Heteropsis imerina Lees

C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, Zootaxa 4118 (1), pp. 1-97 : 41-43

Heteropsis imerina Lees , sp. nov.

LSID: urn:lsid:zoobank.org:act:988AC583-72F8-4CA4-824A-157D9F7C5AEC

Prior references: sp. 22 ( Lees, 1997; Torres et al., 2001: 462);

Sp. 76 ( Torres et al., 2001: 462).

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].

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];

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.

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

Type locality. Madagascar E., Mantadia National Park, Jofa, near bridge, 18.7674°S, 48.4324°E +/- 0.4 km, 945 +/- 15 m.

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.

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.

Androconia: discocellular brush HWD mid brown, to ochreous brown towards tip. Underlying patch lenticular, pale ochreous. No other androconia.

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) (♀).

Palps: mid brown without clear pale striping.

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.

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).

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.

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).

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).

Ecology and distribution.

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.

Behaviour: Low flying, found mostly along ridges but also in riparian areas, perching low off the ground.

Hostplant: occurs among native climbing bamboos, which it might feed on.

Early stages: expressed egg pearly whitish.

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.

Elevational range: 786–1932 m. (n=118; DCL-DB-2901 from Tamatave-Anivorano [estimated at 80 m] is discounted as an unlikely outlier).

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].

Figures

FIGURE 1. Venation and wing patterning diagram of a fairly typical species of Heteropsis from Madagascar (Heteropis antahala group; ♂), DV. Sdbp = Supra-discocellular brush and patch. Cub = Cubital brush. Ventral banding: Cb = Costal bars. Mf = Median fleck (pale highlight). Mb = Median band. PMb = Premedian band. Ore = Ocellus ring extension. Pml = Premarginal line. Sml = Submarginal line.

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

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

FIGURE 9. Adults in DV (left), VV (right) A: HT ♂ of Heteropsis vanewrighti Lees, sp. nov. (Mt. Bekolosy; NHMUK 010289124). B: PT ♀ of Ht. vanewrighti (Mt. Bekolosy; NHMUK 010289125). C: HT ♂ of Ht. imerina Lees, sp. nov. (Mantadia; NHMUK 010289132). D: PT ♀ of Ht. imerina (Ankasoka, route Lakato; DCL-DB- 2906, MNHN). Scale bar: 30 mm.

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