Protoneura tenuis Selys, 1860

Ellenrieder, Natalia Von & Garrison, Rosser W., 2017, A synopsis of the Neotropical genus Protoneura (Odonata: Coenagrionidae), Zootaxa 4361 (1), pp. 1-76 : 39-41

Protoneura tenuis Selys, 1860

Figs. 20 (♂ habitus), 44 (♀ habitus), 66 (♀ mes. plate), 90–92 (gen. lig.), 116 (♂ app.), 124 (map)

Protoneura tenuis Selys, 1860: 461 , 462 [reprint 33, 34] (description of ♂ and ♀ from Pará);—Selys (1886: 210–211, record with doubts from Brazil);—Kirby (1890: 136); — Cowley & Gloyd (1938: 1–18, pl. 1 Figs. 1–6 , pl. 2 Figs. 1–8 , pl. 3 Figs. 1–11 , pl. 4 Figs. 1–8 ; redescription of types ♂ and ♀, species variability, diagnosis from P. calverti );— Cowley (1941: 146, 153, 157, 158, 160-165, 167-170, 172, fig. 24; descriptive comments, measurements legs, key to species in ‘tenuis’ group, map, illustration of ♀ outer valve of ovipositor);— Rácenis (1955: 55; Venezuela);— Garrison (1989: 1; Brazil);— De Marmels (1992: 40; Venezuela);— Butt (1995: 95; Peru);— Belle (2002: 2; Suriname);— Rehn (2003: 212; in phylogenetic analysis);— Lencioni (2005: 224; Brazil);— Hoffmann (2009: 63; Peru);—von Ellenrieder (2009c; IUCN assessment);— Garrison et al. (2010: 379);—von Ellenrieder (2011: 59; Suriname);— Pessacq et al. (2012: 70;Brazil);— Batista Calvão et al. (2014: 305; Brazil);— Michalski (2015: 76–78; Trinidad, photos, key);—von Ellenrieder (2017: 119, 200, 205; Guyana);—von Ellenrieder et al. (2017: 8; Guyana).

Protoneura ? tenuis — Williamson (1915: 619, 627).

Protoneura calverti nec Williamson, 1915 — Geijskes (1932: 238–239; Trinidad).—Misidentification.

Primary types. ♂ Lectotype: Pará, 1850 [OUMNH]. Since the species was described by Selys based on male and female syntypes, the male redescribed by Cowley & Gloyd (1938) as holotype actually constitutes the lectotype.

Specimens examined. 123 ♂ 59 ♀: VENEZUELA, Bolivar State : 6 ♂ 2 ♀, Churún River at entrance to Salto Angel (5°58'3'' N, 62°32'8'' W, 172 m), 10–12 viii 1990, R.W. Garrison leg. [RWG] ; 1 ♂ 1 ♀ (in copula), same data but T.W. Donnelly leg. [ FSCA] ; 1 ♂, Cinco Ranchos River , 15 km E of El Pauji (4°36'4'' N, 61°24'54'' W, 898 m), 7 viii 1990, R.W. Garrison leg. [RWG]; Amazonas State : 1 ♀, Alto Mavaca Base Camp, in primary rain forest understory (2°1'30'' N, 65°7'0'' W, 228 m), 2 ii 1989, D.A. Polhemus leg. [USNM]; 1 ♂ 1 ♀, Siapa 2 Camp, tributary to Siapa River {1°43' N, 64°30' W, 635 m}, 5 ii 1989, D.A. Polhemus leg. [ USNM]. TRINIDAD, St. George Co .: 1 ♂, creek 3.3 mi W of Blanchisseuse {10°48' N, 61°20' W, 31 m}, 15 i 1981, R.W. Garrison leg. [RWG] ; 1 ♀, Yarra River , 1 km SE of Filette {10°47' N, 61°21' W}, 25 vi 1993, O.S. Flint, Jr. & N.E. Adams leg. [ USNM] ; 1 ♂, Arima River tributary, 3 mi S of Arima {10°43' N, 61°17' W, 480 m}, 4 iv 1965, T.W. Donnelly leg. [ FSCA]; St. Andrew Co .: 1 ♂, road 7 mi E Valencia, tributary Oropuche River {10°38' N, 61°6' W, 20 m}, 9 vii 1970, T.W. Donnelly leg. [ FSCA] ; 2 ♂, 1.9 mi W of Cumuto , forest small stream {10°35' N, 61°12' W, 57 m}, 8 iv 1975, K.W. Knopf leg. [RWG]. GUYANA, Potaro-Siparuni Region : 1 ♂ 1 ♀, Elinkwa creek (5°8'55'' N, 59°28'28'' W, 438 m), 28 iii 2014, N. von Ellenrieder & W. Washington leg. [ CSCA] ; 2 ♂, Amerindian village Paramakatoi (4°41'57'' N, 59°42'48'' W, 720 m), 25 viii 1997, O.S. Flint, Jr. leg. [ USNM]. SURINAME, Sipaliwini Dist. : 5 ♂ 3 ♀ (two pairs in tandem), forest creek, Werehpai Camp (2°21'46'' N, 56°41'53'' W, 258 m), 4–6 ix 2010, N. von Ellenrieder leg. [CSCA]; 3 ♂ 2 ♀ (one pair in tandem), same data but [RWG] ; 2 ♂ 1 ♀ (one pair in tandem), same data but [ NZCS] ; 5 ♂ 2 ♀ (one pair in tandem), forest creek, Sipaliwini River Camp (2°17'40'' N, 56°36'13'' W, 208 m), 28 viii–1 ix 2010, N. von Ellenrieder leg. [RWG] ; 3 ♂ 1 ♀ (one pair in tandem), same data but [ CSCA] ; 1 ♂, forest creek, Kutari River Camp (2°10'17'' N, 56°46'40'' W, 239 m), 20 viii 2010, N. von Ellenrieder leg. [RWG] ; 1 ♂, same data but (2°10'21'' N, 56°46'32'' W, 248 m), 21 viii 2010 [ CSCA] ; 1 ♂ 1 ♀, same data but (2°10'31'' N, 56°46'56'' W, 263 m), 23–24 viii 2010 [ CSCA] ; 1 ♂ 1 ♀ (in tandem), forest creek N of Kutari River Camp (2°11'6'' N, 56°47'17'' W, 229 m), 22 viii 2010, N. von Ellenrieder leg. [CSCA]; 1 ♂ 1 ♀ (in tandem), same data but [RWG]. PERU: Madre de Dios Dept .: 3 ♂ 1 ♀, Explorer's Inn on Tambopata River , 30 km SW of Puerto Maldonado, sandy forest stream (12°50'12'' S, 69°17'36'' W, 300 m), 22 26 vii 2002, D.R. Paulson & N. Smith leg. [DRP]. BRAZIL, Para State : 3 ♂ 1 ♀, Benevides {1°22' S, 48°15' W, 21 m}, x 1918, S. M. Klages leg. [FSCA]; 25 ♂ 9 ♀, Belem, Igarape beyond Souza {1°27' S, 48°29' W}, 7–14 viii 1922, J.H. Williamson & J.W. Strohm leg. [UMMZ]; 1 ♂ 1 ♀, no locality given, probably Belem [RBINS]; 1 ♂, Obidos {1°54' S, 55°31' W, 81 m}, 1889 [RBINS]; 4 ♂ 3 ♀, Rio Xingu Camp , Igarapé Jabuti, malaise trap during day {3°39' S, 52°22' W, 138 m}, 3–14 x 1986, P. Spangler & O.S. Flint, Jr. [ USNM] ; 2 ♂, same data but [RWG] ; 1 ♂ 1 ♀ (in copula), same data but jungle stream, 2 x 1986 [ USNM] ; 1 ♂ 1 ♀, same data but [RWG] ; 1 ♂, Floresta Nacional de Carajás , collecting point Buritizal I - parte baixa (riacho) (6°6'14'' S, 50°8'13'' W, 600 m), 25 ix 2007, N. Ferreira Jr. & V. Alecrim leg. [ DZRJ] ; 1 ♂, same data but pique-nique Errado, 16 ix 2006, N. Ferreira Jr. & L.L. Dumas leg. [ DZRJ] ; 1 ♂ 1 ♀, same data but 7 iii 2008, N. Ferreira Jr. & A. Santos leg. [ DZRJ] ; 2 ♂ 1 ♀, Jacareacanga {6°16' S, 57°39' W, 88 m}, iv 1970, F.R. Barbosa leg. [UMMZ]; Amazonas State: 13 ♂ 8 ♀, Porto Velho {8°45' S, 63°54' W, 60 m}, 13 ii–24 iv 1922, J.H. Williamson & J.W. Strohm leg. [UMMZ]; 2 ♂ 2 ♀, same data but [ FSCA] ; 1 ♂ 1 ♀, same data but [ CSCA]; Acre State : 1 ♂ 1 ♀ (in tandem), Senador Guiomard, Igarapé Floresta, Fazenda Experimental Catuaba (10°4'20'' S, 67°36'53'' W, 175 m), 25 vii 2008, P.S. Ferreira Peruquetti leg. [RWG]; Rondonia State: 9 ♂ 2 ♀, Villa Murtinho {10°21' S, 65°19' W, 133 m}, 1–7 iv 1922, J.H. Williamson & J.W. Strohm leg. [UMMZ]; 2 ♂ 1 ♀, same data but [ CSCA] ; 1 ♂ 1 ♀ (in tandem), 16 km W of Fazenda Rancho Grande on C-20 Rio Pardo and Linea C-20 (10°20'30'' S, 62°54'0'' W, 500 m), 14 xi 1991, M.J. Westfall, Jr. leg. [ FSCA] ; 2 ♂ 1 ♀ (one pair in copula), Governador Jorge Teixeira Municipality, Fazenda Rancho Grande, farm of Harald Schmitz and environs within about 15 km radius (10°31'48'' S, 62°48'0'' W, 165 m), 16–24 iii 1989, S.W. Dunkle leg. [RWG] ; 1 ♂ 1 ♀ (in tandem), same data but [ FSCA] ; 3 ♂, same data but stream about 7 km SE of Cacaulandia , 7 xi 1989, R.W. Garrison leg. [RWG] ; 1 ♂, same data but [ CSCA] ; 1 ♂, ponds near Fernandez trail, linea C-16, 1 km N Cacaulandia on B-65, 13 xi 1991, M.J. Westfall, Jr. leg. [FSCA]; 1 ♂ 1 ♀ (in tandem), same data but linea C-20, 7 km E of B-65 (10°32' S, 62°48' W, 160 m), 19 xi 1991, M.J. Westfall Jr. leg. [ FSCA]. BOLIVIA, El Beni Dept. : 3 ♂ 3 ♀, Beni River, Cachuela Esperanza {10°32' S, 65°38' W, 122 m}, 10–13 iv 1922, J.H. Williamson & J.W. Strohm leg. [ UMMZ].

Characterization. Male: Epicranium black with copper to greenish metallic reflections and a small triangular yellow spot adjacent to compound eye behind antenna, dorsum of thorax black with reddish orange spots, and dorsum of abdomen black with narrow light blue basal spots on S3–7; pale colors reddish orange, and pale yellow to yellowish blue ( Fig. 20 ). Pronotum black with large reddish orange spot on center of middle lobe. Mesepisternum black with reddish orange triangular spot on basal 3/4 to 4/5 narrowing distally, which branches at about midlength of mesepisternum into a short stripe directed posteriorly along humeral suture; mesepimeron black; metepisternum basal half pale yellow with ventral, posterior and dorsal corners black; metepimeron anterior half black and posterior half pale bluish yellow. Coxa, trochanter, and base of femur yellow, remainder of leg black; tibial spurs shorter than twice intervening spaces. Genital ligula with distal margin approximately straight to vshaped, with laterodistal corners projected ventrally, and with short lateral lobes at about midlength present to absent ( Figs. 90–92 ). Cercus slightly longer than S10 length, slightly shorter than paraproct, about twice as long as wide, with a depression on external dorsolateral surface, dorsal margin convex and ventral margin concave, and with a small ventrobasal blunt tooth in lateral view ( Fig. 116a ), with medial surface slightly concave, with a small blunt ventrobasal tooth, a partially membranous laminar medial lobe with tip located distal to midlength of cercus, and tip curved medially forming a subapical ridge, visible in dorsal and mediodorsal views ( Figs. 116b, c ). Paraproct longer than S10 length, at basal third narrowing to half its basal width and digitiform ( Fig. 116 ). TL 32– 38.5; Hw 16.5–18.

Female: As male but pale colors pale yellow; pronotum black with margins yellow; mesepisternum black with pale yellow narrow upper humeral stripe along basal 3/4 of suture and middorsal carina yellow; mesepimeron black with lower portion of humeral stripe along distal 1/2 to 1/3 of suture interrupted medially; pale area of S9 extended dorsally medially to about 1/3 of segment height. Middle lobe of pronotum lacking pronounced lateral depressions; posterior lobe entire, directed posterodorsally, with sides straight and tip smoothly convex. Mesostigmal plate approximately triangular and flat, with yellow anterior margin of lateral corner produced dorsally into a blunt tubercle ( Fig. 66 ). TL 31.5–34.5; Hw 19–21.

Diagnosis. Protoneura tenuis shares male cercus longer than wide and with a laminar lobe on medial surface with P. calverti , P. klugi , P. macintyrei , P. scintilla , and P. woytkowskii ( Figs. 101 ; 107–108; 114; 116; 118). Among them, it shares male cerci and paraprocts longer than S10 length with P. calverti and P. woytkowskii ( Figs. 101a ; 116a; 118a), and it can be distinguished from them by the design of the orange areas on mesepisternum; in males of P. tenuis and P. calverti they form a triangular spot on basal portion of mespisternum narrowing distally and branching into a short stripe along humeral suture, and middorsal carina is black ( Figs. 5 ; 20), whereas in male of P. woytkowskii they form a broad reddish orange stripe along humeral suture extending along entire length of sclerite or interrupted subapically by black, and there is an orange stripe over middorsal carina ( Figs. 22–24 ). However, in male of P. tenuis the triangular orange spot extends along basal 3/4 to 4/5 of mesepisternum ( Fig. 20 ), and only along basal 1/2 of mesepisternum in male of P. calverti ( Fig. 5 ). Protoneura tenuis further differs from P. calverti and P. woytkowskii by genital ligula morphology, with distal segment wider than long in ectal view from flexure to tip, and laterodistal process occupying 1/3 or more of distal segment length in lateral view ( Figs. 90–92 ), whereas segment is longer than wide in ectal view from flexure to tip and laterodistal process occupies only 1/4 or less of distal segment length in lateral view in P. calverti ( Fig. 75 ), and the tip of distal segment is projected distally into a long strap-like process, about as long as or longer than remainder of distal segment in P. woytkowskii ( Figs. 94–96 ). Female of P. tenuis shares an approximately triangular and flat mesostigmal plate with P. klugi , P. macintyrei , P. sulfurata , and P. woytkowskii ( Figs. 57–58 ; 65; 68–70 ), but it can be distinguished from them by the yellow anteroexternal corner produced dorsally ( Fig. 66 ).

Habitat and biology. This species is found along clear, slow flowing streams running over sand, silt, clay, and gravel substrates in heavy shade in Peru ( Butt 1995). In Trinidad, P. tenuis can be found mating and guarding territories along lower, medium-gradient stretches of montane rivers where the bed is composed of sand or gravel, along bends of the river where the bedrock outcroppings allow the water to become stiller and there are overhanging Maraval lilies (Spatthiphyllum cannifolium) ( Michalski 2015).

Distribution. Trinidad, Venezuela, Suriname, Guyana, and Brazil to Peru and Bolivia ( Fig. 124 ). Assessed as Least Concern by IUCN (von Ellenrieder 2009c).

Figures

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 1–4. Male body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale). (1) P. ailsa; (2) P. amatoria; (3) P.aurantiaca; (4) P.caligata.

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 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 85–91. Genital ligula in ectal (85–91a), lateral (85–91b), and entolateral (85c) views (to scale).(85) P. rojiza; (86) P. romanae; (87)P.sanguinipes; (88) P.scintilla; (89) P.sulfurata; (90–91) P.tenuis. All by RWG.

FIGURES 92–96. Genital ligula in ectal (92–96a), lateral (92–96b), and entolateral (93c) views (to scale). (92) P. tenuis; (93) P. viridis; (94) P. woytkowskii form a; (95) P. woytkowskiiform b; (96) P. woytkowskii form c. All by RWG.

FIGURES 115–118. Male caudal appendages in lateral (115–117a), posterodorsal (115–117b), and dorsal (115–117b) views (not to scale). (115) P.sulfurata; (116) P. tenuis; (117) P.viridis; (118) P.woytkowskii.

FIGURES 64–70. Female pronotum (67a, c) and mesostigmal plates (64–66; 67b, d), in lateral (66a; 67a, b), dorsal (64; 65; 66b; 67c, d; 68–70) and dorsolateral (an) views (to scale). (64) P. scintilla; (65) P. sulfurata; (66) P. tenuis; (67) P. viridis; (68) P. woytkowskii form a; (69) P. woytkowskii form b; (70) P. woytkowskii form c.

FIGURES 100–102. Male caudal appendages in lateral (100–102a), posterodorsal (100–102b), and dorsal (100–102b) views (not to scale). (100) P.caligata; (101) P. calverti; (102) P.capillaris.

FIGURES 21–24. Male body scan; head in dorsal view, thorax and abdomen in lateral view (not to scale). (21) P. viridis; (22) P. woytkowskii form a; (23)P.woytkowskii form b; (24) P.woytkowskii form c.

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 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 123–124. Distribution maps.