Metaleptobasis quadricornis ( 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 : 66-68

Metaleptobasis quadricornis ( Selys, 1877)

Figs. 1y ; 2y ; 3y ; 4y ; 5y ; 8y ; 9x ; 10y ; 11y ; 12y ; 14b

Leptobasis quadricornis Selys, 1877: 104–105 [10–11 reprint] (description ♀).

Metaleptobasis quadricornis: Calvert (1907: 386 ; suggestion of possible inclusion in genus);— Williamson (1915: 608; mention);— Calvert (1948: 61; mention);— Cumming (1954: 24, 30; mention);— Lencioni (2006: 166, fig. 104 A; mention, illustration mesanepisternal horns);— Heckman (2008: 397, fig. 3.1.457; in key, reproduction of illustration from Lencioni 2006);—von Ellenrieder & Garrison (2007: 25, figs. 11f–h; illustration of female posterior lobe of pronotum, mesanepisternal horns and pretarsal claw);— Garrison & von Ellenrieder (2009: 49; mention);— Garrison et al. (2010: 284, figs. 1826, 1833; mention, reproduction of illustrations from von Ellenrieder & Garrison 2007);—von Ellenrieder (2011: 57–58, 62, 73; mention).

Metaleptobasis brysonima nec Williamson, 1915 —Rácenis (1956: 12–13, 18–19, fig. 2d mislabeled in legends as 2c; description of ♀ based on misidentified ♂ from Venezuela, inclusion in key for species from Venezuela);— Michalski (1988: 45; in part, inclusion of female in key);— De Marmels (1989: 31; mention);— De Marmels (1992a: 337 ; mention);— De Marmels (1992b: 63 ; mention).— Misidentifications.

Metaleptobasis weibezahni nec Rácenis, 1956—Belle (2002: 3; mention);— Garrison & von Ellenrieder (2009: 49; mention).— Misidentification.

Types. Holotype ♀ *: BRAZIL, Pará State , Pará {1°27'S, 48°29'W, 2 m}, Bates leg. [ IRSNB].

Specimens examined. Total: 53 ♂, 64 ♀: 1 ♀ holotype; SURINAME : 1 ♀, No locality, Staudinger & Bang- Haas vendors [ BMNH]; Brokopondo Dis .: 1 ♀, Kabelstation , forest {4°55'N, 55°8'W, 41 m}, 23 ix 1938, D.C. Geijskes leg. [ RMNH] ; 1 ♀, same but Suriname River , 4 i 1960, J. Belle leg. [ RMNH] ; 2 ♂, Stoelmans Island , forest by the river {5°4'N, 54°58'W, 10 m}, 18 viii 1953, D.C. Geijskes leg. [ RMNH]; Para Dis .: 1 ♂, Sapenda to Copie {5°31'N, 54°44'W}, in forest 14 vii 1953, D.C. Geijskes [ RMNH] ; 1 ♀, Zanderij {5°27'N, 55°12'W, 23 m}, 1956, J. Belle leg. [ TWD] ; 1 ♂, 1 ♀, same but Troelinde Creek , 1 viii 1959 [ FSCA] ; 1 ♀, same but [ RWG] ; 1 ♂, same but 8 viii 1959 [ RMNH] ; 1 ♂, same but 19 viii 1962 [ TWD] ; 1 ♂, Zanderij I, forest, 2 ix 1942, D.C. Geijskes leg. [ RMNH] ; 1 ♂, same but Pontjibrug , 10 i 1957, J. Belle leg. [ RMNH] ; 1 ♀, same but 12 i 1957 [ RMNH] ; 1 ♂, same but 23 vi 1957 [ RMNH] ; 1 ♀, Mata , in forest {5°27'N, 55°19'W, 20 m}, 8 i 1963, S. Ligorie leg.[ RMNH] ; 1 ♀, same but Boven Coropino , gallery forest {5°32'N, 55°10'W}, 28 iv 1957 [ RMNH] ; 1 ♂, 3 ♀, same but 29 vii 1958 [ RMNH] ; 1 ♀, same but 4 xi 1958 [ RMNH] ; 1 ♀, same but trail near Hannover , small creek in savannah, 13 xi 1949, D.C. Geijskes leg. [ RMNH] ; 1 ♀, Republiek {5°30'N, 55°12'W, 15 m}, 10 x 1946, D.D. Geijskes leg. [ RMNH] ; 1 ♂, same but forest, 1 xi 1963 [ RMNH]; 2, same but 19 x 1963 [ RMNH] ; 2 ♂, same but Coropina Creek , forest near creek {5°32'N, 55°10'W, 10 m}, 9 iii 1947 [ RMNH]; Commewijne Dis .: 1 ♀, Copie {5°32'N, 54°48'W, 30 m}, 26 xi 1979, C.F. Mees leg. [ RMNH] ; 1 ♀, near Potribo, Macreabo Creek {5°45'N, 54°46'W, 10 m}, 16 x 1954, Radin leg. [ RMNH]; Suriname Dis .: 1 ♂, Lelydorp , forest km 25 {5°42'N, 55°14'W, 8 m}, 25 x 1938, D.C. Geijskes leg. [ RMNH] ; 1 ♀, Rijsdijkweg , 12 i 1957, J. Belle leg. [ RMNH]; Marowijne Dis .: 1 ♂, Donderbari, Baptist Creek {5°14'N, 55°14'W, 50m}, 15 xii 1945, D.C. Geijskes leg. [ RMNH] ; 1 ♂, Mooi Wanna Creek, Weijneweg {5°35'N, 54°15'W}, 4 i 1964, J. Belle leg. [ RWG] ; 1 ♂, 2 ♀, Wane Creek N, Camp 1, forest {5°38'N, 54°2'W, 20 m}, 28 ix 1948, D.C. Geijskes leg. [ RMNH] ; 1 ♂, same but near km 8, 30 ix 1948 [ RMNH] ; 1 ♂, Moengo, Cottio River {5°37'N, 54°24'W, 25 m}, 17 v 1927, E.B. Williamson leg. [ FSCA] ; 1 ♀, same but boermanhill in forest, 18 ix 1961, D.C. Geijskes [ RMNH] ; 1 ♂, 1 ♀, Moengotapoe , trail km 0.5 {5°34'N, 54°14'W, 33 m}, 20 ix 1948, D.C. Geijskes [ RMNH] ; 2 ♂, same but 21 ix 1948 [ RMNH] ; 1 ♀, same but 22 ix 1948 [ RMNH] ; 1 ♂, 1 ♀, same but trail km 1.0–1.5 [ RMNH] ; 1 ♀, same but trail km 3–4 [ RMNH] ; 1 ♂, same but 23 ix 1948 [ RMNH] ; 1 ♀, same but trail km 4.6, 24 ix 1948 [ RMNH] ; 2 ♀, same but Djai Creek , trail 1 km 8.6, forest, 10 x 1948 [ RMNH] ; 4 ♀, same but 3rd camp, trail km 15–15.7, forest on sand ridge parallel to the coast, 13 x 1948 [ RMNH] ; 1 ♀, same but km 14.9, forest on sand ridge parallel to the coast, 14 x 1948 [ RMNH] ; 1 ♂, 4 ♀, same but trail km 14.3–15.5 [ RMNH] ; 2 ♀, same but N trail km 15.7 [ RMNH] ; 5 ♀, same but trail km 14.6–14.9, 18 x 1948 [ RMNH] ; 1 ♂ 1 ♀, same but trail km 15.5 [ RMNH] ; 2 ♀, same but 19 x 1948 [ RMNH] ; 2 ♂, trail km 14.2–14.7, forest on sand ridge parallel to the coast, 14 x 1948 [ RMNH] ; 2 ♂, same but trail km 14.6–14.9, 18 x 1948 [ RMNH]; Sipaliwini Dis .: 1 ♀, Werehpai , forest trail (2°21'46''N, 56°41'53''W, 258 m), 3 ix 2010, NVE [ NVE] ; 1 ♂, Tafelberg Creek , trail km 20 {3°47'N, 56°11'W, 170 m}, 1 xi 1943, D.C. Geijskes [ RMNH] ; 1 ♂, 1 ♀ (in copula), same but 26 iii 1958 [ RMNH] ; 1 ♂, Nassaugebergte , trail km 0.7, forest {4°48'N, 54°29'W}, 10 ii 1949, D.C. Geijskes [ RMNH] ; 1 ♀, same but trail km 6, 14 ii 1949 [ RMNH] ; 1 ♂, 1 ♀, same but km 0.7, 15 ii 1949 [ RMNH] ; 1 ♂, 1 ♀, same but trail km 1 [ RMNH] ; 1 ♂, 1 ♀, same but trail km 2, forest, 16 ii 1949 [ RMNH] ; 1 ♂, same but km 0.7, forest, 17 ii 1949 [ RMNH] ; 1 ♂, same but trail km 1.4, forest [ RMNH] ; 2 ♂, same but trail km 1.2 forest creek, 18 ii 1949 [ RMNH] ; 1 ♀, same but trail km 0.7 [ RMNH] ; 2 ♂, same but trail km 1.3 creek, 22 ii 1949 [ RMNH] ; 1 ♂, same but trail km 1.2 forest creek [ RMNH] ; 1 ♂, same but trail km 1.4, forest [ RMNH] ; 2 ♂, same but trail km 11.5 [ RMNH] ; 3 ♂, same but trail km 1.4–1.5, forest [ RMNH] ; 1 ♂, 1 ♀ (in copula), same but trail km 1.2, forest creek [ RMNH] ; 1 ♂, same but trail km 6, 24 ii 1949 [ RMNH] ; 1 ♀, same but trail km 6, 1 iii 1949 [ RMNH]; FRENCH GUIANA: Saint Laurent-Du-Maroni Dep .: 1 ♂, 3 ♀, Pied Saut, Oyapok river {5°21'N, 53°41'W, 19 m}, xi 1917, S.M. Klages leg. [ FSCA] ; 1 ♀, Mana river {5°40'N, 53°47'W, 1 m}, v 1917 [ FSCA] ; 1 ♂, 1 ♀, Tamanoir , v 1917 [ FSCA]; BRAZIL: Pará State : 1 ♀, Benevides {1°22'S, 48°15'W, 21 m}, x 1918, S.M. Klages leg. [ FSCA] .

Characterization. Head. Labrum mostly pale; black on dorsum of head extensive; postocular lobes markedly angled ( Fig. 1y ). Thorax. Pronotum anterior lobe ( Figs. 4y ; 5y ) smooth; anterior and middle lobes of pronotum separated dorso-laterally by a groove; anterior area of propleuron with a prominent rounded tubercle (t.), a short longitudinal crest (lo.) and a c-shaped crest (c.); middle lobe of pronotum smooth; anterior margin of middle lobe of pronotum forming a prominent crest (cr.) followed by a transverse dorsal depression; pronotum posterior lobe entire to slightly trilobed in male, trilobed with medial lobe with short central point and lateral lobes rounded and longer than medial lobe in female; lateral margins of pronotum posterior lobe free and rounded; bases of mesanepisternal horns joined by a low ridge to separated, horns thin to medium in width, parallel sided to slightly divergent to convergent, directed antero-dorsally at 60°–85° with dorsum in lateral view, with tip bluntly pointed, well developed, equal to two times mesostigmal plate width to slightly longer in male, two to three times mesostigmal plate width in not dimorphic female, and 0.50 of mesostigmal plate width in dimorphic female; mid-dorsal dark stripe wide, 0.50 of mesanepisterna, wider than mesostigmal sinus, narrowing between horn bases, remainder parallel sided ( Fig. 3y ); Pt sub-rectangular, with anterior and posterior sides longer than distal side. Abdomen. Male genital lobe short, less than 0.50 of anterior hamule height, 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, apex with a shallow u- or vshaped incision, lacking an ectal fold ( Fig. 8y ); posterior margin of female S8 sternum smooth, lacking any denticles, spines, or processes; distal end of ovipositor reaching apex of paraproct to apex of cercus ( Fig. 9x ); medial portion of male S10 postero-dorsal margin ( Figs. 10y ; 11y ; 12y ) projected posteriorly, with a small u-shaped incision and lacking a dorsal prominence; male cercus in dorsal view ( Fig. 10y ) about straight, narrowing to tip, which forms a hook directed ventrally; ratio of male cercus length to S10 maximum length in lateral view 0.88–1.0; ratio of male cercus length to paraproct length in lateral view 0.51–0.55; male paraproct in lateral view ( Fig. 12y ) with a marked ventral kink at about mid-length, of about uniform width along distal half; paraproct medial surface with a marked longitudinal crest on inner surface at apex, ending on a single apical tooth directed medially.

Dimensions. Males (n 10): Hw 20.9 ± 0.4 [20.3–21.8]; abdomen 35 ± 1.1 [33.6–37]; total length 41.9 ± 1.2 [40.2–44.1]. Females (n 10): Hw 22.9 ± 0.7 [22–24.1]; abdomen 35.5 ± 1.5 [33.2–37.5]; total length 42.6 ± 1.6 [40–44.7].

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. quadricornis shares only with M. foreli and M. turbinata the antero-dorsal margin of middle lobe of pronotum forming a distinct crest. It differs from M. turbinata by the crest being entire ( Fig. 5y ; bilobate in M. turbinata , Fig. 5 ae), distal segment of genital ligula lacking flagella ( Fig. 8y ; with long coiled paired flagella in M. turbinata , Fig. 8 ae), and male cercus distinctly shorter than paraproct ( Fig. 12y ; slightly longer than paraproct in M. turbinata , Fig. 12 ae). It differs from M. foreli by pronotum laterally smooth ( Fig. 4y ; with a prominent rounded tubercle and a longitudinal crest on middle lobe in M. foreli , Fig. 4h ), posterior lobe of female pronotum deeply trilobed with lateral lobes rounded and longer than medial lobe ( Fig. 5y ; slightly trilobed with lateral lobes shorter than medial lobe in M. foreli , Fig. 5h ), distal segment of genital ligula lacking a sclerotized pointed tubercle at base and lacking an ectal fold ( Fig. 8y ; with a sclerotized pointed tubercle (p.t.) at base and a wide ectal fold (e.f.) in M. foreli , Fig. 8h ), male cercus sub-cylindrical, gradually narrowing distally, with tip curved ventrally forming a hook and male paraproct curved ventrally at an angle, with tip pointed in lateral view ( Fig. 12y ; cercus basal half globose, distal half sub-cylindrical, with only extreme tip pointed ventrally, and paraproct curved ventrally gradually, with distal end rounded in lateral view in M. foreli , Figs. 12h ).

All females of M. quadricornis are dimorphic in shape of pronotal hind lobe, and can share long horns with males (about 67%) or have short horns (about 33%).

Remarks. Selys (1877) described the female pronotum of M. quadricornis as follows (translated from the French): ‘the posterior margin almost straight and each of its sides with a flattened horn, upright and leaning towards the anterior thorax’, which probably reflects Selys name choice for this species. Unfortunately, his description lacked illustrations, and his ‘flattened horns’ refer actually to the lateral lobes of the pronotum posterior lobe ( Figs. 4y iii–vii). This led all later workers who relied solely on the original description, to expect four horns on the thorax ( Cumming 1954; Rácenis 1955; Heckman 2008), and contributed to the idea this species was represented only by the holotype until very recently.

Based on similarity of a male of M. quadricornis collected in Venezuela to the description of the holotype of M. brysonima, Rácenis (1955) described the females collected with the male as M. brysonima . As male cerci and color pattern in M. brysonima and M. quadricornis are very similar, and he did not examine type material and relied on original descriptions for his identifications, unfortunately he misassociated these two names. After examining the holotype female of M. quadricornis and the holotype male of M. brysonima , as well as pairs of males and females of both species, I conclude that the species Rácenis (1955) identified as M. brysonima and for which he described and illustrated the female (figs. 2d, mislabeled as 2c as stated by De Marmels 1989: 32) is in fact M. quadricornis . De Marmels (1989, 1992a, b) followed Rácenis (1955) species concepts and treated M. quadricornis as M. brysonima . Belle (2002), probably based on the mislabeled figure of M. brysonima (= M. quadricornis ) as M. weibezahni in Rácenis (1955: figs. 2d mislabeled as 2c), recorded M. weibezahni from Surinam based on misidentified specimens of M. quadricornis , which I confirmed through specimens with his identification labels in FSCA, RWG, TWD, and RMNH.

Habitat. Forest near rivers and creeks.

Distribution. Venezuela, Surinam, French Guiana, and N Brazil ( Fig. 14b).

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.