Heteropsis avaratra 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 : 52-54

Heteropsis avaratra Lees & Kremen , sp. nov.

LSID: urn:lsid:zoobank.org:act:17C4A71B-C04D-4AC1-9407-4D52B343F803

Prior references: sp. 25 ( Lees 1997: 64, Torres et al., 2001: 462); ‘ H. subsimilis ’ ♂, ( d’Abrera 1980: 182; 1997: 221).

Type material., Deposition BMNH: Holotype: ♂ ( Fig. 13 C), Madagascar N, Montagne d'Ambre, 12.5289o S, 49.1727o E +/- 0.15 km, 1057 +/- 25 m, 18/11/2004, fruit trap MD3, D.C. Lees: DL-05-169, NHMUK 010289141 [ QTR barcode].

Paratypes: Deposition BMNH: ♂, Madagascar N, Montagne d'Ambre, 12.4984o S, 49.1754o E +/- 0.5 km, 812 +/- 5 m 17/11/2004, D.C. Lees: DL-05-69, BMNH (E) #676332; IA82 [isotope voucher], CCDB-02225-B04 [ DNA barcode voucher]; ♂, N, Montagne d'Ambre, 12.5289o S, 49.1727o E, +/- 0.15 km, 1057 +/- 25 m, 16/11/2004, D.C. Lees: DL-05-68, BMNH (E) #676331, 2133 (= DL 2133; DNA extract number), IA91 [isotope voucher]; ♀, N, Montagne d'Ambre, 1200 m, 12.5405o S, 49.1682o E +/- 1 km, 1200 +/- 50 m, 13/1/1995, D.C. Lees: DLMDA 95- 0 0 0 1, NHMUK 010289142 [ QTR barcode]; ♂, N, Montagne d'Ambre, 900 m, 12.525o S, 49.1722o E +/- 5 km, 900 +/- 5 m, 16/1/1995, 0 945 [= DL 0945], BMNH (E) #672310 [ DNA voucher], FJ 819380 , cob 357 bp; ♂, N, Montagne d'Ambre, c. 1000 m, 12.525o S, 49.1722o E +/- 5 km, 900 +/- 5 m 16/1/1995, D.C. Lees, DCLW-0153 [wing prep.]; BMNH (E) #672438 [ DNA voucher; FJ 819358 , cob 357 bp];

Deposition MNHN: ♂ [MNHN], MADAGASCAR NORD Montagne d’Ambre Les Roussettes alt. 1000 m 27.XI.1958 P. Viette|DCL-DB-3364; ♂ [MNHN], MADAGASCAR NORD MONTAGNE D’AMBRE LES ROUSSETTES 1000 m 25-V-1970 P. GRIVEAUD|DCL-DB-3381; ♀ [MNHN] ( Fig. 13 D), MADAGASCAR NORD Montagne d’Ambre Les Roussettes alt. 1000 m 6-7 XII 1958 |DCL-DB-3385;

Deposition ABRI: ♂, Madagascar N, Montagne d'Ambre, 12.5289o S, 49.1727o E +/- 0.15 km, 1057 +/- 25 m, 19/11/2004, R. Ranaivosolo: DL-05-181.

Deposition summary: BMNH (HT♂, 4 PT♂♂, 1 PT♀)), MNHN (2 PT♂♂, 1 PT ♀), ABRI (PT♂).

Type locality. Madagascar N, Montagne d'Ambre, 12.5289o S, 49.1727o E +/- 0.15 km, 1057 +/- 25 m.

Diagnosis. compared with Ht. avaratra , Ht. subsimilis has the Rs ocellus on HWV usually quite prominent and Ht. sogai has the Mb and PMb strongly dentate between M1 and CuA2. Populations of Ht. avaratra in the northern reaches of the main rainforest belt can be hard to distinguish by wing pattern from the sympatric Ht. subsimilis though easily discriminated by DNA barcoding, but Ht. avaratra lacks an inflated M1-M2 vein ( Fig. 12 E), and the inflated vein below the Sdb basad of the fork of Rs+M1 is more symmetric (more rounded on the anteriad edge) in Ht. subsimilis ( Fig. 12 D). Ht. avaratra , especially from the topotypical population, has a more prominently ventral yellow cast than in other species including Ht. subsimilis , and is not variegated with orange as in Ht. kremenae .

Description. Wings: Dorsal surface homogeneous mid-brown as in others of the Ht. subsimilis group. FWD space-CuA1 ocellus with black iris at least spanning CuA1-CuA2, with a fairly wide pale orange roundsubhexagonal ring, not perfectly concentric, often protruding a bit proximad within space-CuA1. FWD space-M1 ocellus usually small but conspicuous and subelliptic, with narrow orange ring. HWD space-CuA1 ocellus subelliptic and a little larger than that of FWD space-M1, usually spanning at least ¾ of HW veins CuA1-CuA2, including the fairly narrow concentric orange ring. Other ocelli not expressed on HWD. Ventral surface a more strongly ochreous background colour, especially in Montagne d’Ambre populations, than in others of the Ht. subsimilis group, and the area between Mb and PMb is finely irrorated in brown to a greater extent than other parts of the wing. FWV space-CuA1 ocellus with black iris same size as upperside and ring fairly subhexagonal and mainly yellow with gradation to orange towards tergal edge. The ring is slightly more pointed towards proximad edge where it hugs a strongly concave rufous brown Mb before it bends back to mid costa and Ore as a yellow hook like the small tail of a comma, representing the end of a spiral arm. FWV space-M1 ocellus as on upperside, slightly more conspicuous with a yellowish ring of background colour. On HWV, space-CuA1 ocellus is the largest as usual and subelliptic and including the concentric yellow ring spanning about 4/5 of veins CuA1-CuA2 (generally smaller than in Ht. kremenae ). HWV space-CuA2 ocellus not evident in the HT ♂. HWV space Rs-M3 ocellus reduced to white point often with narrow black iris and Orng blending with background. The FWV cell area has two unequal brown edged more yellow transverse Cbs, the distad one wider. There is no distinctly orange shoulder to the basal FWV costa, which is strigulated evenly with black. The HWV is highlighted with coronal-like yellow-ochreous flares distad of the Mb, particularly in M2, where the Mb is toothed outwards slightly at vein intersections M2, M3 and CuA1, inflexed proximad of ocellus space-CuA1 and bends back towards 1A. Variation. ♀♀ similar but slightly larger. Referred material from the main rainforest block is sometimes harder to distinguish by wing pattern from Ht. subsimilis , but Ht. avaratra tends to be a larger species (see also Fig. 16 F). Mb well marked and sometimes more jagged-slightly inflexed than in HT, with yellow Mf in space-M2 distad of the Mb sometimes rectangular, although this character is not distinct in the HT specimen. HWV space-CuA2 ocellus pointlike in some specimens, or subelliptic with a pale yellow ring. HWV space-Rs-M3 ocelli usually but not always point-like with that of HWV space-Rs sometimes slightly more evident, as may be that of space-CuA2. FWV Cbs sometimes delineate indistinctly a third yellow band.

Androconia: Sdb dark brown to russet brown, underlying Sdp varying from ‘bullet’-shaped, pointed to flattopped. M3-A2 veins, but not M2, linearly inflated in the HWD (1A from 1/6 and widest at about 40%, tapering thence to margin; 3A from base to margin +/- parallel from ca. 1/8 to 7/8).

Wingspan/fwl: range 32.1–38/ 17.7–21 mm (n=7 ♂♂); mean = 35.8 +/- 2.3 SD/19.7 +/- 0.5 SD mm (n=7 ♂♂), including HT ♂ 33.2/ 19.5 mm. Range 35.8–37.0 (n=2 ♀♀)/ 20.4–20.7 mm (n=4 ♀♀); mean 36.4 (n=2 ♀♀)/20.5 +/ - 0.1 SD mm (n=4 ♀♀).

Palps: penultimate segment on outside face cream ochreous towards compound eye, dark brown away from eye with a few light coloured scales within this dark border (more apparent in the ♀); fringed dark brown where not contacting eye. Face of palp away from compound eye pale ochreous.

Etymology. ‘avaratra’ means ‘north’ in Malagasy, referring to its distribution, as it is currently known.

Additional information. ♂ genitalia: 315DL (referred specimen, Joffreville); Lees (1997: 106, Fig. 7 f; “25”; Fig. 15 B). Miniaturised, about 1.6 mm long, and with configuration typical of the Ht. subsimilis group; there is no 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 Ht. subsimilis group; from LV, the valve bases are rather symmetrically-‘skittle’-shaped ( Lees 1997). Uncus is slightly longer than tegumen (which is shallowly and widely notched proximad from DV) and quite inflated dorsoventrally before the slightly pointed downturned tip (distinctly inflated from DV before tip). Gnathos tapered from small base, fairly straight, not reaching tip of uncus (from DV, the appearance in the examined specimen is quite quadrate, not particularly sinuate or bull’s horn-like). Valve with prominent rounded dorsal ‘shoulder’, valve arm tapering to distinct ‘beak’ that points to uncus base (strongly incurved from DV) with limited serration (more obvious distad from DV) and not much indication of a club at valve end. Saccus relatively small, slightly bulbous and juxta also not prominent proximad. Aedeagus about same length as valve, shallowly recurved distad of ostium, which is slightly bulbous at proximad tip.

DNA divergences: the COI-5P barcode (cluster number BOLD:AAD0195, exemplar BMAD016-09 from Montagne d’Ambre) is 3.46% pairwise divergent to Ht. subsimilis (cluster number BOLD:AAB4493, currently with 27 members on the BOLD database; e.g. exemplar FJ666680 , Linares et al., 2009 dataset) whereas about 6.55% divergent to that of Ht. pauper (BOLD:AAA6758). The COII dataset of Torres et al., (2001) shows 5.27% pairwise divergence (398 bp compared) of Ht. avaratra (‘sp. 25’) to Ht. subsimilis .

Phylogeny/sister species: in the morphological analysis of Lees 1997, this taxon (“TWNFV”) connected to Heteropsis sogai (“THRNN”) but in only one (jackknife) tree with support ( Lees 1997: 157, Fig. 2 ). In their analysis of COII sequences (where Ht. sogai was the only member of the Ht. subsimilis group on Madagascar not included), Torres et al., (2001: 366) had a meager 64% bootstrap support for a sister relation of Ht. avaratra , their “ Hen. sp. 25”, AY 040146 based on one individual from the type locality) with Ht. subsimilis ( AY 040145 based on one individual from Ranomafana National Park). Ht. avaratra (as “sp. 25”) was however strongly supported as sister to Ht. subsimilis in the phylogeographic analysis of Linares et al., (2009, p. 488, their Fig. 2 ) including COI or also EF-1a and wingless (p. 490, their Fig. 4 ). However, neither the latter study nor that of Kodandaramaiah et al., (2010), which found the same relationship, included Ht. kremenae nor Ht. sogai (but that of Torres et al., 2001, who found the same with COII, did include Ht. kremenae as their ‘ Hen. sp. 14A’), in fact only otherwise Ht. pauper was included among the Ht. subsimilis group. Ht. kremenae is more distant to Ht. avaratra by COII data (see also description above). Ht. sogai might be its sister (as in combined tree in Aduse-Poku et al., 2016, in press).

Discussion. This species was first recognized in the field as ‘sp. 25’ by Claire Kremen on 19-20/02/1992 at Montagne d’Ambre, but there were sparse earlier collections in BMNH (Meade-Waldo, 1906) and at MNHN. It is understandable however that these specimens were not described earlier owing to their superficial similarity with Ht. subsimilis . All other available types of the Ht. subsimilis group have been examined and are clearly distinct with their type localities further south in Madagascar (see under Ht. kremenae ).

Ecology and distribution.

Habitat: montane primary rainforest, particularly on ridges/slopes.

Behaviour: unusually for Heteropsis , adults have been seen visiting Impatiens (Balsaminaceae) flowers in Montagne d’Ambre (pers. obs.). Adults fly low and would usually feed on fruits on the ground, but also feed on sap runs. In Anjanaharibe Sud, however, adults were attracted to banana bait and no flower visitation was witnessed.

Hostplant: unknown, probably low grasses.

Early stages: unknown.

Distribution: Montagne d’Ambre is an important locality but the species occurs in rainforest also in the main rainforest block from Tsaratanana to Makira ( Fig. 30 B, red dots). If Ht. sogai , not sequenced, is not more closely related, Ht. avaratra might represent Ht. subsimilis ’s replacement in Montagne d’Ambre, and in that case, Ht. avaratra has achieved sympatry and secondary contact with Ht. subsimilis in northwestern and northeastern parts of the main rainforest belt. Due to a formatting error, the locality was incorrectly given in Table 1 of Torres et al., 2001 as “Ankazomivady, 32 km S. Ambositra” instead of “Montagne d’Ambre”. Lees (1997: 66) assigned a series of specimens in BMNH from Mahassabe [= Mahasoabe], forêt SE Fianarantsoa (“sp. 55”) to this species, but this needs re-examination as they might easily be referable to Ht. subsimilis or another species (plus this location includes a range of material that seems more characteristic of northeastern localities, for example Ht. erebina ( Oberthür, 1916) ; see also Ht. tornado ).

Elevational range: 530–1630 m. (n=64, including referred specimens and observations).

Referred specimens. ♂, Madagascar N, Montagne d'Ambre, path to Petit Cascade, 12.526o S, 49.1749o E +/- 0.2 km, 1084 +/- 75 m, 16/11/2004, D.C. Lees: DL-05-71, BMNH (E) #675489 [ DNA voucher; FJ 819359 ]; ♂, N, Montagne d'Ambre, 12.5289o S, 49.1727o E +/- 0.15 km, 1057 +/- 25 m, D.C. Lees: DL-05-66, BMNH (E) #676329 [ DNA voucher; FJ 819360 ]; ♂, data as above but: DL-05-67, BMNH (E) #676330 [ DNA voucher; FJ 819361 ]; ♂, N, Montagne d'Ambre, 12.51405o S, 49.17385o E +/- 1.6 km, 985 +/- 125 m, R. Ranaivosolo: DL-05-77, BMNH (E) #676336 [ DNA, FJ 819364 ], IA92 [isotope voucher]; ♂, N, Montagne d'Ambre, piste vers petit cascade, 12.5264o S, 49.1761o E +/- 0.25 km, 1064 +/- 25 m, R. Ranaivosolo: DL-05-99, BMNH (E) #676342 [ DNA voucher; FJ 819365 ]; specimen, N, Montagne d’Ambre, 14/01/1995, WG33 [wing imaged], IA138 [isotope voucher]; ♂, N, Montagne d'Ambre, piste petit cascade, 12.5264o S, 49.1761o E +/- 0.25 km, 1064 +/- 25 m, R. Ranaivosolo: DL-05- 100, BMNH (E) #676343 [ DNA voucher; FJ 819366 ]; ♂, N, Montagne d'Ambre, petit cascade, 12.5264o S, 49.1761o E +/- 0.25 km, 1064 +/- 25 m, R. Ranaivosolo: DL-05-101, BMNH (E) #676344 [ DNA voucher; FJ 819367 ]; ♂, N, Montagne d'Ambre, 1100 m, 12.511o S, 49.161o E +/- 0.75 km, 1100 m, 19/2/1992, C. Kremen et al.: Genitalia 146; ♂, N, Montagne d'Ambre, 12.5253o S, 49.1483o E +/- 2.47 km, 950 +/- 350 m, 20/2/1992, C. Kremen et al.: Genitalia 147; ♂, data as above but: Genitalia 149; ♂, N, Montagne d'Ambre, 12.5253o S, 49.1483o E +/- 2.47 km, 1185 +/- 115 m, 20/2/1992, C. Kremen: CK841; ♂, N, Montagne d'Ambre, site MD1, 12.5289o S, 49.1727o E +/- 0.15 km., 1057 +/- 25 m, D.C. Lees: DL-05-176, 19/11/2004, KA521 [=KA-P521; DNA extract voucher]; ♀, NE, Lac Amparihibe, Makira, 15.0349o S, 49.5836o E +/- 0.5 km, 849 +/- 50 m, 31/2/2003, D.C. Lees, BMNH (E) #697130 [ DNA voucher]; ♀, NE, Lac Amparihibe, Makira, 15.039o S, 49.572o E +/- 0.5 km, 900 +/- 50 m, 31/2/2003, D.C. Lees, BMNH (E) #697131 [ DNA voucher], KA528 [=KA-P538; DNA extract number]; ♂, NE, Lac Amparihibe-Ankilandy, walkout, 15.07o S, 49.58o E +/- 5 km, 700 +/- 100 m, 1/3/2003, D.C. Lees, BMNH (E) #697942 [ DNA voucher]; ♀, NE, Sahantaha, 15.23o S, 49.53o E +/- 0.15 km, 550 +/- 100 m, 28/1/2003, D.C. Lees: 1645 [= DL 1645; DNA extract number], BMNH (E) #697949; ♀, NE, Sahantaha, 15.2406o S, 49.5436o E +/- 0.15 km, 697 +/- 100 m, 18/1/2003, D.C. Lees: 1649 [= DL 1649; DNA extract number], BMNH (E) #697953; ♀, NW, Tsaratanana, descent, 14.1898o S, 48.9452o E +/- 0.8 km, 1630 +/- 100 m, 24/12/2004, D.C. Lees: DL-05-855, BMNH (E) #675490 [ DNA voucher]; ♂, data as above but: DL-05-853, BMNH (E) #675491 [ DNA voucher]; ♂, NW, Tsaratanana, descent, near forest margin, 14.2034o S, 48.9532o E +/- 0.25 km, 1441 +/- 50 m, 24/12/2004, D.C. Lees: DL-05-844, BMNH (E) #675492 [ DNA voucher]; ♂, data as above but: DL-05-843, BMNH (E) #675493 [ DNA voucher]; ♂, data as above but: DL-05-842, BMNH (E) #675494 [ DNA voucher]; ♀, NE, Lac Amparihibe, 15.0351o S, 49.5862o E +/- 0.15 km, 870 +/- 50 m, 23/2/2002, D.C. Lees, BMNH (E) #675516 [ DNA voucher]; ♀, NE, Lac Amparihibe, 15.039o S, 49.572o E +/- 0.5 km, 814 +/- 50 m, 19/2/2003, D.C. Lees; BMNH (E) #675520 [ DNA voucher]; specimen, NW, Tsaratanana, Antetykalambazaha, below camp, “fougeres”, fruit low trap, 14.1898o S, 48.9452o E +/- 0.1 km, 1630 +/- 25 m, R. R. Ranaivosolo: DL-05-829, BMNH (E) #676598 [ DNA voucher]; ♀, NW, Tsaratanana, Ampidiranala, 14.20025o S, 48.95175o E +/- 0.54 km, 1542 +/- 85 m, 20/12/2004, R. Ranaivosolo: DL-05-757, 2487 [= DL 2487; DNA extract number], BMNH (E) #676687; ♀, N, Montagne d'Ambre, 12.5289o S, 49.1727o E +/- 0.15 km, 1057 +/- 25 m, 19/11/2004; D.C. Lees: DL-05-182: IA204 [isotope voucher]; ♀, N, Montagne d'Ambre, 12.5289o S, 49.1727o E +/- 0.15 km, 1057 +/- 25 m, 19/11/2004; D.C. Lees: DL-05-183, IA205 [isotope voucher]; ♂ [ BMNH] ( Fig. 15 B), N, Joffreville [ca. 12.483o S, 49.2o E +/- 15 km, 717 +/- 50 m], 1921, E. Brouhard|315 DL [genitalia]; specimen [malaise], N, Parc National Montagne d'Ambre [1st campsite], 12.5144o S, 49.1814o E, 960 m, Irwin, Schlinger, Harin'Hala: MA-01-01A-02; specimen [malaise], data as above but: MA-01-01A-06; specimen [malaise], N, Parc National Montagne d'Ambre [Petit Lac road], 12.5203o S, 49.1792o E, 1125 m, R. Harin’Hala: MA-01-01D-05; specimen [malaise], data as above but: MA-01-01D-06; specimen [malaise], data as above but: MA-01-01D-07; specimen [malaise], data as above but: MA-01-01D-08; series DCL-DL-3369 to DCL-DB-3385 [ MNHN], including: ♂ [ MNHN], Nord, Les Rousettes, Montagne d'Ambre, 1000 m, [ca. 12.525o S, 49.1722o E +/- 5 km], 3/11/1958; ♂ [ MNHN], data as above but: 6/12/1958, P. Viette; 3 ♂♂ [ MNHN], data as above but: 25/5/1970, P. Griveaud; specimen, N, Petit Lac, Montagne d'Ambre, 1115 m, [12.5333o S, 49.1667o E +/- 0.91 km, 1115 m], 26/5/1970, P. Griveaud; ♂ [ MNHN], data as above but ♂ [ MNHN]: 26–27/5/1970, P. Griveaud; ♂, N, Montagne d'Ambre, vers premier petit sommet, route GCASC->Gite, 12.59435o S, 49.1583o E +/- 1 km, 872 +/- 25 m, 17/11/2004, R. Ranaivosolo: DL-05-121; ♂ [ BMNH], N, Forêt d'Ambre, 3000 ft, [ca. 12.6282o S, 49.1448o E +/- 8.51 km, 914 m], 7/3/1906, Meade-Waldo| Photographed by B. D’Abrera 77/78| BMNH (E) 674887; ♂, E, Anjanaharibe Sud, ~ 950 m, 14.7502o S, 49.4998o E +/- 0.8 km, 950 +/- 50 m, 25/11/1995, 11:00–13:15, C. Kremen: CK484, IA336 [isotope voucher]; specimen, NE, Lac Amparihibe, Makira, 15.0301o S, 49.5827o E +/- 0.75 km, 920 +/- 75 m, 19/2/2003, R. Ranaivosolo: DL 03-0002, IA337 [isotope voucher]; ♂, NE, Lohan i' Sahantaha, site SD1, Makira, 15.2287o S, 49.5320o E +/- 0.15 m, 400 +/- 50 m, D.C. Lees, 24/01/2003: 10:00, BMNH (E) #697350, 0 234 [= DL 0234, DNA extract number], KA538 [=KA-P538, DNA extract number]; ♂, E, Anjanaharibe Sud, 14.7502o S, 49.4998o E +/- 0.8 km, 950 +/- 50 m, 25/11/1995, 11:00– 13:15, C. Kremen: CK485, IA180 [isotope voucher]; ♂, NE, Anjanaharibe Sud, 11/2014, D.C. Lees: DL-14A- 0 0 83, IA495 [isotope voucher]; ♂, NE, Anjanaharibe Sud, 11/2014; D.C. Lees: DL-14A-0059, IA496 [isotope voucher]; ♀, NE, Anjanaharibe Sud, Takhtajania trail, 23/11/2014, D.C.Lees: DL 14A-0365, IA561 [isotope voucher]; ♂, NE, Anjanaharibe Sud, summit trail, 19/11/2014: 05:51, D.C.Lees: DL 14A-0170, IA563 [isotope voucher].

Figures

FIGURE 2. Adults in DV (left), VV (right). A: HT ♂ of Heteropsis mimetica Lees & Kremen, sp. nov. (Anjanaharibe Sud; BMNH (E) 1717105), B: PT ♀ of Ht. mimetica (Anjanaharibe Sud; BMNH (E) 1717113). C: HT ♂ of Pseudonympha cowani Butler, 1880 (‘ Fianarantsoa’). D: LT ♂ of Culapa houlbertiana Oberthür, 1923 (‘ Tamatave’, BMNH (E) # 674789). F. LT ♂ of Mycalesis antahala Ward, 1872 ([Madagascar], BMNH (E) 672567). Scale bar: 30 mm. The figure shows apparent contrasting mimicry on the two surfaces across three different Heteropsis clades.

FIGURE 4. Adults in DV (left), VV (right) unless otherwise stated. A: HT ♂ of Heteropsis hazovola Lees & Raharitsimba, sp. nov. (Masoala; NHMUK 010289116). B: PT ♀ of Ht. hazovola (Masoala; NHMUK 010289118). C: Abdomen of HT ♂ of Ht. hazovola (VV, left). D. Abdomen of Mycalesis ankoma LT ♂ (VV, right). E: LT ♂ of Mycalesis ankoma Mabille, 1878 (Madagascar; BMHH (E) 672615). F: PLT ♀ of M. ankoma (BMNH (E) 672618). Scale bar: 30 mm (C – D with separate scale).

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 12. Diagrams of ♂ HW inflated vein system in the Ht. subsimilis group (modified after Lees, 1997: 96); wings in DV and R vein component on the anterior border of the cell in VV (flipped). A: Ht. pauper Oberthür, 1916 (Ankazomivady; “ H. 23 ” in Torres et al., 2001). B: Ht. pauper (Anjanaharibe Sud). C: Ht. kremenae Lees, sp. nov. (Masoala). D: Ht. subsimilis (Butler, 1878) (Ambohitantely), showing M 1 + M 2 inflation and asymmetric swelling above cell. E: Ht. avaratra Lees & Kremen, sp. nov. (Montagne d’Ambre). No wing preparation was available for Ht. sogai.

FIGURE 13. Adults in DV (left), VV (right). A: HT ♂ of Heteropsis kremenae, Lees, sp. nov. (Ambavaony R., Masoala; NHMUK 010289135). B: PT ♀ of Ht. kremenae (Masoala; NHMUK 010289139). C: HT ♂ of Heteropsis avaratra Lees & Kremen, sp. nov. (Montagne d’Ambre; NHMUK 010289141). D: PT ♀ of Ht. avaratra (Montagne d’Ambre, Les Roussettes; DCL-DB- 3385, MNHN).

FIGURE 15. ♂ genitalia in LV (left), DV (right) unless otherwise stated. A: Heteropsis kremenae Lees, sp. nov. (Masoala, Ambanivony; 19 DL). B: Ht. avaratra Lees & Kremen, sp. nov. (315 DL, referred specimen, Joffreville). C: Heteropsis sogai Lees, sp. nov. PT (Tsaratanana; 96 DL, MNHN), LV, SV. Scale bar: 3 mm.

FIGURE 16. Adults in DV (left), VV (right) for mounted specimens and adults in natural posture (photos: D. C. Lees). A: HT ♂ of Heteropsis sogai, sp. nov. (Tsaratanana; DCL-DB- 3386). B: PT ♀ of Ht. sogai (Tsaratanana; DCL-DB- 3387). Scale bar: 30 mm. C: ♀ of Ht. kremenae Lees, sp. nov. (Vohitaly, Makira; 30 / 12 / 2002). D: ♂ of Ht. pauper (Vohitaly, Makira; 30 / 12 / 2002). E: ♀ of Ht. subsimilis (Ambohitantely; 6 / 12 / 2006); F: ♀ of Ht. avaratra Lees & Kremen, sp. nov. (Anjanaharibe Sud; 23 / 11 / 2014).

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.