Heteropsis tianae Lees & Kremen
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publication ID |
https://doi.org/ 10.11646/zootaxa.4118.1.1 |
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publication LSID |
lsid:zoobank.org:pub:CFA586DA-10EE-468B-80EE-35351E3845FD |
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DOI |
https://doi.org/10.5281/zenodo.6086456 |
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https://treatment.plazi.org/id/03874732-4C38-C606-1EB7-2DBAFE8226A3 |
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Plazi |
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Heteropsis tianae Lees & Kremen |
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sp. nov. |
Heteropsis tianae Lees & Kremen , sp. nov.
LSID: urn:lsid:zoobank.org:act:E6864E90-B8CF-4E5F-99C4-AC879E6F70A
Prior references: sp. 49, 69 ( Lees, 1997: Torres et al., 2001: 462).
Type material., Deposition BMNH: Holotype: ♂ ( Fig. 24 View FIGURE 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].
Paratypes: Deposition BMNH: ♂ ( Fig. 25 View FIGURE 25 A), Madagascar C, Anjozorobe 1400 m, 12/11/1994, C. Kremen, 228 DL [genitalia], NHMUK 010289191 [ QTR barcode]; ♀ ( Fig. 24 View FIGURE 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 View FIGURE 24 C) and BMNH (E) #674767;
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;
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.
Deposition summary: BMNH (HT ♂, 2PT ♂♂, 3 PT ♀♀), MNHN (PT ♂), ABRI (PT ♂).
Type locality. Madagascar C, Amboasary-an-ala, vic. Anjozorobe, 18.45o S, 47.95o E +/- 2.45 km, 1375 +/- 75 m.
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 View FIGURE 24 A– C), and the HWV tends to have the space-CuA2 ocellus expressed as small black-ringed ocellus. However, Ht. oberthueri ( Fig. 24 View FIGURE 24 D–E, Fig. 26 View FIGURE 26 A) can be distinguished from the sympatric Ht. andasibe by the distinctly more outangled HWV Mb of that species ( Fig. 22 View FIGURE 22 C; Fig. 26 View FIGURE 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 View FIGURE 22 AB).
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.
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 ♀♀).
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 View FIGURE 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 View FIGURE 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).
Palps: penultimate segment with narrow brown medial strip, flanked by yellow and fringed by dark brown scales, with yellow scales mainly on mesad face.
♂ genitalia: 228DL, PT ( Fig. 25 View FIGURE 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.
Etymology. Etymology. After ‘Tiana’ Raharitsimba (= Heritiana Rahagalala), who found some of the first modern exemplars of this species at Anjozorobe.
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 View FIGURE 22 C–D) were examined and are clearly different, along with Ht. oberthueri , described below.
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 View Materials , based on a ♂ from Anjozorobe, 1400 m.; see correction below) is considerably less pairwise divergent (4.76%), however, to Ht. turbans ( AY 040130 View Materials , based on 2 ♂♂ and 1 ♀ from Ranomafana National Park; which has COI-5P cluster number BOLD:ACD8579).
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).
Ecology and distribution.
Habitat: montane rainforest.
Behaviour: both sexes come readily to fruit bait. Flies in substratum of forest.
Hostplant: unknown, presumed to be grasses.
Early stages: unknown.
Distribution: endemic to the Angavo massif, including Anjozorobe and Andrangoloaka, as far as is known ( Fig. 30 View FIGURE 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.”
Elevational range: 1320–1375 m. (n=27 including referred specimens and observations).
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].
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