Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision Author C, Lees David text Zootaxa 2016 4118 1 1 97 journal article 38930 10.11646/zootaxa.4118.1.1 a4e408c4-2575-4b5c-878b-e1b9010ec619 1175-5326 264597 CFA586DA-10EE-468B-80EE-35351E3845FD Heteropsis pauliani Lees , sp. nov. LSID: urn:lsid:zoobank.org:act:422E5526-8FBB-4AD6-A47D-EC95FF54E16F Prior references : sp. 37 ( Lees, 1997 : 64; Torres et al., 2001 : 462). 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. 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]; 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; 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]. Deposition summary : MNHN (HT ♂, 9 PT ♂♂, PT ♀); BMNH (PT ♂, PT ♀); PBZT (1 PT ♂). 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 ]. 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 . 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. 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. Androconia : small dark brown brush overlying lenticular dark brown or blackish patch on stem of HWD R vein just above cell. 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. ♂ 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. 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. 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. 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) . 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. Ecology and distribution. Habitat : montane rainforest, sclerophyllous forest including ‘matsabory’ (marshy and lake forest areas). Behaviour : known to fly low across gaps between forest. One was caught in the middle of a ‘matsoborimaiky’ (dry lake) at Tsaratanana. Hostplant : unknown. Early stages : unknown. 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. Elevational range : 1778–2320 m . (n= 13 incl. referred specimens and observations). 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. 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].