Metaleptobasis bicornis ( Selys, 1877 )

Ellenrieder, Natalia Von, 2013, <p> <strong> A revision of <em> Metaleptobasis </ em> Calvert (Odonata: Coenagrionidae) with seven synonymies and the description of eighteen new species from South America </ strong> </ p>, Zootaxa 3738 (1), pp. 1-155 : 18-22

Metaleptobasis bicornis ( Selys, 1877)

Figs. 1b ; 2a ; 3b ; 4b ; 5b ; 8b ; 9b ; 10b ; 11b ; 12b ; 14c

Leptobasis bicornis Selys, 1877: 103–104 [9–10 reprint] (description of ♀).

Metaleptobasis bicornis: Calvert (1907: 386 ; suggestion of possible inclusion in genus);— Williamson (1915: 608; mention);— Cumming (1954: 29, 30; inclusion in key based on ♂ of uncertain identity, mention);— Rácenis (1955: 15; diagnosis from M. fernandezi );— Santos (1956: 384; diagnosis from M. selysi );— Daigle (2003: 373; mention);— Lencioni (2006: 164, fig. 102a; mention from Brazil, illustration of ♀ posterior lobe of pronotum and mesanepisternal horns, by NVE);—von Ellenrieder & Garrison (2007: 24, figs. 11a, b; illustration of ♀ posterior lobe of pronotum and mesanepisternal horns);— Heckman (2008: 399, figs. 3.1.462; inclusion in key, illustration of ♀ posterior lobe of pronotum and mesanepisternal horns, by NVE);— Garrison & von Ellenrieder (2009: 48; mention);— Garrison et al. (2010: 285, figs. 1827, 1831; illustration of ♀ posterior lobe of pronotum and mesanepisternal horns).

Metaleptobasis mauritia Williamson, 1915: 603–607 , figs. 9–12 (description of ♂, illustrations mesanepisternal horns and S10);— Cumming (1954; 29, 31; inclusion in key to males, proposed grouping);— Santos (1956; 384; comparison with M. selysi );—Rácenis (1954: 5; mention);— Michalski (1988: 45–46; mention from Trinidad, inclusion in key);—De Marmels (1990: 337; mention);— De Marmels (1992b: 63–64, figs. 41– 45; mention, illustrations of S10 and mesanepisternal horns of ♂ from Trinidad, posterior lobe of pronotum and mesanepisternal horns of andromorphic ♀ from Venezuela);— Heckman (2008: 404–405, fig. 3.1.469; in key, reproduction of Williamson 1915 and De Marmels 1992b illustrations);— Garrison & von Ellenrieder (2009: 49; mention);— Garrison et al. (2010: 284; mention);— Tennessen (2012: 95; comparison with M. gibbosa );—von Ellenrieder (2011: 57–58, 73; mention).— New synonymy.

Metaleptobasis fernandezi nec Rácenis, 1955 — De Marmels (1992b: 63–64, figs. 37–40; comparison with ♂ from Trinidad, illustrations of S10 and mesanepisternal horns of ♂ from Venezuela, posterior lobe of pronotum of ♀ from Venezuela);— Belle (2002: 3; mention).— Misidentifications.

Metaleptobasis tetragena nec Calvert, 1948 —Belle (2002: 3; mention).— Misidentification.

Types. Holotype ♀ *: ‘Amazonas’, Bates leg. [ RMNH].

Specimens examined. Total: 115 ♂, 100 ♀: 1 ♀ (he) holotype; TRINIDAD, St. Andrew Co .: 1 ♂ holotype M. mauritia, Cumuto , swamp {10°35'N, 61°12'W, 57 m}, 10 iii 1912, E.B. Williamson, L.A. Williamson & B.J. Rainey leg. [ UMMZ] ; 1 ♂ paratype M. mauritia same data [ UMMZ] ; 1 ♂ Valencia , forest {10°39'N, 61°13'W}, 10 iv 1965, TWD leg. [ DRP] ; 1 ♂, same but [ RWG] ; 1 ♀ (he), same but SE of Valencia (10°38'24''N, 61°11'24''W), TWD leg. [ TWD] ; 2 ♂, forest cut and forest 2 mi SE of Valencia on Eastern Main road (10°37'26''N, 61°10'55''W, 42 m), 7 iv 1980, J.A. Garrison & RWG leg. [ RWG] ; 1 ♂, stream 7 mi E Valencia , 7 iv 1965, TWD leg. [ FSCA] ; 1 ♂, 1 ♀ (an), same but stream, 16 iv 1965 [ TWD] ; 1 ♂, 3 mi N of Sangre Grande {10°34'N, 61°8'W, 44 m}, 9 iv 1965, TWD leg. [ RMNH]; GUYANA, Cuyuni-Mazaruni Region : 1 ♂, Oko River {6°25'N, 59°5'W, 260 m}, 21 vi 1936, N.A. Weber leg. [ UMMZ] ; 1 ♀ (he), Essequibo River, Picrewane Island, above Wineperu {6°11'N, 58°35'W, 6 m}, 9–15 iii 1969, W.D. Duckworth & R.E. Dietz leg. [ USNM]; Potaro-Siparuni Region : 1 ♂ paratype M. mauritia , right bank of Potaro River near Tumatumari {5°17'N, 58°59', 20 m}, 9 ii 1912, E.B. Williamson, L.A. Williamson & B.J. Rainey leg. [ UMMZ]; Upper Takutu-Upper Essequibo Region : 1 ♂, Kumu River, ca 20 km S Lethem, Kanuku Mountains (3°15'93''N, 59°43'47''W), 30 iv 1995, O.S. Flint, Jr. [ USNM]; SURINAM : 1 ♂, Sector 0, forest, 4 xi 1942, D.C. Geijskes leg. [ RMNH] ; 1 ♂, Coronieweg marker 189, forest trail S km 0.3, forest on sand ridge parallel to coast, 23 xii 1948, D.C. Geijskes leg. [ RMNH]; Commewijne Dis .: 1 ♂, 1 ♀ (he), Tapoeripa Channel {5°59'N, 54°45'W}, 19 iv 1972, D.C. Geijskes leg. [ RMNH] ; 2 ♂, 6 ♀ (an), 2 ♀ (he), Commetewane Creek , E–W connector, forest swamp {5°45'N, 54°45'W, 10 m}, 17 ix 1958, D.C. Geijskes leg. [ RMNH] ; 1 ♀ (he), same but 20 x 1959, J. Belle leg. [ RMNH] ; 1 ♀ (an), Paulus Creek {5°45'N, 55°6'W, 10 m}, 29 v 1960, J. Belle leg. [ RMNH]; Nickerie Dis .: 1 ♀ (an), Nani Creek {5°53'N, 57°5'W}, 21 ix 1941, D.C. Geijskes leg. [ RMNH] ; 1 ♂, same but forest near bird colony [ RMNH] ; 1 ♀ (an), same but forest near small village [ RMNH]; Marowijne Dis .: 2 ♂ 1 ♀ (an), Wia Wia , km 10.3–12.1 {5°53'N, 58°29'W, 0 m}, 25 xi 1948, D.C. Geijskes leg. [ RMNH] ; 1 ♂ 1 ♀ (an) (in copula), same but km 11–12, forest [ RMNH] ; 1 ♀ (an), same but 27 xi 1948 [ RMNH] ; 1 ♀ (he), same but km 4.6, forest on sand ridge parallel to the coast, 19 xi 1948 [ RMNH] ; 1 ♀ (an), same but Perits , km 23.8–24.5, 20 xi 1948 [ RMNH] ; 1 ♂, Galibi {5°45'N, 54°0'W, 0 m}, 30 vii 1959, van Doesburg leg. [ RMNH] ; 3 ♀ (an), 1 ♀ (he), Langamankondre {5°42'N, 54°2'W, 0 m}, 23 x 1963 B. Malkin leg. [ RMNH] ; 1 ♀ (an), Marowijne River, Christiaankondre , forest trail {5°42'N, 54°0'W}, 3 x 1963, B. Malkin leg. [ RMNH] ; 1 ♂, Moengo, Boven {5°37'N, 54°24'W, 25 m}, 17 v 1927, E.B. Williamson leg. [ FSCA] ; 1 ♀ (an), Moengo , boesman hill {5°37'N, 54°24'W, 0 m}, 10 iv 1939 D.C. Geijskes leg. [ RMNH] ; 4 ♂, same but forest, 18 ix 1961 [ RMNH] ; 1 ♀ (he), Thorarica , forest swamp {5°34'N, 54°14'W, 33 m}, 6 ii 1957, D.C. Geijskes [ RMNH] ; 1 ♀ (an), Moengotapoe, Djai Creek , forest trail 1 km 8.6, forest {5°34'N, 54°15'W, 20 m}, 7 x 1948, D.C. Geijskes leg. [ RMNH] ; 1 ♀ (he), forest trail N of Moengotapoe , km 17.2 {5°34'N, 54°14'W, 33 m}, 15 x 1948, D.C. Geijskes leg. [ RMNH] ; 1 ♂, same but 3e Kamp , forest trail km 15, forest on sand ridge parallel to the coast [ RMNH] ; 2 ♂, same but 3 rd camp, forest on sand ridge parallel to the coast, km 16.1–16.3 [ RMNH] ; 2 ♀ (an), same but, 3 rd camp, forest trail km 17.1, forest swamp [ RMNH] ; 1 ♂, 1 ♀ (an), same but km 10.3–10.7, forest on sand ridge parallel to the coast, 13 x 1948 [ RMNH] ; 2 ♀ (an), 1 ♀ (he), same but forest trail N of 3 rd camp, forest on sand ridge parallel to the coast, 14 x 1948 [ RMNH] ; 1 ♀ (an), same but forest trail km 15.5 [ RMNH] ; 1 ♂, same but forest trail km 14.2–14.7, forest on sand ridge parallel to the coast [ RMNH] ; 1 ♂, same but km 15.3, forest on sand ridge parallel to the coast, 15 x 1948, D.C. Geijskes leg. [ RMNH] ; 1 ♂, same but km 15.7–14.9 [ RMNH] ; 2 ♀ (an), same but, 3rd Camp , forest trail km 15.5, forest on sand ridge parallel to the coast, 19 x 1948 [ RMNH] ; 1 ♀ (an), same but 3rd camp, forest trail km 29.6, forest on sand ridge parallel to the coast, 25 xi 1948 [ RMNH] ; 2 ♂, 1 ♀ (an), same but 3 rd camp, forest trail km 10.3–10.7, forest on sand ridge parallel to the coast [ RMNH] ; 1 ♀ (an), same but, 3rd Camp , forest trail km 13, forest on sand ridge parallel to the coast, 6 x 1948, D.C. Geijskes leg. [ RMNH] ; 2 ♀ (he), same but 3rd Camp , forest trail km 14.6–14.9, forest on sand ridge parallel to the coast, 18 x 1948 [ RMNH] ; 1 ♂, 1 ♀ (an), same but 3 rd camp, forest trail km 15–15.7, forest on sand ridge parallel to the coast, 13 x 1948 [ RMNH]; Suriname Dis .: 1 ♀ (an), De Morgenstond Estate {5°52'N, 55°6'W, 2 m}, 25 v 1958, J. Belle leg. [ RMNH] ; 1 ♂, 1 ♀ (an), Charlesburg {5°50'N, 55°10'W, 10 m}, 30 x 1938, D.C. Geijskes leg. [ FSCA] ; 1 ♀ (an) same but 29 i 1940 [ RMNH] ; 2 ♂, same but 2 ii 1940 [ RMNH] ; 1 ♂, same but 5 ii 1940 [ RMNH] ; 2 ♂, same but 12 vi 1958, J. Belle leg. [ RMNH] ; 1 ♂, same but 16 x 1958 [ RMNH] ; 1 ♂, 1 ♀ (an), same but 24 vii 1959 [ RMNH] ; 1 ♂, same but 4 ix 1959 [ RMNH] ; 4 ♂, Cultuurtui , in bamboo {5°50'N, 55°9'W, 9 m}, 6 x 1938, D.C. Geijskes leg. [ RMNH] ; 1 ♂, Paramaribo {5°49'N, 55°10'W, 6 m}, 27 xi 1920, D.C. Geijskes leg. [ RMNH] ; 1 ♂, same but Combe {5°49'N, 55°8'W, 5 m}, 2 vii 1946, D.C. Geijskes leg. [ RMNH] ; 1 ♀ (an), same but Zorg en Hoop {5°48'N, 55°11'W, 3 m}, 21 vi 1958, J. Belle leg. [ RMNH] ; 1 ♂, Lelydorp , forest km 25 {5°42′N, 55°14′W}, 25 x 1938, D.C. Geijskes [ RMNH]; Para Dis .: 1 ♀ (an), Carolina Creek {5°24'N, 55°11'W, 20 m}, 5 ii 1960, J. Belle leg. [ RMNH] ; 1 ♀ (an), same but 1 v 1962, von Doesburg leg. [ RMNH]; Brokopondo Dis .: 4 ♂, Brokopondo {5°4'N, 54°58'W, 72 m}, 1–22 iii 1966 [ RMNH] ; 2 ♂, same but 22 iii 1966, G.F. Mees leg. [ RMNH] ; 1 ♂, 1 ♀ (he) (in copula), same but dry creek bed deep in forest, 30 iii 1966, Balling leg. [ RMNH] ; 1 ♀ (he), same data [ RMNH] ; 2 ♂, Stoelmanseiland , forest tree over river {5°4'N, 54°58'W, 10 m}, 3 iv 1963, J. Belle leg. [ RMNH] ; 1 ♂, Kabel, Suriname River {4°55'N, 55°8'W, 41 m}, 16 v 1961, J. Belle leg. [ RMNH] ; 1 ♀ (he), opposite side of Kabelstation , in forest {4°54'N, 55°8'W}, 24 ix 1938, D.C. Geijskes leg. [ RMNH] ; 3 ♀ (he), Gansee, Suriname River {4°49'N, 55°4'W, 23 m}, 27 xi 1957, J. Belle leg. [ RMNH] ; 1 ♂, Bakra Opposton, Suriname River {4°44'N, 55°6'W, 23 m}, 31 vii 1942, L. Schmidt leg. [ RMNH]; Sipaliwine Dis .: 1 ♂, Coppename, Fossi Bergi , forest by the river {5°17'N, 56°23'W}, 8 vii 1943, D.C. Geijskes leg. [ RMNH] ; 1 ♂, Nickerie River, Lombok Val {4°52'N, 57°0'W}, 8 ii 1971, D.C. Geijskes leg. [ RMNH] ; 8 ♂, 10 ♀ (an), 3 ♀ (he), Kabalebo River {4°25'N, 57°13'W}, 31 viii 1963, J. Belle leg. [ RMNH] ; 2 ♂, 3 ♀ (an), 2 ♀ (he), same but 16 viii 1964 [ RMNH] ; 1 ♂, same but 17 viii 1964 [ RMNH] ; 2 ♀ (an), same but 18 viii 1964 [ RMNH] ; 2 ♂, same but 19 viii 1964 [ RMNH] ; 1 ♂, same but 20 viii 1964 [ RMNH] ; 1 ♀ (an), same but 31 viii 1964 [ RMNH] ; 5 ♂, same but Aranavero valley , in the shade in forest, 6 iv 1971, D.C. Geijskes leg. [ RMNH] ; 1 ♀ (an), Marowijne river, forest near village by river {4°21'N, 54°26'W, 55 m}, 30 viii 1939 D.C. Geijskes leg. [ RMNH] ; 4 ♂, 1 ♀ (an), Tapanahoni, Tapatosso , 5 iv 1958, J. Belle leg. [ RMNH] ; 1 ♀ (an), Tapanahoni, Gran Alosoela , 6 iv 1958, J. Belle leg. [ RMNH] ; 1 ♂, Tapanahoni, Malobbi {4°10'N, 55°26’W}, forest on island, 22 viii 1953, D.C. Geijskes leg. [ RMNH] ; 1 ♂, Wilhelmina Mountain range, Lucie Camp , in forest {3°26'N, 57°13'W, 190m}, 12 vii 1963, Ligorie leg. [ RMNH] ; 2 ♂, 1 ♀ (he), same but 8 viii 1963 [ RMNH] ; 7 ♂, 2 ♀ (an), 5 ♀ (he), same but 10 viii 1963, S. Ligorie leg. [ RMNH] ; 4 ♂, same but 19 ix 1960 / 19 i 1961, J. Beatty leg. [ RMNH] ; 1 ♂, 1 ♀ (he), same but forest swamp, i 1961 [ RMNH] ; 1 ♀ (an), Kayser airstrip, small creek {3°5'N, 56°28'W, 410 m}, 28 vi 1963, S. Ligorie leg. [ RMNH] ; 2 ♂, Lawa river, Anapaike Village , N side in forest along the river {3°25'N, 54°2'W, 84 m}, 20 xi 1963, S. Ligorie leg. [ RMNH] ; 1 ♂, same but forest, 26 xi 1963, B. Malkin leg. [ RMNH] ; 1 ♂, Sipaliwini {2°6'N, 56°2'W}, 14 v 1972, D.C. Geijskes leg. [ RMNH] ; 1 ♂, 1 ♀ (an), same but Pina palm forest, 18 ii 1961 [ RMNH] ; 2 ♂, same but 28 i/ 2 ii 1966 [ RMNH] ; 2 ♂, same but small pool in forest near airport, 12 ii 1961; 1 ♂, same but 15 ii 1961; 1 ♂, same but small creek near airport, 17 ii 1961 [ RMNH] ; 7 ♂, 5 ♀ (he), Coeroeni Island {2°51'N, 56°33'W, 290 m}, 12 viii / 7 x 1959, D.C. Geijskes leg. [ RMNH] ; 1 ♂, Werehpai , forest stream (2°21'54''N, 56°41'45''W, 258 m), 5 ix 2010, NVE leg. [ NVE] ; 1 ♂, 1 ♀ (he), half way between West Paroe and Aleensoe , swamp by small creek, 25 iv 1952 [ RMNH] ; 1 ♂, Grensgebergte , seepage creek, 7 v 1952 [ RMNH] ; 1 ♂, Paroe , 1 i 1941, Schmidt leg. [ RMNH]; FRENCH GUIANA : 5 ♂, 2 ♀ (an), Cayenne , woods 5 km N of Matoury on N2 (4°30'36''N, 52°12'36''W), 14 ii 1988, RWG leg. [ RWG]; BRAZIL, Ceará State : 1 ♂, 1 ♀ (he), Maranhão, Rio Maracaguare , ca. 80 km E of Caninde {4°21'S, 39°19'W}, xii 1964, leg. B. Malkin [ RMNH]; Pará State : 1 ♂, Pará, Floresta Nacional de Carajás, Parauapebas , Buritizal I (6°5'13''S, 50°8'50''W, 682 m), 23 ix 2007, N. Ferreira, Jr. & V. Alecrim leg. [ DZRJ] ; 1 ♂, same but (6°4'57''S, 50°8'5''W, 682 m), 25 iii 2006, N. Ferreira, Jr. leg. [ DZRJ] ; 1 ♂, 1 ♀ (an), 1 ♀ (he), Belem, swamp in forest, Instituto Agronómico do Norte {1°27'S, 48°30'W}, 7–10 xii 1954 [ RMNH]; Roraima State : 1 ♀ (an), Ilha de Maracá, Reserve (4°24'50''N, 61°39'58''W), 12 xi 2007, U. Neiss leg. [ INPA] ; 1 ♂, Ilha de Maracá, Rio Uraricoera , fogging {3°21'50''N, 61°26'0''W}, 21–30 xi 1987, J.A. Rafael leg [ INPA] .

Characterization. Head. Labrum mostly pale; black on dorsum of head limited; postocular lobes rounded ( Fig. 1b ). Thorax. Pronotum ( Figs. 2a ; 4b ; 5b ) anterior lobe with posterior margin of medio-lateral portions with a marginal crest (cr.), and with lateral margins projected postero-ventrally forming a smooth L-shaped ridge (l-v.p.); anterior portion of propleuron with an anterior rounded tubercle (t.) opposite the ridge; anterior and middle lobes of pronotum separated dorso-laterally by a groove; anterior margin of middle lobe of pronotum smooth; posterior lobe trilobed with medial lobe smoothly convex in male, in female trilobed with medial lobe smoothly convex or bilobed, or sinuous, or bilobed with a central shallow u-shaped incision, with a medial dark spot extensive to absent; lateral margins of pronotum posterior lobe relatively long, free and rounded; mesanepisternal horns with bases joined by a low ridge to separated, horns well developed in males, as long as about 1.5–2.5 times mesostigmal plate width, oriented antero-dorsally at an angle of about 35°–45° with dorsum, about parallel sided to slightly diverging, medium in thickness, with rounded tips; in andromorphic females as in males ( Figs. 5b xiii–xiv), in heteromorphic females represented by bases only and as long as about 0.30–0.50 of mesostigmal plate width ( Figs. 5b viii–xii) or represented by bases and a free portion directed laterally, about as long as width of mesostigmal plate ( Figs. 5b vi–vii); mid-dorsal dark stripe slightly wider than 0.33 to slightly narrower than 0.50 of mesanepisternal width, about parallel sided ( Figs. 2a ; 3b ); Pt short rectangular, with anterior and posterior sides slightly longer than distal side, to squarish, with posterior side about equal to distal side. Abdomen. Male genital lobe ( Fig. 8b ) short, less than 0.50 of anterior hamule height; male genital lobe smoothly curved; male posterior hamule digit-like and small, with at most only tip surpassing ventral margin of genital fossa in lateral view; curvature of basal segment of genital ligula marked by a slight concave depression (s.d.); genital ligula distal segment sub-rectangular ( Fig. 8b ), apex slightly concave, with a narrow ectal fold (e.f.); posterior margin of female S8 sternum ( Fig. 9b ) smooth, lacking any denticles, spines, or processes; distal end of ovipositor reaching level of tip of paraproct to slightly longer than cercus; medial portion of male S10 postero-dorsal margin ( Figs. 10b ; 11b ; 12b ) projected posteriorly and bilobate, with rounded lobes separated by a small u-shaped incision or apressed and separated by a slit, with a dorsal prominence surrounding incision or slit; male cercus in dorsal view ( Fig. 10b ) arched medially with a uniform curvature, with distal portion slightly widened sub-apically, depressed dorsoventrally, with tip pointed medio-ventrally; ratio of male cercus length to S10 maximum length in lateral view about 1.0–1.5; ratio of male cercus length to paraproct length in lateral view about 0.66–0.87; distal half of male paraproct in lateral view ( Fig. 12b ) of uniform width, tip bent medio-ventrally, with a ridge on medial surface ending on an apical tooth.

Dimensions. Males (n 10): Hw 22.3 ± 0.8 [21.2–23.4]; abdomen 37.5 ± 1.9 [35.2–40.8]; total length 44.9 ± 2.1 [42.1–48.3]. Females (n 10): Hw 23.7 ± 0.7 [22.3–25]; abdomen 36.6 ± 1.6 [35.3–39.4]; total length 44.1 ± 1.6 [41.4–47].

Diagnosis. Males and females of M. bicornis differ from all species except for M. prostrata , by the anterior lobe of pronotum with lateral margins projected postero-ventrally forming an L-shaped ridge (l-v.p.), ending opposite an anterior tubercle (t.) on propleuron ( Figs. 2a ; 4b ; 5b ). Metaleptobasis bicornis differs from M. prostrata by distal portion of male cercus widened sub-apically ( Fig. 10b ) and depressed dorso-ventrally ( Fig. 12b ; gradually narrowing distally and sub-cylindrical in M. prostrata , Figs. 10x ; 12x ), mesanepisternal horns oriented antero-dorsally at an angle of about 45° with dorsum in males and andromorphic females, or oriented laterally or represented by bases only in heteromorphic females (oriented antero-laterally at an angle of about 15° with dorsum in males and females of M. prostrata ), and medio-lateral portions of anterior lobe of pronotum with posterior margin with a marginal crest (cr., Fig. 4b ; lacking a crest in M. prostrata , Fig. 4x ).

Sixty percent of examined females of M. bicornis are andromorphic, sharing both a trilobed pronotum posterior lobe and long mesanepisternal horns directed antero-dorsally with males ( Fig. 5b xiii). The remaining 40% are heteromorphic, presenting various combinations of shape of pronotum posterior lobe and development and orientation of horns: 26% differ from males by both posterior lobe of pronotum shape and mesanepisternal horn development, with pronotum posterior lobe sinuous and free portion of horns directed laterally (2%; Figs. 5b vi–vii) or pronotum posterior lobe sinuous and horns represented by bases only (24%; Figs. 5b viii, x–xi); 4% differ only by posterior lobe of pronotum indented, with long horns ( Fig. 5b xiv), and 10% only by horn development, with vestigial horns and pronotum posterior lobe trilobed ( Figs. 5b ix, xii).

Remarks. The holotype of Leptobasis bicornis Selys, 1877 , which I examined and illustrated during two visits to IRSNB in 2001 and 2007 ( Fig. 4b vi), is a female currently missing its head, with mesanepisternal horns adjacent at base and with distal free portions oriented laterally.

Therese (1900: 263) recorded a male of ‘ Leptobasis bicornis ’ from Colombia (Mochila, at the middle Río Magdalena). Following this mention, Colombia was given as part of the distribution range of M. bicornis by later workers (i.e. Ris 1918; Lencioni 2006; Heckman 2008; Pérez-Gutiérrez & Palacino-Rodriguez 2011). A specimen with labels matching the information given in Therese (1900) for identification, locality, collector, and date was found at the ZSM by Lazlo Börzsöny, who kindly photographed it at my request. The picture shows that it represents a female of Metaleptobasis foreli .

Calvert (1909: 198, figs. 108–109) assigned a male from Chapada (in Mato Grosso State, Brazil) with mesanepisternal horns directed laterally and lacking the last four abdominal segments to this species with doubt. Examination of the prothoracic and pterothoracic characters and genital ligula of this specimen ( Figs. 4p iv; 5p iv; 8p ii), deposited at the CMNH, revealed that it corresponds to a male of M. lillianae , which furthest known E locality was in the neighboring lowland forest in Bolivia ( Fig. 14 d).

Williamson (1915) described Metaleptobasis mauritia based on three males from Trinidad and one from Guyana. In his diagnosis he discarded the possibility of this species being the same as M. bicornis Selys (1877) based on the color of pale areas of the head, blue and yellow, and the shape of the mesanepisternal horns, directed antero-dorsally in his three males of M. mauritia , vs. brown and directed laterally respectively in the female holotype of M. bicornis . However, I found that both pale color of head and orientation of mesanepisternal horns can vary intraspecifically and differ between males and females of the same species. Photographs of the holotype female of M. bicornis ( Figs. 2a i , iii–iv) show that the lateral margin of anterior lobe of pronotum projects posteroventrally forming an L-shaped ridge (lv.p.) ending opposite a tubercle (t.) located on the anterior portion of propleuron, which was kindly confirmed by Jérôme Constant (in litt) through direct re-examination of the type. The presence of these characters, in addition to the shared basic black pattern of head and width of mid-dorsal dark stripe of pterothorax ( Figs. 2a i–i ), allowed me to associate this name with numerous specimens of both sexes, including the males holotype and paratypes of M. mauritia Williamson, 1915 . Consequently I consider that Metaleptobasis mauritia Williamson, 1915 is a junior subjective synonym of M. bicornis ( Selys, 1877) . Females of this species can be both andromorphic and heteromorphic, the latter differing from males in shape of pronotal hind lobe and/or development and orientation of mesanepisternal horns, the holotype of M. bicornis representing a heteromorphic female dimorphic for both characters. Males collected in copula with andromorphic and heteromorphic females, and together in several localities, including a heteromorphic female from Belem in Pará, Brazil, which shares the exact shape of pronotal hind lobe and development and orientation of mesanepisternal horns ( Fig. 4b vii) with the female holotype, collected at the same locality as a male and an andromorphic female, reinforce this conclusion.

De Marmels (1992b) assigned six males from Amazonas State in Venezuela to M. fernandezi with doubt. His illustrations match the morphology of M. bicornis , and the slight differences he mentioned in morphology of male cercus tip and pronotum color between these males and the male of M. bicornis from Trinidad, identified as M. mauritia , he compared them with fall well within the intraspecific variability observed here for M. bicornis . In specimens of M. bicornis , even considering only the series of specimens examined from Trinidad (n 9 ♂, 1 andromorphic ♀, 1 heteromorphic ♀), the dark spot on posterior lobe of pronotum shows variable degrees of extension to complete absence, and the degree of angulation and color of male cerci tips is also variable. The single female collected alongside the males, identified as M. fernandezi in the same paper ( De Marmels 1992b), corresponds to a heteromorphic female of M. bicornis . Heteromorphic females of M. bicornis and M. diceras (the senior synonym of M. fernandezi ; see under M. diceras account) can both have mesanepisternal horns represented only by their bases and posterior lobe of pronotum bilobed to sinuous, and andromorphic females of both species have a trilobed pronotum and well developed mesanepisternal horns. However, the two species differ from each other in the width of the mid-dorsal pterothoracic stripe, which is slightly wider in M. bicornis ( Figs. 2a , 3b ; mentioned by De Marmels 1992b: 63 for that female), than in M. diceras ( Fig. 3f ), and in the shape of anterior lobe of pronotum and structural characters of propleuron; in M. bicornis lateral corners of anterior lobe project postero-ventrally, forming a smooth L-shaped ridge (l-v.p) that ends opposite a rounded tubercle (t.) on anterior portion of propleuron ( Figs. 2a ; 4b ; 5b ), while in M. diceras the lateral margins of the anterior lobe do not project postero-ventrally, and there are two sub-vertical crests (l.c.) on anterior portion of propleuron: a crest ventral to anterior lobe lateral margin, which can be tall and denticulate to low and smooth, and a low and smooth crest parallel and posterior to it ( Figs. 4f ; 5f ). De Marmels (in litt.) kindly compared the specimens mentioned in his 1992b paper with my illustrations of the prothorax of M. bicornis and M. diceras , confirming the presence of the diagnostic characters of M. bicornis , and provided me with further records of this species from Venezuela (from Bolívar, Monagas, and Amazonas States; Fig. 14c).

Belle’s (2002: 3) records of M. fernandezi and M. tetragena from Surinam correspond to heteromorphic females of M. bicornis with vestigial horns and sinuous posterior lobe of pronotum, and with well developed horns and indented posterior lobe of pronotum respectively, which I concluded after examining specimens with Belle’s identification labels at RMNH.

Habitat. Forests near rivers, flooded forests, small streams, creeks, swamps, and pools.

Distribution. Trinidad, Venezuela, Guyana, Surinam, French Guiana, and N Brazil ( Fig. 14c).

Figures

FIGURE 1. Head, dorsal view. (a) i. M. amazonica, male lectotype; (a) ii. M. amazonica, female paralectotype; (b) i. M. bicornis, male, Surinam, Kabalebo; (b) ii. M. bicornis, male, French Guiana, N of Cayenne; (b) iii. M. bicornis, female, Surinam, Commewetane; (c) i. M. bovilla, male, Costa Rica, La Selva; (c) ii. M. bovilla, female, same data; (d) i. M. brevicauda, male holotype; (d) ii. M. brevicauda, female paratype, Peru, Tingo María; (e) i. M. brysonima, male, paratype of M. incisula, Venezuela, Marawaka; (e) ii. M. brysonima, female, paratype of M. incisula, same data; (e) iii. M. brysonima, female, holotype of M. weibezahni; (e) iv. M. brysonima, female, holotype of M. tetragena; (f) i. M. diceras, male lectotype; (f) ii. M. diceras, male, Brazil, Abuna; (f) iii. M. diceras, female paralectotype; (f) iv. M. diceras, female, holotype of M. fernandezi; (f) iv. M. diceras, female, Venezuela, no locality; (f) v. M. diceras, female, Brazil, Abuna; (g) i. M. falcifera, male holotype; (g) ii. M. falcifera, female paratype, Peru, Pakitza; (h) i. M. foreli, male, Colombia, Puerto Berrio; (h) ii. M. foreli, male, paratype of M. westfalli, Panama, No. 5 on Standley, 22 vii 1950, FSCA; (h) iii. M. foreli, female, same data.

FIGURE 2. Head and thorax. (a) i. M. bicornis, female holotype, dorsal view; (a) ii. M. bicornis, male, holotype of M. mauritia, dorsal view; (a) iii. M. bicornis, female holotype, dorsal view; (a) iv. M. bicornis, female holotype, lateral view; (b) i. M. brysonima, male holotype, dorsal view; (b) ii. M. brysonima, male holotype, lateral view. l-v.p.: latero-ventral margin of anterior lobe projected ventro-posteriorly; t.: tubercle.

FIGURE 3. Pterothorax, dorsal view. (a) i. M. amazonica, male lectotype; (a) ii. M. amazonica, female paralectotype; (b) i. M. bicornis, male, French Guiana, N of Cayenne; (b) ii. M. bicornis, male, Surinam, Kabalebo; (b) iii. M. bicornis, female, Surinam, Moengotapoe; (b) iv. M. bicornis, female, Surinam, Commewetane; (c) i. M. bovilla, male, Costa Rica, La Selva; (c) ii. M. bovilla, female, same data; (d) i. M. brevicauda, male holotype; (d) ii. M. brevicauda, female paratype, Peru, Tingo María; (e) i. M. brysonima, male, paratype of M. incisula, Venezuela, Marawaka; (e) ii. M. brysonima, female, paratype of M. incisula, same data.

FIGURE 4. Prothorax and anterior portion of pterothorax, dorsal view. (a) i. M. amazonica, male lectotype; (a) ii. M. amazonica, male paralectotype; (a) iii. M. amazonica, female paralectotype; (b) i. M. bicornis, male, Surinam, Kabalebo; (b) ii. M. bicornis, male, Brazil, Paranapuebas; (b) iii. M. bicornis, male, French Guiana, N of Cayenne; (b) iv. M. bicornis, male, Guyana, Potaro; (b) v. M. bicornis, male, Trinidad, Cumuto; (b) vi. M. bicornis, female holotype. o.c.: oblique crest; cr.: crest; e-l.: ear-lobe; l-v. p.: latero-ventral projection; p.l.: postero-lateral projection; t.: tubercle.

FIGURE 5. Prothorax and anterior portion of pterothorax, lateral view. (a) i. M. amazonica, male lectotype; (a) ii. M. amazonica, female paralectotype; (b) i. M. bicornis, male, Trinidad, Cumuto; (b) ii. M. bicornis, male, Guyana, Potaro; (b) iii. M. bicornis, male, Surinam, Kabalebo; (b) iv. M. bicornis, male, French Guiana, N of Cayenne; (b) v. M. bicornis, male, Brazil, Paraupebas; (b) vi. M. bicornis, female, Surinam, Langamankondre; (b) vii. M. bicornis, female, Surinam, Kabalebo; (b) viii. M. bicornis, female, Surinam, Gansee; (b) ix. M. bicornis, female, Surinam, Commetewane; (b) x. M. bicornis, female, French Guiana, N of Cayenne. c.: c-shaped crest; e-l.: ear-lobe; l-v.p.: latero-ventral margin projection; p.l.: postero-lateral projection; t.: tubercle.

FIGURE 8. Genital ligula, lateral view (above) and ventral view (below). (a) M. amazonica, lectotype; (b) i. M. bicornis, Surinam, Kabalebo; (b) ii. M. bicornis, French Guiana, N of Cayenne; (c) i. M. bovilla, Costa Rica, La Selva; (c) ii. M. bovilla, same data; (d) M. brevicauda, holotype; (e) i. M. brysonima, paratype of M. incisula, Venezuela, Marawaka; (e) ii. M. brysonima, Peru, Tambopata; (e) iii. M. brysonima, Brazil, Abuna; (f) i. M. diceras, lectotype; (f) ii. M. diceras, Brazil, Abuna; (f) iii. M. diceras, Peru, Manu. e.f.: ectal fold; pr.: prominent protuberance; s.d.: slight depression; t.: terminal fold.

FIGURE 9. Female S8–10, lateral view unless indicated otherwise. (a) M. amazonica, paralectotype; (b) i. M. bicornis, Surinam, Commetewane; (b) ii. M. bicornis, Brazil, Belem; (c) M. bovilla, Costa Rica, Puerto Viejo; (d) M. brevicauda, paratype, Peru, Tingo María; (e) i. M. brysonima, paratype of M. incisula, Venezuela, Marawaka; (e) ii. M. brysonima, holotype of M. weibezahni; (e) iii. M. brysonima, holotype of M. tetragena; (f) i. M. diceras, paralectotype; (f) ii. M. diceras, Brazil, Abuna; (f) iii. M. diceras, Guyana, Kamana; (f) iv. M. diceras, Venezuela; (f) v. M. diceras, Trinidad, N of Sangre Grande; (g) i. M. falcifera, paratype, Peru, Explorer’s Inn, ventral view on right.

FIGURE 10. Male S10, dorsal view. (a) M. amazonica, lectotype; (b) i. M. bicornis, Brazil, Paraupebas; (b) ii. M. bicornis, Surinam, Kabalebo; (b) iii. M. bicornis, Guyana, Kumu River; (b) iv. M. bicornis, French Guiana, N of Cayenne; (b) v. M. bicornis, Trinidad, E of Valencia; (c) M. bovilla, Costa Rica, La Selva; (d) i. M. brevicauda, holotype; (e) M. brysonima, Venezuela, Río Baria; (f) i. M. diceras, Brazil, Paraupebas; (f) ii. M. diceras, Brazil, Abuna; (f) iii. M. diceras, Peru, Pakitza; (f) iv. M. diceras, Peru, Pakitza; (f) v. M. diceras, Trinidad, Cumuto, paratype of M. manicaria; (f) vi. M. diceras, Trinidad, Cumuto, paratype of M. manicaria; (g) M. falcifera, holotype; (h) i. M. foreli, Colombia, Fundación; (h) ii. M. foreli, paratype M. westfalli, Panama, Barro Colorado # 54; (i) i. M. furcifera, holotype; (i) ii. M. furcifera, paratype, Peru, Río Seco. b.t.: bicuspidate tooth; r.: ridge.

FIGURE 11. Male S10, medio-dorsal view. (a) M. amazonica, lectotype; (b) i. M. bicornis, Surinam, Kabalebo; (b) ii. M. bicornis, Guyana, Potaro; (b) iii. M. bicornis, French Guiana, N of Cayenne; (b) iv. M. bicornis, Trinidad, E of Valencia; (c) M. bovilla, Costa Rica, La Selva; (d) M. brevicauda, holotype; (e) i. M. brysonima, Venezuela, Río Baria; (e) ii. M. brysonima, Peru, Tambopata; (e) iii. M. brysonima, Brazil, Abuna; (f) i. M. diceras, Brazil, Abuna; (f) ii. M. diceras, Trinidad, Cumuto, paratype of M. manicaria; (f) iii. M. diceras, Trinidad, E of Valencia; (g) M. falcifera, holotype; (h) M. foreli, Colombia, Fundación; (i) i. M. furcifera, paratype, Peru, Río Seco; (i) ii. M. furcifera, paratype, Peru, Río Seco; (j) M. gabrielae, holotype; (k) M. gibbosa, Ecuador, Copales; (l) M. guillermoi, holotype. b.t.: bicuspidate tooth; p-m.p.: postero-medial projection; pr.: prominence; r.: ridge; s.t.: single tooth; te.: teeth.

FIGURE 12. Male S10, lateral view. (a) M. amazonica, lectotype; (b) i. M. bicornis, Surinam, Kabalebo; (b) ii. M. bicornis, French Guiana, N of Cayenne; (c) M. bovilla, Costa Rica, La Selva; (d) M. brevicauda, holotype; (e) i M. brysonima, paratype of M. incisula, Venezuela, Marawaka; (e) ii M. brysonima, Venezuela, Río Baria; (f) i. M. diceras, Brazil, Paraupebas; (f) ii. M. diceras, Brazil, Abuna; (f) iii. M. diceras, Peru, Pakitza; (f) iv. M. diceras, Trinidad, Cumuto, paratype of M. manicaria; (g) M. falcifera, holotype; (h) i. M. foreli, Colombia, Fundación; (h) ii. M. foreli, paratype M. westfalli, Panama, Barro Colorado # 54; (i) i. M. furcifera, holotype; (i) ii. M. furcifera, paratype, Peru, Río Seco; (j) M. gabrielae, holotype; (k) M. gibbosa, paratype [FSCA], Ecuador, Copales; (l) M. guillermoi, holotype. sw.: swelling.