Argia calverti Garrison & von Ellenrieder

Garrison, Rosser W. & Ellenrieder, Natalia Von, 2017, New species of the damselfly genus Argia from Mexico, Central America and Ecuador with an emphasis on Costa Rica (Insecta: Odonata: Coenagrionidae), Zootaxa 4235 (1), pp. 1-93 : 6-10

Argia calverti Garrison & von Ellenrieder , n. sp.

Figs. 1 (head ♂), 5 (labrum ♀), 13 (head, thorax, S1–4 ♂); 35 (head, thorax, S1–4 ♀); 56 (S7–10 ♂), 80 (S7–10 ♀), 106 (hind margin of prothorax ♀), 107 (mesostigmal plates ♀); 148 (appendages ♂); 165 (map); 172 (field picture of ♂); Table 2 (measurements).

Etymology. Named calverti (Latinized name) in honor of the late Philip Powell Calvert (1871–1961), in recognition to his valuable contributions to the knowledge of Central American Odonata culminating in his contribution to the Neuroptera volume of the Biologia Centrali-Americana (1901–1908).

Specimens examined. 35 ♂, 14 ♀. Types. Holotype ♂: COSTA RICA , Cartago Prov., Reserva Tapantí , 1,310 m, 6 vii 1963, F. G. Thompson leg. [ FSCA] . Paratypes: COSTA RICA , Alajuela Prov.: 1 ♂, Peñas Blancas River Valley, Monteverde Cloud Forest Reserve, Refugio Alemán , small tributary down trail from Refugio (10°18'28'' N, 84°44'25'' W, 1,037 m), 1 viii 2007, W. A. Haber & L. Ramirez leg. [RWG] ; Cartago Prov.: 2 ♂, Río Chiriquí, just N of Peralta (9°58'49'' N, 83°36'25'' W, 348 m), 25 iii 1910, P. P. Calvert leg. [ FSCA] ; 2 ♂, 1 ♀, A.C. Amistad, Turrialba, Tayutic, Grano de Oro , Chirripo (9°49'6'' N, 83°27'33'' W, 1,120 m), 19–30 vi 1993, P. Campos leg. [ INBIO] ; 2 ♂, 1 ♀, Tapantí {9°43' N, 83°46' W}, 1,310 m, 6 vii 1963, F. G. Thompson leg. [ FSCA] ; 1 ♂, same data but [RWG] ; Guanacaste Prov.: 1 ♂, 1 ♀ [in tandem] , Guanacaste National Park, Estación Cacao, SE side of Volcán Cacao (10°55'53'' N, 85°27'43'' W, 1,299 m), 21–29 v 1992, M. A. Zumbado leg. [ INBIO] ; 1 ♂, 1 ♀ [in tandem], same data but Estación Biológica Maritza, trail to Cacao Station , river in secondary forest, perching on rocks near water in rapids (10°57'25'' N, 85°29'42'' W, 590 m), 31 viii 1990, M. A. Zumbado leg. [ INBIO] ; 1 ♂, 1 ♀ [in copula], same data but Quebrada Tempisquito (10°57' N, 85°30' W, 600 m), 25 viii 1990, G. Varela leg. [ INBIO] ; 1 ♂ same data but 29 viii 1990, M. A. Zumbado leg. [ INBIO] ; 1 ♂, 1 ♀ [in tandem], same data but 6 v 1991, J. Zloty leg. [RWG] ; 1 ♂, 1 ♀, Río Góngora , on stones in the river and along river margins (10°52'44'' N, 85°32'19'' W, 600 m), 8 v 1989, CEH leg. [CEH] ; Heredia Prov.: 2 ♂, 2 ♀ [in copula], La Selva Biological Station , 10 km SW of Horquetas (10°18' N, 84°3' W, 600 m), 24–26 iii 1995, TWD & A. Ramírez leg. [TWD] ; 3 ♂, 1 ♀, La Selva altitudinal transect, 1,070 m, 11 iv 2001, R. Vargas leg. [CEH] ; 1 ♂, 1 ♀, Sarapiquí, Magsasay (10°24'6'' N, 84°3'18'' W), 20 ix 1993, J. Benstead leg. [CEH] ; Limón Prov., 1 ♂, 11 km SSW of Pocora, Finca Las Brisas, on main river channel among boulders and rocks, Río Parismina and tributaries (10°4'10'' N, 83°37'58'' W, 821 m), 31 i 2007, WAH, D. Wagner & M. Thomas leg. [RWG] ; San José Prov., 1 ♂, 15 km WNW of San Isidro de El General, on large river below Yoga Ashram of Tamara Budowski, small tributary of Río División , Quebrada Grande (9°24'36' N, 83°49'53'' W, 1,124 m), 2 v 2010, WAH leg. [RWG] ; 1 ♂, 1 ♀, Braulio Carrillo National Park (10°9'44'' N, 83°56'18'' W, 472 m), 24 iv 1983, K. D. leg. [RWG] ; 2 ♂, 1 ♀, Parque Nacional Braulio Carrillo, Quebrada Sanguijuela (10°9'36'' N, 83°57'47'' W, 800 m), 6 vii 1989, CEH leg. [CEH] ; Puntarenas Prov.: 2 ♂, Finca Las Cruces , 4 mi S of San Vito de Java {8°46'11'' N, 82°57'35'' W, 1,220 m}, 21 ii 1975, DRP leg. [RWG] ; 2 ♂, 1 ♀ [one pair in tandem], Monteverde, Río San Luis Arriba, Río San Luis , along river below waterfall (10°16'42'' N, 84°47'3'' W, 1,180 m), 7 viii 2013, L. Ramírez leg. [RWG] ; 1 ♂, same data but 20 ix 2013 [RWG] ; 1 ♂, 1 ♀ [in tandem], same data but [ CSCA]. PANAMA , Chiriquí Prov.: 2 ♂, Potrerillos (8°39' N, 82°29' W, 664 m), 18 ii 1935, J. W. MacSwain leg. [ UMMZ] ; Veraguas Prov.: 1 ♂, near Santa Fe, Quebrada Alto de la Piedra (8°30'53'' N, 81°7'19'' W, 850 m), 18 vi 2011, A. Donnelly leg. [ FSCA] ; 1 ♂, 1♀ (in tandem) about 3 km NW of Alto de Piedra on N.P. Santa Fe Road, tributary to Río Maluba , branches 1 & 2 (8°31'32'' N, 81°7'48'' W, 610 m), 25 v 2016, W. Mauffray leg. [ FSCA] .

A large red-eyed species with a brilliant cupreous thoracic dorsum and with dorsum of S1–7 largely blue, similar to pattern observed in Argia cupraurea Calvert.

Description of male holotype. Head: Entire face with metallic cupreous-red reflections except for the following: narrow apical margin of labrum, anteclypeus, base of mandibles and genae dark orange, small postocular spots blue, small pale spot anterolateral to lateral ocellus (similar to Fig. 1 ); antennae black, rear of head including margin except for narrow pale border along eye margin.

Prothorax entirely metallic cupreous-red except for blue anterior lobe. Mesothorax with mesepisternum brilliant cupreous-red, mesepimeron and mesinfraepisternum metallic dark purple; metepisternum and metepimeron blue interrupted by dark stripe along metapleural suture (as in Figs. 13 , 172 ). Wings slightly smoky with venation black; pterostigma trapezoidal, dark brown, surmounting 1.5 cell in left Fw, 1.0 cells in right Fw, 1 cell in left Hw wing, 1.5 cells in right Hw; postnodals Fw 18/18, Hw 15/16; postquadrangular cells Fw 5/5, Hw 4/ 5; RP2 at Fw 8/8, Hw 7/7. Coxae and trochanters brown anteriorly becoming blue posteriorly; femora, tarsi and armature black, tibiae black, pale externally.

Abdomen (as in Figs. 13 , 56 , 172 ) mostly blue; S1 black at basal 0.40 with ventrolateral extension laterally, remainder blue; S2 blue dorsally, laterally with a black stripe with pointed extension almost meeting dorsally at apical 0.20, below with a complete narrow, largely parallel blue stripe followed by black along ventral margin of tergite, annulus black; S3 blue with a narrow black stripe ventrally with a dorsal pointed extension at apical 0.05 but not meeting above, annulus black; S4–6 similar to S3 but posterior portion of black stripe more extensive; S7 blue with narrow ventral black stripe joining black annulus apically (as in Fig. 56 ); S8–10, blue dorsally with black stripe laterally, on S10 this stripe extending dorsally along posterior margin; torus pale, appendages black.

Genital ligula as in Argia cuprea ( Figs. 126, 127 ), with a pair of small laterally extended lobes at base near flexure (these hidden by membranous terminal fold) followed by a pair of flexible chitinized branches, each arising from a narrow stem at distal segment base extending laterad of the ligula; distal portion with narrow digit-like hood slightly inflated distally and extending over coiled digit-like paired flagella, the latter fused along a common stem basally and arching entally toward first segment.

Torus small, transversely oval, occupying entire ventral margin of torifer and not overlapping bilobed epiproct ( Fig. 148 c); base of epiproct and area around base black, apical portion of epiproct pale; cercus ( Fig. 148 ) robust, longer than wide, about subequal to paraproct, strongly convex dorsally, branched apically with distal margin of shorter outer branch ending at approximate right angle to longer decumbent inner branch; medial margin strongly convex; cercus ventrally strongly concave with a basal lobe externobasally ( Fig. 148 d); paraproct bilobed, its ventral branch narrow, bluntly pointed, its base at about 120° from acutely pointed anteriorly directed dorsal branch.

Dimensions. Hw 26.4, abdomen 33.8, total length 44.1.

Description of female paratype ( Costa Rica: Puntarenas Prov., Monteverde, Río San Luis Arriba, Río San Luis, along river below cataract; ♀ H 9123– 4 in tandem with male). Head ( Fig. 35 ): labrum with distal half ochre (as in Fig. 5 ), basal half metallic cupreous-red; anteclypeus ochre; postclypeus metallic cupreous; antefrons, genae and base of mandibles pale ochre; postfrons and remainder of epicranium metallic cupreous with pair of small spots anterolateral to lateral ocelli and small postocular spots pale bluish green; rear of head including margin except for narrow pale border along eye margin.

Prothorax metallic cupreous, anterior lobe pale blue, lateral 0.20 of posterior margin of hind lobe and propleuron pale ochre; pterothoracic dorsum ( Fig. 35 ) black with cupreous reflections and with a pale ochre antehumeral stripe widening slightly ventrally; entire metepimeron metallic cupreous, divided at upper 0.20 with branches enclosing an irregular elongate pale ochre spot, mesinfraepisternum with anterior half black, remainder ochre; metathorax ochre except for narrow black stripe at metapleural suture; venter of thorax ochre; S1 pale olive with basal ring of black and with narrow posterior black ring; S2 largely black with cupreous metallic reflections with narrow pale middorsal line abruptly widening at apical 0.20 and followed below by a narrower pale longitudinal stripe laterally and bordered ventrally with black, annulus black; S3–7 similar but with a pale basal ring not connected above and with a further narrowing of pale middorsal and lateral stripes (similar to Fig. 87 ); S8 like S7 but with dorsal black expanding laterally and enclosing a pale middorsal spot at apical 0.20 and with additional pale stripe ventrally, posterior portion of sternum pale; S9 similar to S8 but with dorsal black divided middorsally forming a pair of lobes ending at apical 0.40 and with pale blue area connecting with pale lateral stripe, S10 largely black with pale blue spot dorsally; cerci black, ovipositor pale except for black ventral margin ( Fig. 80 ).

Mesostigmal lobe ( Fig. 107 b) forming a small stubby upright lobe occupying medial 0.40 of slightly raised plate, medial margin of lobe well separated from branching middorsal carina by a low carina extending from medial base of lobe, medial margin of lobe strongly recurved in posterior view ( Fig. 107 e); mesostigmal lobe accompanied by a slight mesepisternal tubercle postero-distal to outer margin of erect lobe ( Fig. 107 c).

Variation in paratypes. On one male the pale postocular spot is absent on the right side and present as a small spot on the left side ( Fig. 1 ). The holotype male is more heavily marked with black than majority of other specimens; S2 on most males has a lateral blue stripe extending full length (similar to Fig. 15 ) and in some, posteriorly connecting with dorsal blue; in others, there is a greater lateral extension of dorsal blue on S10. Females show little variation but some have the majority of S10 blue and the degree of metallic coloration on the thorax is tempered by age; older females tend to be less metallic. One female (Estación Cacao) has a greatly reduced dark humeral stripe similar to Fig. 34 . Pterostigma surmounting 1–2 cells in males and females; postnodals: Fw 14–19, Hw 14–16 in males, Fw 15–19, Hw 13–17 in females; postquadrangular cells Fw 5, Hw 4–5 in males, Fw 4–6, Hw 4–5 in females; RP2 at Fw 7–9, Hw 6–8 in males, Fw 7–8, Hw 6–7 in females. Dimensions. ♂: Hw 25.8 ± 1.52 [23.8–27.6], abdomen 33.8 ± 1.71 [31.2–36.1], total length 43.9 ± 2.27 [40.8–47]. ♀: Hw 26.4 ± 1.32 [23.5–28.2], abdomen 31.4 ± 1.77 [29–34], total length 41.3 ± 2.15 [38.2–44.5].

Diagnosis. Male belongs to a distinctive group of species possessing an entirely metallic cupreous thoracic dorsum and, in life, dorsal half of compound eyes red, although postmortem preservation may preclude the practical use of the latter as a diagnostic character. Within its range ( Costa Rica, Panama, Fig. 165 ) male can be distinguished from similar metallic red species with largely blue S2–6 ( A. cupraurea , A. oenea ) by large size (Hw 23.8–27.2 mm; total length 40.8–47.0 mm; N = 10) compared to A. cupraurea (Hw 20.2–22.5 mm; total length 36.1–40.0 mm; N = 10) and A. oenea (Hw 18.5–22.6 mm; total length 33.8–39.7 mm; N = 10) and by the robust and strongly curved cercus ( Fig. 148 a) versus the almost linear ( Fig. 149 a) and slightly arched ( Fig. 150 a) cercus in A. cupraurea and A. oenea respectively. The paraproct of A. oenea in lateral view ( Fig. 150 b) is almost linear with both branches small and of about the same size, almost bilobate; the dorsal or ventral branches of the paraproct of A. calverti ( Fig. 148 b) and A. cupraurea ( Fig. 149 b) are more robust and are strongly divergent. The labrum in A. oenea is largely pale ( Fig. 3 ) but mostly metallic in A. calverti ( Fig. 1 ) and A. cupraurea (as in Fig. 2 ). The hood of the genital ligula in A. oenea ( Fig. 131 ) is quadrate, not linear and digit-like as in A. calverti and A. cupraurea (as in Figs. 126, 127 ).

A cuprea -like dark morph of Argia fulgida known from Costa Rica, western Panama, and Ecuador ( Fig. 165 ) occurs within the range of A. calverti . Male of this color morph is unique among metallic species within the distribution range of A. calverti in having dorsum of S2–6 entirely black except for a narrow basal blue ring ( Fig. 10 ); in the three other sympatric metallic species, A. calverti ( Fig. 13 ), A. cupraurea ( Fig. 14 ), and A. oenea ( Figs. 15–16 ), the dorsum of S2–6 has at least basal half or more blue ( Figs. 13–15 ) or violet ( Fig. 16 ). S 8 in A. fulgida is blue with the posterior half to third black ( Fig. 54 ), differing from the entirely blue S 8 in A. calverti ( Fig. 56 ), A. cupraurea ( Figs. 57–59 ) and A. oenea ( Figs. 61–62 ). The broad explanate hood of the apical segment of the genital ligula in A. fulgida ( Figs. 128–130 ) is shared with that of A. oenea ( Fig. 131 ), and both differ from the largely long linear hood present in A. calverti and A. cupraurea (as in Figs. 126–127 ). The genital ligula of A. fulgida and A. oenea are similar in possessing a broad campanulate ( A. fulgida , Figs. 128–130 ) to quadrate ( A. oenea , Figs. 131 b, c) hood but differ as follows: in A. fulgida the thickened flagella are not fused at base and they do not arch entally toward first segment ( Fig. 128 c); in A. oenea , the bifurcate flagella are fused into a common stem that arches entally toward first segment ( Figs. 131 a, c).

The male appendage, genital ligula, and female mesostigmal plate morphology of dark and pale S2–6 morphs of A. fulgida are essentially the same. Argia fulgida was described from a single male from Muzo, Colombia, which belongs to the pale morph with dorsum of S2–6 with at least the basal half or more blue ( Figs. 11–12 ). Its appendages and genital ligula are illustrated here for comparative purposes ( Figs. 129 , 147 ). We have seen only one other specimen representing the pale morph from Colombia (Valle del Cauca Dept., San Jose, Río Dagua), but pale morphs of A. fulgida occur on the western side of the Andes of Ecuador (Bolivar, Los Ríos, Manabi, Pichincha, Santo Domingo de los Tsáchilas Provinces, Fig. 165 ). William Haber and Fred Morrison collected a pair of A. fulgida at a shallow forest stream in Ecuador (Pichincha Prov., road from highway to Silanche Bird Sanctuary, 10 km NW of Pedro Maldonado) whose male has a largely dark S2–6 as figured for Central American A. fulgida ( Fig. 10 ). Additionally, a molecular analysis comparing mt DNA 16S sequences from one Costa Rican, two Panamanian, and four Ecuadorean specimens of Argia fulgida (including the Pichincha pair) shows that specimens displaying the two color morphs do not segregate into two groups based on this ribosomal gene ( Table 5, Fig. 191 ), which was used to separate Argia species and to measure genetic divergence in previous studies ( Caesar & Wenzel 2009; Nava Bolaños et al. 2016). All sequenced dark specimens from Panama are identical to several pale specimens from Ecuador, and the other sequenced specimens differ from them by 2 to up to 6% of the about 500 mtDNA 16S base pairs sequenced. At the present time we have found no evidence—morphological or molecular—that justifies treating these two color morphs as separate species, although future studies involving other genetic markers might prove otherwise.

Female of A. calverti can be easily distinguished from other similar species by the short, stubby mesostigmal lobe ( Fig. 107 ) compared to the longer more foliate lobe in A. fulgida ( Fig. 103 ), A. cuprea ( Figs. 104, 105 ), A. cupraurea ( Fig. 108 ), and A. oenea ( Figs. 109, 110 ). Thoracic and abdominal coloration for females of all five species are similar ( Figs. 30–38 , 80–85 ).

Males of A. calverti and A. fulgida are two of the five species from North and Central America possessing an entirely metallic cupreous thoracic dorsum. In both of them a pair of chitinized branches arises from a narrow stem at the base genital ligula distal segment. Each branch arches dorsally (ectally), and becomes broader at tip where its outer margin is armed with an irregular series of small spines. These branches, which are best seen in dorsal (ectal) view, are flexible and extend laterally to dorsolaterally depending on preservation ( Figs. 128–130 ). We have not found the armature of these structures to be species specific. Females also have a degree of metallic thoracic coloration, but this is confined to the medial half of the mesepisternum. In addition to their metallic thoracic coloration, they can usually be identified also by the presence of a pair of tuberculate processes or swellings each mediolaterally along the hind lobe of the prothorax, which gives a trilobate appearance to the hind margin ( Fig. 106 ), although this condition may not be so pronounced in some females.

The keys below should allow for the identification of both sexes of these five species, although examination of the male genital ligula and details of the mesostigmal plates and associated mesepisternal tubercles coupled with overall body pattern may be necessary to confirm some identifications.

Figures

FIGURE 191. Bootstrapped Maximum-Likelihood tree of dark and pale morphs of Argia fulgida based on mt 16 S sequences of nine specimens. The tree is drawn to scale, with branch lengths measured in the number of substitutions per site.

FIGURES 1 – 8. Color pattern of head of metallic species. Head, anterior view (1 – 4); labrum, anterior view (5 – 8).

FIGURES 13 – 16. Color pattern of male head, thorax, and S 1 – 4 of metallic species. Head, dorsal view, thorax and S 1 – 4, lateral view.

FIGURES 172 – 174. Field photographs of metallic species. (172) A. calverti, male from Alto San Luís, Puntarenas Prov., Costa Rica, 20 x 2013; (173) A. cupraurea, male at rocky river 4 km S of Route 32 between La Unión and Flores, Limón Prov., Costa Rica, 27 v 2013; (174) male of A. oenea from Hacienda El Rodeo, San José Prov., Costa Rica, 28 v 2013. Photograph (172) by WAH; (173) by NVE; (174) by RWG.

FIGURES 53 – 62. Color pattern of male S 7 – 10 of metallic species, lateral view.

FIGURES 126 – 128. Genital ligula of metallic species. (a) lateral view; (b) ectal view; (c) entolateral view.

FIGURES 148 – 150. Male caudal appendages of metallic species. (a) mediodorsal view; (b) lateral view; (c) dorsal view; (d) left cercus, ventral view.

FIGURES 35 – 38. Color pattern of female head, thorax, and S 1 – 4 of metallic species. Head, dorsal view, thorax and S 1 – 4, lateral view (35 – 37); pterothorax, lateral view (38).

FIGURES 80 – 88. Color pattern of female S 7 – 10 of metallic species, lateral view (80 – 86); S 2 – 5, dorsolateral view (87, 88).

FIGURES 106 – 108. Female thoracic structures of metallic species. Pronotal posterior lobe, dorsal view (106). Mesostigmal plates (107 – 108): (a) frontal (anterior) view; (b) detail of left mesostigmal lobe, frontal view; (c) dorsal view; (d) detail of right mesostigmal lobe, dorsal view; (e) detail of right mesostigmal lobe, posterior view; (f) anterolateral view.

FIGURES 30 – 34. Color pattern of female head, thorax, and S 1 – 4 of metallic species. Head, dorsal view, thorax and S 1 – 4, lateral view (30, 31); pterothorax, lateral view (32).

FIGURES 165 – 166. Distribution of metallic species.

FIGURES 129 – 131. Genital ligula of metallic species. (a) lateral view; (b) ectal view; (c) entolateral view.

FIGURES 9 – 12. Color pattern of male head, thorax, and S 1 – 4 of metallic species. Head, dorsal view, thorax and S 1 – 4, lateral view.

FIGURES 145 – 147. Male caudal appendages of metallic species. (a) mediodorsal view; (b) lateral view; (c) dorsal view.

FIGURES 103 – 105. Female mesostigmal plates of metallic species. (a) frontal (anterior) view; (b) detail of left mesostigmal lobe, frontal view; (c) dorsal view; (d) detail of left mesostigmal lobe, dorsal view; (e) detail of right mesostigmal lobe, posterior view; (f) anterolateral view.

FIGURES 109 – 110. Female mesostigmal plates of metallic species. (a) frontal (anterior) view; (b) detail of left mesostigmal lobe, frontal view; (c) dorsal view; (d) detail of left mesostigmal lobe, dorsal view; (e) detail of right mesostigmal lobe, posterior view.