New species of the damselfly genus Argia from Mexico, Central America and Ecuador with an emphasis on Costa Rica (Insecta: Odonata: Coenagrionidae) Garrison, Rosser W. Ellenrieder, Natalia Von Zootaxa 2017 4235 1 1 93 9X22G Garrison & von Ellenrieder Garrison & von Ellenrieder [151,670,1686,1712] Insecta Coenagrionidae Argia Animalia Odonata 5 6 Arthropoda species calverti sp. nov.   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 Odonataculminating in his contribution to the Neuroptera volume of the Biologia Centrali-Americana( 1901–1908).  Specimens examined. 35 ♂, 14 ♀. Types.   Holotype ♂: COSTA RICA,  CartagoProv., Reserva Tapantí,  1,310 m,  6 vii 1963, F. G. Thompsonleg. [ FSCA].  Paratypes: COSTA RICA,  AlajuelaProv.: 1 ♂, Peñas Blancas River Valley,  Monteverde Cloud ForestReserve, 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. Ramirezleg. [RWG];  CartagoProv.: 2 ♂, Río Chiriquí, just N of Peralta( 9°58'49'' N, 83°36'25'' W,  348 m),  25 iii 1910, P. P. Calvertleg. [ 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. Camposleg. [ INBIO];  2 ♂, 1 ♀, Tapantí{ 9°43' N, 83°46' W},  1,310 m,  6 vii 1963, F. G. Thompsonleg. [ FSCA];  1 ♂, same data but [RWG];  GuanacasteProv.: 1 ♂, 1 ♀[in tandem],  GuanacasteNational Park, Estación Cacao, SEside of Volcán Cacao( 10°55'53'' N, 85°27'43'' W,  1,299 m),  21–29 v 1992, M. A. Zumbadoleg. [ 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. Zumbadoleg. [ INBIO];  1 ♂, 1 ♀[in copula], same data but Quebrada Tempisquito( 10°57' N, 85°30' W,  600 m),  25 viii 1990, G. Varelaleg. [ INBIO];  1 ♂same data but  29 viii 1990, M. A. Zumbadoleg. [ INBIO];  1 ♂, 1 ♀[in tandem], same data but  6 v 1991, J. Zlotyleg. [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];  HerediaProv.: 2 ♂, 2 ♀[in copula],  La Selva BiologicalStation,  10 kmSW of Horquetas( 10°18' N, 84°3' W,  600 m),  24–26 iii 1995, TWD & A. Ramírezleg. [TWD];  3 ♂, 1 ♀,  La Selvaaltitudinal transect,  1,070 m,  11 iv 2001, R. Vargasleg. [CEH];  1 ♂, 1 ♀, Sarapiquí, Magsasay( 10°24'6'' N, 84°3'18'' W),  20 ix 1993, J. Bensteadleg. [CEH];  LimónProv., 1 ♂,  11 kmSSW of Pocora, Finca Las Brisas, on main river channel among boulders and rocks, Río Parisminaand tributaries ( 10°4'10'' N, 83°37'58'' W,  821 m),  31 i 2007, WAH, D. Wagner& M. Thomasleg. [RWG];  San JoséProv., 1 ♂,  15 kmWNW of San Isidro de El General, on large river below Yoga Ashramof 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];  PuntarenasProv.: 2 ♂, Finca Las Cruces,  4 miS 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. Ramírezleg. [RWG];  1 ♂, same data but  20 ix 2013[RWG];  1 ♂, 1 ♀[in tandem], same data but [ CSCA]. PANAMA,  ChiriquíProv.: 2 ♂, Potrerillos( 8°39' N, 82°29' W,  664 m),  18 ii 1935, J. W. MacSwainleg. [ UMMZ];  VeraguasProv.: 1 ♂, near Santa Fe,  Quebrada Alto de la Piedra( 8°30'53'' N, 81°7'19'' W,  850 m),  18 vi 2011, A. Donnellyleg. [ FSCA];  1 ♂, 1♀(in tandem) about  3 kmNW of Alto de Piedraon 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. Mauffrayleg. [ 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 cupraureaCalvert.   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. 148c); 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. 148d); 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(  CostaRica: PuntarenasProv., Monteverde, Río San Luis Arriba, Río San Luis, along river below cataract; ♀H 9123– 4 intandem 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. 107b) 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. 107e); mesostigmal lobe accompanied by a slight mesepisternal tubercle postero-distal to outer margin of erect lobe ( Fig. 107c).  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 holotypemale 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 inmales, Fw 15–19, Hw 13–17 infemales; postquadrangular cells Fw 5, Hw 4–5 inmales, Fw 4–6, Hw 4–5 infemales; RP2 at Fw 7–9, Hw 6–8 inmales, Fw 7–8, Hw 6–7 infemales. 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 (  CostaRica,  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. 148a) versus the almost linear ( Fig. 149a) and slightly arched ( Fig. 150a) cercus in  A. cupraureaand  A. oenearespectively. The paraproct of  A. oeneain lateral view ( Fig. 150b) 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. 148b) and  A. cupraurea( Fig. 149b) are more robust and are strongly divergent. The labrum in  A. oeneais 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. calvertiand  A. cupraurea(as in Figs. 126, 127). A  cuprea-like dark morph of  Argia fulgidaknown from  CostaRica, 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. calvertiin 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. fulgidais 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. calvertiand  A. cupraurea(as in Figs. 126–127). The genital ligula of  A. fulgidaand  A. oeneaare similar in possessing a broad campanulate (  A. fulgida, Figs. 128–130) to quadrate (  A. oenea, Figs. 131b, c) hood but differ as follows: in  A. fulgidathe thickened flagella are not fused at base and they do not arch entally toward first segment ( Fig. 128c); in  A. oenea, the bifurcate flagella are fused into a common stem that arches entally toward first segment ( Figs. 131a, c). The male appendage, genital ligula, and female mesostigmal plate morphology of dark and pale S2–6 morphs of  A. fulgidaare essentially the same.  Argia fulgidawas 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(  Valledel CaucaDept., San Jose, Río Dagua), but pale morphs of  A. fulgidaoccur on the western side of the Andes of Ecuador( Bolivar, Los Ríos, Manabi, Pichincha, Santo Domingo de los TsáchilasProvinces, Fig. 165). William Haber and Fred Morrison collected a pair of  A. fulgidaat a shallow forest stream in Ecuador( PichinchaProv., road from highway to Silanche Bird Sanctuary, 10 kmNW 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 CostaRican, two Panamanian, and four Ecuadorean specimens of  Argia fulgida(including the Pichinchapair) 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  Argiaspecies and to measure genetic divergence in previous studies ( Caesar & Wenzel 2009; Nava Bolaños et al. 2016). All sequenced dark specimens from  Panamaare 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. calvertican 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. calvertiand  A. fulgidaare 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. 1434659053 [716,1049,1973,1998] Costa Rica COSTA RICA 5 6 1 1 holotype 1434659087 1963-07-06 FSCA F. G. Thompson Costa Rica 1310 Reserva Tapanti 5 6 1 Cartago holotype 1434659074 [733,1027,2010,2034] Costa Rica COSTA RICA 5 6 1 paratype 1434659066 2007-08-01 W. A. Haber & L. Ramirez Costa Rica Penas Blancas River Valley 1037 6 7 10.307778 Monteverde Cloud Forest 21 -84.74027 Monteverde Cloud Forest Reserve 5 6 1 1 Alajuela paratype 1434659056 1910-03-25 FSCA P. P. Calvert Costa Rica 348 9.980277 Peralta 21 -83.60694 Rio Chiriqui 6 7 2 2 Cartago paratype 1434659077 1993-06-19 1993-06-30 1993-06-19 INBIO A. C. Amistad & Grano de Oro & P. Campos Costa Rica Turrialba 1120 9.818334 Grano de Oro 21 -83.45916 Tayutic 6 7 3 1 2 Cartago paratype 1434659068 [410,1432,295,320] 1963-07-06 FSCA F. G. Thompson Costa Rica 1310 9.716666 Tapanti 1299 -83.76667 6 7 3 1 2 Cartago paratype 1434659092 [151,462,332,357] 1963-07-06 FSCA F. G. Thompson Costa Rica 1310 9.716666 Tapanti 1299 -83.76667 6 7 1 1 Cartago paratype 1434659088 [475,932,332,357] Costa Rica Guanacaste 6 7 2 1 1 Guanacaste paratype 1434659055 1992-05-21 1992-05-29 1992-05-21 INBIO Estacion Cacao, SE & M. A. Zumbado Costa Rica 1299 10.931389 Volcan Cacao 21 -85.461945 6 7 1 Guanacaste paratype 1434659076 1990-08-31 1992-05-29 1992-05-21 INBIO M. A. Zumbado Costa Rica 590 10.956944 Cacao Station 21 -85.494995 Estacion Biologica Maritza 6 7 2 1 1 Guanacaste paratype 1434659028 1990-08-25 1992-05-29 1992-05-21 INBIO G. Varela Costa Rica 600 10.95 Quebrada Tempisquito 1296 -85.5 Estacion Biologica Maritza 6 7 2 1 1 Guanacaste paratype 1434659091 [396,1108,511,536] 1990-08-29 1992-05-29 1992-05-21 INBIO M. A. Zumbado Costa Rica 600 10.95 Quebrada Tempisquito 1296 -85.5 Estacion Biologica Maritza 6 7 1 1 Guanacaste paratype 1434659082 1991-05-06 1992-05-29 1992-05-21 INBIO J. Zloty Costa Rica 600 10.95 Quebrada Tempisquito 1296 -85.5 Estacion Biologica Maritza 6 7 2 1 1 Guanacaste paratype 1434659086 1989-05-08 Costa Rica 600 10.878889 Rio Gongora 21 -85.53861 6 7 2 1 1 Guanacaste paratype 1434659084 1995-03-24 1995-03-26 1995-03-24 La Selva Biological & A. Ramirez Costa Rica 600 10.3 La Selva Biological Station 1298 -84.05 6 7 4 2 2 Heredia paratype 1434659073 [249,1192,655,680] 2001-04-11 La Selva & R. Vargas Costa Rica 1070 La Selva 6 7 4 1 3 Heredia paratype 1434659075 1993-09-20 J. Benstead Costa Rica 10.401667 Magsasay 21 -84.055 Sarapiqui 6 7 2 1 1 Heredia paratype 1434659057 2007-01-31 D. Wagner & M. Thomas Costa Rica 821 10.069445 Rio Parismina 21 -83.63278 Finca Las Brisas 6 7 1 1 Limon paratype 1434659093 2010-05-02 Costa Rica Yoga Ashram 1124 Rio Division -83.83139 Tamara Budowski 6 7 1 1 San Jose paratype 1434659065 1983-04-24 Costa Rica 472 10.162222 Braulio Carrillo National Park 21 -83.93833 6 7 2 1 1 San Jose paratype 1434659085 1989-07-06 Costa Rica 800 10.16 Quebrada Sanguijuela 21 -83.96305 Parque Nacional Braulio Carrillo 6 7 3 1 2 San Jose paratype 1434659096 1975-02-21 Costa Rica 1220 8.769722 Finca Las Cruces 21 -82.95972 6 7 2 2 Puntarenas paratype 1434659090 2013-08-07 L. Ramirez Costa Rica Monteverde 1180 10.278333 Rio San Luis 21 -84.784164 Rio San Luis Arriba 6 7 3 1 2 Puntarenas paratype 1434659080 2013-09-20 L. Ramirez Costa Rica Monteverde 1180 10.278333 Rio San Luis 21 -84.784164 Rio San Luis Arriba 6 7 1 1 Puntarenas paratype 1434659101 [314,962,1052,1077] 2013-09-20 CSCA L. Ramirez Panama Monteverde 1180 10.278333 Rio San Luis 21 -84.784164 Rio San Luis Arriba 6 7 2 1 1 Puntarenas paratype 1434659098 1935-02-18 UMMZ J. W. MacSwain Panama 664 8.65 Potrerillos 1300 -82.48333 6 7 2 2 Chiriqui paratype 1434659120 2011-06-18 FSCA Quebrada Alto de la Piedra & A. Donnelly Panama 850 8.514722 Quebrada Alto de la Piedra 21 -81.12195 Santa Fe 6 7 1 1 Veraguas paratype 1434659102 2016-05-25 FSCA N. P. Santa Fe Road & W. Mauffray Panama 610 8.525556 Rio Maluba 21 -81.130005 6 7 2 1 1 Veraguas paratype