Protoneura amatoria Calvert 1907
Creators: name: Ellenrieder, Natalia Von
name: Garrison, Rosser W.
Publisher: name: Zenodo
Year: 2017
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Odonata
Protoneuridae
Protoneura
Protoneura amatoria

description: Protoneura amatoria Calvert, 1907
Figs. 2 (♂ habitus), 26 (♀ habitus), 48 (♀ mes. plate), 72 (gen. lig.), 98 (♂ app.), 123 (map)
Protoneura amatoria Calvert, 1907: 394, 395, pl. 10, Figs. 49–52 (in key, description of ♂ and ♀ from Honduras and 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; type information);— 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. Williamson leg. [UMMZ].
Specimens examined. 351 ♂ 165 ♀: BELIZE, Toledo Dist.: 1 ♀, deep river at S Highway {16°14' N, 89°1' W}, 7 vi 1993, S.W. Dunkle leg. [FSCA]. GUATEMALA: Izabal Dept.: 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]; Zacapa Dept.: 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, Cortés Dept.: 9 ♂ 4 ♀ (paratypes), San Pedro Sula, 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, Chontales Dept.: 1 ♂, route 7, km 183, 4.2 mi W of Villa Somosa {12°3' N, 84°59' W}, 29 vii 1967, O.S. Flint, Jr. & M.A. Ortiz B. leg. [USNM]; Granada Dept.: 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, Guanacaste Prov.: 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]; Alajuela Prov.: 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 & Tristán leg. [UMMZ]; 1 ♂, same data but [FSCA]; Heredia Prov.: 1 ♂, Sarapiquí, small flood plain pool at small clear water river 9.7 km SE of Puerto Viejo {10°22' N, 83°55' W}, 2 iii 1987, S.W. Dunkle leg. [FSCA]; Puntarenas Prov.: 1 ♂ 1 ♀ (in tandem), stony muddy creek about 11 km N 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 km N of entrance to Miramar on Panamerican Highway {10°5' N, 84°43' W}, 16 iii 1990, S.W. Dunkle leg. [RWG]; 1 ♀, 8.1 mi WNW 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 km NW 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 km E 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, Panamá Prov.: 2 ♂, Chepo Distr., 2.3 km E 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 mi NW 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 km NW 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 km SE 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 ♂, Trinidad River {8°8' N, 81°14' W}, 1–10 vi 1912, A. Busck leg. [USNM]; Colón Prov.: 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]; Panama Oeste 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, Magdalena Dept.: 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]; Bolivar Dept.: 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]; Choco Dept.: 4 ♂ 1 ♀, Sautata {7°50' N, 77°4' W}, 27 i 1918, M.A. Carriker leg. [UMMZ]; 1 ♂ 1 ♀, same data but [CSCA]; Antioquia Dept.: 2 ♂, Puerto Berrio {6°30' N, 74°24' W}, 31 i 1917, J.H. & E.B. Williamson leg. [UMMZ]; 17 ♂ 4 ♀, Cristalina, Magdalena river, about 60 km S of Puerto Barrio {6°24'43'' N, 74°45'38'' W, 652 m}, 12–20 ii 1917, J.H. & E.B. Williamson leg. [UMMZ]; Tolima Dept.: 2 ♂ 1 ♀, Mariquita {5°12' N, 74°54' W, 450 m}, 4 ii 1917, J.H. & E.B. Williamson leg. [UMMZ]. TRINIDAD, Diego Martin Co.: 2 ♂ 1 ♀, Diego Martin river 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 Grande road (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 ♀, Arima River at Churchill– Roosevelt highway, about 2.25 mi SSE 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 mi W of Navet Dam (10°23'43'' N, 61°16'14'' W, 63 m), 14 i 1981, R.W. Garrison leg. [RWG]. VENEZUELA, Falcón State: 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]; Zulia State: 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]; Yaracuy State: 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]; Carabobo State: 4 ♂ 2 ♀, San Esteban {10°25' N, 68°4' W}, 1–5 ii 1920, W.H. Ditzler, J.H. & E.B.Williamson leg. [UMMZ]; Barinas State: 1 ♂, Barinas {8°38' N, 70°12' W, 196 m}, xii 1942, P.J. Anduze leg. [FSCA]; Táchira State: 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]; Bolivar State: 1 ♂, Corumo River, 11 km E 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 km E 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 km S 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 Essequibo Region: 8 ♂ 1 ♀ (one pair in copula), Moco Moco, about 30 km E of Lethem, Kanuku mountains (3°18' N, 59°39' W, 160 m), 3–6 iv 1994, O.S. Flint, Jr. leg. [USNM]. ECUADOR, Manabi Prov.: 2 ♂, Cojimies {0°22' N, 80°1' W}, iii 1949, W.C. MacIntyre leg. [UMMZ]; 1 ♂, small forested stream 0.5 km N of Chindul, highway E15 (0°14'20'' S, 79°52'27'' W), 1 iii 2010, K.J. Tennessen leg. [FSCA]; Los Ríos Prov.: 2 ♂ 2 ♀, Río Palenque Biological Station, 47 km S of Santo Domingo de los Colorados, forest slough {0°39' S, 79°15' W}, 18 vii 1977, D.R. Paulson leg. [FSCA]; Guayas Prov.: 1 ♂, stream 1.8 km N of highway 491, about 25 km E 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 Oro Prov.: 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, Ayacucho Dept.: La Mar Prov.: 1 ♂, Sivia, Apurimac River {12°30' S, 73°51' W}, 14 vi 1941, F. Woytkowski leg. [UMMZ]; Huanuco Dept.: 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 Dios Dept.: 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 km S 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. amatoria shares 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. peramans and 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. amatoria it is unmodified (Fig. 48).
Habitat and biology. The type series 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 Rica along 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 Belize and Guatemala south to Peru and Brazil (Fig. 123). Assessed as Least Concern by IUCN (Paulson 2009a).
description: Published as part of Ellenrieder, Natalia Von & Garrison, Rosser W., 2017, A synopsis of the Neotropical genus Protoneura (Odonata: Coenagrionidae), pp. 1-76 in Zootaxa 4361 (1) on pages 10-13, DOI: 10.11646/zootaxa.4361.1.1, http://zenodo.org/record/1069337