Metaleptobasis foreli Ris, 1918

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 : 35-37

Metaleptobasis foreli Ris, 1918

Figs. 1h; 3h; 4h; 5h; 8h; 9h; 10h; 11h; 12h; 13d; 14b

Leptobasis bicornis nec Selys, 1877 — Therese (1900: 263; mention from Colombia).— Misidentification.

Metaleptobasis foreli Ris, 1918: 130–131 , fig. 72 (description ♀, illustration of pronotum);— Geijskes (1932: 261; mention);— Cumming (1954: 24, 27, 31; mention);— Rácenis (1955: 6–10, 18, figs. 1a–e; description ♂, illustrations of ♂ mesanepisternal horns and S10, ♀ mesanepisternal horns);—De Marmels (1990: 337; mention);— Heckman (2008: 397, fig. 3.1.458; in key, reproduction of Ris 1918 illustration of ♀ pronotum)— Garrison & von Ellenrieder (2009: 48; mention);— Garrison et al. (2010: 284; mention).

Metaleptobasis bicornis nec Selys, 1877 — Ris (1918: 191; mention based on misidentification in Therese 1900);—Pérez- Gutiérrez & Palacino-Rodríguez (2011: 214; mention based on misidentification in Therese 1900 repeated in Ris 1918).— Misidentifications.

Metaleptobasis westfalli Cumming, 1954: 23–32 , figs. 1–5 (description ♂ ♀, inclusion in key to ♂, illustrations of ♂ mesanepisternal horns, S10, and ♀ S8–10);— Rácenis (1955: 10; comparison with M. foreli );— Ramírez et al. (2000: 4; mention);— Garrison & von Ellenrieder (2009: 49; mention);— Garrison et al. (2010: 284; mention);— Pérez-Gutiérrez & Palacino-Rodríguez (2011: 214; mention).— New synonymy.

Types. Holotype ♀: COLOMBIA, Magdalena Dep., Don Diego {11°14'N, 73°41'W, 229 m}, Forel leg. [ DEI].

Specimens examined. Total: 181 ♂, 98 ♀: COSTA RICA, Guanacaste Prov. : 1 ♀ (he), Estación Biológica Pitilla, 9 km S of Santa Cecilia, Parque Nacional Guanacaste, in regenerated forest {10°59'N, 85°25'W, 700 m}, 19 xi 1991, C. Moraga leg. [ INBIO]; Alajuela Prov. : 1 ♂, Los Chiles {10°25'N, 84°20'W, 60 m}, 13 x 1966, DRP leg. [ FSCA]; Puntarenas Prov. : 1 ♂, near Reserva Biológica Carara, Quebrada Bonita and smaller tributaries, swampy seep in forest along Laguna trail {9°46'N, 84°33'W, 3m}, WAH leg. [ WAH] ; 1 ♀ (he), Villa Colón , 16 mi S of Palmar Norte {8°44'N, 83°10'W, 3 m}, 3 vii 1967, O.S. Flint Jr. & M.A. Ortiz B. leg. [ USNM] ; 2 ♂, vicinity of Rincón de Osa {8°42'N, 83°29'W, 270 m}, 9 iii 1967, DRP & M.L. Paulson [ DRP] ; 1 ♂, same but 20 iii 1967 [ DRP] ; 1 ♂, Parque Nacional Corcovado, Estación Biológica Los Patos a Sirena Pavo , 26 xii 2000, J. Azofeifa leg. [ CE]; PANAMA, Colón Prov. : 2 ♂, W of Portobelo, Río Buenaventura {9°33'N, 79°39'W, 113 m}, 2 viii 1974, T. Abrams leg. [ DRP] ; 1 ♀ (an), Portobelo Dis., road to Portobelo, 13,4 km NE of Colón, Panama City Highway Tropical moist forest {9°26'N, 79°45'W, 5 m}, 21 xii 1993, J. Beierl leg. [ RWG]; Panamá Prov. : 1 ♂ holotype, 1 ♀ (an) allotype of M. westfalli, Barro Colorado Island {9°9'N, 79°51'W, 62 m}, 22 vii 1950, R.B. Cumming leg. [ UMMZ] ; 1 ♂, 1 ♀ (an), paratypes of M. westfalli , same but small pond at No. 5 on Standley trail [ USNM] ; 1 ♂, 1 ♀ (he), paratypes of M. westfalli , in tandem, same but [ FSCA] ; 3 ♂ paratypes of M. westfalli , same but [ FSCA] ; 1 ♂, same but pond, 17 v 1960 [ TWD] ; 1 ♂ paratype of M. westfalli, Summit, Canal Zone Experimental Gardens (9°4'N, 79°39 W, 75 m), 1 ix 1950, R.B. Cumming leg. [ FSCA] ; 2 ♀ (he), same but xi 1946, N.L.H. Krauss leg. [ FSCA] ; 1 ♀ (he), Barro Colorado Island, pond at Standley nr 59 (9°0'N, 79°51'W, 62 m), 18 ii 1972, M. May leg. [ RWG] ; 1 ♂, same but 6 i 1974 [ RWG] ; 1 ♂, same but 6 vi 1974, M.L. May leg. [ FSCA] ; 1 ♂, Lion Hill {9°13'N, 79°53'W, 64 m}, 18 vi 1907, A. Busck [ USNM] ; 1 ♂, Trinidad River {8°8'N, 81°14'W, 217 m}, 1–10 vi 1912, A. Busck [ USNM] ; 3 ♀ (an), 1 ♀ (he), same but 15–30 iii 1912 [ USNM] ; 1 ♂, 2 ♀ (an), same but [ FSCA] ; 1 ♂, same but 2 v 1911 [ FSCA] ; 1 ♀ (he) Canal Zone, Cocoli , ix 1946, N.L.H. Krauss leg. [ FSCA] ; 1 ♂ paratype of M. westfalli, Pacora region {9°5'N, 79°14'W, 6 m}, 26 viii 1950, R.B. Cumming leg. [ FSCA]; Darién Prov. : 1 ♀ (an), Parque Nacional Darién, Río Perrecénega at Estación Rancho Frío ( Pirre Station ), 14 km S of El Real (8°2'24''N, 77°45'0''W, 200 m), 27–31 iii 2002, J.L. Hogue & F. Hertel leg. [ RWG]; Heredia Prov .: 1 ♀ (an), Porto Bello {8°1'N, 80°36'W, 49 m}, 18 iv 1912, A. Busck leg. [ FSCA]; COLOMBIA, Antioquia Dep .: 2 ♂, Las Delicias {6°36'N, 76°6'W, 2059 m}, 28 i 1917, J.H. & E.B. Williamson leg. [ UMMZ] ; 17 ♂, 6 ♀ (an), Puerto Berrío {6°30'N, 74°24'W, 69 m}, 31 i 1917, J.H. & E.B. Williamson leg. [ UMMZ] ; 6 ♂, 4 ♀ (an), same but 8 ii 1917 [ UMMZ] ; 1 ♂, 1 ♀ (an), same but [ CEUA] ; 1 ♀ (an), same but 21 ii 1917 [ UMMZ] ; 1 ♂, Municipio El Bagre, Vereda 505 (70 m), 18 iii 2011, M. Ruiz [ CEUA]; Magdalena Dep .: 2 ♂, 2 ♀ (an), Fundación , S of Aracataca {10°31'N, 74°11'W, 45 m}, 11 i 1917, J.H. & E.B. Williamson leg. [ RWG] ; 21 ♂, 18 ♀ (an), 8 ♀ (he), same but [UMMZ]; 1 ♂, same but 14 i 1917 [ UMMZ] ; 18 ♂, 12 ♀ (an), 1 ♀ (he), El Banco {9°3'N, 73°58'W, 46 m}, 24 i 1917, J.H. & E.B. Williamson leg. [ UMMZ] ; 1 ♂, 1 ♀ (an), same but [ RWG] ; 5 ♂, 3 ♀ (an), same but 23 i 1917 [ UMMZ]; VENEZUELA, Zulia State : 2 ♂, El Guayabo {10°37'N, 71°51'W, 84 m}, 20 iv 1920, J.H. & E.B. Williamson & W.H. Ditzler [ UMMZ] ; 7 ♂, 3 ♀ (an), El Guayabo {10°37'N, 71°51'W, 84 m}, 22 iv 1920, J.H. & E.B. Williamson & W.H. Ditzler [ UMMZ]; Falcón State : 1 ♂, Tucacas {10°47'N, 68°19'W, 2 m}, 25 iii 1920, J.H. & E.B. Williamson & W.H. Ditzler [ UMMZ] ; 55 ♂, 1 ♀ (he), Palma Sola {10°36'N, 68°32'W, 37 m}; 9 iii 1920, J.H. & E.B. Williamson & W.H. Ditzler [ UMMZ] ; 2 ♂, same but [ RWG] ; 2 ♂, same but 8 iii 1920 [ UMMZ]; Yaracuy State : 3 ♂, Boquerón {10°34'N, 68°49'W, 91 m}, 16 iii 1920, J.H. & E.B. Williamson & W.H. Ditzler leg. [ UMMZ]; Miranda State : 1 ♂, San José {10°13'N, 66°23'W, 63 m}, 3 x 1954, J. Rácenis [ FSCA] ; 1 ♂, between El Guapo and Cúpira (10°12'N, 65°48'W), 29 vii 1961, TWD leg. [ TWD]; Barinas State : 1 ♀ (he), San Silvestre {8°16'N, 70°6'W, 119 m}, 23 xii 1957, J. Rácenis [ FSCA]; Táchira State : 11 ♂, 18 ♀ (an), La Fría {8°13'N, 72°14'W, 155 m}, 12–18 iv 1920, J.H. & E.B. Williamson & W.H. Ditzler [ UMMZ] .

Characterization. Head. Labrum mostly pale; black on dorsum of head extensive; postocular lobes rounded ( Fig. 1h). Thorax. Pronotum anterior lobe with a postero-lateral digit-shaped projection on each side (po.): well developed, much longer than wide in 83% of examined females ( Figs. 4h iii–iv; 5h iii, v), short, about as long as wide in 4.5% of examined males ( Figs. 4h; 5h ii) and 4% of examined females ( Fig. 5h vi), fused to middle lobe of pronotum in 12% of examined females ( Fig. 5h vii), or absent in 95.5% of examined males and 1% of examined females ( Figs. 4h i –ii; 5h i, iv); anterior and middle lobes of pronotum separated dorso-laterally by a groove; anterior area of propleuron with a prominent rounded tubercle (t.), which is followed posteriorly by a latero-longitudinal crest (lo.); anterior margin of middle lobe of pronotum forming a crest (cr.) followed by a transverse dorsal depression; pronotum posterior lobe slightly trilobed, with medial lobe smoothly convex to medially pointed, and lateral lobes about as long as medial lobe; mesanepisternal horns with bases separated; in males and andromorphic females ( Figs. 4h i –iii; 5h i–ii, iv–vii) well developed, as long as four to five times mesostigmal plate width, thin, with tips pointed, oriented dorsally at an angle of 85°–105° with dorsum, with distal portions sometimes bent anteriorly or medially or posteriorly, in a few cases asymmetrical in length and orientation or even curled, in heteromorphic females ( Figs. 4h iv; 5h iii) blunt, as long as about 0.30 of mesostigmal plate width; mid-dorsal dark stripe as wide 0.33–0.50 of mesanepisterna and parallel sided along most of its length, narrowing slightly anteriorly between mesanepisternal horns ( Fig. 3h); Pt short rectangular, with posterior and anterior sides slightly longer than distal side, to squarish, with all sides about equal. Abdomen. Male genital lobe ( Fig. 8h) short, less than 0.50 of anterior hamule height; smoothly curved; 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; genital ligula distal segment pear-shaped with a pair of lateral sclerotized tubercles at base (p.t.), apex convex with a wide ectal fold (e.f., Fig. 8h); posterior margin of female S8 sternum smooth, lacking any denticles, spines, or processes ( Fig. 9h); distal end of ovipositor reaching level between tip of paraproct and tip of cercus to tip of cercus; medial portion of male S10 postero-dorsal margin ( Figs. 10h; 11h; 12h) projected posteriorly into a triangular to bifid process, lacking an incision or slit, and lacking a dorsal prominence; male cercus with basal half globose and distal half sub-cylindrical, medial margin about straight in dorsal view ( Fig. 10h), narrowing to pointed tip, with a single tooth directed ventrally; ratio of male cercus length to S10 maximum length in lateral view 1.0– 1.5; ratio of male cercus length to paraproct length in lateral view 0.60–0.66; male paraproct in lateral view ( Fig. 12h) gently arched dorsally, about as wide at tip as at medial section, with medial margin sinuous and tip truncate in dorsal view, with a sub-apical ridge on medial surface ending on a single distal tooth directed medially ( Figs. 10h; 11h).

Dimensions. Males (n 10): Hw 22.1 ± 1 [20.2–23.2]; abdomen 37.8 ± 1.6 [35–40]; total length 45.7 ± 1.8 [42.8–48]. Females (n 10): Hw 23.2 ± 0.6 [22.3–24.2]; abdomen 37.2 ± 1.1 [35.8–38.7]; total length 44.7 ± 1.3 [42.2–46.8].

Diagnosis. Among the species with male posterior hamule small and digit-like, posterior margin of female S8 sternum lacking any denticles, spines, or processes, black on dorsum of head extensive, and labrum mostly pale, M. foreli shares anterior margin of middle lobe of pronotum forming a distinct crest only with M. quadricornis and M. turbinata . It differs from M. turbinata by the crest (cr.) being entire ( Figs. 4h; 5h; bilobate in M. turbinata , Figs. 4 ae; 5ae), distal segment of genital ligula lacking flagella ( Fig. 8h; with long coiled paired flagella in M. turbinata , Fig. 8 ae), and male cercus distinctly shorter than paraproct ( Figs. 10h; 12h; slightly longer than paraproct in M. turbinata , Figs. 10 ae; 12ae). It differs from M. quadricornis by posterior lobe of female pronotum slightly trilobed with lateral lobes shorter than medial lobe ( Figs. 4h iii–iv; deeply trilobed with lateral lobes rounded and longer than medial lobe in M. quadricornis , Figs. 4y iv–vii), distal segment of genital ligula ( Fig. 8h) with a pair of sclerotized pointed tubercles at base (p.t.) and a with a wide ectal fold (e.f.) (lacking sclerotized pointed tubercles at base and lacking an ectal fold in M. quadricornis , Fig. 8y), male cercus basal half globose, distal half sub-cylindrical, with only extreme tip pointed ventrally ( Fig. 12h; sub-cylindrical, gradually narrowing distally, with tip curved ventrally forming a hook in M. quadricornis , Fig. 12y), and male paraproct curved ventrally gradually, with distal end rounded in lateral view ( Fig. 12h; curved ventrally at an angle, with tip pointed in lateral view in M. quadricornis , Fig. 12y).

Most females (83%) differ from males by the presence of a well developed postero-lateral digit-shaped projection on each side of anterior lobe of pronotum (po.); however, this structure is reduced to absent in some females, and also present in a few males (4.5%) in a reduced state, making it unpractical to categorize females as andromorphic based on this feature. About 81% of the females share mesanepisternal horns as long as four to five times mesostigmal plate width, thin, directed dorsally at an angle of 85°–105° with dorsum and with tips pointed with males ( Figs. 4h iii; 5 iv–vii), whereas in heteromorphic females (about 19%) mesanepisternal horns are limited to their blunt bases ( Figs. 4h iv; 5h iii).

Remarks. Ris (1918) described M. foreli based on a single specimen, an andromorphic female, from Colombia. In his description of M. westfalli, Cumming (1954) did not compare his new species with M. foreli , diagnosing it only from the other described species with known males. Even though Cumming (1954) described the female of M. westfalli , he did not mention the lateral digit-shaped projections of the anterior lobe of pronotum which are present in the allotype and female paratypes of M. westfalli ( Figs. 4h iv; 5h iii), and which are shared with the holotype of M. foreli as described and illustrated by Ris (1918: fig. 72). Based on Cumming’s (1954) illustrations of the holotype male of M. westfalli, Rácenis (1955) diagnosed the male of M. westfalli from that of M. foreli by the inner margin of paraprocts in dorsal view entire along most of their length vs. sinuous in M. foreli , projection of postero-medial margin of S10 triangular vs. slightly bifid in M. foreli , and absence of transverse green spot on posterior lobe of pronotum vs. present in M. foreli . However, examination of the type series of M. westfalli shows that the shape of the projection of postero-medial margin of S10 is variable, from triangular to slightly bifid ( Fig. 10h ii), that the illustration of the paraprocts ( Cumming 1954: Fig. 2) is inaccurate, since inner margin of paraprocts in dorsal view is sinuous in all specimens of the type series, and that the transverse green spot on the posterior lobe of pronotum is also present in all specimens. Comparison of the type series of M. westfalli , which includes males and females in tandem, with a large series of specimens of M. foreli from Colombia and Venezuela including males and females, shows them to represent the same species, and in consequence Metaleptobasis foreli Ris 1918 constitutes the senior subjective synonym of Metaleptobasis westfalli Cumming 1954 .

Habitat. Forest near rivers and small ponds. Bota-Sierra (pers. comm.) observed that after a night of heavy rain, the forest at the National Park Tayrona in Colombia ‘was filled with adults of M. foreli ( Fig. 13d i), principally close to the edges of a little stream, appearing close to 7 a.m. and disappearing before the afternoon’. He observed no reproductive activity.

Distribution. Costa Rica along Pacific slope, Panama, Colombia, and Venezuela ( Fig. 14b).