Paraphlebia quinta Calvert, 1901

Ortega-Salas, Héctor, González-Soriano, Enrique & Jocque, Merlijn, 2022, Untangling the waterfall damsels: a review of the Mesoamerican genus Paraphlebia Selys in Hagen, 1861 (Odonata: Thaumatoneuridae) with descriptions of 11 new species, Zootaxa 5089 (1), pp. 1-66 : 45-49

Paraphlebia quinta Calvert, 1901

Figs. 32 (thorax ♂ Bw), 49–53 (wings ♂♂ Bw ♂♂ Hw ♀), 73–76, 91–92 (appendages ♂ Bw ♂ Hw), 115–116, 124 (posterior lobe of prothorax ♂ Bw ♀), 131 (S8– 10 ♀), 138 (mesostigmal plates ♀), 145–147 (types labels), 148 (map), 162 (habitus ♀) 163–166 (habitus ♂♂ Bw) 167–169 (habitus ♂♂ Hw).

Paraphlebia quinta Calvert 1901: 60 (♂ holotype description, Guatemala); Kirby 1890: 122 (cat.); Beatty & Beatty 1962: 6 ( Mexico: Veracruz, Oaxaca [not published]); Kimmins 1970: 198 (types cat.); Paulson 1982: 251 ( Mexico); González- Soriano & Novelo-Gutiérrez 1996: 162 (Oaxaca, Veracruz); González-Soriano 1997: 333 (natural history and male dimorphism description); González-Soriano & Córdoba-Aguilar 2003: 345 (dimorphism and sexual behaviour description); González-Soriano & Córdoba-Aguilar 2005: 379 (male territoriality); Paulson 2009 (IUCN Red List); Cuevas-Yañez et al. 2015 (conservation status); Garrison & von Ellenrieder 2019: 58 (MCZ paratypes data).

Paraphlebia abrogata Calvert 1907: 355 (♂ holotype description, Guatemala; junior subjective synonym); Calvert 1913: 237 (comparison with Phenacolestes Cockerell ); Flint Jr. 1991: 57 (type cat.); González-Soriano & Córdoba-Aguilar 2003: 251 (holotype examined, P. abrogata synonymized with P. quinta ).

Type series. Holotype of P. quinta . GUATEMALA • 1 ♂; [Alta Verapaz], [Purulhá], Panima [Mojón Panimá]; [15.250°N, 90.083°W]; [360 m]; G. C. Champion leg.; [examined, BMNH]; NHMUK 013324347 .

Paratype. GUATEMALA • 1 ♂; [Alta Verapaz], [Purulhá], Panima [Mojón Panimá]; [15.250°N, 90.083°W]; [360 m]; G. C. Champion leg.; [examined, BMNH] .

Other material examined. GUATEMALA • 1 ♂; (holotype of P. abrogata ); [Alta Verapaz], [Senahú], Cacao [Subestación Cacao, Finca Trece Aguas]; [15.416°N, 89.783°W]; 800 m; 12 Apr 1906; Barber leg.; USNM; SNMNH_391500 • 3 ♂♂; Cahabon, Minas de Oxec , 10 km E of Cahabon; [15.580°N, 89.720°W]; 200 m; 12–14 Jun 1977; T. Donnelly leg.; TWD; IBUNAM:CNIN:OD23138 to 23140 • same data but 1 ♂ 1 ♀; 3 Jun 1978; IBUNAM:CNIN:OD23135 to 23136 • same data but 1 ♂; 5 Jun 1978; IBUNAM:CNIN:OD23137 • same data but 2 ♀♀; San Miguel Tucuru, Los Alpes, nr. Pancajché A.V., Polochic R. Valley ; [15.290°N, 90.110°W]; 600 m; 2 Sep 1964; IBUNAM:CNIN:OD22718 • same data but 1 ♂; Izabal, El Estor, Trib. Río Sauce , 5 km N of Las Dantas; [15.546°N, 89.351°W]; 500 m; 1 Jun 1978; IBUNAM:CNIN:OD22716 • MEXICO • 1 ♂; Chiapas, Ixhuatán, Arroyo en camino a Emiliano Zapata , 1.2 km al NE del camino hacia Chapayal Grande ; 17.305°N, 92.946°W; 599 m; 27 Jul 2016; H. Ortega-Salas, C. Raieli & I. Ángeles-Munguía leg.; IBUNAM:CNIN:OD23053 • same data but 1 ♂; Ixtacomitán, Arroyo cruzando la carretera en el km 95.5 de la carretera Mex-195, 3.6 km al NE de Ixtacomitán; 17.450°N, 93.076°W; 233 m; IBUNAM:CNIN:OD23052 • same data but 5 ♂♂ 2 ♀♀; Halfway between Ixtacomitán & Solosuchiapa, waterfall 20’ high 200’ from highway; [17.442°N, 93.067°W]; 450 m; 12 Aug 1969; G.H. & A.F. Beatty leg.; FEM • same data but 8 ♂♂; Ixtapa, Km 47–48 The 500’ of stream above road descend 80’–90’ (despues de El chorreadero, Río escopetazo y El zapotal); [16.710°N, 92.853°W]; 548 m; 21 Aug 1969 • same data but 1 ♂ 1 ♀; Ocozocoautla de Espinosa, Stream 21.4 mi. N Ocozocoautla; [16.910°N, 93.440°W]; 850 m; 20 Jul 1965; D.R. & M.L. Paulson leg.; FSCA; IBUNAM:CNIN:OD22731 • same data but 2 ♂♂; Stream 24.4 mi. N Ocozocoautla; IBUNAM:CNIN:OD22908 to 22909 • same data but 1 ♂; Pichucalco, Stream 4.4 mi. W Pichucalco; [17.531°N, 93.148°W]; 90 m; 4 Jul 1965; IBUNAM:CNIN:OD22910 • same data but 1 ♂; Salto de Agua, Cascada Misol-Ha; [17.391°N, 91.998°W]; 270 m; 17 May 1981; H. Velasco leg.; IBUNAM; IBUNAM:CNIN:OD22897 • same data but 6 ♂♂; Tapalapa, Río 500 m arriba del puente en el Km 12.5 de la carretera estatal 102 Rayón-Tapalapa ; 17.196°N, 93.080°W; 1680 m; 28 Jul 2016; H. Ortega-Salas, C. Raieli & I. Ángeles-Munguía leg.; IBUNAM:CNIN:OD23054 to 23060 • same data but 23 ♂♂ 4 ♀♀; Tapilula, Several miles (prob 15) past Solosuchiapa on road towards Las Casas; [17.268°N, 93.0258°W]; 609 m; 14 Aug 1969; G.H. & A.F. Beatty leg.; FEM • same data but 1 ♂; Tecpatán, Presa Malpaso, 18 km N of end of road at Malpaso according to Km markers on road. 100’ up the road back towards Malpaso ; [17.293°N, 93.562°W]; 120 m; 9 Aug 1969 • same data but 5 ♂♂ 1♀; Presa Malpaso , 18 km N of end of road at Malpaso according to Km markers on road. 50’ back from road, 5’ waterfall • same data but 3 ♂♂; leg.; IBUNAM; IBUNAM:CNIN:OD22742 • same data but 1 ♂; Oaxaca, San Juan Bautista Valle Nacional, “[Waterfall streams crossing road 6.8 mi. S of Valle Nacional (local station No. 12)]”; [17.723°N, 96.321°W]; 720 m; 15 Aug 1968; G.H. & A.F. Beatty leg.; FEM; PSUC_ FEM _69149 • same data but 3 ♂♂ 1 ♀ • same data but 6 ♂♂ 1 ♀; 2 small streams crossing road 5.8 mi. S. of Valle Nacional (local station No. 9); [17.726°N, 96.326°W]; 626 m; 24 Aug 1959; PSUC_ FEM _83113, 83359, 83386, 83486, 83497, 83533, 83574 same data but 6 ♂♂ 1 ♀; 2 small streams crossing road 5.9 mi. S. of Valle Nacional (local station No. 10); [17.725°N, 96.328°W]; PSUC_ FEM _83352, 83378, 83387, 83391, 83393, 83521 • same data but 9 ♂♂ 1 ♀; 6.1 min S of Valle Nacional. Small stream with slight flow & some pools, above & below road to Oaxaca (local station 11); [17.726°N, 96.327°W]; 582 m; 28 May 1962 • same data but 1 ♂; 6.8 mi S of Valle Nacional. Dried-up bed of small stream near waterfall-stream now practicaly destroyed. Crossing road to Oaxaca (local station 12); [17.723°N, 96.321°W]; 720 m; 27 May 1962 • same data but 3 ♂♂ 1 ♀; 28 May 1962 • same data but 6 ♂♂ 2 ♀♀; Km. 83.5carretera federal 175 Tuxtepec-Oaxaca ; [17.610°N, 96.360°W]; 1700 m; 26 May 1990; A. Cadena & E. Barrera leg.; IBUNAM; IBUNAM:CNIN:OD22770 to 22772, 22774 to 22778 • same data but 1 ♂; Mountain road from Tuxtepec to Cd. Oaxaca. Small stream crossing road, 5.4 mi S. of Valle Nacional ; [17.743°N, 96.327°W]; 360 m; 24 Aug 1959; G.H. & A.F. Beatty leg.; IBUNAM:CNIN: OD22744 • same data but 1 ♂; San Mateo Yetla ; [17.756°N, 96.314°W]; 137 m; 27 May 1990; A. Cadena leg.; IBUNAM:CNIN:OD22748 • same data but 1 ♀; A. Cadena & E. Barrera leg.; IBUNAM:CNIN:OD22747 • same data but 3 ♂♂; Small stream crossing road 4.7 mi. S. of Valle Nacional (local station 7); [17.737°N, 96.327°W]; 457 m; 24 Aug 1959; G.H. & A.F. Beatty leg.; FEM • same data but 1 ♂ 1 ♀; Small stream crossing road 5.4 mi S. of Valle Nacional (local station No. 8); [17.729°N, 96.330°W]; 545 m; PSUC_ FEM _83330 • same data but 7 ♂♂ 2 ♀♀ • same data but 8 ♂♂ 3 ♀♀; 27 Aug 1959; PSUC_ FEM _83001, 83097, 83100, 83343, 83376, 83382, 83384, 83476, 83520, 83534, 83538 • same data but 1 ♂; Valle Nacional ; [17.776°N, 96.300°W]; 85 m; 28 Jun 1966; Alfredo B. Lau leg.; IBUNAM; IBUNAM:CNIN:OD22750 • same data but 1 ♂; 31 Jul 1966; IBUNAM:CNIN: OD22749 • same data but 7 ♂♂ 2 ♀♀; Waterfall stream crossing road 6.1 mi. S. of Valle Nacional (local station No. 11); [17.726°N, 96.327°W]; 582 m; 18 Aug 1959; G.H. & A.F. Beatty leg.; FEM • same data but 2 ♂♂ 1 ♀; 26 Aug 1959; PSUC_ FEM _82911, 83508, 83532 • same data but 4 ♂♂ 2 ♀♀; 27 Aug 1959; PSUC_ FEM _83482, 83490, 83494, 83525, 83529 • same data but 7 ♂♂ 2 ♀♀; Waterfall streams crossing road 6.8 mi. S of Valle Nacional (local station No. 12); [17.723°N, 96.321°W]; 720 m; 12 Aug 1959 • same data but 4 ♂♂; 18 Aug 1959 • same data but 1 ♂ 2 ♀♀; 26 Aug 1959; PSUC_ FEM _83091, 83483 • same data but 5 ♂♂; 27 Aug 1959; PSUC_ FEM _83222, 83288, 83492, 83524, 83539 • same data but 1 ♂; 12–17 Aug 1968; PSUC_ FEM _69085 • same data but 11 ♂♂ 2 ♀♀ • same data but 2 ♂♂ 1 ♀; 24 Jul 1969 • same data but 1 ♂; 26 Jul 1969 • same data but 1 ♂ 1 ♀ • same data but 1 ♂ 1 ♀; 25 Jul 1971; PSUC_ FEM _69260, 69268 • same data but 5 ♂♂ 1 ♀ • same data but 9 ♂♂ 1 ♀; Santiago Comáltepec, Km. 56.9 carretera federal 175 Tuxtepec-Oaxaca ; [17.726°N, 96.326°W]; 626 m; 25 May 1981; C.M. Flint & O.S. Flint Jr. leg.; USNM • same data but 2 ♂♂; H. Velasco leg.; IBUNAM; IBUNAM:CNIN:OD22746, 22821 • same data but 8 ♂♂ 1 ♀; 29 Jul 1990; E. González-Soriano & V. García leg.; IBUNAM:CNIN:OD22779 to 22786, 22789 • same data but 10 ♂♂; 17.726°N, 96.326°W; 10 May 2016; H. Ortega-Salas & C. Raieli leg.; IBUNAM:CNIN:OD22995 to 23004 • same data but 3 ♂♂; Tabasco, Huimanguillo, 5 km al O de Chimalapa 1a sección, camino hacia Francisco J. Mújica ; 17.379°N, 93.628°W; 200 m; 2 Aug 2016; H. Ortega-Salas, C. Raieli & I. Ángeles-Munguía leg.; IBUNAM:CNIN:OD23073 to 23075 • same data but 5 ♂♂ 6 ♀♀; 5.5 km al O de Chimalapa 1a sección, camino hacia Francisco J. Mújica ; 17.380°N, 93.630°W; 375 m; 27 Aug 2015; H. Ortega-Salas, M. Trujano-Ortega & U. García-Vázquez leg.; IBUNAM:CNIN:OD22960 to 22962, 22964, 22966 to 22971 • same data but 5 ♂♂ 2 ♀♀; 4 Sep 2015; IBUNAM:CNIN:OD22972 • same data but 2 ♂♂; Veracruz, Angel R. Cabada, Arroyo Frio near Río Tecolapan, 10.1 mi. N.W. of Santiago Tuxtla ; [18.570°N, 95.329°W]; 243 m; 26 Jul 1959; G.H. & A.F. Beatty leg.; FEM • same data but 1 ♂; Arroyo Frio / Tecolapa (at springs in deep shade); 180 m; 30 Jul 1971 • same data but 8 ♂♂ 1 ♀; Rio Tacolapan [Tecolapan], Route 180, km 551; [18.571°N, 95.331°W]; 25–26 Jul 1966; O.S. Flint Jr. & M.A. Ortiz B. leg.; USNM • same data but 46 ♂♂ 9 ♀♀; Catemaco, Arroyo Huasalapa [Río Cuetzalapan], Catemaco ; [18.374°N, 95.015°W]; 350 m; 3 Aug 1969; G.H. & A.F. Beatty leg.; FEM • same data but 1 ♂; Arroyo Rancho de Moreno, Los Tuxtlas; [18.449°N, 94.976°W]; 110 m; 28 Aug 1982; E. González-Soriano leg.; IBUNAM; IBUNAM:CNIN:OD22862 • same data but 1 ♂ 1 ♀; 3 Jul 1986; R. Mendoza leg.; IBUNAM: CNIN:OD23043 • same data but 1 ♀; Río Cuetzalapan ; [18.374°N, 95.015°W]; 350 m; 2 Aug 1971; G.H. & A.F. Beatty leg.; FEM • same data but 2 ♂♂; Stream 6.7 mi. N Los Mangos; [18.324°N, 95.099°W]; 490 m; 2 Jul 1965; D.R. & M.L. Paulson leg.; FSCA; IBUNAM:CNIN:OD22823 • same data but 2 ♂♂; IBUNAM:CNIN:OD22824 • same data but 1 ♂ 1 ♀; Stream at Coyame ; [18.438°N, 95.022°W]; 380 m; 1 Jul 1965; IBUNAM:CNIN:OD22825 • same data but 1 ♂ ♀; USNM • same data but 2 ♂♂ 1 ♀; Fortín, Barranca de Metlac ; 18.911°N, 97.011°W; 984 m; 26Aug 2015; H. Ortega-Salas, M. Trujano-Ortega & U. García-Vázquez leg.; IBUNAM; IBUNAM:CNIN:OD22955 to 22957 • same data but 1 ♀; San Andrés Tuxtla, Arroyo cerca Laguna Escondida ; [18.588°N, 95.089°W]; 160 m; 19 May 1977; P. Villeda leg.; IBUNAM:CNIN:OD22853 • same data but 2 ♂♂; 21 Apr 1979; E. González-Soriano leg.; IBUNAM:CNIN:OD22849 to 22850 • same data but 1 ♂; 11–15 May 1981; IBUNAM:CNIN:OD22851 • same data but 1 ♂ 1 ♀; 20–24 Jun 1981; IBUNAM:CNIN:OD22852 • same data but 1 ♂; Estación de Biología Tropical Los Tuxtlas ; [18.583°N, 95.067°W]; 120 m; 4 Aug 1971; G.H. & A.F. Beatty leg.; FEM • same data but 1 ♀; 5 Aug 1971 • same data but 1 ♂; 26 May 1978; E. González-Soriano leg.; IBUNAM; IBUNAM:CNIN:OD22861 • same data but 1 ♂; IBUNAM:CNIN:OD22868 • same data but 1 ♀; 2 Jun 1978; IBUNAM:CNIN:OD22860 • same data but 1 ♂; 2 Sep 1978; P. Villeda leg.; IBUNAM:CNIN:OD22859 • same data but 12 ♂♂ 1 ♀; 4–15 May 1981; C.M. Flint & O.S. Flint Jr. leg.; USNM • same data but 1 ♂ 1 ♀; 11–15 May 1981; E. González-Soriano leg.; IBUNAM; IBUNAM:CNIN:OD22900 • same data but 1 ♂; 28 Jul 1982; IBUNAM:CNIN:OD22899 • same data but 4 ♂♂ 1♀; 24–25 May 1984; IBUNAM:CNIN:OD1746 to 1748 • same data but 2 ♂♂; 14–18 Jul 1984; IBUNAM:CNIN: OD1735 • same data but 17 ♂♂ 1 ♀; IBUNAM:CNIN:OD1736 to 1738, 1740 to 1745 • same data but 1 ♂; 23 Apr 1985; A. Ibarra leg.; IBUNAM:CNIN:OD23041 • same data but 1 ♀; 30 Apr 1985; P. Sinaca leg.; IBUNAM:CNIN: OD23037 • same data but 1 ♂; 2 May 1985; E. Ramírez leg.; IBUNAM:CNIN:OD23033 • same data but 1 ♂; P. Sinaca leg.; IBUNAM:CNIN:OD23036 • same data but 1 ♂; 7 May 1985; E. Ramírez leg.; IBUNAM:CNIN: OD23034 • same data but 1 ♂; P. Sinaca leg.; IBUNAM:CNIN:OD23035 • same data but 1 ♂; 2 Jul 1985; A. Ibarra leg.; IBUNAM:CNIN:OD23039 • same data but 1 ♂; E. Ramírez leg.; IBUNAM:CNIN:OD23032 • same data but 1 ♂; 10 Jul 1985; A. Ibarra leg.; IBUNAM:CNIN:OD23038 • same data but 1 ♀; 4 Aug 1985; S. Sinaca leg.; IBUNAM:CNIN:OD23042 • same data but 1 ♀; 28 Aug 1985; R. Mendoza leg.; IBUNAM:CNIN:OD23045 • same data but 17 ♂♂; 21 Sep 1986; E. González-Soriano leg.; IBUNAM:CNIN:OD22842 to 22848 • same data but 3 ♂♂; 20 Sep 1987; IBUNAM:CNIN:OD1734 • same data but 1 ♂; 10 Aug 2003; IBUNAM:CNIN:OD22980 • same data but 1 ♂; 20 Sep 2015; M. Madora-Astudillo leg.; IBUNAM:CNIN:OD22985 • same data but 4 ♂♂; 3 Oct 2015; IBUNAM:CNIN:OD22981 to 22984 • same data but 2 ♂♂; Limite S de la Estación de Biología Tropical Los Tuxtlas, proximo a Lázaro Cárdenas ; [18.576°N, 95.093°W]; 230 m; 7 Jul 1986; R. Mendoza & E. Ramírez leg.; IBUNAM:CNIN:OD23044 • same data but 5 ♂♂ 1 ♀; Los Tuxtlas, Arroyo cerca de Laguna Emilia ; [18.583°N, 95.067°W]; 120 m; 11–15 May 1981; E. González-Soriano leg.; IBUNAM:CNIN:OD22828 to 22832, 22839 • same data but 6 ♂♂ 2 ♀♀; 20–24 Jun 1981; IBUNAM:CNIN:OD22833 • same data but 1 ♂; Los Tuxtlas, Arroyo cerca de Playa Escondida ; [18.618°N, 95.064°W]; 2 m; 13 Sep 1980; IBUNAM:CNIN:OD22841 • same data but 1 ♂ 1 ♀; 11–15 May 1981; IBUNAM:CNIN:OD22840 • same data but 1 ♀; Volcán San Martín , S. slope; [18.530°N, 95.190°W]; 990 m; 20 Jun 1961; J.A. Harshaw leg.; IBUNAM:CNIN:OD22822 • same data but 1 ♂; Santiago Tuxtla, Arroyo Frio, for more than 0.5 mi above confluence with Río Tecolapan , 10.1 mi NW of Santiago Tuxtla; [18.529°N, 95.305°W]; 243 m; 1 Jun 1962; G.H. & A.F. Beatty leg.; FEM • same data but 2 ♂♂ 1 ♀; 2 Jun 1962 • same data but 17 ♂♂ 2♀♀; Arroyo Tatocapan, near highway #180, 1.3 mi. N.W. of Santiago Tuxtla ; [18.482°N, 95.297°W]; 265 m; 20 Aug 1958; PSUC_ FEM _37434, 40263, 40540, 40577, 40589, 40626, 40649, 40788, 84547, 84558, 84559, 84597, 84621, 84716, 98969, 98974, 98984, 99044 • same data but 3 ♂♂; 22 Jul 1959; PSUC_ FEM _ 40283, 40389 • same data but 1 ♂; PSUC_ FEM _40599 • same data but 11 ♂♂ 4♀♀ • same data but 5 ♂♂; 23 Jul 1959; PSUC_ FEM _40280, 40566, 40585, 40646, 40652 • same data but 2 ♂♂ 1 ♀ • same data but 1 ♂; 26 Jul 1959 • same data but 14 ♂♂ 3 ♀♀; 24 Aug 1959 • same data but 11 ♂♂; 2 Jun 1962 • same data but 14 ♂♂ 2 ♀♀; 3 Jun 1962 • same data but 9 ♂♂ 1 ♀; 30 Jul 1969 • same data but 3 ♂♂; 6 Aug 1971 • same data but 1 ♂; Río Frio near Río Tecolapan, 10.1 mi. N.W. of Santiago Tuxtla ; [18.529°N, 95.305°W]; 243 m; 8 Aug 1959; PSUC_ FEM _40517 • same data but 1 ♂; Soteapan, Santa Marta, cañada Bastonal , Estación de Biología Tropical Los Tuxtlas ; [18.332°N, 94.943°W]; 860 m; 2 Jul 1985; A. Ibarra leg.; IBUNAM; IBUNAM:CNIN:OD23040 • same data but 1 ♀; 28 Aug 1988; E. González-Soriano leg.; IBUNAM:CNIN:OD15758 to 15759 • same data but 9 ♂♂ 1 ♀; 18.332°N, 94.943°W; IBUNAM:CNIN:OD22854 to 22858 • same data but 2 ♂♂ 3 ♀♀; Tuxpan, Tuxpan [mislabeled]; 15 m; 27 Jul 1978; J. Pérez Higareda leg.; IBUNAM:CNIN:OD22826, 22863 to 22864 .

Description of male. Small to medium-sized chiefly black individuals. This species has the widest known distribution and the greatest variation in colouration of the genus. Males dimorphic: one morph with hyaline wings, the other with a black tip, the black area covering between 20 and 50% of the wing length. Pale areas on the head, thorax and abdomen highly variable, these areas usually more extensive in hyaline individuals.

Head. Chiefly black, labrum dark glossy blue; anteclypeus black or with diffuse pale markings, postclypeus turquoise to pale blue in hyaline wing males with or without lateral and posterior black spots, pale colouration restricted to anterior border or absent in back-tipped males; frons turquoise or pale blue, pale colouration can be reduced to spots on lateral edges, vertex with two pale spots anterolateral to lateral ocelli, occiput with pale occipital bar complete, reduced, or absent; eyes black in life; antennal scape chiefly black, usually with pale spots or ring on distal margin, pedicel black or brown.

Thorax. Prothorax: Completely black or pale above notopleural suture; see Figs. 115–116 for morphology; propleuron black, sometimes with pale medial spot on ventral margin. Pterothorax: with diffuse green metallic reflections over dark colouration on mesepisternum and mesepimeron, can be completely black or with pale areas as follows: mesepisternum entirely black ( Figs. 164, 166 ) to black with narrow pale line above mesopleural suture, line can be complete in hyaline wing males but more often restricted to spot (as in Fig. 162 ) or discontinuous line on posterior third in both morphs (as in Fig. 163 ), in hyaline wing males or young back-tipped males pale markings can extend forming a continuous line on anterior margin and middorsal carina (as in Fig. 32 ); mesinfraepisternum usually with pale posteroventral spot (as in Fig. 169 ), in darker individuals completely black (as in Figs. 164–166 ); mesepimeron chiefly black, sometimes with narrow pale line below mesopleural suture on posterior third (as in Figs. 164 , 167 ), this line usually reduced to pale spot on mesopleural fossa (as in Figs. 165–166 ); second line highly variable, two-tenths of width of metepisternum, fairly uniform along its length, and not reaching antealar carina (as in Figs. 164–166 ), or in some populations widening up to half width of metepisternum above metathoracic spiracle, then narrowing again posteriorly (as in Figs. 162 , 168 ) in Tapalapa population up to half width of metepisternum at lower end and then widening uniformly to posterior eight-tenths, then narrowing after metapleural fossa (as in Fig. 169 ); metepimeron highly variable, usually with pale line below the metapleural suture two-tenths width of metepimeron, then widening to half width next to antealar carina, thus forming L-shaped area posteriorly (as in Figs. 32 , 164–166 ),in the Tapalapa population covering up to half the width of the metepimeron anteriorly and then widening uniformly to eight-tenths posteriorly, thus forming an elongated triangular pale marking (as in Fig. 169 ) and narrow pale line above metapleural carina, these lines sometimes joining anteriorly by a very narrow line on the anterior margin of the metepimeron (as in Fig. 32 ); metinfraepisternum with or without pale ventral spot; sternum usually with black stripes parallel to metapleural carinae; legs pale but coxa usually with diffuse markings on meso- and metacoxae; femora usually with dim dark markings on extensor surfaces and black rings at distal ends (those marks can be reduced to diffuse spots); tibiae usually with diffuse dark marks along their entire surface; tarsi usually light brown to black; claws reddish.

Wings. Hyaline, with or without apical black tip, when black tip absent a smoky area can be present at tip; if present, black tip covering at least 20% of wing length, thus reaching at least proximal edge of pterostigma; in specimens from Los Tuxtlas black tip can cover up to 60% of wing length; vein that descends from the subnodus, from slightly proximal to slightly distal to first post-quadrangular Vx; CuA in HW usually forked, with at least one well-developed extra sector and one supplementary vein in at least one wing. Px: FW 31–38, HW 27–32.

Abdomen. Chiefly black: S1 usually with pale ventrolateral and anterodorsal spots, the latter sometimes absent; S2 with pale ventrolateral horizontal lines; S3–7 with pale basolateral spots, in some individuals pale colouration extending diffusely on dorsum and seemingly forming pale rings; S9–10 and cerci pruinose on dorsum in mature individuals.

Caudal appendages. Cerci: see Figs. 73–76 for structure. Paraprocts: rudimentary, never with well-defined upper and lower branches; superior lobe rudimentary, only recognisable by a transverse groove; inferior lobe in lateral view with acute projection.

Measurements. Abdomen: 39–43, FW: 29–38, HW: 28–37.

Description of female

Head. As in hyaline wing males but pale colouration on frons less extensive, usually restricted to lateral areas above genae.

Thorax. Prothorax: See Figs. 123–24 for morphology (posterior margin of posterior lobe in some specimens from Comaltepec subquadrate as in Fig. 123 or concave towards lateral edges as in Fig. 124 ). Pterothorax: as in male; depression mesad to mesostigmal lobe in mesostigmal plate rounded.

Wings. Hyaline, with a slightly amber tint; Px: FW 31–38; Px: FW 27–32.

S-1–7 pale colouration as in hyaline male or more extensive. S8 dorsal pale spot present. S9 pale dorsal spot subquadrate, lateral edges parallel. Ovipositor ending beyond tip of cerci.

Measurements. Abdomen: 35–43, FW: 29–38, HW: 28–37.

Diagnosis. This species belongs in the group with little-developed mediodorsal flange of the cerci along with P. flinti , P. hunnal , P. itzamna , and P. kinich ( Figs. 73–80 ). In P. quinta the mesal margin is semi straight, with its greatest width distal to the middle ( Figs. 73–76 ), while in P. itzamna it is convex, with greatest width proximal to the middle ( Fig. 80 ). The posterior lobe of the prothorax in P. quinta is directed posteriorly, and its posterior margin is entire or with a medial notch (as in Figs. 100–102 , 115–116 ), while in P. flinti , P. hunnal and P. kinich it is directed dorsally ( Figs. 98–99, 103 ), and it is either convex towards the outer edges ( Fig. 107 ) or with dorsolateral projections ( Figs. 105–106 ).

Natural history. A dimorphic species, with a black and a hyaline male morph ( González-Soriano 1997; González-Soriano & Córdoba-Aguilar 2003, 2005); the two male morphs differ both in morphology and behaviour. The dark form includes individuals of larger size and a black tip, and this morph exhibits more aggressive behaviour and often displays territoriality. Hyaline males comprise smaller individuals without dark wing tips, and although they can also exhibit territorial behaviour, they do so less frequently ( González-Soriano & Córdoba-Aguilar 2003, 2005). This species has been recorded at streams, rivers and isolated seepages inside tropical rainforests and mountain cloud forests. The larvae are found under stones, or rotten leaves or in seepage vegetation and are frequently seen on vertical walls covered only by a narrow film of water. Under experimental conditions, they are able to survive for several days out of water in moist leaf litter ( González-Soriano 1997). At some locations, especially in cloud forests, this species is found along with P. hyalina ( Fig. 148 ).

Distribution. This is the species with the widest known distribution in the genus, occurring from Barranca de Metlac, Veracruz, Mexico to the Izabal Department in Guatemala. This range includes the eastern part of the Trans- Mexican Volcanic Belt, the Sierra Juárez, the Sierra de los Tuxtlas and the Sierra de las Minas mountain ranges. The species occurs in Veracruz, Oaxaca, Tabasco and Chiapas in Mexico and Alta Verapaz and Izabal in Guatemala. It has been collected between near sea level and 1700 m.a.s.l.

Remarks. Calvert described this species from seven males and one female from Panima, Vera Paz [actually Mojón Panimá, Baja Verapaz]. All males of the type series have black-tipped wings, so when Calvert reviewed hyaline material from Cacao, Alta Verapaz, although he recognized that the cerci of both species were similar: “Caudal appendages as described for P. zoe and as figured for quinta “( Calvert 1907: 355), he designated that material as abrogata , sp. N. Later, González-Soriano and García-Castillo compared a paratype of P. quinta with the holotype of P. abrogata and concluded that those two specimens should be considered conspecific. This information was published as a note at the end of a study on the sexual behaviour of P. quinta (González-Soriano and Córdoba- Aguilar 2003).

In most catalogues of species between 1991 and 2003, P. abrogata is treated as a subjective junior synonym of P. quinta due to a personal communication between E. González-Soriano and R.W. Garrison, information that the latter published in a 1991 manuscript “synonymic list of the new Odonata world” (R.W. Garrison, personal communication, 18 Nov 2016). However, in this manuscript and subsequent editions, that author has clarified that the “synonymic list” is not a publication and therefore the taxonomic changes suggested in it are not valid in accordance with Art. 8.2 of the ICNZ (1999).

Although strong variation in male colouration is present across the geographic range of P. quinta , we did not find enough evidence based on structural characters to consider these more than intraspecific variation.

The specimen of P. quinta labelled Tuxpan, Veracruz, listed in Material Examined was delivered to EGS as part of material occasionally collected by herpetologist J. Pérez Higareda, and since the beginning there has been doubt about its origin. Therefore, until further collections are made in the area and this presence of this species can be confirmed, we regard it as mislabelled and exclude it from the species distribution. Carrillo-Muñoz & García- Miranda (2020) listed one female individual from the Chimalapas region in Oaxaca ( Paraphlebia sp. Mexico • 1 ♀; Oaxaca, Santa María Chimalapas; 17.132° N, 94.459° W, 499 m; 11 Oct 2018; A.I. Carrillo-Muñoz, leg.), but from the photos provided by the authors it is likely that this is P. quinta (A.I. García-Miranda personal communication 10 Jun 2020); however, it is not listed here without closer examination.

Figures

FIGURES 28–34. Paraphlebia spp. colour pattern of thorax and abdomen. Thorax lateral view (28–32); abdomen lateral view (33–34).

FIGURES 105–116. Paraphlebia spp. posterior lobe of prothorax frontal view (105–107); dorsal view (108–116).

FIGURES 164–166. Field photographs. Paraphlebia quinta ♂Bw (164), 5.5 km al O de Chimalapa 1ª sección, camino hacia Francisco J. Mújica, Tabasco, Mexico, 4 Sep 2015; P. quinta ♂Bw (165), Km. 56.9 carretera federal 175 Tuxtepec-Oaxaca, Oaxaca, Mexico, 10 May 2016; P. quinta ♂Bw (166), Reserva Ecológica La Otra Opción, Veracruz, Mexico, 1 Aug 2019.

FIGURES 161–163. Field photographs. Paraphlebia kinich ♂ (161), Cascada Misol-Ha, Chiapas, Mexico, 30 Aug 2015; P. quinta ♀ (162), Km. 56.9 carretera federal 175 Tuxtepec-Oaxaca, Oaxaca, Mexico, 10 May 2016; P. quinta ♂Bw, young individual (163), 5.5 km al O de Chimalapa 1ª sección, camino hacia Francisco J. Mújica, Tabasco, Mexico, 4 Sep 2015.

FIGURES 167–169. Field photographs. Paraphlebia quinta ♂Bw, young individual (167), Km. 56.9 carretera federal 175 Tuxtepec-Oaxaca, Oaxaca, Mexico, 10 May 2016; P. quinta ♂ Hw (168), 5.5 km al O de Chimalapa 1a sección, camino hacia Francisco J. Mújica, Tabasco, Mexico, 4 Sep 2015; P. quinta ♂ Hw (169), Río 500 m arriba del puente en el Km 12.5 de la carretera estatal 102 Rayón-Tapalapa, Chiapas, Mexico, 28 Jul 2016.

FIGURES 73–78. Paraphlebia spp. caudal appendages. (a) dorsal view; (b) mediodorsal view. W – maximum width of mediodorsal flange, w – maximum width of the distal lobe.

FIGURES 117–128. Paraphlebia spp. posterior lobe of prothorax dorsal view.

FIGURES 79–84. Paraphlebia spp. caudal appendages (a) dorsal view; (b) mediodorsal view (79–80) W – maximum width of mediodorsal flange, w – maximum width of the distal lobe; dorsal view (81); lateral view (82, 84); posterior view (83).

FIGURES 95–104. Paraphlebia spp. caudal appendages and posterior lobe of prothorax. Caudal appendages lateral view (95–97); posterior lobe of prothorax dorsolateral view (98–100); lateral view (101–104).

FIGURES 148–149. Paraphlebia spp. collection sites.