Heteropsis harveyi Lees & Kremen

C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, Zootaxa 4118 (1), pp. 1-97 : 28-31

Heteropsis harveyi Lees & Kremen , sp. nov.

LSID: urn:lsid:zoobank.org:act:CD7A88FE-87A6-4324-8404-5EDE47D8E9F2

Prior references: Mycalesis andravahana Mabille ♂ “var.” ( Mabille [1885]: Pl. 5 Fig. 8 , [1887: 57–59]);

Henotesia andravahana ab. marmorata Aurivillius, 1911 (unavailable name; holotype examined in Stockholm and conspecific); Gaede 1931: 401; Ackery et al., 1995: 297 (297 [raised from synonymy with Ht. anganavo (Ward, 1871) ]; the name ‘ marmorata ’ was incorrectly linked to “sp. 13” in Lees, 1997: 64;

Henotesia Wardii / Henotesia Andravahana ( Oberthür, 1916: 205) . Charles Oberthür labelled his specimens in his original drawer ambiguously as “ wardii - andravahana ” (see Oberthür, 1916: 204–205; Lees, 1997: 64). He stated ( Oberthür 1916: 205) that he had in his collection “22 ♂♂ et 10 ♀♀ d' Andravahana provenant de Fianarantsoa et d'Antsianaka” (subsequently known as Heteropsis viettei Lees, 2003 ; Lees, 1997: 62; 391–392, but here treated as Heteropsis andravahana ), “et 78 ♂♂ et 45 ♀♀ du supposé Wardii , capturés également à Fianarantsoa et dans l’Antsianaka"; referred to here as Ht. harveyi ; All 78 ♂♂ and 13 of the ♀♀ of Oberthür’s “ wardii ” (most of the latter from Antsianaka) have been located (in BMNH). An additional four ♀♀ placed above the " wardii ♀” label in the Oberthür collection were not referable to this species ( Lees, 1997: 391);

sp. 12 ( Kremen, 1994: 417, as “ Henotesia ? obscura ”; Lees 1997: 61, 64; Torres et al., 2001: 462);

“ Heteropsis (Henotesia) harveyi Lees (MS), nom.nov.” (ms name in Appendix A of Lees, 1997: 391).

Type material., Deposition BMNH: Holotype: ♂ ( Fig. 7 A), Madagascar E, Sahavondronina VOI, Ranomafana National Park, 21.276o S, 47.325o E +/- 0.25 km, 1313 +/- 25 m, 8/11/2014, D.C. Lees: DL 14R-229, NHMUK 010289119 [ QTR barcode].

Paratypes: Deposition BMNH: ♂, same data as HT but 8/11/2014: 10:40, A. Allaoui: DL 14R-204, NHMUK 010289120 [ QTR barcode]; ♀ ( Fig. 7 B), 8/11/2014: 10:57, D.C. Lees: DL 14R-207, NHMUK 010289121 [ QTR barcode]; ♂, same data as HT but 6/11/2014: 13:03, fruit trap, D.C. Lees: DL 14R-126, IA599 [isotope analysis], NHMUK 010289122 [ QTR barcode]; ♂, same data as HT but 8/11/2014, A. Allaoui: DL 14R-104, NHMUK 010289123 [ QTR barcode]; ♀, same data as HT but 8/11/2014, A. Allaoui: DL 14R-206, BMAD 217-15 [ DNA barcode], IA597 [isotope voucher], NHMUK 010289180 [ QTR barcode];

Deposition MNHN: ♂, same data as HT but 6/11/2014, A. Allaoui: DL14R-116; ♀, same data as HT but 7/11/ 2014, D.C. Lees: DL14R-156, E37; ♂, 25-V-69 FIANAR: SENTIER FORESTIER A 3 KM AU S.O. DE, VOHIPARARA, [21.24o S, 47.43o E +/- 1km, 1029 +/- 50 m], J. Minet; ♀, same data as above but: 9-III-69;

Deposition ABRI: ♂, same data as HT but 8/11/2014, D.C. Lees: DL14R-245; ♀, same data at HT but 8/11/ 2014: 10:48, A. Allaoui: DL14R-191;

Deposition PBZT: ♂, same data as HT but 8/11/2014: 11:27, A. Allaoui: DL14R-213.

Deposition summary: BMNH (HT ♂, 3 PT ♂♂ and 2 PT ♀♀), MNHN (2 PT ♂♂ and 2 PT ♀♀); ABRI (1 PT ♂ and 1 PT ♀); PBZT (1 PT ♂).

Type locality. Madagascar E, Sahavondronina VOI, Ranomafana National Park, 21.276o S, 47.325o E +/- 0.25 km, 1313 +/- 25m.

Diagnosis. Ht. vanewrighti and an undescribed species (‘ Ht. sp. 20’) are both generally smaller than Ht. harveyi and the former occurs only in NW Madagascar, whereas Ht. harveyi is generally similar to Ht. vanewrighti in wing colouration, but in all known specimens shows a prominent androconial patch on the FWV. As currently recognized, Ht. harveyi occurs in the centre-east of the montane rainforest biome at latitudes south of about 17.5o S, whereas Ht. vanewrighti is only known from the extreme northwestern part of the rainforest biome, but Ht. sp. 20 appears to occur partially sympatrically, distributed from the NE to CE of Madagascar. Ht. westwoodi is quite distinctive from either of these species, having a more uniform colouration on the FWV towards the costa than any of these species, while Ht. viettei also has a prominent pale or whitish highlighting distal of the HWV Mb. Counting from the start of the 658 bp COI-5P barcode, the following four putatively fixed nucleotide differences in the DNA barcode appear (n=4) to be diagnostic within the Ht. harveyi complex (cluster number BOLD:ACW5694) throughout its range, where in parentheses is specified the un-derived state: 274 C(T); 298 T(C); 466 A(G); 498 C(A). DNA barcoding is currently the best means for determining material of Ht. harveyi among material that exhibit the FWV patch.

Description. Wings: dorsal surface uniform very dark chocolate brown (fresh specimen). FW and HW margin strongly crenate. HW has dark brown Sml following the margin, highlighted distad with slightly paler dark brown (also on ventral surface). FWD space-CuA1 ocellus expressed conspicuously, spanning intervein distance CuA1- CuA2 including the narrow concentric deep reddish orange ring, and HWD space-cuA1 ocellus to a lesser extent, spanning about 2/3 of intervein distance CuA1-CuA2, elliptic, also with a narrow deep orange ring. FWV ocellus space-CuA1 is expressed to a similar extent to the FWD, but more conspicuous with a brighter orange ring, while FWV ocelli space-M2 ocellus is about half the diameter (though barely visible on FWD), the orange ring of that of space-M2 slightly compromising space-M1 and -M3. HWV ocelli of space-Rs-CuA2 are expressed as white points lacking strongly discernible rings, except in the case of space-Rs and space-CuA2 which have small black irises and faint orange rings, and that of CuA1 already referred to. HWV space-CuA1 ocellus is elliptic, same size as on HWD, with a quite narrow bright orange ring. The dark Mb of the HWV is strongly and concavely indented proximad of space-CuA1 ocellus and is quite irregular in shape, sharply convex in space-M2 and space-CuA2 and concave-inflexed below vein CuA2 terminating at 1A; above M2 it is again concave to beyond mid-costa but not so distinct from background. A general irroration of darker brown pervades the HWV background and a generally darker band is formed between the Mb and PMb of the central symmetry system, the PMb not so irregular and somewhat convex. This band is highlighted distad and proximad and towards the anal angle by yellowish background colour. (This marbled ventral colour pattern actually befits Aurivillius’ [unavailable] name for his aberration (marmorata) of ‘ Henotesia andravahana ’). Distad of the central HWV band and highlighting it there is a more obscure dark violet to grey-brown region, and the FWV pattern is generally more dark-obscure and less marbled but with about four ochreous strigulae (Costal bars, henceforth ‘Cbs’) perpendicular to the costa beyond mid-costa. Variation. ♂♂ similar, but vary in the intensity of the darker brown irroration and Mb on the HWV. Sexually dimorphic, but marginally so. The ♂ and ♀ are one of the most similar of the Ht. drepana group and in fact the only pair in the Ht. drepana group to have been correctly matched by Oberthür (1916), but the ♀ is larger and paler. The ♂ and ♀ have been photographed in copula ( Fig. 7 D). The ♀ is similar to the ♂, but rather larger and lighter brown on either surface, with a similar ocellus configuration, although the space-CuA1 ocelli tend to be larger in either wing.

Androconia: HWD discocellular brush is bright yellowish fawn (in the HT) to russet brown at tip where it locates on to an androconial patch on the ventral FW towards the base of space-CuA2 just above vein 1A. HWD discocellular patch is dingy orange to brick red, very large and broad lenticular, bisected by R vein near the top of the cell (not observed on HT).

Wingspan/fwl: range 33–37.1/ 18–19.3 mm (n=6 ♂♂), including HT ♂ = 36.3/ 19.3 mm; mean = 35 +/- 1.7 SD/18.7 +/- 0.5 SD mm (n=6 ♂♂). Range 34.8–38.4 mm (n=3 ♀♀)/18.7–21.6 (n=5 ♀♀); mean= 36.6 +/- 1.8 SD/ 20 +/- 1 mm SD (n=3 ♀♀); all examples from type locality.

Etymology. After Don Harvey, who first pointed out (to Claire Kremen) the FWV androconial patch ( Lees 1997: 64).

Discussion. Apart from Ht. tianae sp. nov., Ht. harveyi is the Heteropsis species described here which has longest been known in historical collections (in BMNH, MNHN, and SNHM, at least). The first modern field specimens of this species were recognized by C. Kremen in 1988-1991 in what is now Ranomafana National Park ( Lees, 1997: 64). Until now, recognition of this species has been taxonomically very confusing (as detailed above). Compared with closely related species, it can readily be distinguished androconially and allopatrically from Ht. vanewrighti . However, as mentioned above, morphological and genetic variation across the potential full range of Ht. harveyi needs further examination and this is amenable to a phylogeographic analysis, to see if its circumscription might eventually comprise one or more potentially allopatric or sympatric morphospecies that also have a ventral androconial patch, ‘sp. 20’ (same as ‘sp. 77’?) known from the northeastern montane rainforests of the island (see also Lees, 1997). As mentioned above, the HT ♂ (NHRS-TOBI000000050; Fig. 7 C) of Ht. andrahahana ab. marmorata Aurivillius was examined in SNHM and is considered to be conspecific, but the name is not nomenclaturally available. Also, the HT ♂ of Henotesia wardii Butler, 1879 (BMNH(E) 697802); now called Heteropsis viettei Lees, 2003 , a species potentially confused with Ht. harveyi in Mabille’s (1878) description of Mycalesis andravahana , is not conspecific and the genitalia are different (P. Viette dissection slide no. 4847). Differences in the lateral shape of the tegumen and the extent of the valve relative to the uncus tip in this slide to other preparations of Ht. viettei is considered to represent distortion towards two dimensions due to slide preparation, with distortion occurring most noticeably in the structures most susceptible to pressing, and not in the aedeagus or its relative length.

Additional information. ♂ genitalia: ( Lees, 1997: 109, Fig. 7 i, “12”, voucher 98DL, Fig. 8 A, specimen of Heteropsis cf. harveyi from Anjozorobe). From LV, a very shallowly domed tegumen with a rather straight dorsal edge profile and a distinctly angled brow leads to a hand-scythe shaped uncus whose ventral edge at the base is downbent as if like a ‘dewlap’ and with a rather narrow and pointed hook. Tegumen dorsal profile features a quite narrow v-shaped proximad notch and lateral profile bends round to constriction at division with vinculum, which 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 and valve base very broad leading abruptly to the narrow quite tapering and sub-sinuate valve arms, which are ventrally elbowed and with only a slight indication of a ventral ‘head’ at the unexpanded club, covered with fine setae and sporting a stout spine, inrecurved from DV, where somewhat serrate on mesad edge, pointing towards the uncus tip, which falls well short of the maximum uncus extension, in fact points to the middle of the ‘hook’. Gnathos from right-angled base pointing in line with the uncus tip from LV where relatively straight blade-like, tapered to tip, recurved inward from DV. Saccus quite extended and inflated proximad, less than a third of total valve length (it is longer in the Ht. vanewrighti specimen described below). Aedeagus particularly thin and dorsoventrally flattened, very gently uprecurved distad of the ostium, also quite extended proximad of it, distad tip quite pointed, about 1.75 x total length of valve. Juxta quite prominent proximad and rounded like a small plate from DV. A specimen referred to Ht. harveyi from Miaranony, 1080 m (156DL) is very similar in proportions to 98DL, with valve terminating in an inwardly pointing spine that falls short of uncus tip, but has a saccus about 1/4 longer and the hook of uncus about 1/3 deeper (from LV). A future more satisfactory resolution of genitalic variation within the Ht. harveyi complex (currently containing four COI- 5P BIN clusters) should be based on DNA barcoded material.

DNA divergences: COI-5P cluster number BOLD:ACW5694 (exemplars BMAD217-15 from type locality and BMAD270-15, DL-4-963 from Ambondrombe) which is 1.52% divergent to Ht. sp. 20 (exemplar BMAD213- 15 DL14A-0387, Anjanaharibe Sud, cluster number BOLD:ACW5777), 1.67% divergent to Ht. sp. cf. harveyi (BMAD268-15, DL14AM-0025 from Anjozorobe, cluster number BOLD:ACW5778) and 1.98% divergent to Ht. vanewrighti (BMAD064-04, Bekolosy, cluster number BOLD:AAE4111). The 357 bp cytochrome b sequence of Ht. harveyi (exemplars BMNH(E) 671648 FJ819453 , DL-05-367 from Ranomafana, and DL-0290, BMNH(E) 697406 from Maromizaha), is about 1.12% divergent both to Ht. sp. 20 (as Ht. sp. 77; exemplar CK678 from Anjanaharibe Sud) and to Ht. harveyi (exemplar BMNH(E) #676467 FJ819454 , DL-05-385 from Ranomafana), and is about 1.56% divergent to Ht. vanewrighti (exemplar DLBEK94-130, BMNH(E) #697875 from Bekolosy), whereas it is about 4.2% pairwise divergent to sequences of Heteropsis imerina (BMNH(E) #676683 FJ819266 , BMNH(E) #676685 FJ819267 ; Monaghan et al., 2009; additional members of the Ht. passandava group in that study included Ht. passandava , Ht. narova , ‘ Ht. difficilis ’ [ Ht. andravahana stat. rev.] and Ht. strato ). No COII sequence is available on GenBank (see below).

Phylogeny/sister species: closely related to Ht. vanewrighti , and to two (?) still undescribed species in the same cryptic complex. In their cladistic analysis of COII sequences, not including Ht. vanewrighti, Torres et al., (2001: 465) had full support for a sister relation of Ht. harveyi (their “ Hen. sp. 12” from Ranomafana National Park); with their “ Hen. sp. 77” from Anjanaharibe Sud, a morphospecies that Lees (1997: 64) also considered the same as “sp. 20” from Ambohitsitondroina Mahalevona; unfortunately there seems to be no sequence on NCBI for the exemplar of “sp. 12”. The “sp. 77” of Torres et al., 2001 [here considered to be the same as “sp. 20” from Masoala] appears from putatively fixed differences in its DNA barcodes (n=10 currently on BOLD; cluster number BOLD:ACW5777) not to be conspecific with Ht. harveyi (as also treated by Lees, 1997: 64).

Ecology and distribution.

Habitat: montane rainforest with fine-leaved climbing bamboos.

Behaviour: ♂♂ fly quite rapidly in the substory in fairly open light areas of the forest. ♀♀ have been found resting quite low on tree trunks and fly less readily except just before rain.

Hostplant: suspected to be fine-leaved climbing bamboos.

Early stages: unknown.

Distribution: from Fianarantsoa/Ranomafana (including type locality) probably at least as far north as Andasibe-Mantadia in the eastern rainforest biome, south to Ambondrombe (according to DNA barcode) and possibly as far South as Chaines Anosyennes (referred specimen) ( Fig. 30 D, lilac dots). Light brown dots in Fig. 30 D represent localities for specimens morphologically similar to Ht. harveyi falling outside the known range of this species, and where the presence of cluster BOLD:ACW5694 has not been confirmed. Populations in the more central Angavo Massif (exemplar BMAD268-15, cluster number BOLD:ACW5778, Anjozorobe) might represent a different species. However, data is currently insufficient to indicate whether the illustrated genitalia from Anjozorobe represent Ht. harveyi or not (hence the labelling ‘ Ht. cf. harveyi ’). Ideally, dissections would be based on barcoded material, a matter to be addressed in future.

Elevational range: 700–2030 m. (n=86, including referred specimens and observations; 1087–1590 m. for DNA barcode-verified specimens).

Referred specimens. 33 ♂♂, Madagascar Fianarantsoa Perrot Frères 2e Semestre 1892/ Ex Oberthür Coll. Brit. Mus. 1927-3; specimen [ MNHN], E, Anjorofozo [Anjororo?], Forêt Tanala, Ranomafana Madagascar, [?ca. 20.933o S, 47.367o E +/- 15 km, 1304 +/- 11 m], 1901, Ch. Allaud; ♂, E, Miaranony, 0.5 km N of, Ranomafana National Park, 1079 m [21.1633o S, 47.5528o E +/- 0.3 km, 1080 m], 10 hi, Station 11 [fruit trap], 23/10/1992, C. Kremen and H. Raharatsimba|156 DL; 2 ♂♂, E, Tsitery, Forêt de, Tsarafidy [Vatobe forest lies to W.], 21.1881o S, 47.2132o E +/- 0.25 km, 1425 +/- 50 m, 2/3/1995, D.C. Lees; ♂, E, Vohiparara, 1.4. km W, 1220 m, 21.2346o S, 47.3943o E +/- 0.9 km, 1220 m, 13/3/1991, C. Kremen et al.: Genitalia 100; ♂, data as above but: Genitalia 102; ♂, E, Vohiparara, 21.2346o S, 47.3943o E +/- 0.92 km, 1190 +/- 5 m 19/2/1995, 11:00, D.C. Lees; specimen [ MNHN], E, Vohiparara, Sentier Forestier a 3 km au S.O. de, Fianarantsoa [21.24o S, 47.43o E +/- 1 km, 1029 +/- 50 m], 25/5/ 1969; specimen [ MNHN], data as above but: 9/3/1969; ♀, E, Vohiparara, 0.6 km S[W], 1220 m, 21.2427o S, 47.3785o E +/- 1 km, 1220 +/- 6 m, 11/3/1991, C. Kremen et al.; specimen, data as above but: 19/2/1995, 14:30– 15:09, D.C. Lees; ♂, E, Ranomafana National Park, Sahamalaotra, 21.2437o S, 47.4019o E +/- 1.77 km, 1087 +/- 50 m, 5/12/2004, D.C. Lees: DL-05-385; BMNH (E) #676467 [ DNA voucher; cytochrome b sequence FJ 819454 : Monaghan et al., 2009]; ♂, E, Ranomafana National Park, Sahamalaotra, 21.2437o S, 47.4019o E +/- 1.77 km, 1087 +/- 50 m, 5/12/2004, R. Ranaivosolo: DL-05-367; BMNH (E) #671648 [ DNA voucher; cytochrome b sequence FJ 819453 : Monaghan et al., 2009]; ♂, E, Ranomafana, Sahamahalaotra?, 21.262o S, 47.4212o E +/- 0.15 km, 991 +/ - 25 m, 5/12/2004: 12:19, Pierre: DL-05-372, IA194 [isotope voucher]; ♂, E, Vatoharanana, Ranomafana, 7 km SW of, 1150 m, VR, 21.2667o S, 47.4333o E +/- 0.87 km, 1150 m, 22/2/1991, C. Kremen et al.: Genitalia vial 103; ♂, E, Mt. Maharira, 21.3307o S, 47.4162o E +/- 3 km, 1288 +/- 87.5 m, 12-Oct-92, C. Kremen: CK822; ♂, E, Andranobazaha forest, Ambondrombe, camp, 21.87816o S, 47.2474o E +/- 0.25 km, 1604 +/- 50 m, 22/3/2004: 11:38, R. Ranaivosolo: DL-4-883; ♂, E, Ambondrombe, 21.878 o S, 47.247 o E +/- 0.5 km, 1590 +/- 50 m, D.C. Lees: DL-4-963, BMAD 270-15 [ DNA barcode, verified]; ♂, E, Ambondrombe, 21.878 o S, 47.247 o E +/- 0.5 km, 1590 +/ - 50 m, D.C. Lees: DL-0614, BMNH (E) 671979 [ DNA voucher], KA509 [=KA-P509, DNA extract number]; ♀, E, Ambavafatra-Vohitrasiva, Andringitra, 22.1579o S, 47.025o E +/- 1.24 km, 1350 m, 17/2/1995, C. Kremen; ♂, E, Vohitrasiva, near ridge, 1390–1450 m, Andringitra, [mountaintop on way to Vohipia (1760 m. pass)], 22.1684o S, 47.0289o E +/- 0.1 km, 1450 +/- 50 m, 17/2/1995, 11:40–13:15, D.C. Lees:, specimen [ MNHN], forêt d’Anjavidilava, Andringitra oriental, Madagascar Centre, 2005 m [ca. 22.21o S, 46.94o E +/- 7.5 km], 1–15/1/1971, P. Griveaud; specimen [ MNHN], forêt d’Imaitso Anjavidilava, Andringitra-Ambalavao, Madagascar Centre, 2030 m [ca. 22.21o S, 46.94o E +/- 7.5 km], 20/1/1958, P. Griveaud; ♂, [ MNHN], SE, Chaines Anosynnes, massif nord, N.-O. de Manentenina, [ca. 24.16o S, 47.01o E +/- 12 km, 1050 m], 22/11/1971, P. Griveaud; [treated with reservation; more northerly records]: ♀, E, Maromizaha, near granite quarry, S side R.N.2 28 km E Moramanga, 3.2 km E of turnoff for Perinet, then right on dirt road to quarry, path to S, 1100–1200 m, 18.9545o S, 48.4477o E +/- 1 km, 1150 +/- 50 m, 29/1/1995, C. Kremen: CK215 [ DNA voucher]; ♂, E, Maromizaha, 18.9668o S, 48.4547o E +/ - 1.49 km, 1109 +/-73 m, 25/2/2002, D.C. Lees: DL 02-516; ♂, data as above but: D.C. Lees: A71, BMNH (E) #697406 [ DNA voucher; cytochrome b], KAP545 [=KA-P545, DNA extract number]; ♀, E, Andasibe-Mantadia, 11/2014, D.C.Lees: DL 14AM-156, IA598 [isotope voucher]; ♂, E, Ankeniheny, 19.15o S, 48.303o E +/- 2.5 km, 1068 +/- 50 m, 7/1/1994, WG16 [wing image], IA56 [isotope voucher]; ♂, E, near Ankosy summit, Zahamena, 17.49405o S, 48.73325o E +/- 0.067 km, 1300 +/- 25 m, 7/11/2006, D.C. Lees: DL 06-157; ♀, E, slope above cascade [Zahamena NW], 17.53o S, 48.7269o E +/- 1.5 km, 1064 +/- 25 m, 6/11/2006, 09:28, D.C. Lees: DL 06-094, IA31 [isotope voucher]; 5 specimens [ BMNH], E, Antsianaka [ca. 17.6182o S, 48.738o E +/- 10.7 km, 1328 m],1892, E. & B. Perrot; 2 specimens, E, Belongoina, Reserve Nat. III, Andranomalaza [ca. 17.65o S, 48.617o E +/- 5 km, 915 m], 10/1956, P. Soga; ♀, E, Analamay, 18.80965o S, 48.34535o E +/- 0.3 km, 1050 +/- 50 m, 13/3/2004 | 16:29, D.C. Lees: DL-4-545; specimen [ MNHN], E, Moramanga, Reg. [18.933o S, 48.2o E +/- 25 km, 902 m]|193- [last digit of year not specified], G. Oulsoufieff; 2 specimens, E, Analamazaotra Forest Station, 18.9384o S, 48.41o E +/- 0.72 km, 1043 +/- 93 m, 27/2/2002, D.C. Lees: DL 02-535A, DL 02-535B; specimen [ MNHN], Ambodiriana, route de Lakato, km 10, Madagascar Est, 1050 m [19.025o S, 48.35o E +/- 1 km, 1050 m], 3/1957, P. Griveaud, R. Vieu; specimen, E, Sandrangato [19.1083o S, 48.2417o E +/- 15 km, 1033 m]; ♀, E, Sandrangate, 19.1083o S, 48.2417o E +/ - 15 km, 1033, 6/11/1994, WG41 [wing image], IA215 [isotope voucher]; specimen [ MNHN], E, Mahatsinjo pres Tananarive Madagascar, [?ca. 19.167o S, 48.033o E +/- 15 km, 913 m], 1913, H. Donckier.

Figures

FIGURE 7. Adults in DV (left), VV (right) or in natural postures. A: HT ♂ of Heteropsis harveyi Lees & Kremen, sp. nov. (NHMUK 010289119). B: PT ♀ of Ht. harveyi (NHMUK 010289121). C: HT ♂ of Henotesia andravahana ab. marmorata Aurivillius, 1911 (NHRS-TOBI 000000050). D: pair of Ht. harveyi in copula (Maromizaha, photo courtesy Lance Featherstone). E: ♂ of Ht. imerina (Ankazomivady, 23 / 03 / 2003, photo: D. C. Lees), for comparison, showing relatively produced ‘ tail’ on HW vein M 3.

FIGURE 8. ♂ genitalia in LV (left), DV (right). A: Heteropsis cf. harveyi (see text) (Anjozorobe; 98 DL). B: Ht. vanewrighti Lees, sp. nov. (Mt. Bekolosy; 233 DL). C: Ht. imerina Lees, sp. nov. (Ankazomivady; 154 DL). Scale bar: 3 mm.

FIGURE 30. Distribution maps. A. Purple: Heteropsis vanewrighti Lees, sp. nov. Green: Ht. pauliani Lees, sp. nov. Yellow: Ht. mimetica Lees & Kremen, sp. nov. Light blue: Ht. hazovola Lees & Raharitsimba, sp. nov. Turquoise: Ht. imerina Lees, sp. nov. [northern populations overlapping with Ht. vanewrighti and Ht. pauliani are not shown]. Pink: Ht. westwoodi Lees, sp. nov. B. Red: Ht. avaratra Lees & Kremen, sp. nov. Light blue: Ht. kremenae Lees, sp. nov. Fawn: Ht. vertigo Lees & Raharitsimba, sp. nov. C. Ochre: Ht. roussettae Lees & Kremen, sp. nov. Slate-blue: Ht. menamenoides Lees, sp. nov. Olive: Ht. lanyvary Lees, sp. nov. Dark green: Ht. borgo Lees, sp. nov. Light green: Ht. barbarae Lees & Kremen, sp. nov. Brown: Ht. oberthueri Lees, sp. nov. Dark pink: Ht. tianae Lees & Kremen, sp. nov. D. Mid green: Ht. sogai Lees, sp. nov. Dark brown: Ht. tornado Lees, Allaoui & Aduse-Poku, sp. nov. Lilac: Ht. harveyi Lees & Kremen, sp. nov. Light brown: centraleastern populations of Ht. harveyi complex. Scale bar: 500 km.