Protoneura woytkowskii Gloyd, 1939

Ellenrieder, Natalia Von & Garrison, Rosser W., 2017, A synopsis of the Neotropical genus Protoneura (Odonata: Coenagrionidae), Zootaxa 4361 (1), pp. 1-76 : 43-50

Protoneura woytkowskii Gloyd, 1939

Figs. 22–24 (♂ habitus), 46 (♀ habitus), 68–70 (♀ mes. plate), 94–96 (gen. lig.), 118 (♂ app.), 125, 126 (map)

Protoneura woytkowskii Gloyd, 1939: 8 –17, pl. 1, Figs. 7–15 ; pl. 2, Figs. 6–10 (description of ♂ and ♀, illustrations color pattern, ♂ S10, and genital ligula, diagnosis);— Cowley (1941: 157, 160, 162, 163; descriptive comments, measurements of legs, key to species in tenuis - group, map); — Soukup (1954: 13; Peru);— Louton et al. (1986: 438; Peru);— Daigle (1996: 11; Ecuador);— Garrison et al. (2003: 24; type information);— Tennessen (2006: 14; Ecuador);— Hoffmann (2009: 63; Peru);— Garrison et al. (2010: 379);—von Ellenrieder & Garrison (2011: 44; Ecuador); Pessacq et al. (2012: 70; Brazil);— Pessacq et al. (2015: 65; Bolivia).

Primary types. Holotype ♂. Peru, San Martin Dep., Vicinity of Rioja, Soritor , 900 m. a.s.l, 13 x 1936, F. Woytkowski leg. [ UMMZ] .

Specimens examined. 276 ♂ 39 ♀: ECUADOR, Sucumbios Prov. : 1 ♂, 2 km NE of Gonzalo Pizarro village, 200 m from Blanco River bridge, small swampy tricke in secondary forest (0°0'11'' S, 77°22'34'' W, 530 m), 23 i 2010, D. Wagner, W. Haber, M. Thomas leg. [RWG] ; 1 ♂, Aguarico (0°5'6'' N, 76°59'33'' W, 458 m), 16 viii 1975, A. Langley leg. [ USNM] ; 1 ♂, stream about 3 km W of Shushufindi (0°9'59'' S, 76°41'30'' W, 290 m), 14 x 2009, J. Johnson leg. [ FSCA] ; 1 ♂, Limoncocha , forest small stream {0°24' S, 76°36' W, 300 m}, 24 viii 1980, S.W. Dunkle leg. [ FSCA] ; 1 ♂ 1 ♀ (in tandem), La Selva, 100 km E Coca, streams in hill forest S of Napo River {0°32' S, 76°32' W, 262 m}, 13 x 1988, S.W. Dunkle leg. [RWG]; Orellana Prov. : 1 ♂, Concepción , Napo River, 28 xii 1939, W. Clarke-MacIntyre leg. [ UMMZ] ; 1 ♂, Indillama River , 19 xii 1938, W. MacIntyre leg. [ UMMZ] ; 1 ♂, forest stream about 12 km NE of Loreto {0°37'18'' S, 77°15'36'' W, 370 m}, 12 x 2009, K.J. Tennessen leg. [ FSCA] ; 2 ♂, Tiputini Biodiversity Station USFQ, stream crossing Matapalo trail (0°37'46'' S, 76°9'16'' W, 246 m), 14– 20 i 2009 [ USFQ] ; 4 ♂, same data but [ CSCA] ; 5 ♂, same data but [RWG] ; 2 ♂ 1 ♀, same data but stream crossing Chorongo trail (0°38'16'' S, 76°8'55'' W, 224 m), 12–16 i 2009 [ CSCA] ; 2 ♂ 1 ♀ (one pair in tandem), same data but [RWG] ; 2 ♂, same data but [ USFQ] ; 3 ♂ 1 ♀ (one pair in tandem), same data but black water stream crossing Guacamayo trail at marker 2050 (0°38'18'' S, 76°8'57'' W, 220 m), 16–17 i 2009, N. von Ellenrieder & R.W. Garrison leg. [ CSCA] ; 4 ♂ 1 ♀ (one pair in tandem), same data but [RWG] ; 2 ♂, same data but [ USFQ] ; 5 ♂ 2 ♀ (one pair in tandem), same data but stream crossing Numa trail at marker 100, muddy bottom, shady (0°38'18'' S, 76°0'37'' W, 220 m), 10 i 2009 [ CSCA] ; 14 ♂ 4 ♀ (one pair in tandem), same data but [RWG] ; 2 ♂, same data but swamp at marker 650 of Parahuaco trail (0°38'23'' S, 76°8'43'' W, 229 m), 21 i 2009 [RWG] ; 2 ♂, Yasuni National Park, small stream, km marker 38 on Yasuni Road (0°38' S, 76°30' W, 12 vi 1995, K.J. Tennessen leg. [ FSCA]; 1 ♂, Yasuni National Park , small stream and pond, 3.2 km W Yasuni Rd on PC 29-30 (0°40'17'' S, 76°24'2'' W, 220 m), 10 vi 1995, S.W. Dunkle leg. [ FSCA] ; 2 ♂ 1 ♀ (one pair in tandem), Yasuni N.P., stream on road to Waorani Reserve, 1.8 km W of Waorani Reserve road, small stream on Markus Oil PC 29-30), 16 xi 1997, W.F. Mauffray leg. [ FSCA] ; 1 ♂ 1 ♀ (in tandem), same data but 18 vii 1996, K.J. Tennessen leg. [ FSCA] ; 1 ♂, Yasuni Biological research Station, along trails and roads within 5 km of Research Station (0°42'S, 76°28' W, 200 m), 17 vi 1996, W.F. Mauffray leg. [ USNM] ; 1 ♂, pond, road to Waorani Reserve, 6.2 km S of turn to Yasuni Research Station (0°42'43'' S, 76°28'28'' W, 230 m), 10 vi 1995, K.J. Tennessen leg. [ FSCA] ; 3 ♂, Shiripuno River, Shiripuno Lodge, 26 km SE of Shiripuno bridge, forest stream 1 m across in a sun fleck (1°6'17'' S, 76°43'54'' W, 235 m), 23–26 i 2009, D. Wagner & W. Haber leg. [RWG]; Napo Prov. : 1 ♂ (paratype), Chucapi River {0°57' S, 77°45' W, 599 m}, 26 ix 1934, W. Clarke-Macintyre leg. [ UMMZ] ; 1 ♂, Cotopino {1°0' S, 78°42' W, 400 m}, 1–28 ii 1950, W. Clarke-Macintyre leg. [ UMMZ] ; 33 ♂ 2 ♀ (paratypes), Jatun Yacu River {1°1' S, 77°50' W, 700 m}, i 1935 – iii 1937, W. Clarke-Macintyre leg. [ UMMZ] ; 2 ♂ 1 ♀ (paratypes), same data but [RWG] ; 3 ♂ 1 ♀ (paratypes), same data but [ CSCA] ; 3 ♂ (paratypes), Napo River watershed, i 1937, W. Clarke-Macintyre leg. [ UMMZ] ; 1 ♂ (paratype), same data but 20 xi 1935 [ UMMZ] ; 1 ♂ 1 ♀, same data but [RWG] ; 1 ♂ (paratype), Anzu River {1°12' S, 77°50' W, 700 m}, iv 1937, W. Clarke-Macintyre leg. [ UMMZ] ; 9 ♂ 2 ♀, small stream half way between Huagra-yacu and Churi-yacu , 500 m within the jungle, W. Clarke-Macintyre leg. [ UMMZ] ; 19 ♂ 2 ♀, Churi-yacu , 18 iii–23 iv 1941, 900 m, W. Clarke-Macintyre leg. [ UMMZ] ; 1 ♂ 1 ♀ (in copula), same data but [RWG] ; 1 ♂ 1 ♀, (in copula), same data but [ CSCA] ; 1 ♂, Arajuno River, Dayuma Camp {1°6' S, 77°35' W, 434 m}, 22–25 ix 1990, J. Belle leg. [RWG] ; 53 ♂ 4 ♀, Headwaters of Arajuno River, Camp Souder , Napo watershed, stream {4°46' S, 77°32' W, 1,000 m}, 22 iv–2 v 1941, W. Clarke-Macintyre leg. [ UMMZ] ; 1 ♂ 1 ♀, same data but [ CSCA] ; 1 ♂ 1 ♀, same data but [RWG]; Morona Santiago Prov. : 17 ♂ 1 ♀, Shicayacu River {1°5' S, 78°4' W, 900 m}, iv–vi 1941, W. Clarke-MacIntyre leg. [ UMMZ] ; 1 ♂ 1 ♀, same data but [ CSCA]; Pastaza Prov. : 5 ♂ 1 ♀ (paratypes), El Partidero, indian village on E bank of Anzu River , junction of trails from Mera and Puyo northward to Napo {1°22' S, 78°0' W, 1,000 m}, 12 ix–14 xi 1935, W. Clarke-Macintyre leg. [ UMMZ] ; 1 ♂ (paratype), same data but [ CSCA] ; 1 ♂, Mera (1°28' S, 78°8' W, 28 ii 1946, W. Clarke-Macintyre leg. [ UMMZ]; 3 ♂, Puyo (1°28' S, 77°59' W, 1,000 m), 2 i–2 iii 1950, W. Clarke-MacIntyre leg. [ UMMZ] ; 1 ♂, same data but [ CSCA] ; 2 ♂, Pastaza River, near Puyo , xii 1948, W. Clarke-Macintyre leg. [ UMMZ] ; 3 ♂, same data but [RWG] ; 1 ♂, Los Copales, rivulet, trails and forest N of Metolo River, between Mera and Shell (1°29'31'' S, 78°4'21'' W, 1,080 m), 20 ix 2005, K.J. Tennessen leg. [ FSCA] ; 1 ♂, same data but 22 ix 2005, W.F. Mauffray leg. [ FSCA]. PERU, Loreto Dept. : 2 ♂, Explorama Lodge, 50 mi NE Iquitos on Amazon River at junction with Yanamono River {3°21' S, 72°47' W, 103 m}, 3 vii 1990, S.W. Dunkle leg. [ FSCA] ; 1 ♂, Aguaitia {9°1' S, 75°31' W, 170 m}, 2 x 1946, F. Woytkowski leg.

[ UMMZ]; San Martin Dept. : 1 ♂ (paratype), Rioja {6°2' S, 77°10' W, 900 m}, P. Martin leg. [UMMZ]; 5 ♂ (paratypes), vicinity of Rioja (Soritor), small muddy brook in shade in jungle, 13 ix 1936, F. Woytkowski leg. [UMMZ]; 1 ♂ (paratype), vicinity of Moyobamba {6°3' S, 76°58' W, 890 m}, 26 xii 1936, F. Woytkowski leg. [UMMZ]; 1 ♂ (paratype), same data but [ FSCA]; Huanuco Dept. : 4 ♂, Pumahuasi, 20 km N of Tingo Maria {8°46' S, 75°49' W}, 12–19 xii 1939, F. Woytkowski leg. [ UMMZ] ; 1 ♂, same data but [ CSCA] ; 11 ♂, Cucharas, tropical jungle in the Huallaga river valley {9°18' S, 75°59' W, 500 m}, ii 1954, F. Woytkowski leg. [ UMMZ] ; 1 ♂ 1 ♀, same data but [ CSCA]; Ucayali Dept. : 3 ♂, Boquerón del Padre Abad, NE of Tingo María {9°4' S, 75°41' W, 440 m}, 23–25 viii 1946, F. Woytkowski leg. [ UMMZ]; Madre De Dios Dept .: 1 ♂, Manu, Pakitza, Aguajal about 3 km E of Pakitza (11°55'48' S, 71°15'18'' W, 250m), 18–19 x 1987, J.A. Louton leg. [RWG] ; 1 ♀, same data but trail 1, stream at marker 14, N.E. Adams leg. [RWG] ; 1 ♂ 1 ♀, same data but trail T2, stream at marker 15 [ USNM] ; 1 ♂ 1 ♀ (in copula), same data but T1 toT2 to basecamp, 11 ix 1989, J.A. Louton leg. [ USNM] ; 1 ♂, Tambopata, Trocha C, pump {13°8' S, 69°37' W}, 27 vi 2008, C. Beatty leg. [ CSCA]. BOLIVIA, Beni Dept. : 2 ♂, National Park and Indian Land Isiboro-Sécure, pedro stream, tributary of Pluma river , road from Coruya to Pluma lodge (16°02'49'' S, 66°15'11'' W), 8–10 xi, P. Pessacq leg. [ CSCA]; Cochabamba Dept. : 1 ♂ 1 ♀ (in tandem), small stream N of Shinahota (16°59'18'' S, 65°13'95'' W, 265 m), 22 xi 1999, K.J. Tennessen leg. [ FSCA]; Dept. Santa Cruz Dept. : 1 ♂ (paratype), Buena Vista {17°27' S, 63°40' W, 400 m}, ii 1933, F. Steinbach leg. [UMMZ]; 1 ♂, Amboro National Park, Yapacani, streams and tributaries along road to Matarao Encampment, S side of Yapacani River (17°37' S, 63°37' W, 457 m), 12 xi 1998, W.F. Mauffray leg. [ FSCA] ; 1 ♂ 1 ♀, same data but Matarac (17°37' S, 63°47' W, 609 m), 12 xi 1998, K.J. Tennessen leg. [ FSCA].

Characterization. Male: Epicranium black with copper to greenish metallic reflections, dorsum of thorax black with reddish orange spots, and dorsum of abdomen black with basal half of S2, distal half of S9 and sometimes laterodorsal spots on S10 orange, narrow yellow to blue basal spots on S3–7; pale colors reddish-orange and yellow to pale blue ( Fig. 22–24 ). Pronotum orange with sides of middle lobe black, and medial portion of posterior lobe sometimes black. Mesepisternum black with broad reddish orange stripe along humeral suture extending along entire length of sclerites or interrupted subapically by black, and orange stripe over middorsal carina; mesepimeron black with orange spot on posterodistal corner; metepisternum orange with posterodistal corner black; metepimeron anterior half black and posterior half yellow. Coxa and trochanter yellow, basal 2/3 of femur yellow, tibia yellow with distal 1/3 to 1/4 black and a diffuse subbasal grayish-brown ring, remainder of leg black; tibial spurs shorter than twice intervening spaces. Genital ligula with well-developed lateral lobes and with tip of distal segment projected distally into a long strap-like process, about as long as or longer than remainder of distal segment distal segment ( Figs. 94–96 ). Cercus slightly longer than S10 length, 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. 118a ), 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 tooth, visible in dorsal and mediodorsal views ( Figs. 118b, c ). Paraproct longer than S10 length, at basal third narrowing to half its basal width and digitiform ( Fig. 118 ). TL 40.5–42.5; Hw 18–20.

Female: As male but pale colors pale yellow; pronotum black with margins yellow; mesepisternum black with pale yellow upper humeral stripe along basal 3/4 to 4/5 of suture and middorsal carina yellow along distal 1/2 to 3/ 4; pale area of S9 extended dorsally medially to about 1/2 of segment height ( Fig. 46 ). 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 a subapical bulge on anterior margin and posteromedial corner adjacent to a small mesepisternal tubercle ( Figs. 68–70 ). TL 34–37; Hw 19–21.

Diagnosis. Protoneura woytkowskii 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. tenuis ( Figs. 101 ; 107–108; 114; 116; 118). Among them, it shares male cercus and paraproct longer than S10 length with P. calverti and P. tenuis ( Figs. 101a ; 116a; 118a), and male can be distinguished from them by the presence of an orange stripe along middorsal carina, tibiae pale with dark apices, and venter of thorax entirely pale ( Figs. 22–24 ); in P. calverti and P. tenuis male lacks a middorsal pale stripe on pterothorax, tibiae are dark, and there are dark markings on venter of thorax ( Figs. 5 ; 20). Protoneura woytkowskii further differs from P. calverti and P. tenuis by genital ligula morphology, with tip of distal segment projected distally into a long strap-like process, about as long as or longer than remainder of distal segment distal segment wider than long in ectal view from flexure to tip ( Figs. 94–96 ), which is straight and not projected distally in the other two species ( Figs. 75 ; 90–92). Female of P. woytkowskii shares an approximately triangular and flat mesostigmal plate with P. klugi , P. macintyrei , P. sulfurata , and P. tenuis ( Figs. 57–58 ; 65; 68– 70 ). It can be distinguished from P. klugi and P. tenuis by tibiae pale with dark apices, and venter of thorax entirely pale, which are entirely dark and with dark markings respectively in the other three species; it differs from P. macintyrei and P. sulphurata by anterior margin of mesostigmal plate with a subapical bulge ( Figs. 68–70 ), which is lacking in the other two species ( Figs. 58 ; 65 ).

Remarks. Gloyd (1939) described this species based on specimens from Peru, Ecuador, and Bolivia, and detailed differences in color pattern between the Bolivian specimen and the others. Later Cowley (1941) noted that the two color forms described by Gloyd probably represented subspecies, and called the darker specimens from Ecuador and N Peru ‘form a’, and the paler Bolivian specimen ‘form b’. He ( Cowley 1941: 163) also noted differences in genital ligula and male appendages between one male from Peru and one from Bolivia.

All examined genital ligulae that are not broken have a long distal strap-like process, in many cases folded unnaturally during preservation, and as Cowley (1941) already pointed out, Gloyd’s illustration (1939: Plate II, Figs. 9–10 ) of the genital ligula of P. woytkowskii was based on a specimen in which the distal strap was broken.

Examination of a large series of specimens allows us to conclude that the morphological differences pointed out by Cowley (1941) in the shape of the medial laminar lobe of cercus do not hold, representing variability in the shape, also partly due to postmortem preservation, of this structure. We did however find differences in the shape of genital ligula, female mesostigmal plate, and color pattern which loosely correlate with distribution ( Fig. 126 ). Specimens from the western portion of Ecuador and northern Peru, corresponding to ‘form a’ of Cowley (1941), have genital ligula with a relatively wider and shorter strap ( Fig. 94 ), female mesostigmal plate narrow triangular without a posterior vertical surface and no mesespisternal tubercle or bump associated with its medioposterior corner ( Fig. 68 ), male pronotum hind lobe entirely orange or with only base or apex of medial portion black, pale stripe over middorsal carina extended along posterior 1/2 to 2/3 of carina, male orange humeral stripe complete to interrupted distally, and dorsum of male S10 lacking pale spots ( Fig. 22 ). Specimens from the eastern portion of Ecuador and Peru, which we are calling ‘form c’, have genital ligula with a relatively longer and narrower strap ( Fig. 96 ), female mesostigmal plate short and wide without a posterior vertical surface and with a mesespisternal tubercle associated with its medioposterior corner ( Fig. 70 ), male pronotum hind lobe with medial portion entirely black, pale stripe over middorsal carina extended along posterior 1/2 of carina, male orange humeral stripe interrupted distally, and dorsum of male S10 usually lacking pale spots ( Fig. 24 ). Specimens from southern Peru and Bolivia, ‘form b’ of Cowley, share the genital ligula with a relatively longer and narrower strap ( Fig. 95 ) and male pronotum hind lobe with medial portion entirely black with form c, pale stripe over middorsal carina extended along posterior 2/3 to almost full length of carina and male orange humeral stripe complete with form a, but differ from both by female mesostigmal plate with a posterior vertical surface and with a low mesespisternal bump associated with its medioposterior corner ( Fig. 69 ) and dorsum of male S10 with orange spots ( Fig. 23 ).

We consider that all specimens represent the same species, and interpret the described differences as intraspecific geographic variability expressed in a mosaic of characters in three morphs. It is unknown if the one female recorded from Brazil ( Pessacq et al. 2015) fits one of these morphs or not.

Habitat and biology. Adults are found at small muddy shaded brooks ( Gloyd 1939) and small streams in primary forest ( Louton et al. 1986; von Ellenrieder & Garrison 2011).

Distribution. Eastern Ecuador, Peru, Brazil, and Bolivia ( Figs. 125–126 ).

Figures

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 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 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 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 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 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 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 125–126. Distribution maps.