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

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https://doi.org/10.11646/zootaxa.4361.1.1

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scientific name

Protoneura woytkowskii Gloyd, 1939
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Protoneura woytkowskii Gloyd, 1939 

Figs. 22–24View FIGURES 21–24 (♂ 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. 7–15View FIGURES 5–8View FIGURES 9–12View FIGURES 13–16; pl. 2, Figs. 6–10View FIGURES 5–8View FIGURES 9–12 (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. [ UMMZAbout 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]GoogleMaps  ; 1 ♂, Aguarico (0°5'6'' N, 76°59'33'' W, 458 m), 16 viii 1975, A. Langley leg. [ USNMAbout USNM]GoogleMaps  ; 1 ♂, stream about 3 km W of Shushufindi (0°9'59'' S, 76°41'30'' W, 290 m), 14 x 2009, J. Johnson leg. [ FSCAAbout FSCA]GoogleMaps  ; 1 ♂, Limoncocha , forest small stream {0°24' S, 76°36' W, 300 m}, 24 viii 1980, S.W. Dunkle leg. [ FSCAAbout FSCA]GoogleMaps  ; 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. GoogleMaps  : 1 ♂, Concepción , Napo River, 28 xii 1939, W. Clarke-MacIntyre leg. [ UMMZAbout UMMZ]  ; 1 ♂, Indillama River , 19 xii 1938, W. MacIntyre leg. [ UMMZAbout 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. [ FSCAAbout FSCA]GoogleMaps  ; 2 ♂, Tiputini Biodiversity Station USFQ, stream crossing Matapalo trail (0°37'46'' S, 76°9'16'' W, 246 m), 14– 20 i 2009 [ USFQ]GoogleMaps  ; 4 ♂, same data but [ CSCAAbout CSCA]GoogleMaps  ; 5 ♂, same data but [RWG]GoogleMaps  ; 2 ♂ 1 ♀, same data but stream crossing Chorongo trail (0°38'16'' S, 76°8'55'' W, 224 m), 12–16 i 2009 [ CSCAAbout CSCA]GoogleMaps  ; 2 ♂ 1 ♀ (one pair in tandem), same data but [RWG]GoogleMaps  ; 2 ♂, same data but [ USFQ]GoogleMaps  ; 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. [ CSCAAbout CSCA]GoogleMaps  ; 4 ♂ 1 ♀ (one pair in tandem), same data but [RWG]GoogleMaps  ; 2 ♂, same data but [ USFQ]GoogleMaps  ; 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 [ CSCAAbout CSCA]GoogleMaps  ; 14 ♂ 4 ♀ (one pair in tandem), same data but [RWG]GoogleMaps  ; 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]GoogleMaps  ; 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. [ FSCAAbout 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. [ FSCAAbout FSCA]GoogleMaps  ; 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. [ FSCAAbout FSCA]  ; 1 ♂ 1 ♀ (in tandem), same data but 18 vii 1996, K.J. Tennessen leg. [ FSCAAbout 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. [ USNMAbout USNM]GoogleMaps  ; 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. [ FSCAAbout FSCA]GoogleMaps  ; 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.GoogleMaps  : 1 ♂ (paratype), Chucapi River {0°57' S, 77°45' W, 599 m}, 26 ix 1934, W. Clarke-Macintyre leg. [ UMMZAbout UMMZ]GoogleMaps  ; 1 ♂, Cotopino {1°0' S, 78°42' W, 400 m}, 1–28 ii 1950, W. Clarke-Macintyre leg. [ UMMZAbout UMMZ]GoogleMaps  ; 33 ♂ 2 ♀ (paratypes), Jatun Yacu River {1°1' S, 77°50' W, 700 m}, i 1935 – iii 1937, W. Clarke-Macintyre leg. [ UMMZAbout UMMZ]GoogleMaps  ; 2 ♂ 1 ♀ (paratypes), same data but [RWG]GoogleMaps  ; 3 ♂ 1 ♀ (paratypes), same data but [ CSCAAbout CSCA]GoogleMaps  ; 3 ♂ (paratypes), Napo River watershed, i 1937, W. Clarke-Macintyre leg. [ UMMZAbout UMMZ]  ; 1 ♂ (paratype), same data but 20 xi 1935 [ UMMZAbout 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. [ UMMZAbout UMMZ]GoogleMaps  ; 9 ♂ 2 ♀, small stream half way between Huagra-yacu and Churi-yacu , 500 m within the jungle, W. Clarke-Macintyre leg. [ UMMZAbout UMMZ]  ; 19 ♂ 2 ♀, Churi-yacu , 18 iii–23 iv 1941, 900 m, W. Clarke-Macintyre leg. [ UMMZAbout UMMZ]  ; 1 ♂ 1 ♀ (in copula), same data but [RWG]  ; 1 ♂ 1 ♀, (in copula), same data but [ CSCAAbout CSCA]  ; 1 ♂, Arajuno River, Dayuma Camp {1°6' S, 77°35' W, 434 m}, 22–25 ix 1990, J. Belle leg. [RWG]GoogleMaps  ; 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. [ UMMZAbout UMMZ]GoogleMaps  ; 1 ♂ 1 ♀, same data but [ CSCAAbout CSCA]GoogleMaps  ; 1 ♂ 1 ♀, same data but [RWG]; Morona Santiago Prov.GoogleMaps  : 17 ♂ 1 ♀, Shicayacu River {1°5' S, 78°4' W, 900 m}, iv–vi 1941, W. Clarke-MacIntyre leg. [ UMMZAbout UMMZ]GoogleMaps  ; 1 ♂ 1 ♀, same data but [ CSCAAbout CSCA]; Pastaza Prov.GoogleMaps  : 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. [ UMMZAbout UMMZ]GoogleMaps  ; 1 ♂ (paratype), same data but [ CSCAAbout CSCA]GoogleMaps  ; 1 ♂, Mera (1°28' S, 78°8' W, 28 ii 1946, W. Clarke-Macintyre leg. [ UMMZAbout UMMZ]; 3 ♂, Puyo (1°28' S, 77°59' W, 1,000 m), 2 i–2 iii 1950, W. Clarke-MacIntyre leg. [ UMMZAbout UMMZ]GoogleMaps  ; 1 ♂, same data but [ CSCAAbout CSCA]GoogleMaps  ; 2 ♂, Pastaza River, near Puyo , xii 1948, W. Clarke-Macintyre leg. [ UMMZAbout 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. [ FSCAAbout FSCA]GoogleMaps  ; 1 ♂, same data but 22 ix 2005, W.F. Mauffray leg. [ FSCAAbout FSCA]. PERUAbout PERU, Loreto Dept.GoogleMaps  : 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. [ FSCAAbout FSCA]GoogleMaps  ; 1 ♂, Aguaitia {9°1' S, 75°31' W, 170 m}, 2 x 1946, F. Woytkowski leg.GoogleMaps 

[ UMMZAbout 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 [ FSCAAbout FSCA]; Huanuco Dept.  : 4 ♂, Pumahuasi, 20 km N of Tingo Maria {8°46' S, 75°49' W}, 12–19 xii 1939, F. Woytkowski leg. [ UMMZAbout UMMZ]GoogleMaps  ; 1 ♂, same data but [ CSCAAbout CSCA]GoogleMaps  ; 11 ♂, Cucharas, tropical jungle in the Huallaga river valley {9°18' S, 75°59' W, 500 m}, ii 1954, F. Woytkowski leg. [ UMMZAbout UMMZ]GoogleMaps  ; 1 ♂ 1 ♀, same data but [ CSCAAbout CSCA]; Ucayali Dept.GoogleMaps  : 3 ♂, Boquerón del Padre Abad, NE of Tingo María {9°4' S, 75°41' W, 440 m}, 23–25 viii 1946, F. Woytkowski leg. [ UMMZAbout UMMZ]; Madre De Dios DeptGoogleMaps  .: 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 [ USNMAbout USNM]  ; 1 ♂ 1 ♀ (in copula), same data but T1 toT2 to basecamp, 11 ix 1989, J.A. Louton leg. [ USNMAbout USNM]  ; 1 ♂, Tambopata, Trocha C, pump {13°8' S, 69°37' W}, 27 vi 2008, C. Beatty leg. [ CSCAAbout CSCA]. BOLIVIA, Beni Dept.GoogleMaps  : 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. [ CSCAAbout CSCA]; Cochabamba Dept.GoogleMaps  : 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. [ FSCAAbout 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. [ FSCAAbout FSCA]GoogleMaps  ; 1 ♂ 1 ♀, same data but Matarac (17°37' S, 63°47' W, 609 m), 12 xi 1998, K.J. Tennessen leg. [ FSCAAbout FSCA].GoogleMaps 

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. 22–24View FIGURES 21–24). 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. 94–96View FIGURES 92–96). 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. 118aView FIGURES 115–118), 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, cView FIGURES 115–118). Paraproct longer than S10 length, at basal third narrowing to half its basal width and digitiform ( Fig. 118View FIGURES 115–118). 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. 46View FIGURES 45, 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. 101View FIGURES 100–102; 107–108; 114; 116; 118). Among them, it shares male cercus and paraproct longer than S10 length with P. calverti  and P. tenuis  ( Figs. 101aView FIGURES 100–102; 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. 22–24View FIGURES 21–24); 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. 5View FIGURES 5–8; 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. 94–96View FIGURES 92–96), which is straight and not projected distally in the other two species ( Figs. 75View FIGURES 71–78; 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. 9–10View FIGURES 9–12) 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. 126View FIGURES 125–126). 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. 94View FIGURES 92–96), 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. 22View FIGURES 21–24). 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. 96View FIGURES 92–96), 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. 24View FIGURES 21–24). Specimens from southern Peru and Bolivia, ‘form b’ of Cowley, share the genital ligula with a relatively longer and narrower strap ( Fig. 95View FIGURES 92–96) 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. 23View FIGURES 21–24).

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. 125–126View FIGURES 125–126).

UMMZ

University of Michigan, Museum of Zoology

USNM

Smithsonian Institution, National Museum of Natural History

FSCA

Florida State Collection of Arthropods, The Museum of Entomology

CSCA

California State Collection of Arthropods

PERU

Universit� di Perugia

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Odonata

Family

Protoneuridae

Genus

Protoneura

Loc

Protoneura woytkowskii Gloyd, 1939

Ellenrieder, Natalia Von & Garrison, Rosser W. 2017

2017
Loc

Protoneura woytkowskii

Pessacq 2015: 65
Pessacq 2012: 70
Garrison 2010: 379
Garrison 2003: 24
Louton 1986: 438Gloyd 1939: 8

1986