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Lees, N G, QX13947
This record was harvested from a previous catalogue system and will be withdrawn in 2025. Information in this record may be superseded or incomplete. Visit this record in UMA's new catalogue at: https://archives.library.unimelb.edu.au/nodes/view/398970Surname: LEES. Given Name(s) or Initials: N G. Military Service Number or Last Known Location: QX13947. Missing, Wounded and Prisoner of War Enquiry Card Index Number: 26141.216376
Item: [2016.0049.31263] "Lees, N G, QX13947
Modificacions de les Propietats Fisicoquímiques de les "Lies" de Segona Fermentació durant la Criança del Cava
[cat] Els caves són vins escumosos que es distingeixen per la seva criança particular en contacte amb els llevats de la segona fermentació per un període no inferior a 9 mesos. Com més es perllonga la criança, se suposa una millor qualitat del vi. Així, la categoria “Gran Reserva” implica com a mínim 30 mesos d’envelliment en rima. Tanmateix, encara no s'han esclarit els mecanismes que involucren l’autòlisi amb la qualitat organolèptica dels vins escumosos.
En aquest treball s’ha proposat identificar possibles mecanismes que expliquin la relació entre la criança del vi escumós en contacte amb les lies i el temps d’envelliment. Per fer-ho, s'han avaluat els canvis a les propietats fisicoquímiques de les lies atenent a les seves modificacions ultraestructurals al llarg de la criança i s'ha estudiat la capacitat de les lies per retenir compostos volàtils que participen en l’aroma del vi, així com la seva possible intervenció en la protecció del vi enfront de l’oxidació.
Les propietats fisicoquímiques de les lies estudiades han estat: hidrofobicitat, capacitat electródonador i electró-receptor i potencial “Xi”; les quals varien amb el període de criança del cava. A més, aquests canvis s'han relacionat amb les modificacions morfològiques de les cèl•lules.
Així, les lies de segona fermentació pateixen profunds canvis a la seva ultraestructura al llarg de la rima, els quals impliquen a tots els orgànuls cel•lulars. Es desestructura profundament el citoplasma i desapareixen els orgànuls. La membrana plasmàtica es trenca, augmenta l’espai periplasmàtic i la paret perd progressivament la part amorfa de la seva estructura, relacionada amb el glucà, i exposa la capa fribril•lar interna, relacionada amb les mannoproteïnes. A més, sembla que el vi on es duu a terme la segona fermentació no influeix en la cinètica de degradació del llevat.
Pel que fa a la hidrofobicitat de les cèl•lules de llevats, disminueix amb el temps de rima, i aquesta propietat està relacionada amb el seu caràcter electró-donador i amb l’increment de potencial “Xi” i del caràcter electró-receptor. La davallada de la capacitat per flocular de les lies de segona fermentació sembla ésser deguda a les proteïnes de superfície i estar correlacionada amb la pèrdua de la hidrofobicitat.
Finalment, les lies en contacte amb el vi retenen compostos volàtils que participen a l’aroma del cava. La desestructuració de la paret cel•lular modificarà els compostos volàtils retinguts així com aquells que es remouran durant el procés de degollament del cava. D'altra banda la superfície de les cèl•lules també ha mostrat tenir una capacitat antioxidant similar a la de fruites i vegetals, després d’ésser analitzades mitjançant els mètodes de FRAP i DPPH. Aquesta capacitat es redueix amb el temps de rima. Els polifenols retinguts semblen ésser els majors contribuïdors al poder reductor de les lies, seguits dels tiols i, finalment, dels mannans i glucans.[eng] Aging on lees after second fermentation is a fundamental stage in the production of some high quality sparkling wines by the traditional method, and it results in an increase in product richness and roundness. During this ageing, lees interact with the wine and undergo important modifications to their structure, due to the self-degradation process known as autolysis.
Cell wall biochemical components confer physicochemical surface properties that enable yeasts and lees to interact among them and with other compounds. These interactions are mainly referred to lees sorptive properties toward organic compounds, to the protective effect of lees toward wine oxidation, and to their flocculation capacity.
Yeast lees have shown the capacity to interact with wine organic compounds such as volatiles and polyphenols. The sorption capacity of yeast surface seems to be related to both chemical properties of the sorbed substances and cell surface characteristics. These sorptive phenomena are thought to influence the chemical composition of wines during their ageing on lees.
The phenomena related with the antioxidant properties of yeast cells is other feature with relevance in wine technology. Lees’ antioxidant properties could be attributed to three main mechanisms: The action of enzymes and biomolecules released during autolysis, the effects of membrane lipids and by means of elements in cell wall (constitutional or retained during aging).
Finally, Flocculation capacity is especially important in the production of sparkling wine by the méthode champenoise, in which the yeast cells can only be removed from the bottle by being settled. The flocculation seems to be related with the presence of flocculins (surface proteins), calcium and mannose. Moreover, the behaviour of cells seems strongly conditioned by their cell surface physicochemical properties. The changes in lees wall structure caused by the autolytic process during sparkling wine ageing could induce relevant modifications in the cell surface structure and physicochemical characteristics and thus in the above mentioned lees properties
Heteropsis oberthueri Lees, sp. nov.
Heteropsis oberthueri Lees, sp. nov. LSID: urn:lsid:zoobank.org:act:FEE 8 CA 6 F-A 53 B- 43 A 5-8 C 9 F-E 5 D 69 AD0E 468 Prior references: sp. 89, sp. 87 (on some envelopes). “KA 518 _Heteopsis_ anceps ” [sic] in Aduse-Poku et al., (2015, Fig. 1). Type material. Deposition BMNH: Holotype: ♂ (Fig. 24 D), Madagascar E, Zahamena NW, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 5 / 11 / 2006, D.C. Lees: DL 06-064, G04 [leg for DNA], NHMUK 010289159 [QTR barcode]. Paratypes: Deposition BMHN: ♂, Madagascar E, Ambavala camp, Zahamena, 17.5451 o S, 48.7237 o E +/- 0.5 km, 1300 +/- 50 m, 6 / 11 /2006, 13: 10, D.C. Lees: DL 06- 130 A, NHMUK 010289160 [QTR barcode]; ♀ (Fig. 24 E), E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: DL 06- 130, IA 585 [isotope voucher], NHMUK 010289161 [QTR barcode]; ♂, E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: 371 DL, NHMUK 010289192 [QTR barcode]; ♂, E, Zahamena NW, 17.53 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 6 / 11 / 2006, D.C. Lees: DL 06-086, IA 586 [isotope voucher], NHMUK 010289162 [QTR barcode]; ♂, E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6- 7 / 11 / 2006, D.C. Lees: DL 06- 1001, IA 324 [isotope voucher], NHMUK 010289163 [QTR barcode]; ♀, E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 /2006, 11: 30, D.C. Lees: DL 06- 112, IA 587 [isotope voucher], NHMUK 010289164 [QTR barcode]; ♀, E, Towards Ambavala camp, Zahamena, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: DL 06- 141, IA 588 [isotope voucher], NHMUK 010289165 [QTR barcode]; ♀, E, returning to Bemoara camp, Zahamena, 17.5356 o S, 48.71875 o E +/- 0.70 km, 1180 +/- 100 m, 5 / 11 /2006, 14: 59, D.C. Lees: DL 06-059, IA 589 [isotope voucher], NHMUK 010289166 [QTR barcode]; Deposition MNHN: ♂, Madagascar E, Ambavala, Zahamena NW, 17.5451 o S, 48.7237 o E +/- 0.25 km, 1325 +/ - 50 m, 6 / 11 /2006, 14: 11, D.C. Lees: DL 06- 117, IA 584 [isotope voucher], DSC 03096.jpg [image]; Deposition ABRI: ♂, Madagascar E, Zahamena NW, 17.5376 o S, 48.71965 o E +/- 0.94 km, 1180 +/- 125 m, 6 / 11 / 2006, D.C. Lees: DL 06- 121. Deposition summary: BMNH (HT ♂, 4 PT ♂♂, 4 PT ♀♀), MNHN (PT ♂), ABRI (PT ♂). Type locality. NW Zahamena, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m. 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-M 3 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-M 3 as well as in space-CuA 1 expressed on the HWD (Fig. 19 C). The ventrally very similar Ht. roussettae has a strong tendency to expression of a small ocellus M 3 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. Description. Wings: description is similar to that of Ht. tianae. Upperside uniform mid brown, with FW space-CuA 1 ocellus featuring an orange ocellus ‘ring’ which is sometimes rather ‘squashed’ along the diagonal from mid-costa to tergal angle. FWD space-M 1 ocellus small with narrow orange ring. HWD space-CuA 1 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 1 A+ 2 A, 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-CuA 1 ocellus with orange ring yellowing proximad, following darkish brown concave curve of Mb before it bends back to mid-costa. Space-M 1 ocellus FWV a white pupil in HT. On HWV, space-CuA 1 ocellus small and slightly elliptic with narrow black iris, also without orange ring trace. HWV space-Rs-M 2 ocelli as white pupil-points and space-CuA 2 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-M 2. 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- M 1 ocellus FWV sometimes features a narrow black iris and space-M 2 sometimes just as a white pupil without narrow black iris nor trace of orange ring. Space-CuA 2 ocellus HWV slightly expressed in some specimens. ♀♀ similar in pattern, but larger and somewhat lighter dorsally with a more rounded HW shape. 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 ♀♀). Androconia: HWD vein 1 A+ 2 A has a narrowly tapered inflation and 3 A is much more swollen distad, from base to mid-vein, covered in thin grey-brown scales. The length of HW veins M 2, M 3, CuA 1 and CuA 2 are also narrowly inflated, and have specialized scales on the dorsal surface. Abdominal black androconia are just visible ventro-laterally around A 4 –A 5. HWD discocellular brush dark brown. Sdp HWD grey, small, lenticular, composed of narrow grey scales). 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. ♂ 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). Etymology. after Charles Oberthür, who in 1916 produced one of the first treatments, a seminal work on the Malagasy Mycalesina. 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). Additional information. DNA divergences: COI- 5 P cluster number BOLD:ACW 4996 (exemplar BMAD 242 - 15, DL 06- 11), about 2.58 % divergent to Ht. tianae (BOLD:AAE 4112, exemplar BMAD 200 - 15, DL 14 Z-051). 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- 5 P barcode. Ecology and distribution. Habitat: primary rainforest. Behaviour: flies low. Hostplant: unknown, but likely to be low-growing grasses. Early stages: unknown. Distribution: as far as is known, endemic to RNI Zahamena (Fig. 30 C, brown dots). Elevational range: 1180–1310 m. (n= 27, including referred specimens). Referred specimens. ♀, Madagascar E, returning to Bemoara camp, Zahamena, 17.5376 o S, 48.71965 o 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, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 5 / 11 / 2006, D.C. Lees: DL 06-061; ♂, E, Zahamena NW, 17.53 o S, 48.7269 o 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, 17.5376 o S, 48.71965 o 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, IA 432 [isotope voucher], KAP 518 [=KA-P 518; 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, 17.5451 o S, 48.7237 o 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, 17.4946 o S, 48.733 o E +/- 1 km, 1321 m, 7 / 11 /2006, 10: 37, D.C. Lees: DL 06- 156; ♂, E, near Ankosy summit, Zahamena, 17.49405 o S, 48.73325 o E +/- 0.067 km, 1300 +/- 25 m, 7 / 11 / 2006, D.C. Lees: DL 06- 188; ♀, E, Zahamena, 17.5131 o S, 48.7269 o E +/- 1.5 km, 1064 +/- 25 m, 6 / 11 / 2006, D.C. Lees: DL 06- 67, IA 69 [isotope voucher]; ♂, E, Zahamena NW, 17.4946 o S, 48.733 o E +/- 1 km, 1321 +/- 25 m, 6 / 11 / 2006, D.C. Lees: 364 DL [genitalia voucher], IA 29 [isotope voucher], leg sample [KAP].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 83-85, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis kremenae Lees, sp. nov.
Heteropsis kremenae Lees, sp. nov. LSID: urn:lsid:zoobank.org:act:D 5567 C 45 - 4 EB 9 - 4 E 62 - 8 F 67 -ABF 2 C 4008 F 4 E Prior references: sp. 14 A (Lees 1997, Torres et al., 2001: 462). Type material., Deposition BMNH: Holotype: ♂ (Fig. 13 A), Madagascar NE, Ambavaony R. [= Ambanivony R.], R. Onive (Ankavanana), E Masoala, 250 m, 15.282 o S, 50.288 o E +/- 0.1 km, 250 +/- 50 m, 28 / 11 / 1993, D.C. Lees: DL 93 -0021, NHMUK 010289135 [QTR barcode]. Paratypes: Deposition BMNH: ♂, Madagascar NE, Ambavaony R., E. Masoala, 125 m, 15.279 o S, 50.288 o 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 14 LO, disturbed forest, NHMUK 010289137 [QTR barcode]; ♂, NE, Ambavaony R., E. Masoala, 280 m, 15.282 o S, 50.288 o 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, 15.784 o S, 50.2164 o E +/- 0.15 km, 45 m, 28 / 1 / 1994, D.C. Lees: DL 94 -0004, E 105 [egg voucher], W 7 L 1-45 M 1 LO [fruit trap], NHMUK 010289139 [QTR barcode]; ♀, NE, Ambavaony R., E Masoala, " 200 m. ", 15.282 o S, 50.288 o E +/- 0.1 km, 200 +/- 50 m, 28 / 11 / 1993; D.C. Lees: DL 93 -0009 [DNA voucher], E 4 [egg voucher: “ 3 greenish eggs expressed 29 / 11 ”], NHMUK 010289140 [QTR barcode]; ♂ (Fig. 15 A), NE, Masoala E, Ambanivony [=Ambavaony], 15.288 o S, 50.288 o E +/- 0.5 km, 50 +/- 25 m, site W 2 -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., 15.295 o S, 50.227 o E +/- 0.15 km, 275 m, 11 / 12 / 1993, H. Raharitsimba: CK 93 -0020; NHMUK 010289184 [QTR barcode]. Deposition MNHN: ♂, NE, Ambavaony R., E. Masoala, ca, 100 m, disturbed streamside, 15.279 o S, 50.288 o E +/- 0.12 km, 180 +/- 75 m, 28 / 11 / 1993, C. Kremen: DL 93 -0007; ♂, NE, Antsamanarana R., Masoala E., "~ 50 m, Piste C", 15.307 o S, 50.233 o E +/- 0.1 km, 115 +/- 75 m, 11 / 12 / 1993; D.C. Lees: DL 93 -0008; ♀, NE, Antafononana lower Anaovandrano R., W 7 -L 2, Masoala E., 15.7458 o S, 50.1858 o E +/- 0.15 km, 40 m, H. Raharitsimba: DL 94 - 0 0 0 3, W 7 L 2-40 M 2 LO [fruit trap], IA 600 [isotope voucher]; Deposition ABRI: ♂, NE, Ambery R., Masoala E., 15.3716 o S, 50.4263 o E +/- 0.98 km, 26 m, 22 / 12 / 1994, D.C. Lees: DL 93 -0006; ♀, NE, Masoala PN, W 7 L 2, Antafonona, 40 m, fruit trap 1 Hi, riparian, 15.738 o S, 50.182 o E +/- 1.85 km, 185 +/- 145 m, 17 / 1 / 1994, H. Raharitsimba: DL 94 -0002. Deposition summary: BMNH (HT ♂, 3 PT ♂♂, 3 PT ♀♀); MNHN (2 PT ♂♂, PT ♀); ABRI (PT ♂, PT ♀). Type locality. Madagascar NE, Ambavaony R., R. Onive, E Masoala, 250 m, 15.282 o S, 50.288 o E +/- 0.1 km. Diagnosis. Apparently most similar to Ht. pauper, from which it differs by rounder wings especially in FW wingshape (as in Ht. subsimilis and Ht. avaratra sp. nov.), and more generally by a larger space-CuA 1 HWD ocellus and more contrasting orange and yellow banding/shading on the underside, as well as a more consistently orange FWV ‘shoulder’ at the base of the costa). ♂♂ can be distinguished by HWD Spdb tending to be blackish (whitish in Ht. pauper). Description. Wings: Dorsal wing surface uniform mid-brown, FWD space-CuA 1 ocellus with black iris at least spanning veins CuA 1 -CuA 2, light orange ring narrow to wide and sub-hexagonal, sometimes compressed along diagonal from tergal angle to mid costa. FWD space-M 1 ocellus very small, with narrow pale orange ring. HWD ocellus is the only one expressed there and fairly round with fairly narrow pale orange ring and spanning about 2 / 3 of vein CuA 1 -CuA 2, on FWV space-CuA 1 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- 1 A ocellus much more strongly expressed than on HWD and sub-elliptic, black iris spanning most of vein CuA 1 -CuA 2 and with a concentric yellow ring that spans the whole space-CuA 1. 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 M 2 and then fairly straight to mid space- CuA 2 and angled back to 1 A. 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- CuA 2, often as white spots. HWV space-Rs ocellus sometimes expressed, even when small, with elliptic black iris and narrow yellow ring. Wingspan/fwl: range 31.0– 33.9 /17.0– 18.6 mm (♂♂, n= 5), mean = 33.2 +/- 2.1 SD/ 17.8 +/- 0.8 SD mm (n= 3 ♂♂), including HT ♂ 33.9 / 18.6 mm; range 33.8–37.5 / 18.1–20.7 mm (♀♀, n= 6); mean = 35.9 +/- 1.4 SD/ 19.2 +/- 1.1 SD mm (n= 3 ♀♀). Androconia: Sdb HWD compact, dark brown to blackish, Sdp HWD fat lenticular, black, CuA 2 inflated vein narrowly swollen to about 2 /3, 1A+ 2 A 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 M 3 for the medial system. Palps: penultimate segment on outside face light creamy white towards compound eye, blackish away from eye with light coloured scales within this border; fringed with dark brown where not contacting eye. Face away from eye whitish ochreous. ♂ genitalia: 19 DL, PT (Fig. 15 A); Lees (1997: 106, Fig. 7 f; “ 14 A”): from LV: miniaturised, about 1.7 mm long and with configuration typical of the Ht. subsimilis group; there is no really obvious difference in shapes of the genitalic components from other members (Lees (1997: 106) and the description below might apply to almost any of the group. From LV, the valve bases are rather symmetrically-‘skittle’-shaped (Lees 1997). Uncus is slightly longer than tegumen and quite inflated dorsoventrally before the slightly pointed downturned tip (inflated before tip from the DV). Gnathos tapered from small base, fairly straight, not sinuate from the SV. Valve with prominent rounded dorsal shoulder, valve arm tapering to distinct ‘beak’ that points to uncus base with limited serration and not much indication of a club at valve end (tip strongly incurved from the SV). Saccus relatively small, slightly bulbous, juxta not very prominent proximad. Aedeagus about 1 / 5 longer than valve, moderately stout and recurved distad of ostium. Etymology. After Claire Kremen, who worked tirelessly for the creation of the Masoala National Park, that was created in 1994. Discussion. No historical museum material has been found and this species was first recognized in the field in 1993 in the survey of what is now Masoala National Park (Lees, 1997: 64). Kremen et al., (2001: 412) considered it a potential endemic there, but it has since been discovered to be more widespread among the remaining northeastern lowland rainforests. All available types in the Ht. subsimilis group were examined, from which it differs. Concerning Culapa pauper Oberthür, 1916 with over 300 STs, a ♂ LT is here designated: BMNH (E) # 674888; Madagascar Antsianaka Perrot Freres 2 e 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 2 e Semestre 1890 | Culapa undulata Obthr. ♂ type |[copy of f. 3064]|P.E.L. Viette det. 1968 Culapa undulata Ch. Obthr. ♂ LECTOTYPE |supposed “ Type ” (syntype) of Culapa undulata Oberthür ”| BMNH (E) # 674883; Oberthür specified as STs 190 other ♂♂ (including BMNH (E) # 674885) and 58 ♀♀ (including BMNH (E) # 674884, Fig. 14 B, illustrated in Oberthür 1916: P. 367, f. 3065 and BMNH (E) # 674886), which are then automatically PLTs of Culapa undulata). Additional information. DNA divergences: cluster number BOLD:ACW 4937 (exemplar BMAD 249 - 15, Marojejy), diverges by 5.47 % from that of H. avaratra (cluster number BOLD:AAD0195, exemplar BMAD 016- 09) and by about 5.5 % from that of H. subsimilis (cluster number BOLD:AAB 4493). In the Torres et al., (2001) dataset, a COII sequence for Ht. kremenae (their “ Hen. sp. 14 A”) from Masoala is 7.03 % pairwise divergent to Ht. avaratra from Montagne d’Ambre (AY 040161 compared to AY 040146 based on single individuals respectively, 398 bp comparable) and 8.14 % pairwise divergent to Ht. subsimilis from Ranomafana National Park (AY 040145), whereas 8.83 % pairwise divergent to Ht. pauper from that site (AY 040133, 414 bp compared). Meanwhile, Ht. avaratra and Ht. subsimilis are more obviously closely related according to COII data (5.27 % pairwise divergent) and share privately at least four SNPs not found elsewhere in the Ht. s ubsimilis group. Note that in the Torres et al., dataset, “ Henotesia 14 ” (AY 040181, cytochrome b) from Masoala is apparently synonymous with Ht. subsimilis (AY 040192), with only 1 bp difference. A similar situation seems to pertain to their grouping of (‘ Hen. sp. 23 ’ + ‘ Hen. sp. 30 ’ [representing opposite sexes of Ht. pauper from Ankazomivady] + Ht. pauper from Ranomafana + “ Hen. sp. 7 ” from Masoala), the latter ‘morphospecies’ exhibiting only 3 bp difference from the first three (COII); see also the COI phylogeography of Ht. pauper in Linares et al., (2009: 488–489). Phylogeny/sister species: Ht. kremenae is likely most closely related to Ht. subsimilis and Ht. avaratra. Lees (1997) did not resolve the position of Ht. kremenae ‘FRTNA’] within the Ht. subsimilis group. The two-gene study of Linares et al., (2009) did not include this species and its exact position was not clear in Aduse-Poku et al., (2016, in press). Ecology and distribution. Habitat: Primary rainforest, preferring riparian areas. Behaviour: flies among low grasses, liking the slopes above streams and rivers. Hostplant: not reared, but presumably low forest grasses. One was caught though in a canopy fruit trap in Marojejy (K. Aduse-Poku, pers. comm.). Early stages: expressed eggs were greenish. Distribution: endemic to lowland rainforest in the northeast of the rainforest belt and Masoala Peninsula (Fig. 30 B, dark blue dots), with a preference for riparian areas. Elevational range: 10–675 m (n= 86, including referred specimens and observations). Referred specimens. ♂, NE, Marojejy PN, riparian forest near camp 1, 14.4377 o S, 49.7756 o 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, 14.4377 o S, 49.7756 o 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, 14.4548 o S, 49.792 o 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 14 M-0037, IA 528 [isotope voucher]; CCDB-02230-E 11 [DNA barcode voucher]; ♀, NE, Marojejy, 700 m, 24 / 1 / 2014: 11: 21, D.C.Lees: DL 14 M-0035, IA 532 [isotope voucher]; ♂, NE, Sahantaha, Makira, 15.2337 o S, 49.5311 o E +/- 0.15 km, 500 +/- 130 m, 29 / 1 / 2003, D.C. Lees: BMNH (E) # 697172 [DNA voucher]; ♀, NE, Andreketa near 17 B 364 steep rainforest with lots of “Tsongolovo” 15.2642 o S, 49.5479 o 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, 15.2642 o S, 49.5479 o E +/- 0.75 km, 375 +/- 25 m, 17 / 1 / 2003, D.C. Lees: BMNH (E) # 697950 [DNA voucher]; specimen, NE, Ambodivoangy forest, 15.29 o S, 49.62 o E +/- 1 km, 50 +/- 25 m, 4 / 12 / 2001; ♂, NE, Ankirindro, 15.2904 o S, 49.5474 o 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, 15.295 o S, 50.227 o E +/- 0.15 km, 275 m, 11 / 12 / 1993, H. Raharitsimba: TR 89; ♂, NE, Antsamanarana R., Masoala E., 50 m, 15.307 o S, 50.233 o E +/- 0.1 km, 50 m, 12 / 12 / 1993; H. Raharitsimba: TR 93; ♂, NE, Vohitaly, 15.4377 o S, 49.5344 o E +/- 0.15 km, 28 / 12 /2002, 15: 15, D.C. Lees; ♂, NE, Vohitaly, Makira, 15.4379 o S, 49.5344 o E +/- 0.15 km, 25 / 12 / 2002, D.C. Lees: DL-SF 3, KA 552 [=KA-P 552, DNA extract number]; ♂, NE, Ambatoavy, Masoala, 630 m, 15.658 o S, 50 o E +/- 1.26 km, 630 m, 14 / 2 / 1993, D.C. Lees: H 14-14293 - 1; ♀, NE, Ambavaony R., E Masoala, 125 m, 15.279 o S, 50.288 o E +/- 0.12 km, 125 m, 30 / 12 / 1993; C. Kremen: E 15 [egg voucher], DNA DCL 32 [DNA voucher]; ♀, NE, Iketry [Iketra], lower Anaovandrano R., W 7 -L 2, Masoala E., 15.7458 o S, 50.1858 o E +/- 0.15 km, 40 m, 16 / 1 / 1994, D.C. Lees, W 7 L 2-40 M 2 LO [fruit trap]; ♀, NE, Masoala, 15 / 1 / 1994, WG 2 [wing voucher], IA 33 [isotope voucher]; ♀, NE, Masoala, 16 / 1 / 1994, IA 36 [isotope voucher]; ♂, NE, Masoala, 50 m, 12 / 12 / 1993, H. Raharitsimba, WG 6, [wing image], IA 39 [isotope voucher]; ♂, NE, Masoala, 26 / 11 / 1993, H. Raharitsimba: TR 47, IA 47 [isotope voucher]; ♂, NE, Ivontaka R. camp, between Ivontaka N and S. reserves, ca. 90 m, 16.294 o S, 49.81 o E +/- 0.7 km, 90 +/- 25 m, 3 / 2 / 1995; D.C. Lees: DL 95 -0003A, H 14 A 3295 - 1 [leg sample]; specimen, NE, Ivontaka R., ~ 120 m, 16.294 o S, 49.81 o 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, 16.297 o S, 49.785 o E +/- 2 km, 230 +/- 130 m, 3 / 2 / 1995, 11: 46 – 16: 10, D.C. Lees: H 14 A 3295 - 2 [leg sample], IA 601 [isotope voucher]; ♀, NE, Ivontaka Sud, 16.294 o S, 49.81 o E, 90 m, 2 / 1995, D.C. Lees, IA 338 [isotope voucher]; ♀, NE, Ivontaka Sud, path to Marokoto, 16.297 o S, 49.785 o E +/- 2 km, 230 +/- 130 m, 3 / 2 / 1995, D.C. Lees: H 14 A 3296 - 2, IA 339 [isotope voucher].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 46-51, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis menamenoides Lees, sp. nov.
Heteropsis menamenoides Lees, sp. nov. LSID: urn:lsid:zoobank.org:act:D 2 FCD 726 -B 198-484 F- 846 C-B 5229 FE 174 E0 Prior references: sp. 62 (Lees 1997; Torres et al., 2001: 462). Type material., Holotype, deposition BMNH: ♂ (Fig. 21 A), Madagascar NW, Bekolosy, RS Manongarivo, 950 [880] m, [14.0487 o S, 48.29495 o E +/- 0.15 km, 95 m +/- 50 m], 15 / 12 / 1994: 09: 50, D.C. Lees: DLBEK 94 _ 160; KAP 16 [=KA-P 16 DNA extract voucher], IA 312 [isotope voucher], NHMUK 010289153 [QTR barcode]. Paratypes: Deposition BMNH: ♂, NW, Bekolosy, RS Manongarivo, ca. 800 m, [14.05135 o S, 48.2937 o 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; HM 40425 [DNA barcode voucher], KA 571, KA 4084 [=KA-P 571, KA-P 4084; DNA extract numbers]; ♂ (Fig. 18 D), Madagascar NW, Bekolosy, RS Manongarivo, 890 m, 14.04936 o S, 48.294 o 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, 14.04894 o S, 48.2945 o E +/- 0.15 km, 925 +/- 50 m, 13 / 12 /1994, 09: 56 – 11: 15, D.C. Lees: DLBEK 94 _ 181, NHMUK 010289187 [QTR barcode]. Deposition summary: BMNH (HT ♂, 3 PT ♂♂). Type locality. Madagascar NW, Bekolosy, RS Manongarivo, 950 m, [14.0465 o S, 48.3 o E +/- 0.2 km, 950 +/-70 m]. 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-CuA 1 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-CuA 1 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. 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-M 3 of FWD and HWD. Space-CuA 1 ocellus expressed more strongly in FWD than in HWD; Orng not evident and black iris considerably smaller diameter than spacing of veins CuA 1 and CuA 2. 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-CuA 1 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-M 2 ocellus as small white point surrounded by narrow black ring in FWV while space-CuA 1 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. 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. Androconia: black ventral abdominal patch between A 3 –A 6 divided centrally by narrow yellow overlying scales, in potential contact with diffusely separated and slightly wavy androconial brush largely emanating from base of space-CuA 2 before its fork with vein CuA 1 and overlying veins 1 A+ 2 A and 3 A; the latter symmetrically inflated before midlength in compact ‘tear’-shape (Lees, 1997: 97, Fig. 3 b), whereas 1 A+ 2 A shows very restricted swelling towards the base of 1 A. 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. 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. ♂ genitalia: 231 DL (DLBEK 94 _ 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. 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). 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. & B. Perrot| Culapa grandis Obthr. ♀ type |[Copy of f. 3074]”). Additional information. DNA divergences: COI- 5 P cluster number BOLD:AAE 4113 (exemplar DLBEK 94 _ 179, BMAD 075-09, HM 404251). The DNA barcode is about 5.5 % divergent to an undescribed species, sp. 28 (cluster number BOLD:AAE 5458), and about 5.8 % pairwise divergent to that of Ht. tianae (BOLD:AAE 4112). Phylogeny/sister species: unknown, new molecular study awaited. Ecology and distribution. Habitat: on a very steep slope with a fine leaved bamboo that was considered to be ‘ Nastus ’ manongarivensis A. Camus, not necessarily the hostplant. Behaviour: flies in the forest substory. Hostplant: unknown, but possibly grasses. Early stages: unknown. Distribution: endemic as far as is known to Mt. Bekolosy, Reserve Speciale de Manongarivo (Fig. 30 C, slateblue dots). Elevational range: 800–1035 m. (n= 8 incl. observations). 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. Referred specimens. None.Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 72-74, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Heteropsis barbarae Lees & Kremen, sp. nov.
Heteropsis barbarae Lees & Kremen, sp. nov. LSID: urn:lsid:zoobank.org:act:DC 87986 A- 3822 - 43 C 9 -B 1 B 9-83 A 2714 D088B Prior references: sp. 16 (Lees 1997; Torres et al., 2001: 462). Type material., Deposition MNHN: Holotype: ♂ (Fig. 20 A), Madagascar NE, Maitsoarongana, Sikiory River, near camp, 15.1885 o S, 49.5163 o 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]. Paratypes (accession BMNH (E) 2008 - 69): ♂, Madagascar NE, Maitsoarongana, Sikiory R., 660–750 m, 15.1907 o S, 49.5309 o 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, 15.1885 o S, 49.5163 o 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, 15.1907 o S, 49.5309 o 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, 15.1848 o S, 49.4659 o 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, 49.5309 o E, +/- 2 km., 750 +/- 100 m., 13 / 12 / 2001, D.C. Lees: DL 01- 248, BMNH (E) 1717095 [QTR barcode]. Deposition summary: BMNH (HT ♂, 4 PT ♂♂, 1 PT ♀). Type locality. Madagascar NE, Maitsoarongana, Sikiory River, 15.1885 o S, 49.5163 o E +/- 0.02 km, 685 +/- 23 m. 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). Description. Wings: Upperside mid-brown with a slight russet brown cast. Space-CuA 1 ocellus expressed strongly in FWD; orange ring sub-hexagonal., M 1 ocellus very small black white pupilled spot surrounded by a narrow orange ring. HWD ocellus CuA 1 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- M 2 to CuA 1. FWV space-CuA 1 ocellus similar size to FWD (in the HT), as is FWV M 1 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-CuA 1 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-CuA 1 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 1 A. 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-CuA 1 ocellus FWD is a little wider proximad in some specimens, and FWD M 1 ocellus sometimes not visible. HW space-CuA 1 ocellus is more circular in some specimens. 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). Androconia: (observations on ♂ specimen MAD 182 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 1 A+ 2 A base, mid brown, dispersed, juxtaposed potentially under abdominal sternal segments A 2 –A 5, 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 2 A swollen within basal half. Palps: Ochreous, with intermingled brown scales on outside face away from compound eye, dark brown scales above where potentially brushing eye. Etymology. After Barbara, the mother of Claire Kremen. 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 |MadagascarNo 10 [?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.&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.&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 ”. Additional information. ♂ genitalia: 141 DL (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. DNA divergences: COI- 5 P cluster number BOLD:ACW 4995 (exemplar BMAD 248 - 15, DL 14 M-0033, Marojejy) is 4.86 % divergent to Ht. tianae (BOLD:AAE 4112), and 5.63 % divergent to Ht. maeva (BOLD:AAE 5488), 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). 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). Ecology and distribution. Habitat: riparian primary rainforest. Behaviour: flies low and both sexes come easily to fruit bait. Hostplant: unknown but almost certainly low grasses. Early stages: unknown but for expressed eggs, which were whitish. Distribution: endemic to the northeastern rainforests of Madagascar, from Marojejy and Masoala south to Ambatovaky (Fig. 30 C, light green dots). Elevational range: 0– 890 m. (n= 153, including referred specimens and observations). Referred specimens. ♀, Madagascar NE, Marojejy PN, Camp II [above], 14.4344 o S, 49.759 o E +/- 0.25 km, 850 +/- 50 m, 12 / 11 / 2006: 10:00, D.C. Lees: DL 06- 298; ♂, NE, Marojejy PN, 14.4344 o S, 49.7606 o 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, 14.4347 o S, 49.7601 o E +/- 0.5 km, 730 +/- 50 m, 10 / 11 / 2006, D.C. Lees: DL 06- 169; specimen, NE, Marojejy PN [between camp 1 and camp 2], 14.4362 o S, 49.7679 o E +/- 0.85 km, 613 +/- 125 m, 10 / 11 /2006, 15: 58, D.C. Lees: DL 06- 170; ♀, NE, Marojejy PN, 14.4377 o S, 49.7756 o E +/- 0.25 km, 400 +/- 50 m, 20 / 11 / 2006: 15:07, D.C. Lees: DL 06- 456; ♂, NE, Marojejy PN, 14.4425 o S, 49.7818 o 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, 14.45015 o S, 49.786 o E +/- 1.78 km, 326 +/- 175 m, 10 / 11 / 2006: 13: 16, D.C. Lees: DL 06- 204; ♂, NE, Marojejy PN [around entrée], 14.4626 o S, 49.7964 o 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], IA 387 [isotope voucher]; ♂ [BMNH], NE, Andranobe, Masoala Peninsula, 15.6816 o S, 49.9573 oE +/- 1.31 km, 0 m, 3 -Dec- 91, C. Kremen: CK 826, BMNH (E) 2008 - 69; ♂, NE, Masoala, Anaovandrano, 14 / 1 / 1994, D.C. Lees: WG 5 [wing image], IA 38 [isotope voucher]; ♂, NE, Masoala, Ambavaony, 125 m, 27 / 11 / 1994, D.C. Lees: DCL 47 a, IA 108 [isotope voucher]; ♂, NE, Masoala, Ambanivony, 25 / 11 / 1993, C. Kremen: CK 26, IA 109 [pheromone voucher]; ♀, NE, Makira, Vohitaly, 15.4414 o S, 49.5309 o E +/- 0.15 km, 645 m, 30 / 12 / 2002, D.C. Lees: DL-SF00, IA 112 [isotope voucher]; ♀, NE, Makira, Ankirindro, 15.2931 o S, 49.5472 o E +/- 0.15 km, 675 +/- 25 m, 15 / 1 / 2003, D.C. Lees: DL- SE00, IA 114 [isotope voucher]; specimen, NE, Ankiatomboka forest, c. 710 m, 15.179 o S, 49.412 o E, 710 +/- 50 m, 9 / 12 / 2001, D.C. Lees: DL 01- 169; ♂, NE, Bivontro, Makira, 15.3984 o S, 49.4477 o E +/- 0.1 km, 762 +/- 25 m, 21 / 12 / 2002, D.C. Lees: DL-SG 77; ♂, NE, Vohitaly-Anjiahely, 15.3989 o S, 49.5211 o E +/- 1.82 km, 435 +/- 50 m, 31 / 12 / 2002, BMNH (E) # 672331 [DNA voucher]; ♀, NE, Anjiahely, 0.37 km E., 15.4120 o S, 49.5022 o E +/- 0.25 km, 380 +/- 5 m 12 / 12 / 2002, D.C. Lees: DL-SG 1, BMNH (E) 2008 - 69, BMNH (E) 1717101; ♂, NE, Sahantaha, near ‘fanihy’ roost area, 15.1963 o S, 49.5564 o E +/- 0.98 km, 517 +/-54 m, 13 / 12 / 2001: 10: 46, D.C. Lees: DL 01- 257; ♂, NE, Anjanaharibe Mt., 15.188 o S, 49.6139 o E +/- 0.1 km, 475 +/- 5 m, 6 / 2 / 2003, D.C. Lees; ♀, NE, Tampolo, W. Masoala, ca. 150 m, primary rainforest, 15.7301 o S, 49.9604 o 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, 15.7279 o S, 49.9751 o E +/- 1.8 km, 32 +/- 25 m, 9 / 11 / 2001, D.C. Lees, DL 01- 27, BMNH (E) # 697188 [DNA voucher], KA 556 [=KA-P 556; DNA extract voucher]; ♀, NE, Andranobe, W. Masoala, 240 m, 15.4 o S, 49.58 o E +/- 1.78 km, 240 m. 3 / 2 / 1991, C. Kremen et al., Genitalia 157 [genitalia voucher]; ♀, NE, Andranobe, W. Masoala, 90 m, 15.6816 o S, 49.9573 o E +/- 1.79 km, 90 m, 14 / 11 / 1991, C. Kremen et al.: Biodiversity voucher; ♀, NE, W. Masoala, Andranobe field station, T 2 S 2 [site], 90 m, 15.6816 o S, 49.9573 o E +/- 1.79 km, 90 m, 14 / 11 / 1991, C. Kremen et al., Biodiversity survey voucher|Gen 158 [genitalia], IA 594 [isotope voucher]; ♂, data as above but: 2 / 11 / 1991, C. Kremen et al., Biodiversity voucher; ♂, NE, Andranobe, W. Masoala, 520 m, 15.65 o S, 49.983 o 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|Gen 156 [genitalia]; ♀, data as above but: 29 / 10 / 1991, C. Kremen et al.: Bio|Gen 159; ♀, NE, Andranobe, W. Masoala, 520 m, 15.65 o S, 49.983 o 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, 15.6733 o S, 49.9885 o E +/- 0.15 km, 500 +/- 50 m, 18 / 2 / 1993, D.C. Lees: DL 93 -0012, IA 593 [isotope voucher]; ♂, NE, Masoala E, Be Dinta, 550 m, 17 /02/ 1993, D.C. Lees: DL 93 -0029, IA 592 [isotope voucher]; ♂, NE, Be Dinta, upper Anaovandrano R., W. Masoala, 500 m, 15.6733 o S, 49.9885 o 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, 15.6733 o S, 49.9885 o 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., W 7 -L 2, Masoala E., 15.7216 o S, 50.1753 o E +/- 0.15 km, 270 m, 18 / 1 / 1994, D.C. Lees; ♀, NE, Manosona, Anaovandrano R., E Masoala, 15.784 o S, 50.2164 o E +/- 0.15 km, 45 m, 1 / 2 / 1994, D.C. Lees; ♂, NE, Masoala E., Antsamanarana R., 275 m, 15.295 o S, 50.227 o E +/- 0.15 km, 275 +/- 50 m, site W 2 L 2-275 M 5 LO, 5 / 12 / 1993, H. Raharitsimba: DL 93 -0028; ♂ [developmental abnormality LHS CuA 1 HWV], NE, E Masoala, Ambavony R. [Ambanivony], W 2 -L 1-125 M [site], disturbed forest, 15.2813 o S, 50.2877 o E +/- 0.72 km, 223 +/- 50 m, 29 / 11 / 1993, C. Kremen: CK 93 -0009; ♀, NE, E Masoala, [Ambavaony R.], bifurcation to 370 m, riparian, W 2 L 1 Saharand. [Sarahandrano] 15.28 o S, 50.2860 o E +/- 1.4 km, 240 +/- 190 m, 25 / 11 / 1993, H. Raharitsimba: TR 46 A; ♂, NE, Masoala E., Antsamanarana R., W 2 L 2 [site], 275 m, 15.2813 o S, 50.2877 o E +/- 0.72 km, 5 LO [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], 15.307 o S, 50.233 o E +/- 0.1 km, 100 m, 29 / 11 / 1993, C. Kremen: CK 30, GEN 141 DL [genitalia]; ♀, data as above but: 15.2968 o S, 50.2271 o E +/- 1.31 km, 294 +/- 225 m, 10 / 12 / 1993, D.C. Lees: E 64 (egg voucher); ♂, NE, Antsamanarana R., Masoala E., 250 m, 15.3 o S, 50.23 o E +/- 0.15 km, 269 m, 12 / 12 / 1993, H. Raharitsimba, GEN 142 DL [genitalia]; ♀, NE, Ambavaony R., E Masoala, [50–380 m], 15.279 o S, 50.288 o E +/- 0.12 km, 150 m, 1 / 12 / 1993, C. Kremen: CK 37, EGG [egg voucher]; ♀, data as above but: 223 m, 29 / 11 / 1993, C. Kremen; ♀, NE, E Masoala, Ambavaony R., riparian, W 2 -L 1-50 M [SITE], 15.2813 o S, 50.2877 o E +/- 0.72 km, 50 m, 2 LO [fruit trap] 28 / 11 / 1993, D.C. Lees: E 3 [egg voucher; “pearly white, smallish, like Ht. pauper ”]; ♂, NE, E Masoala, Ambavaony [Ambanivony], 380 m, W 2 L 1 [SITE] 15.2813 o S, 50.2877 o E +/- 5 km, 30 / 11 / 1993, H. Raharitsimba: TR 61; ♂, NE, Ambavaony R., E Masoala, 125 m, 15.279 o S, 50.288 o E +/- 0.12 km, 125 m, 27 / 11 / 1993, C. Kremen, GEN 124 DL [genitalia]; ♂, NE, Masoala, 1993, C. Kremen: CK 93 -0005, IA 595 [isotope voucher]; ♂ [BMNH], NE, Reserve Speciale d'Ambatovaky, 700 m [16.8 o S, 49.1222 o E, 580 m], 11 / 2 / 1990, M.I. Evans [# 1.16], 254 DL [genitalia].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 67-70, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
Coptotriche alavelona Lees & Stonis, sp. n.
Coptotriche alavelona Lees & Stonis, sp. n. Type material: Holotype: male, “ Madagascar SE: Prov. Fianarantsoa Pref. Ambalavao S/P Ambohimahimasina, Ambondrombe Mt. Near Andranobazaha camp, GPS point “ 22 G 676 ” 21.8817o S 47.2495o E, elev. c. 1700 m. Tropical vegetation by stream, 160 W blended bulb on sheet near ground; vegetation had been disturbed; 23.iii. 2004 c. 23: 36; D.C. Lees. /Genitalia slide No 31441 (BMNH), collection number DL 2028, institute # BMNH (E) # 732343 (voucher specimen and treated genomic DNA from abdomen: plate #MSL 160 H09). Extracted genomic DNA from leg only, plucked in field (BMNH (E) # 678507, plate #MSL079 F06). Holotype deposition: BMNH. Description: Male (Fig. 2). Forewing length: 3.1 mm. Wingspan: 6.7 mm. Head: palpi cream; face smooth and glossy, brownish cream; frontal tuft and collar comprised of glossy grey brown slender lamellar scales; antenna grey brown on upper side, cream on underside, with long piliform sensillae. Thorax and tegulae dark grey brown. Forewing bright ochre (i.e., yellowish, with orange tint) but darkened with fuscous brown scales on basal area and at wing apex; a few fuscous brown scales also along dorsum. Underside of forewing dark brown. Cilia fuscous, paler (grey) at tornus. Hindwing fuscous brown or ochreous brown grey on upper side and underside (hindwing colour depends on angle of view); cilia brown. Legs grey cream, densely covered with fuscous scales on upper side. Abdomen fuscous brown, same colour as tegulae, on sternum whilst light grey covered in sparse brown scales on lateral and ventral parts of tergum. Female unknown. Male genitalia (Figs 3 AB, 4 ABC, 5). Capsule 578–620 µm (including the extended vinculum). Uncus unusually elaborate, with spines on caudal thickening and triangularly shaped pointed ventral lobe (Fig. 4 A). Socii large, as long as uncus, distinctly paired, with tiny setae. Tegumen with deep but asymmetrical anterior excavation. Valva about 390 µm, slender and with median inner lobe in ventral view (Fig. 3 A) but broad and sinuous in lateral view (Fig. 5). Transtilla absent; sublateral processes of valvae situated close to each other but not connected by transverse bar, which is absent. A huge (approx. 50 µm at base, 100 µm at caudal end) membranous anellus-like structure connected with sublateral processes of valvae (Fig. 4 B). Vinculum small but with extremely long (2 / 3 of valva length) rod-like anterior extension (apodeme) (Figs 3 A, 4 C). Aedeagus long (560 µm), folded at the apex, with tiny spines on ovally broadened apical third (Fig. 3 B). Distribution and conservation. The species occurs in high elevation tropical moist forest of SE Madagascar, Ambondrombe Mt. The locality where the species was found is famous in Malagasy legend as the final destination of the spirits of the ancestors (Cousins, 1875), for all 18 ethnic groups. It is thus conserved officially as a historical site of cultural and archaeological significance, rather than a nature reserve (Fig. 6). However, Ambondrombe is further planned as a Site de Conservation within one of the forest corridor zones in the current “Durban Vision” process to triple Madagascar’s protected areas (Rabetaliana 2005). Bionomics. The single adult was found in mid-March. It was attracted to light (to a white sheet using a blended UV/tungsten bulb: Fig. 6 E) but since the surrounding vegetation (Fig. 6 D) had been disturbed, and considering that a usually strictly day-flying micropterigid moth came to the sheet at the same time (Lees, unpublished), it is plausible that the species is not normally active at night. The collection site was beside a river, in riparian tropical montane bamboo forest, with a canopy height> 18 m (Fig. 6 C). Diagnosis: This species differs from all other Coptotriche species in the remarkably elaborate uncus and the extremely extended rod-like vinculum (see also Discussion). DNA extract: in frozen collection at BMNH Entomology, details as above. Etymology: The species name ‘ alavelona ’ means ‘living forest’ in Malagasy and we use it to refer to the remaining Madagascar tropical primary forest with its astonishing biodiversity. The name is composed of two words “ala” (forest) and “velona” (living).Published as part of Lees, David C. & Stonis, Jonas R., 2007, The first record of Tischeriidae (Insecta: Lepidoptera) from Madagascar, with description of Coptotriche alavelona sp. n. and an updated distributional checklist of Afrotropical Tischeriidae, pp. 35-45 in Zootaxa 1645 on pages 38-39, DOI: 10.5281/zenodo.17972
Site-monotonicity properties for reflection positive measures with applications to quantum spin systems
We consider a general statistical mechanics model on a product of local
spaces and prove that, if the corresponding measure is reflection positive,
then several site-monotonicity properties for the two-point function hold. As
an application of such a general theorem, we derive site-monotonicity
properties for the spin-spin correlation of the quantum Heisenberg
antiferromagnet and model, we prove that such spin-spin correlations are
point-wise uniformly positive on vertices with all odd coordinates -- improving
previous positivity results which hold for the Ces\`aro sum -- and we derive
site-monotonicity properties for the probability that a loop connects two
vertices in various random loop models, including the loop representation of
the spin O(N) model, the double-dimer model, the loop O(N) model, lattice
permutations, thus extending the previous results of \textit{Lees and Taggi
(2019)}
Heteropsis tianae Lees & Kremen, sp. nov.
Heteropsis tianae Lees & Kremen, sp. nov. LSID: urn:lsid:zoobank.org:act:E 6864 E 90 -B 8 CF- 4 E 5 F- 99 C 4 -AC 879 E 6 F 70 A Prior references: sp. 49, 69 (Lees, 1997: Torres et al., 2001: 462). Type material., Deposition BMNH: Holotype: ♂ (Fig. 24 A), Madagascar C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 18.45 o S, 47.95 o 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 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, 18.4368 o S, 47.9469 o 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, HM 404245 [DNA barcode voucher], BMNH (E) # 676763 [DNA voucher; cytochrome b], IA 85 [isotope voucher]; ♂, C, Amboasary-an-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 12–14 m canopy forest, flat summit, 18.4497 o S, 47.9404 o 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, IA 122 [isotope voucher]; 2 ♀♀ [BMNH], “Andrangalooka Forest, Madagascar ” [=Andrangoloaka, E. Lac Mantasoa], BMNH (E) # 674766 (Godman-Salvin Coll.; Fig. 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, 18.413 o S, 47.946 o E +/- 0.02 km, 1310 +/- 50 m, 22 / 10 / 2014, D.C. Lees: DL 14 Z-066. Deposition summary: BMNH (HT ♂, 2 PT ♂♂, 3 PT ♀♀), MNHN (PT ♂), ABRI (PT ♂). Type locality. Madagascar C, Amboasary-an-ala, vic. Anjozorobe, 18.45 o S, 47.95 o 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-M 2 (Fig. 24 A– C), and the HWV tends to have the space-CuA 2 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-M 2 and strong tendency to expression of a small ocellus in space-M 3 on the HWD (Fig. 22 AB). Description. Wings: upperside uniform mid brown, FW space-CuA 1 ocellus with an orange ocellus ‘ring’ which is sometimes rather hexagonal in shape and eccentric, being wider at proximad edge. FW space-M 1 ocellus small with narrow orange ring. HW space-CuA 1 ocellus the only one expressed there and somewhat elliptic. Darkish brown diffuse, slightly wavy hair-brush emanating mainly from below vein 1 A+ 2 A 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-CuA 1 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-M 1 ocellus FWV reduced to white pupil with narrow black iris without trace of orange ring. On HWV, space-CuA 1 ocellus slightly elliptic with narrow black iris and small also, without orange ring trace. HWV space-M 1 -M 2 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-M 2. HWV space-CuA 2 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 1 A+ 2 A and 3 A 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 M 2, M 3, CuA 1 and CuA 2 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 A 2. 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, MAD 244; 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: 228 DL, 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. 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 19 th 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. Additional information. DNA divergences: COI- 5 P cluster number BOLD:AAE 4112 (exemplar BMAD 200 - 15, DL 14 Z-051) is about 2.58 % divergent to Ht. oberthueri (cluster number BOLD:ACW 4996, exemplar BMAD 242 - 15, DL 06- 11, RNI Zahamena) and about 4.17 % divergent from ‘sp. 28 ’ (BOLD:AAE 5458), 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- 5 P cluster number BOLD: ACD 8579). Phylogeny/sister species: probably Ht. oberthueri based on the close relationship of the COI- 5 P 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 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. [18.4084 o S, 47.9377 o 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 14 Z-051, BMAD 200 - 15 [DNA barcode voucher]; ♀, C, Anjozorobe, 1400 m, 18.4084 o S, 47.9377 o 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: CK 753, IA 211 [isotope voucher]; ♀, same data but 138 m 14 / 12 / 1994, C. Kremen: CK 754; ♂, C, Anjozorobe, 18.4498 o S, 47.939 o 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, IA 9 [isotope voucher]; ♂, C, Amboasaryan-ala, Tsaracamp Boogie Pilgrim, Anjozorobe, 18.45 o S, 47.95 o 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, 18.45 o S, 47.95 o 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], IA 123 [isotope voucher]; ♂, C, Amboasary-an-ala, Vanjamanitra-Vazimba forest, Anjozorobe, 18.45 o S, 47.95 o 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, 18.45 o S, 47.95 o 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 S 3 ’, 18.4505 o S, 47.9399 o E +/- 0.15 km, 1323 m, 2 / 3 / 2004: 08: 56, D.C. Lees: DL- 4-167, IA 19 [isotope voucher]; ♂, C, Anjozorobe, 18.45 o S, 47.95 o E +/- 2.45 km, 1330 m, 4 / 2 / 2014: 15: 20, D.C. Lees: MAD 240 [pheromone voucher], IA 403 [isotope voucher]; ♂, Anjozorobe, Babakoto trail, c. 1350 m, K. Aduse-Poku, 04/02/ 2014, MAD 243 [pheromone voucher], KA 2059 [=KA-P 2059; DNA extract voucher]; specimen, Anjozorobe, KAP-MAD 1408, KA 1018 [=KA-P 1018; DNA extract number].Published as part of C, Lees David, 2016, Heteropsis (Nymphalidae: Satyrinae: Satyrini: Mycalesina): 19 new species from Madagascar and interim revision, pp. 1-97 in Zootaxa 4118 (1) on pages 79-82, DOI: 10.11646/zootaxa.4118.1.1, http://zenodo.org/record/26459
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