Protoneura cara Calvert, 1903

Ellenrieder, Natalia Von & Garrison, Rosser W., 2017, A synopsis of the Neotropical genus Protoneura (Odonata: Coenagrionidae), Zootaxa 4361 (1), pp. 1-76 : 20-22

Protoneura cara Calvert, 1903

Figs. 7 (♂ habitus), 31 (♀ habitus), 53 (♀ mes. plate), 77 (gen. lig.), 103 (♂ app.), 121 (map)

Protoneura cara Calvert, 1903: 140 , 143–144, pl. 5, Figs. 38 , 45 , pl. 6, fig. 9 (in key, description of ♂ ♀, illustrations of ♂ S10 and body);— Williamson (1915: 619, 624, Figs. 5 , 19 ; in key, illustration of wings and thoracic pattern);— Johnson (1972: 73; characterization, counties in Texas, USA);— Gloyd (1981: 133; El Salvador);— Paulson (1982: 252; USA, Mexico, Guatemala, El Salvador);— Paulson (1984: 51; El Salvador);— González-Soriano & Novelo-Gutiérrez (1996: 162; Mexico);— Abbott & Stewart (1998: 207; USA & Mexico);— Abbott (2001: 196; Texas);— González-Soriano (2001: 335– 339; notes on biology);— Sibley (2003: 6; Nicaragua);— Abbott (2005: 13, 49–51, Figs. 6B , 12 ; characterization, notes on habits and behavior, illustrations of ♂ S10 and ♀ mesostigmal plates, distribution map in Texas);— Urrutia (2005: 33; Colombia);— Abbott (2006: 50, 56, 111; flight period and distribution map Texas);— Westfall & May (2006: 431, 433, 436–438, Figs. 223D, 224B, 226B; in key to northern representatives of genus, characterization of adults, illustrations of ♂ thorax and S10, ♀ pronotum and mesostigmal plates);— Abbott (2007: 20, 32, 89; flight period and distribution map Texas);— Upson et al. (2007: 28; Mexico);— Behrstock et al. (2007: 32; Mexico);— Cuevas-Yañez (2007: 86, 87; Mexico);— Abbott (2008: 23, 35, 93; flight period and distribution map Texas);— Bailowitz et al. (2009: 16; Mexico);— Paulson (2009h: 189 190; Texas, characterization, natural history, flight period, color photos);—González-Soriano et al. (2011: 184; Mexico);— Bailowitz et al. (2012: 9; El Salvador);— Garrison et al. (2010: 379);— Abbott (2011: 100–101; characterization, notes on habits and behavior, color illustration of adults, picture of pair ovipositing, distribution map in Texas);— González-Soriano & Paulson (2011: 303, 307; Mexico);— González-Soriano & Novelo-Gutiérrez (2011: 2, Appendix VIII.21.2; Mexico);— Escoto-Moreno et al. (2014: 1046, 1052; Mexico);— Bailowitz et al. (2016: 210 –211; habitat, characterization, field pictures of male, female, and pair, dot map of distribution in Sonora).

Primary types. Holotype ♂. Mexico, Guerrero, Iguala , ix 1898, O.W. Barrett leg. [ ANSP # 9239 ].

Specimens examined. 84 ♂ 20 ♀: U.S.A., Texas State, Kimble Co. : 1 ♂, North Llano River at Farm road 1674 (30°30'58'' N, 99°48'19'' W, 506 m), 14 vii 2001, J. Miller leg. [ FSCA] ; Kerr Co.: 4 ♂ 1 ♀, South Fork Guadalupe River near Hunt (30°4'14'' N, 99°20'15'' W, 550m) 18 vi 1985, S.W. Dunkle leg. [ FSCA] ; 2 ♂, same data but [RWG] ; 1 ♂, same data but J.J. Daigle leg. [RWG] ; Real Co.: 1 ♂ 1 ♀, Frio River at highway 337, E of Leakey {29°43' N, 99°45' W, 475 m}, 14 vii 2001, K.J. Tennessen leg. [ FSCA] ; 1 ♂, Frio River, Garner State Park {29°34' N, 99°43' W}, 19 vi 1979, C. Cook leg. [ FSCA] ; Uvalde Co.: 3 ♂ 1 ♀, Sabinal River at Texas highway 187, 3.7 mi S of Utopia {29°36' N, 99°31' W, 410 m}, 22 viii 1977, D.R. Paulson & S. Hills leg. [ FSCA] ; Bexar Co.: 1 ♂ 1 ♀ (in tandem), Medina River at Jungman Road, W of San Antonio (29°20'45'' N, 98°47'11'' W, 216 m), 16 v 2001, S.W. Dunkle leg. [RWG] . MEXICO, San Luis Potosí State: 1 ♂, El Salto Falls {22°21'42'' N, 99°15'53'' W, 250 m}, 12 vi 1963, R. E. Woodruff leg. [ FSCA] ; 1 ♂, Coy River headwaters, 28 km S of Ciudad Valles (21°43'59'' N, 98°58'41'' W, 42 m), 21 viii 1999, R.W. Garrison & J.A. Garrison leg. [ CSCA] ; 1 ♂ 1 ♀ (in tandem), Municipio de Huehuetlán, Micos River above Cascada de Micos , 2 km N of Micos (22°7'29'' N, 99°10'46'' W, 240 m), 22 viii 1999, R.W. Garrison & J.A. Garrison leg. [ CSCA] ; Nayarit State: 1 ♂ 1 ♀, Jumatán {21°30'34'' N, 104°52'3'' W, 935 m}, 13 iii 1981, E. González leg. [RWG] ; 4♂ 1 ♀, Tepic {21°28' N, 104°53' W, 966 m}, 5–11 xi 1923, J.H. Williamson leg. [ UMMZ] ; 1 ♂ 1 ♀, same data but [ FSCA] ; Michoacan State: 1 ♂, Apatzingan {19°5' N, 102°26' W, 329 m}, 21 vii 1947, T.H. Hubbellleg. leg. [ UMMZ] ; Morelos State: 2 ♂, Lagunas Zempoala National Park {19°3' N, 99°18' W, 2,800 m}, 10–11 vii 1965, O.S. Flint, Jr. & M.A. Ortiz B. leg. [ USNM] ; 1 ♂, stream in Pueblo Cocoyotla , highway 421 (18°46'4'' N, 99°27'26'' W, 1,070 m), 22 vii 1992, K.J. Tennessen leg. [ FSCA] ; 6 ♂ 1 ♀, Xochitepec {18°45'53'' N, 99°15'39'' W, 1,100 m}, 12–14 vii 1965, O.S. Flint & M.A. Ortiz leg. [ USNM] ; 12 ♂, route 95, km 91 near Xochitepec, 1 viii 1965, O.S. Flint Jr. leg. [ USNM] ; 1 ♂, Municipio Tlaltizapán, Las Estacas, spring and stream (18°43'48'' N, 99°6'48'' W, 980 m), 8 vii 1987, M.J. Westfall, Jr. leg. [FSCA] ; 1 ♂, Amacuzac River at Tehuixtla and stream at Balneario Las Palmas (18°32'51'' N, 99°16'13'' W, 866 m), 20 vii 1992, S.W. Dunkle leg. [ FSCA] ; 1 ♂, 1 ♀, same data but K.J. Tennessen leg. [ FSCA] ; 1 ♂, same data but W.F. Mauffray leg. [ FSCA] ; Chiapas State: 2 ♂, E of Cintalapa , 21 km at route 190 {16°41' N, 93°40' W, 550 m}, 9 xii 1975, C.M. & O.S. Flint Jr. [ USNM] ; 4 ♂ 3 ♀, route 190, km 1024, beyond Cintalapa {16°41' N, 93°40' W, 550 m}, 8–10 vii 1966, O.S. Flint, Jr. & M.A. Ortiz B. leg. [ USNM] ; Oaxaca State: 1 ♂, route 190, km 698, near El Coyul {16°29' N, 95°53' W, 19 m}, 11 vii 1966, O.S. Flint, Jr. & M.A. Ortiz, B. leg. [ USNM] ; 1 ♂, stream 14 mi E of El Camarón {16°29' N, 95°47' W, 850 m}, 26 viii 1967, D.R. Paulson & M.L. Paulson leg. [ FSCA] ; Veracruz State: 2 ♂, Verde River plane, route 140 km 368 {18°56' N, 97°2'0'' W, 1,200 m}, 23 vii 1965, O.S. Flint, Jr. & M.A. Ortiz B. leg. [ USNM] . GUATEMALA, Alta Verapaz Dept.: 1 ♀, Finca Semuc Champey, 5 km SSE of Lanquín {15°32' N, 89°57' W}, 3 v 1980, T.W. Donnelly leg. [ FSCA] ; Zacapa Dept.: 1 ♂, Gualán {15°6' N, 89°21' W, 141 m}, 14 i 1909, E.B. Williamson leg. [ UMMZ] ; El Progreso Dept.: 1 ♂, Estancia de La Virgen {14°56' N, 89°53' W, 320 m}, 11 viii 1965, O.S. Flint, Jr. & M.A. Ortiz B. leg. [RWG] ; Santa Rosa Dept.: 2 ♂, Agua Caliente {14°55' N, 90°24' W, 569 m}, 2 vi 1909, E.B. Williamson leg. [ UMMZ] ; Chiquimula Dept.: 1 ♂, near Tierra Colorada {14°34'37'' N, 89°20'20'' W, 800 m}, 19 vii 1962, T.W. Donnelly leg. [ FSCA] ; 1 ♂, small stream vicinity Santa Elena, 600 m, 12 vii 1962, T.W. Donnelly leg. [ FSCA] ; Escuintla Dept.: 1 ♂, stream 5.7 mi SW of Escuintla {14°22' N, 90°49' W, 671m}, 9 vii 1964, F.G. Thompson leg. [ FSCA] ; Jutiapa Dept.: 1 ♂ 1 ♀, 27.2 mi ESE of Chiquimulilla {14°5' N, 90°22' W, 215 m}, 3 ix 1964, F. G. Thompson leg. [ UMMZ] ; 1 ♂, same data but [ FSCA] . EL SALVADOR, La Libertad Dept.: 4 ♂ 2 ♀ (one pair in tandem), 4.7 mi S of Mizata {13°30' N, 89°33' W, 70 m}, 2 vii 1966, O.S. Flint, Jr. & M.A. Ortiz B. leg. [ USNM] . HONDURAS, Francisco Morazán Dept.: 1 ♀, Zamorano, Quebrada El Gallo , small stream (14°1' N, 87°2' W, 650 m), 7 iii 1990, S.W. Dunkle leg. [ FSCA] ; 1 ♂, Zamorano, Escuela Agrícola Panamericana, 30 km ESE of Tegucigalpa, ponds and Yeguari River with riffles & pools, 14 xii 1987, L. Córdoba leg. [ FSCA] . NICARAGUA: Chontales Dept.: 1 ♂, route 7, km 183, 4.2 mi W of Villa Somosa {12°3' N, 84°59' W, 200 m}, 29 vii 1967, O.S. Flint, Jr. & M.A. Ortiz B. leg. [ USNM] . PANAMA, Canal Zone: 10 ♂ 1 ♀, Sardanilla River {9°7' N, 79°35' W, 145 m}, 5 xii 1916, E.B. Williamson leg. [ UMMZ] .

Characterization. Male: Epicranium black with orange spots along margin of compound eyes, anterior margin of postfrons and occipital bar; black stripes on thorax; dorsum of abdomen brown to black with pale spots on S1–3, basal portion of S4–7, and distal margin of S9–10; pale colors red to orange and yellow ( Fig. 7 ). Pronotum anterior, middle and posterior lobes black with medial and lateral orange spots. Mesepisternum black with middorsal carina orange, a antehumeral stripe along entire sclerite, and a narrow black stripe along distal 1/4 of mesepisternal-mesepimeral suture; mesepimeron black margined in orange; metepisternum orange with black along posterior margin; metepimeron orange to yellow with central dark area to black with only posterior margin orange. Coxa dark brown, remainder of leg mostly pale, yellow to orange, sometimes with two diffuse brownish rings subbasally and subapically on each femur and tibia; tibial spurs shorter than twice intervening spaces. Genital ligula lacking lateral lobes and with a slightly convex distal margin, with laterodistal corners projected ventrally ( Fig. 77 ). Cercus about as long as S10 length, about as long as paraproct, longer than wide, with a depression on external dorsolateral surface, dorsal margin and ventral margin smoothly convex in lateral view ( Fig. 103a ), medial surface with a small blunt ventrobasal tooth, a large medial branch ending on a curved tooth separated by a membranous area from reminder of cercus, and a subapical dorso-ventral ridge, visible in dorsal and mediodorsal views ( Figs. 103b, c ). Paraproct about as long as S10 length, at midlength narrowing to half its basal width and ending in a bluntly pointed tip ( Fig. 103 ). TL 36–38; Hw 16–17.7.

Female: As male but black more extensive on abdomen; pale lateral area of S9 extending along basal 1/3 of segment to about 2/3 of segment height posteriorly ( Fig. 31 ). Middle lobe of pronotum lacking pronounced lateral depressions; posterior lobe directed dorsally and smoothly convex. Mesostigmal plate reniform and convex, with posteromedial corner contiguous through lateral arm of middorsal carina with middorsal carina, which is higher at convergence point than remainder of pterothorax; with a depression on mesepisternum behind plate ( Fig. 53 ). TL 32.5–36.5; Hw 18–19.

Diagnosis. Protoneura cara can be distinguished from all congeners by the morphology of its male cercus, with a large medial branch ending on a curved tooth separated by a membranous area from remainder of cercus ( Figs. 103b, c ), and by female mesostigmal plate reniform and contiguous with elevated middorsal carina via lateral arms ( Fig. 53 ).

Habitat and biology. Adults found at pools and sluggish backwaters bordered by extensive overhanging bush and undercut root-balls ( Upson et al. 2007), at small, slow-flowing streams, sometimes in the open, sometimes with a dense overhead canopy ( Bailowitz et al. 2009), small clear streams with sand bottom in woodland or open ( Paulson 2009h), and along shallow slow clear rivers ( Behrstock et al. 2007). According to González-Soriano (2001), adults are commonly seen flying along streams and rivers in Mexico, either as solitary males or tandem pairs, and males patrol water surface searching for ovipositing females and switch from one perch to another without defending a particular area. González-Soriano (2001) observed only one pair in copula perching on grasses at the edge of a spring-fed pond and reported oviposition as occurring in tandem, with male remaining erect beating its wings, and tandem pairs flying along streams and rivers searching for oviposition sites consisting of submerged grasses, living roots, and floating objects such as small pieces of wood. He also observed males aggregating to form aerial mating swarms, hovering in a stationary fashion 4–5 m above ground level, 2–3 m from the stream edge at a forest gap surrounded by small trees with all hovering individuals facing in the same direction and displaying no foraging activity, apparently waiting for receptive females. Paulson (2009h) also reported seeing occasional mating swarms, with aggregations of males hovering well above head height in sunny clearings near water, and ensuing tandem formation when females entered swarm. He reported males spending much of their time hovering just above water but also perching on slender stems, and slowly rising into trees when disturbed. According to Abbott (2011), egg laying typically occurs in tandem near the margin of slow moving water in floating and submerged vegetation and debris, with females curving abdomen between wings, and males typically hovering just above water for extended periods of time. Paulson (2009h) observed tandem pairs ovipositing in any sort of floating detritus, including tiny wood chips, typically where currents concentrate detritus, but also in grass stems and rootlets of woody plants near shore. Bailowitz et al. (2016) reported its flight period in Sonora, Mexico, as spanning from late April through late October, and Paulson (2009h) from June to October in Texas, USA.

Distribution. Texas State in USA to Colombia ( Fig. 121 ).

Figures

FIGURES 5–8. Male body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale). (5) P. calverti; (6) P. capillaris; (7) P.cara; (8) P.corculum.

FIGURES 37–40. Female body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale). (37) P. paucinervis; (38) P.peramans; (39) P.rojiza; (40) P.romanae. Insert in Fig. 40: pronotum in dorsal view.

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.

FIGURES17–20. Male body scan; head in dorsalview, thorax and abdomen in lateral view (not to scale). (17) P.sanguinipes; (18) P.scintilla; (19) P.sulfurata; (20) P. tenuis.

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 71–78. Genital ligula inectal (71–78a) and lateral (71–78b) views (to scale). (71) P.ailsa; (72) P.amatoria; (73) P. aurantiaca; (74) P.caligata; (75) P.calverti; (76) P.capillaris; (77) P.cara; (78) P.corculum. 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 29–32. Female body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale). (29) P. calverti; (30) P.capillaris; (31) P.cara; (32) P.corculum. Insert in Fig. 29: posterior lobe of pronotum in dorsal view.

FIGURES 51–54. Female pronotum (54c) and mesostigmal plates (51–53; 54a, b), in lateral (51a; 52a; 53a; 54a), dorsal (51b; 52b; 53b; 54b, c), and dorsolateral (51c) views (to scale). (51) P. calverti; (52) P. capillaris; (53) P. cara; (54) P. corculum.

FIGURES 121–122. Distribution maps.