Calvert, 1907 : 394 Louton et al. (1986 : 438 De Marmels (1988 : 97; Venezuela ) Machado et al. (1991 : 169 Garrison et al. (2003 : 20 Larison (2007 : 1021 Garrison et al. (2010 : 379 Pessacq et al. (2012 : 70 A synopsis of the Neotropical genus Protoneura (Odonata: Coenagrionidae) Ellenrieder, Natalia Von Garrison, Rosser W. Zootaxa 2017 2017-12-01 4361 1 1 76 4N6FT Calvert, 1907 Calvert 1907 [151,581,1231,1258] Insecta Protoneuridae Protoneura Animalia Odonata 9 10 Arthropoda species amatoria     Protoneura amatoria  Calvert, 1907: 394, 395, pl. 10, Figs. 49–52(in key, description of ♂and ♀from Hondurasand  Guatemala, illustrations of ♂S10, base of wing, and ♀pronotum);— Williamson (1915: 619, 623–624, Figs. 3, 15, 16; in partial key, notes on biology and variability, illustration of wings and thoracic pattern);— Soukup (1954: 13; Peru);— Campos (1922: 30; Ecuador);— Rácenis (1955: 56; Venezuela);— Rácenis (1958: 180; Venezuela);— Paulson (1982: 252;  Guatemala, Honduras, Costa Rica,  Panama, South America);—  Louton et al.(1986: 438; Peru);— Brooks (1989: 50; Costa Rica);—  De Marmels (1988: 97; Venezuela);— Convey (1989: 57; Venezuela);—  Machado et al.(1991: 169; Brazil);— Donnelly (1992: 82;  Panama);— Boomsma & Dunkle (1996: 25;  Belize);— Daigle (2002: 14; Nicaragua);—O’Grady & May (2003: 2834; in phylogenetic analysis);— Sibley (2003: 6; Nicaragua);—  Garrison et al.(2003: 20; typeinformation);— Lencioni (2005: 221; Brazil);— Botero (2006: 52; Colombia);— Martinez (2006: 61; Colombia);—  Larison (2007: 1021–1028; mating tactics);— Pessacq (2008: 521, 522, 527; in phylogenetic analysis);— Hoffmann (2009: 63; Peru);— Paulson (2009a: IUCN assessment);—  Garrison et al.(2010: 379);—  Pessacq et al.(2012: 70; Brazil);—Garrison & von Ellenrieder (2013: 22; Costa Rica);— Michalski (2015: 76–78; habitat in Trinidad, photos, key);—von Ellenrieder et al.(2017: 7; Guyana).   Primary types. Holotype ♂: Honduras: San Pedro Sula,  28 ii 1905, E.B. Williamsonleg. [ UMMZ].   Specimens examined. 351 ♂ 165 ♀:  BELIZE, ToledoDist.: 1 ♀, deep river at S Highway { 16°14' N, 89°1' W}, 7 vi 1993, S.W. Dunkle leg. [FSCA].  GUATEMALA: IzabalDept.: 1 ♂, Puerto Barrios { 15°43' N, 88°36' W}, 6 iii 1905, J.S. Hines leg. [UMMZ]; 1 ♂, Morales { 15°29' N, 88°49' W}, 27 v 1909, E.B. Williamson leg. [UMMZ]; ZacapaDept.: 2 ♂ 1 ♀, Los Amates { 15°16' N, 89°6' W}, 21 vi 1909, E.B. Williamson leg. [UMMZ]; 1 ♂ 1 ♀, same data but [USNM]. HONDURAS, CortésDept.: 9 ♂ 4 ♀(  paratypes), San PedroSula, in gulch { 15°30' N, 88°2' W, 83 m}, 27–28 ii 1905, E.B. Williamson leg. [UMMZ]; 1 ♂ 1 ♀, same data but [USNM]; 1 ♂, same data but [FSCA]; 1 ♂ 1 ♀, same data but [CSCA]. NICARAGUA, ChontalesDept.: 1 ♂, route 7, km 183, 4.2 miW of Villa Somosa { 12°3' N, 84°59' W}, 29 vii 1967, O.S. Flint, Jr. & M.A. Ortiz B. leg. [USNM];  GranadaDept.: 1 ♂, Domatila Private Reserve, spring area of Dorado River ( 11°42'7'' N, 85°57'6'' W, 50 m), 23 viii 2003, F.C. Sibley leg. [FSCA]; 1 ♀, same data but 27 vii 2003[FSCA]. COSTA RICA, GuanacasteProv.: 1 ♂ 1 ♀, Parque Nacional Santa Rosa, Estación Santa Rosa( 12°51'0'' N, 85°36'36'' W, 280 m), 18 viii 1990, U. Chavarría G. leg. [RWG]; 1 ♂ 1 ♀, Piedras River, N of Bagaces { 10°32' N, 85°15' W}, 27 vii 1967, O.S. Flint & M.A. Ortiz B. leg. [USNM]; AlajuelaProv.: 1 ♂, Los Chiles { 10°25' N, 84°20' W, 60 m}, 30 vii 1966, D.R. Paulson leg. [RWG]; 3 ♂ 1 ♀, same data but 14 x 1966[FSCA]; 3 ♂, S of Turrucares, Tizate River { 9°58' N, 84°19' W, 610 m}, 22–23 xii 1909, P.P. Calvert & Tristánleg. [UMMZ]; 1 ♂, same data but [FSCA]; HerediaProv.: 1 ♂, Sarapiquí, small flood plain pool at small clear water river 9.7 kmSE of Puerto Viejo { 10°22' N, 83°55' W}, 2 iii 1987, S.W. Dunkle leg. [FSCA]; PuntarenasProv.: 1 ♂ 1 ♀(in tandem), stony muddy creek about 11 kmN of Sardinal ( 10°9'38'' N, 84°51'12'' W, 246 m), 8 vi 2013, N. von Ellenrieder & R.W. Garrison leg. [RWG]; 1 ♂, upstream from bridge on Sardinal road, quiet eddy pool ( 10°8'44''N, 84°51'1'' W, 270 m), 14 v 2004, W. Haber & J. Richards leg. [RWG]; 1 ♂ 1 ♀, Bosque Los Negritos, about 2 kmN of entrance to Miramar on Panamerican Highway { 10°5' N, 84°43' W}, 16 iii 1990, S.W. Dunkle leg. [RWG]; 1 ♀, 8.1 miWNW of Esparta { 9°59' N, 84°40' W}, 26 vi 1967, M.J. & D.N. Westfall leg. [FSCA]; 3 ♂ 2 ♀, same data but O.S. Flint, Jr. & M.A. Ortiz B. leg. [USNM]; 1 ♂ 1 ♀, same data but 24 x 1966, D.R. Paulson & R.P. Seifert leg. [FSCA]; 1 ♂, Hacienda Baru, 3 kmNW of Dominical ( 9°16'30'' N, 83°52'57'' W, 10 m), 24 v 2009, F.C. Sibley leg. [FSCA]; 1 ♂ 1 ♀(in tandem), small stream crossing Panamerican Highway a few mi NW of Paso Real ( 9°0'48'' N, 83°13'36'' W, 150 m), 6 i 2007, F.C. Sibley leg. [FSCA]; 1 ♂, Quebrada Diecinueve, 11 kmE of Golfito on route 14; sandy stream ( 8°36'56'' N, 83°3'56'' W, 25 m), 31 v 2013, N. von Ellenrieder & R.W. Garrison leg. [RWG]; 1 ♂, Corcovado National Park { 8°34' N, 83°30' W}, 7 iii 1980, W. Brehme leg. [FSCA]; 1 ♀, same data but Estación Sirena, Camaronal River ( 8°28'51'' N, 83°35'38'' W, 5 m), 12–13 iv 1989, Holzenthal & Blahnik leg. [FSCA].  PANAMA,  PanamáProv.: 2 ♂, Chepo Distr., 2.3 kmE of El Llano ( 9°12'56'' N, 78°56'24'' W, 10 m), 25 ii 1985, O.S. Flint, Jr. & J. Louton leg. [USNM]; 1 ♂, same data but [RWG]; 1 ♂, Canal Zone, Pipeline Road, Agua Salud River near Frijoles { 9°12' N, 79°12' W}, 2 ii 1974, M.L. May leg. [FSCA]; 2 ♂ 1 ♀(one pair in copula), Canal Zone, Pipeline Road, Frijoles River ( 9°12' N, 79°12' W), 26 ii 1985, Flint & Louton leg. [USNM]; 2 ♂, Chagres Valley, Madden Dam { 9°12' N, 78°37' W}, Ward's National Science Expedition [FSCA]; 1 ♂, Pipeline Road, 1.7–4.8 miNW of Gamboa { 9°11'' N, 79°45' W, 140 m}, 8–15 viii 1970, T.W. Donnelly & E.S. Morton leg. [RWG]; 2 ♂, same data but 5 ii 1971[FSCA]; 1 ♂, Barro Colorado Island { 9°9' N, 79°51' W}, 27 xii 1950, R.B. Cumming leg. [FSCA]; 7 ♂ 3 ♀(one pair in tandem), canal Zone 2.2 kmNW of Gamboa, stream at Pipeline road { 9°7' N, 79°42' W, 27 m}, 7 viii 1979, R.W. Garrison & J.A. Garrison leg. [RWG]; 11 ♂ 2 ♀, Sardanilla River { 9°7' N, 79°35' W}, 5 xii 1916, E.B. Williamson leg. [UMMZ]; 3 ♂ 1 ♀, same data but [CSCA]; 1 ♂, Cabima { 9°7' N, 79°31' W}, 20 v 1911, A. Busck leg. [USNM]; 1 ♂, same data but [FSCA]; 1 ♂, Pipeline road, Agua Salud River ( 9°6' N, 79°42' W), 8–13 vii 1967, O.S. Flint, Jr. & M.A. Ortiz B. leg. [USNM]; 3 ♂, forest road at Gaillard highway, at milepost 12, 7.4 kmSE of Gamboa { 9°4' N, 79°40' W, 75 m}, 8 vi 1979, R.W. Garrison & J.A. Garrison leg. [RWG]; 1 ♂, Canal Zone, Chiva Chiva road { 9°1' N, 79°35' W}, 28 xii 1937, Bliss leg. [FSCA]; 6 ♂ 1 ♀, Mazamba River { 9°0' N, 79°36' W}, 6 xii 1916, J.H. & E.B. Williamson leg. [UMMZ]; 5 ♂, TrinidadRiver { 8°8' N, 81°14' W}, 1–10 vi 1912, A. Busck leg. [USNM]; ColónProv.: 1 ♂ 1 ♀(in copula), Canal Area, Pipeline road, Quebrada Juan Grande{ 9°7' N, 79°42' W}, 1 iv 1974, M.L. May leg. [FSCA]; 2 ♂ 1 ♀, same data but 6 iv 2001, B. Larison leg. [RWG];  PanamaOeste Prov.: 1 ♂ 1 ♀, Arraiján { 8°57' N, 79°39' W, 180 m}, 20 i 1946, D.W. Jenkins leg. [UMMZ]; 1 ♂, same data but [FSCA]. COLOMBIA, MagdalenaDept.: 5 ♂ 1 ♀, Santa Marta Mountains { 11°6' N, 74°5' W}, 16 xii 1916, J.H. & E.B. Williamson leg. [UMMZ]; 1 ♂ 1 ♀, same data but 3 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 19 ♂ 6 ♀, Frio River { 10°55' N, 74°10' W, 43 m}, 4– 8 i 1917, E.B. Williamson leg. [UMMZ]; 3 ♂ 2 ♀, same data but [CSCA]; 1 ♂, El Banco { 9°2' N, 73°58' W, 46 m}, 23 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; BolivarDept.: 31 ♂ 11 ♀, Bolivar{ 10°16' N, 75°28' W}, 21–24 xii 1916, J.H. & E.B. Williamson leg. [UMMZ]; 3 ♂ 3 ♀, same data but [CSCA]; ChocoDept.: 4 ♂ 1 ♀, Sautata { 7°50' N, 77°4' W}, 27 i 1918, M.A. Carriker leg. [UMMZ]; 1 ♂ 1 ♀, same data but [CSCA]; AntioquiaDept.: 2 ♂, Puerto Berrio { 6°30' N, 74°24' W}, 31 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 17 ♂ 4 ♀, Cristalina, Magdalenariver, about 60 kmS of Puerto Barrio { 6°24'43'' N, 74°45'38'' W, 652 m}, 12–20 ii 1917, J.H. & E.B. Williamson leg. [UMMZ]; TolimaDept.: 2 ♂ 1 ♀, Mariquita { 5°12' N, 74°54' W, 450 m}, 4 ii 1917, J.H. & E.B. Williamson leg. [UMMZ]. TRINIDAD, Diego MartinCo.: 2 ♂ 1 ♀, Diego Martinriver near Port of Spain{ 10°43' N, 61°33' W, 36 m}, 29 ii 1912, E.B. Williamson leg. [UMMZ]; St. Andrew Co.: 3 ♂, Maturo Ward, Maturo River and rivulets off Río Granderoad ( 10°40' N, 61°3' W, 18 m), 5 v 1988, J.J. Daigle leg. [FSCA]; 1 ♂ 1 ♀(in tandem), Maturo River at Toco main road, 22 viii 1986, S.W. Dunkle leg. [RWG]; 1 ♂ 1 ♀, ArimaRiver at Churchill– Roosevelt highway, about 2.25 miSSE of Arima( 10°35'17'' N, 61°14'48'' W, 35 m), 12 i 1981, R.W. Garrison leg. [RWG]; 1 ♂, Pan, A. Busck leg. [FSCA]; 1 ♂ 1 ♀(in copula), Sangre Grande{ 10°34' N, 61°8' W}, 17 vii 1960, G. Miller, L. Miller & H. Wood leg. [FSCA]; Caroni Co.: 1 ♀, shaded still water area, 1.5 miW of Navet Dam ( 10°23'43'' N, 61°16'14'' W, 63 m), 14 i 1981, R.W. Garrison leg. [RWG]. VENEZUELA, FalcónState: 6 ♂ 3 ♀, Tucacas { 10°47' N, 68°19' W}, 24–25 iii 1920, W.H. Ditzler, J.H. & E.B. Williamson leg. [UMMZ]; 22 ♂ 12 ♀, Palma Sola { 10°36' N, 68°32' W}, 4–10 iii 1920, W.H. Ditzler, J. H. & E.B. Williamson leg. [UMMZ]; 2 ♂ 2 ♀, same data but [CSCA]; ZuliaState: 20 ♂ 8 ♀, El Guayabo { 10°37' N, 71°51' W, 84 m}, 20–22 iv 1920, W.H. Ditzler, J.H. & E.B. Williamson leg. [UMMZ]; 3 ♂ 1 ♀, same data but [CSCA]; YaracuyState: 12 ♂ 10 ♀, Boquerón { 10°34' N, 68°49' W}, 16–21 iii 1920, W.H. Ditzler, J.H. & E.B. Williamson leg. [UMMZ]; 1 ♂ 1 ♀, same data but [RWG]; 5 ♂ 5 ♀, Aroa { 10°26' N, 68°53' W}, 12–13 iii 1920, W.H. Ditzler, J.H. & E.B. Williamson leg. [UMMZ]; 1 ♂ 1 ♀, same data but [CSCA]; CaraboboState: 4 ♂ 2 ♀, San Esteban { 10°25' N, 68°4' W}, 1–5 ii 1920, W.H. Ditzler, J.H. & E.B.Williamson leg. [UMMZ]; BarinasState: 1 ♂, Barinas{ 8°38' N, 70°12' W, 196 m}, xii 1942, P.J. Anduze leg. [FSCA]; TáchiraState: 13 ♂ 12 ♀, La Fria { 8°13' N, 72°14' W}, 12–18 iv 1920, J.H. & E.B. Williamson leg. [UMMZ]; 1 ♂ 1 ♀, same data but [CSCA]; 5 ♂ 5 ♀, Táchira, San Félix { 8°7' N, 72°15' W}, 6–9 iv 1920, W.H. Ditzler, J.H. & E.B. Williamson leg. [UMMZ]; BolivarState: 1 ♂, Corumo River, 11 kmE of Tumeremo { 7°18' N, 61°30' W, 150 m}, 27–30 vii 1987, T.W. Donnelly leg. [FSCA]; 1 ♂ 1 ♀(in copula), tributaries of Botanamo River, 20–50 kmE of Tumeremo { 7°18' N, 61°7' W, 150 m}, 27 vii 1987, T.W. Donnelly leg. [FSCA]; 1 ♂ 1 ♀, Serranía La Cerbatana, Montañita Arca ( 7°1'33'' N, 66°27'3'' W, D. Parkimon leg. [CSCA]; 6 ♂ 2 ♀(two pairs in copula), Caño Curususuk, 63 kmS of Cuyuni River { 6°15' N, 61°20' W}, 11–12 ii 1976, C.M. & O.S. Flint, Jr. leg. [USNM]. GUYANA, Upper Takutu-Upper EssequiboRegion: 8 ♂ 1 ♀(one pair in copula), Moco Moco, about 30 kmE of Lethem, Kanuku mountains ( 3°18' N, 59°39' W, 160 m), 3–6 iv 1994, O.S. Flint, Jr. leg. [USNM]. ECUADOR, ManabiProv.: 2 ♂, Cojimies { 0°22' N, 80°1' W}, iii 1949, W.C. MacIntyre leg. [UMMZ]; 1 ♂, small forested stream 0.5 kmN of Chindul, highway E15 ( 0°14'20'' S, 79°52'27'' W), 1 iii 2010, K.J. Tennessen leg. [FSCA]; Los RíosProv.: 2 ♂ 2 ♀, Río Palenque Biological Station, 47 kmS of Santo Domingo de los Colorados, forest slough { 0°39' S, 79°15' W}, 18 vii 1977, D.R. Paulson leg. [FSCA]; GuayasProv.: 1 ♂, stream 1.8 kmN of highway 491, about 25 kmE of Babahoyo ( 1°48'40'' S, 79°19'8'' W, 7 iii 2010, K.J. Tennessen leg. [FSCA]; 2 ♂, San Rafael { 2°14' S, 80°1' W, 200 m}, F. Campos R. leg. [FSCA]; El OroProv.: 3 ♂ 1 ♀, Piedras { 3°38' S, 79°55' W, 300 m}, 3–6 vii 1941, D.B. Laddey leg. [UMMZ]; 1 ♂ 1 ♀, same data but [CSCA]. PERU, AyacuchoDept.: La Mar Prov.: 1 ♂, Sivia, ApurimacRiver { 12°30' S, 73°51' W}, 14 vi 1941, F. Woytkowski leg. [UMMZ]; HuanucoDept.: 9 ♂ 9 ♀, Shapajilla, N of Tingo María and NE of Naranjillo { 9°14' S, 75°58' W}, 10 vii–2 viii 1938, F. Woytkowski leg. [UMMZ]; 2 ♂ 2 ♀, same data but [CSCA]; Madre De DiosDept.: 5 ♂ 4 ♀(three pairs in copula), Manu National Park, Pakitza, trail ( 11°55'48'' S, 71°15'18'' W, 250 m), 9 ix 1988, O.S. Flint, Jr. leg. [USNM]; 3 ♂ 3 ♀(two pairs in copula), same data but Aguajal, about 5 kmS of Pakitza 19 ix 1988[USNM]; 1 ♀, same data but trail 2, 20 ix 1988[USNM]; 1 ♂ 1 ♀, same data but T2 base camp to S85, 10 ix 1989, J.A. Louton leg. [USNM]; 1 ♀, P.N. Manu, Pakitza, Aguajal, about 4.5 trail km E, 22 ix 1988[USNM]; 1 ♂, same data but [RWG]; 1 ♂ 1 ♀(in copula), same data but bamboo thicket, 4.3 trail km E, 22 ix 1988, J.A. Louton & N.E. Adams leg. [RWG]; 1 ♀, same data but Trail 1, first stream at marker 14, 19 ix 1989, N.E. Adams leg. [RWG].  Characterization.Male: Epicranium black, black stripes on thorax, dorsum of S1, and distal portion of S6– 10; pale colors red to orange ( Fig. 2), with yellow limited to venter. Pronotum anterior lobe black with medial and lateral orange spots, middle lobe with dorsum orange, and posterior lobe black with medial and lateral orange spots. Mesepisternum black with middorsal carina narrowly orange and orange antehumeral stripe extending along basal 2/3 to 3/4 of sclerite, separated from a small distal orange spot; mesepimeron black with orange humeral stripe along distal 2/3 of sclerite; metepisternum orange with black along posterior margin; metepimeron entirely pale or with a black dorsal spot. Base of leg yellow, distal 1/3 of femur and tibia, and entirety of tarsi dark brown to black, basal 2/3 of femur and tibia with one and two diffuse brownish rings respectively, extensor surface of femur and flexor surface of tibia sometimes dark brown to black as well; tibial spurs shorter than twice intervening spaces. Genital ligula lacking lateral lobes and with distal margin slightly concave, with margins of distal segment spiculate, and laterodistal corners not projected ventrally ( Fig. 72). Cercus shorter than S10 length, about half as long as 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. 98a), 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. 98b, c). Paraproct about as long as S10 length, at midlength narrowing to half its basal width and ending in a triangular tip ( Fig. 98). TL 35.5–40; Hw 17–18. Female: As male but pale colors yellow ( Fig. 26); antehumeral stripe only extending along basal 1/4–1/3 of mesepisternum, and S1–10 black dorsally; pale lateral area of S9 extended dorsally medially to about 1/2 of segment height. Middle lobe of pronotum lacking pronounced lateral depressions; posterior lobe entire, with sides straight and tip truncate, directed dorsally. Mesostigmal plate approximately triangular with a central concavity and posteromedial corner adjacent to a mesepisternal horn directed anteriorly ( Fig. 48); no depressions behind mesostigmal plates. TL 36–38; Hw 19.5–22.   Diagnosis.Among species with male paraproct no longer than S10 length,  P. amatoriashares male cercus that is 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. 98; 99; 105; 115); and genital ligula distal segment with lateral margins spiculate only with  P. aurantiaca,  P. cupida, and  P. sulfurata( Figs. 72; 73; 79; 89). It differs from all three by paraproct being about twice as long as cercus ( Fig. 98; paraproct only slightly longer than cercus in the other species, Figs. 99; 105; 115), and by middorsal pterothoracic carina orange ( Fig. 2; black in the other species, Figs. 3; 9; 19). Posteromedial corner of female mesostigmal plate adjacent to a mesepisternal horn directed anteriorly is only shared with  P. peramansand  P. rojiza( Figs. 48; 60–61), but in those other two species the anterior margin of the female mesostigmal plate is expanded into an upright semicircular structure ( Figs. 60–61), whereas in  P. amatoriait is unmodified ( Fig. 48).  Habitat and biology.The typeseries was found in a gulch, on dead twigs over water or on vegetation high above water, and floating or drifting through the air ( Calvert 1907). In Trinidad, adults were found at a small stream, along the shadow of masses of bamboo roots forming vertical banks over pools of deeper water ( Williamson 1915), and in Costa Ricaalong a large, exposed temporary river with shallow riffle and deep pooled area ( Brooks 1989). Calvert (1907)observed that the male fills its seminal vesicle after capturing a female, and female oviposits (attended by male which stands upright, bringing apex of abdomen directly under thorax) in floating woody twigs, usually at broken ends of the twigs, and that many pairs congregate on the same litter. Larison (2007)observed females ovipositing alone on occasion, and determined that males reversibly use two tactics to gain matings: sit and wait in the canopy for passing females or hover over the water and attempt to grab females that are ovipositing in floating debris. She determined that use of the hovering tactic was greater under high-light than low-light conditions and at higher densities of ovipositing females, and density of ovipositing females was correlated with both light conditions and availability of oviposition substrate. The larva is still undescribed.   Distribution.This is one of the most widely distributed species in the genus, ranging from  Belizeand  Guatemalasouth to Peruand Brazil( Fig. 123). Assessed as Least Concern by IUCN ( Paulson 2009a). 1703062539 [151,1270,1773,1798] 1905-02-28 UMMZ E. B. Williamson Honduras San Pedro Sula 9 10 1 1 holotype