Untangling the waterfall damsels: a review of the Mesoamerican genus Paraphlebia Selys in Hagen, 1861 (Odonata: Thaumatoneuridae) with descriptions of 11 new species Author Ortega-Salas, Héctor 0000-0002-5373-4839 hector.ortegasalas@naturalis.nl Author González-Soriano, Enrique 0000-0002-4798-7274 esoriano@ib.unam.mx Author Jocque, Merlijn 0000-0002-7196-7476 merlijn.jocque@binco.eu text Zootaxa 2022 2022-01-11 5089 1 1 66 journal article 53857 10.11646/zootaxa.5089.1.1 67fbf2b0-b944-485d-90b5-976914d338ad 1175-5326 5836060 E12F2B20-A84A-48E2-9C77-B281F1BFC62E Paraphlebia zoe Selys in Hagen, 1861 Figs. 54–55, 58 (wings ♂♂ Bw ♂♂ Hw), 67–68, 88 (appendages ♂ Bw ♂ Hw), 100, 104, 110–111 (posterior lobe of prothorax ♀ ♂♂ Bw), 134 (S8– 10 ♀ ), 139 (mesostigmal plates ♀ ), 140 ( holotype labels), 141 ( holotype ), 148 (map), 170–171 (habitus ♂ Bw ♀ ♂ Hw). Paraphlebia zoe Selys [Selys in Hagen] 1861: 72 ( holotype diagnosis and data); Selys 1860: 435 ( nomen nudum ; comparison with Palaemnema paulina ); Selys 1862: 9–10 ( ♂ holotype description, Veracruz ); Selys 1886: 33 (key); Kirby 1890: 122 (cat.); Calvert 1901: 60 (additional mat; ♀ desc.); Calvert 1913: 237–238 (comparison with Phenacolestes Cockerell ); Beatty & Beatty 1962: 5 ( San Luis Potosí [unpublished]); Bridges 1994 : VIII.54; (cat.); González-Soriano & Novelo- Gutiérrez 1996: 162 ( Hidalgo , Puebla , San Luis Potosí ); Paulson & von Ellenrieder 2006 (IUCN Red List); Novelo- Gutiérrez 2008: 29 (larva); Romo-Beltrán et al. 2009: 699 (dimorphism); Munguía-Steyer et al. 2010: 175 (effect of male morph condition on survival); Escoto-Moreno & Márquez 2011: 416 ( Queretaro ; new records and distribution map); Anderson et al. 2012 (microsatellites characterization); Ruiz-Guzmán et al. 2013: 620 (male dimorphism effects on immune response); Wong-Muñoz et al. 2013: 407 (effects of body size, male morph, morph frequency, and alternative reproductive tactics on copulation duration); Escoto-Moreno et al. 2014: 1048 (additional records); Bello-Bedoy et al. 2015: 327 (sexual traits pattern effects on mating success and survival); Cuevas-Yañez et al. 2015: 517 (conservation status). Type series. Holotype MEXICO • 1 ♀ ; [ Veracruz ] ; A. Sallè leg.; [not examined, RBINS ]. Other material examined. MEXICO • 1 ♂ ; Hidalgo , Chapulhuacán , Waterfall , 1.1 mi. NE Chapulhuacan ; [ 21.175°N , 98.901°W ]; 120 m ; 27 Jun 1965 ; D.R. & M.L. Paulson leg.; USNM ; SNMNH_383658 • same data but 1 ♂ ; Lolotla , Ixtlahuaco ; [ 20.884°N , 98.702°W ]; 1441 m ; 24 Jul 1994 ; Morris Hopkins leg.; INECOL • same data but 1 ♂ ; Molango , Arroyo de la laguna Azteca [Laguna de Atezca]; [ 20.805°N , 98.745°W ]; 1890 m ; 26 Jun 1993 ; R. Novelo-Gutiérrez leg. • same data but 4 ♂♂ ; 24 Jul 1994 • same data but 1 ♂ ; Molango de Escamilla , 5 km . Al NO de Pemuxtitla , al Wde Tetzintla (mina); [ 20.882°N , 98.782°W ]; 1000 m ; 21 May 1984 ; J. Peña leg.; IBUNAM ; IBUNAM :CNIN:OD22895 to 22896 • same data but 8 ♂♂ ; Tepehuacán de Guerrero , Otongo ( Colonia de la Minera Autlán ). Escurridero ; [ 20.960°N , 98.755°W ]; 1090 m ; 26 Jun 1993 ; R. Novelo-Gutiérrez leg.; INECOL • same data but 1 ♂ ; Tlanchinol , 1 km al N de Chapulhuacán ; [ 21.1670°N , 98.899°W ]; 920 m ; 11 May 1989 ; E. González- Soriano leg.; IBUNAM :CNIN:OD1695 • same data but 6 ♂♂ 1 ♀ ; 16 May 1989 ; IBUNAM :CNIN:OD1658, 1691 to 1694 • same data but 1 ♂ ; 2.2 km al S del Centro de Tlanchinol , Km 161 carretera federal Mex 105; 20.979°N , 98.671°W ; 1563 m ; 16 Jun 2016 ; U.O. García-Vázquez leg.; IBUNAM :CNIN:OD23090 • same data but 1 ♂ ; 2.5 km NE de Chapulhuacán , carr. 85, escurridero.; [ 21.175°N , 98.901°W ]; 930 m ; 26 Jun 1993 ; R. Novelo-Gutiérrez leg.; INECOL • same data but 1 ♂ ; 7 Km al NE de Tlanchinol , Km. 170 carretera federal Mex 105 Tlanchinol- Huejutla ; [ 21.023°N , 98.637°W ]; 1450 m ; 20 Jun 1981 ; Oscar Canu leg.; IBUNAM ; IBUNAM :CNIN:OD22773 • same data but 1 ♂ ; 1490 m ; 28 May 1992 ; E. González-Soriano leg.; IBUNAM :CNIN:OD22893 • same data but 7 ♂♂ 2 ♀♀ ; 1509 m ; 26 Jun 1993 ; R. Novelo-Gutiérrez leg.; INECOL • same data but 1 ♂ ; 2 Jul 1994 • same data but 3 ♂♂ ; 22 Jul 1994 • same data but 1 ♂ ; 29 Aug 2007 • same data but 2 ♂♂ 1 ♀ ; 1450 m • same data but 1 ♂ ; Tlalchinol [Tlanchinol]; [ 20.990°N , 98.660°W ]; 1500 m ; 6 Jun 1981 ; G. Cartas leg.; IBUNAM ; IBUNAM :CNIN: OD22894 • same data but 1 ♂ ; Puebla , Cuetzalan , El Cuichat (arroyo); 20.000°N , 97.511°W ; 870 m ; 2 Jul 1994 ; R. Novelo-Gutiérrez leg.; INECOL • same data but 1 ♀ ; 7 Jul 2000 • same data but 1 ♂ ; 29 Aug 2007 • same data but 1 ♂ ; 30 Jul 2014 • same data but 2 ♂♂ 1 ♀ ; Cuetzalan del Progreso , 1 km al SE de Atepatahua ; 20.013°N , 97.484°W ; 669 m ; 13 Jul 2007 • same data but 5 ♂♂ ; 650 m ; IBUNAM ; IBUNAM :CNIN:OD22944 to 22946 • same data but 1 ♂ ; Km 11 despues de Villa Avila Camacho , carretera Puebla 104 Tlaxcaltongo La Ceiba ; [ 20.317°N , 97.864°W ]; 500 m ; 24 Jul 1983 ; G. Ortega , E. Barrera leg.; IBUNAM :CNIN:OD22940 • same data but 4 ♂♂ ; 2 Jul 1994 ; IBUNAM :CNIN:OD22936 to 22939 • same data but 2 ♂♂ ; 7 Jul 2000 ; IBUNAM :CNIN:OD22941, 22943 • same data but 1 ♂ ; 9 Jul 2007 ; IBUNAM :CNIN:OD22942 • same data but 1 ♂ 1 ♀ ; Jopala , Barranca de Patla ; [ 20.242°N , 97.855°W ]; 375 m ; 27 Jul 1981 ; A. Ibarra leg.; IBUNAM :CNIN:OD1690, 22892 • same data but 1 ♂ ; H. Patla , El Pozo ; [20.249°N, 9785°W]; 440 m ; 8 Sep 2002 ; K. Cuevas-Yañez leg.; IBUNAM :CNIN:OD23076 • same data but 1 ♀ ; Xicotepec , Km. 11 despues de Villa Avila Camacho , carretera Puebla 104 Tlaxcaltongo La Ceiba ; [ 20.317°N , 97.864°W ]; 500 m ; 2 Jul 1994 ; G. Ortega , E. Barrera leg.; IBUNAM :CNIN:OD23091 • same data but 3 ♂♂ 1 ♀ ; Zihuateutla , Km. 3 de la carretera Los Caminos-La Unión (escurridero); [ 20.260°N , 97.870°W ]; 650 m ; 21 May 1987 ; R. Novelo-Gutiérrez & A. Gómez-Anaya leg.; IBUNAM :CNIN:OD22891 • same data but 9 ♂♂ ; La Unión (escurridero); IBUNAM :CNIN:OD22888 to 22890 • same data but 13 ♂♂ 1 ♀ ; 23 Jul 1987 ; R. Novelo-Gutiérrez leg.; IBUNAM :CNIN:OD22881 to 22887• same data but 1 ♂ 2 ♀♀ ; San Luis Potosí , Tamazunchale , Rancho Quemado , Route 85, km. 353, near Tamazunchale ; [ 21.256°N , 98.836°W ]; 160 m ; 5 Aug 1966 ; O.S. Flint Jr. leg.; USNM ; SNMNH_383659 to 383660, 383669 • same data but 2 ♂♂ ; Waterfall near (WSW) Tamazunchale ; 880 m ; 27 Jun 1965 ; D.R. & M.L. Paulson leg.; IBUNAM ; IBUNAM :CNIN:OD22866 to 22867 • same data but 2 ♂♂ ; Xilitla , Finca Santa Mónica ; 21.393°N , 98.998°W ; 652 m ; 13 Jun 2016 ; U.O. García-Vázquez leg.; IBUNAM : CNIN:OD23081 to 23082 • same data but 1 ♂ ; La Conchita , 3 km al NE de Xilitla ; [ 21.394°N , 98.990°W ]; 570 m ; 8 May 1950 ; IBUNAM :CNIN:OD1732 • same data but 1 ♂ ; Las Pozas de Xilitla ; [ 21.390°N , 98.995°W ]; 580 m ; 15 Jun 2001 ; E. González-Soriano leg.; IBUNAM :CNIN:OD14618 • same data but 3 ♂♂ ; Las Pozas , Xilitla ; [ 21.395°N , 98.985°W ]; 570 m ; 28 Jun 1990 ; E. González-Soriano & A. Maya leg.; IBUNAM :CNIN:OD1656, 1730 to 1731 • same data but 7 ♂♂ 3 ♀♀ ; 600 m ; 23 Aug 1999 ; IBUNAM :CNIN:OD15760 to 15765, 22902 • same data but 4 ♂♂ ; Veracruz , Atotonilco , Río Jamapa ; [ 19.148°N , 97.188°E ]; 2100 m ; 7 Jul 2000 ; J.A. Gómez-Anaya leg.; INECOL • same data but 3 ♂♂ ; 30 Jul 2014 • same data but 3 ♂♂ ; Atzalan , El Ahuacate ( Ixtatahuia ), km 14 carr. Atzalan-Tlapacoyan ; 19.860°N , 97.220°W ; 1118 m ; 17 Aug 1996 ; V. García & R. Novelo-Gutiérrez leg. • same data but 1 ♂ ; 18 Aug 1996 • same data but 1 ♂ ; 15 Jul 2000 • same data but 1 ♂ ; 30 Jul 2014 • same data but 8 ♂♂ ; V. García & R. Novelo-Gutiérrez leg. • same data but 1 ♂ ; El Muro , km 14–16 carretera 131, Tlapacoya-Altotonga ; [ 19.878°N , 97.228°W ]; 1100 m ; 17 Aug 1996 ; R. Novelo-Gutiérrez leg. • same data but 1 ♂ ; 960 m • same data but 1 ♂ ; 1100 m ; 18 Aug 1996 • same data but 1 ♀ • same data but 1 ♂ ; 960 m ; 15 Jul 2000 • same data but 1 ♂ ; 1100 m • same data but 1 ♂ ; 960 m • same data but 1 ♂ ; 1100 m • same data but 1 ♀ ; Ixtatatahuia , carr. 131 Tlapacoyan- Altotonga ; [ 19.860°N , 97.219°W ]; 1000 m ; 21 May 1993 • same data but 5 ♂♂ ; 15 Jul 2000 • same data but 1 ♂ ; 14 Jul 2002 • same data but 3 ♂♂ 1 ♀ ; 31 Jul 2014 • same data but 1 ♀ ; Reserva el Pezma ; 19.914°N , 97.224°W ; 563 m ; 2 Aug 2001 • same data but 1 ♀ ; 30 Aug 2001 • same data but 1 ♂ ; 25 Jun 2002 • same data but 1 ♂ ; • same data but 1 ♂ ; Calcahualco , Barranca Cuapa ( Río Jamapa ); [ 19.129°N , 97.080°W ]; 1600 m ; 3 Jun 1996 ; L. Delgado leg. • same data but 2 ♂♂ ; Chontla , Ejido San Nicolasillo , Rancho de Don Enrique ; 21.248°N , 97.910°W ; 1064 m ; 13 Aug 2016 ; M. Trujano-Ortega & U. García-Vázquez leg.; IBUNAM ; IBUNAM :CNIN:OD23083 to 23084 • same data but 3 ♂♂ 1 ♀ ; 14 Aug 2016 ; IBUNAM :CNIN:OD23085 to 23088 • same data but 1 ♂ ; Ejido San Nicolasillo , Rancho de Don Ramón ; IBUNAM :CNIN:OD23089 • same data but 1 ♂ ; Coatepec , Río Ahuacatlán , cascada Los Tecajetes (escurridero); [ 19.472°N , 96.981°W ]; 1285 m ; 2 Jul 1994 ; R. Novelo-Gutiérrez leg.; INECOL • same data but 1 ♂ ; 3 Jun 1996 • same data but 1 ♀ ; 17 Jun 2009 • same data but 2 ♂♂ ; Río Huehueyapan ; [ 19.423°N , 96.996°W ]; 1225 m ; • same data but 1 ♂ ; Coscomatepec , 4.8 mi. north of Coscomatapec ; [ 19.093°N , 97.026°W ]; 1400 m ; 22–23 Jul 1966 ; O.S. Flint Jr. & M.A. Ortiz B. leg.; USNM ; SNMNH_383685 • same data but 1 ♀ ; Km 30 carr. Fortín-Huatusco ; 19.100°N , 97.033°W ; 1500 m ; 9 Jul 2007 ; J.A. Gómez-Anaya leg.; INECOL • same data but 1 ♂ 1 ♀ ; 17 Jun 2009 • same data but 2 ♂♂ ; 30 Jul 2014 • same data but 20 ♂♂ 3 ♀♀ ; Huatusco , 4.7 km . north Huatusco , Puente Ruiz Cortines ; [ 19.184°N , 96.959°W ]; 1300 m ; 23–24 Jul 1966 ; O.S. Flint Jr. & M.A. Ortiz B. leg.; USNM ; SNMNH_383661 to 383668, 383670 to 383684 • same data but 6 ♂♂ 2 ♀♀ ; Hotel Las Cañadas , Huatusco ; 19.178°N , 96.971°W ; 1320 m ; 22–23 Aug 2003 ; E. González-Soriano leg.; IBUNAM ; IBUNAM :CNIN:OD1696 to 1701, 22878 to 22879 • same data but 3 ♂♂ ; Teocelo , Teocelo , Puente Texolo ; [ 19.402°N , 96.986°W ]; 1100 m ; 11–13 Sep 1981 ; IBUNAM :CNIN:OD1659, 1689, 22827 • same data but 1 ♀ ; 26 Sep 1981 ; IBUNAM :CNIN:OD1676 • same data but 2 ♂♂ 1 ♀ ; 20 Jun 1982 ; IBUNAM :CNIN:OD1660 to 1661, 22901 • same data but 1 ♂ ; 26 Sep 1982 ; L. Rivera leg.; IBUNAM :CNIN:OD22869 • same data but 1 ♂ ; 23 Jul 1983 ; IBUNAM :CNIN:OD1662 • same data but 5 ♂♂ 1 ♀ ; 24 Jul 1983 ; IBUNAM :CNIN:OD1664 to 1669 • same data but 1 ♂ 4 ♀♀ ; 25 Jul 1983 ; IBUNAM :CNIN:OD1670 to 1674 • same data but 1 ♂ ; 26 Jul 1983 ; IBUNAM : CNIN:OD1675 • same data but 14 ♂♂ 1 ♀ ; 10 Jul 1987 ; E. González-Soriano & V. García leg.; IBUNAM :CNIN: OD1657, 1677 to 1682, 22876 to 22877 • same data but 5 ♂♂ 1 ♀ ; M.J. Westfall leg.; IBUNAM :CNIN:OD22870 to 22875 • same data but 5 ♂♂ 1 ♀ ; 26 Aug 1988 ; E. González-Soriano & V. García leg.; IBUNAM :CNIN:OD1683 to 1688 • same data but 1 ♂ ; 26 Jun 1993 • same data but 2 ♂♂ ; 19.402°N , 96.986°W ; 18 Jun 2002 ; E. González- Soriano leg.; IBUNAM :CNIN:OD22947 to 22948 • same data but 1 ♀ ; 9 Jul 2007 ; R. Novelo-Gutiérrez leg.; INECOL • same data but 1 ♀ • same data but 9 ♂♂ 1 ♀ ; 15 Aug 2015 ; E. González-Soriano leg.; IBUNAM ; IBUNAM :CNIN:OD22949 to 22954, 23077 to 23080 • same data but 1 ♂ ; R. Novelo-Gutiérrez leg.; INECOL • same data but 1 ♂ ; Tepetzintla , Sierra de Otontepec ; por brecha, ca, de 10 km al N de Tepetzintla ; 21.244°N , 97.859°W ; 685 m ; 30 Jul 2010 ; IBUNAM ; IBUNAM :CNIN:OD22880 • same data but 2 ♂♂ 1 ♀ ; Tlapacoyan , Arroyo Colorado , km 14 carr. 131 Tlapacoyan-Altotonga ; 19.882°N , 97.232°W ; 1000 m ; 12 Jul 2002 same data but 8 ♂♂ ; 9 Jul 2007 • same data but 1 ♀ ; Arroyo Colorado , km 18 carr. 131 Tlapacoyan-Altotonga ; [ 19.900°N , 97.235°W ]; 950 m ; 22 Jun 1990 • same data but 1 ♀ ; 26 Jun 1993 • same data but 1 ♂ ; 7 Jul 1994 • same data but 9 ♂♂ ; 12 Jul 2002 • same data but 2 ♂♂ ; Tlapacoyán , Cascada de Tomata ; 19.924°N , 97.220°W ; 535 m ; 6 May 2010 ; E. González-Soriano leg.; IBUNAM ; IBUNAM :CNIN:OD14619, 22865 . Description of male. Large and chiefly black-winged individuals.Males dimorphic, the dark morph characterized by the presence of a black tip on the wings and whitish patch proximal to it; the hyaline morph is smaller and with unmarked wings. Head . Chiefly black, labrum dark glossy blue, in some old specimens looks blackish, pale markings usually more extensive in hyaline, postclypeus can be pale cream in both morphs or blue in the hyaline morph, frons usually black, can have a pale cream or blue transverse line that can be complete or reduced to spots below the antennae, vertex usually with two pale spots anterolateral to lateral ocelli, occiput usually with complete pale occipital bar that can be reduced to lateral spots or absent, eyes black in life. Thorax . Prothorax colouration variable; lobes generally chiefly black, pale markings on lateral edges and around central fossa (in some specimens from San Luis Potosí and Hidalgo lobes mostly pale with black markings reduced to two longitudinal parallel discontinuous lines across medial and posterior lobe); for structure see Figs. 104 , 110–111 . Pterothorax with diffuse green metallic reflections on mesepisternum and mesepimeron, see Figs. 170–171 for typical back-tipped and hyaline morphs colour pattern. mesepisternum with entire or discontinuous narrow line above mesopleural suture reaching seven-tenths of length (young black-tipped and hyaline individuals with more extensive pale colour mainly on anterior half of its length, these markings sometimes forming broad irregular stripe); mesinfraepisternum pale (sometimes with ventral dark spot), pruinose on ventral half in mature individuals; mesepimeron with narrow line below mesopleural suture on posterior third of its length reaching antealar carina, this line seemingly continuous with mesepisternum line; second line usually complete covering 40–50% of mesepimeron width (in darker individuals can be reduced to spots above metathoracic spiracle and interpleural suture); metepimeron usually with lines each one-third of its width under metapleural suture and over metapleural carina, superior line widening at its upper end and thus forming a pale triangular area, these lines joining at the margin of the metepimeron (in some individuals, mostly in immature or hyaline ones, black area can be reduced to curved or boomerang-shaped line as in female, see Fig. 171 ); metinfraepisternum with ventral spot; sternum pale; legs pale but femora usually with dim dark markings on extensor surfaces and black rings at distal end; tibiae with black stripes on flexor and extensor surfaces usually restricted to first pair of legs, in some individuals also present on second pair but more diffuse; tarsi dark brown to black; claws reddish. Wings . Hyaline, with or without black tip; when present, black tip covers 20–30% of wing length, reaching at least proximal edge of pterostigma, and is always preceded by a white area; wings; vein that descends from the subnodus usually at or slightly proximal to the first Vx, in some specimens slightly distal to it; HW field between CuA and posterior margin with at least one supplementary vein in at least one wing; CuA in HW usually not forked, at most with one supplementary sector. Px: FW 39–54, HW 36–51. Abdomen . Black with pale areas as follow: S1 with ventrolateral spots; S2 with ventrolateral horizontal lines; S3–7 with basolateral spots; S9–10 and cerci pruinose on dorsum in mature individuals. Caudal appendages . Cerci: see Figs. 67–68 for structure. Paraprocts: rudimentary, never with well-defined upper and lower branches; superior lobe rudimentary, only distinguished by transverse groove; inferior lobe in lateral view with acute projection. Measurements. Abdomen: 38.5–51, FW: 35.5–49, HW: 33.5–47.5. Description of female Head . As in male but more extensive pale markings; pale stripe across postclypeus and frons always cream, never blue. Thorax . Prothorax as in male. Pterothorax: see Fig. 171 for typical colour pattern (in some individuals mesepimeron with more extensive clear markings, line above the mesopleural suture may be thickened to cover half width of metepimeron, metepimeron can be completely pale); depression mesad to mesostigmal lobe in mesostigmal plate angulated. Wings . Hyaline with slightly amber tint or with dark smoky area at the apex that colour can reach proximal margin of pterostigma; Px: FW 35–55; Px: FW 34–50. Abdomen . S1–7 pale colouration as in hyaline male or more extensive; S8 dorsal pale spot present or absent; S9 pale dorsal spot mushroom shaped, widened to rear. Ovipositor ending beyond tip of cerci. Measurements. Abdomen: 40–43, FW: 37.5–42, HW: 36.5–40. Diagnosis . Paraphlebia zoe , P. duodecima and P. kauil form a group whose cerci possess a well-developed mediodorsal flange armed with a sclerotized tooth on the distal margin ( Figs. 67–70 ). In dorsal view, the position of this tooth relative to the distal lobe of the cerci can form either a semi-straight angle as in P. duodecima ( Fig. 69 ) or a U-shaped gap as in P. zoe and P. kauil ( Figs. 67–68, 70 ). The gap formed between the sclerotized tooth and the distal lobe is wide in P. zoe , clearly larger than the tooth ( Figs. 67–68 ), while in P. kauil this gap is narrow, subequal to the size of the tooth ( Fig. 70 ). Paraphlebia duodecima is the only one of the three that shows completely reduced paraprocts with the posterior margin entire and lacking indication of a superior lobe ( Fig. 85 ), while in P. zoe and P. kauil there is a superior lobe delimited by a shallow groove and the inferior lobe bears an acute projection directed posteriorly ( Figs. 88, 90 ). In P. duodecima the posterior lobe of the prothorax is narrow, with the lateral edges of the posterior margin never reaching the notopleural suture ( Figs. 119, 125 ), while in P. zoe and P. kauil they are at least at the level or lateral to it ( Figs. 110–111 , 117, 126 ). Paraphlebia zoe and P. duodecima males are dimorphic, each with a dark-tipped morph, P. zoe black-tipped males are unique in having a whitish patch proximal to the black tip ( Fig. 54 ), while black-tipped males in P. duodecima are unique in having the smallest black wing area of all known species, the black colouration covering less than 15% of the wing length and never reaching the proximal edge of the pterostigma ( Fig. 40 ). This character should not be used by itself when attempting to identify P. duodecima , as young individuals of P. quinta and likely other black-tipped species can look similar when the black tip has not fully coloured (as in Fig. 163 ). Finally, the allopatric distribution of P. zoe (north of the Orizaba region in Veracruz , Mexico ; Fig. 148 ) helps in separating this species from all others in the genus. Paraphlebia duodecima is sympatric with P. kauila near the Mexico-Guatemala border ( Fig. 150 ). Natural history . A species with male dimorphism with black and hyaline male morphs. Romo-Beltrán et al. (2009) found differences in body size, wing length and thoracic muscular fat-mass volume among the two morphs. They also described the relationship linking territorial status and copulation success (territorial males achieve a higher mating success) but not with morph or size, and finally it was found that characteristics of the territory also explain mating success. This was indicated because in some territories there are a greater number of mating events than in others. Additionally, Bello-Bedoy et al. (2015) and Munguía-Steyer et al. (2010) found no relationship between morph and survival probabilities. Further differences related to allometry of sexual traits, immune response, and copulation time have been studied and discussed in P. zoe ( Bello-Bedoy et al. 2015 ; Ruiz-Guzmán et al. 2013 ; Wong-Muñoz et al. 2013 ). This species has been collected in shaded seepages and waterfall-sprayed walls within mountain cloud forest and tropical rainforest. At some sites in Veracruz (Barranca de Texolo), some individuals were found inhabiting seepages on vertical walls at the roadside. Distribution . This is the second most widespread species of the genus. It occurs from Xilitla in southeast San Luis Potosí state to Coscomatepec, south of Xalapa, in Veracruz state , including Hidalgo , Puebla , and Querétaro states. It has been collected from 120–2100 m .a.s.l. This species is the northernmost known of the genus. Remarks . This species is the most studied of the genus, with specimens deposited in numerous collections in the USA and Europe. It was described by Selys from an individual collected by Emile Sallé with no data regarding location or date of collection beyond “mexico” on the labels ( Fig. 140 ). This specimen is missing abdominal segments S7–10 ( Fig. 141 ), and although the main diagnostic characters are found in the caudal appendages and therefore cannot be corroborated in this specimen, the whitish patches proximal to the black wingtips and the distribution leave no doubt about its identity. In addition to the material studied, we have knowledge of material collected at the Biosphere Preserve El Cielo in the state of Tamaulipas, over 100 km N of the most northern record listed here. As this material is lost, we have decided not to list it as part of the species distribution.