Paraphlebia Selys in Hagen, 1861

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 : 5-54

Paraphlebia Selys in Hagen, 1861

Paraphlebia Selys 1860: 435 (nomen nudum); Hagen [Selys in Hagen] 1861: 71 ( Paraphlebia zoe type species, by monotypy); Selys 1862: 8–9 (genus description); Felder in Felder et al. 1864: Tab. 83, fig. 6 (junior homonym); Brauer 1868: 361 (Agrionina key); Scudder 1882: 233 (cat.); Selys 1886: 33 (key); Kirby 1890: 122 (cat.); Calvert 1901: 59 (addition of generic characters); Higgins 1901: 136 (gizzard formulae); Calvert 1902: 31 (comparison with Thaumatoneura McLachlan, 1897 ); Calvert 1903: 133–134 (discussion on mimicry with Palaemnema Selys ); Calvert 1908: 461, 467 (distribution and biogeographic affinities); Cockerell 1908: 70 (comparison with Megalagrion umbratum (Scudder, 1890) and Trichocnemis aliena (Scudder, 1892) ; Calvert 1913: 260–261, 263 (legion Podagrion key, relationship with Phenacolestes Cockerell, 1908 ); Tillyard 1917: 284 (transferred to Megapodagrioninae); Munz 1919: 28 (fig. 69; Megapodagrionidae key); Kennedy 1925: 303 (comparison with megapodagrionids); Beatty & Beatty 1968: 807 (first mention of males colour and behaviour dimorphism); Paulson 1982: 251 (distribution); Bridges (1994: III.38; cat.); González-Soriano & Novelo-Gutiérrez 2007: 113 (distribution in Mexico); Fogarty et al. 2008 (support as sister group of Thaumatoneura ); Novelo-Gutiérrez 2008: 29 (larva description); Kalkman et al. 2010: 123 (discussion on larval characters and comparison with megapodagrionids); González-Soriano & Paulson 2011: 303 (discussion on endemism in Chiapas); Dijkstra et al. 2013: 20 (transferred to Thaumatoneuridae ); Hämäläinen 2016: 38 (cat.); Cuevas-Yañez et al. 2015: 517 (conservation status of P. zoe , P. hyalina and P. quinta ).

Paraphleoia Hagen 1861, Davies & Tobin 1984: 42 (typo or misprint in cat.).

Note on the authorship of Paraphlebia

Selys (1860) first mentioned Paraphlebia zoe Selys in a note comparing its coloration with that of Palaemnema paulina Drury, 1773 . However, he did so before the completion of the work where he originally intended to describe this species ( Selys 1862) making P. zoe Selys, 1860 a nomen nudum for not complying with the provisions of Art. 12 in the International Code of Zoological Nomenclature ( ICZN 1999). Hagen (1861) introduced Paraphlebia zoe Selys linking the name for the first time to a specimen from Mexico which was deposited in a collection (“Collection of Selys Longchamps”), and providing a (very succinct) diagnosis. This publication year was mainanied in some consequent works with various authorships given as: P. zoe to Hagen, 1861 ( Kirby 1890), P. zoe to Selys in Hagen, 1861 ( Calvert 1901), P. zoe to Hagen, 1861 ( Davies & Tobin 1984). Garrison (1991) discussed the authorship of Paraphlebia and P. zoe and decided that they were not “… adequately described, much less diagnosed.” Therefore, he considered them as nomina nuda, which made P. zoe Selys, 1862 the available name. From this point onwards, the authorship of the genus was attributed in different ways, e.g., Paraphlebia Hagen, 1861 ( Bridges 1994) ; Paraphlebia Selys in Hagen, 1861 ( Garrison et al. 2010; Hämäläinen 2016); Paraphlebia Selys, 1861 ( Dijkstra et al. 2013).

The authorship was revised in the present study. Paraphlebia zoe Selys in Hagen, 1861 was found to be the available name (for the reasons explained above) and therefore, Paraphlebia Selys in Hagen, 1861 should be considered the valid name by monotypy.

Diagnostic (modified from Garrison et al. 2010): Abdomen medium to large size (34.9–48.3 mm). Colouration highly variable: body dark brown to black with metallic green reflections with pale areas blue ( Figs. 6–7 , 26 , 152 , 160–167 ), yellowish (e.g., Fig. 22 ) or cream (e.g., postclypeus in Figs. 170–171 ); in preserved specimens the blue colouration can fade and appear cream or yellowish ( Figs. 2–5 , 15 , 29 ). Head: frons rounded, location of most posterior point of head at level of eyes. Thorax: prothorax posterior margin entire ( Fig. 110 ), with a medial bump ( Fig. 108 ) or deeply cleft ( Fig. 116 ); with ( Figs. 107–108 , 121–122 ) or without ( Figs. 110–120 ) laterally or posterolaterally directed lobes, subquadrate ( Figs. 123 ) or with two corniform dorsolateral projections ( Figs. 103 , 105–106 ); pterothorax second line complete ( Figs. 14–15, 19–20 ) or reduced ( Figs. 16, 18 , 23 ); thorax ( Fig. 18 ) and dorsum of S9–10 and cerci can become pruinose in mature individuals ( Figs. 34–35 , 152–157 , 161 , 164, 166–169 , 171 ); female mesostigmal plate with rounded ( Fig. 138 ) or angulated ( Fig. 139 ) depressions mesad to mesostigmal lobe. Wings: hyaline, often with amber tint ( Figs. 38, 41–44, 46–48 , 52–53, 55 ), with the tip slightly smoky ( Fig. 158 , 170 ♀ –171), or with an apical black tip with metallic blue reflections ( Figs. 39–40 , 45, 49–51 , 54 , 153 , 164– 167 , 170 ♂) in P. zoe preceded by a milky-white patch ( Figs. 54 , 170 ); no accessory crossveins basal to CuP, 1 or rarely 2 distal to it, shared with Thaumatoneura ; CuP closer to antenodal 1 than to 2; petiolation ending well beyond CuP for a distance as long as CuP or longer; vein descending from subnodus proximal to first post-quadrangular Vx to slightly distal to second post-quadrangular Vx; RP 3 slightly proximal to subnodus to slightly distal to subnodus; IR2 arising distal to subnodus; two or more supplementary sectors between IR1 and RP2, one supplementary sector between Rp2 and IR2 and two between IR2 and RP3, field between CuA and posterior margin with ( Figs. 57, 59 ) or without supplementary veins ( Figs. 56, 58, 60–61 ) and with one ( Fig. 60 ) to two ( Figs. 56, 58 ) supplementary sectors or lacking supplementary sector nor vein ( Fig. 61 ); pterostigma as long as four or more underlying cells, with proximal margin subequal to or slightly shorter than distal margin; legs with hind femora reaching mid-length of S2 or shorter. Abdomen: genital ligula lacking paired flagella on distal segment ( Fig. 1a, d ); S1 dark ( Fig. 16 ) or with lateral pale markings ( Figs. 35–37 ); S2 with a pale ventrolateral horizontal lines ( Figs. 33–37 ); S3–7 with pale basal spots or rings ( Figs. 34–37 ); S8 black (figs 33–35) or with a pale basal spot ( Figs. 36–37 ); S8–10 pruinose on dorsum in males ( Figs. 152–154 ), the pruinescence can be absent in young individuals ( Fig. 168 ), females with variable pale markings ( Figs. 129–137 ); dorsum of male S10 approximately flat. Cerci ( Figs. 62–80 ): forcipate, in dorsal view the first third of their length straight and slightly laterad, then bending inwards; from 30-80% two medial flanges and on the distal 0.2–0.35 a distal lobe; mediodorsal flange can be well ( Figs. 62–72 ) or poorly developed ( Figs. 73–80 ) and in dorsal view its mesal margin smoothly curved ( Figs. 63–65 , 70 ), nearly straight ( Figs. 67–69, 71 , 73–79 ), or convex ( Figs. 66 , 72 , 80 ); distal margin of mediodorsal flange variously shaped and armed with a sclerotized tooth ( Figs. 67–71 , 73–80 ), a blade ( Figs. 63, 65–66 , 72 ), or neither ( Fig. 64 ); paraprocts rudimentary, with a single lobe ( Figs. 83–86 ), a poorly developed superior lobe delimited by a shallow transverse groove ( Figs. 87–94 ), or well-developed superior and inferior branches ( Figs. 95–97 ); females with ovipositor surpassing posterior margin of S10 and going beyond the posterior margin of the cerci ( Figs. 131–137 ) or not ( Figs. 129–130 ); valves of ovipositor as in Fig. 1d .

Key to males

1. Mediodorsal flange of cerci well-developed ( Figs. 62–72 ), its maximum width at least 1.5 times width of distal lobe...... 2

1’. Mediodorsal flange of cerci poorly developed ( Figs. 73–80 ), its maximum width less than 1.5 times width of distal lobe … 10

2(1). FW with vein descending from subnodus always closer to first post-quadrangular Vx than to second ( Figs. 57–58 ); posterior margin of posterior lobe of prothorax semi-circular, at most with small medial convex section or notch and slight concavity towards lateral edges ( Figs. 110–120 )..................................................................... 3

2’. FW with vein descending from subnodus always closer to second post-quadrangular Vx than to first ( Figs. 42 , 56 ); posterior margin of posterior lobe of prothorax concave towards lateral edges, seemingly trifoliate, clearly armed with digitiform projections ( Fig. 108 )....................................................................... P. esperanza

3(2). In dorsal view width at base of distal lobe of cerci one-half or more of maximum width of mediodorsal flange Figs. 62–71 ). ................................................................................................... 4

3’. In dorsal view, base of distal lobe of cerci narrow, about one-third of maximum width of mediodorsal flange ( Fig. 72 )................................................................................................... P. chaak

4(3). Mediodorsal flange of cerci nearly straight or smoothly curved, in dorsal view its widest point closer to tip than base ( Figs. 62–65 , 67–70 ); lateral edges of posterior lobe of prothorax variously shaped but never angled........................ 5

4’. Mediodorsal flange of cerci convex, in dorsal view its widest point not surpassing half its length ( Fig. 66 ); lateral edges of posterior lobe of prothorax angulated ( Fig. 112 ).................................................... P. kukulkan

5(4). Distal margin of mediodorsal flange of cerci blade-shaped and with sclerotized tip or tooth ( Figs. 63, 65 , 67–70 )......... 6

5’. Distal margin of mediodorsal flange of cerci smoothly rounded, never blade-shaped and with sclerotized tip or tooth ( Figs. 64 )........................................................................................... P. akan

6(5). Paraprocts well-developed, clearly forked at apex, armed with well-defined upper and lower branches ( Fig. 81–82 )................................................................................................... P. hyalina

6’. Paraprocts rudimentary, never armed with well-defined upper and lower branches, at most with ill-defined superior lobe ( Figs. 85–94 )............................................................................................. 7

7(6’). Posterior lobe of prothorax subequal or wider than middle lobe, lateral edges at same height or laterad to notopleural suture; lateral edges of posterior lobe of prothorax straight or forming acute angle at junction with middle lobe ( Fig. 109–111 ); superior lobe of paraprocts rudimentary, only recognizable by a transverse groove ( Fig. 87–94 ); in lateral view inferior lobe of paraprocts with an acute projection ( Fig. 88–94 )..................................................................... 8

7’. Posterior lobe of prothorax narrower than middle lobe, lateral edges clearly mesad to notopleural suture; lateral edges of posterior lobe of prothorax forming obtuse angle at junction with middle lobe ( Fig. 119 ); superior lobe of paraprocts completely absent ( Figs. 85–86 ), inferior lobe in lateral view smoothly rounded ( Fig. 85 )........................... P. duodecima

8(7). Distal margin of mediodorsal flange of cerci armed with ventrally or postero-ventrally directed sclerotized tooth ( Fig. 67–68, 70 )................................................................................................. 9

8’. Distal margin of mediodorsal flange of cerci blade-shaped with sclerotized tip, wider than long ( Fig. 65 )....... P. chiarae

9(8). In dorsal view gap between distal tooth of mediodorsal flange of cerci and distal lobe broad, clearly larger than tooth ( Fig. 67–68 )......................................................................................... P. zoe

9’. In dorsal view gap between distal tooth of mediodorsal flange of cerci and distal lobe narrow and “U” shaped, subequal to size of the tooth ( Fig. 70 )............................................................................ P. kauil

10(1’). In lateral view lateral edges of posterior lobe of prothorax erect, clearly extending dorsad ( Figs. 98–99, 103 )........... 11

10’. In lateral view lateral edges of posterior lobe of prothorax not erect, extending caudad ( Figs. 100–102, 104 )............ 13

11(10). Posterior margin of posterior lobe of prothorax armed with two laterodorsal corniform projections ( Figs. 105–106 )...... 12

11’. Posterior margin of posterior lobe of prothorax concave towards lateral edges, seemingly trifoliate ( Fig. 107 )..... P. hunnal

12(11). Posterior margin of posterior lobe of prothorax with lateral projections dorsad, in frontal view these projections higher than medial part of posterior margin; paraprocts well-developed, clearly forked at apex, with well-defined upper and lower branches ( Fig. 105 )...................................................................................... P. flinti

12’. Posterior margin of posterior lobe of prothorax with lateral projections laterad, in frontal view these projections lower than medial part of posterior lobe; paraprocts rudimentary and never with well-defined upper and lower branches ( Fig. 106 )................................................................................................. P. kinich

13(10). Mediodorsal flange of cerci nearly straight, in dorsal view its widest point closer to tip than base ( Figs. 73–78 ); inferior lobe of paraprocts, in lateral view with an acute projection ( Fig. 92 )............................................ P. quinta

13’. Mediodorsal flange of cerci convex, in dorsal view its widest point not surpassing half its length ( Fig. 80 ); inferior lobe of paraprocts in lateral view smoothly rounded ( Fig. 87 )................................................ P. itzamna

Key to females

1. Ovipositor ending clearly beyond tip of cerci (without stylus) ( Figs. 131–137 ).................................... 2

1’. Ovipositor short, at most reaching tip of cerci (without stylus) ( Figs. 129–130 ).................................... 8

2(1). Lateral edges of posterior lobe of prothorax erect, extending dorsad in lateral view ( Fig. 99 ).................. P. kinich

2’. Lateral edges of posterior lobe of prothorax not erect, extending caudad in lateral view ( Fig. 100 )..................... 3

3(2). Posterior lobe of prothorax narrower than middle lobe, lateral edges mesad to notopleural suture ( Figs. 123–125 )......... 4

3’. Posterior lobe of prothorax subequal to or wider than middle lobe, lateral edges at same height or laterad to notopleural suture ( Figs. 126–128 )...................................................................................... 5

4(3). Posterior margin of posterior lobe of prothorax subquadrate or slightly concave towards lateral edges ( Figs. 123–124 ).................................................................................................. P. quinta

4’. Posterior margin of posterior lobe of prothorax rounded ( Fig. 125 ).................................... P. duodecima

5(3’). Mesostigmal plate depression rounded ( Fig. 138 )................................................... P. itzamna

5’. Mesostigmal plate depression angulated ( Fig. 139 ).......................................................... 6

6(5). Metepimeron completely pale; Eastern Chiapas and Guatemala ( Fig. 150 ).................................. P. kauil

6’. Metepimeron variously marked but never completely pale; Western Chiapas Sierra Madre and northwest to San Luis Potosí ( Fig. 148 )........................................................................................... 7

7(6’) Postclypeus pale colouration cream ( Figs. 170–171 ); HW field between CuA and posterior border usually with one extra sector ( Fig. 60 ); wings with smoky tip colouration usually clearly reaching distal border of Pt ( Figs. 170–171 ); Mexico from Coscomatepec, Ver., north to Xilitla, SLP, including Puebla, Hidalgo and Querétaro states ( Fig. 148 )............... P. zoe

7’. Postclypeus pale colouration pale blue to turquoise ( Fig. 158 ); HW field between CuA and posterior border usually without extra sectors ( Fig. 61 ); wings with or without smoky tip colouration but when present usually not reaching distal border of Pt ( Fig. 158 ); Mexico from Metlac, Ver., south to El Triunfo, Chis., including Oaxaca and Tabasco states ( Fig. 148 )..................................................................................................... P. hyalina

8(1’). FW with vein descending from subnodus always closer to second post-quadrangular Vx than to first ( Fig. 56 ); in frontal view digitiform projections of posterior lobe of prothorax extending ventrad (as in Fig. 122 ).................... P. esperanza

8’. FW with vein descending from subnodus always closer to first post-quadrangular Vx than to second ( Figs. 57–58 ); in frontal view posterior lobe of prothorax with digitiform projections extending slightly dorsad ( Fig. 121 ) or rectangular without digitiform projections................................................................................. 9

9(8’). Posterior lobe concave with well-developed digitiform projections........................................ P. ixchel

9’. Posterior lobe rectangular without digitiform projections............................................. P. kukulkan

Figures

FIGURE 1. Schematic representation of Paraphlebia’s generic characters; (1a) ♂ genital ligula: a – lateral, a’ – ental, scS – sclerotized shaft, Llb – lateral lobe, iF – internal fold; (1b) ♂ anterior and posterior hamuli lateral view: ah – anterior hamuli, pH – posterior hamuli; (1c) ♂ S2 ventral view; (1d) ♀ S9–10 ventral view, St – stylus, V1/V3 – first/third valves of ovipositor, Lam – basal plate of ovipositor.

FIGURES 2–11. Paraphlebia spp. colour pattern of head, frontal view.

FIGURES 20–27. Paraphlebia spp. colour pattern of thorax.

FIGURES 152–154. Field photographs. Paraphlebia chaak ♂ (152), Barranca a 50 m de la carretera hacia Chicacao, km 155, Suchitepequez, Guatemala, 25 Jul 2018; P. duodecima ♂Bw (153), Escurridero 200 m al N del camino.A 3.8 km al E de la garita municipal de Purulhá, Baja Verapaz, Guatemala, 21 Jul 2016; P. duodecima ♂Hw (154), Refugio del Quetzal, San Marcos, Guatemala, 27 Jul 2016.

FIGURES 158–160. Field photographs. Parabhlebia hyalina ♀ (158), Barranca de Metlac, Veracruz, Mexico, 26 Aug 2015; P. itzamna ♂Bw (159), Cusuco National Park, Cortés, Honduras, 18 Jul 2011; P. kauil ♂ (160), 1.5 km al N del ejido Benito Juárez El Plan, Chiapas, Mexico, 30 Jul 2016. Photograph 159 by MJ.

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 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 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 170–171. Field photographs. Paraphlebia zoe ♂Bw♀ (170), Teocelo, Veracruz, Mexico, 14 Jul 2020; P. zoe ♂ Hw (171), Teocelo, Veracruz, Mexico, 15 Apr 2020. Photograph 170 by C. Bota-Sierra.

FIGURES 12–19. Paraphlebia spp. colour pattern of head and thorax. Head frontal view (12–19); thorax lateral view (14– 19).

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

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 138–143. Paraphlebia spp. mesostigmal plates, type labels and type photograph. Mesostigmal plates dorsal view (138–139); type labels (140, 142–143); P. zoe holotype dorsal view.

FIGURES 35–43. Paraphlebia spp. colour pattern of abdomen and wings. Abdomen lateral view (35–37); left pair of wings (38–43).

FIGURES 44–51. Paraphlebia spp. left wings.

FIGURES52–61.Paraphlebiaspp.leftwings.Leftwingdetail (56–61):grayshade–post-quadrangular cells;sV–supplementary vein; sS – supplementary sector.

FIGURES 129–137. Paraphlebia spp. S8–10 lateral view.

FIGURES 62–66. Paraphlebia spp. caudal appendages (a) dorsal view; (b) mediodorsal view. W – maximum width of mediodorsal flange, w – maximum width of the distal lobe.

FIGURES 67–72. Paraphlebia spp. caudal appendages. (a) dorsal view; (b) mediodorsal view. W – maximum width of mediodorsal flange, w – maximum width of the distal lobe.

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 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 85–94. Paraphlebia spp. caudal appendages lateral view.

FIGURES 150–151. Paraphlebia spp. collection sites.

FIGURES 148–149. Paraphlebia spp. collection sites.