Metaleptobasis Calvert, 1907

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 : 7-17

Metaleptobasis Calvert, 1907

Type species: Metaleptobasis bovilla Calvert, 1907: 386 , by original designation.

Other species included: Metaleptobasis amazonica Sjöstedt, 1918 ; M. bicornis ( Selys, 1877) [= M. mauritia Williamson, 1915 syn. nov.]; M. brevicauda sp. nov.; M. brysonima Williamson, 1915 [= M. tetragena Calvert, 1948 syn. nov.; = M. weibezahni Rácenis, 1955 syn. nov.; = M. incisula De Marmels, 1989 syn. nov.]; M. diceras ( Selys, 1877) [= M. manicaria Williamson, 1915 ; = M. fernandezi Rácenis, 1955 syn. nov.]; M. falcifera sp. nov.; M. foreli Ris, 1918 [= M. westfalli Cumming, 1954 syn. nov.]; M. furcifera sp. nov.; M. gabrielae sp. nov.; M. gibbosa Tennessen, 2012 ; M. guillermoi sp. nov.; M. inermis sp. nov.; M. knopfi Tennessen, 2012 ; M. leniloba sp. nov.; M. longicauda sp. nov.; M. lillianae Daigle, 2004 ; M. mauffrayi Daigle, 2000 ; M. minteri Daigle, 2003 ; M. orthogonia sp. nov.; M. paludicola sp. nov.; M. panguanae sp. nov.; M. peltata sp. nov.; M. prostrata sp. nov.; M. quadricornis ( Selys, 1877) ; M. selysi Santos, 1956 ; M. silvicola sp. nov.; M. spatulata sp. nov.; M. tridentigera sp. nov.; M. truncata sp. nov.; M. turbinata sp. nov.

Generic characterization. Medium to large sized coenagrionids (total length 38–53; Hw length 19–29; abdomen length 32–44). Head pale orange or yellow to pale or dark blue or green, variable intraspecifically and usually paler in females than in males, dorsum with some black markings to mostly black ( Figs. 1–2, 13); black areas usually with metallic reflections; frons angulate. Pronotum pale orange, sometimes with a dark spot on medial area of middle and/or posterior lobes, of variable extension to entirely absent in the same species, bearing a variety of diagnostic lobes, tubercles, and/or crests on anterior and/or middle lobes ( Figs. 4–5); middle lobe with a shallow medio-dorsal depression on each side, and posterior lobe posterior margin of various shapes, from entire to trilobed with lobes inflated and corners strongly projected postero-dorsally ( Figs. 4–5). Pterothorax usually pale orange, but it can also be yellowish, brownish, greenish or bluish in life ( Fig. 13), with venter pale yellow and bearing powdery white pruinescence in both males and females, and dark areas limited to black spurs on legs, reddish brown apical 0.25–0.33 of pretarsus, an oval black spot on prosternellum, a small black spot on dorsoanterior corner of mesepimeron and metepimeron, on dorso-anterior corner of metepimeron postero-dorsal triangular section, and mid-dorsal black stripe, with green, bluish, or purple metallic reflections, the particular color of the metallic shine being variable within the same species, probably according to preservation ( Figs. 2–3, 13). Anterior portion of mesanepisterna with a pair of smooth horns, in several females shorter than in conspecific males or reduced to bases or low protuberances ( Figs. 4–5). Metatibial spurs shorter than twice intervening spaces; pretarsus with vestigial supplementary tooth represented by an obtuse low prominence, which may be very small but is always visible under high magnification. Fw CuA extending for more than six cells distal to vein descending from subnodus; CuP linking CuA to CuP&AA and not reaching posterior wing margin; vein descending from quadrangle forming an unbroken line to wing margin (v.q., Fig. 6). Abdomen orange to yellow, light blue, or green, with brown to black areas as follows ( Fig. 13): S1 with a brown to reddish brown dorso-posterior spot and ventral carina of lateral tergum usually margined with brown or black; S2–7 with reddish-brown to black dorsum, with a basal pale incomplete ring on anterior edge interrupted by dorso-longitudinal dark line, and with a small diffuse pale transverse subapical spot near posterior margin on each side, which can be present or absent on some or all segments, on S3–6 dark dorsal marking posterior to pale transverse spot usually extending ventrally along sides of lateral terga; dorsum of S8–10 ranging from dark reddish brown to yellow or reddish-orange; antero-basal corner of female S8 with a black spot; medio-longitudinal carina on sterna yellow to brown to black; S2–9 with denticles along postero-lateral margin, very small on S2–6, larger on S7–9. Genital ligula distal segment lacking inner fold and lateral lobes, with a membranous terminal fold of variable length, its apparent length related to degree of hydration at preservation, sometimes barely visible but always present under high magnification (t in Fig 8). Posterior margin of female sternum S8 ( Fig. 9) smooth, with some denticles or small spines, or with strong spines or a bifid or trifid process. Male cercus entire and sub-cylindrical in lateral view; male paraproct entire, subcylindrical and directed dorso-posteriorly in lateral view, with a wide squarish base in ventral view ( Figs. 10–12). Ovipositor valves smooth, bearing only some sparse long hairs along ventral margin, surpassing posterior margin of S10 and reaching level between paraproct tip to slightly longer than level of cercus tip ( Fig. 9). Larva unknown.

Generic diagnosis. Among New World coenagrionids, Metaleptobasis shares the vein descending from quadrangle forming an unbroken line to wing margin (v.q., Fig. 6) with only four other genera: Aceratobasis , Calvertagrion St. Quentin , Mesoleptobasis , and Tukanobasis Machado. It differs from all four by the pair of smooth horns on anterior portion of mesepisterna present in most males and some females, and when reduced, represented by at least a pair of low protuberances ( Figs. 4–5), and by its pterothoracic color pattern, pale with dark color limited to a mid-dorsal stripe ( Figs. 3, 13); in the other four genera the anterior portion of mesanepisterna is smooth, and the pterothorax either lacks dark stripes ( Mesoleptobasis ) or includes additional dark stripes or areas (other three genera). Among other characters, it further differs from Aceratobasis by head dorsum including at least some small pale areas ( Fig. 1) and postero-dorsal margin of S9 ( Fig. 9) with denticles (dorsum entirely black and denticles on postero-dorsal margin of S9 absent in Aceratobasis ), from Calvertagrion by supplementary tooth of pretarsal claw vestigial and three cells between quadrangle and vein descending from subnodus in Hw (well developed pretarsal tooth and two cells between quadrangle and vein descending from subnodus in Hw in Calvertagrion ), from Mesoleptobasis by its angulate frons, pterothoracic dark mid-dorsal stripe, and Fw CuA extending for seven or more cells distal to vein descending from subnodus (frons rounded, no dark mid-dorsal stripe, and Fw CuA extending for six or less cells distal to vein descending from subnodus in Mesoleptobasis ), and from Tukanobasis by head dorsum including some pale areas, frons angulate, venter of pterothorax flat, and genital ligula lacking inner fold and lateral lobes (dorsum entirely black, frons rounded, venter of pterothorax with a mound-like medial tubercle, and genital ligula with inner fold and lateral lobes in Tukanobasis ).

Characters analyzed. Characters listed below were found to be useful for species recognition. Character states used in species diagnoses and descriptions throughout the text, with variability noted and illustrated when observed, are as follows:

Head.—Labrum color: mostly pale ( Figs. 1a–b, d –ae); mostly black ( Fig. 1c).

Extension of black on dorsum of head: limited to isolated spots and stripes covering less than 0.50 of dorsum ( Figs. 1a–b, d, f, m–q, s, x, z –aa, ad); extensive, covering about 0.50 to most of dorsum ( Figs. 1c, e, g–l, r, t–w, y, ab–ac, ae).

Postocular lobes: rounded ( Figs. 1a–b, g–h, j–p, r–t, v–x, ab–ac, ae) to slightly or markedly angled medially ( Figs. 1c–f, i, q, u, y –aa, ad); in some species intermediates between two states occur, i.e. rounded to slightly angled in M. inermis and M. paludicola , and slightly to markedly angled in M. selysi .

Thorax.—Pronotum anterior lobe dorsally [NOTE: to properly see the ultrastructures of pronotum high microscopy magnification (at least 50 x) and good illumination are necessary, since small relief structures usually have the same color as the surface and are not necessarily evident; softening of specimens to rotate the head might also be necessary in order to properly view these structures]: inflated laterally (in., Figs. 4z, 5z); with medio-lateral portions of posterior margin with a marginal crest (cr., Fig. 4b); with a postero-lateral projection on each side, free (po., Fig. 4h iii–iv), or fused to middle lobe of pronotum (p-l., Fig. 4a); with lateral margin projected posteroventrally forming a smooth L-shaped ridge (l-v. p., Figs. 4b, x); with lateral margin projected ventrally forming a denticulated ridge ending on a rounded tip (d.r., Fig. 4k ii); smooth ( Figs. 4c–g, h i –ii, i–j, k i, l–p, r–w, y–ae).

Anterior and middle lobes of pronotum dorso-laterally (best seen in dorso-lateral view examined from outer side): separated by an evident groove (g., Figs. 5a–o, r–t, v–z, ab–ac, ae); almost touching, separated by a narrow fissure (f., Figs. 5q, u, aa, ad).

Propleuron: smooth ( Figs. 5c, g, t, w, ac); with a denticulated anterior oblique crest (c., Fig. 5a); with a longitudinal crest (lo., Figs. 5h, n, y–z); with sub-vertical crest/s or ridge/s (l.c., Figs. 5f, o–q, aa, ad); with a cshaped crest (c., Figs. 5e, i, m, u, y); with a rounded tubercle or plate, low or prominent (t., Figs. 5b, d–e, h–l, n, r, u–v, x–z, ab, ae).

Anterior margin of middle lobe of pronotum: smooth ( Figs. 4a–g, i–x, z –ad); forming a crest (cr.), which can be entire ( Figs. 4h, y; 5h, y) or bilobate ( Fig. 4 ae; 5ae).

Dorsum of middle lobe of pronotum: with a latero-dorsal c-shaped longitudinal depression (de., Fig. 5c); lacking a latero-dorsal c-shaped longitudinal depression ( Figs. 5. a–b, d –ae).

Paired ear-lobe projections on anterior portion of middle lobe of pronotum: digit like or tongue-shaped, oriented laterally or postero-laterally ( Figs. 4a iii, g, t, v–w, ac); flap-like with a dorsal concavity, oriented dorso-ventrally ( Fig. 4l); flap-like and upright, wide and sinuous ( Fig. 4r iii–iv, vi, viii–ix) or narrow ( Fig. 4r v, vii); absent ( Figs. 4b–f, h–k, m–q, s, u, x –ab, ad–ae).

Shape of pronotum posterior lobe: entire, smoothly convex ( Figs. 4k, l i, s iii, y ii, ab iii–iv, vi) or with a medial point ( Figs. 4l ii–iii, ab x); bilobed, with a slight medial concavity shallower than 0.30 of posterior lobe length ( Figs. 4b xiv, d iv, f iv–vii, p viii–x), with a medial u- or v-shaped incision about as deep or deeper than 0.50 of posterior lobe length ( Figs. 4e v –xiii, i iii–vi), and with lateral lobes entire ( Figs. 4b xiv, d iv, e vi–x, i iii–vi, p viii, x) or bilobed ( Figs. 4b vi, e v, ix, xi–xiii, f iv–vii, p ix); trilobed, with medial lobe bilobed ( Figs. 4b iii, vii–viii, x–xi, g i, p v, s ii, iv–v, u iii, x v, y iii, vi–vii, ab i–ii, ix, ac iv), smoothly convex ( Figs. 4a–b i –ii, iv–v, ix, xii–xiii, c v, d ii–iii, f i–iii, viii–ix, g ii, h ii–iv, j, m–o, p ii–vi, xi, q i, r i–iii, s i, vi–viii, t, u i, iv, vi, y i, iv–v, z i–iii, aa i, iii, ab v, vii–viii, ac i–iii, ad i–ii, ae), with a central point ( Figs. 4c i, iii, d i, e i–iv, h i, i i–ii, p i, vii, q ii, r iv–ix, u ii, v, v–w, x i–ii, z iv, aa ii, ad iii), with a central posterior or postero-dorsal projection ( Figs. 4c vi–vii, r iv–v, ix), and lateral lobes entire ( Figs. 4a i –ii, b i–v, vii–xiii, c v–vii, f i–iii, h, i i–ii, m–o, p ii–vi, xi, r, t i–iii, v–ad, ae i) or bilobed ( Figs. 4a iii, t iv, ae ii) [in some species shape is variable, especially in females, presenting two or more possible states].

Lateral lobes: shorter ( Figs. 4a–b, c vi–vii, e iii, v, ix, xi–xiii, f–g, h i, i i, iii–iv, j–m, n i–ii, o i, iii–iv, p i–vii, q i, r–x, y i–ii, z, ab i–v, vii–viii, x, ac, ad i–ii, ae i), as long as (4c i, iii, v, e i–ii, iv, vi–viii, x, h ii–iv, i ii, v–vi, n iii, o ii, p viii, q ii, y, ab vi, ix), or longer ( Figs. 4y iii–vii, ad iii, ae ii) than medial lobe or central area; smoothly convex ( Figs. 4a–f, h–v, w i, x –ab, ac i–iii, ad–ae) or dorsally globose ( Figs. 4g ii, w ii, ac iv); not projected ( Figs. 4a–f, g i, h–i, j i, k, l i, m–q, s, t i –ii, u, v i, w–ae) or projected dorsally ( Figs. 4g ii, j ii, l ii–iii, t iii–iv, v ii).

Mesanepisternal horns ( Figs. 4–5) length: about as long as 0.30–0.50 of mesostigmal plate width ( Figs. 4a ii, b vi–viii, c iii–iv, f vi–ix, g ii, h iii, j ii–iii, k ii, m i, q ii, r ii–iii, s i–iii, t iii, v ii, y iv–v, aa ii,) to well developed, from about equal to mesostigmal plate width to as long as 4–5 times mesostigmal plate width [even though length can be variable range of values can aid in diagnosis].

Relative width of mesanepisternal horns (width can be variable, but approximate ranges can aid in diagnosis): thin ( Figs. 4d–e, g–k, n, s–t, v–x, y iii–v, z iii–iv, ac i) to thick ( Figs. 4c).

Orientation of free portion of mesanepisternal horns in dorsal view (orientation can be variable, but approximate ranges can aid in diagnosis): lateral ( Figs. 4p, s) to antero-lateral ( Figs. 4a, b vi–vii, xiii–xiv, d, e v–xiii, f–g, i iii, vi, j, l i, n i, o ii–iii, q–r, t, v, x, z iii–iv, aa, ab i–iv, ac ii–iii, ad–ae), antero-dorsal ( Figs. 4b i –iv, e i–ii, k, l ii, n ii, o, v, u, w, y, z, ab v–x, ac i), and dorsal ( Figs. 4h).

i, iv

i–ii

Angle of mesanepisternal horns to mesepisternal surface in lateral view (angle can be variable, but range of values can aid in diagnosis): parallel to surface ( Figs. 5b vii–vii, d, p, s) or at an angle with surface.

Bases of mesanepisternal horns (both states can occur in some species, i.e. M. bicornis , M. leniloba ): adjacent or almost touching ( Figs. 4e–f, p–q, aa, ad) to separated.

Shape of tip of mesanepisternal horns: rounded ( Figs. 4a–b, l–m, n ii–iii, z); pointed or bluntly pointed ( Figs. 4c– d, e–h, n i, o–y, ac); with a transverse dorsal slit ( Fig. 4c v); antero-posteriorly compressed ( Figs. 4 ab, ae).

Mid-dorsal dark stripe ( Fig. 3): parallel sided; widening medially; widening posteriorly; narrowing posteriorly.

Pt shape: rectangular, with anterior and posterior sides longer than distal side ( Fig. 6a ii–iii); squarish, with anterior and posterior sides about equal to distal side; trapezoidal, with anterior side shorter than distal side ( Figs. 6b, c) [in some species intermediates between two states occur].

Abdomen.—Male genital lobe height: distinctly shorter than 0.50 of anterior hamule height ( Figs. 8a–f, h–v, x – ab, ad–ae); about as high as 0.50 of anterior hamule height and shorter than 0.50 of genital lobe’s basal width ( Fig. 8g, ac); about as high as anterior hamule and as high as 0.50 of genital lobe’s basal width ( Fig. 8w).

Male genital lobe contour: smoothly curved ( Figs. 8a–s, u –ae); with a distinct anterior angle ( Fig. 8t).

Male posterior hamule: digit-like and small, with at most only tip surpassing ventral margin of genital fossa in lateral view ( Figs. 8a–f, h–i, m–q, s, u, x –ab, ad–ae); laminar and large, clearly surpassing ventral margin of genital fossa in lateral view ( Figs. 8g, j–l, r, t, v–w, ac), and shorter than ( Figs. 8j–l, v) or as high as or higher than anterior hamule ( Figs. 8g, r, t, w, ac), with posterior side smoothly convex ( Figs. 8g, j–l, r, w, ac) or angled ( Figs. 8t, v).

Tip of posterior hamule: not differentiated ( Figs. 8a–f, h–i, m–q, s, u, x –ab, ad–ae); bluntly pointed ( Figs. 8j–l, r, v); or bent at about 90° and triangular ( Figs. 8g, t, ac) or rounded ( Figs. 8w).

Curvature of basal segment of genital ligula: marked by a slight depression followed by a slight convex prominence ( Figs. 8a–f, h–i, m–q, s, u, x –ab, ad–ae); marked by a slight depression followed by a pronounced prominence ( Fig. 8c ii), with tip of ligula laying between depression and prominence when folded ( Fig. 8c i); marked by a deep depression, followed by a pronounced prominence ( Figs. 8g, j–l, r, t, v–w, ac), with tip of ligula laying in the depression when folded. Shape of genital ligula distal segment: sub-quadrate ( Figs. 8a, c, m, o); sub-rectangular ( Figs. 8b, d, k–l, n, p–s, x, z –aa, ad–ae); pear-shaped, distinctly widened sub-apically ( Figs. 8e–f, g–j, t–w, y, ab– ac).

Terminal fold (ectal fold at base of genital ligula distal segment): short to long (t., Fig. 8) [the length of this fold can be variable according to the degree of hydration of the membrane, which can extend considerably when placed in water (i.e. Figs. 8c i – ii), and should therefore not be used as a diagnostic feature].

Base of genital ligula distal segment: with a lateral sclerotized pointed tubercle (p.t., Fig. 8h); smooth ( Figs. 8a–g, i – ae).

Shape of apex of genital ligula distal segment: convex ( Figs. 8c, h); transverse or with a shallow concavity ( Figs. 8a–b, d–g, i–r, t –aa, ac–ad); with a deep concavity or cleft, about as deep as wide or deeper ( Figs. 8s, ab); with lateral corners projected into long coiled flagella ( Fig. 8 ae).

Ectal fold on apex of genital ligula distal segment: present, narrow ( Figs. 8b, f–g, j–l, p, r, t–w) or wide ( Figs. 8c, h); absent ( Figs. 8a, i, m–o, q, s, x –ae).

Posterior margin of female S8 sternum: smooth, lacking any denticles, spines or processes ( Figs. 9a–f, h–i, m–p, r–s, w–z, ab–ac); with 1 to 6 denticles or small spines ( Figs. 9k–l, q); with strong spines ( Figs. 9j, u); with a two ( Figs. 9g, t, v) or three ( Fig. 9 aa) pronged process, which can be parallel to the sternum ( Figs. 9t, aa), or to the sides of the lateral terga ( Figs. 9g, v).

Distal end of ovipositor reaching ( Fig. 9): apex of paraproct to level slightly distal to cercus [length of ovipositor is variable].

Male S10 postero-dorsal margin: with a small medial u-shaped incision or longitudinal slit ( Figs. 10a–b, d–f, i, m–q, s, u, x –aa, ad); smoothly convex ( Figs. 10c, g–h, j–l, r, t, v–w, ab–ac, ae).

Male S10 postero-dorsal surface: with a medial prominence, which can be entire ( Figs. 11a–d, f–g, j, l–r, t–x, z – ae; 12a–d, f–g, j, l–r, t–x, z–ae) or divided in two lateral portions ( Figs. 11k; 12k); lacking a prominence ( Figs. 11e, h–i, s, y; 12e, h–i, s, y).

Male S10 postero-dorsal margin: with a medial projection ( Figs. 10a–b, d–j, l–r, t –ad; 11a–b, d–j, l–r, t–ad); lacking a projection ( Figs. 10c, k, s, ae; 11c, k, s, ae).

Male cercus shape in dorsal view: curved medially ( Figs. 10a–d, f–g, j–o, q–x, z –ae); about straight ( Figs. 10e, h–i, p, y).

Male cercus width in dorsal view: about uniform ( Figs. 10q, r, t, w, z, ad–ae); narrowing to tip ( Figs. 10a, c–e, h–i, m–p, s, u–v, x–y, aa–ac); widening sub-apically ( Figs. 10b, f–g, j–l).

Shape of male cercus tip: pointed ( Figs. 10a–e, g–n, p, r–y, aa–ac); rounded or blunt ( Figs. 10f, o, q, z, ad–ae).

Ratio of male cercus length to S10 maximum length in lateral view ( Fig. 12): from about 0.5 to about 2.

Ratio of male cercus length to paraproct length in lateral view ( Fig. 12): from about 0.5 to about 1.5.

Width of male paraproct distal third in lateral view: (0) narrower than medial third ( Figs. 12a, c, m–n, q–r, z); about as wide as medial third ( Figs. 12b, h, k–l, o, s, x–y, ab, ae); wider than medial third ( Figs. 12d–g, i–j, p, t–w, aa, ac–ad).

Male paraproct tip: with smooth medial surface and no apical tooth ( Figs. 11 aa, ae); with smooth medial surface and ending on an apical tooth ( Figs. 11a, c, g–h, j–k, m, o, q–w, y, aa, ac–ad); with a ridge on medial surface ending on an apical tooth ( Figs. 11b, d–f, n, x, z); with a ridge on medial surface with basal and apical teeth ( Figs. 11i, l, p).

Remarks. Several male characters are species specific: shape of genital ligula distal segment, posterior hamule in species where it is well developed, dorsum of S10, cerci, and paraprocts. Conspecific males and females share the same ultrastructural features of the pronotal and propleural surface (presence and position of various crests, ridges, tubercles, and projections), the position of the bases of the mesanepisternal horns, and the basic black pattern of head and width and shape of dark mid-dorsal stripe of pterothorax. However, the latter two characters show considerable intraspecific plasticity in a few species, i.e. the extension of the black pattern on head dorsum observed in M. paludicola and M. spatulata ( Figs. 1u, ab) and on the back of the head in M. minteri , and the width of the mid-dorsal stripe in M. minteri ( Fig. 3s) vary considerably even among specimens within the same population.

The shape of the posterior lobe of the pronotum and the exact length, width, orientation, and shape of mesanepisternal horns show considerable intraspecific variability, especially in females. In addition, in females one or both of these structures can be similar to those of conspecific males (species with no secondary sexual dimorphism: M. leniloba , M. minteri , and M. paludicola ) or different (species with secondary female dimorphism: M. bovilla , M. brysonima , M. falcifera , M. furcifera , M. gabrielae , M. gibbosa , M. knopfi , M. lillianae , M. mauffrayi , M. longicauda , M. panguanae , M. peltata , M. quadricornis , M. selysi , M. tridentigera , M. truncata , and M. turbinata ), and several species include both dimorphic and not dimorphic females (species with andromorphic and heteromorphic females: M. bicornis , M. brevicauda , M. diceras , M. foreli , M. guillermoi , M. orthogonia , M. silvicola , and M. spatulata ). Female dimorphism can include just the mesanepisternal horns (with shape of posterior lobe of pronotum the same as in male: M. amazonica , M. foreli , M. knopfi , M. truncata ), just the shape of posterior lobe of pronotum (with mesanepisternal horns similar to those of male: M. brevicauda , M. lillianae , M. prostrata ), both mesanepisternal horns and/or posterior lobe of pronotum ( M. bicornis , M. diceras , M. guillermoi , M. orthogonia , M. spatulata , M. truncata ), or involve other structures of pronotum (i.e. antero-dorsal ear-like lobes on middle lobe of pronotum in M. mauffrayi ). Dimorphic females may display more than one distinct morph in terms of the characters in which they differ from their males (i.e. the upright ear-like lobe on dorsum of middle lobe of pronotum in females of M. mauffrayi , absent in males, can be either narrow or wide, with no gradation between these two states). Species that were represented by small series of available females in this study (see Table 2) might be found to present more morphs when more specimens become available in the future.

I was able to reliably confirm conspecifity of males and dimorphic or polymorphic females in species for which pairs in copula and/or tandem were available, from which I was able to identify diagnostic characters shared by males and females, i.e. ultrastructural features on propleural and pronotal surface, position of bases of mesanepisternal horns, basic black pattern of head, and width and shape of dark mid-dorsal stripe of pterothorax. Conspecificity was reinforced by the examination of series of specimens from the same locality that included males and different female morphs. After discovering the existence of female dimorphism and polymorphism in some species and recognizing which characters of diagnostic value are shared by conspecific males and females, I was accordingly able to associate males and females of species not found in copula but which share states of these diagnostic characters.

structures. Polymorphic species including both andromorphic and heteromorphic females are indicated in bold, and dimorphic species including more than one distinct female morph are underscored.

Key to species of Metaleptobasis

1. Male posterior hamule (p.h.) digit-like, small, with at most only tip surpassing ventral margin of genital fossa in lateral view ( Figs. 8a–f, h–i, m–q, s, u, x –ab, ad–ae); curvature of basal segment of genital ligula marked by a slight concave depression (s.d., Figs. 8a–f, h–i, m–q, s, u, x –ab, ad–ae); posterior margin of female S8 sternum smooth ( Figs. 9a–f, h–i, m–p, r–s, w–z, aa, ac–ad)........................................................................................ 2

1’. Male posterior hamule (p.h.) laminar and curved posteriorly or postero-medially, large, clearly surpassing ventral margin of genital fossa in lateral view ( Figs. 8g, j–l, r, t, v–w, ac); curvature of basal segment of genital ligula marked by a deep concave depression (d.d., Figs. 8g, j–l, r, t, v–w, ac); posterior margin of female S8 sternum with either denticles or small spines ( Figs. 9 k-l, q), strong spines ( Figs. 9j, u), or a bifid or trifid process ( Figs. 9g, t, v, ab)............................ 23

2(1). Black on head dorsum covering half to most of dorsal surface (1c, e, g–l, r, t–w, y, ab–ac, ae)....................... 3

2’. Black on head dorsum covering less than half of dorsum, limited to isolated spots and stripes ( Figs. 1a–b, d, f, m–q, s, x, z – aa, ad) .............................................................................................10

3(2). Labrum mostly black, with only distal margin narrowly pale ( Figs. 1c); lateral surface of middle lobe of pronotum with a cshaped latero-dorsal longitudinal depression (de.), shallow in males ( Figs. 4c I, 5c I –II), deep in females ( Figs. 4c V –VI, 5 c III–IV); mesanepisternal horns thick in male ( Figs. 4c II–IV, 5c I–II), reduced to tubercles with a dorsal transverse slit in females ( Figs. 4c V, 5c III–IV); male cercus about as long as 0.33 of paraproct ( Figs. 10c, 12c); male paraproct with a dorsal swelling at basal 0.40 of its length (sw., Fig. 12c); Guatemala to Costa Rica ( Fig. 14a)........................................ M. bovilla

3’. Labrum mostly pale, with marginal black stripe along base and sides ( Figs. 1e, h–i, u, y, ab, ae); lateral surface of middle lobe of pronotum smooth ( Figs. 4e, h–i, u, y, ab, ae; 5e, h–i, u, y, ab, ae); mesanepisternal horns thin in males, and never with a dorsal transverse slit in females ( Figs. 4e, h–i, u, y, ab, ae; 5e, h–i, u, y, ab, ae); male cercus about as long as 0.50 of paraproct or longer; male paraproct lacking a dorsal swelling at basal 0.40 of its length ( Figs. 12e, h–i, u, y, ab, ae)........ 4

4(3’). Anterior margin of middle lobe of pronotum forming a distinct crest (cr., Figs. 4h, y, ae; 5h, y, ae)............... 5

4’. Anterior margin of middle lobe of pronotum smooth ( Figs. 4e, i, u, ab; 5e, i, u, ab)............................. 7

5(4’). Black on head dorsum covering about half of dorsal surface ( Figs. 1 ae); crest on anterior margin of middle lobe of pronotum bilobate ( Figs. 4 ae; 5ae); distal segment of genital ligula with long paired coiled flagella (fl., Fig. 8 ae); N Peru ( Fig. 14a)..................................................................................... M. turbinata

5’. Black on head dorsum covering most of dorsal surface ( Figs. 1h, y); crest on anterior margin of middle lobe of pronotum entire (cr., Figs. 4h, y; 5h, y); distal segment of genital ligula lacking flagella ( Figs. 8h, y)........................ 6

6(4’). Anterior lobe of female pronotum with a lateral digit-shaped projection directed posteriorly (po., Figs. 4h III–IV, 5h V), rarely reduced ( Fig. 5h VI), absent (as in Fig. 5h I, IV), or fused to middle-lobe of pronotum ( Fig. 5h VII); posterior lobe of female pronotum slightly trilobed with lateral lobes shorter than medial lobe length ( Figs. 4h III–IV); distal segment of genital ligula with a pair of sclerotized pointed tubercles at base (p.t.) and with a wide ectal fold (e.f., Fig. 8h); male cercus basal half globose, distal half sub-cylindrical, with only extreme tip pointed ventrally; male paraproct curved ventrally gradually, with distal end rounded in lateral view ( Figs. 12h I –II); Costa Rica to N Colombia and Venezuela ( Fig. 14b).................... M. foreli

6’. Anterior lobe of female pronotum smooth ( Figs. 4y IV, 5y IV–VII); posterior lobe of female pronotum deeply trilobed with lateral lobes rounded and longer than medial lobe length ( Figs. 4y III–VII); distal segment of genital ligula lacking paired sclerotized pointed tubercles at base and lacking an ectal fold ( Fig. 8y); male cercus sub-cylindrical, gradually narrowing distally, with tip curved ventrally forming a hook; male paraproct curved ventrally at an angle, with tip pointed in lateral view ( Figs. 12y I –II); Venezuela, Surinam, French Guiana, and N Brazil ( Fig. 14b)................................... M. quadricornis

7(4’). Propleuron with a short triangular anterior laminar ridge (t., Fig. 5 ab); mesanepisternal horns wide at base and compressed antero-posteriorly ( Figs. 4 ab I–VI,VIII, 5ab I–II,IV–V) or reduced to low blunt prominences ( Figs. 4 ab VII, 5ab III); Ecuador, Peru, and W Brazil ( Fig. 14a)......................................................................... M. spatulata

7’. Propleuron lacking a laminar ridge, with an anterior rounded tubercle (t., Figs. 4e, i, u; 5e, i, u); mesanepisternal horns sub-cylindrical ( Figs. 4e, i, u; 5e, i, u).................................................................. 8

8(7’). Female pronotum trilobed with smoothly convex medial lobe ( Figs. 4u V –VI); dark mid-dorsal pterothoracic stripe about as wide as 0.33 of mesepisterna width ( Fig. 3u); male S 10 in lateral view with a well developed medial prominence on postero-dorsal margin ( Fig. 12u); Amazon forest of N Peru and N Brazil ( Fig. 14e).............................. M. paludicola

8’. Female pronotum unlobed to slightly trilobed always with a pronounced medial u or v-shaped incision ( Figs. 4e V –XIII, i III–VI); dark mid-dorsal pterothoracic stripe about as wide as 0.50 of mesepisterna width ( Figs. 3e, i); male S 10 in lateral view flat, lacking a medial prominence on postero-dorsal margin ( Figs. 12e, i).......................................... 9

9(8’). Apex of male genital ligula distal segment with a u-shaped incision ( Fig. 8e), embracing sides of basal segment when ligula is folded; tip of male paraproct with a poorly developed longitudinal crest on inner surface, terminating as a medially directed single apical tooth (s.t., Figs. 10e, 11e); Trinidad and Venezuela to E Peru and N Bolivia ( Fig. 14b)....... M. brysonima

9’ Apex of male genital ligula distal segment transverse ( Fig. 8i), only touching ventral surface of basal segment without extending over its sides when ligula is folded; tip of male paraproct with a well developed longitudinal crest on inner surface, with two medially directed teeth (te.), one on its basal end (may be small) and one on its apical end ( Figs. 10i, 11i); N Peru ( Fig. 14b)......................................................................... M. furcifera

10(2’). Middle lobe of pronotum with an antero-dorsal rounded thumb-like process (e-l., Figs. 4a; 5a) directed laterally on each side; Pará State in Brazil ( Fig. 14e)......................................................... M. amazonica

10’. Middle lobe of pronotum lacking an antero-dorsal thumb-like process directed laterally on each side ( Figs. 4b, d, f, m–q, s, x, z –aa, ad; 5b, d, f, m–q, s, x, z–aa, ad).............................................................. 11

11(10’). Lateral margin of anterior lobe of pronotum extended postero-ventrally forming a ridge (l-v. p.) opposite an anterior tubercle (t.) on propleuron ( Figs. 4b, x; 5b, x)............................................................ 12

11’. Lateral margin of anterior lobe of pronotum not extended postero-ventrally forming a ridge ( Figs. 4d, f, m–q, s, z –aa, ad; 5d, f, m–q, s, z–aa, ad)......................................................................... 13

12(11). Lateral portions of posterior margin of pronotum anterior lobe with a marginal crest (cr., Fig. 4b); mesanepisternal horns oriented antero-dorsally at an angle of about 45° with dorsum ( Figs. 5b I–V,IX–X) or represented by bases oriented laterally ( Figs. 5b VI–VIII); distal portion of male cercus depressed dorso-ventrally in lateral view ( Fig. 12b) and slightly widened sub-apically in dorsal view ( Fig. 10b); Trinidad, Venezuela, Guyana, Surinam, French Guiana, and N Brazil ( Fig. 14c)... M. bicornis

12’. Lateral portions of posterior margin of pronotum anterior lobe lacking a crest ( Fig. 4x); mesanepisternal horns directed antero-laterally at an angle of about 15° with dorsum ( Fig. 5x); distal portion of male cercus sub-cylindrical in lateral view ( Fig. 12x) and gradually narrowing distally in dorsal view ( Fig. 10x); Junín Dep. in Peru ( Fig. 14c)....... M. prostrata

13(11’). Mesanepisternal horns directed antero-laterally or laterally forming an angle of 0°–15° with dorsum ( Figs. 4d, p, s; 5d, p, s), and if limited to horn bases these arising at level of mesostigmal plates bases or more externally ( Figs. 4s I, III,V,VIII)...... 14

13’. Mesanepisternal horns directed antero-dorsally forming an angle of 35°–90° with dorsum ( Figs. 4f, m–o, q, z –aa, ad; 5f, m– o, q, z–aa, ad), and if limited to horn bases these arising medially to level of mesostigmal plates bases ( Figs. 4f IV–VIII, m, q II, aa III, ad II–III)....................................................................................... 16

14(11). Mesanepisternal horns distinctly separated at bases, arising at level of base of mesostigmal plates or more externally ( Fig. 4s); male S10 lacking a postero-dorsal prominence and a postero-medial projection ( Fig. 12s); male paraproct sub-apically about as wide as at medial third ( Fig. 12s) and lacking an apical ridge on medial surface ( Figs. 10s; 11s); S Colombia, Ecuador, Peru, and W Brazil ( Fig. 14e)............................................................ M. minteri

14’. Mesanepisternal horns adjacent at base ( Fig. 4p) or if separated, arising medially to level of mesostigmal plates bases ( Fig. 4d); male S10 with a postero-dorsal prominence and a postero-medial projection ( Figs. 12d, p); male paraproct sub-apically wider than at medial third ( Figs. 12d, p) and with an apical ridge on medial surface (r., Figs. 10d, p; 11d, p).......... 15

15 (14’). Bases of mesanepisternal horns separated ( Fig. 4d); male paraproct shorter than 0.50 of cercus length ( Figs. 10d; 12d); Huánuco Dep. in Peru ( Fig. 14c)............................................................ M. brevicauda

15’. Bases of mesanepisternal horns adjacent ( Fig. 4p); male paraproct about as long as twice cercus length ( Figs. 10p; 12p); Ecuador, Peru, Bolivia, and W Brazil ( Fig. 14d)................................................. M. lillianae

16(15’). Anterior area of propleuron with two sub-vertical crests (l.c., Figs. 5f, q, aa, ad; CAUTION: anterior crest can be low and rounded as in Figs. 5f IV,IX).............................................................................17

16’. Anterior area of propleuron with only one crest, sub-vertical, transverse, or c-shaped (c., l.c., lo., Figs. 5m –n, o, z)..... 20

17(16’). Pt trapezoidal, with anterior side shorter than distal side ( Figs. 6b–c); tip of male cercus sub-cylindrical ( Figs. 11q, ad; 12q, ad)..............................................................................................18

17’. Pt sub-rectangular, with anterior and posterior sides longer than distal side ( Fig. 6a); tip of male cercus depressed dorso-ventrally ( Figs. 11f, aa; 12f, aa)......................................................................... 19

18(17’). Mid-dorsal dark stripe on pterothorax at mid-length as wide as about 0.20 of mesepisterna width ( Fig. 3q); male cercus about as long as 2 times maximum length of S 10 in lateral view, about as long as paraproct, with tip rounded ( Fig. 12q); Mato Grosso and Goias States in Brazil ( Fig. 14d).............................................. M. longicauda

18’. Mid-dorsal dark stripe on pterothorax at mid-length about as wide as 0.25 of mesepisterna width ( Fig. 3 ad); male cercus about as long as 1.5 times maximum length of S 10 in lateral view, shorter than paraproct, with tip truncate posteriorly ( Fig. 12 ad); Pará State in Brazil ( Fig. 14d)..................................................... M. truncata

19(17’). Anterior and middle lobes of pronotum separated dorso-laterally by a groove (g., Fig. 5f); mesanepisternal horns of males and andromorphic females about as long as 1.5 times the width of a mesostigmal plate ( Figs. 4f; 5f); pterothoracic mid-dorsal stripe about as wide as 0.25 of mesepisterna width ( Fig. 3f); distal segment of genital ligula widened from base to tip ( Fig. 8f); male cercus in dorsal view slightly widened distally and constricted sub-apically ( Fig. 10f); tip of male paraproct with ventral bicuspidate tooth pointed medially (b.t.), and with an inner transverse sub-apical ridge (r., Figs. 10f; 11f); Trinidad, Venezuela, Surinam, Guyana, Brazil, and Peru ( Fig. 14d)....................................... M. diceras

19’. Anterior and middle lobes of pronotum almost touching dorsally, separated by a fissure (f., Fig. 5 aa); mesanepisternal horns of males and andromorphic females about as long as two times the width of a mesostigmal plate ( Figs. 4 aa; 5aa); pterothoracic mid-dorsal stripe about as wide as 0.18 of mesepisterna ( Fig. 3 aa); distal segment of genital ligula sub-rectangular, with sides almost straight, slightly narrowing from base to tip ( Fig. 8 aa); male cercus in dorsal view of uniform width sub-apically ( Fig. 10 aa); tip of male paraproct with a single triangular medio-ventral tooth (s.t.), and with two inner longitudinal sub-apical ridges (rs., Figs. 10 aa; 11aa); SE Peru ( Fig. 14d)................................... M. silvicola

20(16’). Latero-ventral area of middle lobe of pronotum with a short c-shaped crest, located opposite a short c-shaped crest on antero-dorsal area of propleuron (c., Fig. 5m); male mesanepisternal horns sub-triangular, about as long as 0.50 of mesostigmal plate width ( Figs. 4m; 5m); Pará State in Brazil ( Fig. 14e) [female unknown]................ M. inermis

20’. Latero-ventral area of middle lobe of pronotum lacking any crest or ridge ( Figs. 5n, o, z); male mesanepisternal horns sub-cylindrical, about as long as mesostigmal plate width or longer ( Figs. 5n, o, z)...............................21

21(20’). Pterothoracic mid-dorsal dark stripe narrower than 0.20 of mesepisterna width, slightly widening posteriorly ( Fig. 3o); anterior area of propleuron with a short comma-shaped vertical crest (l.c., Fig. 5o); distal segment of genital ligula sub-quadrate ( Fig. 8o); N Peru and N Brazil ( Fig. 14e)....................................................... M. leniloba

21’. Pterothoracic mid-dorsal dark stripe as wide or wider than 0.33 of mesepisterna width, about parallel sided ( Figs. 3n, z); anterior area of propleuron with a short longitudinal ridge (lo., Figs. 5n, z); distal segment of genital ligula sub-rectangular ( Figs. 8n, z).......................................................................................22

22(21’). Lateral margin of pronotum anterior lobe not inflated ( Figs. 4n; 5n); mesanepisternal horns thin, narrowing from base to bluntly pointed tip, forming an angle of 80°–95° with dorsum in lateral view ( Fig. 5n); male cercus in dorsal view only slightly curved medially, sub-cylindrical, gradually narrowing to tip ( Figs. 10n; 11n), in lateral view about as long as 0.50 of paraproct length ( Fig. 12n); Ecuador ( Fig. 14c).................................................... M. knopfi

22’. Lateral margin of pronotum anterior lobe inflated (in., Figs. 4z; 5z); mesanepisternal horns of medium thickness, subcylindrical with rounded tips, forming an angle of 35°–45° with dorsum in lateral view ( Fig. 5z); male cercus in dorsal view abruptly bent medially at about mid-length, with distal half about parallel-sided ( Fig. 10z), and depressed dorso-ventrally ( Figs. 11z; 12z), in lateral view distinctly longer than 0.50 of paraproct length ( Fig. 12z); E Brazil and NE Argentina ( Fig. 14c)....................................................................................... M. selysi

23(1’). Males........................................................................................... 24 23’. Females......................................................................................... 32

24(23’). Medially bent distal portion of cercus widened, wider than medial third of cercus in dorsal view ( Figs. 10j–l)........ 25

24’. Medially bent distal portion of cercus not widened, about as wide as or narrower than medial third of cercus in dorsal view ( Figs. 10g, r, t, v–w, ac)............................................................................ 27

25(24). Cercus about as long as paraproct in lateral view ( Fig. 12l); paraproct ending on apical dorsal and ventral teeth ( Figs. 10l; 11l); N Peru ( Fig. 14g)................................................................... M. guillermoi

25’. Cercus shorter than paraproct in lateral view ( Figs. 12j–k); paraproct ending on a single medio-ventral apical tooth ( Figs. 10j–k; 11j–k) .............. .. ........................... ............. ......... .. ................. 26

26(25’). Posterior hamule tip in same plane as hamule axis; axis narrow, about as wide as width of medial sclerotized portion of genital ligula basal segment in ventral view ( Fig. 8k); genital ligula distal segment sub-rectangular in ventral view ( Fig. 8k); dorsum of S10 with sub-apical prominence (pr.) divided into two lateral mounds, and lacking a postero-medial projection ( Figs. 10k; 11k); Ecuador ( Fig. 14g)........................................................ M. gibbosa

26’. Posterior hamule tip bent medially at an angle of about 90° with hamule axis; axis wide, about as wide as 1.5 times width of medial sclerotized portion of genital ligula basal segment in ventral view ( Fig. 8j); dorsum of S10 with sub-apical prominence (pr.) entire and about transverse, and with a postero-medial convex projection (p-m.p., Figs. 10j; 11j); Loreto Dep. in Peru ( Fig. 14g).................................................................... M. gabrielae

27(24’). Genital lobe (g.l.) as tall as or taller than 0.50 of anterior hamule (a.h., Figs. 8g, w, ac)........................... 28

27’. Genital lobe shorter (g.l.) than 0.50 of anterior hamule (a.h., Figs. 8r, t, v)..................................... 30

28(27). Genital lobe (g.l.) as tall as anterior hamule (a.h.), and slightly taller than 0.50 of genital lobe basal width ( Fig. 8w); N Peru ( Fig. 14f)....................................................................... M. peltata

28’. Genital lobe (g.l.) as tall as 0.50 of anterior hamule (a.h.), and shorter than 0.50 of genital lobe basal width ( Figs. 8g, ac) 29

29(28’). Male posterior hamule (p.h.) about as wide as 1.5 times width of medial sclerotized portion of genital ligula basal segment in ventral view ( Fig. 8g); distal portion of male cercus about as wide as medial portion ( Fig. 10g); S Peru and W Brazil ( Fig. 14f).................................................................................... M. falcifera

29’. Male posterior hamule (p.h.) about as wide as width of medial sclerotized portion of genital ligula basal segment in ventral view ( Fig. 8 ac); distal portion of male cercus narrower than medial portion ( Fig. 10 ac); N and W Brazil ( Fig. 14f)............................................................................................. M. tridentigera

30(27’). Posterior hamule slightly (p.h.) taller than 0.50 of anterior hamule (a.h.), with anterior margin smoothly curved and posterior margin widened into a heel, with tip curved gradually medially and rounded in ventral view ( Fig. 8v); Huánuco Dep. in Peru ( Fig. 14g)............................................................................. M. panguanae

30’. Posterior hamule (p.h.) about as tall as anterior hamule (a.h.) or slightly taller, about semicircular with both anterior and posterior margins smoothly curved and tip in same plane as axis ( Figs. 8r), or with anterior margin straight and posterior margin angled near base and hamule tip bent medially at about 90° with hamule axis and triangular in ventral view ( Figs. 8t).. 31

31(30’). Posterior hamule (p.h.) elongated crescent shaped with both anterior and posterior margins smoothly curved and laminar, with tip in same plane as axis ( Fig. 8r); distal segment of male genital ligula rectangular in ventral view ( Fig. 8r); N Peru, Ecuador, and S Colombia ( Fig. 14g)......................................................... M. mauffrayi

31’. Posterior hamule (p.h.) with posterior margin angled near base, and hamule tip bent medially at about 90° with hamule axis and triangular in ventral view ( Fig. 8t); distal segment of male genital ligula pear-shaped, distinctly widened sub-apically in ventral view ( Fig. 8t); N Peru and Ecuador ( Fig. 14f)...................................... M. orthogonia

32(23’). Posterior margin of sternum S8 with 1–6 denticles or small spines ( Figs. 9k–l, q)............................... 33

32’. Posterior margin of sternum S8 with strong spines or processes ( Figs. 9g, j, t, u–v, ab)........................... 35

33(32’). Latero-ventral corners of anterior lobe of pronotum projected ventrally forming a ridge with rounded tip (d.r., Fig. 5k II); middle lobe of pronotum smooth; hind lobe of pronotum entire, smoothly convex with lateral corners fused to middle lobe of pronotum ( Figs. 4k; 5k); Ecuador ( Fig. 14g)................................................... M. gibbosa

33’. Latero-ventral corners of anterior lobe of pronotum not projected ventro-laterally ( Figs. 5l, r); middle lobe of pronotum with antero-dorsal ear-like lobes (e.l., Figs. 4l; 5l, r); female hind lobe of pronotum more or less trilobed, with lateral corners angled ( Figs. 4l; 5l, r).......................................................................... 34

34(33’). Middle lobe of pronotum with antero-dorsal ear-like lobes (e.l.) directed laterally, with surface oriented dorso-ventrally and with a dorso-basal concave depression ( Figs. 4l; 5l); posterior margin of female sternum S8 with 5 small denticles ( Fig. 9l); N Peru ( Fig. 14g)................................................................ M. guillermoi

34’. Middle lobe of pronotum with antero-dorsal ear-like lobes (e.l.) upright, narrow or wide and sinuous ( Figs. 4r; 5r); posterior margin of female sternum S8 with 1–3 denticles or spines ( Fig. 9q); N Peru, Ecuador, and S Colombia ( Fig. 14g)................................................................................. M. mauffrayi

35(32’). Posterior margin of sternum S8 with two strong straight sub-cylindrical spines separated at their bases ( Figs. 9j, u).... 36

35’. Posterior margin of sternum S8 with two or three prongs joined at a common base ( Figs. 9g, t, v, ab)................ 37

36(35). Middle lobe of female pronotum with tongue-shaped antero-lateral lobes (e.l.) directed latero-ventrally ( Figs. 4v; 5v); spines of distal margin of female sternum S8 separated by a distance less than their individual width at their base ( Fig. 9u); Huánuco Dep. in Peru ( Fig. 14g)........................................................... M. panguanae

36’. Middle lobe of female pronotum smooth ( Figs. 4j; 5j); spines of distal margin of female sternum S8 separated by a distance greater than their individual width at their base ( Fig. 9j); Loreto Dep. in Peru ( Fig. 14g)................. M. gabrielae

37(35’). Posterior margin of sternum S8 projected into a bifid process with lateral prongs sub-cylindrical and parallel to sternum ( Fig. 9t); N Peru and Ecuador ( Fig. 14f).......................................................... M. orthogonia

37’. Posterior margin of sternum S8 projected into a trifid process with a central spine and laminar lateral prongs ( Figs. 9g, v, ab) ..................... .. ....................... .. .. .. ............ .................... ............. 38

38(37’). Lateral laminar prongs oriented parallel to sternum, semicircular with medial margin straight, narrowing into slender posterior spines, and central spine about as long as 0.33 of lateral prongs ( Fig. 9 ab); N and W Brazil ( Fig. 14f)................................................................................................... M. tridentigera

38’. Lateral laminar prongs oriented parallel to sides of ovipositor, scythe-shaped, and central spine about as long as 0.10–0.12 of lateral prongs ( Figs. 9g, v).......................................................................... 39

39(38’). N Peru ( Fig. 14f)........................................................................... M. peltata

39’. S Peru and W Brazil ( Fig. 14f)............................................................... M. falcifera