Protoneura cupida Calvert, 1903

Ellenrieder, Natalia Von & Garrison, Rosser W., 2017, A synopsis of the Neotropical genus Protoneura (Odonata: Coenagrionidae), Zootaxa 4361 (1), pp. 1-76 : 24-25

Protoneura cupida Calvert, 1903

Figs. 9 (♂ habitus), 33 (♀ habitus), 55 (♀ mes. plate), 79 (gen. lig.), 105 (♂ app.), 122 (map)

Protoneura cupida Calvert, 1903: 140 , 142–143, pl. 5, Figs. 46 , 47 (in key, description of ♂ ♀, illustrations of S10);— Calvert (1907: 394, 395; in key);— Williamson (1915: 619, 624, fig. 18; in key, illustration of thoracic pattern);— Paulson (1982: 252; Mexico and Guatemala); — Novelo-Gutiérrez & Peña-Olmedo (1991: 130; Mexico);— Measey (1994: 45; Belize);— Novelo-Gutiérrez (1994: 1994: 29–31, Figs. 9–14 ; description of larva, illustrations of habitus, prementum, labial palp, mandibles, antenna, gonapophysis, cerci, and caudal lamellae, diagnosis from larvae of P. ailsa and P. cupida );— Boomsma & Dunkle (1996: 25; Belize);— González-Soriano & Novelo-Gutiérrez (1996: 162; Mexico);—González- Soriano (1997: 253; Mexico);— González-Soriano (2001: 338; notes on biology);— Paulson (2009d; IUCN assessment);— Garrison et al. (2010: 379);— González-Soriano & Paulson (2011: 302, 307; Mexico);—González-Soriano et al. (2011: 184; Mexico);— González-Soriano & Novelo-Gutiérrez (2011: 2, Appendix VIII.21.2; Mexico).

Primary types. Holotype ♂. Guatemala: Livingston, Wilson leg. [ MCZ].

Specimens examined. 162 ♂ 9 ♀: MEXICO , San Luis Potosí State: 1 ♂, 8 km N of Tamazunchale , 5 ix 1963, T.W. Donnelly leg. [ FSCA] ; 7 ♂, Palictla {21°17' N, 98°47' W, 180 m}, 6 viii 1966, O.S. Flint, Jr. leg. [USNM]; 9 ♂ 1♀, stream at Palitla, 5.5 NW of Tamazunchale, 240 m, 19 vi 1966, D.R. Paulson & M.L. Paulson leg. [FSCA]; 1 ♂, same data but 26 v 1965 [RWG] ; 4 ♂, same data but [ FSCA] ; 2 ♂, same data but 26 vi 1965 [ USNM] ; 1 ♂ same data but [ UMMZ] ; 1 ♂, same data but 28 vi 1990, K.J. Tennessen leg. [ FSCA] ; Veracruz State: 2 ♂, Puente Nacional , Jalapa to Veracruz on route 125 {19°19'43'' N, 96°29'4'' W, 60 m}, 11 vii 1987, M.J. Westfall leg. [ FSCA] ; 11 ♂, Dos Ríos, route 140, km 347 {18°56' N, 97°2' W, 1,200 m}, 22 vii 1965, O.S. Flint, Jr & M.A. Ortiz B. leg. [USNM]; 12 ♂ 2 ♀, same data but 1 viii 1966 [ USNM] ; 1 ♂, Atoyac {18°54' N, 96°46' W, 646 m}, 28 viii 1966, A.B. Lau leg. [USNM]; 4 ♂, Otapa River , 8 km S of La Tinaja (18°41'9'' N, 96°26'10'' W, 90 m), 13–20 viii 1976, R.W. Garrison & J.A. Garrison leg. [RWG] ; 1 ♂ Los Tuxtlas Biological Station near Catemaco (18°40'46'' N, 95°9'16'' W, 35 m), 14 vii 1992, S.W. Dunkle leg. [ FSCA] ; 1 ♂, Máquinas River , Villa Cariño (18°37'27'' N, 95°5'31'' W, 18 m), 14 vi 2009, R.W. Garrison & N. von Ellenrieder leg. [RWG] ; 2 ♂, same data but 13–15 v 1981, C.M. & O.S. Flint Jr. leg. [RWG] ; 1 ♂, same data but [ USNM] ; 1 ♂ vicinity Lagunas Azul and Emilia, Los Tuxtlas Biological Station , about 30 km NE of Catemaco (18°25'12'' N, 95°6'36'' W, 600 m), 27 viii 1988, R.W. Garrison leg. [RWG] ; 1 ♂, same data but [ CSCA] ; 3 ♂, same data but 7–14 viii 1982, R.W. Garrison leg. [RWG] ; 1 ♂, same data but [ FSCA] ; 2 ♂, Laguna Azul, Emilia, and Arroyo de Laguna Escondida near Los Tuxtlas Biological Station , 27 viii 1988, S.W. Dunkle leg. [ FSCA] ; 1 ♂, 2 km W of Las Cabañas, Los Tuxtlas area {18°19' N, 95°4' W, 690 m}, 4–7 v 1981, C.M. Flint & O.S. Flint Jr. leg. [ USNM] ; 4 ♂, near Balzapote, Los Tuxtlas area , 15 v 1981, C.M. Flint & O.S. Flint Jr. leg. [ USNM] ; 4 ♂, Hondo River, on road to Colonia La Apachital , 6 mi E of highway from Tierra Blanca to Alemán, 18 viii 1957, G.H. Beatty III leg. [ FSCA] ; 1 ♂, stream at km 212 of highway 180, Ayacuyan-Catemaco, 8 vii 1980, E. González leg. [FSCA]; Oaxaca State: 1 ♂ , Valle Nacional {17°48' N, 96°18' W, 10 vii 1965, A.B. Lau leg. [ USNM] ; Tabasco State: 1 ♂, Tepescuintle, Tenosique {17°27' N, 91°24' W, 12 ix 1962, E.C. Welling M. leg. [ UMMZ] ; Chiapas State: 5 ♂ 1 ♀, Cascada Mizolja, 20 km S of Palenque {17°20' N, 91°58' W, 94 m}, 17–18 v 1981, C.M. Flint & O.S. Flint Jr. leg. [ USNM] ; 1 ♀, same data but [RWG] ; 4 ♂, same data but 25 ix 1983, R.W. Garrison leg. [RWG] ; 1 ♂, same data but [ FSCA] ; 1 ♂, Agua Azul {16°45' N, 90°46' W, 134 m}, 25 ix 1983 [RWG]. BELIZE , Cayo Dist.: 2 ♂, Little Vaquero Creek on Chiquibul Road, N of Augustine, Mountain Pine Ridge {17°3' N, 88°57' W, 500 m}, 22–25 vii 1983, T.W. Donnelly leg. [ FSCA] ; 2 ♂, Frío River at Augustine, Mountain Pine Ridge {16°58' N, 89°59' W, 500 m}, 22–25 vii 1983, T.W. Donnelly leg. [ FSCA] ; 1 ♂, Caves Branch, 1 mile S on Hummingbird highway and tributaries to Caves Branch, 31 v 1993, S.W. Dunkle leg. [FSCA]; Toledo Dist.: 1 ♂ 1 ♀ (pair in tandem), stream just W of San Antonio at waterfall (16°14'9'' N, 89°2'4'' W, 118 m), 6 vi 1993, S.W. Dunkle leg. [RWG] ; 2 ♂, same data but [ FSCA] ; 1 ♂, San Antonio, W of Jute stream, below the falls, W.F. Mauffray leg. [FSCA]; 2 ♂, Aguacate River , 3.6 mi NW of Blue Creek, W of San Antonio {16°13' N, 89°4' W, 169 m}, 5 vi 1993, S.W. Dunkle leg. [ USNM] ; 4 ♂, same data but [ FSCA] ; 2 ♂, Aguacate, 1 mi E of Aguacate Creek , 5 vi 1993, W.F. Mauffray leg. [ FSCA] ; 3 ♂, Blue Creek Village S of San Antonio {16°12' N, 89°3' W, 40 m}, 5 vi 1993, S.W. Dunkle leg. [FSCA]. GUATEMALA: 1 ♂, Ferrocarril del Norte, Section 11, 13 ii 1905, E.B. Williamson leg. [ UMMZ] ; Izabal Dept.: 7 ♂, Las Escobas, near Matías de Galvez {15°41' N, 88°38' W, 56 m}, 14–16 viii 1965, O.S. Flint Jr. & M.A. Ortiz leg. [ USNM] ; 9 ♂, same data but 27 vi 1966 [ USNM] ; Zacapa Dept.: 1 ♂, Los Amates, along San Francisco River {15°16' N, 89°6' W, 80 m}, 19 vi 1909, E.B. Williamson leg. [ UMMZ] ; Chiquimula Dept.: 6 ♂, Lucía Sazo River , 5 km S of Quetzaltepeque {14°35' N, 89°27' W, 800 m}, 9 vii 1962, T.W. Donnelly leg. [ FSCA] ; 24 ♂ 2 ♀, Padre Miguel {14°34' N, 89°27' W, 889 m}, 19 viii 1965, O.S. Flint, Jr. & M.A. Ortiz B. leg. [USNM]; 1 ♀, same data but [RWG] ; 2 ♂, near Tierra Colorada {14°34' N, 89°20' W, 800 m}, 19 vii 1962, T.W. Donnelly leg. [USNM].

Characterization. Male: Epicranium black, black stripes on thorax and dorsum of abdomen, which has pale spots on dorsum of S1–2 and base of S3–7; pale colors light blue to pale yellow ( Fig. 9 ). Pronotum anterior lobe pale blue, middle lobe pale blue with middorsal black line and sides black, and posterior lobe entirely pale blue. Mesepisternum pale blue with broad black stripe over middorsal carina and a narrow black stripe along mesepisternal-mesepimeral suture; mesepimeron pale blue with posterior half black; metepisternum pale blue with black stripe along metepisternal-metepimeral suture; metepimeron entirely pale blue to pale yellow. Leg pale yellow with tips of femur, tibia, and tarsi dark brown to black; tibial spurs shorter than twice intervening spaces. Genital ligula lacking lateral lobes and with a slightly concave distal margin, with margins of distal segment spiculate, and laterodistal corners not projected ventrally ( Fig. 79 ). Cercus shorter than S10 length, slightly shorter than paraproct, about as long as wide, with a depression on external dorsolateral surface and a longitudinal split along ventrodistal portion, dorsal margin with a distal concavity and ventral margin slightly convex in lateral view ( Fig. 105a ), with medial surface concave and with a small blunt ventrobasal tooth and two apical pointed dorsal teeth, one on distal margin and one on medial surface, visible in dorsal and mediodorsal views ( Figs. 105b, c ). Paraproct shorter than S10 length, at midlength narrowing to half its basal width and ending in a triangular tip ( Fig. 105 ). TL 38–40; Hw 18.5–20.

Female: As male but middle lobe of pronotum with pale blue restricted to a lateral band, posterior lobe black with pale blue medial and lateral spots; mesepisternum black with pale blue antehumeral stripe along most of segment; pale lateral area of S9 extended dorsally medially to about 1/2 of segment height ( Fig. 33 ). Middle lobe of pronotum lacking pronounced lateral depressions; posterior lobe entire, directed dorsally, with sides straight and tip smoothly convex. Mesostigmal plate approximately triangular with a central concavity and posteromedial corner contiguous with a carina extended midway to middorsal carina ( Fig. 55b ) and triangular in profile ( Fig. 55a ). TL 35.5–38.5; Hw 20–21.5.

Diagnosis. Among the species with male paraproct no longer than S10 length, P. cupida shares male cercus approximately quadrangular, about as long as wide, with a ventrodistal longitudinal split and depression on external dorsolateral surface, concave medial surface, with two short apical teeth in dorsal view ( Figs. 105 ), and genital ligula distal segment with lateral margins spiculate ( Fig. 79 ) with P. amatoria , P. aurantiaca , and P. sulfurata ( Figs. 98 ; 99; 115; and 72; 73; 89 respectively). It differs from all three by its pale blue color ( Fig. 9 ). Female posteromedial corner of mesostigmal plate contiguous with a carina of triangular profile ( Fig. 55 ) is unique for this species.

Habitat and biology. Adults found along fast-flowing rivers and streams surrounded by forest, open to shady, and trickles in swamps ( Measey 1994), at the pools but not the riffles (D. R. Paulson pers. comm.). Larvae were described by supposition based on specimens found at small pools and backwaters, among bottom leaf litter and shoreline vegetation of a rocky stream running through high elevation perennial forest in Mexico (Novelo- Gutiérrez 1994), where one individual was documented emerging on a small rock sticking out of the water. González-Soriano (2001) observed large aggregations of males flying around bushes over small islets below a waterfall, and a female ovipositing alone in non-aquatic vegetation growing at the edge of a small shaded stream in Mexico.

Distribution. Eastern Mexico, Belize, and Guatemala ( Fig. 122 ). Assessed as Least Concern by IUCN ( Paulson 2009d).

Figures

FIGURES 9–12. Male body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale).(9) P.cupida; (10) P. dunklei; (11) P.klugi; (l2) P.macintyrei.

FIGURES 45, 46. Female body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale). (45) P. viridis; (46) P.woytkowskii form b. Insert in Fig.46 above: pterothorax in dorsal view; below: thorax in ventral view.

FIGURES 47–50. Female pronotum (50a, c) and mesostigmal plates (47–49; 50b, d), in lateral (48a; 50 a, b) and dorsal (47; 48b; 49; 50c, d) views (to scale). (47) P.ailsa; (48) P.amatoria; (49) P.aurantiaca; (50) P. caligata. Figure 50 by RWG.

FIGURES 13–16. Male body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale).(13) P. paucinervis; (14) P.peramans; (15) P.rojiza; (16) P. romanae.

FIGURES 79–84. Genitalligula in ectal (79–84a), lateral (79–84b), and entolateral (81c; 84c) views (to scale).(79) P.cupida; (80) P.dunklei; (81) P.klugi; (82) P.macintyrei; (83) P.paucinervis; (84) P. peramans. All by RWG.

FIGURES 103–105. Male caudal appendages in lateral (103–105a), posterodorsal (103–105b), and dorsal (103–105b) views (not to scale). (103) P.cara; (104)P.corculum; (105)P.cupida.

FIGURES 33–36. Female body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale). (33) P. cupida; (34) P.dunklei; (35)P.klugi; (36) P.macintyrei. Insert in Fig.35: thorax in ventral view.

FIGURES 55–59. Female pronotum (58b) and mesostigmal plates (55–58; 59a, c), in lateral (55a; 56a; 59a) and dorsal (55b; 56b; 57; 58; 59b, c) views (to scale). (55) P. cupida; (56) P. dunklei; (57) P. klugi; (58) P. macintyrei; (59) P. paucinervis.

FIGURES 97–99. Male caudal appendages in lateral (97–99a), posterodorsal (97–99b), and dorsal (97–99b) views (not to scale).(97) P.ailsa; (98) P.amatoria; (99) P.aurantiaca.

FIGURES 121–122. Distribution maps.