Guérin-Méneville, 1844 : 86 Triodontus nitidulus (Guérin-Méneville) Westwood 1845 : 173 Paulian 1937 : 13 Paulian, 1977 : 1220 Brancsik, 1893 : 224 Review of the Madagascan Orphninae (Coleoptera: Scarabaeidae) with a revision of the genus Triodontus Westwood Frolov, Andrey V. Montreuil, Olivier Akhmetova, Lilia A. Zootaxa 2016 4207 1 1 93 Guerin-Meneville, 1844 Guerin-Meneville 1844 [151,719,1492,1518] Insecta Orphnidae Triodontus Animalia Coleoptera 60 61 Arthropoda species nitidulus     Orphnus nitidulus  Guérin-Méneville, 1844: 86.    Triodontus nitidulus(Guérin-Méneville):  Westwood 1845: 173;  Paulian 1937: 13, 1977: 1220.  Triodontus occidentalis  Paulian, 1977: 1220,  new synonym    Orphnus obsoletus  Brancsik, 1893: 224,  new synonym     Typematerial examined.  Orphnus nitidulus: lectotype(here designated, Fig. 32A), male, “ TypeGuerin Men/  Type/ Coll. R. I. Sc. N. B. Madagascar  Orphnus nitidulusGuer.[...] ♂ Madag/ Coll. R. I. Sc. N. B. Madagascarex coll. Guerinex coll. Monchicourtex coll. de Bonneuil Le Moultvendit /  TYPE/  OrphnusM.Leay MadagascarensisDup. Madag” ( IRSNB).   Orphnus obsoletus: one syntype ( Fig. 32C), male, “Nosy Be” (FMNH).     Triodontus occidentalis: holotype( Fig. 32B), male, “ Berati D. Ananalava P.Griveaud  XII.1960/ MUSEUM PARIS Madagascar/  Triodontus occidentalis n. sp. R. Pauliandet. /  TYPE”;  paratypes, one male and four females with the same locality data as the holotype.    Additional material examined. MADAGASCAR: Antsiranana:three males, 15 females, “ Madagascar-Estdct. Sambava Marojejy Ambinanitelo  500m  XII-58 Raharizonina[ Sambava District, Ambinanitelo]” ( MNHN); six males, four females, “ MadagascarEst 17.XIII.72 Marojejy   300 m.[ Marojejy Range,  300 m, 17.XIII.1972]” ( MNHN); two males, two females, “ Andranovolo/ MadagascarR-on Antalaha  XII.38 Vadon[R-on Antalaha, Andranovolo, 15°01′59″S, 50°07′00″E]” ( MNHN); one male, “ MadagascarEst m-if du Marojejyenv-ons Manantenina   150 m.,  II.1974 A.Peyrieras[ Marojejy Range, environs of Manantenina, 14°28′59″S, 49°49′00″E]” ( MNHN);  21 males, 35 females, “ MadagascarRon Antalaha [ Antalaha District]  XI.35 Vadon” ( MNHN); one male, “ Marojejy  400mf[or]et Ambatomitatao A.Peyrieras  I.1973[ Ambatomitatao Forest, Marojejy Nature Reserve, 14°25′59″S, 49°43′59″E]” ( MNHN); three males, “ MadagascarRon Antalaha  XII.35 Vadon Andapa[ Antalaha District, Andapa, 14°39′00″S, 49°39′00″E]” ( MNHN); eight unsexed specimens, “ Sambirano[ 13°43′00″S, 48°22′00″E] N.O. Madagasc.” ( SMTFD); three females, “ MadagascarAndapa 1.69 [ Andapa, 14°39′00″S, 49°39′00″E]” ( IRSNB);   Toamasina:one male, “ Reservenat III Ambatovositra Andranomalaza  I-57 P Soga[ Ambatovositra, Andranomalaza, " Reservenaturelle III", 17°39′00″S, 48°37′00″E]” ( MNHN); seven males, four females, “ MadagascarEst Moramanga -57 Gruvel[ Moramanga, 18°55′59″S, 48°12′00″E]” ( MNHN);  42 males, 16 females, “ Nandihizana/ MadagascarR-ne Maroantsetra  XII.1938 Vadon! [ Maroantsetra District, Nandihizana, 15°45′00″S, 49°19′00″E]” ( MNHN); one male, “ MadagascarBeanana  XII.45 Vadon! [ Beanana, 15°43′59″S, 49°28′00″E]” ( MNHN); six males, six females, “Museum Paris MadagascarAndevorante A. Mathiaux1900 [ Andevoranto, 18°57′00″S, 49°06′00″E, 1900, A. Mathiauxleg.]” ( MNHN);  52 males, 16 females, “ Madacascar Antsianakaet lac Alaotra2e Trimestre1889 Perrot Freres[Antsianaka and Alaotra Lake, 17°30′00″S, 48°30′00″E]” ( MNHN); three males, five females, “ MadagascarFenerive E.Perrot[ Fenerive, 17°22′00″S, 49°25′00″E]” ( MNHN); two males, three females, “ MadagascarAnalamazoatra / Museum Paris1931 Lasere” ( MNHN); four males, two females, “ Madacascar Antsianaka Perrot Freres2 Semestre1890 [ Antsianaka, 17°30′00″S, 48°30′00″E]” ( MNHN); one male, two females, “ Madagascar-Estdct. Moramanga Fanovana X  I-58 R.Vieu[ Moramanga District, Fanovana, 18°55′00″S, 48°34′00″E]” ( MNHN); one male, “ MadagascarEst distr. Mananara- N. Antanambe Vadonet Peyrieras [ Mananara District, Antanambe, 16°25′59″S, 49°51′00″E]” ( MNHN); two females, “Museum Paris MadagascarProv. de Fenerive Reg. de Soanierana A. Mathiaux1905 [ Districtof Soanierana(= Soanierana Ivongo)]” ( MNHN); one female, “ MadagascarBrickaville [ 18°49′00″S, 49°04′00″E] Cap. Refroigney[?].” ( MNHN); one female, “ Perinet[ 18°55′59″S, 48°25′00″E]” ( MNHN); one male, “Museum Paris MadagascarEnv. de Tamatave A. Raffrey1884 [ Tamatave(= Toamasina), 18°10′00″S, 49°22′59″E]” ( MNHN); one male, “ MadagascarTamatave prov. Moramangaenv. [ Moramangaenvirons, 18°55′59″S, 48°12′00″E] 27–  30.12.1996 Ivo. Jenisleg.” ( MHNG); one male, two females, “ MadagascarTamatave prov. Andasibe( Maromizaha) [ Forest Maromizaha, Anevoka, 18°57′00″S, 48°28′00″E] 19–   20.12.1996 Ivo. Jenisleg.”; one male, “ MadagascarTamatave prov. Andasibe( Maromizaha) [ Forest Maromizaha, Anevoka, 18°57′00″S, 48°28′00″E] 21– 24.11.1995Ivo. Jenis leg.” (MHNG);  14 males, five females, “Rogez MadagascarMus Praha  800–990 m[ Rogez,  800–990 m, 18°47′53″S, 48°36′05″E]” ( NMPC); 71 unsexed specimens, “ Madagascar C. Moramangaenv. [ Moramanga, 18°55′59″S, 48°12′00″E]  10–18.XII.1997 P.Pacholatkoleg.” ( NHMB); 41 unsexed specimens, “ Tamatave[ Toamasina, 18°10′00″S, 49°22′59″E], 1884, A. Raffreyleg., ( SMTFD); one male, “ Maroantsetra[ 15°25′59″S, 49°43′59″E] Madagasc.” ( SMTFD); one male, “ Moramanga[ 18°55′59″S, 48°12′00″E] Madagasc.or.” ( SMTFD); one male, “mittl. O.-Madag. Ambalarond[r]a Dr.Denso 1931.11 [ Ambalarondra, 18°28′06″S, 49°00′04″E,  XI.1931, Densoleg.]” ( SMTFD); eight males, two females, “ Tamatavo[ Tamatave(= Toamasina), 18°10′00″S, 49°22′59″E] Madagascar” ( IRSNB);  Antananarivo: eight males, four females, “Museum Paris MadagascarAntsirabe Waterlot 1916 [ Antsirabe, 19°51′00″S, 47°01′59″E, 1916, Waterlotleg.]” ( MNHN); one male, “ Andrangoloaka  1600 m O.S.O. de Tananarive[ Andrangoloaka,  1600 m, 19°01′00″S, 47°52′00″E]” ( MNHN); three males, 10 females, “ MadagascarTanarive  III.33[ Antananarivo, 18°55′00″S, 47°31′00″E] / Museum Paris1933 J.Vadonet E.Lebis” ( MNHN); two females, “ MadagascarEst Ankadimanga Manjakandriana  XII-57 Jean-Elie[ Ankadimanga, Manjakandriana, 18°58′00″S, 47°45′00″E]” ( MNHN); one male, one female, “ MadagascarAmbatolampy [ 19°22′59″S, 47°25′00″E]” ( MNHN); one female, “ MadagascarTananarive Vadon! [ Antananarive, 18°55′00″S, 47°31′00″E]” ( MNHN); eight males, five females, “ MadagascarLa Mandraka Vadon!  XII-36[  La Mandraka, 18°55′00″S, 47°55′59″E]” ( MNHN); one male, “ Tananarive  III.33[ Antananarive, 18°55′00″S, 47°31′00″E]” ( MNHN); one female, “ Madagascar Antananarivoprov. Manankazoenv. [ 18°09′00″S, 47°12′00″E] 15–  17.12.1996 Ivo. Jenisleg.” ( MHNG); 22 unsexed specimens, “  La Mandraka[ 18°55′00″S, 47°55′59″E]  II.1953 Madagascar” ( NHMB);   Fianarantsoa:eight males, 14 females, “ MadagascarRegion de Mananjary [ 21°13′00″S, 48°19′59″E] A. Mathiaux” ( MNHN); two males, “ Madagascar: Prov. Fianarantsoa, Ranomafana,   600 m.[ Ranomafana,  600 m, 21°18′00″S, 47°30′00″E]  21 October 1988 W.E.Steiner& R. Van Eppscollrs.” ( MNHN); five females, “ Madagascar Centredct. Ambohimahasoaforet Tsarafidy  1450 m  XII-59& I.  60 P.Griveaud [ Ambohimahasoa Disctrict, Ankafina Tsarafidy, forest Tsarafidy,  1450 m, 21°12′00″S, 47°15′00″E]” ( MNHN); two females, “ MadagascarVohilava   60 m. Faraony[ Vohilava Faraony,  60 m, 21°46′00″S, 47°55′00″E]” ( MNHN); four males, two females, “ Ambositra[ 20°31′00″S, 47°15′00″E] Madagascar” ( MNHN); six males, six females, “ Madagascar Fianarantsoaprov. Ranomafanaenv. [ 21°18′00″S, 47°30′00″E] 29.11.–  2.12.1995 Ivo. Jenisleg.” ( MHNG); one female, “MADAGASCAR -   CE2010 Ranomafana NP~  1120 m Vohipararaenv., at light on river bank [ Ranomafana National Park, Vohipararaenvirons, ca.  1120 m, 21°13′35″S, 47°22′11″E];  17.XI.local coll. lgt., BMNH(E) 2010-29” (BMNH); two unsexed specimens, “Ambositra Madagascar  14.11.55 G.Minet[ Ambositra, 20°31′00″S, 47°15′00″E]” ( NHMB); one male, “ MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Vohiparara area,  1170 m, mixed tropical forest.  2–22 January 2001, 21.22644°S/ 47.36979°E. Stop# DHK-01-004 D.H. & K.M. Kavanaugh. R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo, E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Ravelosonleg.” ( CASC); one female, “ MADAGASCAR, Fianarantsoa Province, Ranomafana National Park, Talatakely area,  900 m, mixed tropical forest  2–22 January 2001, 21.25041°S/ 47.41945°E, COL-DHK-2001-001 D.H. & K.M. Kavanaugh. R.L. Brett, E. Elsom, F. Vargas, R. Ranaivosolo, E.F. Randrianirina, N. Rasoamananana, T.J. Ravelomanana, and H.C. Ravelosonleg.“ ( CASC); one male, “ MADAGASCAR, 2007, Ranomafana Nat. Park near Ranomafanavill. [ 21°18′00″S, 47°30′00″E], M. Tryznaleg., 26–31.i., BMNH(E) 2010-29[ 26–31.I.2007, M. Tryzna leg.]” (BMNH); three males, three females, “ MADAGASCAR  CE2011, Ranomafana N. P.[ Ranomafana National Park] 17–21.xi.S 21°15'22.6'' E47°25'17.8''  958 m; at light; M. Tryznaleg. BMNH(E) 2010-29” (BMNH); one male, “Deans Cowan Betsileo Madagascar/ 1881  3000 ft 4000 ft[  3000–4000 ft, 1881]” ( SMTFD); one male, “ Ifanadiana[ 21°18′12″S, 47°38′19″E] Madagascar Mus. Praha” ( NMPC); one male, three females, “ Masagascar Midongy[ Midongy-du-Sud, 23°35′13″S, 47°01′00″E]” ( NMPC);   Mahajanga: 57 males, 14 females, “S te Marie de MadagascarPerrot Freres  X–XII 1896[Sainte Marie de Madagascar, near Marovoay, 16°06′00″S, 46°37′59″E]” ( MNHN); one male, one female, “ Berati D. Ananalava P.Griveaud  XII.1960[ Ananalava District, Beraty, 14°01′00″S, 48°15′00″E]” ( MNHN); one female, “ Randiantsileo/ MadagascarRon Mandritsara  XII-37 Vadon! [ Randiantsileo, Mandritsara District]” ( MNHN); three females, “ Morafenobe[unclear locality] Madag. occ.” ( SMTFD); one male, “Maromandia [ 14°13′00″S, 48°04′59″E] Masagascar Mus.  Praha” ( NMPC);  79 males, 20 females, “Coll. IRScNB MADAGASCAR  Sainte Marie de Madagascar[ 16°06′00″S, 46°37′59″E] 1897 Ex. Coll. Oberthur” ( IRSNB);   Toliara:one male, one female, “ MadagascarOuest Bongolava  1300 m. [ Bongolava, 18°30′00″S, 45°30′00″E] / A.Peyrieras  XII.1974” ( MNHN); one female, “Betroka  XI.31[ Betroka, 23°16′00″S, 46°04′59″E,  XI.1931]” ( MNHN);   Madagascar(no precise locality):two females, “ Madagascar E.Perrot” ( MNHN); six unsexed specimens, “ Madagascar” ( SMTFD).   Diagnosis.  Triodontus nitidulusis most similar to  T. bicavatus.Males of the former species can be separated by having abdominal sternite 8 narrower and more sinuate medially, without oval transverse concavity ( Fig. 31F), and by having the lateral lobes of the parameres without small keel-shaped processes on proximal parts ( Fig. 31E). Females of these species are difficult to separate.   Description. Male.Color of head, pronotum and elytra dark brown; legs, antennae, and underside of the body brown ( Fig. 31A). Frontoclypeus convex anteriorly, rounded laterally, anterior margin crenulate and setose in dorsal view. Eyes relatively small (diameter slightly smaller than the distance between eye and gula in ventral view), incompletely divided by canthus into small, dorsal and large, ventral parts. Frontoclypeus with small tubercles mediad of each eye and in some specimens with a horn in the center of the frontoclypeus. The horn, when the most developed, is about as long as the width of the head, acutely rounded apically, straight to somewhat curved caudally at the apex, slightly rugose on posterior side. Pronotum with a bulge medially, with excavations aside the bulge, and with 2 ridge-shaped tubercles laterally of each excavation. The median bulge has 3 small but distinct tubercles. Lateral margins with wide border appearing somewhat crenulate in dorsal view, with a row of brown setae. Anterior margin with wide, smooth border. Posterior margin with fine border, not crenulate, punctate with narrow longitudinal punctures. Surface of most pronotum smooth, anterior and posterior angles with sparse and coarse punctation. Scutellum rounded apically, visible part about 1/12 length of elytra. Elytra convex, with distinct humeral humps, their maximum width is approximately at the middle. Elytra with 10 distinct striae on disc and with oblique line from base of 6th stria to approximately middle of 8th stria. Stria with small punctures. Intervals with minute punctures, almost smooth. Base of elytra with an irregular row of coarse punctures. Sternite 8 medially deeply sinuate (in ventral view) and somewhat excavated ( Fig. 31F). Parameres ( Fig. 31E) with large outer lobes covering inner lobes (in lateral view). Proximal parts of outer lobes without keel-shaped processes. Internal sac of the aedeagus with 1 strongly sclerotized symmetrical sclerite, somewhat horseshoe shaped, and with 2 similar protuberances of the membrane ( Fig. 31D).  Female.Females ( Fig. 31C) differ from males in having a long, apical spur on the protibiae; absence of the head and pronotum armature including tubercles mediad of each eye; densely punctate dorsal side of the head; and longer abdominal sternites including 6th sternite, which is evenly rounded apically.  Variation.Pronotal and head armature of males is subject to allometric variability from fully developed (as described above) to almost undeveloped (with the males having small frontoclypeal tubercles in the middle and small fossa on the pronotum anteriorly. Body length of examined specimens varied from 6.5–9.0 mm (males) and from 6.0– 7.5 mm(females).   Remarks.The lectotypeof  T. nitiduluswas unknown to Paulian (1977). We found one male specimen originated from the collection of Guérin-Méneville and deposited now in IRSNB. This specimen agrees with the original description of the species. Since Guérin-Méneville mentioned both sexes, evidently he examined more than one specimen although he provided no exact data on this account. To ensure the stability of the nomenclature, we designate this specimen ( Fig. 32A) the lectotypeof  Orphnus nitidulus Guérin-Méneville, 1844.  Brancsik (1893), in the original description of  O. obsoletus, noted that it was similar to  O. nitidulus(=  T. nitidulus). From the short diagnosis of the former, it can be inferred that it differs from the later mostly in the less developed head and pronotum armature. Paulian (1977)considered  O. obsoletusa junior synonym of  T. bicavatus, however he did not discuss this synonymy. Examination of the male syntypeof  O. obsoletusshowed that it has all the diagnostic characters of  T. nitidulus, including the characteristic shape of the abdominal sternite 8, therefore the new synonymy is here proposed.  Paulian (1977)distinguished  T. occidentalisPaulianfrom  T. nitidulusby the absence of the fossette on the abdominal sternite 8 inthe males of the former (as opposed to the presence of a “vague fossette” in the later). However  T. nitidulushas no distinct fossette or fovea on the abdominal sternite 8 (as in  T. bicavatus) but rather its abdominal sternite 8 is apically sinuate and somewhat excavated, that may appear “foveate” depending on the angle of view. We found no noticeable differences between the typeof  T. occidentalisand available material of  T. nitidulusand hence we propose the new synonymy.   Distribution and habitat.This is the most widespread and the only relatively abundant species of the Orphninaein Madagascar. It is currently known from 50 localities throughout the island ( Fig. 31G). Most of the localities are in the eastern Madagascar, in the area occupied by primary and secondary forests. The relatively wide distribution and high abundance of this species may be, at least partly, a result of the secondary adaptation to feeding on cultivated crops. Larvae of  T. nitiduluswere collected in upland rice fields in the Central Plateau ( Randriamanantsoa et al.2010). 1416610247 TYPE, IRSNB Coll. R. I. & Sc. N. B. & Madag & Guerin & Monchicourt & de Bonneuil Le Moult Madagascar Type 60 61 1 1 Men lectotype 1416610243 1960-12 MUSEUM, PARIS, TYPE Berati D. & Ananalava P. Madagascar R. Paulian Madagascar 60 61 2 holotype 1416610203 1960-12 MUSEUM, PARIS, TYPE Berati D. & Ananalava P. Madagascar R. Paulian Madagascar 60 61 2 paratype 1416610240 1958-12 MNHN Raharizonina & Ambinanitelo Madagascar Madagascar-Est 500 Sambava District Sambava Marojejy Ambinanitelo 62 63 15 15 Antsiranana 1416610229 MNHN Madagascar 300 Marojejy Range Marojejy 62 63 300 1416610250 1938-12 MNHN Madagascar Andranovolo -15.033055 Vadon 21 50.116665 Antalaha 62 63 1 1416610249 1974-02 MNHN du Marojejy & A. Peyrieras & Marojejy Range & Manantenina Madagascar 150 -14.483055 Manantenina 21 49.816666 62 63 150 1416610245 [269,1317,1309,1334] 1935-11 MNHN Vadon Madagascar Antalaha District 62 63 56 35 21 1416610235 1973-01 MNHN Ambatomitatao A. & Marojejy Nature Reserve Madagascar 400 -14.433056 Ambatomitatao Forest 21 49.733055 Marojejy 62 63 1 1416610215 1935-12 MNHN Madagascar Vadon Andapa -14.65 Andapa 21 49.65 Antalaha District 62 63 1 1416610202 SMTFD N. O. Madagasc. Madagascar -13.716666 Sambirano 21 48.366665 62 63 1 1416610237 IRSNB Madagascar -14.65 Andapa 21 49.65 62 63 1 1416610251 1957-01 MNHN P Soga Madagascar Reserve -17.65 Ambatovositra 21 48.616665 Ambatovositra Andranomalaza 62 63 1 Toamasina 1416610238 MNHN Madagascar -18.933054 Moramanga 21 48.2 Gruvel 62 63 1 1416610234 1938-12 MNHN Madagascar Nandihizana -15.75 Vadon 21 49.316666 Maroantsetra 62 63 58 16 42 1416610230 1945-12 MNHN Madagascar -15.733055 Beanana 21 49.466667 Vadon 62 63 1 1416610241 MNHN A. Mathiaux Madagascar -18.95 Andevoranto 21 49.1 62 63 1 1416610224 MNHN Madagascar Madacascar Antsianaka -17.5 Trimestre 21 48.5 Alaotra 62 63 68 16 52 1416610254 MNHN E. Perrot Madagascar -17.366667 Fenerive 21 49.416668 62 63 1 1416610246 MNHN Madagascar Lasere Museum Paris 62 63 1 1416610258 MNHN Madagascar Madacascar Antsianaka Perrot Freres -17.5 Antsianaka 21 48.5 Semestre 62 63 1 1416610263 1958-01 MNHN Moramanga Fanovana X & R. Vieu Madagascar Madagascar-Est -18.916666 Fanovana 21 48.566666 Moramanga District 62 63 1 1416610226 MNHN N. Antanambe Vadon Madagascar Mananara -16.433054 Antanambe 21 49.85 Mananara District 62 63 1 1416610259 MNHN A. Mathiaux Madagascar District 63 64 Soanierana Ivongo Soanierana 62 63 1 1416610244 MNHN Cap. Refroigney Madagascar -18.816668 Madagascar 21 49.066666 63 64 1 1416610274 [660,1316,188,213] MNHN Madagascar -18.933054 Perinet 21 48.416668 63 64 1 1416610264 MNHN A. Raffrey Madagascar -18.166666 Tamatave 21 49.383053 63 64 1 Toamasina 1416610233 1996-12-30 MHNG Ivo. Jenis Madagascar -18.933054 Moramanga 21 48.2 Moramanga 63 64 1 1416610267 Madagascar Andasibe -18.95 Forest Maromizaha 21 48.466667 Maromizaha 63 64 1 1416610257 1996-12-20 Ivo. Jenis Madagascar Andasibe -18.95 Forest Maromizaha 21 48.466667 Maromizaha 63 64 1 1416610275 NMPC Madagascar 895 -18.798056 Rogez 21 48.601387 63 64 14 14 Praha 1416610272 1997-12-10 1997-12-18 1997-12-10 NHMB C. Moramanga & P. Pacholatko Madagascar -18.933054 Moramanga 21 48.2 63 64 1 1416610270 SMTFD A. Raffrey Madagascar -18.166666 Tamatave 21 49.383053 63 64 1 Toamasina 1416610260 SMTFD Madagasc. Madagascar -15.433056 Maroantsetra 21 49.733055 63 64 1 Toamasina 1416610242 SMTFD Madagasc. Madagascar -18.933054 Moramanga 21 48.2 63 64 1 Toamasina 1416610231 1931-11 SMTFD Denso Madagascar -18.468334 Ambalarondra 21 49.00111 63 64 1 Toamasina 1416610262 IRSNB Madagascar -18.166666 Tamatave 21 49.383053 Tamatavo 63 64 1 Toamasina 1416610276 MNHN Waterlot Madagascar -19.85 Antsirabe 21 47.033054 63 64 1 Antananarivo 1416610268 MNHN O. S. O. de Tananarive Madagascar 1600 -19.016666 Andrangoloaka 21 47.866665 Andrangoloaka 63 64 1 Antananarivo 1416610261 1933-03 MNHN Museum Paris & J. Vadon & E. Lebis Madagascar -18.916666 Antananarivo 21 47.516666 63 64 10 10 Antananarivo 1416610255 1957-12 MNHN Madagascar Jean-Elie -18.966667 Manjakandriana 21 47.75 Ankadimanga 63 64 1 1416610266 MNHN Madagascar -19.383055 Madagascar 21 47.416668 63 64 1 1416610248 MNHN Madagascar -18.916666 Antananarive 21 47.516666 63 64 1 1416610280 1936-12 MNHN La Mandraka Madagascar -18.916666 La Mandraka 21 47.933056 63 64 1 1416610271 1933-03 MNHN Madagascar -18.916666 Antananarive 21 47.516666 Tananarive 63 64 1 1416610282 1996-12-17 MHNG Ivo. Jenis Madagascar -18.15 Manankazo 21 47.2 63 64 1 Antananarivo 1416610289 1953-02 NHMB La Mandraka Madagascar -18.916666 La Mandraka 21 47.933056 63 64 1 1416610253 MNHN A. Mathiaux Madagascar -21.216667 Fianarantsoa 21 48.333054 63 64 14 14 Fianarantsoa 1416610285 1988-10-21 MNHN W. E. Steiner & R. Van Epps Madagascar 600 -21.3 Ranomafana 21 47.5 Ranomafana 63 64 600 Fianarantsoa 1416610269 1959-12 Madagascar 1450 Tsarafidy Ambohimahasoa 63 64 1 Centre 1416610298 MNHN Madagascar Ambohimahasoa Disctrict 1450 -21.2 Tsarafidy 21 47.25 Ankafina Tsarafidy 63 64 60 Centre 1416610288 MNHN Madagascar 60 -21.766666 Vohilava Faraony 21 47.916668 Faraony 63 64 60 1416610256 [433,1432,1268,1293] MNHN Madagascar -20.516666 Ambositra 21 47.25 63 64 1 1416610292 1995-02-12 MHNG Ivo. Jenis Madagascar -21.3 Ranomafana 21 47.5 63 64 1 Fianarantsoa 1416610283 17. XI. CE Ranomafana NP Madagascar Vohiparara 1120 -21.226389 Vohiparara 21 47.36972 Ranomafana National Park 63 64 CE 2010 1 Fianarantsoa 1416610301 55-11-14 NHMB G. Minet Madagascar -20.516666 Ambositra 21 47.25 63 64 1 1416610299 2001-01-02 2001-01-22 2001-01-02 CASC D. H. & K. M. Kavanaugh. & R. L. Brett & E. Elsom & F. Vargas & R. Ranaivosolo & E. F. Randrianirina & N. Rasoamananana & T. J. Ravelomanana & H. C. Raveloson Madagascar 1170 -21.22644 Vohiparara area 1 47.36979 Ranomafana National Park 63 64 1 Fianarantsoa Province 1416610291 2001-01-02 2001-01-22 2001-01-02 CASC D. H. & K. M. Kavanaugh. & R. L. Brett & E. Elsom & F. Vargas & R. Ranaivosolo & E. F. Randrianirina & N. Rasoamananana & T. J. Ravelomanana & H. C. Raveloson Madagascar 900 -21.25041 Talatakely area 1 47.41945 Ranomafana National Park 63 64 1 Fianarantsoa Province 1416610284 M. Tryzna Madagascar -21.3 Ranomafana 21 47.5 63 64 1 1416610265 CE Ranomafana N. P. & M. Tryzna Madagascar 958 Ranomafana National Park 21.256277 63 64 CE 2011 1 1416610252 SMTFD Madagascar 0 Madagascar 63 64 1 1416610287 NMPC Mus. Madagascar -21.303333 Ifanadiana 21 47.638615 63 64 1 Praha 1416610296 NMPC Madagascar -23.586945 Midongy-du-Sud 21 47.016666 Masagascar Midongy 63 64 1 Praha 1416610295 1896-10 1896-12-31 1896-10 MNHN Madagascar -16.1 Marovoay 21 46.633053 63 64 71 14 57 Mahajanga 1416610286 1960-12 MNHN Berati D. & Ananalava P. Madagascar 64 65 -14.016666 Beraty 21 48.25 Ananalava District 63 64 1 Mahajanga 1416610279 1937-12 MNHN Vadon & Randiantsileo Madagascar Mandritsara District Randiantsileo 64 65 1 1416610290 SMTFD Madagascar -14.216666 Morafenobe 21 48.083054 64 65 1 1416610273 [1084,1273,223,248] NMPC Madagascar Praha 64 65 1 Praha 1416610304 IRSNB Sainte Marie de Madagascar & Ex. Coll. Oberthur Madagascar -16.1 Sainte Marie de Madagascar 21 46.633053 64 65 99 20 79 1416610293 1974-12 MNHN A. Peyrieras Madagascar 1300 -18.5 Bongolava 21 45.5 64 65 1 Toliara 1416610305 1931-11 MNHN Madagascar -23.266666 Betroka 21 46.083054 64 65 1 Toliara 1416610310 MNHN E. Perrot Madagascar Madagascar 64 65 1 1416610278 [270,822,404,429] SMTFD Madagascar Madagascar 64 65 1