Paracordulia calcarulata, Fleck & Haber, 2022

Fleck, Günther & Haber, William A., 2022, Paracordulia calcarulata, new species from Ecuador and notes on the genus Paracordulia Martin, 1907 (Odonata: Anisoptera: Corduliidae s. str.), Zootaxa 5124 (5), pp. 551-564 : 552-561

Paracordulia calcarulata sp. nov.

( Figures 1–14 )

Material. Ecuador, Sucumbíos Province. Holotype ♂, Playas de Cuyabeno village, Río San Francisco, 850 m N of Nicky Lodge on Río Cuyabeno , 0.19774, -75.92435, ca 210 masl; 31.I.2016; collectors Santiago Villamarin, William Haber, Fred Morrison, and Jeff Boettner; specimen dry stored ( INABIO, Haber EC-489) . Paratype ♂, Río Libertad, 8 km W Tarapoa, ca 240 masl; 0.12255, -76,41886, 12.I.2014; collectors Eli Wyman, David Wagner, Michael Veit; specimen stored in ethanol ( FSCA, Haber EC-848) .

Etymology. The species name is derived from the Latin calcar (spur or dewclaw) in reference to the two small spurs or dewclaw-like structures present on the male cerci. Clearly visible in dorsal view, they allow easy differentiation of the new species from P. sericea .

Description of holotype. A medium-sized dragonfly with long legs and long cerci relative to other body dimensions and with body showing strong green metallic reflections on head, thorax, and abdomen, and lacking lemonyellow markings ( Fig. 1 ).

Head. Face, frons and vertex covered by long bristle-like black setae except ventral margin of labrum and ventral margin of clypeal lobes bordered by long pale setae ( Fig. 2 ). Labium yellowish light brown, labrum and clypeus orange-brown, frons with lower margin yellowish brown to brownish orange, its upper portion turning progressively dark brown and with strong green or greenish blue metallic reflections (depending on light incidence) ( Figs 2–3 ). Vertex dark brown with green or bluish green metallic reflections (depending on light incidence), strongly bilobed dorsally thus defining a pair of blunt tubercles ( Figs 2–3 ). Antennae dark brown with long flagellum. Eyes green in life, in contact dorsally for a short distance (ca 0.8 mm) in their anterior part ( Fig. 3 ). Posterolateral margin of eyes with a small indentation (remnant of larval eye, often encountered in corduliids) (visible on Fig. 3a ). Occiput dark brown, large in dorsal view with a rounded posterior margin and with a rather long triangular anterior development ( Fig. 3 ).

Thorax. Abundantly covered by pale hair-like setae. Legs long compared to other body dimensions ( Figs 1 , 4 ), hind leg (18.5 mm) being distinctly longer than half of HW length (ca 64% of HW) and distinctly longer than half of abdominal length excluding anal appendages (ca 59% of abdomen). Legs with tibiae and tarsi blackish to dark brown, profemora and mesofemora brown at base becoming gradually dark brown toward apex, metafemora blackish/dark brown ( Fig. 4 ). Tibial keels present only on fore and hind legs, occupying 20% on the protibiae and 86% on the metatibiae. Ventral tooth of tarsal claws well developed and situated at about 3/5 to 3/4 of claw length. Prothorax light brown to yellowish, hind lobe with scattered long setae, the margin rounded with a row of short setae of uniform length ( Fig. 4 ). Synthorax brown with strong green metallic reflections, these last slightly less marked on posterior metepisternum, posteroventral metepimeron, and infraepisterna ( Figs 1 , 4 ).

Wings ( Figs 1 , 5 ): hyaline with light saffron tinge occupying an ill-defined costal fringe from base to Pt, and with apices slightly smoky; two areas with diffuse and inconspicuous light saffron tinge also present: the first occupying basal 2/3 of FW and in HW a surface grossly delimited by anal triangle, the fork of RP and the anal loop, and the second much smaller, centered on oblique vein (distal vein of the bridge). Veins black to dark brown. Membranulae well developed, brown, as long or nearly as long as anal triangle in HW ( Fig. 4 ). Pt dark brown, short (ca 5% of wing length) and trapezoidal with distal margin more oblique than proximal one. FW nodus located distally with nodal ratio of ca 1.24 (we define the nodal ratio as: distance from base of wing to nodus / distance of nodus to wing apex; a ratio of 1 means nodus at mid-length of the wing, ratio <1 means nodus shifted basally, ratio> 1 means nodus shifted distally); HW nodus shifted basally with nodal ratio ca 0.82; FW Ax 12 (left wing)—11 (right wing) of the first rank (between C and ScP) and 12 (left wing)—11 (right wing) of the second rank (between ScP and R); FW Ax of the first rank aligned (left wing) or sub-aligned (distal penultimate and antepenultimate crossveins of the right wing) with those of the second rank but only Ax1 and Ax2 distinctly reinforced and bracketlike HW Ax 7–8 of first rank and 7–8 of second rank with those of first rank and those of second rank well aligned, reinforced and bracketlike to Ax4; FW Px 6–7; HW Px 8; cross veins distal to Pt 3 or 4 (left FW); base of pseudo-IR1 distinctly distal to Pt in FW and below distal margin of Pt in HW; posterior closing of arculus at the level of Ax 2 in FW and slightly proximal to Ax 2 in HW; sectors of arculus meeting at base in FW and very shortly united in HW; bridges with two crossveins placed respectively just below and distal to subnodal transverse; hypertriangles and HW discoidal triangles free; FW discoidal triangles 2-celled, FW subtriangles large, 3-celled; no HW subtriangles (submedian space crossed only by basal CuP between anal triangle and anal loop); position of anteroproximal angle of HW discoidal triangle shortly but distinctly proximal to posterior crossvein of arculus; FW and HW Rspl well defined and covering one row of 6 cells; FW and HW Mspl well defined, covering in FW one row of 4 cells and in HW one row of 5–6 cells; right FW discoidal field with two rows of cells from distal angle of triangle to proximal part of Mspl, with three rows of cells at Mspl and again with 2 rows of cells to wing margin, left FW discoidal field with two rows of cells from distal angle of triangle to proximal part of Mspl, with three rows of cells at Mspl and 2 cells distal, again with two rows of cells of a distance of three cells, and with three cells at wing margin; MA and MP grossly parallel up to the distal part of Mspl, distally slightly convergent (left wing) or distinctly convergent (right wing); anal loop with 18–17 cells and with distinct midrib forking and vanishing distally; anal loop with 3 cells at submedian space margin and distally dilated with distinct toe/sole (posterior evagination) with sole (CuAb) made by 3 cells; posterior margin of anal loop (first branch of CuP&AA) and posterior wing margin separated by 2 rows of cells except postero-distal cell of the loop separated by one cell from posterior margin; anal triangle well defined, rather well elongated and 2-celled; anal angle well defined but tornus not strongly angulated.

Abdomen. Slender, slightly longer than HW ( Fig. 1 ). Brown on S1–2 and lateral anterior part of S3, remainder of S3 to S10 with dorsolateral coverage dark brown to black and with ventral part brown to dark brown; distinct metallic green reflections and some light cupreous gloss on dorsum of S2, most of dorsolateral part of S3 (except an ill-defined triangular spot at the antero-lateral fourth where metallic gloss can be considered absent or inconspicuous), and from dorsolateral part of S4 to anterior dorsolateral half of S7; metallic intensity diminishing progressively from S4 to S7; no trace of pale spots on lateral parts of S8–10 ( Figs 1 , 12 ); anal appendages blackish to dark brown, base of epiproct rufous-brown. In lateral view abdomen moderately swollen at S1–2, narrowing from S3 to anterior half of S5, then slightly expanding again from distal half of S6 to S8 ( Fig. 1 ); in dorsal view slightly constricted on S3, expanding again slightly and progressively from distal S6 to posterior margin of S7, then slightly constricted again on S9. Lateral carina extending from anterior margin of S4 to posterior margin of S8 and progressively better marked from S4 to S8. Secondary accessory genitalia: anterior lamina small, occupying approximately the anterior 2/10 of genital fossa and presenting a posterior margin with lateral low concavities and a median weakly bulbous convex lobe; hamule simple, strongly directed posteroventrally– but not bent at right angle in lateral view—distinctly narrowing distally and ending as a pointed process slightly curved dorsally and laterally; S2 genital lobe long, narrow, distinctly directed backward, blunt at apex, and bearing on ventro-apical edge a clump of long and strong setae ( Figs 6–7 ). Vesica spermalis: as in Figures 8–9 , with two apical asymmetric flagella, median flagellum stronger, much sclerotized and longer than right lateral flagellum, and with sclerotized part of distal segment (V4) slightly asymmetric and ending as a distinct distal horn [note: little sclerotized structures like right flagellum and to a lesser extent balloons present a different shape when dry or immersed in ethanol (compare Figs 8a–c and Figs 9a–c )]. S7 sternite with a sharp median longitudinal carina; S8 sternite bearing a distal rudimentary pilose complex (see Machado & Costa 1995 and Fleck 2017), appearing as a short sclerotized, T-shaped structure occupying ca 2/10 of sternal length followed distally by a short apical field of setae occupying ca 2/10 of sternal length ( Figs 10, 13 ); Tshaped structure with short low rounded median carina bearing a few aligned distinct setae and with low transverse fold bearing on posterior margin strong setae sparsely and irregularly distributed ( Fig. 10a ). Cerci blackish to dark brown, relatively thin and long, much longer than S9+10 (ratio of cerci length / length of S9+10 = 1.4) and nearly as long as S8+9 ( Fig. 11 ); pilosity of their inner margin well marked with: (i) a median fringe, extending from basal 1/10 to basal 5/10 of the length, made by long to very long setae [length reaching 20-25% that of cercus] directed distinctly backward and slightly dorsally and (ii) a dense subapical clump of setae directed medially and ventrally ( Figs 11–12 ). posterior half of cerci in dorsal view distinctly forcipate ( Fig. 11 ) and shaped as follows: (i) outer margin distinctly tripartite, basally straight (at about 3/10 of the length) with latero-ventral carinae clearly distinct from above (these occupying basal 1/10 to basal 3/10), and distally with two inverted curvatures, thus distinctly S-shaped, with proximal curvature less marked and shorter than distal curvature; (ii) inner margin bipartite, with a basal portion long and almost straight (very slightly S-shaped and extending on basal 6/10) ending as a short small spur, and with a shorter distal part strongly forcipate. Cerci in lateral view showing: (i) a ventral margin distinctly undulated and biconvex with basal convexity extending from base to about proximal 3/10 of length and bearing the strong lateral carina close to ventral margin and with a longer distal convexity occupying 5/10 of the length and lacking carina ( Fig. 12 ); (ii) dorsal margin slightly undulate with shallow median concavity at mid-length and occupying about 2/10 of cercus length ( Fig. 12 ). Epiproct distinctly triangular, relatively narrow (in ventral view not significantly overhanging the cerci), its apex shortly truncated and reaching dewclaws of the cerci (at 6/10 of cerci length) ( Fig. 13 ).

Measurements. Total length (including caudal appendages) 42.5, HW length 28.0, FW length 29.0, FW Pt 1.8 (on C vein), abdomen length (including caudal appendages) 31.5, cerci 3.6, epiproct 2.2.

Variations in paratype. Very similar to holotype differing as follows: slightly larger, bluish metallic gloss on frons slightly more pronounced ( Fig. 3b ), saffron and smoky tinges of the wings slightly more intensive and more extended, FW Ax 11 of the first rank and 11 (left wing)—12 (right wing) of second rank, HW Ax 7–6 of first rank and 7 (the fifth forked)—6 of second rank, FW Px 6–6, HW Px 8–7, MA and MP distally parallel or only very slightly convergent in both FW due to discoidal field with three rows of cells distal to Mspl ( Fig. 14 , compare with Figs 1 , 5 ). Posterior margin of S2 genital lobe more strongly curved ( Fig. 14 , compare with Fig. 4 ).

Measurements. Total length (including caudal appendages) 45.0, HW length 30.0, FW length 31.0, FW Pt 2.0 (on C vein), abdomen length (including caudal appendages) 33.5, cerci 3.8, epiproct 2.4.

Differential diagnosis. Paracordulia calcarulata is similar to P. sericea ; both species have almost identical wings—the few venational differences being probably due to individual variability—and have similar general habitus (compare Figs 1 , 5 , 14 and Figs 15–16 ).

They differ as follows:

- (1) P. sericea is slightly more robust (and probably also slightly larger on average with total length 45–46 mm and HW length 31–34 mm for the known males vs respectively 42.5–45 mm and 28–30 mm), with abdomen slightly more stoutly built; for example, S8 of P. sericea is about as long as wide whereas it is distinctly longer in P. calcarulata with ratio length/width ca 1.25 (compare Fig. 13 and Fig. 17 );

- (2) S2 genital lobe of P. sericea is slightly narrower and more sharply pointed;

- (3) the posterior hamule of P. sericea is strongly bent at a right angle so that most of the ventral margin is parallel to the body axis ( Fig. 16 , detail; also, Geijskes 1970: p. 16 and fig. 5). The posterior hamule of P. calcarulata is not bent at a right angle and the ventral margin is distinctly not parallel to the body axis ( Figs 1 , 4 , 7 , 14 );

- (4) pale spots are reported to be present on S8–10 of P. sericea ( Geijskes 1970) [not mentioned by Selys (1871) or Martin (1907, 1914)]; based on the photographs of this species at our disposal they are hardy identifiable, possibly strongly faded (some inconspicuous ill-defined blotches seem to be still faintly visible on Fig. 16 ). No pale spots are present in P. calcarulata ( Figs 1 and 14 , images taken shortly after death and still showing living colors);

- (5) the pilose complex of S8 is rudimentary in P. sericea , appearing as a longitudinal weak carina ending at basal 7/10 of sternite and with a field of long setae occupying the distal 3/10 of the sternite ( Fig. 17 ). In P. calcarulata the field of setae occupies the distal 2/10 and the sclerotized structure is shorter and T-shaped ( Fig. 10 );

- (6) the cerci are distinctly different in both species permitting easy separation (compare Figs 11–12 and Figs 18–22 ; note that the apex of both cerci of the P. sericea lectotype are broken); in P. sericea the basal part bearing the lateral carina is longer, occupying about 1/2 of the cercus length (about 1/ 3 in P. calcarulata ), whereas the distal curved portion is distinctly shorter, occupying about 2.5/10 of cercus length (about 4/ 10 in P. calcarulata ); P. sericea lacks the inner small projections, which resemble spur/dewclaw-like structures but instead has rounded pads with striae; in lateral view the ventral margin of the cercus appears bipartite with a basal half linear or slightly convex and the distal half concave ventrally in P. sericea , thus differing from the smooth undulations present in P. calcarulata .

A note on drawings of the anal appendages of P. sericea published in Garrison et al. (2006): the figures published by these authors show a rather strange distal part of the cerci compared to those in Martin (1907) and those of the paralectotype ( Figs 20–22 ), since their apices appear less convergent and their distal margins less rounded in dorsal view, and with a ventral indentation in lateral view ( Fig. 23 ). The drawings were made from photos of the lectotype, which has broken apices (unknown to the authors at that time), and with a quality of photos not showing the finely broken extremity of the right cercus (Garrison, pers. comm.).

Amended diagnosis of the genus Paracordulia . Geijskes (1970), in his study on the South American genera of corduliids, published a diagnosis of Paracordulia based exclusively on wing venation. Later Garrison et al. (2006) revised the diagnosis and included corporal characters. With the addition of this new species and with two new female specimens of Paracordulia , some previous generic characters merit discussion:

- FW discoidal field narrowing distally. The paratype of the new species and also one undescribed female from Ecuador present parallel MA and MP, therefore the strict character “FW discoidal field narrowing distally” can no longer be used.

- Copper reflections on abdomen and lateral pale spots on last three segments. The new species in addition to light copper reflections on abdomen also shows distinct green metallic reflections; moreover, no trace of pale spots is visible on S8–10. Apparently in some females the abdomen does not present metallic reflections even if some parts are shiny.

- Male tibial keels on all legs. Mesotibial keels are absent in the new species. The mesotibial keels seem to be absent in the lectotype of P. sericea ( Fig. 16 ), as shown on the oriented left mesotibia (a slight dilatation is visible near apex, but this is a common feature observed in many Odonata ).

- Female sternum 9 projected distally to about posterior margin of S10 [unique character after Garrison et al. (2006)]. This character suffers exceptions and cannot be applied to all species of the genus: the female described by De Marmels (1983) under “ Paracordulia sp. 2 ” presents the sternum of S9 not strongly covering S10, and a female from Ecuador (probably not conspecific with “ Paracordulia sp. 2 ” even if possibly closely related) also has the sternum of S9 not strongly projected–not more than many other corduliids species–and ending far from the posterior margin of S10 ( Fig. 24 ).

We propose the following amended diagnosis for the genus Paracordulia based on morphology of both adults and larvae:

Adults. Brown to black corduliids with at least some metallic reflections on part of head, part of thorax, and generally part of abdomen (at least in males), and lacking contrasting lemon yellow markings; vertex with two distinct tubercles; legs long compared to other body dimensions with hind legs distinctly longer than half of HW length; FW discoidal triangle with 2 cells, FW subtriangle with three cells, FW MA and MP converging distally or sub-parallel throughout, FW nodus distinctly located distally, and HW nodus located basally (FW nodal ratio> 1.20, HW nodal ratio <0.90); anal loop elongate with midrib and sole well developed (midrib not reaching postero-distal margin of the loop) and with three cells at anterior margin; male tibial keels on fore and hind legs, absent or extremely reduced on middle legs; anterior lamina small, confined to about anterior 2/10 of genital fossa, its posterior margin with median weakly bulbous convex lobe; posterior hamule entire, bent caudally and hook-like; distal segment of vesica spermalis with distal horn and two flagella; sternum of male S9 with rudimentary pilose complex; vulvar lamina triangular/acuminate; sternum of female S9 variably projected beneath S10, able to reach distal margin of S10.

Larvae (based on Fleck & Neiss 2012a and a photo of a F-0 larva from French Guiana). Somewhat lightly built with spidery aspect and slightly depressed; body almost entirely covered by numerous spinelike setae of modest to moderate size and robust but softer curved setae of various sizes; functional part of the eyes small, globular and very slightly upturned; ventral side of prementum with premental groove deep and very distinct; distal margin of prementum slightly and irregularly crenelated, each crenelation being itself finely serrated and separated from adjacent crenelation by a pair of spinelike setae; distal margin of labial palps with strong and finely serrated indentations, each bearing several raptorial setae (about 5–6 for median indentations) separated into two rows, distal one made by short setae, proximal one made by long setae; legs long and spindly with hind leg distinctly longer than body length; dorsal abdominal hooks present (on S5–9) and backward directed; lateral abdominal spines present on S8 and S9, strongly developed (at least as long as length of respective tergites at sagittal line), divergent (those of S8 distinctly) with their apices upturned, and with those of S9 overtaking the apex of the anal pyramid; S10 small and embedded in S9; anal pyramid relatively small with epiproct slightly shorter than paraprocts in dorsal view, and with cerci well developed, only slightly shorter than epiproct; antepleurites present and well developed on S4–6 and S8.

The keys presented by Martin (1907, 1914) and Geijskes (1970) are outdated since some genera are absent. In Garrison et al. (2006) the couplet keying Paracordulia males (key to males) is based on the shape of HW membranula and shape of S2 genital lobe. In some Paracordulia specimens the HW membranula reaches the end of anal triangle and S2 genital lobe is not clearly triangular, it is therefore preferable to complete the couplet 7 (p. 146) of Garrison et al. as follow: “7(6)—Combination of the two following characters present: vertex with pair of tubercles and one HW cubito-anal crossvein; HW membranule not surpassing end of anal triangle; genital lobe not distinctly rounded”, “7’—Combination of the two following characters absent: vertex with pair of tubercles and one HW cubito-anal crossvein; HW membranule surpassing end of anal triangle; genital lobe rounded or quadrangulate”; in the key to females (p. 151), the character related to the shape of S9 sternum must be removed from couplet 16.

Figures

FIGURE 1. Paracordulia calcarulata sp. nov., holotype. General habitus in left lateral view. Scale bar 10 mm.

FIGURES 2–3. Paracordulia calcarulata sp. nov., 2. paratype. Head in frontal view. 3. a) holotype, head in dorsal view slightly tilted forward and prothorax in lateral view; b) paratype, head in dorsal view.

FIGURES 4–5. 4. Paracordulia calcarulata sp. nov., holotype. Head, thorax, base of wings legs and first abdominal segments in left lateral view. 5. Paracordulia calcarulata sp. nov., holotype. Wings in ventral view.

FIGURES 6–9. 6. Paracordulia calcarulata sp. nov., paratype. Secondary genitalia in ventral view: a) overview of the structure; b) detail of hamule and genital lobes. 7. Paracordulia calcarulata sp. nov., holotype. Secondary genitalia in left lateral view. 8. Paracordulia calcarulata sp. nov., paratype. Vesica spermalis air dried: a) ventral view; b) left lateral view; d) right lateral view. 9. Paracordulia calcarulata sp. nov., paratype. Vesica spermalis in ethanol: a) ventral view; b) left lateral view; d) right lateral view.

FIGURES10–14. 10. Paracordulia calcarulata sp. nov., paratype. Abdominal segment 8 in ventral view: a) with light incidence suitable for a better view of the chaetotaxy; b) with light incidence suitable for a better view of sclerotized T-shaped structure. White arrows indicating T-shaped sclerotized structure, black arrow indicating field of long setae. 11. Paracordulia calcarulata sp. nov., holotype. Two last abdominal segments and anal appendages in dorsal view. 12. Paracordulia calcarulata sp. nov., paratype. Two last abdominal segments and anal appendages in lateral view. 13. Paracordulia calcarulata sp. nov., paratype. Four last abdominal segments and anal appendages in ventral view. 14. Paracordulia calcarulata sp. nov., paratype. General habitus in left lateral view. Scale bar 10 mm. Rectangle: detail of secondary genitalia.

FIGURES 15–16. 15. Paracordulia sericea, lectotype with labels. General habitus in dorsal view. Scale bar 10 mm. 16. Paracordulia sericea, lectotype. General habitus in left lateral view. Scale bar 10 mm. Circle: detail of the secondary genitalia.

FIGURES 17–24. 17. Paracordulia sericea, lectotype. S8 (left) and S9 (right) in ventral view. Note the S8 almost as long as wide and with a rudimentary “pilose complex” (see text). Scale bar 1 mm; white arrow indicating longitudinal weak carina, black arrow indicating field of long setae. 18. Paracordulia sericea, lectotype. Last abdominal segment and anal appendages in dorsal view. Note the broken apex of cerci. Scale bar 2 mm. 19. Paracordulia sericea, lectotype. Last abdominal segment and anal appendages in dorsal view. Scale bar 2 mm. 20. Paracordulia sericea, paralectotype. Last abdominal segment and anal appendages in dorsal view. Scale bar 2 mm. 21. Paracordulia sericea, paralectotype. Last abdominal segment and anal appendages in dorsal view. Scale bar 2 mm. 22. Paracordulia sericea. Last abdominal segment and anal appendages in dorsal view published in the work of Martin (1907); a) dorsal view; b) lateral view. 23. Paracordulia sericea. Last abdominal segment and anal appendages in dorsal view published in the work of Garrison et al. (2006); a) dorsal view; b) lateral view. 24. Paracordulia sp. female. Last abdominal segments in lateral view. Note the sternite 9 not reaching ventral posterior margin of S10.