Mesamphiagrion gaudiimontanum Bota-Sierra

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 : 406-409

Mesamphiagrion gaudiimontanum Bota-Sierra sp. nov.

Figs. 3 c–d (habitus ♂), 5a–c (ontogenic changes ♂), 6a–d (ontogenic changes ♀), 7a–f (diagnostic traits), 13a–b (S7– 10 ♂), 14k–l (S7–10 ♀), 16 (map)

Etymology. From Latin gaudium, happiness, and montanum, of the mountains. The name refers to its beauty and conspicuous presence in the highlands where it lives.

Specimens examined (50 specimens: male holotype, allotype, 48 paratypes).

CEUA: Antioquia: 1 ♂, Municipality Bello: Corregimiento San Félix: Township Sabanalarga: Subpáramo Las Baldías: N6°19'53" W 75°38'58" 3100m, 16.vii.2013, Leg: J. R. Albertino & A. L. Montoya. Municipality Belmira: Township Río Arriba: Páramo El Morro, N6°38'59.3" W 75°40'8.8" 3270 m: 1♂, 04.v.2008, Leg: C. Bota & A.L. Montoya. 7 ♂ and 3 ♀, 2–4.iv.2010, Leg: C. Bota. 2 ♂, 15.i.2011, Leg: L. Ríos, A. L. Montoya & J. Cardona. Holotype, Allotype and 3 ♂, 10.v.2011, Leg: C. Bota & J. D. Castaño. Alto del Indio, N6°37'34.4 " W75°42' 22.0" 2950 m: 6 ♀ and 1 ♂, 3.iii.2012; 2 ♂ and 4 ♀, 7–8.vii.2012, Leg: A. Clavijo. Páramo de Sabanas, N6°37'23,3" W75°38'44.5" 3130 m: 5 ♂ and 1 ♀, 1–3.x.2011, Leg: C. Bota & J.D. Castaño. 2 ♂ and 1 ♀, path from the town toward El Morro, 6°37'N 75°40'W 2950 m approx., 16.vii.2012. Leg: C. Bota, C. Moreno, C. Flórez, J. Caly, J. del Río, J. Sierra, K. Mejía & M. Moreno. 2 ♂, Corregimiento Labores, Finca El Paraíso, N 6°40' W 75°38', 5–7.iv.2012, Leg: C. Moreno, S. Bota, T. Bota & C. Bota. Municipality San José de la Montaña, Páramo El Congo, N6°45'37.6" W75°35'24.0" 3200m: 5 ♀ and 3 ♂, 10–14.ix.2011, Leg: L. Ríos & J. Zapata. 1 ♀, 05.xi.2011, Leg: C. Bota.

Description. Holotype. Head. Labium cream along edges becoming light blue toward center. Mandible base blue. Labrum blue with a black small medio-basal spot and dorsolateral black margins. Gena blue. Anteclypeus blue. Postclypeus black. Frons black, with blue postocular spots touching eye margin, with pruinescence, in anterior view area bordering gena and clypeus on each side light blue, divided by a black medial stripe over clypeus. Black antennae. Rear of head dorsally cream and ventrally pale blue ( Figs. 3 c–d, 5b–c).

Thorax. Prothorax black, with anterior lobe of pronotum light blue, propleura light blue. Medial lobe of posterior prothoracic lobe developed into a caudally projected squarish plate with smoothly rounded margins and posterior edge slightly concave, each lateral edge approximately half length of distal edge. Pterothorax black with blue antehumeral stripe, wide blue stripe covering mesepimeron and metepisternum, metepimeron light blue, and venter cream. Coxa light blue. External side of remainder of leg black and medial side light blue with blue rounded spot near base of femur. Tarsi, spurs, and claws black ( Figs. 3 c–d, 5b–c). Six external spurs on right metafemur and seven on left metafemur, as long as space between them or shorter, gradually increasing in size toward apex. Six external spurs on right metatibia and seven on left metatibia, as long as space between them or shorter, gradually decreasing in size toward apex. Tarsal claws with well developed supplementary tooth. Wings hyaline. Pt blue, ratio between distal and proximal length about 1:1. CuP reaching CuPAA slightly distal to confluence of CuPAA with hind margin of wing. Px 12 in FW, Px 9 in right HW, 10 in left HW. RP 2 branching between Px 4 and 5 in FW, between Px 3 and 4 in HW ( Fig. 5 b).

Abdomen. Dorsum black and lateral terga cream except: distal half of S1 lateral terga light blue, S2 lateral terga bluish, S7 dorsum with a blue spot occupying distal 5/6, S3–7 with a proximal incomplete blue ring, S8–9 dorsum blue ( Fig. 3 c–d, 5b–c, 12b). Genital ligula (as in figures 7d–e) with apex concave with a pair of lateroapical lobes with blunt distal portion and proximal portion terminating in a sharp hook, lateromedial lobes rounded in ectal view, and inner medial process as long as lateromedial lobes in lateral view. Cercus ( Figs. 7 a–c) external side black, medial side blue, shorter than S10 length, with elongated dorsal process finishing in hook directed ventrally, this being most distal point of animal, apex of external ventroapical process blunt, inner ventro-apical process rising as concave carina in laterodorsal view, ventrobasal process terminating in blunt point, ventroapical processes located midway between dorsal and ventrobasal processes in posteromedial view. Paraproct light blue with black tip and distally directed cylindrical apical process, which is about half cercus length or slightly longer. Total length 30 mm. Abdomen length 23 mm. FW length 18 mm. HW length 17 mm.

Variations in male paratypes. Mature. Pruinescence on head present or absent. Rear of head cream. Metapleural black stripe continuous or interrupted once or twice. Six to nine external metafemoral spurs. Four to seven external metatibial spurs. 11–13 Px in FW, 10–11 in HW. 11–13 Px in FW, 10–11 Px in HW. RP 2 branching between Px 4 and 6 in FW, between Px 3 and 5 in HW. In one specimen, S7 pale blue dorsal spot extending along distal 2/3 without reaching distal margin ( Fig. 12 a). S10 sometimes with small proximal light blue spot and medial distal tip blue. Paraproct cream, sometimes with process bluish. Apex of ligula more or less markedly concave, in some cases almost straight. Total length 29–31 mm. Abdominal length 22–24 mm. FW length 18–19 mm. HW length 16–18 mm.

Juvenile. Labium cream. Base of mandible cream. Labrum blue. Gena blue. Anteclypeus light blue and postclypeus reddish brown. Frons as in mature males but paler. Prothorax brown. Pterothorax as in mature males but black zones brown. Coxa light brown. Pt red. S1–4 as in mature males but red instead of black. S4 with a distal black spot. S5–6 black dorsally and red laterally ( Fig. 5 a–b).

Allotype (androchrome female). Head. Color pattern as in holotype but paler, postocular spots not reaching eye margin and occipital bar cream ( Fig. 6 b).

Thorax. Coloration as in mature males but dark brown instead of black ( Fig. 6 b). Medial lobe of posterior lobe of pronotum well developed, with lateral margin as long as about 1/3 of posterior margin, its distal edge strongly concave medially. Mesepisternal plate approximately flat and triangular, black with blue apex ( Fig. 7 f). Coxa bluish cream. Remainder of leg as in holotype but brown instead of black and cream instead of blue. Six external spurs on right metafemur and seven on left metafemur. Five external spurs on right metatibia and six on left metatibia. Pt reddish brown. 12 Px in right FW, 13 in left FW, 10 Px in HW. RP 2 branching between Px 5 and 6 in FW, between Px 4 and 5 in HW ( Fig. 6 b).

Abdomen. Dorsum black and venter cream, except for blue lateral spots on S1–2, distal margin of S1 light blue, S2–3 dorsally brown, S4–5 same but with black distal 1/4, S3–7 with incomplete cream ring along proximal margin, S7 with blue spot located on middle of segment dorsum covering approximately half, S8 with light blue spot covering basal 3/4, S9 with longitudinal media blue line running from center to distal margin. S10 with median blue stripe widening distally ( Figs. 6 b, 13k). Black line from S2 proximal margin to vulvar spine, vulvar spine black. Ovipositor, cercus, and paraproct cream. Subbasal plate of ovipositor triangular. Ovipositor apex reaching level of apex of paraproct, stylus reaching apex of cercus. Cercus brown, conical, slightly longer than half of S10. Paraproct creamy blue, truncate posteriorly. Total length 30 mm. Abdomen length 24 mm. FW length 19 mm. HW length 18 mm.

Variation in mature female androchrome paratypes. Five to eight metafemoral external spurs. Five to eight metatibial external spurs. 10–14 Px in FW, 9–12 Px in HW. RP 2 branching between the Px 4 and 6 in FWs, in HWs between Px 3 and 5. Total length 28–32 mm. Abdomen length 21–25 mm. FW 18–20 mm. HW length 16–19 mm.

Variation in female gynochrome paratypes. Head. As in androchrome females ( Fig. 6 c–d) but labium cream, sometimes blue toward center. Base of mandibles light yellow, becoming yellow toward apex. Labrum brown, with small brown to black medio-basal spot. Gena yellow. Anteclypeus bluish gray to gray. Postclypeus dark brown with or without pruinescence. Frons with or without pruinescence, in anterior view with area above clypeus black, in dorsal view brown, occipital edge cream, ocelli surrounded by black rings, postocular spots iridescent green or absent.

Thorax. Prothorax dark brown with gray propleura, with or without pruinescence. Pterothorax as in holotype but brown instead of black and green instead of blue, becoming darker with age and looking completely brown when covered with a thick layer of pruinescence ( Fig. 6 c–d). Coxa brown with or without pruinescence. Remainder of leg brown with black spines or brown with femur medial surface gray. Pt red or brown to bluish.

Abdomen. Dorsum black and sides cream, except proximal 3/4 of S3–4 and S1–2 which are brown on mature females and red on juvenile females ( Figs. 6 c–d, 13l). Juveniles with small blue spot on dorsum of S10 and sometimes also on S9.

Diagnosis. Although in juvenile specimens the pt is red, in mature males it is blue and distinguishes this species from all other males of the genus. The cercus dorsal process is longer than ventrobasal external process, a character state that ( Figs 7 a, c) groups this species with M. tamaense , M. nataliae , M. gaianii , M. laterale , M. demarmelsi , and M. dunklei . The elongate ventrobasal process ending in a blunt apex ( Figs. 7 b–c) separates it from M. laterale , M. tamaense , M. nataliae , and M. gaianii , being similar to M. dunklei . The cercus of M. gaianii in posterior view is also narrower than the cercus of M. gaudiimontanum , in which the internal carina is more developed ( Fig. 7 b). The wider medial lobe of posterior prothoracic lobe separates M. gaudiimontanum from M. demarmelsi . Shape of genital ligula ( Figs. 7 d–e) clearly separates it from M. nataliae , M. tamaense , M. laterale , and M. dunklei but resembles M. gaianii . Females are distinguished by the medial lobe of posterior lobe of pronotum well developed, with lateral margin as long as about 1/3 of posterior margin and distal edge strongly concave medially ( Fig. 7 f). Androchrome females differ further from females of M. gaianii ( Fig. 13 r), M. demarmelsi ( Fig. 13s ), and some females of M. laterale ( Fig. 13 o–q) by having a blue spot on S7 ( Figs. 6 a–b, 13k), and from M. nataliae ( Fig. 13 a–c) and M. dunklei by presenting different shaped spots on S7–10 ( Figs. 6 a–b, 13k). Juveniles of M. laterale , M. nataliae , M. tamaense , and M. gaianii have red coloration only on S1–3, whereas in M. gaudiimontanum the red coloration extends through S1–6 ( Figs. 5 a, 6a).

Remarks. This species is difficult to diagnose because of its similarity to M. gaianii . R. W. Garrison (pers. comm.) suggested, mainly based on diagnostic traits of the males ( Figs. 7 a–d), that populations of this species found in the northern Cordillera Central could correspond to a population of M. gaianii with certain variations. However, we found subtle differences in these characters that are preserved in the Santa Inés populations, as well as strong differences in the females, which lead us to propose the hypothesis that populations present in this part of Colombia are a lineage whose evolutionary history is sufficiently different from M. gaianii as to provide reproductive isolation, not only due to geographical and ecological separation but also to variation of several morphological traits directly involved in mating, and therefore they can be considered different species.

Color polymorphism was observed in females of this species ( Figs 6 a–d), with gynochrome and androchrome individuals found at a ratio of approximately 3:1 ( Table 1).

Androchrome teneral females Gynochrome teneral females Androchrome mature females Gynochrome mature females 3 7 4 10

Habitat and biology. Adults inhabit páramos (high mountain wet tropical ecosystems unique to the Neotropics) and montane forest edges dominated by the oak Quercus humboldti . Mature males were found near peatlands or lakes, immatures and females from the edge of the oak forest to open areas. Larvae were collected in ponds, peatlands, and small pools with Sphagnum . Larvae in captivity were observed using Sphagnum for camouflage.

Mature males were found close to peatlands perched on vegetation and Sphagnum , usually in great numbers. Fights between males were common at sunny times, sometimes, after which males perched with their wings open at about 45 degrees ( Fig. 5 b), while sometimes bobbing their abdomen up to the level of thorax and down again repeatedly, a behavior also observed in M. tamaense . Machado (2012) suggested blue pterostigmata could have a territorial defense function, nevertheless field studies will need to be done in order to determine if this hypothesis has merit (Corbet 1999). Some fights between males were observed, in which they chased each other until one or both left the area. It was also common to find two males on the same or a nearby perch without fighting. Females were observed both far from and near water.

Alejandra Clavijo (pers. comm.) observed some pairs in copula, around noon, in sunny weather in July 2012 near a small stream with Sphagnum ; all females were gynochrome. On one occasion, CAB observed a male trying to capture a gynochrome female on a sunny morning, both perched on Sphagnum ; the male was performing short flights toward the female, and she was avoiding him until finally the female flew away. An androchrome female with wings quite deteriorated was collected with S7–10 clearly muddy, which indicated it was laying eggs.

Specimens perched on vegetation with rosette-like leaves, including frailejon ( Espeletia sp.), or hedgehog-like leaves as some grasses. When disturbed, individuals often dropped into the center of the plant or tried to hind behind a leaf (also observed on M. laterale and M. tamaense ). Some adults were parasitized by mites ( Figs. 5 c, 6c).

This species co-occurs with the odonate species listed in Table 2, plus M. santainense ; M. santainense has not been recorded at Páramo ecosystems. Other odonates collected here were new species of Ischnura , Oxyallagma , and Rhionaeschna , the descriptions of which are in preparation by the authors. Oxyallagma has thus far been recorded only from Peru and Ecuador (Garrison et al. 2010) and Sympetrum paramo thus far only from Venezuela.

Distribution. Known only from the mountain range of Santa Inés, and Las Badías in the northwestern Cordillera Central, Department of Antioquia, between 2,700 and 3,270 m.a.s.l. ( Fig. 16 ).

Family Genus Species

Libellulidae Erythrodiplax E. cf. cauca

Sympetrum S. gilvum (Selys)

S. paramo DeMarmels *

Megapodagrionidae Teinopodagrion T. oscillans (Selys)

Coenagrionidae Oxyallagma Oxyallagma sp. nov. Ischnura Ischnura sp. nov.

Aeshnidae Rhionaeschna R. brevicercia (Muzón & von Ellenrieder)

R. marchali

Rhionaeschna sp. nov. *

*Not collected with M. santainense ( Páramo inhabitans).

Figures

FIGURE 3. Mesamphiagrion santainense (Antioquia, Belmira): (a) Male holotype; (b) Female paratype. M. gaudiimontanum (Antioquia, Belmira) Mature males: (c) Dorsal view; (d) Lateral view.

FIGURE 5. Ontogenic changes in Mesamphiagrion gaudiimontanum males (Antioquia, Belmira): (a) Teneral; (b) Mature; (c) Middle age parasitized by mites (Photograph by Camilo Flórez-V.).

FIGURE 6. Ontogenic changes in both morphs of Mesamphiagrion gaudiimontanum females (Antioquia, Belmira): Androchrome: (a) Juvenile; (b) Mature. Gynochrome: (c) Juvenile parasitized by mites (Photograph by Cintia Moreno); (d) Mature.

FIGURE 7. Mesamphiagrion gaianii (Paratype, Venezuela, Trujillo, Páramo La Cristalina) vs. M. gaudiimontanum (Antioquia, Belmira) (traits of M. gaianii on left): (a) Lateral view of caudal appendages; (b) Posterior view of left cercus; (c) Medio-dorsal view of cercus; (d) Ectal view of genital ligula; (e) M. gaudiimontanum lateral view of genital ligula; Female paratype of M. gaudiimontanum: (f) dorsal view of posterior lobe of pronotum and mesostigmal plates. p. m.: Mesostigmal plates. l. p.: Ponotum posterior lobe. l. l-ap.: Lateroapical lobe. l. l-m.: Lateromedial 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: Rosser Garrison (a to d), Gloria Mora (e), and Natalia Uribe (f).

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.