs for the two outermost pairs of arcs. The basal half of the FWV costa (‘shoulder’) that is bisected by darker strigulae is strongly tinted with orange, as less prominently shown in most specimens of Ht. pauper. Variation. Sexes similar in upperside and underside colouration but ♀ larger. In ♀ HWV especially, two ocelli may be expressed in space- CuA 2, often as white spots. HWV space-Rs ocellus sometimes expressed, even when small, with elliptic black iris and narrow yellow ring. Wingspan/fwl: range 31.0– 33.9 /17.0– 18.6 mm (♂♂, n= 5), mean = 33.2 +/- 2.1 SD/ 17.8 +/- 0.8 SD mm (n= 3 ♂♂), including HT ♂ 33.9 / 18.6 mm; range 33.8–37.5 / 18.1–20.7 mm (♀♀, n= 6); mean = 35.9 +/- 1.4 SD/ 19.2 +/- 1.1 SD mm (n= 3 ♀♀). Androconia: Sdb HWD compact, dark brown to blackish, Sdp HWD fat lenticular, black, CuA 2 inflated vein narrowly swollen to about 2 /3, 1A+ 2 A vein also inflated. The degree of HW inflation may be variable, but these veins do not exhibit discrete ‘balloons’ (swellings); Lees (1997: 96) showed for one specimen that the veins that were significantly inflated along their length comprised only M 3 for the medial system. Palps: penultimate segment on outside face light creamy white towards compound eye, blackish away from eye with light coloured scales within this border; fringed with dark brown where not contacting eye. Face away from eye whitish ochreous. ♂ genitalia: 19 DL, PT (Fig. 15 A); Lees (1997: 106, Fig. 7 f; “ 14 A”): from LV: miniaturised, about 1.7 mm long and with configuration typical of the Ht. subsimilis group; there is no really obvious difference in shapes of the genitalic components from other members (Lees (1997: 106) and the description below might apply to almost any of the group. From LV, the valve bases are rather symmetrically-‘skittle’-shaped (Lees 1997). Uncus is slightly longer than tegumen and quite inflated dorsoventrally before the slightly pointed downturned tip (inflated before tip from the DV). Gnathos tapered from small base, fairly straight, not sinuate from the SV. Valve with prominent rounded dorsal shoulder, valve arm tapering to distinct ‘beak’ that points to uncus base with limited serration and not much indication of a club at valve end (tip strongly incurved from the SV). Saccus relatively small, slightly bulbous, juxta not very prominent proximad. Aedeagus about 1 / 5 longer than valve, moderately stout and recurved distad of ostium. Etymology. After Claire Kremen, who worked tirelessly for the creation of the Masoala National Park, that was created in 1994. Discussion. No historical museum material has been found and this species was first recognized in the field in 1993 in the survey of what is now Masoala National Park (Lees, 1997: 64). Kremen et al., (2001: 412) considered it a potential endemic there, but it has since been discovered to be more widespread among the remaining northeastern lowland rainforests. All available types in the Ht. subsimilis group were examined, from which it differs. Concerning Culapa pauper Oberthür, 1916 with over 300 STs, a ♂ LT is here designated: BMNH (E) # 674888; Madagascar Antsianaka Perrot Freres 2 e Semestre 1890, which was illustrated by Oberthür, 1916 (Pl. 367: f. 3066); the PLTs then include the ♀ BMNH (E) # 674889, also illustrated in Oberthür, 1916 (Pl. 367: f. 3067). The STs of Culapa comorana Oberthür, 1916 were also examined (LT ♂, here designated, that illustrated by Oberthür, 1916: Pl. 367, f. 3061). The HT ♂ of Pseudonympha subsimilis Butler, 1879 (BMNH (E) # 674882) was examined (Fig. 14 A), with which species Culapa undulata Oberthür, 1916 (a taxon that d’Abrera 1980: 185 was unable to locate) has been synonymised (Lees et al., 2003) (LT ♂, here designated, Fig. 14 C, bearing labels “ Lectotype | Madagascar Antsianaka Perrot Freres 2 e Semestre 1890 | Culapa undulata Obthr. ♂ type |[copy of f. 3064]|P.E.L. Viette det. 1968 Culapa undulata Ch. Obthr. ♂ LECTOTYPE |supposed “ Type ” (syntype) of Culapa undulata Oberthür ”| BMNH (E) # 674883; Oberthür specified as STs 190 other ♂♂ (including BMNH (E) # 674885) and 58 ♀♀ (including BMNH (E) # 674884, Fig. 14 B, illustrated in Oberthür 1916: P. 367, f. 3065 and BMNH (E) # 674886), which are then automatically PLTs of Culapa undulata). Additional information. DNA divergences: cluster number BOLD:ACW 4937 (exemplar BMAD 249 - 15, Marojejy), diverges by 5.47 % from that of H. avaratra (cluster number BOLD:AAD0195, exemplar BMAD 016- 09) and by about 5.5 % from that of H. subsimilis (cluster number BOLD:AAB 4493). In the Torres et al., (2001) dataset, a COII sequence for Ht. kremenae (their “ Hen. sp. 14 A”) from Masoala is 7.03 % pairwise divergent to Ht. avaratra from Montagne d’Ambre (AY 040161 compared to AY 040146 based on single individuals respectively, 398 bp comparable) and 8.14 % pairwise divergent to Ht. subsimilis from Ranomafana National Park (AY 040145), whereas 8.83 % pairwise divergent to Ht. pauper from that site (AY 040133, 414 bp compared). Meanwhile, Ht. avaratra and Ht. subsimilis are more obviously closely related according to COII data (5.27 % pairwise divergent) and share privately at least four SNPs not found elsewhere in the Ht. s ubsimilis group. Note that in the Torres et al., dataset, “ Henotesia 14 ” (AY 040181, cytochrome b) from Masoala is apparently synonymous with Ht. subsimilis (AY 040192), with only 1 bp difference. A similar situation seems to pertain to their grouping of (‘ Hen. sp. 23 ’ + ‘ Hen. sp. 30 ’ [representing opposite sexes of Ht. pauper from Ankazomivady] + Ht. pauper from Ranomafana + “ Hen. sp. 7 ” from Masoala), the latter ‘morphospecies’ exhibiting only 3 bp difference from the first three (COII); see also the COI phylogeography of Ht. pauper in Linares et al., (2009: 488–489). Phylogeny/sister species: Ht. kremenae is likely most closely related to Ht. subsimilis and Ht. avaratra. Lees (1997) did not resolve the position of Ht. kremenae ‘FRTNA’] within the Ht. subsimilis group. The two-gene study of Linares et al., (2009) did not include this species and its exact position was not clear in Aduse-Poku et al., (2016, in press). Ecology and distribution. Habitat: Primary rainforest, preferring riparian areas. Behaviour: flies among low grasses, liking the slopes above streams and rivers. Hostplant: not reared, but presumably low forest grasses. One was caught though in a canopy fruit trap in Marojejy (K. Aduse-Poku, pers. comm.). Early stages: expressed eggs were greenish. Distribution: endemic to lowland rainforest in the northeast of the rainforest belt and Masoala Peninsula (Fig. 30 B, dark blue dots), with a preference for riparian areas. Elevational range: 10–675 m (n= 86, including referred specimens and observations). Referred specimens. ♂, NE, Marojejy PN, riparian forest near camp 1, 14.4377 o S, 49.7756 o E +/- 0.5 km, 420 +/- 50 m, 20 / 11 / 2006: 16:06, D.C. Lees: DL 06- 454; ♂, NE, Marojejy PN, camp 1, riparian forest, 450 m, 14.4377 o S, 49.7756 o E +/- 0.5 km, 450 +/- 50 m, 21 / 11 / 2006: 15: 54, D.C. Lees: DL 06- 485; ♂, data as above but: 21 / 11 /2006, 15: 54, D.C. Lees: DL 06- 484; ♂, NE, Marojejy PN, 1 km d'entrée, 14.4548 o S, 49.792 o E +/- 0.5 km, 258 +/- 50 m, 22 / 11 / 2006: 09: 51, D.C. Lees: DL 06- 510; ♂, NE, Marojejy, 650 m, 24 / 1 / 2014: 11: 37, D.C.Lees: DL 14 M-0037, IA 528 [isotope voucher]; CCDB-02230-E 11 [DNA barcode voucher]; ♀, NE, Marojejy, 700 m, 24 / 1 / 2014: 11: 21, D.C.Lees: DL 14 M-0035, IA 532 [isotope voucher]; ♂, NE, Sahantaha, Makira, 15.2337 o S, 49.5311 o E +/- 0.15 km, 500 +/- 130 m, 29 / 1 / 2003, D.C. Lees: BMNH (E) # 697172 [DNA voucher]; ♀, NE, Andreketa near 17 B 364 steep rainforest with lots of “Tsongolovo” 15.2642 o S, 49.5479 o E +/- 0.75 km, 364 +/- 25 m,? 17 / 1 / 2003, BMNH (E) # 672340 [DNA voucher, cytochrome b]; ♀, data as above but: 17 / 1 / 2003, D.C. Lees, BMNH (E) # 672434 [DNA voucher]; ♀, NE, Andreketa, 15.2642 o S, 49.5479 o E +/- 0.75 km, 375 +/- 25 m, 17 / 1 / 2003, D.C. Lees: BMNH (E) # 697950 [DNA voucher]; specimen, NE, Ambodivoangy forest, 15.29 o S, 49.62 o E +/- 1 km, 50 +/- 25 m, 4 / 12 / 2001; ♂, NE, Ankirindro, 15.2904 o S, 49.5474 o E +/- 0.25 km, 636 +/- 25 m, 16 / 1 / 2003; D.C. Lees: BMNH (E) # 697302 [DNA voucher]; ♀, NE, Antsamanarana R., Masoala E., 275 m, 15.295 o S, 50.227 o E +/- 0.15 km, 275 m, 11 / 12 / 1993, H. Raharitsimba: TR 89; ♂, NE, Antsamanarana R., Masoala E., 50 m, 15.307 o S, 50.233 o E +/- 0.1 km, 50 m, 12 / 12 / 1993; H. Raharitsimba: TR 93; ♂, NE, Vohitaly, 15.4377 o S, 49.5344 o E +/- 0.15 km, 28 / 12 /2002, 15: 15, D.C. Lees; ♂, NE, Vohitaly, Makira, 15.4379 o S, 49.5344 o E +/- 0.15 km, 25 / 12 / 2002, D.C. Lees: DL-SF 3, KA 552 [=KA-P 552, DNA extract number]; ♂, NE, Ambatoavy, Masoala, 630 m, 15.658 o S, 50 o E +/- 1.26 km, 630 m, 14 / 2 / 1993, D.C. Lees: H 14-14293 - 1; ♀, NE, Ambavaony R., E Masoala, 125 m, 15.279 o S, 50.288 o E +/- 0.12 km, 125 m, 30 / 12 / 1993; C. Kremen: E 15 [egg voucher], DNA DCL 32 [DNA voucher]; ♀, NE, Iketry [Iketra], lower Anaovandrano R., W 7 -L 2, Masoala E., 15.7458 o S, 50.1858 o E +/- 0.15 km, 40 m, 16 / 1 / 1994, D.C. Lees, W 7 L 2-40 M 2 LO [fruit trap]; ♀, NE, Masoala, 15 / 1 / 1994, WG 2 [wing voucher], IA 33 [isotope voucher]; ♀, NE, Masoala, 16 / 1 / 1994, IA 36 [isotope voucher]; ♂, NE, Masoala, 50 m, 12 / 12 / 1993, H. Raharitsimba, WG 6, [wing image], IA 39 [isotope voucher]; ♂, NE, Masoala, 26 / 11 / 1993, H. Raharitsimba: TR 47, IA 47 [isotope voucher]; ♂, NE, Ivontaka R. camp, between Ivontaka N and S. reserves, ca. 90 m, 16.294 o S, 49.81 o E +/- 0.7 km, 90 +/- 25 m, 3 / 2 / 1995; D.C. Lees: DL 95 -0003A, H 14 A 3295 - 1 [leg sample]; specimen, NE, Ivontaka R., ~ 120 m, 16.294 o S, 49.81 o E +/- 1.5 km, 120 +/- 25 m, 5 / 2 / 1995; D.C. Lees: DL 95 -0002A; ♂, NE, Ivontaka R., path to Marokoto, border of river, ca. 100–360 m, 16.297 o S, 49.785 o E +/- 2 km, 230 +/- 130 m, 3 / 2 / 1995, 11: 46 – 16: 10, D.C. Lees: H 14 A 3295 - 2 [leg sample], IA 601 [isotope voucher]; ♀, NE, Ivontaka Sud, 16.294 o S, 49.81 o E, 90 m, 2 / 1995, D.C. Lees, IA 338 [isotope voucher]; ♀, NE, Ivontaka Sud, path to Marokoto, 16.297 o S, 49.785 o E +/- 2 km, 230 +/- 130 m, 3 / 2 / 1995, D.C. Lees: H 14 A 3296 - 2, IA 339 [isotope voucher].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 46-51, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis menamenoides Lees, sp. nov.
No full textHeteropsis menamenoides Lees, sp. nov. LSID: urn:lsid:zoobank.org:act:D 2 FCD 726 -B 198-484 F- 846 C-B 5229 FE 174 E0 Prior references: sp. 62 (Lees 1997; Torres et al., 2001: 462). Type material., Holotype, deposition BMNH: ♂ (Fig. 21 A), Madagascar NW, Bekolosy, RS Manongarivo, 950 [880] m, [14.0487 o S, 48.29495 o E +/- 0.15 km, 95 m +/- 50 m], 15 / 12 / 1994: 09: 50, D.C. Lees: DLBEK 94 _ 160; KAP 16 [=KA-P 16 DNA extract voucher], IA 312 [isotope voucher], NHMUK 010289153 [QTR barcode]. Paratypes: Deposition BMNH: ♂, NW, Bekolosy, RS Manongarivo, ca. 800 m, [14.05135 o S, 48.2937 o E +/- 0.15 km, 800 +/- 50 m], 16 / 12 / 1994, D.C. Lees: DLBEK 94 _ 179, DL 9639 [DNA voucher], 0 443 [= DL 0443; DNA extract number], BMNH (E) # 697857 [cytochrome b]; CCDB-02225-G03, BMAD 075-09; HM 40425 [DNA barcode voucher], KA 571, KA 4084 [=KA-P 571, KA-P 4084; DNA extract numbers]; ♂ (Fig. 18 D), Madagascar NW, Bekolosy, RS Manongarivo, 890 m, 14.04936 o S, 48.294 o E +/- 0.15 km, 890 +/- 60 m, 15 / 12 /1994, 10: 59, D.C. Lees: DLBEK 94 _ 180, 231 DL (genitalia), BMNH (E) # 1053956; ♂, NW, Bekolosy, RS Manongarivo, 925 m, 14.04894 o S, 48.2945 o E +/- 0.15 km, 925 +/- 50 m, 13 / 12 /1994, 09: 56 – 11: 15, D.C. Lees: DLBEK 94 _ 181, NHMUK 010289187 [QTR barcode]. Deposition summary: BMNH (HT ♂, 3 PT ♂♂). Type locality. Madagascar NW, Bekolosy, RS Manongarivo, 950 m, [14.0465 o S, 48.3 o E +/- 0.2 km, 950 +/-70 m]. Diagnosis. Heteropsis maeva is similar on the dorsal wing surface. In Asia, a group that includes Mydosama anapita (Moore, 1858), M. marginata (Moore, 1881) and M. patiana Eliot (1869) (see Brattström et al., 2014) are superficially similar on both surfaces. Like M. anapita, of which Mycalesis menamena Mabille 1877 (“ Madagascar ”; see d’Abrera 1980: 186 for a photograph) is a synonym according to Lees et al., (2003), Ht. menamenoides has a dark orange Mb and Smb ventrally, with contiguous pale yellow shading distad to the Mb, and dark orange somewhat triangular blotches distal to that. However, in Ht. menamenoides, only the space-CuA 1 ocellus is expressed as typical of the Ht. strigula group, and the HW margin is less crenulated, whereas in the Mycalesis anapita group, a full complement of ocelli are expressed, especially on the HWV. Within the ‘true’ Malagasy mycalesine fauna, the upperside of Ht. menamenoides is similar to that of Ht. maeva, to which the species is apparently closely related, and shares a smooth HW margin, but differs by a much lighter and more uneven orange colouration on both wing surfaces, whereas the HW-crenulate Ht. barbarae is not known from the northwest of Madagascar, and has a almost entirely mid brown dorsal wing colouration. Among the only other distinctly ‘orange’ Heteropsis (in both sexes) known from Madagascar, the Malagasy Region complex of Ht. narcissus (Fabricius, 1798) lacks ventral abdominal dark androconial scales in the ♂, while Ht. laetifica and Ht. laeta which are similar in dorsal wing pattern, have them, but are paler yellow on underside. Orange ♀ forms of Ht. erebina (‘ Culapa grandis Oberthür 1916 ’) and Ht. antahala (‘ Mycalesis benacus Mabille, 1884 ’) (see, e.g., Lees, 1997: 56 and d’Abrera, 1997) are also not confusable because the Mb shows through to the upperside more strongly and delineates an area just proximad of the space-CuA 1 ocellus which is highlighted more contrastingly in paler orange. Ht. erebina and Ht. antahala belong to different clades with different wing shapes and particularly ventral wing patterning. In particular, none of these species have such a distinctly zigzagged orange Sml, that is especially characteristic of the FWV of Ht. menamenoides. Description. Wings: Upperside orange with wide darkish brown border along margin that widens from costa towards apex in FWD and that is more diffuse on HWD, narrowest at margin and broader at costa and tergal edge. Lighter orange is evident around space-M 3 of FWD and HWD. Space-CuA 1 ocellus expressed more strongly in FWD than in HWD; Orng not evident and black iris considerably smaller diameter than spacing of veins CuA 1 and CuA 2. Underside generally of lighter orange cast especially distad of Mb of central symmetry system, and there is a diffuse darker orange band running through arc of ocelli and a very wavy darker orange line before another darker orange Sml that closely tracks the margin on both wings; the margin is fairly smoothly cut and only very gently crenate. The dark orange Mb is relatively straight in the HWV and curves towards the tergal angle of the FWV, tergad of the space-CuA 1 ocellus, which is very small in the HT. In FWV, Cbs delineated by two sets of approximately parallel darker orange transverse lines in the FWV cell area, not as evident as in the (anyway brown) species Ht. roussettae. Space-M 2 ocellus as small white point surrounded by narrow black ring in FWV while space-CuA 1 ocellus of HWV only slightly larger; no other ocelli expressed. Basal area strongly strigulated with darker orange lines in both wings, and PMb is more evident in the HWV than FWV. Variation. ♂♂ similar to each other, but seasonal variation has not been quantified, as the type series were collected over a few adjacent days in mid December. ♀ unknown. Wingspan/fwl: range 36.2–45.5 / 19.7–24.6 mm (n= 4 ♂♂); mean 39.4 +/- 5.3 SD/ 21.9 +/- 2.0 SD mm (n= 4 ♂♂), including HT ♂ 36.2 / 19.7 mm. ♀ unknown. Androconia: black ventral abdominal patch between A 3 –A 6 divided centrally by narrow yellow overlying scales, in potential contact with diffusely separated and slightly wavy androconial brush largely emanating from base of space-CuA 2 before its fork with vein CuA 1 and overlying veins 1 A+ 2 A and 3 A; the latter symmetrically inflated before midlength in compact ‘tear’-shape (Lees, 1997: 97, Fig. 3 b), whereas 1 A+ 2 A shows very restricted swelling towards the base of 1 A. Fairly short and compact Sdb brownish at base and strongly light yellow towards tip overlying small Sdp with small underlying swelling in HW vein R towards end of cell (Lees, 1997: 97, Fig. 3 b); not described in detail here due to scarcity of material. Palps: penultimate (medial) segment mainly yellow with a narrow medial dark brown streak, fringed by dark scales on outside face and by light yellow scales on inside face of palp; last segment similarly patterned. ♂ genitalia: 231 DL (DLBEK 94 _ 180; Fig. 18 D, PT): from LV, tegumen broader dorsally and very angled proximad (scarcely any notch from DV), very narrow at hinge with vinculum; dorsal edge of tegumen forming a fairly straight line to uncus featuring fairly narrow and evenly deep ‘neck’ distad to hook-tip with distinct dorsal ‘head’ as fairly typical for Ht. strigula group; uncus quite strongly inflated before tip from DV. Gnathos particularly straight and tapering (but slightly sinuate from DV and inrecurved at tip), emanating from rounded base and in the specimen examined, transversing uncus at about 40 degrees. Valve with a long straight dorsal ‘shoulder’. Valve base leads without strong constriction to quite broad and flat valve arm covered in spinoid setae along entirety of fairly truncate tip, featuring a slight distal lobe (incurved from DV), with uncus hook-tip protruding to middle of extension of this truncate tip (from LV). Saccus not very long and aedeagus about same length as valve, strongly uprecurved from midlength and featuring a long proximad ostium, slightly bulbous at tip. Juxta somewhat ‘plate-lipped’ at proximad tip. Etymology. A direct reference to the quite superficial similarity of this new species to Mycalesis menamena Mabille, 1877 (see d’Abrera, 1980), supposedly from Madagascar but actually a synonym of the SE Asian (Mycalesis) anapita Moore, [1858] (Lees, 1997; Lees et al., 2003). Discussion. This distinctive species I first recognized in the field in December 1994 at Bekolosy mountain in the RS Manongarivo (Lees: 1997: 65). It was not detected in my 2011 expedition to the adjacent mountain Antsatrotro and no historic museum material is known. All relevant types of the ‘orange’ species in Madagascar in the Ht. strigula group were examined (see under Ht. barbarae), as well as the ♀ HT of Culapa grandis Oberthür, 1916 (bearing labels “Nord-Madagascar (Antakares) Isokitra a Diego-Suarez Mai a Octobre 1891 E. & B. Perrot| Culapa grandis Obthr. ♀ type |[Copy of f. 3074]”). Additional information. DNA divergences: COI- 5 P cluster number BOLD:AAE 4113 (exemplar DLBEK 94 _ 179, BMAD 075-09, HM 404251). The DNA barcode is about 5.5 % divergent to an undescribed species, sp. 28 (cluster number BOLD:AAE 5458), and about 5.8 % pairwise divergent to that of Ht. tianae (BOLD:AAE 4112). Phylogeny/sister species: unknown, new molecular study awaited. Ecology and distribution. Habitat: on a very steep slope with a fine leaved bamboo that was considered to be ‘ Nastus ’ manongarivensis A. Camus, not necessarily the hostplant. Behaviour: flies in the forest substory. Hostplant: unknown, but possibly grasses. Early stages: unknown. Distribution: endemic as far as is known to Mt. Bekolosy, Reserve Speciale de Manongarivo (Fig. 30 C, slateblue dots). Elevational range: 800–1035 m. (n= 8 incl. observations). Conservation: one of the mycalesines with the smallest known ranges in Madagascar, and likely to be confined to a narrow ‘ecotonal’ bioclimate. The four known specimens were found on a steep ridge in a radius of less than 0.2 km. Referred specimens. None.Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 72-74, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
FIGURE 27 in Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision
No full textFIGURE 27. Adults in DV (left), VV (right). A: HT ♂ of Heteropsis borgo Lees, sp. nov. (Anjanaharibe Sud; NHMUK010289167). B: LT ♂ of Mycalesis butleri Mabille, 1880 ('Madgscr', BMNH(E) 674771). C: HT ♂ of Pseudonympha angulifascia Butler, 1879 ('Antananarivo', BMNH(E) 674769). Scale bar: 30 mm. D: ♂ genitalia of Ht. borgo (Anjanaharibe Sud; CK712, 284DL), DV, SV. Scale bar: 3 mm.Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on page 87, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis westwoodi Lees, sp. nov.
No full textHeteropsis westwoodi Lees, sp. nov. LSID: urn:lsid:zoobank.org:act: 422 E 5526 - 8 FBB- 4 AD 6 -A 47 D-EC 95 FF 54 E 16 F Prior references: sp. 13 B (Lees, 1997: 64). “KA 522 _Heteopsis_ andravahana ” [sic] (Aduse-Poku et al., 2015: Fig. 1). Type material., Deposition BMNH: Holotype: ♂ (Fig. 10 A), Madagascar SE, Vohitrasiva, near ridge, " 1390 m ", Andringitra, [mountaintop on way to Vohipia (1760 m pass)], 22.1684 o S, 47.0289 o E +/- 0.1 km, 1450 +/- 50 m, 17 / 2 /1995, 11: 40 – 13: 15, D.C. Lees: DL 95 -0002; KAP 20 [=KA-P 20; DNA extract number], NHMUK 010289130 [QTR barcode]. Paratypes: Deposition BMNH: ♂, data as HT but DL 95 -0001, IA 288 [isotope voucher], KA 751 [=KA-P 751; DNA extract number], NHMUK 010289131 [QTR barcode]; ♂, data as HT but DL 95 -0003, 236 DL [genitalia], IA 290 [isotope voucher], NHMUK 010289182 [QTR barcode]. Deposition MNHN: ♂, Madagascar, forêt d’Anjavidilava, Andringitra Oriental, 2005 m, 1 / 15 -I- 1971, P. Griveaud et P. Soga|DCL-DB- 2923; six ♂♂, data as above but: representing the rest of series from DCL-DB- 2919 to DCL-DB- 2925. Deposition summary: BMNH (HT ♂, 2 PT ♂♂), MNHN (7 PT ♂♂). Type locality. Madagascar E, Vohitrasiva, Andringitra, 22.1684 o S, 47.0289 o E +/- 0.1 km, 1450 +/- 50 m. Diagnosis. There is only one particularly similar species to Ht. westwoodi: Ht. viettei has a similar white band distad of the HWV Mb, but is generally smaller and much more widely distributed (sympatric in Fianarantsoa province) in Madagascar. Among related species, Ht. harveyi ♂♂ have yellow rather than whitish highlighting distad of the Mb in some forms. Description. Wings: dorsal surface fairly uniform dark brown, darker in area of cell and above towards costa. FWD space-CuA 1 ocellus expressed, roughly circular, its black iris spanning inter-vein distance CuA 1 -CuA 2, with a narrow concentric dark orange ring. HWD space-cuA 1 ocellus elliptic, with a black iris spanning more than half intervein distance CuA 1 -CuA 2, and a dark orange ring relatively slightly broader than in the FWD and nearly spanning the intervein distance. No other ocelli expressed on FWD. A dark Sml quite closely hugs the margin in both wings and both surfaces. HW margin moderately and evenly crenulate, with rather blunt dark blackish brown tails on the otherwise violet-grey-brown fringe. Ventral surface with a light grey-violet cast, especially on the HW. FWV ocellus space-CuA 1 is circular and same size as on the FWD with a concentric quite narrow orange ring. M 1 ocellus FWV fairly small and circular with black iris and faint narrow orange ring. A small ocellus with very small black iris is expressed just below the M 1 ocellus FWV. In the HWV the space-CuA 1 ocellus is as on the HWD and space-R 5 -M 3 ocelli are expressed as very small white points. Mb irregular, strongly concave from the margin to M 2 where it forms a cusp, then strongly and irregularly indented and maximally concave in space-CuA 1, convex again in space-CuA 2 and concave again to vein 1 A+ 2 A where it terminates. The Mb HWV delineates a relatively darker brown irrorated area proximad to the PMb that only slightly repeats the shape of the Mb. Distad of the Mb from veins M 2 - 1 A is a clear white band, also irregular, with prominence in space M 2 and maximum width proximad of space-CuA 1 ocellus, tapering to 1 A+ 2 A with a slight expansion in Space-CuA 2. Distad of this white band is a slightly paler brown area irrorated less densely with darker brown than the band in the more basal area. Patterning on the FWV is fairly uniform, darker proximad of the Mb that is concave around the space-CuA 1 ocellus and recurves back towards mid-costa, with indications of three or four paler stripes above the cell. There is slightly paler brown highlighting in premarginal area just proximad of the dark Sml. Variation. A PT ♂ (DL 95 - 0001) has a much stronger violet-grey cast, especially to the HWV. Sexually dimorphic (Fig. 10 A–B), ♀ larger and lighter in referred specimen BMNH (E) # 674842 (Fig. 10 B), but similar in ventral patterning, although with a more contrasting darker band between the HWV PMb and Mb. Wingspan/fwl: range 37.5–40.3 / 19.7–20.7 mm (n= 3 type ♂♂); mean = 37.4 +/- 1.1 SD/ 20.7 +/- 0.3 SD mm (n= 3 ♂♂), including HT ♂ 38.6 / 20.7 mm. 44.1 / 22.8 mm (n= 1 referred ♀). Androconia: HWD discocellular brush light brown. HWD discocellular patch ‘bullet’-shaped, peach pink coloured. Palps: outside face ochreous with distinct dark brown medial strip, fringed with thin blackish scales except where palps brush eye, and sandwiching ochreous hair scales away from eye. ♂ genitalia: 236 DL, PT ♂ from Vohitraseva (Lees, 1997: 109, Fig. 7 i, “ 13 B”; see Fig. 11 A). From LV, a very shallowly domed tegumen with a rather straight dorsal edge profile and slight brow leads to a ‘hand-scythe’-shaped uncus, which is fairly evenly deep except towards the tip of the hook, where it features more of a ‘head’ dorsad; very thin from DV. Tegumen with wide ‘U’-shaped proximad notch, from LV proximad profile bending round sharply to a constriction at junction with vinculum. This ‘waist’ is over half the maximum dimension of the tegumen (and thus the tegumen does not have a quadrate shape from LV). Valve with a very prominent convex dorsal ‘shoulder’ making up part of an oblong shape which leads to the narrow quite tapering and fairly straight valve arms, which are ventrally slightly elbowed and with a very slightly expanded club extending beyond the uncus tip, with a dorsal tooth and with a slightly incurved, rather blunt and serrate distal region (pointed and somewhat incurved from DV); this club covered in an array of spinoid setae on its mesad face. Gnathos from rightangled base pointing in line with the uncus tip in its porrect position, relatively straight and tapered to pointed tip and with negligible serration; from DV slightly sinuate and inrecurved at tip. Saccus moderately extended and markedly inflated proximad, less than a third of total valve length. Aedeagus very thin, dorsoventrally flattened and uprecurved in distad half, with a long ostium proximad, proximad tip rather semicircular from DV. Juxta quite prominent proximad and keel-like from LV, bilobed from DV. Etymology. After John Obadiah Westwood, founder of the genus Heteropsis and describer of the remarkable Heteropsis drepana. The Hope Department of Entomology at Oxford, that Westwood founded, contains a number of other interesting species from Madagascar. Discussion. No historical museum material is known and the first field specimens of this species were recognized by C. Kremen in 1988–1991 in what is now Ranomafana National Park (Lees, 1997: 64). The sole surviving ST ♀ (in BMNH; now a LT) of Mycalesis andravahana was examined and belongs to a not closely related species, as discussed above. The HT of Henotesia wardii Butler, 1879 was also examined (BMNH (E) # 674838) together with its genitalic slide dissection (P. Viette no. 4847; see also above) and this species (now known as Heteropsis viettei Lees, 2003) is clearly not conspecific, despite the superficial similarity to Ht. westwoodi of Pl. 5, Figs. 6, 7 of Mabille ([1885]). The specimen represents Ht. viettei, as discussed above. Additional information. DNA divergences: COI- 5 P cluster number BOLD:ACW 4998 (exemplars BMAD 134 - 15 DL 06- 980 from RS Kalambatritra and BMAD 243 - 15 CK 92 -0005 from PN Andohahela; also KA- P 510 (see Aduse-Poku et al., 2015) from Ambondrombe). Phylogeny/sister species: apparently most closely related to Ht. harveyi (cluster number BOLD:ACW 5694 and to Ht. viettei (BOLDAAK 5839). Lees (1997) recovered Ht. westwoodi (as ‘THRTB’) as sister to Ht. viettei (‘THRTN’), based on all morphological evidence, genitalic characters, and non-genitalic characters, but without any jackknife support. Aduse-Poku et al., (2016, in press) confirm it as sister to Ht. viettei. Ecology and distribution. Habitat: montane rainforest with fine-leaved climbing bamboos. Behaviour: ♂♂ fly quite rapidly in the substory. Hostplant: suspected to be fine-leaved climbing bamboos. Early stages: unknown. Distribution: endemic to the southeastern mountains of Madagascar from Andohahela north to Andringitra, Mt. Ambondrombe and Mt Maharira, Ranomafana National Park (Fig. 30 A, pink dots). Elevational range: 785–1817 m. (n= 27 including referred specimens and observational data). Referred specimens. ♂, Madagascar E, Ambondrombe, 21.8782 o S, 47.2474 o E +/- 0.5 km, 1604m, +/- 50 m, 22 / 3 / 2004, D.C. Lees et al.: DL- 4-882, 638 [= DL 0638; DNA extract number], BMNH (E) # 672003; ♂, E, Ambondrombe, 21.87874 o S, 47.24969 o E +/- 1 km, 170 +/- 50 m, 17 / 2 / 1995, D.C. Lees, DL- 4-863, KA 522 [=KA- P 522; extract number; sequenced specimen in Aduse-Poku et al., 2015]; ♀, E, Andringitra, ridge, 1800 m, 20 / 11 / 2011, D.C.Lees: DL- 11 -0001, IA 645 [isotope voucher]; ♂, SE, Andohahela, Antananandava, 1200 m, 48 54 'E 24 34 'S, [24.5735 °S, 46.2 °E +/-km, 1200 m], 29 / 11 / 1992; ♂, data as above but BMNH (E) # 674844; ♂, SE, Andohahela, Antananandava, site S 10, 1200 m, 48 54 'E 24 34 'S, [24.5735 o S, 46.2 o E 1200 m], 29 / 11 / 1992, CK 92 - 0 0 0 5, BMAD 243 - 15 [DNA barcode number]; ♀ (Fig. 10 B), SE, PN Andohahela, Antananandava, 1200 m, 48 54 'E 24 34 'S, [24.5735 o S, 46.2 o E, 1200 m], 29 / 11 / 1992, BMNH (E) # 674842; ♂, SE, Andohahela, C. Kremen, BMNH (E) # 697865, 238 BDL [genitalia], IA 636 [isotope voucher]; specimen, SE, Befarara, RS Kalambatritra (western), 23.417 o S, 46.43 o E, 1507 m +/- 140 m, D.C. Lees: DL 06-980, 15-Dec- 2006: 11: 59, BMAD 134 - 15 [DNA barcode voucher]; ♂, SE, Befarara, RS Kalambatritra (western), 23.4186 o S, 46.44025 o E +/- 0.042 km, 1569 +/- 25 m, 12 / 12 / 2006: 11: 47, R. Ranaivosolo: DL 06- 767; ♂, SE, Befarara, RS Kalambatritra (western), 23.418 o S, 46.44 o E +/- 0.15 km, 1557 +/- 25 m, 12 / 12 / 2006, M. Linares: DL 06- 811, IA 161 [isotope voucher]; right wings imaged.Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 35-38, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis lanyvary Lees, sp. nov.
No full textHeteropsis lanyvary Lees, sp. nov. LSID: urn:lsid:zoobank.org:act: 264464 C 2 -C 4 E 2-4470 -A 209-348 CDCCBDF 85 Prior references: sp. 67 (Lees 1997: 66, where considered the same species as Ht. ankova; sp. 68 [label on specimen in PBZT; “sp. 68 ” was an unrelated morphospecies in Lees, 1997: 66]. Type material., Deposition BMNH: Holotype: ♂ (Fig. 19 A), Madagascar NW, Tsaratanana, piste Mangindrano-Antetykalambazaha, 14.187 o S, 48.945 o E +/- 1 km, 1615 +/- 75 m, flat grass-bamboo area, 20 / 12 / 2004 12:04, D.C. Lees: DL-05- 744, TF 19 [leg for DNA], NHMUK 010289144 [QTR barcode]. Paratypes: Deposition BMNH: ♀ ( Fig. 19 B), Madagascar NW, Tsaratanana, piste Mangindrano- Antetykalambazaha, flat grass-bamboo area, 14.187 o S, 48.945 o E +/- 1 km, 1615 +/- 75 m, 20 / 12 / 2004 12:04, D.C. Lees: DL-05- 745, TF 14 [leg for DNA], NHMUK 010289145 [QTR barcode]; ♂, same data as HT but DL-05- 849, TF [legs for DNA] [abdomen lysed], IA 624 [isotope voucher], MAD 233 - 15 [DNA barcode voucher], NHMUK 010289146 [QTR barcode]; ♂, same data as HT but DL-05- 846, TF 12 [tissue sample for DNA], BMAD 232 - 15 [DNA barcode], IA 294 [isotope voucher], NHMUK 010289147 [QTR barcode]; ♂ (Fig. 18 B, right), same data as HT but DL-05- 845, IA 192 [isotope voucher], [thoracic fragment for KAP], NHMUK 010289148 [QTR barcode]; ♂, same data as HT but DL-05- 856, IA 623 [isotope voucher], NHMUK 010289149 [QTR barcode]; ♂, NW, Tsaratanana, near forest margin, 14.2034 o S, 48.9532 o E +/- 2.5 km, 1441 +/- 25 m, 24 / 12 / 2004, D.C. Lees: DL-05- 851, IA 622 [isotope voucher], NHMUK 010289150 [QTR barcode]; ♂, NW, Tsaratanana, Antetykalambazaha, near camp, 14.1958 o S, 48.9497 o E +/- 3.5 km, 1628 +/- 100 m, 20 / 12 / 2004 20: 17, R. Ranaivosolo: DL-05- 763, BMNH (E) # 676586 [DNA voucher; cytochrome b], BMAD 223 - 15 [DNA barcode voucher]; B 43 [‘andromap’]. ♂ (Fig. 18 B, left), NW, Tsaratanana, [Ampidiranala], 14.1958 o S, 48.9497 o E +/- 2.5 km, 1628 +/- 50 m, 20 / 12 / 2004, R. Ranaivosolo: DL-05-761, 2384 [= DL 2384; DNA extract; cytochrome b] MSL 058:H02 [genomic DNA; abdomen lysed] BMNH (E) # 676584; ♂, NW, Tsaratanana, Matsiborimaika, 2035 m, 14.1526 o S, 48.9578 o E +/- 0.015 km, 2035 +/- 25 m, 22 / 12 / 2004, D.C. Lees: DL-05- 781, SM006, NHMUK 010289185 [QTR barcode]; ♂, NW, Tsaratanana, piste Mangindrano-Antetykalambazaha, c. 1550 m, 14.199 o S, 48.952 o E +/- 0.5 km, 1550 +/- 65 m, 20 / 12 / 2004, D.C. Lees: DL-05- 747, TF 13 [leg for DNA], NHMUK 010289151 [QTR barcode]; ♂, NW, Tsaratanana, Ampidiranala, 14.1958 o S, 48.9497 o E +/- 0.5 km, 1628 +/- 50 m, 20 / 12 / 2004, R. Ranaivosolo: DL-05- 758, BMNH (E) # 676582 [DNA voucher; cytochrome b]; ♂, NW, Tsaratanana, [Ampidiranala], 14.1958 o S, 48.9497 o E +/- 3 km, 1628 +/- 75 m, 20 / 12 / 2004, R. Ranaivosolo: DL-05- 762, BMNH (E) # 676585 [DNA voucher; cytochrome b]; ♂, NW, Tsaratanana, piste Mangindrano- Antetykalambazaha, just into primary forest, 14.2047 o S, 48.9538 o E +/- 0.35 km, 1456 +/- 25 m, just into primary forest, 20 / 12 / 2004: 20: 15, D.C. Lees: DL-05- 734 A, TF 17 [leg for DNA], NHMUK 010289152 [QTR barcode]; ♂, NW, RNI Tsaratanana, route Mangindrano-Andohanisambinano, 19 / 12 / 2004, D.C. Lees: DL-05- 731, BMAD 262 - 15, CCDB-02230-F 12 [DNA barcode voucher]; Deposition MNHN: ♂ [MNHN], Madagascar Nord, TSARATANANA. 1,800 m (Forêt de Mousses), [14.17 o S, 48.94 o E +/- 4 km, 1800 m] 10 / 1949, R. Paulian: DCL-DB- 3301; Deposition ABRI: ♂ [ABRI], Madagascar NW, Tsaratanana Mntn N. madagascar below Andohanisambirano, S 14 o 12.486 ' E 0 48 o 58.225 ' 4167 ft, [14.2081 o S, 48.9704 o E +/- 1 km, 1291 +/- 50 m], 24 / 10 / 2003, S. C. Collins: 1204 [leg for DNA]; Deposition PBZT: 2 ♂♂ [PBZT], Madagascar NW, Mt Tsaratanana, 750 a 1400 m, [14.21 o S, 48.96 o E +/- 5 km, 1400 m.; elevation as low as 750 m not found in region] 2 / 1951, R. Paulian. Deposition summary: BMNH (HT ♂, 15 PT ♂♂, 1 PT ♀), MNHN (1 PT ♂); ABRI (1 PT ♂); PBZT (2 PT ♂♂). Type locality. Madagascar NW, Tsaratanana, near forest margin, 14.2034 o S, 48.9532 o E +/- 2.5 km, 1441 +/- 25 m. Diagnosis. The species resembles most closely Ht. ankova from which there are no obvious ♂ genitalic differences. Ht. turbata and Ht. pallida can be told apart by a more irregular HWV Mb. The most obvious difference from Ht. ankova is the darker, much more sombre diffuse dark grey colouration, especially on the underside. Description. Wings: upperside uniform dark brown, with FW space-CuA 1 ocellus with a wide orange ring which is generally quite concentric but apparently a little wider and more angled proximad. FWD space-M 1 ocellus small and slightly elliptic with very narrow orange ring. HWD space-CuA 1 ocellus is the main one expressed there, as well as to lesser extents those of space-M 3 and space-M 2, as in Ht. ankova. Hairbrush around space- 1 A region lacking. Underside dark greyish brown, darker than typical of Ht. ankova, with fine brown and ochreous irroration, especially in basal parts of wings. FWV space-CuA 1 ocellus with large orange ring yellowing slightly proximad following the reddish brown concave curve of the Mb before it bends back indistinctly to mid-costa, terminating as if pinched at the base of vein M 3 (a feature also displayed by Ht. ankova). FWV space-M 1 ocellus rather small and slightly elliptic with a white pupil with narrow black iris and with a narrow pale orange ring. On HWV, space- CuA 1 ocellus quite small and slightly elliptic with a narrow black iris also and pale orange concentric ring. In HT, no other ocelli expressed on HWV. HWV Mb reddish brown, slightly broader than in Ht. ankova and fairly straight, but finely irregular, with a yellow Mf distad of it in lower basal part of space-M 2. A reddish PMb mirrors shape of Mb at least as far as vein 1 A. HWV area distad of Mb lighter due to a higher concentration of ochreous scales than the basal area, but as in Ht. ankova, more fuscous brown scales predominate in the marginal area from veins Rs to M 3 particularly, giving the wing margin a slightly ‘singed’ appearance. FWV cell area with four brown rather equally spaced transverse Cbs delineating spaces highlighted by ochreous-yellow hair-scales. Variation. ♂♂ vary quite a bit in size and wing patterning. HWV ocelli in space M 1 and M 2 often visible as white points. Sexes similar. Wingspan/fwl: range 29.7–35.7 / 16.1–18.2 mm (n= 5 ♂♂), including HT ♂: 35.2 / 18.1 mm. Mean = 32.3 +/- 2.01 SD/ 17.1 +/- 0.95 mm (n= 5 ♂♂). 33 / 18.7 mm (n= 1 ♀). Androconia: Sdb dark reddish brown with grey tip. Androconial patch directly above the end of the R stem small, blackish and lenticular. Veins 1 A+ 2 A narrowly inflated for most of length, that in 3 A slightly more so (more so than in Ht. ankova) and covered in narrow greyish brown scales. No other androconia obvious. Palps: penultimate segment with distinct brown medial strip, flanked by yellowish scales and fringed by dark brown scales except where wiping compound eye, with yellow scales mainly on mesad face. ♂ genitalia: DL-05- 761, PT (Fig. 18 B, left); DL 05- 845, PT (Fig. 18 B, right). Tegumen with fairly rounded proximad profile (LV) and rather flat dorsal profile (LV) and very slight brow leading to uncus which is about 1 / 3 longer than the dorsal edge of the tegumen whose DV features a large proximad rather ‘v’-shaped notch, and an uncus whose depth (LV) increases slightly towards midpoint and features a distinct ‘head’ and a small toothed down-hook at tip, while it is slightly inflated medially (DV). The gnathos emerges from a right-angled base and is widest at its attachment and smoothly downrecurved until not quite in line with tegumen-uncus (LV) and is sinuate and slightly outsplayed (DV), with pointed tips slightly incurved. The valve has a small angled dorsal ‘shoulder’ (LV) and is roughly broad-triangular featuring a stubby and uprecurved arm leading to a small club covered with spinoid setae and with inpointing spine from DV. Saccus bulbous, widening proximad (LV), rather long; aedeagus quite deep and relatively straight with about 3–5 small lateral teeth post-medially (also found in Ht. ankova whose male genitalia are not noticeably different). Etymology. Refers to the practical consideration in finding this species that a trip to Tsaratanana may involve such a long walk with porters that the rice (‘vary’) risks being exhausted (‘lany’) well before the end of the expedition. Discussion. Sparse historical material of Ht. lanyvary collected by R. Paulian at the type locality existed in MNHN (♂ collected in 1949) and in PBZT (two ♂♂ collected in 1951); these specimens are included in the type series, but the species was discovered again recently in the field (in December 2004) at the type locality. All available types in the Ht. strigula group were examined (see also under Ht. roussettae, and elsewhere below). The most relevant is the HT ♂ of Mycalesis ankova Ward, 1870 (BMNH (E) 673767) (Fig. 19 C), lacking locality, from the Chris Ward collection, which differs in ventral wing colouration from all known samples of Ht. lanyvary. See also Fig. 19 and under Ht. tornado regarding type material for Pseudonympha turbata and Culapa pallida (Fig. 19 D) and Culapa ornata (Fig. 19 E). Additional information. DNA divergences: COI- 5 P barcode cluster number BOLD:ACW 4935, about 3.2 % divergent to Ht. ankova (cluster number BOLD:AAK 5841, exemplar BMNH (E) 676678, BMAD 048-09, HM 404225 from Analamazaotra). Phylogeny/sister species: the species was not analysed in Lees 1997 (see though p. 62, where “sp. 67 ” was considered the same species as Ht. ankova; nor was it treated in any prior molecular works but Ht. lanyvary is very likely to be the (at Tsaratanana, sympatric) sister species of Ht. ankova, as found by Aduse-Poku et al., (2016, in press). Ecology and distribution. Habitat: montane rainforest, forest interior, close to margins. Behaviour: flies low. Hostplant: unknown, possibly low grasses. Early stages: unknown. Distribution: endemic as far as is known, to the Tsaratanana massif (Fig. 30 C, olive dots). Elevational range: 1290–1800 m. (n= 17 including referred specimens and observations). Referred specimens. ♂, Madagascar N, Tsaratanana, 14.2034 o S, 48.9532 o E, 1441 m, 24 / 12 / 2004, D.C.Lees: DL-05- 859, IA 624 [isotope voucher]; ♂, NW, Tsaratanana, 14.2034 o S, 48.9532 o E +/- 2.5 km, 1441 +/- 25 m, 12 / 2004; BMNH (E) # 671598 [DNA voucher; cytochrome b], KA 504 [=KA-P 504; DNA extract number]; ♂, NW, Tsaratanana, Ampidiranala, 14.20025 o S, 48.95175 o E +/- 0.54 km, 1542 +/- 85 m, 20 / 12 / 2004, R. Ranaivosolo: DL-05- 756, BMNH (E) # 676686 [DNA voucher; cytochrome b].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 63-66, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis vertigo Lees & Raharitsimba, sp. nov.
No full text<i>Heteropsis vertigo</i> Lees & Raharitsimba, sp. nov. <p>LSID: urn:lsid:zoobank.org:act:38AC77EE-A10A-4DCA-9444-F55C845FFA23</p> <p> <b>Prior references</b>: sp. 61 (on envelopes and/or datalabels but not mentioned in the list of Lees, 1997: 64–65).</p> <p> <b>Type material., Deposition BMNH: Holotype:</b> ♂ (Fig. 28 A), Madagascar NW, Bekolosy Mt., RS Manongarivo, c. 1163 m, 14.04575o S, 48.29623o E +/- 0.5 km, 1130 +/- 35 m, 16/12/1994, soleil/lo [fruit trap], H. Raharitsimba: DLBEK 94_77, KA2049 [=KA-P2049; extract number], NHMUK 010289173 [QTR barcode].</p> <p> <b>Paratypes</b>: <b>Deposition BMNH</b>: ♂, data as HT but 12–22/12/1994, D.C. Lees: DLBEK 94_83, IA354 [isotope voucher]; BMAD 241-15 [DNA barcoded specimen], NHMUK 010289174 [QTR barcode]; ♀ (Fig. 28 B), Madagascar E, Bekolosy Mt., RS Manongarivo, Madagascar NW, 1310 m, 14.0385o S, 48.3073o E +/- 0.55 km, 1310 +/- 50 m, 21/12/1994, lo lt 1310 [fruit trap], D.C. Lees: DLBEK 94_129, IA 356 [isotope voucher], NHMUK 010289175 [QTR barcode]; ♂ (Fig. 29 A, genitalia), Madagascar NW, Bekolosy, RS Manongarivo, ~ 1200 m, 14.0436o S, 48.299o E +/- 0.43 km, 120 m +/- 50 m, 13/12/1994, D.C. Lees: DLBEK 94_159, DCLW-0104 [wing prep.], KA3035 [=KA-P3035, DNA extract number]; IA355 [isotope voucher]; BMAD 240-15 [DNA barcoded specimen] 366 DL [genitalia], NHMUK 010289176 [QTR barcode]; ♀, NW, Bekolosy Mt., RS Manongarivo, Madagascar NW,> 1250 m, 14.0427o S, 48.3028o E +/- 0.5 km, 125 m +/- 50 m, 19/12/1994, D.C. Lees: DLBEK 94_110, 1035 [= DL 1035; DNA extract number], BMNH (E) #672400 [DNA voucher; cytochrome b], 372 DL [♀ genitalia]; ‘leg for KAP’; IA357 [isotope voucher]; ♀, NW, Bekolosy Mt., RS Manongarivo, 1200– 1250 m, 14.04334o S, 48.3013o E +/- 1 km, 1225 +/- 50 m, 12/12/1994, D.C. Lees: DLBEK 94_23, Egg exp. [egg expression voucher], IA353 [isotope voucher], NHMUK 010289177 [QTR barcode]; ♀, NW, Bekolosy Mt., RS Manongarivo, c. 1245 m, 14.0427o S, 48.3028o E +/- 0.5 km, 1250 m, +/- 50 m, 12/12/1994, 10:40, D.C. Lees: DLBEK 94_22 [Egg voucher]; IA352 [isotope voucher], KA2050 [=KA-P2050; extract number], NHMUK 010289178 [QTR barcode].</p> <p> <b>Deposition summary</b>: BMNH (HT ♂, 2 PT ♂♂, 4 PT ♀♀).</p> <p> <b>Type locality.</b> Bekolosy Mt., RS Manongarivo, c. 1163 m, 14.0458o S, 48.2962o E +/- 0.5 km, 1130 +/- 35 m.</p> <p> <b>Diagnosis.</b> In its translucence, the extended FWD space-CuA1 Orng of <i>Ht. vertigo</i> is intermediate between <i>Ht. turbans</i> and <i>Ht. sabas</i>. <i>Ht. vertigo</i> in wing pattern resembles <i>Ht. turbans</i> more than it does <i>Ht. sabas</i>, but the FWD space-CuA1 Orng is largely suffused with orange and only pale whitish-translucent in the anteriad half, especially proximad, rather than throughout it, as in <i>Ht. sabas</i>, or largely orange as in <i>Ht. turbans. Ht. vertigo</i> lacks the extreme ear-like cupping of the gnathos seen in <i>Ht. sabas</i> (Lees, 1997: 107) and <i>Ht. turbans</i> (Fig. 29 B; NW Zahamena) as well as some related species (e.g. <i>Ht. angulifascia, Ht. borgo</i>), and the HWV Mb is much less kinked than in any of these species.</p> <p> <b>Description. Wings</b>: Similar to <i>Heteropsis turbans</i>. Upperside uniform mid brown, except with conspicuous orange Mf/crescent (as in a few other of the <i>Ht. strigula</i> -group, notably <i>Ht. strigula</i>) at base of HWD space-M2 following curve at base of vein M3. FW space-CuA1 ocellus with orange to pale yellow/translucent ‘ring,’ more like a hexagon that has been compressed along a diagonal running from mid-costa to tergal angle. FWD space-M1 ocellus quite small and elliptic with narrow orange ring. HWD space-CuA1 ocellus the only one expressed there and slightly elliptic, with narrow orange ring. Darkish brown, diffuse, slightly wavy, uprecurved hair brush emanates mainly from space-1A as far as mid-vein. Underside greyish brown, similar to <i>Ht. turbans</i> but with a less kinked HWV Mb, and limited irroration, especially in basal area. Space-CuA1 ocellus with orange ring yellowing proximad following reddish brown concave curve of Mb before it bends back to mid-costa. Space-M1 ocellus fairly elliptic, with white pupil and wide narrow black iris, but narrow orange ring. HWV space-CuA1 ocellus large and round to elliptic with narrow pale orange ring. HWV space-Rs-M2 ocelli generally not expressed and space- CuA2 ocellus small and elliptic with faint orange ring. HWV Mb reddish brown and fairly straight but outflexed around M2, generally with a yellow patch (Mf) distad of it in space-M2, and curved back towards the anal angle in space-CuA2. Slight, wavy brown Pml tracks the wing margin and Sml more closely follows the fairly crenate (notably much less so than in the related <i>Ht. sabas</i>) HW margin. FWV cell area with four darker brown rather equally spaced transverse Cbs, the outermost of which flank area that is paler than for the inner pair. Basal areas not much darker than areas distad of the Mb, and darker brown PMb weakly represented. <b>Variation.</b> ♂♂ similar to each other, but seasonal variation is not known, as the entire type series was caught near the start of the rainy season in mid December. ♀♀ similar, but dorsally lighter and larger with more rounded wings.</p> <p> <b>Wingspan/fwl</b>: range 32.2–35.6/17.3–19.6 (n=2 ♂♂), including HT ♂ 35.2/ 19.6 mm. Range 35.3–36.3/ 19.4– 19.5 mm (n=2 ♀♀).</p> <p> <b>Androconia</b>: 1A+2A has a narrowly tapered inflation and 3A is much more swollen distad, from base to midvein, covered in thin grey-brown scales. As for <i>Ht. turbans</i> and <i>Ht. sabas</i>, along their length HW veins M2, M3, CuA1 and CuA2 are also narrowly inflated and have specialized scales on the dorsal surface. Abdominal black androconia just visible ventro-laterally around A4–A5. HWD Sdb dark brown, Sdp grey, small, lenticular, composed of narrow grey scales.</p> <p> <b>Palps</b>: ochreous brown with darker brown thin medial stripe and with dark brown border away from compound eye and with dark brown scales above where potentially brushing eye, sandwiching ochreous scales away from eye.</p> <p> <b>♂ genitalia</b>: 366DL, PT (Fig. 29 A): from LV, uncus fairly straight in line with tegumen in non-deflexed position and recurved down to a sharply hooked point, with a distinct ‘head’ before tip; tegumen about same length as uncus and constriction at fusion line with vinculum only moderate, such that tegumen from LV tapers only slightly from a rectangular frame; tegumen features wide and shallow proximad notch from DV. Gnathos on angled base, flattened distinctly before tip into broadened structure (not nearly as earlike as for example in <i>Ht. turbans</i> and <i>Ht. angulifascia</i>), with drawn out, tapered and pointed tip, most noticeable in DV. Valves fairly stout and uprecurved arms fairly short leading to a ‘bird’s head’ (rather ‘cassowary head’-like, from DV) tip with spinoid setae and with inwardly projecting ‘beak’. Saccus quite short and inflated towards proximad tip and aedeagus about 1/5 longer than valve. Juxta protruding considerably proximad.</p> <p> <b>Etymology.</b> A direct reference to the vertiginous waterfall directly below the camp at Bekolosy Mt. (14.04514°S, 48.29616°E, 1130 m, from Google Earth) that marks the lower elevational limit of this species. Returning after a storm at the summit, I narrowly avoided falling into this cascade, when jumping between two rocks, my net bag caught the strength of the current, and it was thanks to the alertness of the camp assistant, who rushed over, and holding on to my leg, ensured I reached the rock on other side (netless, however). The net was even found in the morning among the rocks at the top of the fall. Alludes also to the marvellous Hitchcock film of this name, with its subliminal spiralling motif.</p> <p> <b>Discussion.</b> No historical museum collections of <i>Ht. vertigo</i> are known. When first found in the field in December 1994, the species was given a morphospecies number (sp. 61) but was nevertheless thought to represent <i>Ht. turbans</i> (indeed Lees 1997: 225 Fig. 3 K erroneously shows the distribution of that species extending to the northwest of Madagascar). <i>Ht. vertigo</i> has since been found on the adjacent mountain of Antsatrotro in November 2011. The species is confusable with a number of described taxa. Examined were the STs of <i>Culapa turbans</i> Oberthür 1916 (LT ♂, here designated, Fig. 28 C, specimen bearing labels “ Madagascar Antsianaka Perrot Freres 2e Semestre 1893| Culapa turbans Obthr ♂ type |P.E.L. Viette det. 1968 Culapa turbans Ch. Obthr. LECTOTYPE |supposed “ type ” of Culapa turbans Oberthür |[copy of Oberthür 1916, Pl. 356, f. 3050]|BMNH(E) #674780”) and of <i>Culapa curvatula</i> Oberthür, 1916 (LT, here designated, Fig. 28 D, specimen bearing labels: “Antsianaka et lac Alaotra 2e Trimestre 1889 Perrot Freres|[copy of Oberthür 1916, Pl. 356, f. 3057]|BMNH(E) #674784; automatically then, PLT ♂: BMNH(E) #674786; PLT ♀♀: BMNH(E) #674785) and BMNH(E) #674787, with same locality data). <i>Ht. curvatula</i> was considered a synonym of <i>Ht. turbans</i> by Lees <i>et al.,</i> 2003 (see also below). Also examined (Fig. 28 E) was the HT ♂ of <i>Culapa sabas</i> Oberthür, 1923 (‘Tamatave’|BMNH(E) #674789), in addition to extensive field material of <i>Ht. turbans</i> and <i>Ht. sabas</i> across their ranges. As well as being allopatric, the species is clearly separated from types of both species by a consistently differing FWD ocellus colour pattern and in field collected material of those species, by divergent ♂ genitalia.</p> <p> <b>Additional information. DNA divergences:</b> COI-5P barcode cluster number BOLD:ACD8579. Bekolosy and Antsatrotro specimens have an identical haplotype; the mountains are not far apart. The COI-5P barcode is about 1.06% divergent to <i>Ht. turbans</i> and about 1.7% divergent to <i>Ht. sabas</i> with six apparently fixed nucleotide differences to <i>Ht. turbans</i>; all three species share the same cluster number. Sister to <i>H. sabas</i> in Aduse-Poku <i>et al.,</i> (2016).</p> <p> <b>Phylogeny/sister species</b>: sister species unclear, but closely related both to <i>Ht. sabas</i> and <i>Ht. turbans</i>, according to the COI-5P barcode. A ♀ of the ‘ <i>curvatula</i> ’ phenotype (DL06-1099, BMAD259-15) from NW Zahamena was one nucleotide different (COI-5P) to other barcoded members of <i>H. turbans</i> (cluster number BOLD:ACD8579), and so the above synonymy appears correct.</p> <p> <b>Ecology and distribution.</b></p> <p> <b>Habitat</b>: primary montane rainforest.</p> <p> <b>Behaviour</b>: one was caught in a low hung fruit trap at the waterfall camp.</p> <p> <b>Hostplant</b>: unknown.</p> <p> <b>Early stages</b>: unknown.</p> <p> <b>Distribution</b>: as far as is known endemic to Réserve Speciale de Manongarivo, Mts. Bekolosy and Antsatrotro (Fig. 30 B, fawn dots).</p> <p> <b>Elevational range</b>: 1130–1410 m. (n=45; including referred specimens and observational data, such as fruit trap data).</p> <p> <b>Referred specimens.</b> ♂, Madagascar NW, RS Manongarivo, Bekolosy, 1215 m, 14.034o S, 48.31o E +/- 1 km, 1215 +/- 50 m, 18/12/1994, 09:30, D.C. Lees: DLBEK 94_165, 46; ♂, NW, Antsatrotro Mt., RS Manongarivo, [1250] m, near camp, 14.0824o S, 48.3663o E +/- 0.35 km, 1240 +/- 25 m, 2/12/2011, 14:51, D.C. Lees: DL 11A- 0001/ BMAD 245-15 [DNA barcoded specimen], IA609 [isotope voucher]; ♀, data as above but: 3/12/2011, D.C. Lees: DL 11A-0002/ BMAD 244-15 [DNA barcoded specimen].</p>Published as part of <i>C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1)</i> on pages 89-92, DOI: 10.11646/zootaxa.4118.1.1, <a href="http://zenodo.org/record/264597">http://zenodo.org/record/264597</a>
Heteropsis borgo Lees, sp. nov.
No full textHeteropsis borgo Lees, sp. nov. LSID: urn:lsid:zoobank.org:act: 45 DC 90 D 0-4167 - 4047 - 89 DC-A 9 C 45 D 523 E 49 Prior references: sp. 73 (Lees 1997; Torres et al., 2001: 462). Type material., Deposition BMNH: Holotype: ♂ (Fig. 27 A), Anjanaharibe Sud, summit Anjavidibe, 14.7396 °S, 49.4605 °E +/- 0.25 m, 1543 +/- 15 m, waypoint 136. Fruit trap 1. 23 / 11 / 2014: 10: 36. Ahamadi Allaoui. DL 14 A-0350, IA 522 [isotope voucher], NHMUK 010289167 [QTR barcode]. Paratypes: Deposition BMNH: ♂, data as HT but 23 / 11 / 2014: 12: 21. D.C. Lees. DL 14 A-0361, BMAD 221 - 15 [DNA barcode], IA 519 [isotope voucher], NHMUK 010289168 [QTR barcode]; ♂, Anjanaharibe Sud, summit Anjavidibe, 14.7396 °S, 49.4605 °E +/- 0.25 m, 1543 +/- 15 m. 23 / 11 / 2014: 13:00, Ahamadi Allaoui: DL 14 A-0363, IA 521 [isotope voucher], BMAD 235 - 15 [DNA barcode], NHMUK 010289169 [QTR barcode]; ♂, NE, Anjanaharibe Sud, ANJAHA- 6 [GPS], 14.7482 o S, 49.4659 o E +/- 0.48 km, 1320 +/- 60 m, 6 / 2 / 1996, C. Kremen: CK 724, rainforest on slope, trap 14, IA 358 [isotope voucher], NHMUK 010289170 [QTR barcode]; ♂, NE, Anjanaharibe Sud, 1530 m, elfin forest just before summit, ANJAHA- 7 [GPS], 14.7426 o S, 49.4608 o E +/- 0.37 km, 1450 m +/- 7 m, 4 / 2 / 1996: 11: 49, H. Raharitsimba: TR 553 A, IA 359 [isotope voucher], NHMUK 010289171 [QTR barcode]; ♂ [left wings and head], Madagascar NE, Anjanaharibe Sud, ANJAHA- 7 [GPS], 14.7406 o S, 49.4505 o E +/- 0.96 km, 1455 +/- 145 m, 12 / 2 / 1996: 08: 59, C. Kremen: CK 729; DL 9659 [DNA voucher], IA 360 [isotope voucher], NHMUK 010289172 [QTR barcode]; Deposition MNHN: ♂, data as HT but 21 / 11 / 2014: 12:02, Ahamadi Allaoui: DL 14 A-0308, IA 520 [isotope voucher]; ♂, data as HT but 23 / 11 / 2014. Ahamadi Allaoui. DL 14 A-0192, 116 [leg in ethanol], IA 626 [isotope voucher]. Deposition summary: BMNH (HT ♂, 5 PT ♂♂); MNHN (2 PT ♂♂). Type locality. Anjanaharibe Sud, summit Anjavidibe, 14.7396 °S, 49.4605 °E +/- 0.25 m, 1543 +/- 15 m. Diagnosis. Ht. angulifascia is particularly similar, but there are no other closely comparable species. HWV Mb, however, much less indented that in Ht. angulifascia, scribing a lobe outward between M 2 and M 3, running for only a short rather than a long distance just below M 2. Description. Wings: upperside uniform dark brown. FWD space-CuA 1 ocellus fairly large, occupying more than the span of this space taking into account the concentric deep orange ring. FWD space-M 1 ocellus small and subelliptic with small white pupil and wider black iris, with only a faint trace of Orng. HWD space-CuA 1 ocellus moderately large, considerably smaller than that in FWD and elliptic, with deep orange ring spanning more than half of the space between veins. No other ocelli expressed nor is there an orange Mf on HWD. Underside dark greyish brown. Darker grey-brown irroration not very noticeable even in the basal area where more pronounced, due to sombre colouration. FWV space-CuA 1 ocellus very slightly larger than on FWD, its black iris spanning the veins CuA 1 and CuA 2, with fairly wide, more or less concentric, deep orange ring. FWV space-M 1 ocellus very small and only slightly elliptic with white pupil and black iris, but very narrow pale orange ring. Small white point in M 2 just under this ocellus. HWV space-Rs ocellus as distinct white spot without obvious black ring. FWV space-M 1 -M 2 ocelli as minute white spots and ocellus in space-CuA 2 not expressed. HWV space-CuA 1 ocellus roughly circular, small, similar in size to HWD, not nearly spanning space, with faint concentric pale orange ring. HWV Mb irregular, reddish brown and most outflexed (convex) around the base of space-M 2 -M 3, bent distad to the base of space-M 2, concave distad of space-M 1 and space-CuA 1, and slightly concave near intersection with vein CuA 2. Weakly expressed lighter brown highlight (flare) distad of Mb. PMb also reddish brown and quite distinct, somewhat irregular, more or less parallel to Mb, gently convex. Four transverse darker bands in FWV cell (Cbs) forming two quite diffuse slightly lighter grey areas and diverging costad between the last two Cbs. Slightly marked, wavy, darker brown Sml more obvious in the HW, with darker finer line closely following the margin, which is only slightly crenate in HW. Basal area not much darker than distad portions of wing, while band between the PMb and Mb slightly more shaded, especially distad. Variation. ♂♂ very similar to each other, without marked variation in eyespot size between the two collection periods (third week of November, late dry season, and second week of February, wet season). The HWV Mb varies in the angularity with which it is kinked out around space- M 2 –M 3, but this ‘kinking’ is not as extreme as in Ht. angulifascia. ♀ unknown, but likely to be similar to the ♂, as for the closely related Ht. angulifascia. Wingspan/fwl: range 34.3–36.7 mm./ 17.6–19.3 mm; mean = 35.5 +/- 0.7 SD/ 18.9 +/- 0.5 SD mm (n= 7 ♂♂), including HT 36.7 / 19.3 mm. ♀ unknown. Androconia: Sdp small, oval and black. Darkish brown hair-brush emanating mainly from above 1 A+ 2 A near the base and from space- 1 A as far as mid-vein. Blackish androconial scales appear to be present ventro-laterally around abdominal A 4, but are indistinct in the material available. Palps: outfacing surface covered in ochreous grey scales, with thin dark blackish brown medial stripe and dark blackish brown scales composing borders, sandwiching ochreous grey scales away from compound eyes. ♂ genitalia: 284 DL, PT (Fig. 27 D): from LV, uncus slightly upbent before middle and recurved down to a sharply hooked point, without a distinct ‘head’ before tip (slightly inflated before tip from DV); tegumen shorter than uncus and constriction at fusion line with vinculum not as pronounced as in some other species. Gnathos flattened distinctly into an ear-like shape (as for, e.g., Ht. angulifascia) and serrate at end/ventrally with a distinct thin projection at dorsal edge, most obviously inrecurved and pointed from DV. Valves fairly stout and uprecurved arms leading to bilobed tip (crest) covered in spinoid setae and with inwardly projecting ‘beak’ at proximad lobe; ostium quite long. Saccus not particularly long and aedeagus shallowly up-reflexed, about 1 / 5 longer than valve. Juxta prominent proximad, with downcurved lip. Etymology. A very dark and still rather mysterious species of mountain passes, in this case the one on the way to the summit of Anjanaharibe Sud, as in the Borgó pass made famous in F.W. Murnau’s film classic ‘Nosferatu’ and its compelling remake by Werner Herzog. Discussion. No historical material has been found in museums. This species was first found in the field as morphospecies 73 in C. Kremen and H. Raharitsimba’s survey of Anjanaharibe Sud which included the summit of ‘Anjavidibe’ in February, 1996 (not in November, 1995 as stated in Lees, 1997: 65). The species was refound (around the type locality, only) in November, 2014 by myself and A. Allaoui. The HT ♂ in BMNH of Pseudonympha angulifascia Butler, 1879 (‘Antananarivo’, Kingdon, BMNH (E) 674769) was examined (Fig. 27 C) as well as the ♂ and ♀ STs of its synonym Mycalesis butleri Mabille, 1880 in BMNH (LT ♂, here designated, Fig. 27 B, bearing data: “ Lectotype / Type /Madgscr/Ex Grose-Smith 1910 / Henotesia butleri Mab. ♂ H.T. selected by G.T. Nov. 1930 | BMNH (E) 674771 ”); these two specimens clearly belong to Ht. angulifascia. Additional information. DNA divergences: COI- 5 P barcode cluster no. BOLD:AAK 5842 (exemplars BMAD 221 - 15 DL 14 A-0361, BMAD 235 - 15 DL 14 A-0363) is shared with, although 1.67 % divergent to, Ht. angulifascia (exemplar BMAD 049-09, CCDB-02225-E01 from Ranomafana) with seven potentially fixed nucleotide differences. In their dataset for the COII marker (Torres et al., 2001), there was an about 2.15 % divergence (8 bp different in 415) for a single sample of Ht. borgo (as “ Hen. sp. 73 ”, 9659 S compared with Ht. angulifascia samples ANG 8 M, ANG 7 M, ANGCK and ANG 11). Phylogeny/sister species: Torres et al., (2001: 466) found 94 % support for a sister relation with Ht. angulifascia in their cladistic analysis of COII sequences, as fully supported by Aduse-Poku et al., (2016, in press). Closely related to Ht. angulifascia, for which this species probably constitutes a northern replacement and allospecies. Ecology and distribution. Habitat: riparian primary rainforest, and rainforest on ridges and slopes. Behaviour: attracted to fruit bait. Flies fairly close to the ground and ♂♂ frequent small sunspots near the known summit in the understory of elfin montane forest. Hostplant: unknown, possibly a small leaved grass that was found downslope of the type locality. Early stages: unknown. Distribution: endemic to rainforest at Anjanaharibe Sud, as far as is known (Fig. 30 C, dark green dot). Conservation: in November 2014 ♂♂ of this species were only seen at the actual summit of Anjavidibe, whereas in the 1996 survey, C. Kremen found them over 220 m. downslope, and it is not known if the species occurs on the adjacent real summit of Anjanaharibe Sud. It is likely then the species could be vulnerable to changes due to global warming. The main local threat to this area is the quarrying of quartzite seams. Elevational range: 1320–1543 m. (n= 8). Referred specimens. ♂ (Fig. 27 D, male genitalia, rest of voucher lost), Madagascar NE, Anjanaharibe Sud, 1520 m, ANJAHA- 8 [GPS point] 14.74265 o S, 49.46065 o E +/- 0.39 km, 1520 +/- 100 m, 2 / 2 / 1996, C. Kremen: CK 712; 284 DL [♂ genitalia], 470 [= DL 0470; DNA extract number], BMNH (E) # 697884, KA 580 [=KA-P 580; DNA extract number].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 86-88, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
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