identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
038747324C7CC65D1EB72CBFFA5F22B2.text	038747324C7CC65D1EB72CBFFA5F22B2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis exocellata	<div><p>Heteropsis exocellata group</p><p>A small clade supported by multiple genes (Aduse-Poku et al., 2015, Fig 1; Suppl. S4) for two sampled species, Ht. exocellata and Ht. cowani (Butler, 1880), with an intriguing published topology (albeit with moderate bootstrap support) as sister to all other Malagasy Region Heteropsis . This clade is also morphologically characterised (Lees, 1997) by the following combination of features: HW ocellus strongly expressed in dorsal space-M3 as well as space-CuA1; spatulate valves, toothed mesad near the slightly folded tip which projects beyond the uncus; gnathos with basal curved projection which is unique among Malagasy Heteropsis and developed (at least in Ht. exocellata and Ht. cowani) with its main arm downward arching over the uncus (Lees, 1997: 106). The vinculum is strongly bowed proximad from LV. See Paulian (1951) for variation in the ♂ genitalia of Ht. exocellata (under its synonyms Henotesia aberrans Paulian, 1951 and He. benedicta Paulian, 1951), but see also the phylogeography of Linares et al., (2009: 488–489), who include representatives from the type localities of both nominal taxa and find no evidence for reciprocal monophyly among the respective populations. The biology of the H. exocellata group is unknown, but these are low flying forest interior butterflies likely to be associated with low grasses, which seem rarely attracted to fruit. They tend to occur in localised and bright forest pockets with a very humid aspect such as near streams, up to around 1800 m elevation. Ht. exocellata has been observed hilltopping on a ridge (pers. obs.).</p></div>	https://treatment.plazi.org/id/038747324C7CC65D1EB72CBFFA5F22B2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C7CC65A1EB72934FCB3240A.text	038747324C7CC65A1EB72934FCB3240A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis mimetica Lees and Kremen	<div><p>Heteropsis mimetica Lees and Kremen, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:365D604C-EAE3-46D4-AAD7-B715062FE056</p><p>Prior references: “sp. 72” (Lees 1997: 56, Fig. 7 G, 7H).</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 2 A), Madagascar NE, Anjanaharibe Sud, 14.7502o S, 49.4998o E +/- 0.8 km, 95 +/- 5 m, 27/11/1995, 8:00–16:00, C. Kremen: CK513, DNA [DNA voucher], BMNH (E) 2008-69 [accession number], BMNH (E) 1717105 [QTR barcode].</p><p>Paratypes: Deposition BMNH (accession BMNH (E) 2008-69): ♂, data as HT but: CK510, DL 9676 [DNA voucher], BMAD 261-15 [DNA barcode voucher], BMNH (E) 1717106 [QTR barcode]; ♂, data as HT but CK528, BMNH (E) 1717107 [QTR barcode]; ♂, data as HT but CK511, DNA [DNA voucher], BMNH (E) 1717108 [QTR barcode]; ♂, data as HT but CK529, DCLW-0136 [wing prep.], 278 DL [genitalia voucher], BMNH (E) 1717109 [QTR barcode]; ♂, data as HT but CK559, DNA [DNA voucher; whole thorax used], BMNH (E) 1717110 [QTR barcode]; ♂, data as HT but CK512b, DL 9678 [DNA voucher, whole thorax and abdomen used], BMNH (E) 1717111 [QTR barcode]; ♂, data as HT but: CK512a, DL 9679 [DNA voucher]; BMNH (E) 1717112 [QTR barcode]; ♀ (Fig. 2 B), data as HT but: CK517, DNA [DNA voucher], BMNH (E) 1717113 [QTR barcode]; ♀, data as HT but CK518, DL 9677 [DNA voucher], BMNH (E) 1717114 [QTR barcode]; ♀, data as HT but CK519, 1 egg expressed [egg voucher], BMNH (E) 1717115 [QTR barcode].</p><p>Deposition MNHN: ♂, data as HT but 25/11/1995, 11:00–13:15, C. Kremen: CK474, IA309 [isotope voucher].</p><p>Deposition summary: BMNH (HT ♂, 7 PT ♂♂, 3 PT ♀♀), MNHN (PT ♂).</p><p>Type locality. Madagascar NE, Anjanaharibe Sud, 14.7502o S, 49.4998o E +/- 0.8 km, 95 m +/- 50 m.</p><p>Diagnosis. The combination in Ht. mimetica of the space-M3 as well as space-CuA1 ocellus expressed on the HWD and a white-translucent ocellus ring (hereafter ‘Orng’) extended close to the base of space M3 and CuA 1 in the FWD, forming a rectangle, as in some forms of Ht. turbans, is diagnostic for this species for both known populations. In the closely related allospecies Ht. cowani, the Orng is nearly circular for all known populations. This trait is consistent in all Ht. mimetica .</p><p>Description. Wings. Upperside uniform mid brown, except light brown in the area around the expressed ocelli of HWD where the ventral white area shows through. On FW, space-MI ocellus expressed as conspicuous white spot. Space-CuA1 ocellus has narrow whitish-orange ring set in a somewhat rectangular white-translucent area whose lower proximad corner extends close to the base of space-CuA1, as in Ht. sabas (Oberthür, 1923) (Fig. 2 E), rather than Ht. cowani (Fig. 2 C-D Fianarantsoa material). Only two ocelli are conspicuously expressed in HWD, that of space-M3 and space-CuA1, both subelliptic and surrounded by a similarly shaped light orange ring. HW margin has two conspicuous mid brown Smls (submarginal lines) following the somewhat crenate margin, slightly more crenate than typical in Ht. cowani . Ocellus expression on ventral surface similar but additional one in HW space-Rs of similar size to those of space-M3 and space-CuA1, and in each case surrounded by a pale orange and a wider light brown ring, the three set in a white-translucent and irregularly ‘X’-shaped area extending from the HW costa to mid space-CuA2, one outer arm of the ‘X’ petering out distad of ocellus space-M3. Basal area of HW mid brown, irrorated with areas of whitish scaling, and forming a right angle where the basal pattern meets vein M2. Two bands of light brown and whitish irroration are found in the basal area of the FW together with a large whitish blotch proximad of white spot in space-M1 and tapering towards the costa from around 2/3 along FW costa. Ht. cowani is almost identical in patterning on either surface except for the proximad extension of the whitish part of the Orng, which is fairly circular in Ht. cowani . Variation. Not much variation is evident between individuals of Ht. mimetica and there only one ♂ was dissected. Sexes similar, but ♀ larger, and may have more extensive white area near FW ventral apex.</p><p>Wingspan/fwl: range 38–43.5 (n=5 ♂♂)/ 20.3–24.3 mm (n=7 ♂♂); mean= 40.5 +/- 2.2 SD (n=6 ♂♂)/22.6 +/- 1.3 SD mm (n=8 ♂♂), including referred specimen and HT ♂, 43.5/ 23.5 mm. Range 43–47/ 24.2–25.4 mm (n=3, ♀♀); mean = 44.7 +/- 2.08 SD/ 24.7 +/- 0.62 mm (n=3 ♀♀).</p><p>Androconia: simple mid brown discocellular brush extending only slightly beyond the fork of Rs and M1, underlying patch light ochreous grey and ‘bullet’-shaped (distally truncate to barely beyond the fork), surrounded by glossy dark scales which extend half way to costa and a similar extent in the other direction, as in Ht. cowani .</p><p>Palps: from LV, light brown stripe in middle, bordered by two white stripes, and dark brown striped fringes of scales to edges, the edge potentially brushing the interommatidial setae consisting of longer scales.</p><p>♂ genitalia: 278DL, PT (Fig. 5 A, LV, DV). Tegumen with fairly straight dorsal profile from LV, where it is relatively quadrate (slight proximad notch from DV). Gnathos narrow with small rounded projection distad at base arising almost perpendicular to base of uncus in its porrect position and curving strongly towards its forwardprojecting and pointed tips (from DV gnathos bends in and out strongly from its stout base). Vinculum strongly arched proximad with moderate constriction with tegumen. Valves with strongly rounded ‘shoulder’ at base, broad throughout with valve arm not much less broad than valve base, fairly parallel sided and spatulate, tips extending further than maximum extension of uncus, with an area of spinoid setae on inner face of tip (not as extensive as in Ht. cowani and Ht. exocellata; Lees, 1997: 104) and with tip incurved from DV. Saccus fairly long (longer than in Ht. cowani and Ht. exocellata) and inflated towards tip, while juxta is quite broad and down-lipped proximad. Aedeagus narrowly cupped beyond ostium and quite recurved away from body, fairly thin and parallel-sided.</p><p>Etymology. Refers to the seemingly dual mimetic colour pattern (Fig. 2), on the upperside to Ht. sabas (Fig. 2 E), and on the underside to some forms of Ht. antahala (Ward, 1872) (Fig. 2 F); some Strabena species such as S. consobrina Oberthür 1916 and S. nepos Oberthür 1916 exhibit similar HWV patterns (see e.g. d’Abrera, 1997). In these butterflies, there is a similar configuration on the underside of ocelli set in a mainly white area. Unlike in most of the Ht. strigula group, but as in others of the Ht. exocellata group, and in some other more early diverging clades of Heteropsis, the HW space-M3 ocellus is expressed.</p><p>Discussion. This species was first recognized in the field by Claire Kremen at Anjanaharibe Sud in November 1995 (Lees 1997: 65). All specimens including the one referred specimen from Makira show the same wing pattern morphology, with no tendency to variation towards all known specimens of Ht. cowani in BMNH and MNHN from Fianarantsoa (Fig. 2 C-D), Ranomafana, Andrambovato or other collected localities of author or others (Vohiparara/ Sahavondronina, Ambondrombe, Anjozorobe). In Ht. cowani, the shape of the space-CuA1 ocellus is always with a near-concentric orange ocellus ring. All relevant types were examined. The LT ♂ of Pseudonympha cowani Butler, 1880 is here designated as the specimen (Fig. 2 C) bearing labels “ Lectotype | Type /♂ Pseudonympha cowani Butl. |Male Fianarantsoa Madagascar, Coll. &amp; pres. By W. D. Cowan 8-23.|B.M. TYPE No. Rh 3070 Pseudonympha cowani, Butl. ♂/ Pseudonympha Cowani Butler Type ”; the PLT becomes the ♀ with the same locality data that was photographed by B. d’Abrera 77/78 (d’Abrera 1980: 183). The two syntypes of Culapa houlbertiana Oberthür, 1923:127, pl. 569, figs 4909, 4910 from ‘Fianarantsoa’, were also examined; the ♂ specimen, that is here designated the lectotype (Fig. 2 D), bears the data “ Lectotype | Madagascar Fianarantsoa ex Lamberton, 1922|A servi de modele a J. Culot, pour le No. 4909 de la Pl. DLXIX Vol. XXI. Etudes de Lepidopterologie comparee|Ex Oberthür Coll. Brit. Mus. 1927-3.”. Automatically PLTs become two ♀♀ bearing labels “ Madagascar Fianarantsoa ex Lamberton, 1922|Ex Oberthür Coll. Brit. Mus. 1927-3.”. The nominal species C. houlbertiana is clearly synonymous with Ht. cowani (Lees, 1997: 60–61; Lees et al., 2003: note 31).</p><p>Additional information. DNA divergences: COI-5P cluster number BOLD:ACW4993 (BMAD261-15, CK510) is shared with and 0.052% divergent to that of Ht. cowani (BMAD122-15, DL14Z-050, Anjozorobe), whereas about 10% divergent to Ht. exocellata (FJ666704, 549 bp compared). There is limited evidence so far of haplotype fixation. Where in the 434th and 439th nucleotide position of the DNA barcode, Ht. cowani has a G and a C (as in Ht. exocellata), Ht. mimetica has an A and a C (n= 2 in both cases). This is then a case of virtual barcode sharing with morphological divergence. More detailed genetic studies are required to investigate further this species pair.</p><p>Phylogeny/sister species: sister species is Ht. cowani .</p><p>Ecology and distribution.</p><p>Habitat: found in Makira in a rather open forest structure dominated by Uapaca Baill. (Phyllanthaceae) growing on quartzite substrate, next to a stream (pers. obs.). Similar habitat on quartzite exists in Anjanaharibe Sud, and it is possible that such forest on poor substrates has an unusually light-penetrating nature.</p><p>Behaviour: flies low, apparently mimetic of Ht. sabas in flight, as first noted by Claire Kremen (pers. comm.). All individuals were caught low down along or near the path; Ht. mimetica, like Ht. cowani, seems reluctant to come to fruit bait. The key functional aspect of this whitening seems to be translucence, which in sunspots seems to make the space-CuA1 ocellus stand out in a more (frightening?) way. Translucence may also be an aspect to the wing pattern convergence for the HW, considering the HWV pattern is visible on the upperside.</p><p>Hostplant: unknown, but presumed to be grasses (rather than bamboos), which are also frequented by Ht. cowani .</p><p>Early stages: unknown, apart from one expressed egg (not described here).</p><p>Distribution: replacement species of Ht. cowani endemic to the rainforest of northeast Madagascar and apparently extremely localized, known from only two very localised sites (Fig. 30 A, yellow dots). The species was searched for in its type locality in November 2014, at a similar date to the HT specimen, without success.</p><p>Conservation: highly range restricted, but the species is currently protected by two parks.</p><p>Elevational range: 775–950 m (n=18 incl. observations).</p><p>Referred specimens. ♂, NE Madagascar, Ankiatomboka-Vohibola, descending to river from O3, c 760 m, 15.1726o S, 49.3913o E +/- 1.5 km, 775 +/- 5 m 10/12/2001, 09:46, D.C. Lees: DL 01-175, 267 [= DL 0267, DNA extract number; cytochrome b], BMNH (E) 697383, KA561 [=KA-P561, DNA extract number]; ♀, data as HT but 22/3/1996, H. Raharitsimba: TR 617; ♀, data as HT but CK520.</p></div>	https://treatment.plazi.org/id/038747324C7CC65A1EB72934FCB3240A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C7BC6591EB728FCFDFA2509.text	038747324C7BC6591EB728FCFDFA2509.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis antahala	<div><p>Heteropsis antahala group (subgenera Masoura and Admiratio Hemming, 1964)</p><p>A new species is described here in the Ht. antahala group, and so it is convenient to characterise its nominal species. The LT ♂ of Mycalesis antahala Ward, 1872 is here designated as the specimen (Fig. 2 F) bearing data “ Lectotype | antahala Ward [in pencil]|Plate 12 Upperside Underside|Ex Coll. CHRIS WARD| Ex Oberthür Coll. Brit. Mus. 1927-3.|P.E.L. Viette det. 1968 Mycalesis antahala LECTOTYPE | SYNTYPE of Mycalesis antahala Ward |BMNH(E) # 672567”; a second ♂ becomes PLT (neither bear a locality label, but the type locality is Madagascar; the LT measures 46.5 mm total span, not quite “two inches”). This group of mycalesines endemic to Madagascar now should include the crenate-HW Admiratio Hemming, 1964 as well as the more strongly HWtailed mycalesines traditionally placed in the subgenus Masoura Hemming 1964 (see below). Here, the status of Ht. alaokola (Oberthür, 1916) is reassessed, and this is germane to new descriptions. A few of the species in the Ht. antahala group ( Ht. ankoma (Mabille, 1878), Ht. alaokola, Ht. mabillei (Butler, 1879), and the species described below) are hard to distinguish on the basis of wing pattern alone, while their genitalia are quite distinctive in most cases, but they are nevertheless extremely different in appearance to that of Ht. paradoxa . The last species might be inferred to exhibit miniaturisation and possibly simplification of the ♂ genitalia. The saccus tip to valve tip distance is about 1.9 mm as opposed to 3–4 mm in Ht. antahala group; see also Ht. subsimilis group, in respect of possible breakdown in pre-mating isolating system. It may be necessary to inspect androconial features of ♂♂ closely to assist in identification. This group are canopy specialists attracted to ripe fruit as adults (sometimes feeding on fruit in situ, sometimes descending to the ground) and probably all are associated with forest bamboos and include at least one species with gregarious larvae (C. Kremen, pers. comm.) and another ( Ht. antahala) observed to roost gregariously as adults (Lees, 1997). The ♂♂ indulge in vigorous hilltopping behaviour on ridgetops, sitting atop bushes just like Ht. drepana, and making canopy chases. Unfortunately there are as yet no biological observations on the large and obvious monomorphic Mylothris mimic, Ht. masoura; its female is one of the largest mycalesines globally (around 37 mm fwl). ♂♂ and ♀♀ are generally phenotypically similar except in size, but the species Ht. antahala exhibits a great deal of polymorphism in both sexes and on both wing surfaces. The Ht. antahala group occurs from sea level up to montane elevations of around 2050 m, and Ht. antahala is distributed also in the west and southwest as far as the ‘sacred’ forest of Analavelona.</p><p>Molecular data now provide good support for monophyly of Ht. paradoxa and four sampled members of the Ht. antahala group: Aduse-Poku et al., 2015, Fig. 1; which was submarginally supported (pp=0.91) for the two exemplars in Kodandaramaiah et al., 2010). A few morphological characters also supported this, notably (1) the presence of a brush emanating from near the base of space CuA 2 in the ♂ HW (Lees, 1997: 76) and overlying at least, vein 1A, a character which is only sporadically seen elsewhere, e.g. in the African ‘ Heteropsis ’ peitho (Plötz, 1880) group and in Ht. drepana and (2) the ocelli in hwv space M2, CuA1 and particularly CuA2, where the ventral pattern is pronounced (i.e. not in Ht. masoura, but in H. paradoxa), are usually shifted markedly inward relative to brown Sml, along with a weak to strong proximad inflection of the HWV Mb, particularly in space-CuA2 towards the base of the cell. HW strongly crenate to tailed. The FWD tends to be much darker costad, extremely so in Ht. paradoxa, except in Ht. masoura, which has the upper surface suffused with white. This FWD trend can also be seen in ♀♀ of some Heteropsis (e.g. Ht. andravahana stat. rev.), but not in the ♂♂; in some other species, darker suffusion is concentrated towards the apex in either sex, rather than the costa. The wing pattern of Ht. paradoxa can be interpreted as an extreme ocellus reduction/loss in space CuA1, combined with loss of melanism in the tergad area of the FW (rather than expansion of the white pupil), and adaptation to resting on lichen covered tree trunks and a mid-high canopy lifestyle. It is stated that this clade is specialised to feed on bamboos (Lees 1997: 137), but there are no published life histories.</p></div>	https://treatment.plazi.org/id/038747324C7BC6591EB728FCFDFA2509	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C78C6471EB729E2FC722384.text	038747324C78C6471EB729E2FC722384.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Gallienia alaokola	<div><p>Gallienia alaokola problem</p><p>The left image (#3043) in Fig. 3 A represents the specimen (BMNH(E) 672621) that P. Viette intended to represent the LT of Gallienia alaokola Oberthür, 1916, according to a label under the specimen (but it is certain that Viette never published this). That on the right (#3044) represents the specimen that d’Abrera (1980, 1997) illustrated as [a ♀ of] ‘ M.? alaokola ’. I contend, considering dorsal colouration, ocellus expression in space-M3 and relatively more pointed HW tails, that both specimens in fact represent Ht. mabillei (both surfaces of the HT ♂ of Strabena mabillei Butler, 1879, BMNH(E) 672624, are figured in Fig. 3 C). Moreover, although d’Abrera (1980: 190) had placed a red label next to photograph of “ M. alaokola ”, the accompanying text says “♀ (? type)”, and by his next edition, d’Abrera (1997) had removed the red label. Without DNA data, choice of a ♀ lectotype would be extremely unsatisfactory in this difficult-to-distinguish species group, and thus unlikely to promote stability. It appears that d’Abrera’s (1980) action does not, however, represent a valid lectotypification, since it is not an unambiguous selection of the name-bearing type, considering in particular his qualification of use of the word 'type' by a question mark (Art. 74.5). A proportion of Oberthür’s syntype specimens of Gallienia alaokola from Fianarantsoa and some also from Antsianaka (the current region around RNI Zahamena) have apparently generally shorter HW tails than in the HT of Strabena mabillei Butler (Fig. 3 C) and a much darker upperside. They lack the ocellus expression in both space-M3 and CuA1 of the ♂ HW upperside, with sometimes neither ocellus expressed (this variation in his mixed series is alluded to in Oberthür’s 1916: 205–206 description). Among Oberthür’s 75 syntypes, a LT ♂ specimen of Gallienia alaokola is here therefore chosen to resolve the problem. This LT ♂ is here designated as the specimen (Fig. 3 B) bearing data labels “ Madagascar, Fianarantsoa, Perrot Frères, 2e Semestre 1892|Ex Oberthür Coll. Brit. Mus. 1927-3|Photographed by B. d'Abrera 77/78|BMNH(E) 1717102 [QTR label]”. This is also the “ Masoura ? sp.” illustrated by d’Abrera (1997: 229) in DV/VV, following preference for a previously illustrated specimen (Recommendation 74B of the code), although in this case, not the one that the original author intended. This action is also in the spirit of the Code as it promotes stability and obviates the need to describe a new species within Oberthür’s syntypic series. The majority of the series of Oberthür (1916) ’s G. alaokola STs from “Fianarantsoa” belong to Ht. mabillei, and are then automatically among the PLTs of Oberthür’s nominal species. Although not illustrated by Oberthür (1916), the chosen LT represents the more widespread/ common and generally darker-upperside species (formerly known as sp. 56; Lees 1997: 65), which tends strongly either to lack space-M3 ocellus expression, or to lack expression in both space-M3 and CuA1 on HWD. This species also lacks a ventral abdominal black androconial patch (usually bilateral of midline, sometimes obscured by overlying ochreous scales) that is characteristic of Ht. mabillei (see Lees, 1997: 99) and also, more markedly, Ht. masoura . It (“sp. 56”; Lees, 1997: 105) also has multiply toothed long valves, which do not have such a projected sharply inbent tip, as does the genitalia of Oberthür’s ♂ ‘iconotype’ of G. alaokola (= Ht. mabillei) (Fig. 3 D). This feature is not in fact evident in the HT specimen of Strabena mabillei Butler, although the abdomen is worn and may even not be the original., The strong expression of the space-M3 ocellus in ♂♂ (in contrast to its strong reduction or absence) confirms the identity of specimens currently referred to Ht. mabillei (in fresh specimens, Ht. alaokola, as here fixed, also has a darker dorsal colouration). Note the androconial scales on abdominal sternum third segment and the inwards recurved valve tips characteristic of Ht. mabillei (Fig. 3 D) and also (Fig. 3 A,C) that Ht. mabillei has relatively long tails especially at M3, a lighter brown dorsal colouration, and three or four white stripes on the FWV tending to be more strongly represented as marks on the FWD costa, than the species/specimen here selected to represent Ht. alaokola (Fig. 3 B). It may be noted that samples respectively representing Ht. mabillei and Ht. alaokola (KAP524 [KA-P524] = DL-4-233, Anjozorobe; KAP537 [KA-P537] = DL-02-20, Makira, in the study of Aduse-Poku et al., 2015) are shallowly diverged considering the morphological (mainly androconial and genitalic) differences between these species.</p></div>	https://treatment.plazi.org/id/038747324C78C6471EB729E2FC722384	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C66C6431EB72E14FCDB240A.text	038747324C66C6431EB72E14FCDB240A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis hazovola Lees & Raharitsimba	<div><p>Heteropsis hazovola Lees &amp; Raharitsimba, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:758D1560-AB02-44F5-B0B8-5AAA92715C2F</p><p>Prior references: sp. 53 (Lees 1997: 65).</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 4 A), Madagascar NE, Ambery R., Masoala E., 15.3743o S, 50.417o E +/- 0.15 km, 100 m, W2L3-100 M 12HI [fruit trap], 20/12/1993, H. Raharitsimba: DL 93- 0 0 0 2, KAP19 [=KA-P19; DNA voucher]|IA300 [isotope analysis voucher], BMAD 229-15 [DNA barcode], NHMUK 010289116 [QTR barcode].</p><p>Paratypes: ♂, same data as HT but Hi12 [fruit trap], H. Raharitsimba and D.C. Lees: DL 93-0003, IA614 [isotope analysis], NHMUK 010289117 [QTR barcode]; ♀ (Fig. 4 B), Madagascar NE, Antsamanarana R., Masoala E., [50–520 m], 15.3o S, 50.23o E +/- 0.15 km, 250 m, 11/12/1993, H. Raharitsimba, D.C. Lees: DL 93-0001, IA301 [isotope analysis voucher], BMAD 228-15 [DNA barcode voucher], KA2054 [=KA-P2054; DNA extract number]; NHMUK 010289118 [QTR barcode]; ♂, NE, Antsamananarana W2L 2, 520 m. 9 hi [fruit trap], 15.2867o S, 50.2184o E +/- 0.15 km, 520 +/- 50 m, 9/12/1993, 16:24, D.C. Lees: DL 93-0018; DL 9719 [DNA voucher], BMNH (E) #697866 [DNA voucher; cytochrome b], IA302 [isotope voucher], BMAD 230-15 [DNA barcode voucher], KA2053 [=KA-P2053; DNA extract number]; ♂ (Fig. 5 B), data as above but: 17:02, D.C. Lees: DL 93- 0017; 26 DL [genitalia]; DCLW-0086 [wing prep.]; NHMUK 010289179 [QTR barcode].</p><p>Deposition summary: BMNH (HT ♂, 3 PT ♂♂, PT ♀).</p><p>Type locality. Madagascar NE, Masoala E., Ambery R., 15.3743o S, 50.417o E +/- 0.15 km, 100 m.</p><p>Diagnosis. this species has rather short and less pointed HW tails compared with Ht. alaokola and Ht. mabillei, but not quite as short as Ht. ankoma . ♂♂ of Ht. hazovola have a dense black latero-ventral androconial patch on the abdominal sternum, whereas Ht. alaokola and Ht. ankoma (particularly similar superficially) lack obvious black ventral abdominal scales in the ♂. Ventral wing pattern is very similar to other species in the group including Ht. alaokola, Ht. hazovola, and to a lesser extent, Ht. mabillei and Ht. ankoma; a violet-lilac cast is, however, particularly prominent in the type series of Ht. hazovola . On the dorsum, in fresh specimens, Ht. alaokola has the darkest wing colouration of these species, whereas in Ht. hazovola, the dorsal wing background colour tends to be reddish brown. Ht. mabillei has an even lighter mid-brown dorsal colour than Ht. hazovola, but generally has space-M3 ocellus HWD strongly expressed in the ♂. Ht. hazovola shares with Ht. ankoma a particularly wide dark costal area on the FWD. ♀♀ are hard to tell apart among the three longer tailed species referred to. The tails are slightly more pointed than typical in Ht. ankoma .</p><p>Description. Wings. Dorsal surface smoky brown along costad third of FW to fairly uniform dull reddish brown on the rest of the wings (among described species. only Ht. ankoma has a darker FWD margin). HWD this uniform dull reddish brown colour with a dark brown double Sml, the outermost darkest and more distinct. HW strongly tailed at vein extremities, prominently so at the end of vein M3; tails not quite as pointed as in Ht.</p><p>alaokola, more similar to the configuration in Ht. ankoma where they are relatively short. FW space-CuA1 ocellus weakly expressed. FWD space-M1 ocellus not expressed and even HWD space-M3 ocellus not expressed. The underside background has a strong dark lilac-grey cast. FWV space-CuA1 ocellus expressed as on upperside its black iris nearly spanning veins CuA1-CuA2, but is more strongly marked, as also for the FWV space-M1 ocellus whose black iris also spans the separation of veins M1-M2 and is similarly surrounded by a dull orange Omg, more or less concentric, or slightly wider tergad in the case of that of space-CuA1. On HWV, space-Rs, M3 and CuA1 ocelli are expressed to the greatest and roughly equal extent, and are subelliptic, followed by much smaller ones in space-M1-M2 and CuA2 and 1A; space-Rs-CuA1 ocelli fall in a separate more proximad arc to those of space- CuA2-1A, as typical for Ht. paradoxa + rest of the Ht. antahala group. Highly irrorated pattern of dark brown strigulae and yellow flecks on a lilac-washed light brown background, but the lilac wash is only strongly evident in the HT and one of the PT ♂♂, and to a lesser extent in the PT ♀, which has a generally lighter greyer-brown colouration on either surface. A striking pattern of ventral irrorations (dark strigulae) which are most finely divided proximad of the jagged dark brown Mb, but continue distad of it in both wings except around the ocelli. The very thick Mb is sharply inflexed around the CuA1 ocelli in both wings, quite jagged, strongly concave in space-M3- CuA1, and again in space-1A, fairly straight to mid costa from just below vein M3. FWV Mb forms an angle distad near vein M2, and there is another dark FWV band slightly proximad but along the irroration, the PMb is not distinct in either wing. There is a prominent yellow patch (highlight) at the base of space-M2 just distad of the Mb. Such underside patterns are not particularly special for Ht. hazovola: in particular, Ht. alaokola and Ht. ankoma are very similar and confusable ventrally, but the type material of Ht. hazovola seems more deeply washed with violet colour. Variation. No ocellus expression on ♂ HWD, neither in the HT nor PT ♂♂, except for M3 slightly in one ♂ PT, while the ♀ PT has ocelli expressed in CuA1, and to a much lesser extent in CuA2. However all the known specimens were taken over a few days in mid December into the start of the wet season, so seasonal variation cannot be quantified. ♀ larger, FWD space-CuA1 ocellus much more strongly expressed in ♀ and HWD space-M3 ocellus expressed along with that of space-CuA1 and most weakly that in space-M 2 in the ♀ PT. ♀ PT HWV ocellus expression similar to the ♂ HT but larger, except that that of space-Rs is lacking. The HWV Mb is more smoothly irregular in the ♀ specimen.</p><p>Wingspan/fwl: range 42.7–43.6/ 22.6–24.9 mm (n=3 ♂♂); mean= 43.3 +/- 0.52 SD (n=4 ♂♂)/23.1 +/- 0.45 SD mm (n=4 ♂♂) including HT♂, 42.7/ 24.9 mm. 44.8/ 25.4 mm (n=1 ♀).</p><p>Androconia: dark brown/blackish Sdbp (supra-discocellular brush and patch). There is no patch on the FWV, nor indeed in any of the Ht. antahala group (the black spot in Fig. 4 A, right, FWV, is a broken wing fragment). Blackish brush emanating near base of space-CuA1, as usual for Ht. paradoxa (very weakly expressed) + Ht. antahala group, over reflexed veins 1A+2A + 3A, and potentially contacting a broad and conspicuous black androconial scale area covering abdominal tergites approximately A3-A6 (not evident though in Ht. paradoxa, despite a weakly developed hindwing brush arising in the cubital region), that is present in Ht. hazovola (Fig. 4 C, PT ♂) but not evident in Ht. ankoma (Fig. 4 D, LT ♂ of M. ankoma) and indeed in Ht. hazovola it is more extensive and obvious than in others of the group that exhibit ventral blackish androconial scales, namely Ht. mabillei, Ht. antahala and Ht. masoura .</p><p>Palps: penultimate segment from LV, with wide dark grey stripe in middle, elegant cream stripes towards the eyeward edge and on ventral side, and dark brown fringes on borders. There is thus only one cream stripe from LV (the second if evident is very thin), rather than the usual two in Ht. mabillei and Ht. alaokola .</p><p>♂ genitalia: 26DL (PT, Fig. 5 B, LV, DV): tegumen shallowly domed from LV (wide concave proximad notch from DV), with strong brow, tegumen much longer than sickle-shaped uncus with strong ventrad ‘dewlap’-like cusp and angularly raised ‘head’ (characteristic of Ht. antahala group), leading to hook with fairly straight edge (uncus is narrow to hook from DV). Gnathos from small base, sinuate and tapering from LV, uprecurved at pointed tip, very narrow and slightly sinuate with a fairly square frame from DV. There is a fairly narrow waist along division with strongly arched vinculum that also broadens towards the saccus, and ‘shoulder’ at valve base also enlarged and convexly round in other direction, fused with narrow base of valve which is long and fairly evenly thin. Much longer valve than in Ht. ankoma in particular (Lees 1997: 105), protruding well beyond uncus tip at tip, slightly inbent towards midline with a short infacing spine, and a few serrae mesad towards tip (unlike Ht. alaokola; 146DL, Fianarantsoa, illustrated in Fig. 5 C, which has a large number of teeth along the inside edge of the valves). Saccus strongly uprecurved, inflated towards tip. Aedeagus almost the same length as valves, slightly uprecurved towards blunt distal end, and broadening towards a cup-like ostium. Juxta narrow, not prominent proximad.</p><p>Etymology. From ‘ hazovola’ (Money Wood in Malagasy, vernacular name for Dalbergia L. spp., ‘palissandre’ or ‘bois de rose’) is an ironic reference to the rosewood plundering and resulting degradation of the primary forest of Masoala National Park, in particular, to which this new species is apparent endemic, that has been brazingly allowed to take place since the coup in 2009, and still apparently continues following the installation of a democratically elected government in a clandestine manner.</p><p>Discussion. This species was first collected in the field by DCL and Heritiana Raharitsimba on 9/12/1993 at R. Antsamanarana, in the survey of the area that would become Masoala National Park (Lees, 1997: 65). All relevant types in the Ht. antahala group were examined and are clearly distinct (HT of Strabena mabillei Butler and four ♂ and two ♀ STs of Mycalesis ankoma Mabille, 1878 in BMNH, plus an additional ST in MNHN ex Chris Ward from the Galichon collection, DCL-DB-2982). A LT has already been designated for Gallienia alaokola (see above), and as there is more possibility for confusion in this group, it is necessary to select another LT. The ♂ LT of Mycalesis ankoma is here designated as that illustrated here, Fig. 4 D–E (there are several very similar STs but it is possible this is also the ♂ illustrated in Pl. 6 Figs 11, 12 of Mabille [1885]), bearing the labels “ ankoma ♂ [apparently in Mabille’s handwriting; no other ♂ bears this label]| Madagascar |Ex Musaeo P. Mabille 1923|Ex Oberthür Coll. Brit. Bus. 1927-3”|BMHH(E) 672615. The apex to apex dimension of this specimen is 48 mm, exactly as mentioned for wingspan in Mabille’s ([1885]) redescription, so it is likely he had this specimen at hand during the original description. A ♀ PLT of M. ankoma from the Grose-Smith collection (BMNH(E) 672618) is illustrated in Fig. 4 F. Ht. hazovola now constitutes the first described butterfly species apparently endemic to the Masoala National Park (Kremen et al., 2001).</p><p>Additional information. DNA divergences: COI-5P cluster number BOLD:ACW4994 (exemplars BMAD228-15 to BMAD230-15 from Masoala), 5.02% divergent to a still undescribed species (BMAD147-15, DL95-0005, Vohitraseva) which is in turn about 2–2.4% divergent to other members, whether conspecific or not, of cluster BOLD:ACW4938. Ht. hazovola appears more closely related to Ht. mabillei in fact than to Ht. alaokola (see above regarding the shallow divergence of exemplars in Aduse-Poku et al., 2010) but only 140 bp are currently available (BMAD131-15, DL14M0-0003, Anjozorobe, for which there is a G rather than A or T in the 517th nucleotide position) and this stretch is 3.1% divergent from the barcode of the HT ♂ of Ht. hazovola (BMAD229-15, DL93-0002).</p><p>Phylogeny/sister species: the sister taxon of Ht. hazovola (as “sp. 53”) was unresolved in Lees (1997), but at least one character suggested a relationship with Ht. ankoma and Ht. alaokola (“sp. 56”; Lees 1997: 168). Sister taxon supported as Ht. ankoma (Aduse-Poku et al., 2016, in press).</p><p>Ecology and distribution.</p><p>Habitat: lowland tropical rainforest.</p><p>Behaviour: comes into canopy fruit traps.</p><p>Hostplant: unknown, presumed to be a bamboo.</p><p>Early stages: unknown.</p><p>Distribution: endemic as far as is known to the eastern lowlands of the Masoala Peninsula (Fig. 30 A, light blue dots).</p><p>Elevational range: 15– 531 m. (n=11 incl. referred specimen and observations).</p><p>Conservation: the lowland rainforest of eastern Masoala where this species survives is under severe threat from illegal logging.</p><p>Referred specimen: ♀, NE, Andranomainty, E. Masoala, 15.7825o S, 50.2979o E +/- 0.15 km, 9/12/1993, H. Raharitsimba: E112 (egg voucher); 247DL (dissection).</p></div>	https://treatment.plazi.org/id/038747324C66C6431EB72E14FCDB240A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C62C6401EB728FCFB6F2681.text	038747324C62C6401EB728FCFB6F2681.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis drepana	<div><p>Heteropsis drepana group (including subgenus Henotesia Butler, 1879)</p><p>The Ht. drepana group (comprising the ‘ narova -group’ of Torres et al., 2001, otherwise known as subgenus Henotesia sensu Lees et al., 2003, but referred to here as the ‘ Henotesia clade’, with the addition of H. drepana and Ht. fuliginosa; see Aduse-Poku et al., 2015) also includes the prior genus Houlbertia. The type species of the last genus is Houlbertia erebennis Oberthür, 1916, as selected by Viette, 1961, which has priority over Hemming’s, 1964, selection of Erebia passandava Ward, 1871 as the type species; NHM’s Lepindex needs to be updated in this regard. The (subgenus) Henotesia as circumscribed by Lees et al., (2003), who synonymised Houlbertia with it, is based on He. wardii Butler, 1879, which became on combination a junior subjective homonym of Ht. wardii (Mabille, 1877) . The Ht. drepana group, or at least the Henotesia clade, is supported by both morphological (Lees, 1997) and molecular data (e.g. Kodandaramaiah et al., 2010). Perhaps the most important morphological feature of the Henotesia clade is the dorso-ventrally flattened aedeagus (Lees, 1997). However, the entire Ht. drepana group includes moderately to highly sexually dimorphic species (Lees, 1997). Aduse-Poku et al., (2015, Suppl. Fig. S4) using ten genes found that their 11 sampled members of the Henotesia clade, had 92% bootstrap support as sister to Ht. drepana, and this clade 87% support as sister to the hitherto hard to place Ht. fuliginosa . The sister of Ht.</p><p>drepana group is recovered with significant bootstrap support (Aduse-Poku et al., 2016, in press) as sister to a ‘ Ht. iboina group’ of Malagasy Heteropsis that included Ht. vola, that was previously (and erroneously) placed in (subgenus) Telinga by Lees et al., 2003. The iboina group also comprised Ht. iboina (Ward, 1870) + Ht. parva (Butler, 1879) and members of the Ht. avelona group (= bicristata group of Torres et al., 2001), including Ht. avelona (Ward, 1870) + Ht. uniformis (Oberthür, 1916), Ht. parvidens (Mabille, 1880), and Ht. bicristata (Mabille, 1878) . No undescribed species are known in the Ht. iboina group. In general, the question of the number and limits of subgenera in Heteropsis is left open here. If the name Houlbertia Oberthür was restricted to a clade of highly sexually dimorphic species that include Ht. narova, excluding Ht. passandava according to Aduse-Poku et al., (2015, Fig. 1), with Henotesia Butler retained as a separate subgenus, this would leave Ht. passandava without a subgeneric name. While more faithful to traditional taxonomy in Madagascar, this also seems complicated and a pragmatic solution might be to synonymise both Henotesia and Houlbertia with the nominotypical subgenus Heteropsis Westwood, perhaps with the addition of the previously enigmatic Ht. fuliginosa, forming together the well-supported clade in Aduse-Poku et al., (2015, Fig. 1). However, such an action, that would abandon names in prior use as historic curiosities, awaits a forthcoming study. It should also be noted that without explicit justification (perhaps merely because it seemed different to other African hairy-eyed mycalesines), d’Abrera (1997: 226) included the African species ‘ peitho Plötz, 1880’ in ‘ Houlbertia ’ (but see new genus in Aduse-Poku et al., 2016, in press).</p><p>Species of the Ht. drepana group with the most vivid blue upperside ♂♂ such as Ht. erebennis occur at lower elevations than those with dark ♂♂, further discussed below. The last group tend to be found in more montane areas, up to over 2300 m, in the case of Ht. pauliani, which is as high as any mycalesines are found in Madagascar. ♂♂ of most species are quite rarely attracted to ripe fruit and Ht. andravahana stat. rev. (see also below) has been observed to feed on flowers of Dichaetanthera Endl. ( Melastomataceae; Lees, 1997), whereas the opposite sex is regularly attracted to fruit. Adults sometimes feed on in situ fruits. Species fly from understory to lower canopy, males following lightgaps, sometimes resting on top of clumps of bamboo, whereas Ht. drepana males hilltop and rest on top of ridge bushes in sunny weather. Many if not all species seems to be closely associated with different types of forest bamboo, but early stage data is very sparse. Stands of bamboo (such as of the giant Cathariostachys S.Dransf. in the lowlands) can sometimes support large sympatrically flying populations of several closely related species. ♀♀ of the more sexually dimorphic members of the Ht. drepana group tend to be very sedentary resting low in the undergrowth on leaf litter (where they are cryptic with fallen leaves including those of bamboo), and may not be noticed at all when not venturing off paths. The ♀♀ of Ht. passandava itself closely resemble those of the distantly related Ht. iboina (Ward, 1870) and Oberthür (1916) not only described its ♀ as a form (fitensis) of ‘ iboina ’, but in a different genus than Culapa Moore, 1878, as Henotesia andravahana var. macrophthalma Oberthür, 1916, in which he also included ♀♀ of Ht. passandava along with Ht. obscura (Oberthür, 1916; see Lees, 1997). The more montane members of the Ht. drepana group have ♀♀ that tend to be vertical trunk-resting (as do often the ♂♂, likely explaining their less sexually dimorphic wing patterns).</p><p>Within the Ht. drepana group are two or more groups of phenetically similar, moderately sexually dimorphic species, which inhabit montane forest ridges between 1000–2000 m. Two sampled members in the phylogeny of Aduse-Poku et al., (2015, Fig. 1) are Ht. viettei Lees, 2003 and Ht. ‘ andravahana ’ (= Ht. westwoodi sp. nov., see below), which were sister to Ht. obscura + Ht. difficilis (Mabille, 1880) (= Ht. andravahana stat. rev., see also below). Unlike the last two species, the ♂♂ of the first pair have dark, non-reflective uppersides, and correspond to sp. 13 (‘ Henotesia sp. 13’ Kremen, 1994: 417; now including sp. 74), and sp. 13B, respectively, of Lees (1997: 64). This species pair exhibit varying degrees of whitish or cream highlighting distal of the HWV Mb. The Ht. harveyi sub-group of the Ht. drepana group represents a tightly knit species complex with dark-uppersides and includes Ht. harveyi sp. nov. and Ht. vanewrighti sp. nov. (sp. 12, and sp. 63, respectively), as well as at least one undescribed species (sp. 20, which is the same as sp. 77; see Lees, 1997; Torres et al., 2001). The Ht. harveyi sub-group exhibits the presence or absence of a russet or dark patch on the underside of the FW along vein 1A+2A (in Madagascar, only the Ht. avelona group of mycalesines also exhibit a FWV patch). Another dark non-reflective upperside pair, more similar to Ht. viettei + Ht. westwoodi, although not represented in the phylogeny of Aduse-Poku et al., (2015), and so with relationship unresolved, is Ht. imerina sp. nov. (sp. 22; same as sp. 76 of Lees, 1997; Torres et al., 2001) and Ht. pauliani (sp. 37 of Lees, 1997). In order to describe these five new species, it is necessary to go into some detail first to clarify the identity of Mycalesis andravahana Mabille, 1878, which potentially blurs the identity of some of these species, and then to lectotypify it.</p><p>Mycalesis andravahana problem. Although Mabille’s original (1878) short unillustrated description of Mycalesis andravahana, in Latin, does not mention the number of specimens, citing only Madagascar as type locality, the first and main part of the short description matches at least the (apparently lost, see below) ♂ specimen illustrated in Mabille ([1885]: Pl. 5, Fig. 6.7). However, this first part does not match the ♂ of the taxon represented by the “var.” in the latter work and illustrated in Mabille [1885]: Pl. 5, Fig. 8). The additional details in Mabille ([1885 -7]) suggests that the original description had been based on at least three phenotypically divergent specimens, two ♂ and one ♀ (that illustrated dorsally by Mabille, [1885]: Pl. 5, Fig. 9).</p><p>Despite several searches over two decades and no doubt by previous potential authors interested in resolving the problem (including P. Viette, who had planned to revise the Malagasy satyrines; all of his ms lectotypes of this group are unpublished), no possible ♂ syntypes of Mycalesis andravahana had been traced in the three main feasible repositories for Mabille material, BMNH and MNHN (DCL, pers. obs.), or ZHMU (W. Mey, pers. comm.). Although the ♂ (Fig. 6A, ‘6, 7’) might have been a good choice for the lectotype of Mycalesis andravahana Mabille, 1878, unfortunately both it and the “var.” appear to have been lost or destroyed. However, Lees (1997: 33, 62) and Lees et al., (2003: [789], note 45) already suggested a possible solution to the vexing problem of typification of this nominal taxon by fixing as LT a ♀ specimen that came from the Henley Grose-Smith collection (via the Joicey bequest to BMNH). This specimen bears a historical rectangular “ TYPE ” label, without labelling of its type identity and is now numbered as BMNH(E) #674841. Because of its wide and distinctive HWV white postmedial band, which is strongly indented around the HWV space-CuA1 ocellus, this specimen had been thought (Lees et al., (2003: [789], note 45; where “ H. 13” was a lapsus for ‘ H. 13B’), to correspond to ‘sp. 13B’ (described below as Ht. westwoodi, a species apparently endemic to the southeastern mountains of Madagascar, from Andohahela to Ranomafana and Andringitra National Parks).</p><p>A close re-examination of this specimen (BMNH(E) #674841) now reveals diagnostic features (a highly crenate HW, and HWV ocelli, apart from that in space-CuA1, expressed as strong white spots, as well as a strongly indented near-right-angle of the Mb around CuA1), that together identify it as a potentially dry season form of a species that was also described by Mabille as Mycalesis difficilis Mabille, 1880 . This last taxon is illustrated in Mabille, ([1885], Pl. 7B, ‘2’, ‘2a’) and reproduced here (Fig. 6A, ‘9’). M. difficilis is based on a single individual, for which a matching ♀ HT (DCL-DB-2872), previously unlabelled as a type, was already located in MNHN (Lees 1997: 389; Lees et al., 2003). Its (metallic blue upperside) ♂ was described by Oberthür 1916 as Henotesia undulosa (together with a non-reflective form¸ var. luctuosa), and its ♀ described independently in the same work as He. andravahana var. macrophthalma (see Lees, 1997). Although it is has not previously been clear what species is represented by the ♀ whose dorsal surface only is illustrated in Mabille ([1885]: Pl. 5, Fig. 9) as an example of Mycalesis andravahana, indeed it cannot be ruled out that this illustrates (considering its small ocelli), a potentially dry seasonal form of Mabille’s nominal species M. difficilis . In fact the resemblance to a recent photograph of a living ♀ from Anjanaharibe Sud is striking (Fig. 6 C). About the original ♀ specimen, Mabille ([1887]: 58) stated “La femelle est d'une couleur plus claire que le mâle, surtout sur le disque. L'ocelle des ailes supérieures est très grand, d'un fauve clair; l'ocelle des inférieures est très petit. En dessous, les premières ailes ont les zébrures blanchâtres jusque dans la cellule; l'espace terminal est d'un gris cendré. Aux inférieures, il y a les mêmes dessins, mais plus délayés; la bandelette blanche est d'un gris sale. Les autres parties de l'insecte sont comme dans le type mâle que nous avons décrit plus haut”. Just below this he stated “L'envergure est de 36 millimetres chez le ♂, de 44 chez la ♀. Mycalesis Andravahana à été envoyé a le Henley Grose Smith des parties boisées du nord-est de l'île”. The HT of Mycalesis difficilis had the same provenance (Mabille, 1880), but as implied above, would represent a likely wet seasonal form of the same species. The potential ♀ ST of M. andravahana coming from the Grose-Smith collection (BMNH(E) #674841; Fig. 6 D) measures 50.2 mm absolute wingspan and between 23–23.3 mm FWL. However, all four wings are fixed to the body with the FW tergal edges angled in a non-orthogonal orientation, with blobs of red sealing wax. My digital reconstruction of the wings shows that the apex to apex dimension, when reconstructed as Mabille ([1885]: Pl. 5, Fig. 9) actually would measure 44– 45 mm apex to apex. It should be acknowledged that in general, Heteropsis are in general hard to identify from uppersides, more so for a painting of a ♀. Oberthür (1916: 205) was convinced that it represented the ♀ of the taxon/taxa in his collection that he referred to as “ wardii - andravahana ”. This usage might reflect his insight into the likely compound nature of Mabille’s description (and redescription) of M. andravahana; indeed, Oberthür labelled one specimen (now numbered BMNH(E) 647848) with the above compound taxon name. Perhaps (or not), Oberthür was aware of the underside pattern of the specimen in the Grose-Smith collection (whether or not he ever visited the Grose-Smith or the Joicey collection, housed in Surrey, where the specimen would have gone in 1910).</p><p>Oberthür nevertheless named his form ‘ macrophthalma ’ as a “var.” of “ Henotesia andravahana ”. In any case, it now transpires that the only surviving potential syntype of Mycalesis andravahana (BMNH(E) #674841) that was previously located (Lees, 1997, Lees et al., 2003) does not disagree with Mabille’s ([1885-7]) redescription in any significant aspect. The dorsal illustration would indeed represent a quite accurate portrayal of the original specimen. There is indication, especially on the non hand-coloured copy of Mabille’s work available on Biodiversity Heritage Library (Fig. 6 B), of the pale HWV postmedian area showing through to the dorsal surface, just as it does in BMNH(E) #674841, and as clearly indicated in the illustration of Mycalesis difficilis in Mabille ([1885: Pl. 7B) (Fig. 6A). The relative size of dorsal FW and HW ocelli and their rings in these two versions of Mabille’s illustration (Pl. 5, ‘9’) is also convincingly accurate as corresponding.</p><p>The specimen labelled “BMNH(E) #674841” is therefore now designated as the lectotype ♀ of Mycalesis andravahana Mabille 1878 .</p><p>As mentioned above, Oberthür (1916) also included ♀♀ of what has been known (Lees et al., 2003) as ‘ Ht. difficilis ’, as well as of Ht. obscura and Ht. passandava, in his series of Henotesia andravahana var. macrophthalma . As it was originally curated (pers. obs.), Oberthür’s series of “ wardii - andravahana ”, however, came mainly from Fianarantsoa, with a few specimens from Antsianaka, and essentially comprised ♂♂ and a few ♀♀ of just one species, previously known as morphospecies 12 (‘ Henotesia ? obscura (sp. 12)’ of Kremen, 1994: 417) from Ranomafana; actually, the unrelated Ht. obscura, which like the nominal species Henotesia undulosa Oberthür, 1916, has dull metallic blue upperside ♂♂, is not known from the Southeast of Madagascar). ‘Sp. 12’, as known from Ranomafana, is now described below as Ht. harveyi (a manuscript name of Lees, 1997). Ht. harveyi not only has ♂♂ with very dark brown uppersides, but also lacks the distinctly white postmedial band which as mentioned above is present in Ht. viettei . It is not certain if the species without HWV whitish postmedian highlighting represented as “var.” of Mycalesis andravahana in Mabille’s description and redescription (1878; [1885]: Pl. 5, Fig. 8; [1887], text) and reproduced here in Fig. 6A is referable to Ht. harveyi, but it might be. However, Aurivillius (1911) described a similar ‘var.’ lacking the white postmedian HWV band as ‘ Henotesia andravahana ab. marmorata ’ (specimen represented in Fig. 7 C). Its HT ♂ (‘TYPUS’, specimen NHRS- TOBI000000050) was examined by DCL in SNHM. This is an unavailable name (clearly infrasubspecific, Art 45.6.2), but the specimen matches well Ht. harveyi, in fact much more clearly than does that illustrated in Mabille ([1885]: Pl. 5, 8). As judged by Mabille’s [1885] plate, Ht. harveyi may have been one of the species under Mabille’s eyes when he described M. andravahana, even though not the first mentioned, or focal, form of the description. In Ht. harveyi, although the ventral irrorated pattern and general colouration resemble the ventral illustration in Mabille ([1885]: Pl. 5, Fig. 8), generally the median and pre-median HWV bands are much more clearly delineated. This is the case in the HT of Aurivillius’ aberration, and in most specimens of Ht. harveyi, but not so for Mabille’s ♂ ‘var.’ (Fig. 6A, ‘8’).</p><p>To resolve the Mycalesis andravahana problem, the following synonymy of Heteropsis andravahana is thus proposed:</p><p>Heteropsis andravahana (Mabille, 1878) . LT ♀, here designated (Fig. 6 D), lacking abdomen (the only known surviving potential ST, from the Grose-Smith collection): BMNH(E) #674841, bearing labels “LT (round purplebordered label)/ Type HT (round red-bordered label)/ TYPE. [white red printed label]/Ex Grose-Smith, 1910/Joicey Bequest Brit. Mus. 1934-120./[Colour photocopy of Fig. 9 from plate 5 of Mabille, 1887]”;</p><p>Mycalesis difficilis Mabille, 1880, syn. nov. HT ♀ (Lees et al 2003: note 42, [p. 789]): MNHN (DCL-DB- 2872);</p><p>Henotesia undulosa Oberthür, 1916, syn. nov. LT ♂, here designated: BMNH(E) #674825, LT (round purplebordered label)/ Type (round red-bordered label)/ Madagascar, Antsianaka, Perrot Freres, 2e Semestre 1893/Ex Oberthür Coll. Brit. Mus. 1927-3/[copy of f. 2830 in Oberthür, 1916];</p><p>Henotesia undulosa var. luctuosa Oberthür, 1916, syn. nov. LT ♂, here designated (of 4 potential STs), lacking abdomen: BMNH(E) #674827, bearing labels “LT (round purple-bordered label)/ Type (round red-bordered label)/ Madagascar, Antsianaka, Perrot Freres, 2e Semestre 1893/Ex Oberthür Coll. Brit. Mus. 1927-3/ Gallienia undulosa var. luctuosa Obthr. [label in Oberthür’s handwriting]”;</p><p>Henotesia andravahana var. macrophthalma Oberthür, 1916, syn. nov. LT ♀, here designated, lacking abdomen (of 34 original STs, mixed series): BMNH(E) #674826, bearing labels: “LT (round purple-bordered label)/ Type (round red-bordered label)/ Madagascar, Antsianaka, Perrot Freres, 2e Semestre 1893/Ex Oberthür Coll. Brit. Mus. 1927-3/[copy of f. 3042 in Oberthür, 1916]”;</p><p>The above lectotypification releases the four montane species below to be described anew.</p></div>	https://treatment.plazi.org/id/038747324C62C6401EB728FCFB6F2681	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C6DC6481EB72DBAFBA8218A.text	038747324C6DC6481EB72DBAFBA8218A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis harveyi Lees & Kremen	<div><p>Heteropsis harveyi Lees &amp; Kremen, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:CD7A88FE-87A6-4324-8404-5EDE47D8E9F2</p><p>Prior references: Mycalesis andravahana Mabille ♂ “var.” (Mabille [1885]: Pl. 5 Fig. 8, [1887: 57–59]);</p><p>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;</p><p>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);</p><p>sp. 12 (Kremen, 1994: 417, as “ Henotesia ? obscura ”; Lees 1997: 61, 64; Torres et al., 2001: 462);</p><p>“ Heteropsis (Henotesia) harveyi Lees (MS), nom.nov.” (ms name in Appendix A of Lees, 1997: 391).</p><p>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].</p><p>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];</p><p>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;</p><p>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;</p><p>Deposition PBZT: ♂, same data as HT but 8/11/2014: 11:27, A. Allaoui: DL14R-213.</p><p>Deposition summary: BMNH (HT ♂, 3 PT ♂♂ and 2 PT ♀♀), MNHN (2 PT ♂♂ and 2 PT ♀♀); ABRI (1 PT ♂ and 1 PT ♀); PBZT (1 PT ♂).</p><p>Type locality. Madagascar E, Sahavondronina VOI, Ranomafana National Park, 21.276o S, 47.325o E +/- 0.25 km, 1313 +/- 25m.</p><p>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.</p><p>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.</p><p>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).</p><p>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.</p><p>Etymology. After Don Harvey, who first pointed out (to Claire Kremen) the FWV androconial patch (Lees 1997: 64).</p><p>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.</p><p>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.</p><p>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).</p><p>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).</p><p>Ecology and distribution.</p><p>Habitat: montane rainforest with fine-leaved climbing bamboos.</p><p>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.</p><p>Hostplant: suspected to be fine-leaved climbing bamboos.</p><p>Early stages: unknown.</p><p>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.</p><p>Elevational range: 700–2030 m. (n=86, including referred specimens and observations; 1087–1590 m. for DNA barcode-verified specimens).</p><p>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. &amp; 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.</p></div>	https://treatment.plazi.org/id/038747324C6DC6481EB72DBAFBA8218A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C69C6751EB72C61FB15228C.text	038747324C69C6751EB72C61FB15228C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis vanewrighti Lees	<div><p>Heteropsis vanewrighti Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:6CE17307-FD07-47EB-A07A-70C0C94D3FB8</p><p>Prior references: sp. 63 (Lees 1997).</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 9 A), Madagascar NW, Bekolosy, RS Manongarivo, 1280 m, low stature forest, 14.0295o S, 48.3095o E +/- 1 km, 1280 +/- 50 m, 18/12/1994: 10:30, D.C. Lees: DLBEK 94_173, NHMUK 010289124 [QTR barcode];</p><p>Paratypes: Deposition BMNH: ♀ (Fig. 9 B), Madagascar NW, Bekolosy Mt., RS Manongarivo, 14.0427o S, 48.3028o E +/- 0.25 km, 1250 +/- 50 m, 23/12/1994, D.C. Lees: DLBEK 94_175, NHMUK 010289125 [QTR barcode]; ♂, NW, Bekolosy Mt., RS Manongarivo, 14.045o S, 48.306o E, 1162 m, at fruit, 22/12/1994, D.C. Lees: DLBEK 94_130 [barcoded specimen: CCDB-02225-F04; BMAD 064-09; HM404241], IA219 [isotope voucher], BMNH (E) #697875, KA579 [=KA-P579, DNA extract voucher]; ♂, NW, Bekolosy, RS Manongarivo, Bekolosy, 1162 m, at fruit, 19/12/1994: 10:45, D.C. Lees: DLBEK 94_163, IA328 [isotope voucher], NHMUK 010289126 [QTR barcode]; ♂, NW, Bekolosy, RS Manongarivo, 1200–1230 m, 14.034o S, 48.31o E +/- 1 km, 1215 +/- 50 m, 20/12/1994, 09:40–10:18, D.C. Lees: DLBEK 94_182, KA2055 [=KA-P2055; DNA extract number], NHMUK 010289127 [QTR barcode]; ♂, NW, Bekolosy, RS Manongarivo, 1215 m, 14.034o S, 48.31o E +/- 1 km, 1215 +/- 50 m, 16/12/1994, 07:00–07:52, D.C. Lees: DLBEK 94_162; IA329 [isotope voucher], NHMUK 010289128 [QTR barcode]; ♀, NW, Bekolosy, RS Manongarivo, ridge ~ 1250 m medium stature forest, 14.0295o S, 48.3095o E +/- 1 km, 1295 +/- 50 m, 18/12/1994, D.C. Lees: DLBEK 94_167, 1 greenish egg expressed 18/12: E53, NHMUK 010289129 [QTR barcode]; ♂, NW, Bekolosy Mt., Manongarivo RS, 1200–1300 m, 14.0295o S, 48.3095o E +/- 1 km, 1250 +/- 50 m, D.C. Lees: 233 DL [genitalia], DL 9703 [DNA voucher], IA466 [isotope voucher], NHMUK 010289181 [QTR barcode];</p><p>Deposition MNHN: ♂, Madagascar NW, Bekolosy, RS Manongarivo, slope above camp, 1215 m, 14.034o S, 48.31o E +/- 1.5 km, 1215 m, 16/12/1994, 07:34, D.C. Lees: DLBEK94_171; ♂, NW, Bekolosy, RS Manongarivo, near camp, 1163 m, 14.04575o S, 48.29623o E +/- 0.5 km, 1130 +/- 35 m, 12/12/1994: 08:00–09:00, D.C. Lees: DLBEK94_176; ♂, data as above but: DLBEK94_172; ♂, NW, Bekolosy, RS Manongarivo, 1250 m, 14.0427o S, 48.3028o E +/- 0.25 km, 1250 +/- 50 m, 18/12/1994: 09:20–10:30, D.C. Lees: DLBEK94_161;</p><p>Deposition ABRI: ♂, Madagascar NW, Bekolosy, RS Manongarivo, 1250 m, 14.0427o S, 48.3028o E +/- 0.25 km, 1250 +/- 50 m, 12/12/1994, 10:16–12:08, D.C. Lees: DLBEK94_168;</p><p>Deposition PBZT: ♂, Madagascar NW, Bekolosy Mt., summit, RS Manongarivo, Madagascar NW, 1487 m, 14.0275o S, 48.3048o E +/- 0.5 km, 1482 +/- 25 m, 19/12/1994, D.C. Lees: DLBEK94_118, IA602 [isotope voucher]; ♂, NW, Bekolosy, RS Manongarivo, 1350–1400 m, 14.0304o S, 48.3049o E +/- 0.16 km, 1375 +/- 25 m, 19/12/1994, D.C. Lees: DLBEK94_111.</p><p>Deposition summary: BMNH (HT, 5 PT ♂♂, 2 PT ♀♀), MNHN (3 PT ♂♂), ABRI (PT ♂), PBZT (2 PT ♂♂).</p><p>Type locality. Madagascar NW, Bekolosy, RS Manongarivo, 14.0295o S, 48.3095o E +/- 1 km, 1280 +/- 50 m.</p><p>Diagnosis. Especially in ♂♂, Ht. vanewrighti is the smallest species of Malagasy Heteropsis so far known (only in Africa are found smaller mycalesines such as Bicyclus nobilis (Aurivillius, 1893): Condamin, 1973). It can normally be distinguished from Ht. harveyi by the lack of an androconial patch on the FWV, and by its restriction, as presently known, to the extreme northwestern part of the rainforest biome. In Marojejy, northeastern Madagascar, I found a population of a putatively undescribed species (equivalent to “ Hen. sp. 77” of Torres et al., 2001), which could be confused with Ht. vanewrighti, which sometimes exhibits extremely reduced FWV patch. However, counting from the start of the standard 358 bp COI-5P barcode, a barcoded individual of Ht. vanewrighti from Bekolosy (CCDB-02225-F04, BMAD064-09) has the following five potentially autapomorphic nucleotide states relative to other members (in parentheses) of the Ht. harveyi group: 268, 292, 547: T(C); 352: A(G); 451: C(T).</p><p>Description. Wings: Upperside dark brown, darkening towards FW costa and apex. The only ocellus strongly expressed is that of FWD CuA1, whose round black iris spans vein CuA1-CuA2 and has a very dull reddish-orange and fairly narrow concentric ring. HWD ocellus space-CuA1 faintly indicated as a black point, white pupil barely visible. A dark brown line flanked by slightly paler mid brown follows the strongly crenate HW border and the FW margin is also quite crenate. Underside mid brown with ocellus space-CuA1 a similar size to FWD with black iris also spanning veins CuA1-CuA2, more or less circular with a fairly narrow dull orange concentric ring. FWV space-M1 ocellus quite small, with black iris and faint very narrow orange ring. HWV space-CuA1 ocellus about half size of FWV one and elliptic with narrow dull orange ring. Rs-M3 and CuA2-1A ocelli expressed as minute white dots. Area between PMb and Mb dark brown-edged outside and inside respectively. HWV Mb irregular, a similar shape to Ht. andravahana as revised here, with a near right angle inflexion around vein CuA1, curving convexly around vein CuA2, and sharply convex above vein M2 to mid-costa. Distad of Mb there is mottling of mid and darker brown grey, with irrorated strigulae pronounced as on FWV. Four brown Cbs in FWV cell area do not stand out as a particularly distinct pattern from the general brown irroration, but tend to be parallel to one another and perpendicular to costa. A wavy Sml is indistinct but flanked distad with paler mid brown, while the distalmost Sml is dark brown and more distinct, following the crenate margin. Five short and convex-subtriangular dark brown tails at vein ends are interspersed with a few paler brown scales elsewhere in the fringe. Tails at M3- CuA2 are slightly more strongly produced than the others. Variation: ♂♂ similar, but on ventral surface, ocelli are sometimes expressed by stronger white dots (notably those of space-Rs-M3), and FWV space-CuA2 ocellus often includes a small black iris with faint orange ring, as in the ♀. ♀♀ tend to be larger with a paler mid-brown upperside and space-CuA1 ocellus tends to be slightly elliptic on HWD and strongly expressed in both wings, while FWD space-M1 ocellus is small and has a narrow dull orange ring. FWV space-M1 ocellus relatively large and circular to elliptic with a pale orange ring. HWV space-CuA1 Orng wider, dull red (in at least one ♀). The ventral surface of the ♀ has an especially mottled, ‘bark-cryptic’ pattern, consisting of intense russet brown irroration with ochreous on the HWV that is more strongly contrasting than in the ♂.</p><p>Wingspan/fwl: range 30.2–33.7/ 15.8–17.4 mm (n=8, ♂♂), mean 32.05 +/- 1.4 SD/16.76 +/- 0.56 SD mm (n= 7 ♂♂), including HT ♂= 30.2/ 16.26 mm. Range 30.6–37.1/17.0– 18.7 mm (n=3, ♀♀); mean = 33.78 +/- 3.23 SD/ 17.99 +/- 0.86 SD mm (n=3 ♀♀).</p><p>Androconia: discocellular brush ochreous-grey overlying a lenticular orange-brown patch. There is no androconial patch in FWV near 1A+2A as in related species. Variation not quantified.</p><p>Palps: penultimate segment on outside face with narrow grey stripe, flanked by greyish cream scales, bordered by narrow dark brown scales; inside face greyish-ochreous, flanked by dark brown-black scales; ♀ similar.</p><p>Male genitalia (Fig. 8 B, 233DL, PT): from LV: tegumen shallowly domed (with small concave proximad notch from DV), with fairly distinct brow down to scythe-hooked uncus. Gnathos fairly straight, thin and tapered, in line with up-hook of uncus in specimen examined (slightly incurved and not sinuate from DV bending round from stout oblong base). Tegumen-vinculum division slightly more than maximum dimension of tegumen. Valve base fits in rather oblong frame with rather angular ‘shoulder’ at base of dorsal edge, leading abruptly to gently elbowed rather parallel-sided valve arm which has a very slender ‘head’ for club with stout small spine at tip, uprecurved and significantly proud of uncus tip (strongly incurved at tip from DV and with a few short distad teeth). Aedeagus very long and thin, dorso-ventrally flattened, more than one and a half times length of valve, gently and smoothly uprecurved distad from ostium, which is itself long and cupped proximad, winged into two lobes proximad from DV. Saccus relatively long, and juxta featuring a prominent down-lip, which also has small side lobes.</p><p>Etymology. After R.I. Vane-Wright, who published the serious SEM study of mycalesine androconia (Vane- Wright, 1971), revised the subtribe in Sulawesi (Vane-Wright &amp; Fermon (2003)), and who inspired the modern study of the subtribe in Madagascar.</p><p>Discussion. The species was first recognized in the field by DCL as sp. 63 in December 1994 at Bekolosy mountain in the Réserve Spéciale de Manongarivo (Lees: 1997: 65), and subsequently found at the adjacent Mt. Antsatrotro in November 2011. Although of a similar size, the HT of Henotesia wardii Butler, 1879 (= Heteropsis viettei Lees, 2003) has whitish shading distal of the HWV Mb. The only similar specimen actually described (but as detailed above not nomenclaturally available) is the HT of He. andravahana ab. marmorata Aurivillius, 1911. This and a large series of specimens referred to Heteropsis harveyi in the Oberthür collection are clearly distinct from Ht. vanewrighti in being generally larger in size, possessing a FWV androconial patch, and occurring much further south in the central to southeastern parts of the rainforest biome.</p><p>Additional information. DNA divergences: CO1-5P cluster number BOLD:AAE4111, with two haplotypes (exemplars BMAD064-09 (Bekolosy), BMAD212-15 and BMAD216-15 from Antsatrotro) is about 1.52% divergent to ‘ H. sp. 20’ (“sp. 77” of Torres et al., 2001; exemplars of cluster number BOLD:ACW5777 including HM 240620); for more details see under Ht. harveyi .</p><p>Phylogeny/sister species: without a detailed phylogeographic analysis of the Ht. harveyi -complex, the sister taxon of Ht. vanewrighti is not yet certain, but the closest currently described relative is Ht. harveyi . Aduse-Poku et al., (2016, in press) find it sister to Ht. sp. 20.</p><p>Ecology and distribution.</p><p>Habitat: montane rainforest with fine leaved climbing bamboos.</p><p>Behaviour: at rest and in rainy weather, both sexes rest on tree trunks.</p><p>Hostplant: unknown, assumed to be fine-leaved climbing bamboos among which the adults fly.</p><p>Early stages: unknown.</p><p>Distribution: Bekolosy and Antsatrotro mountains (where found in 2011), Réserve Spéciale de Manongarivo, to where, as far as is known, Ht. vanewrighti i s endemic (Fig. 30 A, purple dots).</p><p>Elevational range: 1045–1487 m. (n=73 including referred specimens and observational data).</p><p>Referred specimens. ♂, Madagascar NW, Bekolosy Mt., RS Manongarivo, 14.045o S, 48.306o E +/- 3.5 km, 1150 +/- 350 m, 23/12/1994, D.C. Lees: DLBEK 94_144A; ♂, NW, Bekolosy Mt., RS Manongarivo, 1370 m, 14.0263o S, 48.3075o E +/- 0.16 km, 1370 +/- 50 m, 16/12/1994, 11:20, D.C. Lees: DLBEK 94_99; ♂, data as above but: DLBEK 94_100; ♂, NW, Bekolosy Mt., RS Manongarivo, 1163 m, 14.0458o S, 48.2962o E +/- 0.5 km, 1130 +/- 35 m, 12/12/1994: 08:00–09:00, 11:20, D.C. Lees: DLBEK 94_169; ♀, NW, Bekolosy Mt., RS Manongarivo, 1225 m, 14.0335o S, 48.3098o E +/- 1 km, 1250 +/- 50 m, over river, 18/12/1994: 09:30–10:20, 11:20, D.C. Lees: DLBEK 94_164; ♂, NW, Antsatrotro, c. 1250 m, 3/12/2011: 13:52, D.C.Lees, IA610 [isotope voucher]; ♂, NW, Antsatrotro, c. 1260 m, 3/12/2011, D.C.Lees, IA611 [isotope voucher]; ♀, NW Antsatrotro, c. 1250 m, 3/12/2011, D.C.Lees, IA612 [isotope voucher]; ♂, NW, Antsatrotro, c. 1250 m, 3/12/2011: 10:24, D.C.Lees, IA613 [isotope voucher]; ♂, NW, Antsatrotro, Manongarivo, 3/12/2011: 09:25, WG46 [wing image], IA230 [isotope voucher]; ♂, NW, Antsatrotro, Manongarivo, 12/2011, DL 0033, KA753 [=KA-P753; DNA extract voucher].</p></div>	https://treatment.plazi.org/id/038747324C69C6751EB72C61FB15228C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C54C6711EB72966FF2221A3.text	038747324C54C6711EB72966FF2221A3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis westwoodi Lees	<div><p>Heteropsis westwoodi Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:422E5526-8FBB-4AD6-A47D-EC95FF54E16F</p><p>Prior references: sp. 13B (Lees, 1997: 64). “KA522_Heteopsis_ andravahana ” [sic] (Aduse-Poku et al., 2015: Fig. 1).</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 10 A), Madagascar SE, Vohitrasiva, near ridge, " 1390 m ", Andringitra, [mountaintop on way to Vohipia (1760 m pass)], 22.1684o S, 47.0289o E +/- 0.1 km, 1450 +/- 50 m, 17/2/1995, 11:40–13:15, D.C. Lees: DL 95-0002; KAP20 [=KA-P20; DNA extract number], NHMUK 010289130 [QTR barcode].</p><p>Paratypes: Deposition BMNH: ♂, data as HT but DL 95-0001, IA288 [isotope voucher], KA751 [=KA-P751; DNA extract number], NHMUK 010289131 [QTR barcode]; ♂, data as HT but DL 95-0003, 236 DL [genitalia], IA290 [isotope voucher], NHMUK 010289182 [QTR barcode].</p><p>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.</p><p>Deposition summary: BMNH (HT ♂, 2 PT ♂♂), MNHN (7 PT ♂♂).</p><p>Type locality. Madagascar E, Vohitrasiva, Andringitra, 22.1684o S, 47.0289o E +/- 0.1 km, 1450 +/- 50 m.</p><p>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.</p><p>Description. Wings: dorsal surface fairly uniform dark brown, darker in area of cell and above towards costa. FWD space-CuA1 ocellus expressed, roughly circular, its black iris spanning inter-vein distance CuA1-CuA2, with a narrow concentric dark orange ring. HWD space-cuA1 ocellus elliptic, with a black iris spanning more than half intervein distance CuA1-CuA2, 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-CuA1 is circular and same size as on the FWD with a concentric quite narrow orange ring. M1 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 M1 ocellus FWV. In the HWV the space-CuA1 ocellus is as on the HWD and space-R5-M3 ocelli are expressed as very small white points. Mb irregular, strongly concave from the margin to M2 where it forms a cusp, then strongly and irregularly indented and maximally concave in space-CuA1, convex again in space-CuA2 and concave again to vein 1A+2A 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 M2-1A is a clear white band, also irregular, with prominence in space M2 and maximum width proximad of space-CuA1 ocellus, tapering to 1A+2A with a slight expansion in Space-CuA2. 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-CuA1 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 ♂ (DL95- 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.</p><p>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 ♀).</p><p>Androconia: HWD discocellular brush light brown. HWD discocellular patch ‘bullet’-shaped, peach pink coloured.</p><p>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.</p><p>♂ genitalia: 236DL, PT ♂ from Vohitraseva (Lees, 1997: 109, Fig. 7 i, “13B”; 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.</p><p>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.</p><p>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.</p><p>Additional information. DNA divergences: COI-5P cluster number BOLD:ACW4998 (exemplars BMAD134-15 DL06-980 from RS Kalambatritra and BMAD243-15 CK92-0005 from PN Andohahela; also KA- P510 (see Aduse-Poku et al., 2015) from Ambondrombe).</p><p>Phylogeny/sister species: apparently most closely related to Ht. harveyi (cluster number BOLD:ACW5694 and to Ht. viettei (BOLDAAK5839). 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 .</p><p>Ecology and distribution.</p><p>Habitat: montane rainforest with fine-leaved climbing bamboos.</p><p>Behaviour: ♂♂ fly quite rapidly in the substory.</p><p>Hostplant: suspected to be fine-leaved climbing bamboos.</p><p>Early stages: unknown.</p><p>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).</p><p>Elevational range: 785–1817 m. (n=27 including referred specimens and observational data).</p><p>Referred specimens. ♂, Madagascar E, Ambondrombe, 21.8782o S, 47.2474o 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, 21.87874o S, 47.24969o E +/- 1 km, 170 +/- 50 m, 17/2/1995, D.C. Lees, DL-4-863, KA522 [=KA- P522; extract number; sequenced specimen in Aduse-Poku et al., 2015]; ♀, E, Andringitra, ridge, 1800 m, 20/11/ 2011, D.C.Lees: DL-11-0001, IA645 [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 S10, 1200 m, 48 54'E 24 34'S, [24.5735o S, 46.2o E 1200 m], 29/11/1992, CK92- 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, [24.5735o S, 46.2o E, 1200 m], 29/11/1992, BMNH (E) #674842; ♂, SE, Andohahela, C. Kremen, BMNH (E) #697865, 238BDL [genitalia], IA636 [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), 23.4186o S, 46.44025o E +/- 0.042 km, 1569 +/- 25 m, 12/12/2006: 11:47, R. Ranaivosolo: DL 06-767; ♂, SE, Befarara, RS Kalambatritra (western), 23.418o S, 46.44o E +/- 0.15 km, 1557 +/- 25 m, 12/12/2006, M. Linares: DL 06-811, IA161 [isotope voucher]; right wings imaged.</p></div>	https://treatment.plazi.org/id/038747324C54C6711EB72966FF2221A3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C50C67F1EB72C0FFB4522CC.text	038747324C50C67F1EB72C0FFB4522CC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis pauliani Lees	<div><p>Heteropsis pauliani Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:422E5526-8FBB-4AD6-A47D-EC95FF54E16F</p><p>Prior references: sp. 37 (Lees, 1997: 64; Torres et al., 2001: 462).</p><p>Type material., Deposition MNHN: Holotype: ♂ (Fig. 10 C, MNHN), Madagascar Nord [NW], massif du Tsaratanana, en dessous de l’Andohanisambirano [position estimated at 14.14o S, 48.97o E +/- 1 km, 2050 m], 5–8/ 11/1966, 2050 m|mission du Tsaratanana XI-1966 Camp n o 1 P. Griveaud, P. Soga, P. Viette, D. Wintrebert| HOLOTYPE | DCL-DB-2966.</p><p>Paratypes: Deposition BMNH (accession BMNH (E) 2008-69): ♂, Madagascar NW, Ampitsinjobana, Tsaratanana, below Andohanisambirano, 14.1478o S, 48.9679o E +/- 0.15 km, 2320 +/- 25 m, 23/12/2004, D.C. Lees et al.: DL-05-804, BMNH (E) #671691 [DNA voucher; cytochrome b], KA507 [=KA-P507; DNA extract voucher]; ♀, NW, Tsaratanana, Matsoborimaiky, in middle of dry lake, 14.1526o S, 48.9578o E +/- 0.015 km, 2035 +/- 25 m, 23/12/2004, Alain: DL-05-818, BMNH (E) #671694 [DNA extract, cytochrome b];</p><p>Deposition MNHN: ♀ (Fig. 10 D), [Madagascar NW], Mt. Tsaratanana, 2000 m. [estimated at 13.9755o S, 48.8381o E +/- 12.88 km, 2000 m], II/1951; R. Paulian, DCL-DB-2975; ♂, M’car. Nord [Madagascar NW], massif du Tsaratanana Matsabory - en dessous de l’Andohasambirano [Andohanisambirano, position estimated at 14.14o S, 48.97o E +/- 1 km, 2050 m], 5–8/11/1966 [P. Griveaud, P. Soga, P. Viette, D. Wintrebert]| INSTITUT SCIENTIFIQUE MADAGASCAR | DCL-DB-2967; 8 ♂♂, same data as above but: DCL-DB-02968-DCL-DB- 02975;</p><p>Deposition PBZT: ♂, Madagascar NW, matsabory (lake) in Antsaralasy, S14°11'51,8'', E49°04'06,6'' [14.1977o S, 49.0685o E], 2153 m. 7/10/2013. Ravo Ranaivosolo: MTSBR, BMAD234-15 [DNA barcode], IA291 [isotope voucher].</p><p>Deposition summary: MNHN (HT ♂, 9 PT ♂♂, PT ♀); BMNH (PT ♂, PT ♀); PBZT (1 PT ♂).</p><p>Type locality. Madagascar Nord, massif du Tsaratanana, en dessous de l’Andohanisambirano [position estimated at 14.14o S, 48.97o E +/- 1 km, 2050 m].</p><p>Diagnosis. No really similar species even in the Ht. drepana group. This species is very distinctive in possessing tails on the HW margin, giving it a superficial resemblance at least dorsally to a species of the Ht. antahala group. The long tails distinguish it also from the perhaps quite closely related Ht. imerina .</p><p>Description. Wings: upperside fairly uniform dark brown (noticeably faded in the HT), grading to smoky brown, especially towards costa. FWD space-CuA1 ocellus spanning veins CuA1-CuA2, with approximately concentric, reddish (faded in the HT) Orng. M1 ocellus very small, not obvious. In HWD, space-CuA1 ocellus small and with reddish ring, slightly elliptic. Basal area of HWD covered in long dark appressed hairscales, unlikely to be androconial., Dark brown Sml of HWD highlighted with ochreous scales on either flank, the proximad band being considerably wider. HW margin pointedly crenate with acutely pointed dark brown tails (faded in the HT) at veins M3 (longest in the HT) and CuA1 and with lesser tails at veins CuA2 and 1A+2A (where tail quite rounded). FWV dark brown with space-CuA1 ocellus again spanning veins CuA1-CuA2 and with more or less concentric (in the HT) Orng of intermingled mosaic of reddish and ochreous-yellow scales, extended proximad where following concave dark brown trace of Mb which borders generally darker scaling proximad. FWV M1 ocellus as distinct white spot (in the HT), lacking a distinct black border and red Orng. HWV very elegantly patterned. Space-CuA1 ocellus a bit smaller than that in FWV and with quite wide dark red Orng. Lacking a distinct PMb, basal area of HWV irrorated dark violet-lilac-brown (faded in the HT) and glossy light brown to ochreous (towards anal area). Mb forming sharp edge to the basal area, dark brown and irregular, concave-inflexed towards base of space-M3 and CuA1 (maximally concave between the corresponding veins) and then with a fairly straight line towards anal angle, then slightly bent back towards 1A+2A straight also above its distad cusp at vein M3 to 60% along costa. Distad of the HWV Mb, mid brown from above vein Rs to M2 and deep cream scales pervading rest of distal area, especially proximad of ocellus space-CuA1. A narrow dark brown Sml is highlighted more in cream proximad than it is on the distad flank, forming in effect an irregular wide cream band that terminates costad at vein M2. Slightly above, below and marginad of space-CuA1 ocellus, the scales are a relatively uniform greyish to violet grey. In the HT, ocelli of space-Rs and M1, and less distinctly so M2 and M3, are expressed as small white points. Fringes dark brown, thicker towards tails. Variation. Sexes similar but the only known ♀ larger and generally lighter in colouration.</p><p>Wingspan/fwl: range 33.2–38.6/18.7–21 (n=3 ♂♂), mean = 39.1 +/- 1.8 SD (n=8 ♂♂), including HT ♂ 33.2/ 18.7 mm. PT ♀ about 42 mm wingspan.</p><p>Androconia: small dark brown brush overlying lenticular dark brown or blackish patch on stem of HWD R vein just above cell.</p><p>Palps: penultimate segment with narrow cream stripe fringed with black hair-scales beyond compound eye, away from eye dark grey or black with lighter greyish scales between these two stripes. Mesad and inner face dark grey.</p><p>♂ genitalia: 97DL (PT, Fig. 11 B): from LV, tegumen with distinct brow towards scythe-hooked uncus which is relatively broad at base and narrow in middle. Uncus slightly longer than dorsal profile of tegumen. Constriction at hinge with vinculus is relatively broad, about 2/3 length of tegumen. Gnathos fairly straight and tapered and slightly downcurved pointing towards dorsal edge of uncus. Valves quite long with a slight ventral elbow at base of arm and flattened blades strongly covered in spinoid setae on mesad face of distal club, those at the edge appearing like saw-tooth serrae. Valves protrude almost the full length of club beyond uncus tip. Saccus not especially long and up-bent. Aedeagus nearly full length of valve, quite long proximad of ostium and slightly recurved distad of it.</p><p>Etymology. After the pioneering French entomologist the late Rector Renaud Paulian (1913–2003), who described two Heteropsis taxa and discovered the first known specimens of this species on Madagascar’s highest mountain a lifetime ago.</p><p>Discussion. This species was first recognized by DCL in 1994 (Lees, 1997: 54) at MNHN with collections from Tsaratanana by R. Paulian in 1951 and by P. Griveaud et al., in November 1966. It was since found at Anjanaharibe Sud in 1996 by Claire Kremen and by R. Ranaivosolo just to the east of RNI Tsaratanana in October 2013. All available types within the Ht. drepana group have been examined and the species is distinct from any Heteropsis hitherto described.</p><p>Additional information. DNA divergences: COI-5P cluster number BOLD:ACW4997 (exemplar MTSBR, BMAD234-15), 6.08% divergent to Ht. viettei (Anjanaharibe Sud; exemplar BMAD137-15, DL14A-0196, cluster number BOLD:AAK5839) and with a similar divergence to a few other related species. In their COII dataset (Torres et al., 2001), with 422 bp comparable, Ht. pauliani (accession AY 040165), based on a single individual, is closest (5% pairwise divergent) to “ H. sp. 32” (?also = H. viettei; AY 040143), based on one ♂ from Andohahela. It is considerably more pairwise divergent, about 7.67%, in the same COII dataset (417 bp compared) to Ht. imerina (their ‘ Hen. sp. 22’; AY 040150, based on two ♂♂ from Ankazomivady, an exemplar of which is illustrated in Fig. 7 E). This last taxon is also only marginally different from, and undoubtedly conspecific with (see Ht. imerina description below) ‘ Hen. sp. 76’, (AY 040174) of Torres et al., (2001).</p><p>Phylogeny/sister species: sister group uncertain, without agreement so far between molecules and morphology (but see Aduse-Poku et al., 2015; 2016, in press, where sister to Ht. imerina in combined tree). Based on parsimony analysis of morphological characters, Lees (1997: 156, Fig. 1) had over 96% jackknife support for a sister relationship between Ht. pauliani sp. nov. (“THRSV”) and Ht. 76 (“SVTSX”) + Ht. imerina (“TWNTW”), while Ht. westwoodi sp. nov. (THRTB) and Ht. viettei (THRTN) fell in the next branch down. In their cladistic parsimony analysis of COII sequences, Torres et al., (2001: 467) found a topological relationship, although with meagre 51% bootstrap support, for a clade consisting of Ht. pauliani (their Hen. sp. 37, from Anjanaharibe Sud) and ‘ H. sp. 74’ + ‘ H. sp. 32’ (taxa from Andohahela and Anjanaharibe Sud, respectively that were fully supported as sisters; but they did not include Ht. viettei (H. sp. 13) nor Ht. westwoodi (H. sp. 13B)). Apparently, however, there was a problem with the GenBank submission and this result cannot be verified (it must be based on an unsubmitted sequence), because on GenBank, AY 040173 (“ Henotesia sp. Torres-74”) is apparently mislabelled and cannot correspond to its position in the tree. That is because sequence AY 040173 is identical to AY 040169 ( Heteropsis laetifica (Oberthür, 1916), Analalava, a species in the Ht. strigula group), whereas the real morphospecies 74 should be either closely related, if not conspecific, with Ht. viettei Lees, 2003). This last taxon (then ‘sp. 13’) was not [otherwise] included in their study.</p><p>Ecology and distribution.</p><p>Habitat: montane rainforest, sclerophyllous forest including ‘matsabory’ (marshy and lake forest areas).</p><p>Behaviour: known to fly low across gaps between forest. One was caught in the middle of a ‘matsoborimaiky’ (dry lake) at Tsaratanana.</p><p>Hostplant: unknown.</p><p>Early stages: unknown.</p><p>Distribution: endemic to the mountain ranges between Tsaratanana and Anjanaharibe Sud, as far as is known (Fig. 30 A, dark green dots). Due to a formatting error, the locality was incorrectly given in Table 1 of Torres et al., 2001 as Andohahela, instead of its true provenance in the line below, Anjanaharibe Sud.</p><p>Elevational range: 1778–2320 m. (n= 13 incl. referred specimens and observations).</p><p>Conservation: probably the highest elevation Heteropsis in Madagascar. Not likely in imminent danger by habitat destruction. Ht. pauliani likely occurs on a number of high and rather inaccessible peaks or ridges across its range. It might however be endangered by global warming. No individuals were found in a lower elevational transect up to 1550 m on Anjanaharibe Sud in 2014, although this may be too low.</p><p>Referred specimens. ♂ [head and right wings only], Madagascar NE, RS Anjanaharibe Sud, 14.7431o S, 49.4371o E +/- 0.51 km, 1788 +/- 238 m, 8/2/1996, 16:05–17:15, C. Kremen: CK720 IA293 [isotope voucher]; ♂, NE, RS Anjanaharibe Sud, 2000 m, 14.7431o S, 49.4374o E +/- 0.5 km, 2000 m, C. Kremen; BMNH (E) 2008-69; ♂ [worn], NE, Anjanaharibe Sud, 14.7431o S, 49.4371o E +/- 0.51 km, 1788 +/- 240 m, 8/2/1996: 16:05-17:15, C. Kremen: CK722, DL 9686 [DNA voucher], IA292 [isotope voucher]; ♂, NE, Anjanaharibe Sud, 2025 m, 14.7431o S, 49.43735o E +/- 0.495 km, 2025 +/- 50 m, 8/2/1995, C. Kremen: CK715; 307 DL [genitalia], 335 [= DL 0335; DNA extract number], BMNH (E) #697749; ♂, NE, Anjanaharibe [Sud], 14.7431o S, 49.4371o E +/- 0.51 km, 1788 +/- 238 m, 8/2/1996: 16:05–17:15, C. Kremen: CK726 [DNA voucher]; ♂, NE, Anjanaharibe Sud, 2000 m, 14.7431o S, 49.4374o E +/- 0.5 km, 2000 m, C. Kremen; ♀, NE, Anjanaharibe Sud, 14.7431o S, 49.4371o E +/- 0.51 km, 1788 +/- 238 m, 8/2/1996: 16:05–17:15, C. Kremen: CK720, IA293 [isotope voucher].</p></div>	https://treatment.plazi.org/id/038747324C50C67F1EB72C0FFB4522CC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C5EC67C1EB7292BFD3E27E9.text	038747324C5EC67C1EB7292BFD3E27E9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis imerina Lees	<div><p>Heteropsis imerina Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:988AC583-72F8-4CA4-824A-157D9F7C5AEC</p><p>Prior references: sp. 22 (Lees, 1997; Torres et al., 2001: 462);</p><p>Sp. 76 (Torres et al., 2001: 462).</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 9 C), Madagascar E, Mantadia National Park, Jofa, near bridge, 18.7674°S, 48.4324°E +/- 0.4 km, 945 +/- 15 m. 16/01/2014. D.C. Lees: DL 16114-22-1, NHMUK 010289132 [QTR barcode].</p><p>Paratypes: Deposition BMNH: ♂, data as HT but DL 16114-22-2, NHMUK 010289133 [QTR barcode]; ♂, Madagascar E., Mantadia National Park, Mitsinjo site PK 17.5 from Vakona Forest Lodge, 1100 m, 11/12/2011, D.C. Lees: DL 11M-0001, NHMUK 010289134 [QTR barcode];</p><p>Deposition MNHN: ♂, Madagascar E. route d’Anosibe VI-1966 /MUSEUM PARIS COLL. G. DUJARDIN DEL. ET R. VIOSSAT|DCL-DB-2908; ♀ (Fig. 9 D), Madagascar Est, route de Lakato km 15, Ankasoka 1100 m 17/ 21-X-1963 P. Viette|DCL-DB-2906.</p><p>Deposition summary: BMNH (HT ♂, 2 PT ♂♂), MNHN (PT ♂, PT ♀).</p><p>Type locality. Madagascar E., Mantadia National Park, Jofa, near bridge, 18.7674°S, 48.4324°E +/- 0.4 km, 945 +/- 15 m.</p><p>Diagnosis. Very distinct by its shorter HW tails from the somewhat similar Ht. pauliani sp. nov., where potentially (see discussion) sympatric (from RS Anjanaharibe Sud to RNI Tsaratanana). Heteropsis harveyi is the species most likely to be confused in the main central-eastern rainforest block, from which Ht. imerina differs by lack of an androconial patch on the FWV, by the proximad inflexion of the HWV medial band around ocellus space-CuA1 being rather sharply angled, and by the more evenly pronounced HW tails of the former, as for the other small dark species (in the case of ♂♂). Ht. vanewrighti sp. nov., Ht. viettei and Ht. westwoodi are easily distinguished from Ht. imerina by their whitish highlighting distad of the HWV Mb, which in Ht. imerina ♂♂ is yellowish, sometimes with a reddish suffusion proximad of the space-CuA1 ocellus.</p><p>Description. Wings: upperside uniform mid to dark brown (costad area not much darker) with space-CuA1 ocellus well expressed on FWD, with a round concentric orange-reddish ring, its black iris almost spanning or spanning CuA1-CuA2, and the Orng compromising these veins and moderately narrow and eccentric. FWD space- M1 ocellus not evident. HWD space-CuA1 ocellus also hardly evident. FWV space-CuA1 ocellus similar size to FWD and marginally more eccentric with mid orange ring slightly skewed to tergum. Space-M1 ocellus much smaller and with a small black iris and a faint mid orange ring; that in space-M2 almost reduced to a small white spot. HWV space-CuA1 ocellus relatively small compared with others of Ht. drepana group and subelliptic with a narrow mid orange ring. Other HWV ocelli, particularly those of spaces Rs-M3, also of space-CuA2-1A, expressed as white spots. Ventral wing background colour with a distinct dark reddish brown cast. FWV more uniform than HWV but paler towards tergal angle with a pale ochreous mark near costa towards apex, HWV lighter brown distad of Mb. This dark russet brown Mb is irregularly snaking, concave around its intersection with M1, convex around that with M2, curving back to the deepest concavity in space-CuA1 and convex again near vein CuA2 before terminating at 1A. This line is highlighted with ochreous admixture with reddish brown scales distad of it and sombrely shaded proximad of it, with a more consistent irroration with russet brown contrasting with more ochreous brown scales in the basal areas; the basal HWV areas are thus relatively dark in the ♂, while the PMb is indistinct and basad of it is also ochreous highlighted. Before the margin in both wings is a slightly more ochreous highlighting and a double thin russet-brown Pml (pre-marginal line), the outer of which follows the margin, which is lightly crenate. The slightly more prominent but short dark brown ‘tail’ is at the end of M3, which can be a field character distinguishing from similar members of the Ht. drepana group, particularly in the ♀ (Fig. 9 D). Variation. ♂♂ vary quite a lot in their underside colouration, and the HWD space-CuA1 ocellus is sometimes expressed. White spots not always evident on HWV. ♀ is similar but larger, lighter brown on upperside and with more ochreous cast on upperside with PMb and Mb russet brown on HWV, thus more pronounced. In apparent dry seasonal forms, the HWV ocelli can be very reduced.</p><p>Androconia: discocellular brush HWD mid brown, to ochreous brown towards tip. Underlying patch lenticular, pale ochreous. No other androconia.</p><p>Wingspan/fwl: 35.2 (n=1)/ 18.4–19.3 mm (n=20); 35.4–38.1/19.7–22.0 mm (n=4); mean = 33.8 +/- 1.44 SD/ 17.87+/- 0.54 SD (n=6 ♂♂, including HT, 35.4/ 18.4 mm); mean = 36.2 +/- 3.03 SD (n=4) /18.75 +/- 1 SD mm (n=4) (♀).</p><p>Palps: mid brown without clear pale striping.</p><p>Etymology. Refers to its potentially wide distribution across the Madagascan haut-plateaux also including surrounding mountain ranges, as for the principal Merian tribe of the Malagasy people, which gives this species its name.</p><p>Discussion. There is a certain amount of historical material, but perhaps surprisingly the species is not implicated in any pre-existing descriptions. This species was first recognized in the field as new as morphospecies 22 at Ankazomivady forest on 12/01/1992 by Claire Kremen (Lees, 1997: 64) and as sp. 76 by the same worker in 1995–1996 at Anjanaharibe Sud, but there were earlier collections, mostly represented in MNHN. The morphospecies ‘sp. 76’ (Anjanaharibe Sud) is definitely referable to Ht. imerina, although its ♂ genitalia had been considered distinct in Lees (1997: 65); spp. ‘22’ and ‘76’ were recovered anyway as sister taxa in Lees (1997: 156), but the recent discovery by myself of additional populations with adult phenotype similar to Ht. imerina from Manongarivo through Tsaratanana to Marojejy and Anjanaharibe Sud suggest the species is indeed quite widespread. A deeper study of variation in the genitalic variation would be interesting, especially if backed up by molecular sequences; the study of Torres et al., (2001) also showed minimal differences in COII (see below).</p><p>Additional information. ♂ genitalia: 154DL (not in type series, Fig. 8 C): from LV, tegumen with fairly straight dorsal profile (strong v-shaped notch proximad from DV), leading to narrowly scythe-hooked uncus without a strong ventral ‘dewlap’. Gnathos relatively thin and sinuate, tapering, first downcurved and then upcurved (also sinuate from DV). Constriction in division with vinculum about half maximum dimension of tegumen. Vinculum not strongly bowed and broadening towards base of saccus. Valves relatively long, with distinctly angled shoulder, and stretched out valve base narrowing gradually to valve arm which is slightly longer than valve base with distinct ventral ‘elbow’, leading to spatulate tips with a limited number of spinoid setae on inner face/at tip (valve arms gently incurved from DV without a strong terminal spine but with a small number of teeth on mesad edge towards tip). Valve tips protrude considerably proud of maximum extension of uncus tip (Lees 1997: 109, Fig. 7 i). Aedeagus very thin and gently recurved distad of ostium, as long as valve. Saccus not particularly long and bulbous. Juxta prominent and narrowly cupped proximad.</p><p>DNA divergences: the COI-5P cluster number (BOLD:AAE4110) includes the same specimen as sequenced for cytochrome b as “sp. 22” in the Monaghan et al., (2001) study (DL-4-515, BMNH(E) #676683, BMAD046-09, Analamay [18.828o S, 48.339o E, 978 m]), as well as exemplars BMAD231-15 from RNI Tsaratanana and BMAD246-15 from RS Anjanaharibe Sud. This last exemplar ‘sp. 76’ has an identical sequence to that from Analamay. The 357 bp cytochrome b sequence of the last specimen (BMNH(E) #676683 FJ819266, BMNH(E) #676685 FJ819267) is about 4.2% pairwise divergent to those of Ht. harveyi described here (BMNH(E) #671648 FJ819453, BMNH(E) #676467 FJ819454; Monaghan et al., 2009; additional members of the Ht. drepana group in that study include Ht. passandava, Ht. narova, Ht. difficilis and Ht. strato). A 706-bp relatively conserved 28SDDFF sequence (BMNH(E) #676640, FJ817836) as aligned is identical to those of Ht. strato (BMNH(E) #671544 FJ817810) and Ht. andravahana (BMNH(E) #671565 FJ817806, BMNH(E) #676629 FJ817807, BMNH(E) #676641 FJ817808; ‘ difficilis ’: Monaghan et al., 2009).</p><p>Phylogeny/sister species: Based on parsimony analysis of COII sequences, Torres et al., 2001 had ‘sp. 76’ as sister to Ht. imerina (their ‘ Hen. sp. 22’) with 100% bootstrap support. In their dataset, with 417 bp compared, ignoring two differences at the 5’ end, Ht. imerina (AY 040150 based on 2 ♂♂ from Ankazomivady Forest 32 km S. Ambositra, see correction below) is only 2 bp (0.48% pairwise divergent) from their ‘ Hen. sp. 76’ (AY 040174, based on 2 ♂♂ from RS Anjanaharibe Sud). According to the ‘total evidence’ morphological analysis of Lees (1997: 156, Fig. 1), there was 0.963 jackknife support for a sister relationship of these exemplars with Ht. pauliani (“sp. 37”). There is some support for Ht. imerina as the sister of Ht. pauliani in the combined tree of Aduse-Poku et al., (2016).</p><p>Ecology and distribution.</p><p>Habitat: primary rainforest. This species was (remarkably) once found in Parc de Tsimbazaza in 1951 (MNHN), so it may persist or may have persisted until recently in other central habitats well separated from the current eastern rainforest block.</p><p>Behaviour: Low flying, found mostly along ridges but also in riparian areas, perching low off the ground.</p><p>Hostplant: occurs among native climbing bamboos, which it might feed on.</p><p>Early stages: expressed egg pearly whitish.</p><p>Distribution: widespread in the principal eastern rainforest belt and also in high plateau relics notably Ankazomivady south of Ambositra (Fig. 30 A, turquoise dots). The locality was incorrectly given in Table 1 of Torres et al., 2001 (which contains a formatting glitch) as ‘Anjanaharibe Sud, 1240 m’ instead of the line below, ‘ 32 km S. of Ambositra’. The species extends to the northern mountain range of the rainforest biome. ‘Sp. 76’ from RS Anjanaharibe Sud is doubtless referable to Ht. imerina, as for material (‘sp. 65’) from Bekolosy (RS Manongarivo) and RNI Tsaratanana, while this is a species for which phylogeographic study is desirable.</p><p>Elevational range: 786–1932 m. (n=118; DCL-DB-2901 from Tamatave-Anivorano [estimated at 80 m] is discounted as an unlikely outlier).</p><p>Referred specimens: ♂, Madagascar E, Bemoara camp (going out from to Ambavala), Zahamena PN [northwestern sector], 17.51875o S, 48.72455o E +/- 0.27 km, 1100 +/- 25 m, 4/11/2006, D.C. Lees: DL 06-023; ♂, Imerina Madagascar |1910 E. La Moult | COLL. BOULLET MUSEUM PARIS | H. andravahana ab. marmorata, Auriv. [sic]|DCL-DB-2897; ♀, Imerina Madagascar |1910 E. La Moult |DCL-DB-2911; ♂ [MNHN], Centre, forêt d'Andranobe, route d'Andriamena, O. of Lac Alaotra [17.755o S, 47.98o E +/- 1.5 km, 1250 m], 1/11/1972 |DCL-DB- 2914; ♂ [MNHN], data as above but: 1–6/9/1972, A. Peyrieras|DCL-DB-2915; ♀, Centre, forêt d'Andranobe, route d'Andriamena O. du lac Alaotra [ca. 17.755o S, 47.98o E +/- 1.5 km, 1250 m], 29-1/ 6-II-1970, P. Griveaud|DCL-DB- 2913; ♀, data as above but DCL-DB-2913; ♀, [MNHN], data as above but|DCL-DB-2916; ♂, E, Tamatave et Anivorano, [ca. 18.4585o S, 49.1832o E +/- 39 km, 80 m]|1915|MUSEUM PARIS DON. DE H. UNGEMACH/ GENITALIA m. P. VIETTE PREP. NO. 4864|DCL-DB-2901; ♂, E, Analamay, 18.82795oS, 48.3388oE +/- 2.1 km, 978 +/- 50 m, 11/3/2004, D.C. Lees: DL-4-515, BMNH (E) #676683, BMAD 046-09, CCDB-02225-D10 [DNA barcode; cluster number BOLD:AAE4110; ♂, E, Ambatovy-Berano, 18.848845o S, 48.337055o E +/- 0.65 km, 965 +/- 7 m, 12/3/2004: 10:43, R. Ranaivosolo: DL-4-643; ♀, E, Ambatovy, 18.84935o S, 48.32399o E +/- 0.3 km, 1024 +/- 25 m, 25/2/2004: 15:33, R. Ranaivosolo: DL-4-99; ♂, E, Ambatovy-Berano, 18.85o S, 48.335o E +/- 1.7 km, 1000 +/- 90 m, 12/3/2004, D.C. Lees: DL-4-590, IA27 [isotope voucher]; ♂, data as above but: R. Ranaivosolo: DL-4-666; ♂, E, Ambatovy ‘second site’, 18.8511o S, 48.3221o E +/- 0.15 km, 1055 +/- 25 m, 25/2/ 2004: after 12:00, D.C. Lees: DL-4-139, 547 [= DL 0547; DNA extract number], BMNH (E) #671912; ♀, data as above but: 21/3/2004: 09:29, V. Rakotomalala: VFR_1329 [photo, verified DCL]; ♀, data as above but: 19/3/2004: 09:54, V. Rakotomalala: VFR_1254 [photo, verified DCL]; ♂, E, Ambatovy, 18.8522o S, 48.3177o E +/- 0.5 km, 1085 +/- 30 m, 25/2/2004: 12:59, R. Ranaivosolo: DL-4-84; ♂, E, Ambatovy-Sahavarina, 18.8577o S, 48.32579o E +/- 0.3 km, 1090 +/- 40 m, 13/3/2004: 09:23, R. Ranaivosolo: DL-4-707; ♀, E, Ambatovy-Sahavarina, 18.8590o S, 48.3252o E +/- 0.4 km, 1090 +/- 45 m, 13/3/2004: 10:40, R. Ranaivosolo: DL-4-681; ♂, data as above but: DL-4- 682; ♀, E, Ambatovy- Sahamaloto-Torotorofotsy path, S. to Sahavarina, 18.86313o S, 48.3425o E +/- 0.15 km, 976 +/- 25 m, 14/3/2004: 10:39, D.C. Lees: DSC _0496-7 [photo]; ♂, E, Torotorofotsy-Sahavarina, 18.8657o S, 48.34547o E +/- 0.25 km, 969 +/- 25 m, 14/3/2004: 10:52–13:54, D.C. Lees: DL-4-730; ♀, E, Torotorofotsy (Sahavarina), ilot, 18.8665o S, 48.3492o E +/- 0.015 km, 959 +/- 25 m, 14/3/2004, D.C. Lees: DL-4-769, 2485 [= DL 2485; DNA extract number], BMNH (E) #676685; ♀, data as above but: 14:30, D.C. Lees: DL-4-774; ♂, E, Ambatovy-Sahavarina, 18.8620o S, 48.3408o E +/- 0.2 km, 1004 +/- 28 m, 14/3/2004: 10:30, R. Ranaivosolo: DL-4- 737; ♂ [MNHN], C, Tsimbazaza, [ca. 18.933o S, 47.533o E +/- 0.9 km, 1317 m], 7/1/1951, Villette; ♂ [CMN], Ambatolaona [ca. 18.9333o S, 47.9o E +/- 15 km, 1372 m], 12/1932, M.E.H. Fountaine; ♂ [MNHN], CE, Perinet ANALAMAZAOTRA, [ca. 18.9347o S, 48.4305o E +/- 1.68 km, 925 +/- 20 m], 7/1967, P. Griveaud; 4 ♂♂ [BMNH], CE, Perinet [ca. 18.9347o S, 48.4305o E +/- 1.68 km, 925 +/- 50 m], 4/1973, A. E. Wright; 3 ♂♂, E, Perinet, forêt de l'Est, Madagascar 1910. E. Le Moult [ca. 18.9347o S, 48.4305o E +/- 1.68 km], DCL-DB- 2899- 2901; 1 ♀, E, Perinet, forêt de l'Est, Madagascar 1910. E. Le Moult [ca. 18.9347o S, 48.4305o E +/- 1.68 km], DCL- DB-2912; ♂, E, Mantadia, Jofa, 18.78o S, 48.43o E +/- 1 km, 950 +/- 50 m, 16/1/2014: 12:29, D.C. Lees: MAD 121 [pheromone voucher], IA374 [isotope voucher]; ♂, E, Maromizaha, 18.9668o S, 48.4547o E +/- 1.49 km, 1109 +/- 73 m, 24/2/2002, D.C. Lees; ♂, CE, Antanandava: Sandrangato Forest, 19.1083o S, 48.2417o E +/- 0.44 km, 880 +/- 50 m, 6/11/1994, D.C. Lees; ♂ [MNHN], CE, Anosibe, route d' km 45 [ca. 19.13o S, 48.34o E +/- 5 km, 1041 m], 25/11/ 1966, P. Viette|DCL-DB-2907; 2 ♂♂ [MNHN], C, Pays Betsileo route du Sud, km 292; ♂ [MNHN], MADAGASCAR CENTRE, 1700 m. [ca. 20.7667o S, 47.1833o E +/- 5 km] 14/2/1974, P. Viette et A. Peyrieras; ♂, E, Amboditavy, Ankazomivady, 20.77188o S, 47.16522o E +/- 1 km, 1760 +/- 50 m, 21/3/2003, D.C. Lees: BMNH (E) #697284 [DNA voucher; cytochrome b], KA535 [=KA-P535; DNA extract voucher]; ♂, C, 32 km S. Ambositra, 1400 m. [should be 1600+], 20.8167o S, 47.1833o E +/- 0.87 km, 1670 m, 12/1/1992, C. Kremen et al.: Genitalia 162; ♂, C, Ankazomivady, 20.8167o S, 47.1833o E +/- 0.87 km, 1670 m, 29/10/1992, C. Kremen: CK816; ♂, C, Ankazomivady, 20.769o S, 47.182o E +/- 0.25 km, 1750 +/- 50 m, 22/3/2003: 10:55; WG25 [wing image]; IA117 [isotope voucher]; ♂, C, Ankazomivady, 33 km S Ambositra, 20.769o S, 47.182o E +/- 0.25 km, 1750 +/- 50 m, 21/2/1995; D.C. Lees: DLKAZ 95-5A, IA185 [isotope voucher]; ♀, C, Ankazomivady, 35km S Ambositra, 20.769o S, 47.182o E +/- 0.25 km, 1730 m, 5/3/1995, D.C. Lees: DLKAZ 95-26; 1 egg expressed 5/3, IA186 [isotope voucher]; ♀, C, Ankazomivady, 20.769o S, 47.182o E +/- 0.25 km, 1750 +/- 50 m, 8/12/2004, DL-05-497, IA202 [isotope voucher]; ♀, data as above but DL-05-505, IA203 [isotope voucher]; ♀, E, Ranomafana National Park: Sahamalaotra, 21.2357o S, 47.3977o E, 1154 +/- 50 m, 20/12/2004, D.C. Lees: DL-05-922, BMNH (E) #676640 [DNA voucher]; ♀, NW, Bekolosy, 14.0458o S, 48.2962o E, 1163 m, 22/12/1994, D.C. Lees: DLBEK 94_183, IA334 [isotope voucher]; ♀, data as above but: 1130 m, 15/12/1994, D.C. Lees: DLBEK 94_170, IA335 [isotope voucher]; ♂, N, Tsaratanana, 7/10/2013, Ravo: DCL-14-028, IA330 [isotope voucher]; ♂, N, Tsaratanana, 14.158o S, 48.9525o E, 1932 m, 22/12/2004, D.C. Lees: BMNH (E) #671601; IA331 [isotope voucher]; ♀, NE, Anjanaharibe Sud, 1260 m, 4/2/1996, C. Kremen: CK703, IA135 [isotope voucher]; ♂, NE, Anjanaharibe Sud, 3/2/ 1996. H. Raharitsimba: TR 511, IA171 [isotope voucher]; ♀, NE, Anjanaharibe Sud, 4/2/1996: 09:15, C. Kremen: CK666, 1 egg, IA172 [isotope voucher]; ♂, NE, Anjanaharibe Sud, 14.7482o S, 49.4659o E, 1320, 3/2/1996, C. Kremen: CK662; DL 9667 [DNA voucher], IA332 [isotope voucher]; ♂, NE, Anjanaharibe Sud, 14.7524o S, 49.4777o E, 1130 m, 4/2/1996, C. Kremen: CK689, IA333 [isotope voucher]; ♂, NE, Anjanaharibe Sud, 14.7454o S, 49.4632o E +/- 0.4 km, 1400 +/- 50 m, C. Kremen: CK732, BMNH (E) 697885, 471 [= DL 0471, DNA extract number], KA581 [=KA-P581, DNA extract number].</p></div>	https://treatment.plazi.org/id/038747324C5EC67C1EB7292BFD3E27E9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C5AC67A1EB729EDFCB32348.text	038747324C5AC67A1EB729EDFCB32348.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis subsimilis	<div><p>Heteropsis subsimilis group</p><p>The ‘ subsimilis 7’ species group (after Pseudonympha subsimilis Butler, 1878) of Torres et al., (2001) resemble each other not only in wing pattern, but also in ♂ genitalia. This Ht. subsimilis group, which occurs within Malagasy rainforest, has also colonised the Comoros, as H. comorana (Oberthür, 1916), a species closely related to the generally abundant Ht. pauper (Oberthür, 1916) (Lees, 1997: 168; Torres et al., 2001). The sister grouping of the Ht. subsimilis group was unresolved in several earlier studies (Lees, 1997, Torres et al., 2001, Linares et al., 2009, Kodandaramaiah et al., 2010), although these studies found good support that the Ht. subsimilis species group forms a clade (97% based on COII parsimony analysis: Torres et al., 2001; 100% based on maximum likelihood analysis of COI, Ef1a and wingless: Kodandaramaiah et al., 2010). Linares et al., (2009) presented a supported phylogeography of relationships of three species. Recently though, Aduse-Poku et al., (2015, Fig. 1) found strong support for three sampled species namely Ht. subsimilis, Ht. pauper and Ht. comorana, which clade they recovered as sister to 13 sampled species of the Ht. strigula group, including Ht. tornado sp. nov. and the Ht. ankova subgroup (see below). ♂ genitalia have valve tips with a stout inward projecting spine that fall far short of the uncus extension; from LV, the uncus is inflated before its tip. The ♂ genitalia are in fact remarkably miniaturised compared to other Heteropsis, only around 1.7 mm in length (apart from those of Ht. tornado which is about 2 mm long, it can be seen in figures here that those of the Ht. strigula group are generally between 2–3 mm long and the Ht. drepana and Ht. antahala groups are more than three mm long and exhibit extension of particular parts). It may be hypothesised that such reduction comes with a relaxation of importance of this character system as a pre-mating isolation mechanism, as apparent in the Ht. subsimilis group, yet only in the Ht. drepana group is there a tendency to reduction or even loss (the supra-discocellular patch of Ht. narova) of parts of the androconial system (perhaps regained in Ht. drepana only). Atrophication of the genitalic system may be compensated for in a different, possibly androconial character system. Phenotypically, adult ♂♂ and ♀♀ of the Ht. subsimilis group are similar, nearly monomorphic except in size, but for simple androconia in the ♂ discocellular region and an inflated HW vein system also found in the Ht. strigula group. In the Ht. subsimilis group, this system features a set of linearly rather than bulbously inflated veins in the ♂ HW (Fig. 12; Lees, 1997: 96). As predominant in the Ht. strigula group, all species have the reduced ocellus configuration of the HWD, with only that in space CuA1 expressed as well as often that in space Rs; the morphological ‘total evidence’ phylogenetic hypotheses of Lees (1997: 156) that the Ht. subsimilis group is sister to only part of the Ht. strigula group (see also Torres et al., 2001) is presumably biased by the inclusion of this character system, whereas the unsupported Bayesian topology in Kodandaramaiah et al., (2010, Fig. 3) would imply that this configuration of HWD reduction is convergent between these two sister species groups (only with a gain of expression in HWD space-M 3 in Ht. roussettae sp. nov.). In any case, the two groups are sisters (Aduse-Poku et al., 2016, in press). Both sexes are attracted to ripe fruit as adults, as well as sometimes other sugar sources, and these low flying butterflies seem to specialise on grasses in the forest interior. There seem to be ridge and riparian specialists (Kremen, 1994).</p></div>	https://treatment.plazi.org/id/038747324C5AC67A1EB729EDFCB32348	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C5BC6651EB72FAAFA9326FE.text	038747324C5BC6651EB72FAAFA9326FE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis kremenae Lees	<div><p>Heteropsis kremenae Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:D5567C45-4EB9-4E62-8F67-ABF2C4008F4E</p><p>Prior references: sp. 14A (Lees 1997, Torres et al., 2001: 462).</p><p>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].</p><p>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].</p><p>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];</p><p>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.</p><p>Deposition summary: BMNH (HT ♂, 3 PT ♂♂, 3PT ♀♀); MNHN (2PT ♂♂, PT ♀); ABRI (PT ♂, PT ♀).</p><p>Type locality. Madagascar NE, Ambavaony R., R. Onive, E Masoala, 250 m, 15.282o S, 50.288o E +/- 0.1 km.</p><p>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).</p><p>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.</p><p>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.</p><p>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 ♀♀).</p><p>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.</p><p>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.</p><p>♂ 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.</p><p>Etymology. After Claire Kremen, who worked tirelessly for the creation of the Masoala National Park, that was created in 1994.</p><p>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).</p><p>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).</p><p>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).</p><p>Ecology and distribution.</p><p>Habitat: Primary rainforest, preferring riparian areas.</p><p>Behaviour: flies among low grasses, liking the slopes above streams and rivers.</p><p>Hostplant: not reared, but presumably low forest grasses. One was caught though in a canopy fruit trap in Marojejy (K. Aduse-Poku, pers. comm.).</p><p>Early stages: expressed eggs were greenish.</p><p>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.</p><p>Elevational range: 10–675 m (n=86, including referred specimens and observations).</p><p>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].</p></div>	https://treatment.plazi.org/id/038747324C5BC6651EB72FAAFA9326FE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C45C6611EB72DBAFED12521.text	038747324C45C6611EB72DBAFED12521.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis avaratra Lees & Kremen	<div><p>Heteropsis avaratra Lees &amp; Kremen, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:17C4A71B-C04D-4AC1-9407-4D52B343F803</p><p>Prior references: sp. 25 (Lees 1997: 64, Torres et al., 2001: 462); ‘ H. subsimilis ’ ♂, (d’Abrera 1980: 182; 1997: 221).</p><p>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].</p><p>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];</p><p>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;</p><p>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.</p><p>Deposition summary: BMNH (HT♂, 4 PT♂♂, 1 PT♀)), MNHN (2 PT♂♂, 1 PT ♀), ABRI (PT♂).</p><p>Type locality. Madagascar N, Montagne d'Ambre, 12.5289o S, 49.1727o E +/- 0.15 km, 1057 +/- 25 m.</p><p>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 .</p><p>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.</p><p>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).</p><p>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 ♀♀).</p><p>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.</p><p>Etymology. ‘avaratra’ means ‘north’ in Malagasy, referring to its distribution, as it is currently known.</p><p>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.</p><p>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 .</p><p>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).</p><p>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).</p><p>Ecology and distribution.</p><p>Habitat: montane primary rainforest, particularly on ridges/slopes.</p><p>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.</p><p>Hostplant: unknown, probably low grasses.</p><p>Early stages: unknown.</p><p>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).</p><p>Elevational range: 530–1630 m. (n=64, including referred specimens and observations).</p><p>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-&gt;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].</p></div>	https://treatment.plazi.org/id/038747324C45C6611EB72DBAFED12521	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C40C66E1EB72985FAE523DC.text	038747324C40C66E1EB72985FAE523DC.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis sogai Lees	<div><p>Heteropsis sogai Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:F0C54C40-DF74-4124-96B8-364E2F983C4E</p><p>Prior references: sp. 39 (Lees 1997, Torres et al., 2001: 462).</p><p>Type material., Deposition MNHN: Holotype: ♂ (Fig. 16 A), Madagascar Nord, massif du Tsaratanana, Matsabory, en dessous de l’Andohanisambirano [14.14o S, 48.97o E +/- 1 km, 2050 m], 5/ 8-XI-1966, 2050 m|mission au Tsaratatana XI-1966 Camp n o 1 P. Griveaud, P. Soga, P. Viette et D. Wintrebert|DCL-DB-3386.</p><p>Paratypes: Deposition MNHN: ♀ (Fig. 16 B), data as HT but DCL-DB-3387;</p><p>♂ (Fig. 15 C, genitalia), data as HT but DCL-DB-3388| 96 DL [genitalia]; ♀, data as HT but DCL-DB-3389.</p><p>Deposition summary: MNHN (HT ♂, PT ♂, 2 PT ♀♀).</p><p>Type locality. Madagascar Nord, massif du Tsaratanana, Matsabory, en dessous de l’Andohanisambirano [estimated at 14.14o S, 48.97o E +/- 1 km, 2050 m].</p><p>Diagnosis. Heteropsis subsimilis and Ht. avaratra are very similar, ventrally variegated species, whereas Ht. kremenae and Ht. pauper tend to have the base of FWV costa orange. However, the HWV Mb is much more jagged in Ht. sogai than any other species of the Ht. subsimilis group, whereas Ht. avaratra tends to have a yellowish ventral cast. HWV darker band of the central symmetry system (area between Mb and PMb) narrows more distinctly costad than in any other species of the Ht. subsimilis group. Two of FWV Cbs taper and meet tergad, demarcating pale ochreous area that is this relatively triangular rather than relatively oblong as in Ht. avaratra, for example. See below regarding androconial organs.</p><p>Description. Wings: Upperside with rather uniform mid brown background colour. FWD space-CuA1 ocellus with black iris spanning about ¾ of CuA1-CuA2 distance and with a large strongly hexagonal yellow to pale orange (grading to more orange tergad) ring that closely hugs the concave, rather sharply inflexed trace of the ventral Mb, and this ring is wider at its proximad, yellowest margin. FWD space-M1 ocellus small and surrounded by narrow black iris with a trace of an orange ring. FWD space-M2 ocellus expressed as a white spot. On the HWD, the space-CuA1 ocellus is the only one expressed there and is elliptic, occupying only about half of CuA1- CuA2, surrounded by a narrow yellowish ring. Underside with yellowish-ochreous background strongly variegated with brown irroration as detailed below. FWV space-CuA1 ocellus same size and shape as on upperside but with a trace of a comma-like tail in the inner, fused arm (Ore) of the yellow ring. FWV space-M1 and M2 and space-M3 also as on upperside but with yellower Orng, but HWV space-Rs ocellus in particular may be expressed as a white spot with narrow black iris where it is not noticeable on upper surface. FWV brown Mb irregular, jaggedly toothed where it bends back to mid costa. The HWV Mb is even more jagged, sharply inflexed in space-Rs, outdented at intersection with both veins M2 and M3, bending back sharply towards mid anal margin in space M3 and CuA1 and finally straighter towards about 4/5 along vein 1A. The HWV brown PMb somewhat mirrors Mb, but the jagged profile is relatively smoothed out. These bands/lines demarcate an area with much more intense fine brown irroration than either in the most basal section or distad of the Mb, where there is little brown irroration grading to a lot more in spaces M1-M2 (appearing scorched) and in marginad region around and beneath HWV space-CuA1 ocellus. The irregular medial brownish central symmetry system band so formed narrows distinctly towards the costa. A wavy diffuse line before the margin appears more obvious in the HWV and a narrow brown Sml closely follows the quite crenate margin which is alternately fringed brown at vein ends and ochreous at interveins (as evident in PTs). On the FWV the Mb and PMb similarly demarcate an intense brown irroration pattern that towards the margin is similarly suffused as in the HWV, whereas the transverse Cbs in the FWV cell form four or five double brown bands that around 1/3 distad along costa demarcate a roughly V-shaped (on the right side approximately ‘Africa’-shaped) yellow ‘triangle’.</p><p>Androconia: Sdb HWD dark brown but underlying patch has not been examined in detail. There is a not particularly bulbous anteriad inflation (but with 3–4 septa visible) in R near the anterior portion of the HWD cell, unlike for Ht. subsimilis and Ht. avaratra (sp. “25”). M2-2A veins in ♂ HW are all linearly inflated (Lees 1997: 96, Fig. 3 a, “39”).</p><p>Wingspan/fwl: range 30.5–34.7/ 18.1–19 mm (n=2 ♂♂), including HT ♂: 34.7/ 19 mm; range 33.8–34/ 19.3– 20.8 mm (n=2 ♀♀).</p><p>♂ genitalia: 96DL, PT, MNHN (Tsaratanana; Fig. 15 C). Small, about 1.6 mm long, and with configuration typical of the Ht. subsimilis group; no obvious difference in shape of the genitalic components from other group members (Lees, 1997: 106) and the description below might apply to almost any of the group. From LV (DV not available), the valve bases are rather symmetrically-‘skittle’-shaped (Lees 1997). Uncus is slightly longer than tegumen, its dorsal profile relatively downbent, and slightly convex on ventral margin before the slightly pointed downturned tip. Gnathos with little dorsal serration, tapered from small base inbent at right angled, unlobed, ‘support member’, fairly straight, not reaching tip of uncus, and oriented about 30 degrees to it. Valve with rounded dorsal shoulder, arm about half length of valve, tapering to distinct ‘beak’ that points to mid-uncus with limited serration on small, not much expanded ‘club’ with serrae at end. Saccus relatively small, tapered to obliquely truncated (slightly bulbous in another preparation, 134BDL). Aedeagus slightly longer than valve, not particularly thin, only slightly uprecurved recurved distad of ostium.</p><p>Etymology. After the remarkable Malagasy collector Pierre Soga, who climbed many mountains in search of Lepidoptera, joining several expeditions of French lepidopterists, and was involved in collecting these paratypes. To my astonishment, I managed to find and meet P. Soga down on the way back from Mangindrano (S. of RNI Tsaratanana) on Christmas Eve, 2004; he was infirm at this time.</p><p>Discussion. This species was first recognized by myself in April 1994 in MNHN from specimens collected at Tsaratanana in 1966 by P. Griveaud et al., (Lees, 1997: 65). As a possible endemic of Tsaratanana, it is understandable that there is such little material of this species, all from this single collection in MNHN, except for one specimen in BMNH dating from before 1924, which is not included in the type series as it has a dubious locality label (see below). Due to its similarity to the relatively widespread Ht. subsimilis, it is also not surprising the species has not been described until now. All other available types of the Ht. subsimilis group have been examined (see under Ht. kremenae). As well as the two ♂♂ and two ♀♀ of Ht. sogai from Tsaratanana, there is a single ♂ specimen labelled Beforo SE Madagascar in BMNH. When I revisited the type locality (now, of Ht. sogai) in December 2004, I failed to refind this species, but neither did I rediscover another apparent endemic to the Tsaratanana massif, Strabena tsaratananae Paulian, 1951, for which, suspiciously, the BMNH collection includes two specimens with the same locality and accession label. It is not clear that these Beforo-labelled specimens were really collected in SE Madagascar and no place name has been traced exactly as such (in any case, the similar sounding locality ‘Beforona’ is at far too low elevation). Furthermore, another apparently narrowly localized species, S. perrieri Paulian, 1951 bears the same locality and accession label.</p><p>Additional information. DNA divergences: unsequenced. No specimens have been found since 1966 (expedition in 2004 of the author to RNI Tsaratanana including the type locality; and that in late 2013 of R. Ranaivosolo, pers. comm, to an adjacent area, provided no exemplars).</p><p>Phylogeny/sister species: in the morphological analysis of Lees (1997) this taxon (“THTNN”) connected to Ht. avaratra (“TWNFV”), which might possibly be its sister (as in combined tree in Aduse-Poku et al., 2016, in press).</p><p>Ecology and distribution.</p><p>Habitat: ‘Matsabory’: Malagasy for a marshy area or small lake within forest.</p><p>Behaviour: unreported.</p><p>Hostplant: unknown.</p><p>Early stages: unknown.</p><p>Distribution: RNI Tsaratanana, where putatively endemic (Fig. 30 D, mid green dots). See above regarding other localities.</p><p>Elevational range: 2000–2050 m (n=4).</p><p>Referred specimens. ♂ [BMNH], Beforo, SE Madagascar, 61.23 Rothschild coll.|134 DL [genitalia].</p></div>	https://treatment.plazi.org/id/038747324C40C66E1EB72985FAE523DC	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C4FC66E1EB72FD1FAB4267D.text	038747324C4FC66E1EB72FD1FAB4267D.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis strigula	<div><p>Heteropsis strigula group</p><p>The Ht. strigula species group (the ‘ strigula 15 ’ group of Torres et al., 2001), now including Ht. tornado described here (Aduse-Poku et al., 2015) is a monophyletic clade, consisting of a diverse, primarily rainforest-interior, radiation of butterflies 100% endemic to Madagascar (a few species occur in mesic parts of deciduous forest, or in swampland at the margins of forest). This clade is at least partially supported by morphology (Lees, 1997: 156) although it was weakly or not at all (Torres et al., 2001) supported by mitochondrial DNA (COII and cytochrome b). Eight species sampled within it that include Ht. ankova (Ward, 1870) placed by Lees, 1997 in a separate group, (the Ht. ankova clade of the Ht. strigula group that sometimes included Ht. tornado) are united by a single C-&gt;T transversion in the highly conserved 28SDDFF region that is not observed elsewhere in the Malagasy satyrines (dataset of Monaghan et al., 2009). Now with the ten gene study of Aduse-Poku et al., (2015, see Fig. 1 and Suppl. File S4) that included 13 sampled members, the slightly expanded Ht. strigula group is shown to be the wellsupported sister of the Ht. subsimilis clade, which might possibly be (with submarginal bootstrap support) sister to a clade consisting of Ht. narcissus ( sensu Balletto &amp; Lees, 2012) + Ht. ankaratra . Five sampled members of the Ht. strigula group were submarginally supported (pp=0.86) in the three-gene study of Kodandaramaiah et al., (2010). As for the Ht. subsimilis group, ♂♂ and ♀♀ are similar, apart mainly from size and androconial and/or inflated HW vein differences. ♀♀ of some species are as yet unknown. Both sexes are attracted strongly to fruit as adults, do not fly high, and seem to specialise on forest (or sometimes marshland) grasses. Most species (apart from Ht. tornado, and from the members Ht. ankova, Ht. turbata (Butler, 1880) and Ht. pallida (Oberthür, 1916) that were recovered as sister to the rest of the group in Lees, 1997: 156, although equivocally, see Lees, 1997: 168, and also apart from Ht. roussettae described below) show reduction in ocellus expression in the HW, so that only the space- CuA1 ocellus is expressed, as apparently parallelled in the Ht. subsimilis group, as described above. Var i ou s species show from LV a combination of relatively straight or gently sinuate ‘neck’ to the uncus before the apical hook, a relatively asymmetric valve base and the extensive development of spinoid setae on the valve tips, and a few members (belonging to Ht. angulifascia clade) show a striking planification of the gnathos into an ear-like structure, and wavy androconial tufts from HW space-CuA2 sometimes correlate with ventral androconial scale patches. The main radiation occurs throughout the eastern rainforest biome, but one member, Ht. maeva (Mabille, 1878), also occurs in Western Madagascar. Ht. turbata and Ht. pallida are unusual in that they occur outside forest margins in marshy areas, whereas Ht. ankova as well as Ht. lanyvary sp. nov. fly just inside the forest margin.</p></div>	https://treatment.plazi.org/id/038747324C4FC66E1EB72FD1FAB4267D	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C4FC6691EB72A7CFAA72543.text	038747324C4FC6691EB72A7CFAA72543.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis tornado Lees, Allaoui & Aduse-Poku	<div><p>Heteropsis tornado Lees, Allaoui &amp; Aduse-Poku, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:22EB2840-C7CE-42D5-B26F-3DDD0DF3D763</p><p>Prior references: sp. 54 (Lees 1997, Torres et al., 2001: 462—where considered to belong to an unsampled clade); “Heteopsis_sp.nov_ tornado ” [sic, nomen nudum] in Aduse-Poku et al., 2015.</p><p>Type material., Deposition BMNH: Holotype: ♀ (Fig. 17 A), Madagascar NE, R.S. Anjanaharibe Sud, slope above R. Marolakana, road between Befingotra and Analamazava, 14.7676o S, 49.4815o E +/- 0.01 km, 1000 +/- 5 m, 20/11/2014: 09:07, Ahamadi Allaoui. DL 14A-0245, IA518 [isotope voucher], NHMUK 010289143 [QTR barcode].</p><p>Paratype: Deposition ABRI: ♀ (Fig. 17 B), Madagascar NE, P.N. Marojejy, 670 m, [14.4354o S, 49.7647o E; using GPS point 30B672], ridge above “Vallée de Brookesia” below “Camp 2 Marojejy” and above Manantenina river, primary rainforest, on path, 29/01/2014, Kwaku Aduse-Poku, MAD 1403 [=KAP-MAD1403; DNA voucher]|KA1015 [=KA-P1015; extract number]|IA313 [isotope voucher].</p><p>Deposition summary: BMNH (HT ♀); ABRI (PT ♀).</p><p>Type locality. Anjanaharibe Sud, slope above R. Marolakana, 14.7676o S, 49.4815o E +/- 0.01 km, 1000 +/- 5 m.</p><p>Diagnosis. The species is very distinctive especially on the underside (see Fig. 17) and not readily confused with any other species. It has a superficial resemblance though on the dorsal surface to Ht. iboina . Ht. tornado is significantly larger than the typical small size of the four known members of (see Lees, 1997) of the Ht. strigula group ( Ht. ankova, Ht. lanyvary, Ht. turbata and Ht. pallida) (three members of which formed a paraphylum in the study of Aduse-Poku et al., 2015), which like Ht. tornado routinely show expression of HWD ocellus M3. Including referred specimens, the ♂ and ♀ are similar, but the three known ♀♀ are larger than the one known ♂.</p><p>Description. Wings: upperside uniform mid brown, with a reddish-orange flush in the basal areas of both wings. FWD space-CuA1 ocellus with black iris more than spanning veins CuA1-CuA2. Orng very large, subhexagonal, orange, spanning from upper part of space-M3 down to upper half of 1A. Space-M1 ocellus is slightly elliptic (especially on the HT right FW) and quite large in comparison with Malagasy Heteropsis, with a quite wide black iris and narrow orange ring, which extends slightly into space-M2. In HWD three subelliptic ocelli are expressed, the smallest in space-M2, but whose ring is almost contiguous with the ring of the largest of these ocelli in space-M3 whose ring nearly spans veins M3-CuA1, the second largest and most elliptic below it in space-CuA1 spanning about ¾ of inter-vein distance CuA1-CuA2. This is an eyespot expression configuration seen in the Ht. ankova clade, which is reminiscent also of ‘ Telinga ’ oculus (Marshall, 1880) in the Western Ghats. Proximad of the smallest HWD ocellus, whose ring is about half the diameter of the largest, an orange Mf (median fleck) or crescent mark hugs vein M3 as far as its intersection with Mb (faintly visible on HWD). This Mf is a characteristic of quite a few species of the Ht. strigula group, most markedly so in Ht. strigula . HW distinctly crenate (as strongly so as in Ht. bicristata), outwardly reflexed to each darker part of the fringe (forming in effect very convex tails) at the end of veins Rs-CuA1. Underside has an ochreous yellow cast, irrorated with rufous brown especially proximad of the Mb. FWV space-CuA1 Orng varying from light orange to darker orange towards the tergum, where occupying the width of space-1A, its Ore (ocellus ring extension) forming a ‘tornado’ pattern comprising the pale light orange-yellow outer arm of, in effect, a spiral., The paler yellowish colour of this outer arm of the spiral contrasts with the eccentric, elliptic to sub-hexagonal shape of the FWV Orng; such a pattern is evident, with the proximad yellow arm not usually as well separated costad, in a few other Malagasy Heteropsis which would be placed in different clades, such as Ht. iboina, Ht. kremenae, and Ht. undulans (Oberthür, 1916) . This yellow arm follows a strongly concave russet-brown Mb before it bends back sharply towards near-mid costa. FWV space-M1 ocellus smaller than on upperside, with narrow yellowish ring. HWV space-Rs-M2 ocelli not expressed and HWD orange crescent in space-M2 at base of space near the Mb appears as a prominent light Mf. Space-M3 and CuA1 ocelli about equal sized occupying slightly less than half of respective intervein-spans and with narrow yellowish rings faintly bordered by a rufous brown ring, subelliptic. Space-CuA2 ocellus not obviously expressed. HWV Mb russet, irregular, most angularly convex in space-M2, fairly smoothly concave in space-M3-CuA1, and convexly angled returning towards near mid-anal margin, but terminating just below vein CuA2. Ochreous background more evident distad of HWV Mb, while strong brown irroration is slightly more intense in basal areas of both wings than in distad areas. The russet PMb in HWV crudely mirrors the Mb, but is slightly straighter. A diffuse wavy brown line shadows margin in both wings, but is stronger on FWV, and a narrower brown Sml closely follows it, then closely delineating the crenate margin; before chequered brown and ochreous fringe runs a still darker brown line along the margin in both wings. Four transverse brown Cbs in the FWV cell delineate a more ochreous and weakly irrorated band in the middle of the cell, while the costa itself is quite evenly strigulated. Variation. The ♀ paratype very similar in pattern to the HT ♀. In this specimen, the HWD space-M2 orange Mf is even more obvious. It has a somewhat rounder FWD Orng and a small black spot at the anterior part of this ring; this ‘satellite’ eyespot not seen in either the HT nor referred specimens. On the FWV of PT, ‘tornado’ effect (for Ore) is more extreme, and Mb angularly inflexed rather than smoothly concave proximad of the yellow Ore. The HT was collected effectively in a late dry season, whereas the PT collected well into the wet season, so there is no evidence as yet for seasonal variation. The ♂ (not included in type series) is similar to the HT ♀ but slightly smaller. Like the referred ♀, the referred ♂ is very consistent in wing shape and pattern, although slightly smaller (35 mm wsp/ 18.5 mm fwl). However on its HWV, ocelli spaces Rs-M2 are expressed as white points, so perhaps they were collected in a drier season. In the referred specimens also, no orange Mf is obvious in HWD space-M2.</p><p>Androconia: no ♂♂ in type series/not examined.</p><p>Wingspan/fwl: 37.15/ 20.07 mm (HT ♀); 34.9 mm / 19.5 mm (PT ♀).</p><p>Palps: penultimate segment with whitish stripe close to compound eye, fringed with darker brown hairscales above where in potential contact. A light ochreous stripe away from eye is fringed by brown hair-scales. Mesad face of labial palps whitish.</p><p>Etymology. Refers in particular to the tornado-like markings, the sporadic rarity of this species in collections, and also alludes to the chance survival of an apparently deeply branching species in cyclone-prone Madagascar. The impression is of the discovery in Madagascar of a relictual member of a greatly diversified lineage (Aduse- Poku et al., 2015, Fig. 1). In different endemic radiating mycalesine clades in Madagascar, there is a trend towards reduction of the number of ocelli expressed in the upperside HW, so its configuration might also be correlated with a typically more open habitat, such as the marshy areas typical for Ht. turbata .</p><p>Discussion. This species was first discovered by myself in an uncurated drawer of the Oberthür collection (as it was arranged at BMNH in May 1994), with a unique ♂ and ♀ labelled as from ‘Mahassabe’ (see Referred specimens), collection dating pre-1923 (Lees, 1997: 65). Because of the distance of the locality of these specimens from recent material and the uncertainty of the labels (see also above under Ht. sogai) they are not actually included in the type series. The exceptional rarity of this distinctive species, despite intensive prior sampling from the 1890s to the present in the Fianarantsoa (?) and Marojejy-Anjanaharibe Sud areas, is noteworthy. Perhaps this species has naturally small populations, or normally occupies a particular niche that has not been properly sampled and difficult of access, such as open cliff slopes, like the spectacular habitat facing ‘Camp II Marojejya’. Mahasoabe (‘Mahassabe’; Fig. 30 D, lower brown dot) is a village located at around 1100 m, and its surroundings are largely deforested today, but the nearest primary forest is over 20 km to the east, and assumed plantations or secondary forest can be viewed on Google Earth from 4 km to the west. The corresponding printed label is perhaps anyway as mentioned before unreliable, considering that that the same label is placed on series of characteristically northern and lowland species like Ht. erebina . The species is distinctive enough to be unmistakable, but all available types in the Ht. subsimilis and Ht. strigula groups were examined. In particular I checked the STs of Culapa ornata Oberthür, 1916 (which was synonymised by d’Abrera, 1980: 182 with Ht. turbata (Butler, 1880); HT ♀: ‘Madagascar’, Cowan, BMNH(E) 673769). The LT ♂ of Culapa ornata (Fig. 19 E, representing Ht. turbata), here designated: BMNH(E) 673678, Madagascar; PLT ♂, Antananarivo, R. Toy: BMNH(E) 673670)) and syntypes of Culapa pallida Oberthür, 1916 (LT ♂, missing abdomen, Fig. 19 D, here designated, bearing label: “ Madagascar, Antsianaka, Perrot Freres, 2e Semestre 1890”|BMNH(E) 673775”; PLT ♂, BMNH(E) 673779 and ♀♀ BMNH(E) 673776 (illustrated in Oberthür, 1916, Pl. 356: f. 3054), BMNH(E) 673778 and BMNH(E) 673780, all from Madagascar, Antsianaka, Perrot Freres, 2e Semestre 1893; there are also two potential ST specimens of C. pallida not labelled ‘Antsianaka’), were also examined. Also, considering a superficial resemblance, the syntypes of Mycalesis iboina Ward, 1870 in BMNH (Coll. Ward) (LT ♂, here designated: BMNH(E) #674853, PLTs, BMNH(E) #674854-674856) and MNHN (Coll. Galichon) (PLT ♂♂) were examined; also the “HT” ♂ of Culapa parva Butler, 1879 (BMNH(E) #674859, ex Grose-Smith) and the ♂ and ♀ STs of its synonym (Lees, 1997; Lees et al., 2003), Mycalesis irrorata Mabille, 1880 (LT ♂, here designated: BMNH(E) #674860; ex Coll. Grose-Smith; PLT ♀, BMNH(E) #674861, “Madag” ex Coll. P. Mabille).</p><p>Additional information. ♂ genitalia: 362DL (referred specimen, Fig. 18 A): about 2 mm long; from LV, tegumen with bent down to fairly straight uncus leading to downward hook at tip without a distinct ‘head’. Gnathos from square base narrow and tapered, slightly uprecurved to tip. Hinge with vinculum wider than half maximum dimension of tegumen. Saccus rather short, stubby and bulbous. Aedeagus distinctly longer than valve, fairly stout, uprecurved distad of ostium. Valves fairly short, arms uprecurved and leading to club, which has spinoid setae at the end and a distinct ‘beak’ pointing dorsad/mesad. The overall impression is a similarity to genitalia of some of the Ht. strigula group (valves not symmetrically ‘skittle’-shaped’ from LV as in Ht. subsimilis group and rather uncus is more straight from LV; Lees 1997: 106, Fig. 7 f). The genitalia of Ht. turbata are actually quite similar to those of Ht. tornado .</p><p>DNA divergences: COI-5P cluster number BOLD:ACW4939 (exemplar BMAD192-15, DL14A-0245), closest to members of Ht. strigula group, e.g. 6.7% divergent to Ht. barbarae sp. nov and 8.3% to Ht. turbata and a similar distance to Ht. ankova .</p><p>Phylogeny/sister species: Lees (1997: 156) had over 96% jackknife support for a sister relation of Ht. tornado (“FFTFR”) with the Ht. ankova ‘clade’ of (then three) species, but this resolution appeared only in the total evidence parsimony analysis. The species would therefore be tentatively placed in the Ht. strigula group based on morphological features and support. Molecular sequencing of multiple genes provides evidence Ht. tornado is sister to all others of the Ht. strigula group, including the ‘ ‘ Ht. ankova ’ sub-group (Aduse-Poku et al., 2015).</p><p>Ecology and distribution.</p><p>Behaviour: largely unknown. The HT was found resting on the ground in the morning at the edge of a road close to primary forest; the paratype ♀ was sitting on a path above a steep slope in primary forest, presumably attracted by fallen fruit.</p><p>Hostplant: unknown.</p><p>Early stages: unknown.</p><p>Distribution: Marojejy, Anjanaharibe Sud; (?) Mahasoabe, SE of Fianarantsoa (Fig. 30 D, dark brown dots).</p><p>Elevational range: 675 (PT ♀)– 1000 m. (HT ♀) [possibly to ca. 1100 m. at “Mahassabe”]?</p><p>Habitat: one of the PT ♀♀ was caught in tall canopy primary forest on a path above a steep slope and small stream, about 100 m above the Manantenina river at Marojejy. The HT ♀ was caught on a slope at the edge of a currently largely impassable road, close to an island in the road and next to some primary vegetation and ginger, above a steep stream draining into marshland bordering the Marolakana River which was only around 0.15 km away. No other individuals could be found despite repeated searches in these surrounding habitats.</p><p>Referred specimens. ♂ [BMNH], Mahassabe [Mahasoabe], forêt au S.-E de Fianarantsoa Madagascar, ex Lamberton, 1922|Ex Oberthür Coll. Brit. Mus. 1927-3|362 DL; ♀ [BMNH], same data but BMNH (E) #674881.</p></div>	https://treatment.plazi.org/id/038747324C4FC6691EB72A7CFAA72543	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C48C6691EB729A7FABD24EA.text	038747324C48C6691EB729A7FABD24EA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis ankova	<div><p>Ht. ankova sub-group</p><p>Aduse-Poku et al., (2015, Fig. 1) recovered Ht. ankova and Ht. turbata + Ht. pallida within a grade at the base of the Ht. strigula group, so it is not certain that this sub-group is monophyletic, unlike the Ht. strigula sub-group (see below). In Lees (1997: 156), these ‘ Ht. ankova subgroup’ species formed a morphological clade with Ht. tornado (‘FFTFR’). All these species feature expression of an ocellus in space-M3 and sometimes space-M2 of the HWD. The species occur in marshy areas near rainforest margins ( Ht. turbata and Ht. pallida) or in rainforest, sometimes within metres of forest margins, always flying and perching quite low, from sealevel to over 2000 m elevation.</p></div>	https://treatment.plazi.org/id/038747324C48C6691EB729A7FABD24EA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C48C6161EB728DCFCB92691.text	038747324C48C6161EB728DCFCB92691.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis lanyvary Lees	<div><p>Heteropsis lanyvary Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:264464C2-C4E2-4470-A209-348CDCCBDF85</p><p>Prior references: sp. 67 (Lees 1997: 66, where considered the same species as Ht. ankova;</p><p>sp. 68 [label on specimen in PBZT; “sp. 68” was an unrelated morphospecies in Lees, 1997: 66].</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 19 A), Madagascar NW, Tsaratanana, piste Mangindrano-Antetykalambazaha, 14.187o S, 48.945o E +/- 1 km, 1615 +/- 75 m, flat grass-bamboo area, 20/12/ 2004 12:04, D.C. Lees: DL-05-744, TF19 [leg for DNA], NHMUK 010289144 [QTR barcode].</p><p>Paratypes: Deposition BMNH: ♀ (Fig. 19 B), Madagascar NW, Tsaratanana, piste Mangindrano- Antetykalambazaha, flat grass-bamboo area, 14.187o S, 48.945o E +/- 1 km, 1615 +/- 75 m, 20/12/2004 12:04, D.C. Lees: DL-05-745, TF14 [leg for DNA], NHMUK 010289145 [QTR barcode]; ♂, same data as HT but DL-05-849, TF [legs for DNA] [abdomen lysed], IA624 [isotope voucher], MAD 233-15 [DNA barcode voucher], NHMUK 010289146 [QTR barcode]; ♂, same data as HT but DL-05-846, TF12 [tissue sample for DNA], BMAD 232-15 [DNA barcode], IA294 [isotope voucher], NHMUK 010289147 [QTR barcode]; ♂ (Fig. 18 B, right), same data as HT but DL-05-845, IA192 [isotope voucher], [thoracic fragment for KAP], NHMUK 010289148 [QTR barcode]; ♂, same data as HT but DL-05-856, IA623 [isotope voucher], NHMUK 010289149 [QTR barcode]; ♂, NW, Tsaratanana, near forest margin, 14.2034o S, 48.9532o E +/- 2.5 km, 1441 +/- 25 m, 24/12/2004, D.C. Lees: DL-05-851, IA622 [isotope voucher], NHMUK 010289150 [QTR barcode]; ♂, NW, Tsaratanana, Antetykalambazaha, near camp, 14.1958o S, 48.9497o 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]; B43 [‘andromap’]. ♂ (Fig. 18 B, left), NW, Tsaratanana, [Ampidiranala], 14.1958o S, 48.9497o 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, 14.1526o S, 48.9578o 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, 14.199o S, 48.952o E +/- 0.5 km, 1550 +/- 65 m, 20/12/2004, D.C. Lees: DL-05-747, TF13 [leg for DNA], NHMUK 010289151 [QTR barcode]; ♂, NW, Tsaratanana, Ampidiranala, 14.1958o S, 48.9497o 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], 14.1958o S, 48.9497o 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, 14.2047o S, 48.9538o E +/- 0.35 km, 1456 +/- 25 m, just into primary forest, 20/12/2004: 20:15, D.C. Lees: DL-05-734A, TF17 [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-F12 [DNA barcode voucher];</p><p>Deposition MNHN: ♂ [MNHN], Madagascar Nord, TSARATANANA. 1,800 m (Forêt de Mousses), [14.17o S, 48.94o E +/- 4 km, 1800 m] 10/1949, R. Paulian: DCL-DB-3301;</p><p>Deposition ABRI: ♂ [ABRI], Madagascar NW, Tsaratanana Mntn N. madagascar below Andohanisambirano, S 14o 12.486' E 0 48o 58.225' 4167 ft, [14.2081o S, 48.9704o E +/- 1 km, 1291 +/- 50 m], 24/10/2003, S. C. Collins: 1204 [leg for DNA];</p><p>Deposition PBZT: 2 ♂♂ [PBZT], Madagascar NW, Mt Tsaratanana, 750 a 1400 m, [14.21o S, 48.96o E +/- 5 km, 1400 m.; elevation as low as 750 m not found in region] 2/1951, R. Paulian.</p><p>Deposition summary: BMNH (HT ♂, 15 PT ♂♂, 1 PT♀), MNHN (1 PT ♂); ABRI (1 PT ♂); PBZT (2 PT ♂♂).</p><p>Type locality. Madagascar NW, Tsaratanana, near forest margin, 14.2034o S, 48.9532o E +/- 2.5 km, 1441 +/- 25 m.</p><p>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.</p><p>Description. Wings: upperside uniform dark brown, with FW space-CuA1 ocellus with a wide orange ring which is generally quite concentric but apparently a little wider and more angled proximad. FWD space-M1 ocellus small and slightly elliptic with very narrow orange ring. HWD space-CuA1 ocellus is the main one expressed there, as well as to lesser extents those of space-M3 and space-M2, as in Ht. ankova . Hairbrush around space-1A 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-CuA1 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 M3 (a feature also displayed by Ht. ankova). FWV space-M1 ocellus rather small and slightly elliptic with a white pupil with narrow black iris and with a narrow pale orange ring. On HWV, space- CuA1 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-M2. A reddish PMb mirrors shape of Mb at least as far as vein 1A. 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 M3 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 M1 and M2 often visible as white points. Sexes similar.</p><p>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♀).</p><p>Androconia: Sdb dark reddish brown with grey tip. Androconial patch directly above the end of the R stem small, blackish and lenticular. Veins 1A+2A narrowly inflated for most of length, that in 3A slightly more so (more so than in Ht. ankova) and covered in narrow greyish brown scales. No other androconia obvious.</p><p>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.</p><p>♂ genitalia: DL-05-761, PT (Fig. 18 B, left); DL05-845, 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).</p><p>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.</p><p>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).</p><p>Additional information. DNA divergences: COI-5P barcode cluster number BOLD:ACW4935, about 3.2% divergent to Ht. ankova (cluster number BOLD:AAK5841, exemplar BMNH(E) 676678, BMAD048-09, HM404225 from Analamazaotra).</p><p>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).</p><p>Ecology and distribution.</p><p>Habitat: montane rainforest, forest interior, close to margins.</p><p>Behaviour: flies low.</p><p>Hostplant: unknown, possibly low grasses.</p><p>Early stages: unknown.</p><p>Distribution: endemic as far as is known, to the Tsaratanana massif (Fig. 30 C, olive dots).</p><p>Elevational range: 1290–1800 m. (n=17 including referred specimens and observations).</p><p>Referred specimens. ♂, Madagascar N, Tsaratanana, 14.2034o S, 48.9532o E, 1441 m, 24/12/2004, D.C.Lees: DL-05-859, IA624 [isotope voucher]; ♂, NW, Tsaratanana, 14.2034o S, 48.9532o E +/- 2.5 km, 1441 +/- 25 m, 12/ 2004; BMNH (E) #671598 [DNA voucher; cytochrome b], KA504 [=KA-P504; DNA extract number]; ♂, NW, Tsaratanana, Ampidiranala, 14.20025o S, 48.95175o E +/- 0.54 km, 1542 +/- 85 m, 20/12/ 2004, R. Ranaivosolo: DL-05-756, BMNH (E) #676686 [DNA voucher; cytochrome b].</p></div>	https://treatment.plazi.org/id/038747324C48C6161EB728DCFCB92691	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C34C6151EB72DBAFC3D202F.text	038747324C34C6151EB72DBAFC3D202F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis strigula	<div><p>Heteropsis strigula sub-group</p><p>This sub-group of the Ht. strigula group includes all the members of the clade comprising both Ht. strigula (Mabille, 1877) and Ht. maeva but not Ht. ankova in Aduse-Poku et al., (2015) . Its species exhibit loss of expression of HWD space-M3 ocellus together with various degrees of broadening of the valve tip into a club like, truncate or cleft structure bearing numerous spinoid setae (Lees, 1997: 107).</p></div>	https://treatment.plazi.org/id/038747324C34C6151EB72DBAFC3D202F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C34C6111EB72C8FFDC9260E.text	038747324C34C6111EB72C8FFDC9260E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis barbarae Lees & Kremen	<div><p>Heteropsis barbarae Lees &amp; Kremen, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:DC87986A-3822-43C9-B1B9-83A2714D088B</p><p>Prior references: sp. 16 (Lees 1997; Torres et al., 2001: 462).</p><p>Type material., Deposition MNHN: Holotype: ♂ (Fig. 20 A), Madagascar NE, Maitsoarongana, Sikiory River, near camp, 15.1885o S, 49.5163o 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].</p><p>Paratypes (accession BMNH (E) 2008-69): ♂, Madagascar NE, Maitsoarongana, Sikiory R., 660–750 m, 15.1907o S, 49.5309o 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, 15.1885o S, 49.5163o 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, 15.1907o S, 49.5309o 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, 15.1848o S, 49.4659o 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, 49.5309o E, +/- 2 km., 750 +/- 100 m., 13/12/2001, D.C. Lees: DL 01-248, BMNH (E) 1717095 [QTR barcode].</p><p>Deposition summary: BMNH (HT ♂, 4 PT ♂♂, 1 PT ♀).</p><p>Type locality. Madagascar NE, Maitsoarongana, Sikiory River, 15.1885o S, 49.5163o E +/- 0.02 km, 685 +/- 23 m.</p><p>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).</p><p>Description. Wings: Upperside mid-brown with a slight russet brown cast. Space-CuA1 ocellus expressed strongly in FWD; orange ring sub-hexagonal., M1 ocellus very small black white pupilled spot surrounded by a narrow orange ring. HWD ocellus CuA1 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- M2 to CuA1. FWV space-CuA1 ocellus similar size to FWD (in the HT), as is FWV M1 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-CuA1 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-CuA1 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 1A. 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-CuA1 ocellus FWD is a little wider proximad in some specimens, and FWD M1 ocellus sometimes not visible. HW space-CuA1 ocellus is more circular in some specimens.</p><p>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).</p><p>Androconia: (observations on ♂ specimen MAD182 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 1A+2A base, mid brown, dispersed, juxtaposed potentially under abdominal sternal segments A2–A5, 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 2A swollen within basal half.</p><p>Palps: Ochreous, with intermingled brown scales on outside face away from compound eye, dark brown scales above where potentially brushing eye.</p><p>Etymology. After Barbara, the mother of Claire Kremen.</p><p>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 |MadagascarNo10 [?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.&amp;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.&amp;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 ”.</p><p>Additional information. ♂ genitalia: 141DL (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.</p><p>DNA divergences: COI-5P cluster number BOLD:ACW4995 (exemplar BMAD248-15, DL14M-0033, Marojejy) is 4.86% divergent to Ht. tianae (BOLD:AAE4112), and 5.63% divergent to Ht. maeva (BOLD:AAE5488), 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).</p><p>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).</p><p>Ecology and distribution.</p><p>Habitat: riparian primary rainforest.</p><p>Behaviour: flies low and both sexes come easily to fruit bait.</p><p>Hostplant: unknown but almost certainly low grasses.</p><p>Early stages: unknown but for expressed eggs, which were whitish.</p><p>Distribution: endemic to the northeastern rainforests of Madagascar, from Marojejy and Masoala south to Ambatovaky (Fig. 30 C, light green dots).</p><p>Elevational range: 0– 890 m. (n= 153, including referred specimens and observations).</p><p>Referred specimens. ♀, Madagascar NE, Marojejy PN, Camp II [above], 14.4344o S, 49.759o E +/- 0.25 km, 850 +/- 50 m, 12/11/2006: 10:00, D.C. Lees: DL 06-298; ♂, NE, Marojejy PN, 14.4344o S, 49.7606o 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, 14.4347o S, 49.7601o E +/- 0.5 km, 730 +/- 50 m, 10/11/2006, D.C. Lees: DL 06-169; specimen, NE, Marojejy PN [between camp 1 and camp2], 14.4362o S, 49.7679o E +/- 0.85 km, 613 +/- 125 m, 10/11/2006, 15:58, D.C. Lees: DL 06-170; ♀, NE, Marojejy PN, 14.4377o S, 49.7756o E +/- 0.25 km, 400 +/- 50 m, 20/11/2006: 15:07, D.C. Lees: DL 06-456; ♂, NE, Marojejy PN, 14.4425o S, 49.7818o 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, 14.45015o S, 49.786o E +/- 1.78 km, 326 +/- 175 m, 10/11/2006: 13:16, D.C. Lees: DL 06-204; ♂, NE, Marojejy PN [around entrée], 14.4626o S, 49.7964o 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], IA387 [isotope voucher]; ♂ [BMNH], NE, Andranobe, Masoala Peninsula, 15.6816o S, 49.9573oE +/- 1.31 km, 0 m, 3-Dec-91, C. Kremen: CK826, BMNH (E) 2008-69; ♂, NE, Masoala, Anaovandrano, 14/1/1994, D.C. Lees: WG5 [wing image], IA38 [isotope voucher]; ♂, NE, Masoala, Ambavaony, 125 m, 27/11/1994, D.C. Lees: DCL 47a, IA108 [isotope voucher]; ♂, NE, Masoala, Ambanivony, 25/11/1993, C. Kremen: CK26, IA109 [pheromone voucher]; ♀, NE, Makira, Vohitaly, 15.4414o S, 49.5309o E +/- 0.15 km, 645 m, 30/12/2002, D.C. Lees: DL-SF00, IA112 [isotope voucher]; ♀, NE, Makira, Ankirindro, 15.2931o S, 49.5472o E +/- 0.15 km, 675 +/- 25 m, 15/1/2003, D.C. Lees: DL- SE00, IA114 [isotope voucher]; specimen, NE, Ankiatomboka forest, c. 710 m, 15.179o S, 49.412o E, 710 +/- 50 m, 9/12/2001, D.C. Lees: DL 01-169; ♂, NE, Bivontro, Makira, 15.3984o S, 49.4477o E +/- 0.1 km, 762 +/- 25 m, 21/12/ 2002, D.C. Lees: DL-SG77; ♂, NE, Vohitaly-Anjiahely, 15.3989o S, 49.5211o E +/- 1.82 km, 435 +/- 50 m, 31/12/ 2002, BMNH (E) #672331 [DNA voucher]; ♀, NE, Anjiahely, 0.37 km E., 15.4120o S, 49.5022o E +/- 0.25 km, 380 +/- 5 m 12/12/2002, D.C. Lees: DL-SG1, BMNH (E) 2008-69, BMNH (E) 1717101; ♂, NE, Sahantaha, near ‘fanihy’ roost area, 15.1963o S, 49.5564o E +/- 0.98 km, 517 +/-54 m, 13/12/2001: 10:46, D.C. Lees: DL 01-257; ♂, NE, Anjanaharibe Mt., 15.188o S, 49.6139o E +/- 0.1 km, 475 +/- 5 m, 6/2/2003, D.C. Lees; ♀, NE, Tampolo, W. Masoala, ca. 150 m, primary rainforest, 15.7301o S, 49.9604o 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, 15.7279o S, 49.9751o E +/- 1.8 km, 32 +/- 25 m, 9/ 11/2001, D.C. Lees, DL 01-27, BMNH (E) #697188 [DNA voucher], KA556 [=KA-P556; DNA extract voucher]; ♀, NE, Andranobe, W. Masoala, 240 m, 15.4o S, 49.58o E +/- 1.78 km, 240 m. 3/2/1991, C. Kremen et al., Genitalia157 [genitalia voucher]; ♀, NE, Andranobe, W. Masoala, 90 m, 15.6816o S, 49.9573o E +/- 1.79 km, 90 m, 14/11/1991, C. Kremen et al.: Biodiversity voucher; ♀, NE, W. Masoala, Andranobe field station, T2S2 [site], 90 m, 15.6816o S, 49.9573o E +/- 1.79 km, 90 m, 14/11/1991, C. Kremen et al., Biodiversity survey voucher|Gen158 [genitalia], IA594 [isotope voucher]; ♂, data as above but: 2/11/1991, C. Kremen et al., Biodiversity voucher; ♂, NE, Andranobe, W. Masoala, 520 m, 15.65o S, 49.983o 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|Gen156 [genitalia]; ♀, data as above but: 29/10/1991, C. Kremen et al.: Bio|Gen159; ♀, NE, Andranobe, W. Masoala, 520 m, 15.65o S, 49.983o 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, 15.6733o S, 49.9885o E +/- 0.15 km, 500 +/- 50 m, 18/2/ 1993, D.C. Lees: DL 93-0012, IA593 [isotope voucher]; ♂, NE, Masoala E, Be Dinta, 550 m, 17/02/1993, D.C. Lees: DL 93-0029, IA592 [isotope voucher]; ♂, NE, Be Dinta, upper Anaovandrano R., W. Masoala, 500 m, 15.6733o S, 49.9885o 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, 15.6733o S, 49.9885o 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., W7-L2, Masoala E., 15.7216o S, 50.1753o E +/- 0.15 km, 270 m, 18/1/1994, D.C. Lees; ♀, NE, Manosona, Anaovandrano R., E Masoala, 15.784o S, 50.2164o E +/- 0.15 km, 45 m, 1/2/1994, D.C. Lees; ♂, NE, Masoala E., Antsamanarana R., 275 m, 15.295o S, 50.227o E +/- 0.15 km, 275 +/- 50 m, site W2L2-275M 5 LO, 5/12/1993, H. Raharitsimba: DL 93-0028; ♂ [developmental abnormality LHS CuA1 HWV], NE, E Masoala, Ambavony R. [Ambanivony], W2-L1-125M [site], disturbed forest, 15.2813o S, 50.2877o E +/- 0.72 km, 223 +/- 50 m, 29/11/1993, C. Kremen: CK93-0009; ♀, NE, E Masoala, [Ambavaony R.], bifurcation to 370 m, riparian, W2L1 Saharand. [Sarahandrano] 15.28o S, 50.2860o E +/- 1.4 km, 240 +/- 190 m, 25/11/1993, H. Raharitsimba: TR 46A; ♂, NE, Masoala E., Antsamanarana R., W2L2 [site], 275 m, 15.2813o S, 50.2877o E +/- 0.72 km, 5LO [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], 15.307o S, 50.233o E +/- 0.1 km, 100 m, 29/11/1993, C. Kremen: CK30, GEN 141 DL [genitalia]; ♀, data as above but: 15.2968o S, 50.2271o E +/- 1.31 km, 294 +/- 225 m, 10/12/1993, D.C. Lees: E64 (egg voucher); ♂, NE, Antsamanarana R., Masoala E., 250 m, 15.3o S, 50.23o E +/- 0.15 km, 269 m, 12/12/1993, H. Raharitsimba, GEN 142 DL [genitalia]; ♀, NE, Ambavaony R., E Masoala, [50–380 m], 15.279o S, 50.288o E +/- 0.12 km, 150 m, 1/12/1993, C. Kremen: CK37, EGG [egg voucher]; ♀, data as above but: 223 m, 29/11/1993, C. Kremen; ♀, NE, E Masoala, Ambavaony R., riparian, W2-L1-50M [SITE], 15.2813o S, 50.2877o E +/- 0.72 km, 50 m, 2LO [fruit trap] 28/11/1993, D.C. Lees: E3 [egg voucher; “pearly white, smallish, like Ht. pauper ”]; ♂, NE, E Masoala, Ambavaony [Ambanivony], 380 m, W2L1 [SITE] 15.2813o S, 50.2877o E +/- 5 km, 30/11/1993, H. Raharitsimba: TR 61; ♂, NE, Ambavaony R., E Masoala, 125 m, 15.279o S, 50.288o E +/- 0.12 km, 125 m, 27/11/ 1993, C. Kremen, GEN 124 DL [genitalia]; ♂, NE, Masoala, 1993, C. Kremen: CK93-0005, IA595 [isotope voucher]; ♂ [BMNH], NE, Reserve Speciale d'Ambatovaky, 700 m [16.8o S, 49.1222o E, 580 m], 11/2/1990, M.I. Evans [#1.16], 254 DL [genitalia].</p></div>	https://treatment.plazi.org/id/038747324C34C6111EB72C8FFDC9260E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C31C61E1EB72DBAFDD727ED.text	038747324C31C61E1EB72DBAFDD727ED.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis menamenoides Lees	<div><p>Heteropsis menamenoides Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:D2FCD726-B198-484F-846C-B5229FE174E0</p><p>Prior references: sp. 62 (Lees 1997; Torres et al., 2001: 462).</p><p>Type material., Holotype, deposition BMNH: ♂ (Fig. 21 A), Madagascar NW, Bekolosy, RS Manongarivo, 950 [880] m, [14.0487o S, 48.29495o E +/- 0.15 km, 95 m +/- 50 m], 15/12/1994: 09:50, D.C. Lees: DLBEK 94_160; KAP16 [=KA-P16 DNA extract voucher], IA312 [isotope voucher], NHMUK 010289153 [QTR barcode].</p><p>Paratypes: Deposition BMNH: ♂, NW, Bekolosy, RS Manongarivo, ca. 800 m, [14.05135o S, 48.2937o 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; HM40425 [DNA barcode voucher], KA571, KA4084 [=KA-P571, KA-P4084; DNA extract numbers]; ♂ (Fig. 18 D), Madagascar NW, Bekolosy, RS Manongarivo, 890 m, 14.04936o S, 48.294o 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, 14.04894o S, 48.2945o E +/- 0.15 km, 925 +/- 50 m, 13/12/1994, 09:56–11:15, D.C. Lees: DLBEK 94_181, NHMUK 010289187 [QTR barcode].</p><p>Deposition summary: BMNH (HT ♂, 3 PT ♂♂).</p><p>Type locality. Madagascar NW, Bekolosy, RS Manongarivo, 950 m, [14.0465o S, 48.3o E +/- 0.2 km, 950 +/-70 m].</p><p>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-CuA1 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-CuA1 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 .</p><p>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-M3 of FWD and HWD. Space-CuA1 ocellus expressed more strongly in FWD than in HWD; Orng not evident and black iris considerably smaller diameter than spacing of veins CuA1 and CuA2. 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-CuA1 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-M2 ocellus as small white point surrounded by narrow black ring in FWV while space-CuA1 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.</p><p>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.</p><p>Androconia: black ventral abdominal patch between A3–A6 divided centrally by narrow yellow overlying scales, in potential contact with diffusely separated and slightly wavy androconial brush largely emanating from base of space-CuA2 before its fork with vein CuA1 and overlying veins 1A+2A and 3A; the latter symmetrically inflated before midlength in compact ‘tear’-shape (Lees, 1997: 97, Fig. 3 b), whereas 1A+2A shows very restricted swelling towards the base of 1A. 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.</p><p>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.</p><p>♂ genitalia: 231DL (DLBEK94_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.</p><p>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).</p><p>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. &amp; B. Perrot| Culapa grandis Obthr. ♀ type |[Copy of f. 3074]”).</p><p>Additional information. DNA divergences: COI-5P cluster number BOLD:AAE4113 (exemplar DLBEK94_179, BMAD075-09, HM404251). The DNA barcode is about 5.5% divergent to an undescribed species, sp. 28 (cluster number BOLD:AAE5458), and about 5.8% pairwise divergent to that of Ht. tianae (BOLD:AAE4112).</p><p>Phylogeny/sister species: unknown, new molecular study awaited.</p><p>Ecology and distribution.</p><p>Habitat: on a very steep slope with a fine leaved bamboo that was considered to be ‘ Nastus ’ manongarivensis A. Camus, not necessarily the hostplant.</p><p>Behaviour: flies in the forest substory.</p><p>Hostplant: unknown, but possibly grasses.</p><p>Early stages: unknown.</p><p>Distribution: endemic as far as is known to Mt. Bekolosy, Reserve Speciale de Manongarivo (Fig. 30 C, slateblue dots).</p><p>Elevational range: 800–1035 m. (n=8 incl. observations).</p><p>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.</p><p>Referred specimens. None.</p></div>	https://treatment.plazi.org/id/038747324C31C61E1EB72DBAFDD727ED	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C3FC61A1EB72BCFFAD227D5.text	038747324C3FC61A1EB72BCFFAD227D5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis roussettae Lees & Kremen	<div><p>Heteropsis roussettae Lees &amp; Kremen, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:7F5B27F5-6625-4B55-83D8-B92869CF1545</p><p>Prior references: sp. 26 (Lees, 1997).</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 22 A): ♂, Madagascar, Montagne d'Ambre, 12.5262o S, 49.1710o E, 1060 m, forestry station, GPS point AMBRE-2 [used max] 20/2/1992, C.Kremen: CK840, IA314 [isotope voucher], NHMUK 010289154 [QTR barcode].</p><p>Paratypes: Deposition BMNH: ♂, Madagascar, Montagne d'Ambre, 12.525o S, 49.1722o E +/- 5 km, 900 +/- 50 m, 16/1/1995, D.C. Lees: DLMDA 95-0002, IA362 [isotope voucher], DCLW _0085 [wing prep.], KA2068, KA4085 [=KA-P2068, KA-P4085 [extract numbers], NHMUK 010289155 [QTR barcode]; ♂, N, Montagne d'Ambre, 900 m, 12.525o S, 49.1722o E +/- 5 km, 900 +/- 50 m, 16/1/1995, D.C. Lees: DLMDA 95-0003, KAP18 [=KA-P18; DNA extract number], IA316 [isotope voucher], NHMUK 010289156 [QTR barcode]; ♀ (Fig. 22 B), Madagascar N, Montagne d'Ambre, tourist campsite, c. 1000 m, 12.5268o S, 49.1721o E +/- 0.15 km, 1055 +/- 50 m, 14/1/1995, D.C. Lees: KAP17 [=KA-P17; DNA extract number], DLMDA 95-0004; IA315 [isotope sample], NHMUK 010289157 [QTR barcode]; ♂, N, Montagne d'Ambre [12.5253o S, 49.1483o E +/- 2.47 km, 1185 +/- 115 m], 20/2/1992, C. Kremen: CK92-0002, NHMUK 010289188 [QTR barcode]; ♂ (Fig. 23 A), Madagascar N, Montagne d'Ambre, W of divide, 1080 m, [12.5253o S, 49.1483o E +/- 2.47 km, 1185 +/- 115 m], 20/2/1992, C. Kremen: CK92-0003, 33 DL [genitalia], NHMUK 010289189 [QTR barcode]; ♂, N, Montagne d’Ambre, vers premier petit sommet, route GCASC-&gt;Gite, 12.59435o S, 49.1583o E +/- 0.26 km, 872 +/- 25 m, 17/11/2004, R. Ranaivosolo: DL-05-122, BMNH (E) #671644 [DNA voucher; FJ 819174, 357 bp cytochrome b];</p><p>Deposition PBZT: ♂ [PBZT], 9/12/1958, Madagascar Nord, Montagne d'Ambre, Les Roussettes, 1100 m, IX &amp; XII-58 A. Robinson.</p><p>Deposition summary: BMNH (HT ♂, 5 PT ♂♂, 1 PT ♀); PBZT (PT ♂).</p><p>Type locality. Montagne d'Ambre, ‘Les Roussettes’, 12.5262o S, 49.1710 o E, 1060 m.</p><p>Diagnosis. With a white point emphasised in space-M2 and often M3 of the FW and in M1-M 3 in the HWV, Ht. roussettae looks very like Ht. andasibe . It is however allopatric to Ht. andasibe, as far as known confined to the northern rainforest of Montagne d’Ambre, and likewise it is allopatric to Ht. anceps (Oberthür, 1916), Ht. tianae and Ht. oberthueri sp. nov., which like Ht. andasibe are distributed at more central latitudes of the eastern rainforests. Ht. roussettae has a smoother outcurve to the HWV Mb than those species at M3, and in known material there is a strong tendency to expression of an ocellus, albeit small, in space-M3 on the HWD (Fig. 22 A- B); these features distinguish it from the other four generally similar species. In the ♂ genitalia from LV, compared to Ht. andasibe, the tegumen has a relatively straight dorsal edge (and wide and shallow proximad notch from DV), appearing quite extended proximad and the uncus tip extends towards or slightly beyond maximum extension (distad lobe) of valve tips; the reliability of these distinguishing features is unknown. The HWV has a relatively straight Mb (compared to Ht. strigula (Mabille, 1877) and to Ht. ankova, where it is outangled at M2), and this characteristic is similar to Ht. anceps and the light orange Ht. menamenoides described above, distinguishing it strongly from members of the Ht. angulifascia clade ( Ht. strigula group).</p><p>Description. Wings: Upperside uniform dark brown. FW space-CuA1 ocellus with black iris spanning veins CuA1 and CuA2; Orng not perfectly circular, tending to be angled towards base of vein CuA1. FW space-M1 ocellus small, with faint orange ring; M2 ocellus white pupil only evident. In HW, space-CuA1 ocellus elliptic, with narrow dark orange ring, spanning about 1/3 of vein CuA1-CuA2 distance; space-M3 ocellus expressed (unusually for a member of the Ht. strigula group), but smaller, less elliptic. White pupils as points only, in HW spaces-M1-M3 particularly. Underside has ochreous-olive brown cast. FW space-CuA1 ocellus with orange ring that is yellower towards costa tending to be blocked by vein M3, and forming Ore that follows concave curve of dark brown FWV Mb, which as usual curves back towards mid costa. Background colour is slightly yellower olivebrown distad of Mb. Mb in HW relatively straight, as in Ht. menamenoides, and there is a distinct dark brownish convex PMb. Fine irroration of small darker brown line fragments throughout wing and concentrated in basal area, with Cbs on FW comprising two sets of approximately parallel transverse lines in the FW cell area. Very wavy darker diffuse brown line runs submarginally in both wings especially FW (somewhat toothed outward at each vein) and narrow darker brown line tracks wing margin, which is fairly smooth, only very slightly crenate. Variation. ♂♂ similar to each other, but seasonal variation has not been quantified. ♀♀ similar in pattern, but larger and somewhat lighter dorsally with a more rounded HW shape.</p><p>Wingspan/fwl: range 39.4–40 mm (n=2 ♂♂)/ 19.7–21.7 mm (n=3 ♂♂), including HT ♂ 40/ 22 mm; 40/ 20.89 mm (n=1 ♀).</p><p>Androconia. Aside from Sdb, dark dispersed Cub consists of brownish hairs emanating from above 1A+2A and 3A that may contact a blackish patch of specialized scales located on abdominal sternite A3–A5 (Lees, 1997: 99, Fig. 5). Note that an androconial abdominal patch in this position is not very evident, and does not appear distinctly from superficial examination. The cubital/anal brush(es) partially overlie two ‘balloon’ inflations (swellings) terminating around mid-vein in 1A+2A and 3A, tapering towards base, covered in very small thin grey scales, that in 3A are rather larger than that in 1A+2A (Lees 1997: 96, Fig. 3 A).</p><p>Palps: penultimate segment with medial narrow brown strip, flanked by yellowish scales, and fringed with browner scales; yellowish scales prominent on mesad face of labial palp.</p><p>♂ genitalia: 33DL (Fig. 23 A, PT): from LV, very similar to Ht. andasibe (as “ antsianakana ” in Lees 1997: 107, Figure 7 g), but tegumen, which has fairly straight dorsal edge (and wide and shallow proximad notch from DV), appears quite extended proximad from LV and uncus tip extends towards or slightly beyond maximum extension (the distad lobe) of valve tip. Uncus slightly longer than tegumen and parallel sided from LV with distinct dorsal ‘head’ at tip; slightly inflated before tip from DV. Valves stout and asymmetric at base which also has a small and gently curved ‘shoulder’; dorsal valve margin is the much more recurved one, with a recurved spine facing proximad/mesad. As in Ht. andasibe, the shallowly bifid ends of valves impart a somewhat ‘crab pincer’-like appearance, heavily armed with spinoid setae along wide, uncus-facing margin. Saccus not long and aedeagus similar length to valve, quite deep and strongly uprecurved distad. The gnathos emanates from a distinctly rectangular base, recurved downward as crossing the fairly narrow and parallel-necked uncus, at an angle of around 40 degrees in the specimen examined. The juxta is narrow proximad and slightly down-lipped.</p><p>Etymology. Named after Les Roussettes (a name likely derived from Rousettus, a genus of fruit bat; note that the derivation used here though is feminine singular from the French word ‘roussette’, which might approximate the root of both names). Les Roussettes is a rather famous touristic site at PN Montagne d’Ambre and when visited this former forestry station was provided with a ‘ gite ’ around which this low-flying species could be found. The name also reflects that of the closely related species Ht. andasibe, also named after a familiar tourist destination in Madagascar, and again also emphasing the high local endemicity of much of the fauna.</p><p>Discussion. This species was first recognized in the field as “sp. 26” by Claire Kremen on 20/02/1992 (two ♂♂). There is a single historical specimen of this species in PBZT included in the type series. All relevant Heteropsis types in the Ht. strigula group were examined to ensure recognition of this species as new. The former syntypes of Culapa antsianakana Oberthür 1916 were examined: LT ♂, according to the Code, was first designated by d’Abrera (1980: 184) who illustrated a specimen of the first species with a red label next to it and a text which stated “♂ (holotype) as illustrated” (Fig. 22 C); bearing labels “BMNH(E) #674757; Madagascar Antsianaka Perrot Freres 2e Semestre 1890”. Also examined were the STs of Culapa anceps Oberthür 1916; LT ♂, here designated (Fig. 22 D), specimen bearing labels “BMNH(E) #674762; Madagascar Antsianaka Perrot Freres 2e Semestre 1893”. Regarding Mycalesis strigula Mabille, 1877, not clearly a HT by monotypy, a LT ♂ is here also designated (not illustrated here): BMNH(E) 673784. Among the two synonyms of the last species (Lees 1997; Lees et al., 2003; illustrated in d’Abrera, 1997), also examined were Culapa ankovana Oberthür, 1916 (HT ♂, by monotypy: “BMNH(E) 673783; Madagascar Fito Mai à Aout 1897 Perrot Freres”; specimen illustrated by Oberthür, 1916, Pl. 367, f. 3060 and by d’Abrera 1980: 184 (“♂ (? holotype) as illustrated”)) and Culapa wardiana Oberthür, 1916 (HT ♂: BMNH(E) 673785); illustrated by Oberthür 1916, Pl. 356, f. 3056. Types were also examined of Mycalesis ankova Ward, 1870 (HT ♂: “BMNH(E) 673767; Madagascar Ex COLL. CHRIS WARD”). See also under Ht. tornado regarding types of Pseudonympha turbata Butler, 1880, Culapa ornata Oberthür, 1916 and Culapa pallida Oberthür, 1916 .</p><p>Additional information. DNA divergences: no COI-5P barcode yet available. The cytochrome b sequence, 357 bp (BMNH(E) #671644, FJ819174) is about 4.75% divergent from Ht. andasibe (comparison sequences BMNH(E) #676684 FJ819448 from Ambatovy; BMNH(E) #676666 FJ819446, BMNH(E) #676679 FJ819447 and BMNH(E) #697771 FJ819449; dataset of Monaghan et al., 2009).</p><p>Phylogeny/sister species: Lees (1997: 156) found morphological support for a clade consisting of this species (“TWNSX”), Ht. anceps (“ ANCEPS ”) and Ht. andasibe (“ANTSI”), with a topology retained in analysis of ♂ genitalic characters. Although it resembles more closely Ht. andasibe, either species might potentially be its allospecific replacement in Northern Madagascar, but new collections of the narrowly endemic Ht. anceps are also needed to provide DNA evidence (see also combined tree in Aduse-Poku et al., 2016).</p><p>Ecology and distribution.</p><p>Habitat: fairly open areas in rainforest.</p><p>Behaviour: flies low over the ground.</p><p>Hostplant: unknown, but assumed to be low growing grasses.</p><p>Early stages: unknown.</p><p>Distribution: only known from Montagne d’Ambre, in the vicinity of Les Roussettes (Fig. 30 C. ochre dots).</p><p>Elevational range: 872–1185 m. (n=16 including referred specimens and observations).</p><p>Referred specimens. ♂, N, Parc National Montagne d'Ambre [Petit Lac road], [12.5203o S, 49.1792o E +/- 0.15 km, 1125 +/- 50 m], 4/3/2001, R. Harin'Hala; MSL 024: BO8 [genomic DNA]; BMNH (E) #672502; 1137 [= DL 1137; DNA extract number, not amplified]; ♂, data as above but: 1125 +/- 50 m], 4/3/2001, R. Harin'Hala; MSL 024: CO8 [genomic DNA]; BMNH (E) #672503; 1138 [= DL 1138; DNA extract, not amplified].</p></div>	https://treatment.plazi.org/id/038747324C3FC61A1EB72BCFFAD227D5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C3BC61A1EB72BC9FD9D2680.text	038747324C3BC61A1EB72BC9FD9D2680.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis angulifascia	<div><p>Heteropsis angulifascia clade</p><p>In the sampling of Aduse-Poku et al., (2015, Fig. 1), Ht. angulifascia (Butler, 1879), Ht. ‘ anceps ’ (= Ht. oberthueri), Ht. turbans and Ht. sabas formed a well-supported molecular clade. This clade, nested within the Ht. strigula sub-group, and named after Ht. angulifascia, also includes the described species Ht. anceps, as well as at least five undescribed species, four of which are introduced below. Morphologically, the group features some degree of dorso-ventral flattening of the gnathos, in its extreme forming bizarre, cupped, ear-like structures unique in the Mycalesina, either side of uncus.</p></div>	https://treatment.plazi.org/id/038747324C3BC61A1EB72BC9FD9D2680	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C38C6061EB72DBAFE8226A3.text	038747324C38C6061EB72DBAFE8226A3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis tianae Lees & Kremen	<div><p>Heteropsis tianae Lees &amp; Kremen, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:E6864E90-B8CF-4E5F-99C4-AC879E6F70A</p><p>Prior references: sp. 49, 69 (Lees, 1997: Torres et al., 2001: 462).</p><p>Type material., Deposition BMNH: Holotype: ♂ (Fig. 24 A), Madagascar C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 18.45o S, 47.95o E +/- 2.45 km, 1375 +/- 75 m, 2/3/2004, D.C. Lees: DL-4-182, NHMUK 010289158 [QTR barcode].</p><p>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, 18.4368o S, 47.9469o 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, HM404245 [DNA barcode voucher], BMNH (E) #676763 [DNA voucher; cytochrome b], IA85 [isotope voucher]; ♂, C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 12–14 m canopy forest, flat summit, 18.4497o S, 47.9404o 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, IA122 [isotope voucher]; 2 ♀♀ [BMNH], “Andrangalooka Forest, Madagascar ” [=Andrangoloaka, E. Lac Mantasoa], BMNH (E) #674766 (Godman-Salvin Coll.; Fig. 24 C) and BMNH (E) #674767;</p><p>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;</p><p>Deposition ABRI: ♂, Madagascar C, Anjozorobe, 18.413o S, 47.946o E +/- 0.02 km, 1310 +/- 50 m, 22/10/ 2014, D.C. Lees: DL14Z-066.</p><p>Deposition summary: BMNH (HT ♂, 2PT ♂♂, 3 PT ♀♀), MNHN (PT ♂), ABRI (PT ♂).</p><p>Type locality. Madagascar C, Amboasary-an-ala, vic. Anjozorobe, 18.45o S, 47.95o E +/- 2.45 km, 1375 +/- 75 m.</p><p>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-M2 (Fig. 24 A– C), and the HWV tends to have the space-CuA2 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-M2 and strong tendency to expression of a small ocellus in space-M3 on the HWD (Fig. 22 AB).</p><p>Description. Wings: upperside uniform mid brown, FW space-CuA1 ocellus with an orange ocellus ‘ring’ which is sometimes rather hexagonal in shape and eccentric, being wider at proximad edge. FW space-M1 ocellus small with narrow orange ring. HW space-CuA1 ocellus the only one expressed there and somewhat elliptic. Darkish brown diffuse, slightly wavy hair-brush emanating mainly from below vein 1A+2A 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-CuA1 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-M1 ocellus FWV reduced to white pupil with narrow black iris without trace of orange ring. On HWV, space-CuA1 ocellus slightly elliptic with narrow black iris and small also, without orange ring trace. HWV space-M1-M2 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-M2. HWV space-CuA2 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.</p><p>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 ♀♀).</p><p>Androconia: both 1A+2A and 3A 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 M2, M3, CuA1 and CuA2 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 A2. 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, MAD244; Anjozorobe, n=5).</p><p>Palps: penultimate segment with narrow brown medial strip, flanked by yellow and fringed by dark brown scales, with yellow scales mainly on mesad face.</p><p>♂ genitalia: 228DL, 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.</p><p>Etymology. Etymology. After ‘Tiana’ Raharitsimba (= Heritiana Rahagalala), who found some of the first modern exemplars of this species at Anjozorobe.</p><p>Discussion. Historical specimens, one from Andrangoloaka forest in the Angavo massif, probably late 19th 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.</p><p>Additional information. DNA divergences: COI-5P cluster number BOLD:AAE4112 (exemplar BMAD200- 15, DL14Z-051) is about 2.58% divergent to Ht. oberthueri (cluster number BOLD:ACW4996, exemplar BMAD242-15, DL06-11, RNI Zahamena) and about 4.17% divergent from ‘sp. 28’ (BOLD:AAE5458), 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-5P cluster number BOLD:ACD8579).</p><p>Phylogeny/sister species: probably Ht. oberthueri based on the close relationship of the COI-5P 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).</p><p>Ecology and distribution.</p><p>Habitat: montane rainforest.</p><p>Behaviour: both sexes come readily to fruit bait. Flies in substratum of forest.</p><p>Hostplant: unknown, presumed to be grasses.</p><p>Early stages: unknown.</p><p>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.”</p><p>Elevational range: 1320–1375 m. (n=27 including referred specimens and observations).</p><p>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. [18.4084o S, 47.9377o 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 14Z-051, BMAD 200-15 [DNA barcode voucher]; ♀, C, Anjozorobe, 1400 m, 18.4084o S, 47.9377o 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: CK753, IA211 [isotope voucher]; ♀, same data but 138 m 14/12/1994, C. Kremen: CK754; ♂, C, Anjozorobe, 18.4498o S, 47.939o 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, IA9 [isotope voucher]; ♂, C, Amboasaryan-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 18.45o S, 47.95o 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, 18.45o S, 47.95o 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], IA123 [isotope voucher]; ♂, C, Amboasary-an-ala, Vanjamanitra-Vazimba forest, Anjozorobe, 18.45o S, 47.95o 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, 18.45o S, 47.95o 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 S3’, 18.4505o S, 47.9399o E +/- 0.15 km, 1323 m, 2/3/2004: 08:56, D.C. Lees: DL-4-167, IA19 [isotope voucher]; ♂, C, Anjozorobe, 18.45o S, 47.95o E +/- 2.45 km, 1330 m, 4/2/2014: 15:20, D.C. Lees: MAD 240 [pheromone voucher], IA403 [isotope voucher]; ♂, Anjozorobe, Babakoto trail, c. 1350 m, K. Aduse-Poku, 04/02/2014, MAD 243 [pheromone voucher], KA2059 [=KA-P2059; DNA extract voucher]; specimen, Anjozorobe, KAP-MAD1408, KA1018 [=KA-P1018; DNA extract number].</p></div>	https://treatment.plazi.org/id/038747324C38C6061EB72DBAFE8226A3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C24C6021EB72BF9FBA92005.text	038747324C24C6021EB72BF9FBA92005.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis oberthueri Lees	<div><p>Heteropsis oberthueri Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:FEE8CA6F-A53B-43A5-8C9F-E5D69AD0E468</p><p>Prior references: sp. 89, sp. 87 (on some envelopes). “KA518_Heteopsis_ anceps ” [sic] in Aduse-Poku et al., (2015, Fig. 1).</p><p>Type material. Deposition BMNH: Holotype: ♂ (Fig. 24 D), Madagascar E, Zahamena NW, 17.5131o S, 48.7269o E +/- 1.5 km, 1064 +/- 25 m, 5/11/2006, D.C. Lees: DL 06-064, G04 [leg for DNA], NHMUK 010289159 [QTR barcode].</p><p>Paratypes: Deposition BMHN: ♂, Madagascar E, Ambavala camp, Zahamena, 17.5451o S, 48.7237o E +/- 0.5 km, 1300 +/- 50 m, 6/11/2006, 13:10, D.C. Lees: DL 06-130A, NHMUK 010289160 [QTR barcode]; ♀ (Fig. 24 E), E, Zahamena NW, 17.5376o S, 48.71965o E +/- 0.94 km, 1180 +/- 125 m, 6/11/2006, D.C. Lees: DL 06-130, IA585 [isotope voucher], NHMUK 010289161 [QTR barcode]; ♂, E, Zahamena NW, 17.5376o S, 48.71965o E +/- 0.94 km, 1180 +/- 125 m, 6/11/2006, D.C. Lees: 371 DL, NHMUK 010289192 [QTR barcode]; ♂, E, Zahamena NW, 17.53o S, 48.7269o E +/- 1.5 km, 1064 +/- 25 m, 6/11/2006, D.C. Lees: DL 06-086, IA586 [isotope voucher], NHMUK 010289162 [QTR barcode]; ♂, E, Zahamena NW, 17.5376o S, 48.71965o E +/- 0.94 km, 1180 +/- 125 m, 6- 7/11/2006, D.C. Lees: DL 06-1001, IA324 [isotope voucher], NHMUK 010289163 [QTR barcode]; ♀, E, Zahamena NW, 17.5376o S, 48.71965o E +/- 0.94 km, 1180 +/- 125 m, 6/11/2006, 11:30, D.C. Lees: DL 06-112, IA587 [isotope voucher], NHMUK 010289164 [QTR barcode]; ♀, E, Towards Ambavala camp, Zahamena, 17.5376o S, 48.71965o E +/- 0.94 km, 1180 +/- 125 m, 6/11/2006, D.C. Lees: DL 06-141, IA588 [isotope voucher], NHMUK 010289165 [QTR barcode]; ♀, E, returning to Bemoara camp, Zahamena, 17.5356o S, 48.71875o E +/- 0.70 km, 1180 +/- 100 m, 5/11/2006, 14:59, D.C. Lees: DL 06-059, IA589 [isotope voucher], NHMUK 010289166 [QTR barcode];</p><p>Deposition MNHN: ♂, Madagascar E, Ambavala, Zahamena NW, 17.5451o S, 48.7237o E +/- 0.25 km, 1325 +/ - 50 m, 6/11/2006, 14:11, D.C. Lees: DL06-117, IA584 [isotope voucher], DSC03096.jpg [image];</p><p>Deposition ABRI: ♂, Madagascar E, Zahamena NW, 17.5376o S, 48.71965o E +/- 0.94 km, 1180 +/- 125 m, 6/ 11/2006, D.C. Lees: DL06-121.</p><p>Deposition summary: BMNH (HT ♂, 4 PT ♂♂, 4 PT ♀♀), MNHN (PT ♂), ABRI (PT ♂).</p><p>Type locality. NW Zahamena, 17.5131o S, 48.7269o E +/- 1.5 km, 1064 +/- 25 m.</p><p>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-M3 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-M3 as well as in space-CuA1 expressed on the HWD (Fig. 19 C). The ventrally very similar Ht. roussettae has a strong tendency to expression of a small ocellus M3 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.</p><p>Description. Wings: description is similar to that of Ht. tianae . Upperside uniform mid brown, with FW space-CuA1 ocellus featuring an orange ocellus ‘ring’ which is sometimes rather ‘squashed’ along the diagonal from mid-costa to tergal angle. FWD space-M1 ocellus small with narrow orange ring. HWD space-CuA1 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 1A+2A, 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-CuA1 ocellus with orange ring yellowing proximad, following darkish brown concave curve of Mb before it bends back to mid-costa. Space-M1 ocellus FWV a white pupil in HT. On HWV, space-CuA1 ocellus small and slightly elliptic with narrow black iris, also without orange ring trace. HWV space-Rs-M2 ocelli as white pupil-points and space-CuA2 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-M2. 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- M1 ocellus FWV sometimes features a narrow black iris and space-M2 sometimes just as a white pupil without narrow black iris nor trace of orange ring. Space-CuA2 ocellus HWV slightly expressed in some specimens. ♀♀ similar in pattern, but larger and somewhat lighter dorsally with a more rounded HW shape.</p><p>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 ♀♀).</p><p>Androconia: HWD vein 1A+2A has a narrowly tapered inflation and 3A is much more swollen distad, from base to mid-vein, covered in thin grey-brown scales. The length of HW veins M2, M3, CuA1 and CuA2 are also narrowly inflated, and have specialized scales on the dorsal surface. Abdominal black androconia are just visible ventro-laterally around A4–A5. HWD discocellular brush dark brown. Sdp HWD grey, small, lenticular, composed of narrow grey scales).</p><p>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.</p><p>♂ 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).</p><p>Etymology. after Charles Oberthür, who in 1916 produced one of the first treatments, a seminal work on the Malagasy Mycalesina .</p><p>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).</p><p>Additional information. DNA divergences: COI-5P cluster number BOLD:ACW4996 (exemplar BMAD242-15, DL06-11), about 2.58% divergent to Ht. tianae (BOLD:AAE4112, exemplar BMAD200-15, DL14Z-051).</p><p>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-5P barcode.</p><p>Ecology and distribution.</p><p>Habitat: primary rainforest.</p><p>Behaviour: flies low.</p><p>Hostplant: unknown, but likely to be low-growing grasses.</p><p>Early stages: unknown.</p><p>Distribution: as far as is known, endemic to RNI Zahamena (Fig. 30 C, brown dots).</p><p>Elevational range: 1180–1310 m. (n=27, including referred specimens).</p><p>Referred specimens. ♀, Madagascar E, returning to Bemoara camp, Zahamena, 17.5376o S, 48.71965o 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, 17.5131o S, 48.7269o E +/- 1.5 km, 1064 +/- 25 m, 5/11/2006, D.C. Lees: DL 06-061; ♂, E, Zahamena NW, 17.53o S, 48.7269o 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, 17.5376o S, 48.71965o 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, IA432 [isotope voucher], KAP518 [=KA-P518; 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, 17.5451o S, 48.7237o 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, 17.4946o S, 48.733o E +/- 1 km, 1321 m, 7/11/2006, 10:37, D.C. Lees: DL 06-156; ♂, E, near Ankosy summit, Zahamena, 17.49405o S, 48.73325o E +/- 0.067 km, 1300 +/- 25 m, 7/ 11/2006, D.C. Lees: DL 06-188; ♀, E, Zahamena, 17.5131o S, 48.7269o E +/- 1.5 km, 1064 +/- 25 m, 6/11/2006, D.C. Lees: DL 06-67, IA69 [isotope voucher]; ♂, E, Zahamena NW, 17.4946o S, 48.733o E +/- 1 km, 1321 +/- 25 m, 6/11/ 2006, D.C. Lees: 364 DL [genitalia voucher], IA29 [isotope voucher], leg sample [KAP].</p></div>	https://treatment.plazi.org/id/038747324C24C6021EB72BF9FBA92005	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C23C6001EB72C94FE4726AB.text	038747324C23C6001EB72C94FE4726AB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis borgo Lees	<div><p>Heteropsis borgo Lees, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:45DC90D0-4167-4047-89DC-A9C45D523E49</p><p>Prior references: sp. 73 (Lees 1997; Torres et al., 2001: 462).</p><p>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 14A-0350, IA522 [isotope voucher], NHMUK 010289167 [QTR barcode].</p><p>Paratypes: Deposition BMNH: ♂, data as HT but 23/11/2014: 12:21. D.C. Lees. DL 14A-0361, BMAD 221- 15 [DNA barcode], IA519 [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 14A-0363, IA521 [isotope voucher], BMAD 235-15 [DNA barcode], NHMUK 010289169 [QTR barcode]; ♂, NE, Anjanaharibe Sud, ANJAHA-6 [GPS], 14.7482o S, 49.4659o E +/- 0.48 km, 1320 +/- 60 m, 6/2/1996, C. Kremen: CK724, rainforest on slope, trap 14, IA358 [isotope voucher], NHMUK 010289170 [QTR barcode]; ♂, NE, Anjanaharibe Sud, 1530 m, elfin forest just before summit, ANJAHA-7 [GPS], 14.7426o S, 49.4608o E +/- 0.37 km, 1450 m +/- 7 m, 4/2/1996: 11:49, H. Raharitsimba: TR 553A, IA359 [isotope voucher], NHMUK 010289171 [QTR barcode]; ♂ [left wings and head], Madagascar NE, Anjanaharibe Sud, ANJAHA-7 [GPS], 14.7406o S, 49.4505o E +/- 0.96 km, 1455 +/- 145 m, 12/2/1996: 08:59, C. Kremen: CK729; DL 9659 [DNA voucher], IA360 [isotope voucher], NHMUK 010289172 [QTR barcode];</p><p>Deposition MNHN: ♂, data as HT but 21/11/2014: 12:02, Ahamadi Allaoui: DL14A-0308, IA520 [isotope voucher]; ♂, data as HT but 23/11/2014. Ahamadi Allaoui. DL14A-0192, 116 [leg in ethanol], IA626 [isotope voucher].</p><p>Deposition summary: BMNH (HT ♂, 5 PT ♂♂); MNHN (2 PT ♂♂).</p><p>Type locality. Anjanaharibe Sud, summit Anjavidibe, 14.7396°S, 49.4605°E +/- 0.25 m, 1543 +/- 15 m.</p><p>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 M2 and M3, running for only a short rather than a long distance just below M2.</p><p>Description. Wings: upperside uniform dark brown. FWD space-CuA1 ocellus fairly large, occupying more than the span of this space taking into account the concentric deep orange ring. FWD space-M1 ocellus small and subelliptic with small white pupil and wider black iris, with only a faint trace of Orng. HWD space-CuA1 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-CuA1 ocellus very slightly larger than on FWD, its black iris spanning the veins CuA1 and CuA2, with fairly wide, more or less concentric, deep orange ring. FWV space-M1 ocellus very small and only slightly elliptic with white pupil and black iris, but very narrow pale orange ring. Small white point in M2 just under this ocellus. HWV space-Rs ocellus as distinct white spot without obvious black ring. FWV space-M1-M2 ocelli as minute white spots and ocellus in space-CuA2 not expressed. HWV space-CuA1 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-M2-M3, bent distad to the base of space-M2, concave distad of space-M1 and space-CuA1, and slightly concave near intersection with vein CuA2. 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- M2–M3, 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 .</p><p>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.</p><p>Androconia: Sdp small, oval and black. Darkish brown hair-brush emanating mainly from above 1A+2A near the base and from space-1A as far as mid-vein. Blackish androconial scales appear to be present ventro-laterally around abdominal A4, but are indistinct in the material available.</p><p>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.</p><p>♂ genitalia: 284DL, 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.</p><p>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.</p><p>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 .</p><p>Additional information. DNA divergences: COI-5P barcode cluster no. BOLD:AAK5842 (exemplars BMAD221-15 DL14A-0361, BMAD235-15 DL14A-0363) is shared with, although 1.67% divergent to, Ht. angulifascia (exemplar BMAD049-09, 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 (8bp different in 415) for a single sample of Ht. borgo (as “ Hen. sp. 73”, 9659S compared with Ht. angulifascia samples ANG8M, ANG7M, ANGCK and ANG11).</p><p>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.</p><p>Ecology and distribution.</p><p>Habitat: riparian primary rainforest, and rainforest on ridges and slopes.</p><p>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.</p><p>Hostplant: unknown, possibly a small leaved grass that was found downslope of the type locality.</p><p>Early stages: unknown.</p><p>Distribution: endemic to rainforest at Anjanaharibe Sud, as far as is known (Fig. 30 C, dark green dot).</p><p>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.</p><p>Elevational range: 1320–1543 m. (n=8).</p><p>Referred specimens. ♂ (Fig. 27 D, male genitalia, rest of voucher lost), Madagascar NE, Anjanaharibe Sud, 1520 m, ANJAHA-8 [GPS point] 14.74265o S, 49.46065o E +/- 0.39 km, 1520 +/- 100 m, 2/2/1996, C. Kremen: CK712; 284 DL [♂ genitalia], 470 [= DL 0470; DNA extract number], BMNH (E) #697884, KA580 [=KA-P580; DNA extract number].</p></div>	https://treatment.plazi.org/id/038747324C23C6001EB72C94FE4726AB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
038747324C2EC60C1EB72DBAFC8B2602.text	038747324C2EC60C1EB72DBAFC8B2602.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Heteropsis vertigo Lees & Raharitsimba	<div><p>Heteropsis vertigo Lees &amp; Raharitsimba, sp. nov.</p><p>LSID: urn:lsid:zoobank.org:act:38AC77EE-A10A-4DCA-9444-F55C845FFA23</p><p>Prior references: sp. 61 (on envelopes and/or datalabels but not mentioned in the list of Lees, 1997: 64–65).</p><p>Type material., Deposition BMNH: Holotype: ♂ (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>Paratypes: Deposition BMNH: ♂, 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,&gt; 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>Deposition summary: BMNH (HT ♂, 2 PT ♂♂, 4 PT ♀♀).</p><p>Type locality. Bekolosy Mt., RS Manongarivo, c. 1163 m, 14.0458o S, 48.2962o E +/- 0.5 km, 1130 +/- 35 m.</p><p>Diagnosis. In its translucence, the extended FWD space-CuA1 Orng of Ht. vertigo is intermediate between Ht. turbans and Ht. sabas . Ht. vertigo in wing pattern resembles Ht. turbans more than it does Ht. sabas, 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 Ht. sabas, or largely orange as in Ht. turbans . Ht. vertigo lacks the extreme ear-like cupping of the gnathos seen in Ht. sabas (Lees, 1997: 107) and Ht. turbans (Fig. 29 B; NW Zahamena) as well as some related species (e.g. Ht. angulifascia, Ht. borgo), and the HWV Mb is much less kinked than in any of these species.</p><p>Description. Wings: Similar to Heteropsis turbans . Upperside uniform mid brown, except with conspicuous orange Mf/crescent (as in a few other of the Ht. strigula -group, notably Ht. strigula) 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 Ht. turbans 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 Ht. sabas) 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. Variation. ♂♂ 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>Wingspan/fwl: 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>Androconia: 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 Ht. turbans and Ht. sabas, 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>Palps: 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>♂ genitalia: 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 Ht. turbans and Ht. angulifascia), 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>Etymology. 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>Discussion. No historical museum collections of Ht. vertigo 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 Ht. turbans (indeed Lees 1997: 225 Fig. 3 K erroneously shows the distribution of that species extending to the northwest of Madagascar). Ht. vertigo 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 Culapa turbans 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 Culapa curvatula 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). Ht. curvatula was considered a synonym of Ht. turbans by Lees et al., 2003 (see also below). Also examined (Fig. 28 E) was the HT ♂ of Culapa sabas Oberthür, 1923 (‘Tamatave’|BMNH(E) #674789), in addition to extensive field material of Ht. turbans and Ht. sabas 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>Additional information. DNA divergences: 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 Ht. turbans and about 1.7% divergent to Ht. sabas with six apparently fixed nucleotide differences to Ht. turbans; all three species share the same cluster number. Sister to H. sabas in Aduse-Poku et al., (2016) .</p><p>Phylogeny/sister species: sister species unclear, but closely related both to Ht. sabas and Ht. turbans, according to the COI-5P barcode. A ♀ of the ‘ curvatula ’ phenotype (DL06-1099, BMAD259-15) from NW Zahamena was one nucleotide different (COI-5P) to other barcoded members of H. turbans (cluster number BOLD:ACD8579), and so the above synonymy appears correct.</p><p>Ecology and distribution.</p><p>Habitat: primary montane rainforest.</p><p>Behaviour: one was caught in a low hung fruit trap at the waterfall camp.</p><p>Hostplant: unknown.</p><p>Early stages: unknown.</p><p>Distribution: as far as is known endemic to Réserve Speciale de Manongarivo, Mts. Bekolosy and Antsatrotro (Fig. 30 B, fawn dots).</p><p>Elevational range: 1130–1410 m. (n=45; including referred specimens and observational data, such as fruit trap data).</p><p>Referred specimens. ♂, 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></div>	https://treatment.plazi.org/id/038747324C2EC60C1EB72DBAFC8B2602	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	C, Lees David	C, Lees David (2016): Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision. Zootaxa 4118 (1): 1-97, DOI: 10.11646/zootaxa.4118.1.1
