Mesamphiagrion risi (De Marmels, 1997)

Bota-Sierra, Cornelio Andrés & Echeverri, Martha Isabel Wolff, 2013, Taxonomic revision of Mesamphiagrion Kennedy, 1920 from Colombia (Odonata: Coenagrionidae), with the description of four new species, Zootaxa 3718 (5), pp. 401-440 : 409-413

Mesamphiagrion risi (De Marmels, 1997)

Figs. 1 c (habitus ♂), 1d (habitus ♀), 8a–e (diagnostic traits), 13d (S7– 10 ♂), 14h (S7–10 ♀), 16 (map)

Enallagma ovigerum: Ris 1918: 117 , fig. 58; (description of male and female, illustration of male S10);—De Marmels 1989: 250-251, figs. 16–23 (description and illustrations of male diagnostic characters);—Steinmann 1997: 245 (synonymic list).

" Coenagrionidae A." Arango & Roldán 1983: 102, fig. 26 (larva figured).

Cyanallagma ovigerum: De Marmels 1990a: 74 , fig. 2 (illustration of male genital ligula).

Cyanallagma risi De Marmels 1997: 135 –157, fig. 84 (description, key, map, and illustrations);—Tsuda 2000: 31 ( Colombia);—Heckman 2008: 541, fig. 637 (in part, key, and illustrations).

Mesamphiagrion risi : von Ellenrieder & Garrison 2008: 17, Figs. 17, 42, 63, 74 (key, map, and illustrations of male diagnostic characters);—Garrison et al. 2010: 275–279, fig. 1770 (in part, synonymic list, and illustrations);—Pérez-Gutérrez & Palacino-Rodríguez 2011: 213 (recorded from Cundinamarca Dept., Colombia).

Mesamphiagrion ovigerum: Altamiranda 2009: 13 (record from Antioquia, misidentification);—Bota-Sierra et al. 2010: 334 (record from Antioquia, misidentification).

Examined material (80 specimens). Antioquia: CEUA: 1 ♂, Municipality Caldas, Township La Valeria, N6°5' W75°38' 2000 m. iv.2009, Leg: C. Bota. Municipality Medellín, Corregimiento San Antonio de Prado: Township Yarumalito: 1♂, N6°13'34" W75°41'22" 2300 m, 4.vii.2008, Leg: J. Cardona-D. & C. Bota. 1 ♀, 1.ix.2008, Leg: C. Bota. Township Astilleros: 2 ♂ and 1 ♀ N6°13' W75°39' 2100 m, 3.ix.2009, Leg: C. Bota. Corregimiento de Santa Elena: 2 ♂, 21/viii/2008, Leg: A. Bustamante. 5 ♂, Municipality Envigado, Quebrada El Salado N6°8' W75°34' 1850m, 27.xii.99 Leg: M. Restrepo & A. Botero. 1 ♂, Municipality Girardota, Township El Palmar, N6°19' W75°25', 2300m, 24.vii.2009, Leg: N. Uribe & G. Valencia. 5 ♂, Municipality Amalfi, Township La Quiebra, Finca El Edén, Potrero El Hueso, N6°52'13,8" W75°6'30,2" 1770m, 15.i.2010, Leg: C. Bota, J. Cardona-D. & L. Urrea. 1 ♂, Municipality Valdivia, Township San Fermín, Finca Montenegro, 2000m, 15.vii.2009, Leg: L. Urrea. Municipality San Vicente, Township Chaparral, Finca La Mosca, N6°15'56,9" W75°21'34,7" 2200m: 1♀, 8/xii/ 2007, Leg: C. Bota. 2 ♂ and 1 ♀, 20.iii.2010. 2 ♂ and 2 ♀, 12/ii/2011 Leg: C. Flórez & N. Urquijo. 2 ♂ and 1 ♀, 6.iv.2008, Leg: C. Bota. 5 ♂ and 2 ♀, 28.vi.2009 Leg: A. Bustamante, J. Ortiz & A. Clavijo. 1 Larva, 4.xi.2012, Leg: C. Bota, C. Flórez & K. Mejia. 1 ♂, Municipality Ríonegro, Township Vilachuaga, N6°6' O75°24' 2140m, 6.ix.2011, Leg: J. D'Leon. 1 M, Municipality Carmen de Viboral, Universidad de Antioquia, Seccional de Oriente, N6°6’20,6’’ W75°23’11,9” 2000m, 22.ix.2011, Leg: J. D'Leon. 3♂ and 3 ♀, Municipality Cocorná, Township El Viao, N6°3'46,8" W75°13'11,3" 1955 m. 29.vii.2009, Leg: C. Bota. 1 ♂ and 1 ♀, Municipality Marinilla, Township Altos del Mercado, Finca Los Orillos, N6°11' W75°18' 2100m, 23.iv.2010, Leg: C. Bota. 1 ♂ and 1 ♀, Municipality Guatapé, Township La Peña, N6°12' W75°10' 1920m, 30.ix.2008 Leg: C. Bota. 3♂, Municipality Anorí, Township El Roble, Reserve Arrierito Antioqueño, N6°59' W75°6' 1680m, 26–27.v.12. Leg: J. Zapata, W. Valencia, A. Vélez, J. A. Cogollo & C. Bota. Boyaca: 1♀, Municipality Arcabuco, vii.2011, Leg: P. Lizarazu. Huila: 10 ♂ and 3♀, Municipality El Pital, Vereda El Recreo, Quebrada EL Burro, N2°16'27" W 75°53'32.81" 2250m. 5.vii.2013, Leg: C. Bota & C. Flórez. Santander: 3♂, Municipality Zapatoca, Vereda Alto de las Aguilas, N 6°52'35" O 73°19'56" 2300m. 13.viii.2013, Leg: C. Bota, C. Flórez & G. Valencia. MEFLG: 1♂, Municipality Medellín. 1♂, Municipality Bello. Cundinamarca: ICN: 1♂, La Cadilla. ANDES: Municipality San Francisco de Sales, Township San Miguel, 1600m: 3♂, 8.v.2011, Leg: E. Realpe. 1 ♂, vii/2006, Leg: L. Pérez & E. Realpe. 3♂ and 1♀, Municipality Alban, 04°53'53" N 74°25'31" O, 2000msnm, 13.xi.2004, Leg: L. Pérez. 1♂, Municipality Guaduas, Township La Playa, N5°04'25" W74°33'48", 1539m, 20.ii.2005, Leg: L. Pérez.

Redescription. Mature males. Head. Labium cream with yellow teeth. Prementum light green. Base of mandible green with brown spot at top quarter. Labrum bright green to light green with medio-apical black spot or three apical black spots, one median and two lateral,. Gena bright green to gray. Anteclypeus bright green to gray with two amber spots on ventral half on each side. Postclypeus black. Antefrons dark green to light blue with black median stripe extending to margin with postclypeus and widening dorsally toward postfrons. Postfrons black, postocular spots lilac or light blue reaching or not the eyes. Pair of light green spots behind antenna and brown spot on each side of mediun ocellus present or absent. Antenna black. Rear of head greenish cream.

Thorax. Anterior lobe of prothorax black with light blue lateral apices, middle lobe entirely black or with distal 1/4 light brown, propleura light blue, posterior lobe black with lateral apices cream, trilobed with medial lobe projecting backwards, with a central depression and apical edge concave, more concave and sinuous in specimens from the Cordillera Oriental, length of lateral edge about 3/5 length of apical edge. Mesostigmal plate approximately triangular and flat with blue lateral apex. Pterothorax with black dorsal medial stripe reaching half of mesepisternum at which point a clear blue stripe begins that reaches humeral suture, mesepimeron upper stripe clear coffee, remaining of mesepimeron and metepisternum blue and metepimeron and venter greenish cream. Mesocoxa and procoxa light blue, metacoxa cream ( Fig. 1 c). Remainder of legs black with apex of femur light blue and a light blue stripe near its base, sometimes medial surface of femur completely blue. Seven to ten metafemoral external spurs, as long as space between them or shorter, gradually increasing in size toward apex. Six to nine external metatibial spurs, as long as space between them or shorter, gradually decreasing in size toward apex. Pretarsal claw with well developed supplementary tooth. Wings hyaline with brown pt, ratio of proximal and distal sides of pt approximately 1:1. CuP reaching CuPAA slightly distal to confluence of CuPAA with hind margin of wing, but never for a distance greater than length of CuP. 13–16 Px in FW, 11–13 Px in HW. RP 2 branching between Px 5 and 8 in FW, between Px 5 and 7 in HW ( Fig. 1 c).

Abdomen. Dorsum black, with distal half of S1 blue, distal 3/4 of S7 dark blue, and S8–9 entirely blue. Lateral terga and venter cream, with sides of S1, S2, proximal portion of S3 green, and S10 light green to cream ( Figs. 1 c, 12c–d). Genital ligula (as in Figs. 8 a–b) with apex concave, with latero-apical lobes extended antero-laterally, finishing in small pointed recurved tip, inner medial process arising between lateromedial lobes, latero-medial lobes located at about mid-length of distal segment of ligula, of hemispheric contour in ectal view. Cercus ( Figs.

8c–d) black, about half length of S10 or less, with dorsal process ending in small apical tooth directed ventrally, inner ventroapical process hemispherical, arising between dorsal process and external ventroapical process, representing most distal point of body, external ventroapical process convex and about as long as dorsal process, ventrobasal process extending to about mid-length of paraproct and ending in sharp tip directed proximally. Paraproct entirely black, in specimens from Cordillera Oriental ventral half pale, with convex dorsal process finishing in short spine that reaches about mid-length of cercus. Total length 32–45 mm. Abdomen length 24–37 mm. FW length 20–26 mm. HW length 19–24 mm.

Variation in Cordillera Central populations. Ventral half of paraprocts pale. Pronotum more concave and more sinuous.

Mature females ( Fig. 1 d). Head. As mature males but base of mandible light blue, with small brown spot at upper 1/4. Basal half of labrum blue turning to cream near apex, with three black basal spots, one at center and two lateral. Gena blue turning cream ventrally. Anteclypeus light blue with two amber spots at ventral half on each side. Postclypeus black, in some cases with a clear point on each side of center. Frons may have pruinescence, antefrons as in males but light blue. Postfrons color pattern very variable, entirely black with pale areas represented only by blue postocular spots reaching eye margin or with light spot between ocelli, anterior ocellus with a spot on each side extending to base of antenna, or with a pale green spot between ocelli, antenna base pale blue, and pale spot behind eye margin reaching antenna.

Thorax. Prothorax with anterior lobe black or black with median pale blue stripe, middle lobe black with two brown spots on each side, posterior lobe black with light blue lateral apex, trilobed with median lobe distally projected, approximately flat and with apical edge slightly concave, length of lateral edge about 3/5 of apical edge length ( Fig. 8 e), pleura light blue. Black mesepisternal plate with lateral apex light blue. Pterothorax cream with black mid-dorsal stripe, light blue mesepisternal stripe, brown humeral stripe, and distal 4/5 of metepisternum and mesepimeron blue ( Fig. 1 d). Coxa cream. Inner surface of remainder of leg cream, outer surface black except femur apex cream or blue. Seven to nine external metafemoral spurs. Six to eight external metatibial spurs. 13–15 Px in FW, 12–13 Px in HW. RP 2 branching between Px 6 and 7 in FW, between Px 5 and 6 in HW ( Fig. 1 d).

Abdomen. Dorsum black, S1 distal half blue, S7 distal margin blue, S8 distal third blue, S9 distal half blue, S10 entirely blue. Lateral terga and venter cream with black medio-ventral line, sides of S–2 blue, S3–7 with incomplete apical cream ring ( Figs. 1 d, 13g –h). Well developed dark brown vulvar spine. Sub-basal plate of ovipositor triangular. One row of teeth on ovipositor outer valve. Ovipositor cream, without stylus, reaching about distal margin of paraproct.Stylus black, being most distal point of body. Paraproct and cercus black, cercus conical and slightly shorter than S10. Total length 35–38 mm. Abdomen length 28–31 mm. FW length 23–25 mm. HW length 22–24 mm.

Diagnosis. Males of this species are characterized by cercus with tip of subapical dorsal process and inner ventroapical process rounded ( Figs. 8 c–d), which groups M. risi with M. ovigerum , M. occultum , M. ecuatoriale , M. rosseri , and M. santainense . With the exception of M. ovigerum , the presence of a blue spot covering the distal three quarters of S7 dorsum ( Fig. 12 c–d) separates M. risi from all these species, which lack a blue spot on S7. M. ovigerum has, however, a black stripe between the distal edge of S7 and the blue spot ( Fig. 12 m), lacking in M. risi . The shape of the genital ligula ( Figs. 8 a–b) is also different within this group of species with the exception of M. ovigerum . However, the external ventroapical process in male cercus of M. ovigerum is much longer than in M. risi . The females can be distinguished from females of M. rosseri , M. santainense , M. occultum , M. ovigerum , and M. ecuatoriale by the unique posterior lobe of prothorax, which extends dorsally ( Fig. 8 e), with side edge about 3/ 5 length of the slightly concave apical edge ( Fig. 8 e). The female color pattern of S7–10 is unique to this species ( Fig. 13 g–h). Male and female teneral specimens of M. risi do not exhibit red coloration.

Remarks. Ris (1918) examined specimens of M. risi , which he found agreed overall with Calvert’s description of M. ovigerum (Calvert 1909) , and he assigned them to that species without comparison with Calvert’s type. He mentioned, however, some differences between his specimens and the description of M. ovigerum and provided a brief description of the female. De Marmels (1989, 1990a) reexamined Ris´material, together with additional individuals collected by Fernando Cruz in Colombia, and illustrated their diagnostic traits, but he could not compare them with the type of M. ovigerum . Garrison (De Marmels 1997) later illustrated the caudal appendages of the type of M. ovigerum and sent his illustrations to De Marmels, who then realized that he was dealing with an undescribed species and published these findings (De Marmels 1997). However, at that time De Marmels could not reexamine the material studied by Ris in 1918 (returned to the Senckenberg collection in Frankfurt) or the Colombian material (returned to Fernando Cruz in Bogotá) he had examined earlier (De Marmels 1997). When von Ellenrieder and Garrison (pers. comm.) redefined the genus in 2008 they also tried to borrow and examine the material studied by Ris and De Marmels, but the type specimens were not found and are apparently lost. However, Fernando Cruz (pers. comm.) still keeps his collection at his home in Bogotá ( Colombia) and the paratype of M. risi examined by De Marmels in 1989 could be still extant there.

We examined material collected near the type locality and from various localities from the Cordillera Central and concluded that this is the only species of Mesamphiagrion distributed in both the Central and the Oriental Cordillera.

Distribution. Widely distributed, from Huila to Antioquia in the Cordillera Central, and from Cundinamarca to Santander in the Cordillera Oriental, between 1,000 and 2,300 m.a.s.l. ( Fig. 16 ).

Family Genus Species

Polythoridae Polythore P. gigantea (Selys) Cora C. lugubris Navás Euthore E. fasciata (Hagen in Selys) E. fastigiata (Selys) Calopterygidae Hetaerina H. cruentata ( Rambur) H. aurora

H. capitalis Selys Lestidae Lestes L. apollinaris Navás Megapodagrionidae Philogenia Philogenia sp.*

Teinopodagrion T. mercenarium (Hagen) T. oscillans

T. temporale (Selys)

Coenagrionidae Argia Argia spp. At least three different species

Oreiallagma Oreiallagma sp. nov. O. oreas (Ris)

Homeoura H. chelifera (Selys) Mesamphiagrion M. laterale (Selys) Aeshnidae Rhionaeschna R. marchali

R. cornigera (Brauer) R. psilus

Libellulidae Libellula L.herculea Karsch Cannaphila C.vibex (Hagen)

Sympetrum S.gilvum

Brechmorhoga B. vivax Calvert

B. rapax Calvert

Erythrodiplax E. connata- group Gomphomacromia G. fallax McLachlan Gomphidae Progomphus P. pygmaeus Selys Epigomphus E. pechumani Belle

*Only females have been collected.

Habitat and biology. This species was collected at large dams and marshes, forested or with forest fragments. Adults fly close to the ground or water surface. Males are more often found near wetlands, and females tend to be solitary and can be found far from wetlands. Several pairs were seen flying in tandem on a hot day in July 2009 at around 10:00 hrs.

Other odonate species collected with M. risi are given in Table 4 . Among others they include the rediscovery of Epigomphus pechumani , Oreiallagma oreas , and Cora lugubris , all three species known so far only from their types, and a recently discovered undescribed species of the genus Oreiallagma , as well as Rhionaeschna psilus from the Valle del Aburrá, which constitutes the first record of this species from Colombia.

Figures

FIGURE 1. Mesamphiagrion rosseri (Antioquia, Sonsón): (a) Male holotype; (b) last larval instar resting with half body out of water. M. risi (Antioquia): (c) Male (Amalfi); (d) Female (San Vicente).

FIGURE 8. Mesamphiagrion risi (Antioquia, San Vicente): (a) Ectal view of genital ligula, (b) Lateral view of genital ligula, (c) Lateral view of caudal appendages, (d) Medio-dorsal view of right cercus. Female: (e) Dorsal view of posterior lobe of pronotum. l. p.: Ponotum posterior lobe. l. l-ap.: Lateroapical lobe. p. d.: Dorsal process. p. v-a. e.: Ventroapical external process. p. v-a. i.: Ventroapical inner process. p. v-b.: Ventrobasal process. p. i.: Inner process. Illustrations drawn by Gloria Mora.

FIGURE 12. Mesamphiagrion tamaense (Santander, Suratá): (a) Mature female; (b) Juvenile female; (c) Mature male; (d) Juvenile male.

FIGURE 16. New distributional records of Mesamphiagrion demarmelsi, M. gaudiimontanum, M. ovigerum and M. risi in Colombia.

FIGURE 13. S 7 – 10 color patterns of males: (a) and (b) Mesamphiagrion gaudiimontanum; (c) and (d) M. risi; (e) M. rosseri; (f) and (g) M. santainense; (h) M. tamaense; (i) and (j) M. occultum; (k) M. demarmelsi; (l) M. laterale; (m) M. ovigerum; (n) M. nataliae.