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 hyalina Brauer, 1871 Figs. 9 (head ♂ ), 22 (thorax ♂ ), 37 (abdomen ♂ ), 44, 56 (wings ♂ ), 62–63, 81–82 (appendages ♂ ), 120, 127 (posterior lobe of prothorax ♂ ♀ ), 135 (S8– 10 ♀ ), 142–144 ( types labels), 148 (map), 157–158 (habitus ♂ ♀ ). Paraphlebia hyalina , Brauer 1871: 175 ( ♂ and ♀ description; Veracruz ); Selys 1886: 34 (key and Brauer’s original description translation into French); Kirby 1890: 122 (cat.); Calvert 1901: 61 (key); Calvert 1913: 259 (legion Podagrion key); St. Quentin 1970: 258 ( lectotype , allotypoid and paratypoid designation [invalid]); Cuevas-Yañez et al. 2015 (conservation status); Ortega-Salas et al. 2021 (IUCN Red List) Paraphlebia quinta , Dijkstra 2014: 73 , 78 (misidentification; phylogeny) Paraphlebia n. sp. Beatty & Beatty 1962: 15 ( P. aliciae m.s.; Oaxaca ) Type series. Holotype . MEXICO • 1 ♂ ; [ Veracruz ] , [ Xalapa ], “Jalappa” [ Xalapa ]; [ 11–19 May 1865 ]; D. Bilimek leg.; NHMW ; [examined]. Paratypes . MEXICO • 1 ♂ ; [ Veracruz ] , [ Xalapa ], “Jalapa, Cuernavaca” [ Xalapa ]; [ 11–19 May 1865 ]; D. Bilimek leg. NHMW ; [not examined], “Cotypus” ; • 1 ♀ ; [ Veracruz ] , [ Xalapa ], “Jalappa” [ Xalapa ]; [ 11–19 May 1865 ]; NHMW ; [not examined], “ Allotypus ” . Other material examined. MEXICO • 1 ♀ ; Chiapas , Ángel Albino Corzo , Reserva de la Biósfera El Triunfo. Poligono III, El Quetzal. Campamento ; 15.717°N , 92.934°W ; 1818 m ; 19 Oct 2013 ; H. Ortega-Salas leg.; IBUNAM ; IBUNAM : CNIN:OD22727 • same data but 2 ♂♂ 1 ♀ ; 15.721°N , 92.939°W ; 1679 m ; 21 Oct 2013 ; IBUNAM : CNIN:OD22728 to 22730 • same data but 1 ♂ ; Oaxaca , San Bartolomé Ayautla , Sendero a la cascada Ayautla ; 18.059°N , 96.646°W ; 427 m ; 9 May 2016 ; H. Ortega-Salas & C. Raieli leg.; IBUNAM : CNIN:OD22994 • same data but 10 ♂♂ ; San Felipe Usila , Arroyo 1 km al S de Santa Cruz Tepetotutla sobre la carretera; 17.731°N , 96.560°W ; 1252 m ; 14 May 2016 ; IBUNAM : CNIN:OD23011 to 23020 • same data but 4 ♂♂ 2 ♀♀ ; 16 May 2016 ; IBUNAM : CNIN:OD23021 to 23025 • same data but 1 ♂ 2 ♀♀ ; Arroyo 3.4 km al S de Santa Cruz Tepetotutla sobre la carretera; 17.721°N , 96.552°W ; 1278 m ; IBUNAM : CNIN:OD23026 to 23028 • same data but 2 ♂♂ 1 ♀ ; Arroyo 5.8 km al S de Santa Cruz Tepetotutla sobre la carretera; 17.711°N , 96.552°W ; 1370 m ; IBUNAM : CNIN:OD23029 to 23031 • same data but 1 ♂ ; San Juan Bautista Valle Nacional , 2 small streams crossing road 5.9 mi. S. of Valle Nacional (local station No. 10); [ 17.725°N , 96.327°W ]; 626 m ; 24 Aug 1959 ; G.H. & A.F. Beatty leg.; FEM ; PSUC_ FEM _ 83396 • same data but 1 ♂ ; Km. 60 carretera federal 175 Tuxtepec-Oaxaca ; [ 17.719°N , 96.312°W ]; 820 m ; 28 Jul 1982 ; E. González-Soriano & V. García leg.; IBUNAM ; IBUNAM : CNIN:OD22809 • same data but 1 ♀ ; Km. 78.5 carretera federal 175 Tuxtepec-Oaxaca ; [ 17.630°N , 96.340°W ]; 1500 m ; 24 Sep 1989 ; E. González- Soriano leg.; IBUNAM : CNIN:OD22751 • same data but 3 ♂♂ 3 ♀♀ ; Mountain road from Tuxtepec to Cd. Oaxaca , waterfall streams crossing road, 13.4 mi. S of Valle Nacional ; [ 17.686°N , 96.330°W ]; 900 m ; 18 Aug 1959 ; G.H. & A.F. Beatty leg.; IBUNAM : CNIN:OD22903 to 22907 • same data but 1 ♂ ; Small waterfall streams crossing road 8.1 mi. S of Valle Nacional (local station 16); [ 17.718°N , 96.310°W ]; 762 m ; FEM ; PSUC_ FEM _ 83418 • same data but 1 ♂ ; PSUC_ FEM _ 83437 • same data but 1 ♂ ; 26 Aug 1959 ; PSUC_ FEM _ 83315 • same data but 1 ♀ ; PSUC_ FEM _ 83353 • same data but 1 ♂ ; 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 1 ♂ ; PSUC_ FEM _ 83414 • same data but 1 ♀ ; PSUC_ FEM _ 83417 • same data but 2 ♂♂ ; PSUC_ FEM _ 83425 to 83426 • same data but 1 ♀ ; 18 Aug 1959 ; PSUC_ FEM _ 83412 • same data but 1 ♂ ; 26 Aug 1959 ; PSUC_ FEM _ 83406 • same data but 1 ♀ ; 27 Aug 1959 ; PSUC_ FEM _ 83415 • same data but 21 ♂♂ 4 ♀♀ ; 12–17 Aug 1968 ; PSUC_ FEM _ 69087 • same data but 1 ♂ ; 24 Jul 1969 • same data but 3 ♂♂ ; 26 Jul 1969 • same data but 2 ♂♂ ; 25 Jul 1971 ; PSUC_ FEM _ 69260 • same data but 1 ♂ ; Santiago Comáltepec , La Esperanza ; [ 17.628°N , 96.368°W ]; 1630 m ; 5 Apr 1982 ; A. Ibarra leg.; IBUNAM ; IBUNAM : CNIN:OD22759 • same data but 12 ♂♂ 3 ♀♀ ; [ 17.627°N , 96.369°W ]; 1600 m ; 2 Jul 1995 ; J. Blackaller & A. Pérez leg.; IBUNAM : CNIN:OD22760 to 22767 • same data but 1 ♂ 1 ♀ ; IBUNAM : CNIN:OD8111 • same data but 2 ♂♂ ; 17.628°N , 96.368°W ; 1630 m ; R. Mendoza leg.; IBUNAM : CNIN:OD22752 to 22753 • same data but 5 ♂♂ 3 ♀♀ ; Waterfall stream crossing road 14.2 mi. S. of Valle Nacional “[local station No. 20]”; [ 17.668°N , 96.328°W ]; 1127 m ; 18 Aug 1959 ; G.H. & A.F. Beatty leg.; FEM • same data but 3 ♂♂ ; 26 Aug 1959 • same data but 8 ♂♂ 1 ♀ ; 17 Aug 1968 ; PSUC_ FEM _ 68950 • same data but 5 ♂♂ 3 ♀♀ ; Waterfall stream crossing road 8.5 mi. S of Valle Nacional “[local station No. 15]”; [ 17.716°N , 96.305°W ]; 731 m ; 18 Aug 1959 ; PSUC_ FEM _ 83029, 83305, 83409, 83413, 83422, 83432, 83459, 83469 • same data but 3 ♂♂ 3 ♀♀ ; 25 Aug 1959 ; PSUC_ FEM _ 83118, 83310, 83361, 83416, 83419, 83471 • same data but 2 ♂♂ ; 26 Aug 1959 ; PSUC_ FEM _ 83349, 83458 • same data but 3 ♂♂ 1 ♀ ; Waterfall streams crossing road 13.4 mi. S. of Valle Nacional “[local station 17]”; [ 17.674°N , 96.327°W ]; 883 m ; 18 Aug 1959 ; PSUC_ FEM _ 83317, 83456, 83463, 83474 • same data but 1 ♂ ; 26 Aug 1959 ; PSUC_ FEM _ 83317 • same data but 1 ♂ ; PSUC_ FEM _ 83408 • same data but 1 ♀ ; Santiago Comáltepec , Km. 56.9 carretera federal 175 Tuxtepec-Oaxaca ; [ 17.726°N , 96.326°W ]; 626 m ; 29 Jul 1990 ; E. González-Soriano & V. García leg.; IBUNAM ; IBUNAM : CNIN:OD1727 • same data but 1 ♀ ; IBUNAM : CNIN:OD22787 • same data but 1 ♀ ; IBUNAM : CNIN:OD22788 • same data but 8 ♂♂ 2 ♀♀ ; Santiago Zacatepec , Arroyo en carretera estatal Santa María Yacochi Santiago Zacatepec. 5 km al O de San Juan Metaltepec ; 17.178°N , 95.942°W ; 1380 m ; 26 Jun 2020 ; H. Ortega-Salas leg.; IBUNAM : CNIN:OD23109 to 23118 • same data but 1 ♂ ; Tabasco , Huimanguillo , 5.5 km al O de Chimalapa 1a sección, camino hacia Francisco J. Mújica ; 17.38°N , 93.63°W ; 375 m ; 27 Aug 2015 ; H. Ortega- Salas , M. Trujano-Ortega & U. García-Vázquez leg.; IBUNAM :CNIN:OD22963 • same data but 1 ♂ ; IBUNAM : CNIN:OD22965 • same data but 1 ♀ ; 4 Sep 2015 ; IBUNAM :CNIN:OD22974 • same data but 1 ♂ ; Veracruz , Fortín 2 mi. W. on Hw. 150, in Barranca de Metlac ; [ 18.910°N , 97.010°W ]; 984 m ; 26 Jul 1958 ; G.H. & A.F. Beatty leg.; FEM ; PSUC_ FEM _40817 • same data but 1 ♀ ; Barranca de Metlac ; 18.911°N , 97.011°W ; 26 Aug 2015 ; H. Ortega-Salas , M. Trujano-Ortega & U. García-Vázquez leg.; IBUNAM ; IBUNAM :CNIN:OD22958 • same data but 1 ♀ ; IBUNAM :CNIN:OD22959 • same data but 9 ♂♂ 2 ♀♀ ; Barranca de Metlac ; [ 18.910°N , 97.010°W ]; 28 Jun 1989 ; E. González-Soriano & V. García leg.; IBUNAM :CNIN:OD22798 to 22808 • same data but 1 ♂ ; USNM • same data but 4 ♂♂ 2 ♀♀ ; 8 Jul 2000 ; E. González-Soriano & L.E. González leg.; IBUNAM ; IBUNAM :CNIN: OD22810 to 22813 • same data but 1 ♂ 1 ♀ ; 18.911°N , 97.011°W ; 1 Aug 2011 ; E. González-Soriano & R. Novelo- Gutiérrez leg.; IBUNAM :CNIN:OD1702 • same data but 2 ♂ ; IBUNAM :CNIN:OD22814 • same data but 6 ♂♂ 2 ♀♀ ; IBUNAM :CNIN:OD22815 to 22820 • same data but 2 ♂♂ ; Fortín de las Flores , Barranca de Metlac. Río San Miguel ; INECOL • same data but 1 ♂ ; Stream above Río Metlac , 1.9 mi. SW & 0.5 mi. N Fortín ; [ 18.911°N , 97.011°W ]; 10 Aug 1965 ; D.R. & M.L. Paulson leg.; IBUNAM ; IBUNAM :CNIN:OD1708 . Description of male. Medium to large and chiefly black species; from the four species formerly described, this species is the only one in which males are strictly hyaline with no black morph known. The pale areas on the head are cream in immatures and turquoise or pale blue on mature individuals. Head . Chiefly black, labrum dark glossy blue, postclypeus turquoise to pale blue, in some individuals restricted to lateral third, maxillae black or pale, frons turquoise to pale blue, in some individuals restricted to lateral edges, vertex with two pale spots anterolateral to lateral ocelli, eyes black in life; antennae black, scape can have pale line on frontal surface or be completely black with pale distal margin, pedicel with pale distal margin. Thorax . Prothorax: chiefly black with variable pale markings on dorsum; see Fig. 120 for morphology. Pterothorax: chiefly black, usually with diffuse green metallic reflections over dark colouration on mesepisternum and mesepimeron; see Figs. 22 , 157 for colour pattern (mediodorsal carina can be pale, narrow pale line above mesopleural suture on mesepisternum can be entire, second line can be reduced to a short line of about one-fifth of the mesepimeron length starting at its posterior three-fourths, in young specimens the black mark in metepimeron reduced or absent); sternum pale; legs pale, femora with extensor surfaces and distal joints black; tibiae dark brown to black on flexor surfaces; tarsi dark brown to black; claws reddish. Wings . Hyaline, without an apical black tip; FW with the vein that descends from the nodus from slightly proximal to slightly distal to the first post-quadrangular Vx, always closer to the first than to the second postquadrangular Vx; 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 31–41, HW 28–35. Abdomen . Black with pale areas as follow: S1 usually with lateral or ventrolateral spots; S2 usually with ventrolateral horizontal lines, can be completely black; S3–7 usually with basolateral rings, in some males rings not completely closed on dorsum; S8 usually with basolateral spots; S9–10 and cerci pruinose on dorsum in mature individuals. Caudal appendages . Cerci: see Figs. 62–63 for structure. Paraprocts: well-developed, forked at apex, armed with well-defined upper and lower branches. Measurements . Abdomen: 40–48.3, FW: 34–40, HW: 33–38. Description of female Head . As in male. Thorax . See Fig. 158 for colour pattern. Prothorax: see Fig. 127 for morphology. Wings . Hyaline, with a slightly amber tint; Px: FW 33–43; Px: FW 29–38. Abdomen . S1–7 as in male but pale markings more extensive. S8 dorsal pale spot absent. S9 pale dorsal spot mushroom shaped, widened to rear. Ovipositor ending beyond tip of cerci. Measurements. Abdomen: 35–43, FW: 37.5–42, HW: 36.5–40. Diagnosis . This species is closely related to P. kukulkan , P. chiarae and P. akan and is diagnosed under the latter. Natural history . This species has been collected at small to medium-sized streams and seepages within mountain cloud forest and tropical rainforest. Found in sympatry with P. quinta at several locations. At Barranca de Metlac, Veracruz , it has been collected at seepages at vertical walls in heavily shaded microsites. Distribution . Our original hypothesis about the restricted distribution of this species changed with time as we visited more sites in cloud forest locations. Our results show that it is more widely distributed than was originally supposed. It is presently known from east-central and southeast Mexico , widespread from the eastern part of the Trans-Mexican Volcanic Belt to the Chiapas Sierra Madre. Veracruz , Tabasco , Oaxaca and Chiapas states. It has been collected from 375–1800 m .a.s.l. Remarks . The specimens used by Brauer to describe this species are labelled “Jalappa” and “Jalapa Cuernavaca”. The reference to Cuernavaca, located 250 km to the W, is undoubtedly an error, and the type locality is here restricted to Veracruz between Orizaba and Xalapa. The labels hold no information on when this material was collected, although we do know that Dominik Bilimek stayed in Mexico from May 1865 to June 1867 ( Bueno Bravo 2014 ), and thanks to correspondence published in his hometown newspaper Beilage zur Biene , we also know that he travelled and collected material on the way from Orizaba to Xalapa [sic! Chalappa] from the 11–19 May 1865 ( Bilimek 1865 ). This species has long been misinterpreted in the past because male dimorphism had not been suspected, so it is common to find hyaline specimens of P. quinta identified as P. hyalina as well as some of the previously undescribed species. St. Quentin (1970) designated a lectotype , an allotypoid and a paratypoid: “Von der ursprünglichen Series liegen noch 2 ♂♂ und ein ♀ vor, von denen ein ♂ als Lectotypus , das ♀ als Allotypoid und das zweite ♂ als Paratypoid betrachtet werden. Inv.-Nr. 605. “ [“From the original series there are still 2 ♂♂ and one ♀ , of which one ♂ is regarded as a lectotype , the ♀ as an allotypoid and the second ♂ as a paratypoid. Inv no. 605.”] These designations are unjustified, since the three specimens of the type series hold the labels with the original Brauer’s designations as: Holotypus , Allotypus and Cotype [actually a paratype ] ( Figs. 142–144 ). Given that this species has been abundantly and widely collected, it is easily distinguishable. Among the material examined (n =195), no specimens with black tips on the wings were found, so it is to be considered strictly hyaline. Recently, Y. Vega-Sánchez sent HOS pictures of Paraphlebia specimens collected in Zongolica, Veracruz ; 18.6414 , -96.9838 (Vega-Sánchez personal communication, October 2, 2021 ). They were confirmed to be P. hyalina which close the gap in between the nearest known localities: Barranca de Metlac to the north and San Bartolomé Ayautla to the south which makes finding this species in the southeast corner of Puebla state more likely.