Phyllogomphoides duodentatus 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 : 21-23

Phyllogomphoides duodentatus Donnelly, 1979

Phyllogomphoides duodentatus Donnelly, 1979: 259–261 [Odonatologica 8 (4)] (♂ and ♀)

Phyllogomphoides duodentatus . Novelo-Gutiérrez (1993: 18–21) [Odonatologica 22 (1)] (larva ♀, reared)

Type. Holotype 1♂: Mexico: Veracruz, Salto Eyipantla, 8.0 kilometers south of San Andres Tuxtla Mex. Hwy. 180, 15 August 1976, R.W. & J.A. Garrison leg. ( FSCA) ; Allotype 1♀: México, Veracruz, Rio Otapa , 8 km S of La Tinaja, 20 August 1976, R.W. Garrison et al. leg. ( FCSA) ; Paratype 2♂♂, 2♀♀: México, Veracruz, Rio Otapa , 8 km S of La Tinaja, 13 August 1976, R.W. Garrison et al. leg. ( RWG) ; 1♂: México, Veracruz, Rio Coyolucan , 8.6 mi. S La Tinaja, 14 August 1965, D. Paulson leg. ( FSCA) ; 1♀: México: Chiapas, Tuxtla Gutierrez , 12 July 1965, D.R. Paulson leg. ( FSCA) ; 1♀: México: Veracruz, Colonial Apachital , 16 km S, 10 km E of Tierra Blanca, 10 September 1965, T. Donnelly leg. ( FSCA) ; 1♂: Guatemala, Zacapa, nr. Teculutan , 17 August 1969, T. Donnelly leg. ( FSCA) (Material examined) . 1♂: Guatemala, El Progreso, Finca La Cajeta, Estancia de la Virgen , 27 August 1964, T. Donnelly leg. ( FSCA) (Material examined) . 1♂: Honduras, Cortez Prov. river 2.2 mi. N Rio Lindo, 29 August 1964, F. G. Thompson leg. ( FSCA) ( Material examined) .

Type repository. ( FCSA) (Material examined).

Material studied: Total specimens: 72♂♂, 25♀♀, distributed like thus: 24♂♂, 8♀♀ ( FSCA), 13♂♂, 5♀♀ ( IEXA), 4♂♂ ( PSUC), 2♂♂, 2♀♀ ( RWG), 29♂♂, 10♀♀ ( UNAM). MEXICO: Campeche; Mun. Calakmul, Arroyo Escondido San Antonio Soda, Ejido Díaz Ordaz (18º25’15.38’’N, 89º09’20.0’’W), elevation 141m, 22 September 1997, E. Gonzalez Soriano leg., 1♂ . Chiapas; Cascada de Mizolza [Mizol-ha], 17 May 1980, leg:?, 1♂ ; Cascada de Mizol-ha 20 km sur-oeste de Palenque, Km 47 carr Palenque , elevation 260m, 25 September 1983, E. González et al. leg., 2♂♂ ; Mun. Frontera Comalapa, Km 224, Tuxtla Gutiérrez - Cd. Cuauhtémoc (15º50’56.6’’N, 91º57’55.9’’W), elevation 600m, 9 July 1988, A. Cadena et al. leg., 1♂ ; Tuxtla Gutierrez, 12 July 1965, D.R. Paul- son leg., 1♀ (Paratype). Oaxaca; Soyolapan , 16 August 2017, R. Novelo leg., 1♂ . Puebla; Piedras Negras , 23 August 1987, A. Gómez leg., 2♂♂ . San Luis Potosí; 13.2 mi. N. of Tamazunchale , arroyo parallel to H’way 85, 1 September 1958, G. H. Beatty et al. leg., 1♂ ; Agua Buena, Tamasopo , 26 June 1990, E. González leg., 1♂ ; cascadas in park, Tamasopo , 26 June 1990, K.J. Tennessen leg., 3♂♂, 1♀ ; Cascadas Micos , 25 June 1990, G. A. Tennessen leg., 2♂♂ ; Mpio. de Ciudad Valles, Nacimiento del Río Coy , elevation 60m, 24 June 1990, E. González et al. leg., 20♂♂, 12♀♀ ; Rio Micos , aproximadamente a 22 km al N.O. de Ciudad Valles, 25 June 1990, A. Maya et al. leg., 1♂ ; stream at Palitla , 5.5 mi. NW Tamazunchale, 19 June 1966, D. Paulson et al. legs., 1♀ ; Tamain, Las Palmas, Rio Choy , 7 July 2004, R. Novelo leg., 1♀ . Veracruz; Acayucan, 43.7 mi. S. of Acayucan , stream crossing H’way 185, 5 August 1959, G. H. Beatty et al. leg., 2♂♂; (puente) - J. Díaz Covarrubias, Km 212 carr. 180, 8 July 1980, E. González leg., 1♂ ; Colonial Apachital, 16 km S, 10 km E of Tierra Blanca, 10 September 1965, T. Donnelly leg., 1♀ (Paratype); Laguna del Cacahuateno , 5 September 1977, E. Gonzalez leg., 2♂♂ ; Las Choapas, Arroyo Blanco 10 Km N, 5 June 1980, E. Gonzalez leg., 1♂, 1♀ ; Municipio de Tlapacoyan, Ixtacuaco , 28 June 1992, R. Novelo leg., 1♂ ; same locality but: 13 July 2000, 1♂ ; Plan del Río , 11 July 1987, E. Gonzalez et al. leg., 6♂♂, 2♀♀ ; Plan del Rio, 280 m near Rinconada, about 34 Km SE of Xalapa (stream), 11 July 1987, M.J. Westfall leg., 2♂♂ ; same locality but: 26 August 1991, R. Novelo leg., 1♂ ; Puenta Nationale [Puente Nacional], E on Mix-140 Hotel Bal- neario, rocky flowing streamwith pools, 13 July 1992, W.F. Mauffray et al. leg., 10♂♂, 2♀♀ ; Rio Coyolucan, 8.6 mi. S La Tinaja, 14 August 1965, D. Paulson leg., 1♂ (Paratype); Rio Otapa, 8 km S of La Tinaja, 20 August 1976, R.W. Garrison et al. leg., 1♀ (Allotype); Rio Otapa , 8 km S of La Tinaja (18º41’9’’N, 96º26’10’’W), elevation 90m, 13 August 1976, R.W. Garrison et al. leg., 4♂♂, 2♀♀ (Paratypes); San Andres Tuxtla, Arroyo 0.8 mi. S. of H’way 180 at Sihuapan, 13 August 1958, G. H. Beatty et al. leg., 1♂ ; Sayula, carretera nueva Sayula-Almagres, Arroyo bajo el puente, 12 June 2003, R. Novelo leg., 1♂ ; Tebanca , 17 June 1992, E. González et al. leg., 2♂♂, 1♀ ; Salto Eyipantla, 8.0 kilometers south of San Andres Tuxtla ¬ Mex. Hwy. 180, 15 August 1976, R.W. & J.A. Garrison leg. 1♂ (Paratype); Salto Eyipantla , 8.0 kilometers south of San Andres Tuxtla Mex. Hwy. 180, 15 August 1976, R.W. & J.A. Garrison leg. 1♂ ( Holotyp e) .

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

Head: Face mostly pale; labium and submentum pale; labrum mostly brown with two pale spots, anterior and posterior borders brown; mandibles pale basally, tips reddish-brown; anteclypeus pale completely, postclypeus brown with two superior pale spots; lower and middle surface of antefrons brown, upper surface of antefrons with two pale spots, and postfrons pale, a blackish-brown stripe at the union with vertex; antennal scape dark brown with apical rim creamy pale, pedicel dark brown, flagellum brown; vertex dark brown, depressed area between ocelli with a median to large, quadrate pale spot, ocelli yellow; occiput dark brown with a small to median, trapezoid, central pale spot, posterior border fringed with long, stiff, reddish-brown setae ( Fig. 21 ).

Thorax: Pronotum reddish-brown, anterior lobe pale completely, middle and posterior lobes with a pale middorsal twin-spot on posterior margin. Pterothorax reddish-brown, with five pale stripes as shown in Fig. 21 ; middorsal thoracic carina pale; first antehumeral stripe disconnected or connected to pale mesothoracic collar forming an inverted “7” and reaching border of antealar crest superiorly; second antehumeral stripe continuous, ending superiorly in a circular spot close to antealar crest; mesepimeral stripe wider than second antehumeral stripe, covering most of mesepimeron (90–95%), more or less parallel-sided; metepisternal stripe discontinuous, sometimes divided into three fragments: a lower and upper spot, and in between stripe covering 40–50% the length of metepisternum, spots covering 20–30% the length of metepisternum, rarely the lower spot encircling the spiracle; metepimeral stripe slightly forming and inverted “L” reaching subalar carina; metaposternum pale to light brown; pectus brown. Legs: Coxae and trocanters ventrally pale, dorsally light brown with pale streakings extending outward to sides of trocanters; femora mostly light brown, distally dark reddish-brown, anterior femora broadly pale ventrally; tibiae black brown to black, armature black; tarsi and pretarsal claws black. Wings: Hyaline, tinged with yellowish-brown at extreme base, venation light brown, costae with a continuous well-defined yellow line in all the wings; second primary antenodal crossveins the 6th in FW, the 7th in HW; antenodal crossveins: FW 18–23, HW 14–18; postnodal crossveins: FW 11–15, HW 11–17; second serie antenodal crossveins: FW 21–24, HW 16–19; triangles 3-celled, rarely 2-celled; subtriangles 2-celled in both wings, rarely 3-celled; supratriangles 3-celled in both wings, rarely 2 or 4-celled; anal triangle 4-celled; pterostigma dark brown.

Abdomen: Reddish-brown on S1–2, black on S3–7, S8–10 blackish-brown on dorsum and sometimes light brown at sides. Pale coloration creamy yellow as follows: a middorsal stripe and ventral 0.50–0.70 of tergum on S1; a middorsal stripe, auricles, and a posteroventral, narrow, vertical spot on S2; a middorsal stripe on basal 0.80–0.95 tapering posteriorly, and a broad basoventral spot on basal 0.25–0.30 of S3; a middorsal stripe on basal 0.25–0.30 tapering posteriorly, a broad basoventral spot on basal 0.15–0.20 of S4; a middorsal stripe on basal 0.25–0.30 tapering posteriorly, a broad basoventral spot on basal 0.10–0.15 of S5; a middorsal spot on basal 0.10–0.20, a broad basoventral spot on basal 0.10–0.15 of S6; basal half of S7, sometimes 0.70–0.80 length of S7; S8 with lateral spots on basal 0.40–0.50 tapering posteriorly; S9 with lateral spots on basal 0.20–0.30 tapering posteriorly; S10 light brown. Foliation on S8–9 moderately developed and scalloped, foliation on S8 increasing gradually in width caudally, ending in a rounded lobe which surpasses by 0.20–0.30 mm the anterior margin of S9, its edge with a row of small spines on apical 0.30–0.40 of its length, maximum width foliation 0.25–0.45 mm; foliation on S9 0.15–0.30 mm wide, same width along the entire, smooth margin ( Fig. 34 ). Accessory genitalia: Anterior lamina thick, entire, anterior margin almost straight; in ventral view anterior hamuli tumid anteriorly, apically cleft forming two subequal branches, internal branch shorter ending in a short tip dorsally directed, external branch stouter, ending in a wide rounded tip which border is dorsally bent, mesal margin entire, straight, posteroexternal surface with a semicircular, excavated area; posterior hamuli subcylindrical, ending in a short, thick, blunt tip directed mesally, with abundant long and short brown hairs, inner edge slightly convex or almost straight ( Figs. 51–52 ). Vesica spermalis : V1 wide and bifid with a very small tooth on middle, V2–3 of usual type, V4 with 2 relatively short flagella not reaching posterior margin of V1. Caudal appendages: Cerci, with abundant long setae, dorsally pale on apical 0.70–0.80 its length, basal 0.20–0.30 and apices mostly blackish-brown; two dorsal teeth directed mesally, the most basal tooth bluntly pointed, 0.14–0.16 mm length, located at basal 0.70–0.75, the distal tooth roundly pointed, 0.12–0.15 mm length, located at basal 0.85–0.93; in lateral view tip of cercus produced dorsally into a large, stout, wide spine, ventrally rounded; in ventrolateral view with a very tiny to small subbasal spine 0.04–0.09 mm length, sometimes little visible, at basal 0.20–0.25. Epiproct brown with few, short, whitish hairs; in dorsal view widely U-shaped, tips blunt, separate from each other by a distance of 0.65–0.94 mm; in lateral view, branches slightly up-curved, almost straight ( Figs. 81–83 ).

Measurements (average in parenthesis): TL, 55.35–62.76 (55.44); AL, 39–46 (41.92); MWh, 7.4–7.9 (7.7); FwL, 32–38 (35.95); HwL, 31–36 (34.37); FwW, 6.16–9.0 (7.47); HwW, 8.66–10.0 (9.43); HfL, 5.9–6.1 (6.0); cerci length, 2.33–3.06 (2.64).

Female. Similar to male, with the following differences: Head: Postclypeus light brown with two large, superior, pale spots. Thorax: Mesepimeral stripe wider similar that second stripe, more or less parallel sided. Wings: Hyaline, darker in old specimens, tinged with yellowish-brown at extreme base, venation light brown; antenodal crossveins: FW 20–24, HW 15–18; postnodal crossveins: FW 13–17, HW 13–16; anal triangle 4 to 6-celled. Abdomen: Pale as follows: a middorsal stripe and the ventral 0.90–0.95 of tergum on S1; a middorsal stripe, and a basoventral spot on basal 0.90 on S2; basal spot on S4–6 more width aproximatly 10–15% wider than in male; S9–10 without pale spots. Foliation on S8–9 slightly developed and scalloped, that on S8 increasing gradually in width caudally, ending in a tiny rounded lobe which surpasses by 0.15–0.25 mm the anterior margin of S9, its edge with a row of small spines on apical 0.50–0.60 of its length, maximum width of foliation 0.3–0.43 mm; foliation on S9 0.20–0.30 mm wide, the same width along the entire margin, inferior margin with some tiny spine ( Fig. 112 ). Vulvar lamina: Small, 0.16 the length of S9; in ventral view ( Fig. 127 ) widely V-shaped, lobes triangular, reddish-brown, with the most ventral margin reddish-yellow and beset with stiff light brown setae, tips separated each other by a distance shorter than the basal width of each lobe, dorsal margin black, straight, with a mesal, dorsally directed blunt tubercle of 0.08 mm length; in lateroventral view as in Fig. 128 . Caudal appendages: Cerci long, longer than S10, conical, sharply pointed, with abundant short setae, mostly pale, basal 0.05–0.10 dark brown ( Fig. 112 ). Epiproct brown, laminar ending in widely roundedtip, directed ventrally,with few, short, whitish hairs, in dorsal view little visible.

Measurements (average in parenthesis): TL, 52–60.8 (58.27); AL, 41–45 (42.85); MWh, 8.1–8.5 (8.3); FwL, 34–41 (38.25); HwL, 34–38 (36.56); FwW, 7.33–9.0 (8.16); HwW, 9–11 (10.36); HfL, 5.7–6.4 (6.13); VlL, 0.32– 0.54 (0.35); cerci length, 1.76–2.60 (1.84).

Comparative diagnostic notes. This species closely resembles P. bifasciatus (see notes under P. bifasciatus ). Similarily, P. duodentatus can be separated from other species by the external surface of posterior hamulus rounded (external surface of posterior hamulus keeled at tip [ P. pugnifer ], 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 ]. Anterior hamulus with mesal margin entire, with the apex cleft forming two subequal branches with a similar shape (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. Campeche, Chiapas, Oaxaca, Puebla, San Luis Potosí, Veracruz ( Fig. 146 ). In other countries: Belize, El Salvador, Guatemala, Honduras, and Nicaragua ( Paulson & Gozález-Soriano 2018).

Natural history. This species was collected at small streams in relatively open locations rather than in forested regions ( Donnelly 1979). According to the records, this species has an altitudinal distribution in Mexico from 35 to 600masl.

Figures

FIGURES 17–22. Head and thorax, lateral view. 17) P. albrighti, male; 18) P. apiculatus, male; 19) P. bifasciatus, female; 20) P. danieli, male; 21) P. duodentatus, male; 22) P. enriquei, male.

FIGURES 30–36. Abdominal segments S8–10 and caudal appendages, left lateral view. 30) P. albrighti, male; 31) P. apiculatus, male; 32) P. bifasciatus, male; 33) P. danieli, male; 34) P. duodentatus, male; 35) P. enriquei, male; 36) P. indicatrix, male (alcoholic teneral specimen).

FIGURES 43–52. Male accessory genitalia, ventral and ventrolateral view. 43–44) P. albrighti; 45–46) P. apiculatus; 47–48) P. bifasciatus; 49–50) P. danieli; 51–52) P. duodentatus.

FIGURES 69–83. Male caudal appendages, dorsal, lateral and ventral view. 69–71) P. albrighti; 72–74) P. apiculatus; 75–77) P. bifasciatus; 78–80) P. danieli; 81–83) P. duodentatus.

FIGURES 108–113. Female abdominal segments S8–10 and caudal appendages, left lateral view. 108) P. albrighti; 109) P. apiculatus; 110) P. bifasciatus; 111) P. danieli; 112) P. duodentatus; 113) P. luisi.

FIGURES 119–130. Female vulvar lamina, ventral and ventrolateral view. 119–120) P. albrighti; 121–122) P. apiculatus; 123–124) P. bifasciatus; 125–126) P. danieli; 127–128) P. duodentatus; 129–130) P. luisi.

FIGURES 145–146. Maps of geographical distribution. 145) P. duodentatus; 146) P. enriquei.