Phyllogomphoides pugnifer Donnelly, 1979

Torres-Pachón, Mónica, Novelo-Gutiérrez, Rodolfo & Ruiz-Sanchez, Eduardo, 2019, A synopsis of Phyllogomphoides Belle, 1970 (Odonata: Gomphidae) of Mexico: taxonomy and distribution, Zootaxa 4634 (1), pp. 1-67 : 35-37

Phyllogomphoides pugnifer Donnelly, 1979

Phyllogomphoides pugnifer Donnelly, 1979: 261–262 [Odonatologica 8 (4)] (♂)

Phyllogomphoides pugnifer . González-Soriano & Novelo-Gutiérrez (1985: 147–150) [Odonatologica 14 (2)] (♀)

Type. Holotype 1♂: México, Veracruz, 3 km. N. of Santiago Tuxtla , 13–14 September 1965, T. Donnelly leg. ( FSCA) ; Paratype 6♂♂: México, Veracruz, 3 km. N. of Santiago Tuxtla , 13–14 September 1965, T. Donnelly leg. ( FSCA) ; 1♂: Mexico, Chiapas, stream 21.4 mi. N Ocozocoautla , 21 July 1965, D. Paulson leg. ( FSCA) ; 2♂♂: same locality but, 25 August 1965, D. Paulson leg. ( FSCA) .

Type repository. FSCA (Material examined).

Material studied: Total specimens: 73♂♂, 18♀♀, distributed like thus: 31♂♂, 7♀♀ ( FSCA), 1♂ ( IEXA), 9♂♂, 6♀♀ ( PSUC), 32♂♂, 5♀♀ ( UNAM). MEXICO: Chiapas ; stream 21.4 mi. N Ocozocoautla , 21 July 1965, D. Paulson leg., 1♂ (Paratype) ; stream 20.1 mi. N Ocozocoautla , 25 August 1967, D. Paulson et al. leg., 2♂♂ (Paratype) ; El Zapotale 2 mi S of Tuxtla Gutiérrez, 12 July 1957 Chemsak leg., 1♂ . San Luis Potosí; 4.4 mi. N. of Tamazunchale , small arroyo in wooded ravine, crossing H’way #85, 30 July1958, leg?., 1♀ ; Arroyo de los Pa- tos, 7.9 mi. (E.) of Tamazunchale, on road to San Felipe Orizatlan , 8 June 1962, leg?., 1♀ . Veracruz; 3 km. N. of Santiago Tuxtla , 13–14 September 1965, T. Donnelly leg., 7♂♂ (Holotype and Paratypes) ; Arroyo aquas [Agua] Caliente, Sontecomapan , “Los Tuxtlas”, 30 August 1988, J. Daigle et al. leg., 5♂♂, 1♀ ; Agua Caliente , 1 km E of Sontecomapan, 18 July 1992, K.J. Tennessen leg., 2♂♂ ; same locality but: 19 July 1992, S.W. Dunkle leg., 2♂♂ ; Arroyo Playa Escondida , 11 June 1979, G. Jiménez leg., 1♀ ; same locality but: 13 July 1979, E. Gonzalez leg., 1♂ ; Arroyo Laguna Escondida , 23 May 1980, R. Novelo leg., 1♂ ; same locality but: 24 May 1980, 1♀ ; same locality but: 25 May 1980, 1♀ ; same locality but: 7 June 1980, 3♂♂ ; same locality but: 9 June 1980, 2♂♂ ; same locality but: 10 June 1980, 4♂♂ ; same locality but: 11 June 1980, 1♂ ; same locality but: 9 July 1980, 4♂♂ ; same locality but: 18–24 June 1981, 1 ♂ ; same locality but: 20–24 June 1981, 7 ♂♂ ; same locality but: 17–24 July 1981, 3 ♂♂ ; same locality but: 16–30 July 1981, 1 ♂ ; same locality but: 7 August 1982, 3♂♂, 1♀ ; same locality but: 14 July 1992, K.J. Tennessen, 1♂, 1♀ ; Arroyo on road to Jicacal Beach, and pond, near Rancho La Esperanza , 28 August 1988, M.J. Westfall et al. leg., 3♂♂, 2♀♀ ; Arroyo Tatocapan, parallel to H’way #180, 1.3 mi. N.W. of Santiago Tuxtla , 2 June1962, Beatty et al. leg., 7♂♂, 3♀♀ ; Fortin, arroyo below crossing of Mexico-Veracruz H’way at km. 333, 30 July 1958, Beatty et al. leg., 1♂, 1♀ ; La Palma , 4 August 1982, E. Gonzalez leg., 1♂ ; Laguna Azul, Laguna Emilia, and Arroyo de la Laguna Escondida, near Los Tuxtlas Biol. Sta , 27 August 1988, S.W. Dunkle leg., 1♂, 1♀ ; Lake Catemaco, Hotel Playa Azul , 17 August 1963, H.V. Weems leg., 1♀ ; Los Tuxtlas Biological Station nr Catemaco , 14 July 1992, S.W. Dunkle leg., 1♀ ; San Andres Tuxtla, Balzapote, Río Máquinas ( Villa Cariño ), 14 June 2009, R. Novelo leg., 1♂ ; Sontecomapan , 7 July 1979, G. Jiménez leg., 1♀ ; same locality but: 10 July 1979, E. Gonzalez leg., 1♂ ; same locality but: near Los Tuxlas Biological Research stat, arroyo de laguna escondida, 15–16 July 1992, W.F. Mauffray leg., 3♂♂ ; same locality but: tributary of Rio Coscoapan , 6 km E of Sontecomapan, 16 July 1992, K.J. Tennessen leg., 4♂♂ .

Description of male. Body dark brown, with five pale stripes on pterothorax.

Head: Face mostly pale; labium pale, submentum pale to light brown; labrum brown sometimes with two pale spots laterally; mandibles pale basally, tips reddish-brown; anteclypeus mostly pale with a brown band posteriorly; postclypeus brown with pale spots laterally; lower surface of antefrons brown, upper of antefrons and postfrons mostly pale, a blackish-brown stripe at the union with vertex, postfrons with a middle depression giving the appearance of two mounds; antennal scape dark brown with apical rim creamy pale, pedicel dark brown, flagellum brown; vertex dark brown, depressed area between ocelli with a median, semicircular pale spot; occiput dark brown with a median, trapezoid, central pale spot, posterior border fringed with long, stiff, reddish-brown setae ( Fig. 27 ).

Thorax: Pronotum reddish-brown, anterior, middle and posterior lobes with a pale middorsal, and sometimes middle and posterior lobes with a pale spots laterally. Pterothorax reddish-brown, with five pale stripes as show in Fig. 27 ; middorsal thoracic carina brown and sometimes with a tiny pale spot; first antehumeral stripe connected to pale mesothoracic collar forming an inverted “7” reaching border of antealar crest superiorly; second antehumeral stripe continuous, covering 0.90–0.95 of mesepisternum’s length, ending at upper end in a rounded tip close to antealar crest; mesepimeral stripe similar in width than second antehumeral stripe, covering most of mesepimeron (85–90%), more or less parallel sided and gradually widening at upper end; metepisternal stripe continuous covering 90–95% of metepisternum, this stripe encircling spiracle inferiorly, and forming a circular spot superiorly; rarely this stripe is discontinuous; metepimeral stripe forming and inverted “L” reaching subalar carina, widening from the middle to posterior end; metaposternum brown with a small pale spot on middle; pectus brown. Legs: Femora mostly brown, distally dark reddish-brown, anterior femora broadly pale ventrally; tibiae, tarsi, and pretarsal claws reddish-black, armature black. Wings: Hyaline, tinged with brown at extreme base, venation blackish-brown, anterior margin of costae brown and sometimes on anterior margin of costae of HW with a yellow line from base to nodus; second primary antenodal crossveins the 7th in FW, the 8th in HW; antenodal crossveins: FW 21–24, HW 13–19; postnodal crossveins: FW 13–17, HW 13–18; second series of antenodal crossvein in FW 24–26, HW 18–21; triangles 3-celled and sometimes 2-celled; subtriangles 2-celled in both wings; supratriangles 3-celled and rarely 4-celled; anal triangle 4-celled; pterostigma dark brown.

Abdomen: Reddish-brown on S1–2, black on S3–7, S8–9 black on dorsum, light brown at sides, S10 dark brown, with small windings on posterior border. Pale coloration creamy yellow as follows: a middorsal stripe and the ventral 0.5–0.70 of tergum on S1; a middorsal stripe, auricles, and a posteroventral, narrow, vertical spot on S2; a middorsal stripe on basal 0.60–0.80 constricted at basal 0.85 of its length, and a broad basoventral spot on basal 0.15–0.20 of S3; a middorsal spot on basal 0.25–0.35 tapering posteroirly, a broad basoventral spot on basal 0.15–0.20 of S4; a middorsal stripe on basal 0.10–0.15 tapering posteriorly, a broad basoventral spot on basal 0.10–0.15 of S5; a middorsal spot on basal 0.10–0.15, a broad basoventral spot on basal 0.10–0.15 of S6; basal half of S7 and sometimes 0.90 its length laterally; S8 with pale strikes on basoventral 0.50–0.70; S9–10 without pale spots. Foliation on S8–9 moderately to poorly developed and scalloped, foliation S8 increasing gradually in width caudally, ending in a small rounded lobe which surpasses by 0.20 mm the anterior margin of S9, its edge with a row of small spines on apical 0.40–0.50 its length, maximum width of foliation 0.40–0.50 mm; foliation on S9 poorly developed, the same width along the entire and smooth margin, 0.25–0.30 mm wide ( Fig. 40 ). Accessory genitalia: Anterior lamina thick, entire, anterior margin concave and posterior margin straight; in ventral view anterior hamuli tumid anteriorly, cleaft apically forming two subequal branches, internal branch widely rounded at tip, external branch ending into a short incurved hook, the tip directed mesally; mesal margin entire, convex. Posterior hamuli blackish-brown, subcylindrical, slightly widenedon distal half, with a tiny, bluntly tip directed mesally, with abundant long and short, brown, bristle-like setae ( Figs. 63–64 ). Vesica spermalis : V1 bifid with a big tooth on middle, V2–3 of usual type, V4 with 2 relatively short flagella that light reaching posterior margin of V1. Caudal appendages: Cerci with moderately long setae, basal 0.30–0.40 blackish-brown, pale on apical 0.60–0.70 its length, apices mostly black brown; a dorsomesal bluntly tipped tooth at basal 0.64–0.70, 0.20–0.30 mm length, directed medially, followed by a small, subapical, shelf-like carina; tips of cerci slightly flattened laterally and produced dorsally into a spine; in lateral view tip of cercus produced dorsally into a large, wide spine; ventral subbasal spine absent. Epiproct black brown with a large pale spot on each branch; in dorsal view widely U-shaped, tips blunt, separated from each other by a distance of 0.70–0.90 mm,; in lateral view, branches straight, in dorsal view ( Figs. 99–101 ).

Measurements (measures are an average): TL, 61.1–64.7 (62.76); AL, 44–48 (45.66); MWh, 8.3–9.1; FwL, 38–41 (39); HwL, 36.5–40 (37.5); FwW, 8–9 (8.41); HwW, 10–11 (10.33); HfL, 6.7–7.6; cerci length, 2.5–3.2 (3).

Female. Similar to male, with the following differences: Head: Occiput dark brown with a small, central pale to light brown spot (sometimes, posterior border fringed with long, stiff, reddish-brown setae). Thorax: First antehumeral stripe not connected to pale mesothoracic collar. Wings: antenodal crossveins: FW 21–25, HW 17–19; postnodal crossveins: FW 14–17, HW 16–17; triangles 4-celled and sometimes 3-celled; subtriangles 2 or 3-celled in both wings; anal triangle 5 to 8-celled. Abdomen: A middorsal stripe on basal 0.60–0.70 constricted at basal 0.85 of its length, and a broad basoventral spot on basal 0.10–0.15 of S3; S8–10 sometimes with a lateral pale spots anteriorly. Foliation on S8–9 moderately to poorly developed and scalloped, foliation S8 increasing gradually in width caudally, not surpassing the anterior margin of S9, its edge with a row of small spines on apical 0.30–0.40 its length, maximum width of foliation 0.25–0.32 mm; foliation on S9 poorly developed, the same width along the entire, smooth margin, 0.12–0.15 mm wide ( Fig. 116 ). Vulvar lamina: Short occupying 0.18 the length of S9; in ventral view ( Fig. 135 ) widely V-shaped, lobes triangular, reddish-black, with the most ventral margin black and beset with stiff yellow setae, tips separated each other by a distance greater than the basal width of each lobe, dorsal margin without a posterodorsal tubercle; in lateroventral view as in Fig. 136 . Caudal appendages: Cerci long, longer than S10, conical, sharply pointed, with abundant short setae, basal 0.20–0.30 blackish-brown, pale on apical 0.70–0.80 its length ( Fig. 116 ). Epiproct black, laminar-shaped, widely rounded at tip, slightly directed ventrally; in ventral and dorsal views with many, short, whitish hairs; epiproct slightly visible dorsally.

Measurements (average in parenthesis): TL, 60–65.33 (62.91); AL, 45–46 (45.5); FwL, 39–42 (41.16); HwL, 38–40 (39.33); FwW, 8.66–9.50 (9.05); HwW, 10.12–12 (11.04); VlL, 0.36–0.48 (0.42); cerci length, 2.0–2.4 (2.24).

Comparative diagnostic notes. This species closely resembles P. bifasciatus . Phyllogomphoides pugnifer is of medium size; while P. bifasciatus are from small to medium size. Males can be differentiated from P. bifasciatus by the (features of the later in parentheses): the external surface of posterior hamulus keeled at tip and the epiproct in lateral view straight (the external surface of posterior hamulus rounded and the epiproct in lateral view strongly up-curved). On the other hand, P. pugnifer and P. nayaritensis females are most similar. Females can be differentiated from P. nayaritensis by the following features (in parentheses): the lobes of vulvar lamina widely V-shaped, triangular, reddish-black, tips separated each other by a distance greater than basal width of each lobe (the lobes of vulvar lamina widely U-shaped, lobes black, trapezoid, tips truncate, separated each other by a distance shorter than basal width of each lobe). On the other hand, the geographic distribution pattern of P. pugnifer and P. bifasciatus it is more towards the south of the Gulf of Mexico; while in P. bifasciatus it is towards the south of the Gulf of Mexico and P. duodentatus it is more towards to center and south of the Gulf of Mexico. Similarily, P. pugnifer can be separated from other species by the external surface of posterior hamulus keeled at tip (the external surface of posterior hamulus rounded [ P. duodentatus ], posterior hamulus strongly expanded at middle, tapering posteriorly ending in a stout, sharply pointed hook [ P. nayaritensis ]. Anterior hamulus with mesal margin entire, with the apex cleft forming two subequal branches with a similar shape (anterior hamulus with mesal margin notched or broadly emarginate, the tip widely rounded or narrowly hooked [ P. apiculatus , P. danieli , P. enriquei , P. indicatrix , P. luisi , and P. pacificus ]. The mesal margin entire but apex cleft forming two branches (anterior hamulus with mesal margin and apex entire [ P. suasus ]. And from the remaining species by abdominal segments S8–10 mainly black (abdominal segments 8–10 brown with extensive yellow or orange [ P. albrighti and P. stigmatus ].

Flight season. May, June, July, August, and September.

Distribution. Chiapas, San Luis Potosí, Veracruz ( Fig. 151 ). In other countries: Belize, Costa Rica, and Guatemala ( Paulson & Gozález-Soriano 2018).

Natural history. This species lives in small streams and little is known on its habits. Some specimens have been collected at a shaded forest stream while others in hilly places. Likewise, males have been taken foraging at dusk in a clearing between forest and river. Other specimens were found in very small shaded forest streams in hilly country ( Donnelly 1979). Males follow the course of the streams hovering frequently in backwaters, where they remain for brief periods. Males perch on twigs or grasses at the stream edge and make pursuit flights when conspecifics pass near their perches ( González-Soriano & Novelo-Gutiérrez 1985). According to the records, this species has an altitudinal distribution in Mexico from 18 to 975masl.

Figures

FIGURES 23–29. Head and thorax, lateral view. 23) P. indicatrix, male (alcoholic teneral specimen); 24) P. luisi, male; 25) P. nayaritensis, male; 26) P. pacificus, male; 27) P. pugnifer, male; 28) P. stigmatus, male; 29) P. suasus, male.

FIGURES 37–42. Abdominal segments S8–10 and caudal appendages, left lateral view. 37) P. luisi, male; 38) P. nayaritensis, male; 39) P. pacificus, male; 40) P. pugnifer, male; 41) P. stigmatus, male; 42) P. suasus, male.

FIGURES 63–68. Male accessory genitalia, ventral and ventrolateral view. 63–64) P. pugnifer; 65–66) P. stigmatus; 67–68) P. suasus.

FIGURES 96–107. Male caudal appendages, dorsal, lateral and ventral view. 96–98) P. pacificus; 99–101) P. pugnifer; 102– 104) P. stigmatus; 105–107) P. suasus.

FIGURES 114–118. Female abdominal segments S8–10 and caudal appendages, left lateral view. 114) P. nayaritensis; 115) P. pacificus; 116) P. pugnifer; 117) P. stigmatus; 118) P. suasus.

FIGURES 131–140. Female vulvar lamina, ventral and ventrolateral view. 131–132) P. nayaritensis; 133–134) P. pacificus; 135–136) P. pugnifer; 137–138) P. stigmatus; 139–140) P. suasus.

FIGURES 151–152. Maps of geographical distribution. 151) P. pugnifer; 152) P. stigmatus.