Heteropsis kremenae Lees

C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, Zootaxa 4118 (1), pp. 1-97 : 46-51

Heteropsis kremenae Lees , sp. nov.

LSID: urn:lsid:zoobank.org:act:D5567C45-4EB9-4E62-8F67-ABF2C4008F4E

Prior references: sp. 14A ( 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, 15.282o S, 50.288o 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, 15.279o S, 50.288o 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 14LO, disturbed forest, NHMUK 010289137 [ QTR barcode]; ♂, NE, Ambavaony R., E. Masoala, 280 m, 15.282o S, 50.288o 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, 15.784o S, 50.2164o E +/- 0.15 km, 45 m, 28/1/1994, D.C. Lees: DL 94-0004, E105 [egg voucher], W7L1-45M 1 LO [fruit trap], NHMUK 010289139 [ QTR barcode]; ♀, NE, Ambavaony R., E Masoala, " 200 m. ", 15.282o S, 50.288o E +/- 0.1 km, 200 +/- 50 m, 28/11/1993; D.C. Lees: DL 93-0009 [ DNA voucher], E4 [egg voucher: “3 greenish eggs expressed 29/11”], NHMUK 010289140 [ QTR barcode]; ♂ ( Fig. 15 A), NE, Masoala E, Ambanivony [=Ambavaony], 15.288o S, 50.288o E +/- 0.5 km, 50 +/- 25 m, site W2-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., 15.295o S, 50.227o E +/- 0.15 km, 275 m, 11/12/1993, H. Raharitsimba: CK93-0020; NHMUK 010289184 [ QTR barcode].

Deposition MNHN: ♂, NE, Ambavaony R., E. Masoala, ca, 100 m, disturbed streamside, 15.279o S, 50.288o E +/- 0.12 km, 180 +/- 75 m, 28/11/1993, C. Kremen: DL93-0007; ♂, NE, Antsamanarana R., Masoala E., "~ 50 m, Piste C", 15.307o S, 50.233o E +/- 0.1 km, 115 +/- 75 m, 11/12/1993; D.C. Lees: DL93-0008; ♀, NE, Antafononana lower Anaovandrano R., W7-L2, Masoala E., 15.7458o S, 50.1858o E +/- 0.15 km, 40 m, H. Raharitsimba: DL94- 0 0 0 3, W7L2-40M 2 LO [fruit trap], IA600 [isotope voucher];

Deposition ABRI: ♂, NE, Ambery R., Masoala E., 15.3716o S, 50.4263o E +/- 0.98 km, 26 m, 22/12/1994, D.C. Lees: DL93-0006; ♀, NE, Masoala PN, W7L2, Antafonona, 40 m, fruit trap 1 Hi, riparian, 15.738o S, 50.182o E +/- 1.85 km, 185 +/- 145 m, 17/1/1994, H. Raharitsimba: DL94-0002.

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

Type locality. Madagascar NE, Ambavaony R., R. Onive, E Masoala, 250 m, 15.282o S, 50.288o 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-CuA1 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-CuA1 ocellus with black iris at least spanning veins CuA1-CuA2, light orange ring narrow to wide and sub-hexagonal, sometimes compressed along diagonal from tergal angle to mid costa. FWD space-M1 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 CuA1-CuA2, on FWV space-CuA1 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-1A ocellus much more strongly expressed than on HWD and sub-elliptic, black iris spanning most of vein CuA1-CuA2 and with a concentric yellow ring that spans the whole space-CuA1. 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 M2 and then fairly straight to mid space- CuA2 and angled back to 1A. 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- CuA2, 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, CuA2 inflated vein narrowly swollen to about 2/3, 1A+2A 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 M3 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: 19DL, PT ( Fig. 15 A); Lees (1997: 106, Fig. 7 f; “14A”): 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 2e 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 2e 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:ACW4937 (exemplar BMAD249-15, Marojejy), diverges by 5.47% from that of H. avaratra (cluster number BOLD:AAD0195, exemplar BMAD016- 09) and by about 5.5% from that of H. subsimilis (cluster number BOLD:AAB4493). In the Torres et al., (2001) dataset, a COII sequence for Ht. kremenae (their “ Hen. sp. 14A”) 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, 14.4377o S, 49.7756o 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, 14.4377o S, 49.7756o 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, 14.4548o S, 49.792o 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 14M-0037, IA528 [isotope voucher]; CCDB-02230-E11 [ DNA barcode voucher]; ♀, NE, Marojejy, 700 m, 24/ 1/2014: 11:21, D.C.Lees: DL 14M-0035, IA532 [isotope voucher]; ♂, NE, Sahantaha, Makira, 15.2337o S, 49.5311o E +/- 0.15 km, 500 +/- 130 m, 29/1/2003, D.C. Lees: BMNH (E) #697172 [ DNA voucher]; ♀, NE, Andreketa near 17B 364 steep rainforest with lots of “Tsongolovo” 15.2642o S, 49.5479o 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, 15.2642o S, 49.5479o E +/- 0.75 km, 375 +/- 25 m, 17/1/ 2003, D.C. Lees: BMNH (E) #697950 [ DNA voucher]; specimen, NE, Ambodivoangy forest, 15.29o S, 49.62o E +/- 1 km, 50 +/- 25 m, 4/12/2001; ♂, NE, Ankirindro, 15.2904o S, 49.5474o 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, 15.295o S, 50.227o E +/- 0.15 km, 275 m, 11/12/1993, H. Raharitsimba: TR 89; ♂, NE, Antsamanarana R., Masoala E., 50 m, 15.307o S, 50.233o E +/- 0.1 km, 50 m, 12/12/1993; H. Raharitsimba: TR 93; ♂, NE, Vohitaly, 15.4377o S, 49.5344o E +/- 0.15 km, 28/12/2002, 15:15, D.C. Lees; ♂, NE, Vohitaly, Makira, 15.4379o S, 49.5344o E +/- 0.15 km, 25/12/ 2002, D.C. Lees: DL-SF3, KA552 [=KA-P552, DNA extract number]; ♂, NE, Ambatoavy, Masoala, 630 m, 15.658o S, 50o E +/- 1.26 km, 630 m, 14/2/1993, D.C. Lees: H14-14293-1; ♀, NE, Ambavaony R., E Masoala, 125 m, 15.279o S, 50.288o E +/- 0.12 km, 125 m, 30/12/1993; C. Kremen: E15 [egg voucher], DNA DCL 32 [ DNA voucher]; ♀, NE, Iketry [Iketra], lower Anaovandrano R., W7-L2, Masoala E., 15.7458o S, 50.1858o E +/- 0.15 km, 40 m, 16/1/1994, D.C. Lees, W7L2-40M 2 LO [fruit trap]; ♀, NE, Masoala, 15/1/1994, WG2 [wing voucher], IA33 [isotope voucher]; ♀, NE, Masoala, 16/1/1994, IA36 [isotope voucher]; ♂, NE, Masoala, 50 m, 12/12/1993, H. Raharitsimba, WG6, [wing image], IA39 [isotope voucher]; ♂, NE, Masoala, 26/11/1993, H. Raharitsimba: TR 47, IA47 [isotope voucher]; ♂, NE, Ivontaka R. camp, between Ivontaka N and S. reserves, ca. 90 m, 16.294o S, 49.81o E +/- 0.7 km, 90 +/- 25 m, 3/2/1995; D.C. Lees: DL 95-0003A, H14A3295-1 [leg sample]; specimen, NE, Ivontaka R., ~ 120 m, 16.294o S, 49.81o 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, 16.297o S, 49.785o E +/- 2 km, 230 +/- 130 m, 3/2/ 1995, 11:46–16:10, D.C. Lees: H14A3295-2 [leg sample], IA601 [isotope voucher]; ♀, NE, Ivontaka Sud, 16.294o S, 49.81o E, 90 m, 2/1995, D.C. Lees, IA338 [isotope voucher]; ♀, NE, Ivontaka Sud, path to Marokoto, 16.297o S, 49.785o E +/- 2 km, 230 +/- 130 m, 3/2/1995, D.C. Lees: H14A3296-2, IA339 [isotope voucher].

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 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 14. Adults in DV (left), VV (right). A: HT ♂ of Pseudonympha subsimilis Butler, 1879 (‘ Antananarivo’; BMNH (E) # 674882). B: PLT ♀ of Culapa undulata Oberthür, 1916 (‘ Antsianaka’; BMNH (E) # 674884). C: LT ♂ of Culapa undulata Oberthür, 1916 (‘ Antsianaka’; BMNH (E) # 674883). D: LT ♂ of Culapa pauper Oberthür, 1916 (‘ Antsianaka’; BMNH (E) # 674888). E: LT ♀ of Culapa pauper ‘ sp. or var. ’ pseudonarcissus Oberthür, 1916 (‘ Antakares’; BMNH (E) # 674890). Scale bar: 30 mm.

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 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.