The larva of Hetaerina duplex Selys, 1853 (Odonata: Calopterygidae) with a key to known Colombian larvae species

Montes-Fontalvo & Stand-Pérez The larva of Hetaerina duplex with a key to known Colombian larvae species 16 International Journal of Odonatology │ Volume 28 │ pp. 16–22 International Journal of Odonatology 2025, Vol. 28, pp. 16–22 doi:10.48156/1388.2025.1917310 Research Article OPEN ACCESS This article is distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Published: 27 March 2025 Received: 4 February 2025 Accepted: 17 March 2025 Citation: Montes-Fontalvo & Stand-Pérez (2025): The larva of Hetaerina duplex Selys, 1853 (Odonata: Calopterygidae) with a key to known Colombian larvae species. International Journal of Odonatology, 28, 16–22 doi:10.48156/1388.2025.1917310 Data Availability Statement: All relevant data are within the paper. The larva of Hetaerina duplex Selys, 1853 (Odonata: Calopterygidae) with a key to known Colombian larvae species Jenilee Montes-Fontalvo a & Miguel Stand-Pérez b a Sección Entomología, Colecciones Biológicas, Centro Colecciones y Gestión de Especies, Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Villa de Leyva, Boyacá, Colombia b Red de Biología Evolutiva, Instituto de Ecología A.C., Xalapa, Veracruz, México *Corresponding author. Email: jmon[email protected]g.co Abstract. The larva of Hetaerina duplex is described and illustrated for the first time based on specimens from Quindío and Risaralda departments of Colombia. Larvae of H. duplex differ from other species of the genus by well-developed spines on the lateral carina from S2 to S10 and S10 with one well-developed mid-dorsal spine on the posterior border. In addition, habitat and habits information are commented on. Finally, a comparative key for known larvae of all Colombian species is presented with their distribution maps. Keywords. Zygoptera, Hetaerininae, Colombia, damselfly, neotropical, nymph, rubyspot, taxonomy Introduction Like most insects, damselflies spend most of their lives as larvae, and consequently, much ecological research is still impossible unless the larvae are matched with the adult stages (Zloty et al., 1993). Knowing the larval stage is crucial not only for understanding their ecology, but also for assessing the environments in which they develop. Odonates, which remain in their larval phase in aquatic environments, are particularly effective as bioindicators, as these insects occupy the interface between aquatic and terrestrial ecosystems and are highly sensitive to environmental changes (Miguel et al., 2017; Oliveira Junior et al., 2015). In countries like Colombia, the immature stages of damselflies are often considered by environmental consultants when assessing freshwater quality. However, the level of knowledge regarding the immature stages of odonates needs improvement, with nearly 60% of larvae remaining undescribed (Bota-Sierra et al., 2024). Therefore, it is essential to intensify research on odonate larvae and develop taxonomic tools to aid in identifying more specific taxonomic levels. The New World genus Hetaerina comprises 37 valid species, 13 (and two subspecies) of which occur in Colombia (Bota-Sierra et al., 2024). Of these, only seven have published larval descriptions (Table 1), the latest dating back to Zloty et al. (1993). In this study, we describe and compare the final instar larva of Het aerina duplex Selys, 1853, a species recently synonymized with H. aurora Ris, 1918 due to characteristics such as significant variability in wing tip coloration and similarity in caudal appendages (Garrison & Von Ellenrieder, 2024) and also, provide distribution maps and a key to separate all Colombian Hetaerina larvae known to date. Montes-Fontalvo & Stand-Pérez The larva of Hetaerina duplex with a key to known Colombian larvae species 17 International Journal of Odonatology │ Volume 28 │ pp. 16–22 Materials and methods The conspecifity of the larva and the adult is confirmed thanks to the fact that the exuvia was collected just after the emergence of the adult (Figure 1). Exuviae and final instar larvae were preserved in 70% alcohol. Drawings were made with the aid of photos taken with a Leica MC120HD camera integrated to a Leica S8APO stereoscope. Mandibular formulae terminology follows Watson (1956). Abdominal segments are given as S1–10; all measurements are given in millimeters (mm). Maps were made using QGIS Geographic Information System, including locality data taken from Bota-Sierra et al. (2024) for all Hetaerina species in Colombia and the localities provided in the material examined section. Figure 1. Hetaerina duplex (a) tandem, (b) female teneral and exuvia in situ. Photos: J. Montes. Montes-Fontalvo & Stand-Pérez The larva of Hetaerina duplex with a key to known Colombian larvae species 18 International Journal of Odonatology │ Volume 28 │ pp. 16–22 Table 1. Hetaerina species present in Colombia, its conservation status and distribution. LC = Least Concern; Am = Amazonas; An = Antioquia; At = Atlántico; By = Boyacá; Cau = Cauca; Ch = Chocó; Cl = Caldas; Cq = Caquetá; Cs = Casanare; Cu = Cundinamarca; Gu = Guainía; Gv = Guaviare; Gj = La Guajira; Hu = Huila; Ma = Magdalena; Me = Meta; Na = Nariño; Ns = Norte de Santander; Pu = Putumayo; Qu = Quindío; Ri = Risaralda; St = Santander; To = Tolima; Va = Vaupés; Vl = Valle del Cauca; *Information about the two reported subspecies H. caja caja (Drury, 1773) and H. caja dominula Hagen, 1853, is included. **The distribution of this species is reported in Garrison, 1990, and the specific localities for Colombia are not described. Species IUCN Status Larva description Distribution in Colombia Hetaerina amazonica Sjöstedt, 1918 LC No Am, Gu, Va Hetaerina caja* (Drury, 1773) LC Geijskes, 1943 An, Ch, Cl, Cs, Gv, Gj, Hu, Ma, Me, Na, St, To, Va, Vl Hetaerina capitalis Selys, 1873 LC De Marmels, 1985 An, By, Cau, Ch, Cl, Cu, Gj, Hu, Ma, Na, Ns, Qu, Ri, St, To, Vl Hetaerina charca Calvert, 1909 LC No Cq, Gv Hetaerina cruentata (Rambur, 1842) LC Zloty et al., 1993 An, By, Cau, Ch, Cl, Cu, Gj, Hu, Ma, Na, Ns, Qu, Ri, St, To, Vl Hetaerina duplex Selys, 1869 LC This study An, By, Cu, Cl, Hu, Me, Na, Qu, Ri, Vl Hetaerina fuscoguttata Selys, 1878 LC Zloty et al., 1993 Ch, Vl Hetaerina laesa Hagen, 1853 LC No Am Hetaerina miniata Selys, 1879 LC Zloty et al., 1993 An, Ch, Cl, St, To Hetaerina occisa Hagen, 1853 LC Zloty et al., 1993 An, At, By, Cl, Cau, Ch, Cs, Cu, Gj, Hu, Ma, Me, Ns, Pu, Ri, To, Vl Hetaerina sanguinea Selys, 1853 LC No Am, Cq, Gu, Gv, Me, Pu Hetaerina sempronia Hagen, 1853** LC Zloty et al., 1993 NA Hetaerina westfalli Rácenis, 1968 LC No Cs, Gu, Gv, Me, Va Figure 2. Details of morphology of Hetaerina duplex. (a) Final instar larva in dorsal view; (b) labium; (c) left mandible; (d) right mandible, both mesal view; (e) S8–10 and caudal gills, dorsal view; (f) S8–10 and caudal gills, ventral view; (g) S8–10 showing the ovipositor, lateral view. Montes-Fontalvo & Stand-Pérez The larva of Hetaerina duplex with a key to known Colombian larvae species 19 International Journal of Odonatology │ Volume 28 │ pp. 16–22 Results Hetaerina duplex Selys, 1853 (Figure 2) Material examined Colombia, 1♀, 9 x 2012, Quindío Department, Filandia Municipality, Barbas Bremen Nature Reserve, 4.69694444°N, -75.65472222°W, 1650 m.a.s.l., Leg: J. Montes, L. Perez; 1♀, 13 v 2022, Risaralda Department, Pueblo Rico Municipality, Monte Bello Township, Tatamá National Park, 5.2357126°N, -76.0871193°W, M. Stand leg. Description of final instar larva Coloration of head and thorax mainly brownish with dark brownish dots, legs yellowish with dark brown dots. Head. Brownish with dark brown pattern (Figure 2a), with prominent and rounded posterolateral angles; antenna without setae. Labium: prementum-postmentum articulation reaching the anterior margin of mesocoxae; prementum with two setae on internal side of each lobe of ligula, external margins of ligular lobes serrate; labial palp ending in three well-developed and Figure 3. Distribution of Hetaerina species in Colombia, part I. Montes-Fontalvo & Stand-Pérez The larva of Hetaerina duplex with a key to known Colombian larvae species 20 International Journal of Odonatology │ Volume 28 │ pp. 16–22 incurved hooks forming an 120° angle with it, median hook longest, inner one smallest; movable hook shorter than palp; inner margin of palp with small spines; four short apical palpal setae nearouter margin as in Figure 2b. Mandibles formula: L 12345 0 a(m1234567)b/R 12345 y ab (Figure 2c–d). Thorax. Anterior margin of pronotum widely concave, with one prominent midlateral projection on each side pointing forward, posterior margin convex at middle; wing sheaths extending to posterior margin of S3; Legs: pale with three dark bands on femur, three protibial dark and faded bands; no setae or spines on femur and Figure 4. Distribution of Hetaerina species in Colombia, part II. tibia. Abdomen: Lateral margins of S2–10 carinate, with well-developed posterolateral spine; posterolateral spine of S10 with five small spinules (Figure 2a); posterior margin of S10 with one mid-dorsal spine, and a group of four small spines each side of postero-ventral border (Figure 2e–f). Female ovipositor not reaching distal margin of S10 (Figure 2g) and, in ventral view, with three ventral rows of spines (Figure 2f); cercus conical and straight, 0.5 the length of S10. Caudal gills: lateral gills triquetral and lanceolate, with spines along lateral margins; central gill foliaceous, rounded at tip, with spines along dorsal and ventral margins about 0.7 the length of lateral gills. Montes-Fontalvo & Stand-Pérez The larva of Hetaerina duplex with a key to known Colombian larvae species 21 International Journal of Odonatology │ Volume 28 │ pp. 16–22 Measurements (mm) Total body length (without caudal gills): 23.1. Head: maximum length: 2.8, maximum width: 4; prementum maximum length: 4.2, prementum maximum width: 2.9; cleft length: 1.7; movable hook: 1.3. Thorax and legs: inner wing pads length: 6, outer wing pads length: 5.9; femur I: 4.5; femur II: 5.8; femur III: 6.8; tibia I: 4; tibia II: 5.1; tibia III: 5.9. Abdomen: total length: 13.7; S1 length: 1.7; S2 length: 1.7; S3 length: 1.4; S4 length: 1.7; S5 length: 1.6; S6 length: 1.2; S7 length: 1.4; S8 length: 1.2; S9 length: 1; S10 length: 0.8; cerci length: 0.4; female ovipositor length: 1.8; lateral caudal gills length: 6.7, median caudal gill length: 4.9. Diagnosis Hetaerina duplex occurs in Colombia in the Quindío and Risaralda departments (Figure 3) and larvae for the present description were collected in fast-flowing streams, where they were found partially submerged among grassy vegetation. In Tatamá National Park, where one specimen was collected, H. duplex is sympatric with H. capitalis and shares the following characters: lateral pronotal protuberance rounded, posterior border of S8–9 without row of spines on the dorsal surface, and two premental setae on each side of cleft, but differs from H. capitalis in: lateral well-developed spine on each side of the lateral carina of S2–10 (one row of small spines in the lateral carina of S8–9 in H. capitalis) and S10 with one well-developed mid-dorsal spine on the posterior border (S10 with three small spines in H. capitalis). Remarks Like other Hetaerina species inhabit lotic evironments, with or without abundant riparian vegetation (Guillermo-Ferreira & Bispo, 2012), Hetaerina duplex is recorded in streams and small rivers with abundant vegetation, it has been found in sympatry with H. capitalis and H. cruentata (Bota-Sierra et al., 2024), interestingly, recent studies show that the surrounding vegetation determines the presence of the wing rubyspot (Stranding et al., 2022), the females have been seen ovipositing in dead wood inside river ponds (Bota-Sierra et al., 2019). Its distribution is limited to Colombia and Ecuador, and its conservation status, according to the International Union for Conservation of Nature (IUCN), is classified as Least Concern (Bota-Sierra & Palacino, 2016; BotaSierra & Rache, 2016). Key to the Hetaerina larva known of Colombia This key follows Zloty et al. (1993), but was modified to include only Colombian species (Figures 3–4): 1. Two pairs of premental setae ................................. 2 1’. One pair of premental setae .................................. 5 2. Posterior borders of S8–9 with a row of spines ...... 3 2’. Posterior borders of S8–9 without rows of spines . 4 3. Lateral pronotal protuberances finger-like . H. miniata 3’. Lateral pronotal protuberances lacking . H. sempronia 4. Lateral margins of S2–10 with a posterolateral, well-developed spine; S10 with one mid-dorsal spine on the posterior border .................. H. duplex 4’. Lateral margins of S2–7 without posterolateral spine; S10 with three small mid-dorsal spines on the posterior border ............................... H. capitalis 5. Middorsal abdominal spines on S1–9 or S2–9 ....... 6 5’. Middorsal abdominal spines lacking ...................... 7 6. Postero-lateral spines on abdominal segments 8–10; length of first antennal segment about equal to head´s width ...................... H. fuscoguttata 6’. Postero-lateral spines on abdominal segments 9–10; length of first antennal segment about 70% of head´s width ............................. H. cruentata 7. Anterior border of the first femora with a few, short, thorn-like spines .................................. H. caja 7’. Anterior border of the first femora without conspicuous spines .................................... H. occisa Acknowledgments We thank Andrea Ballen Guapacha for her support in illustrating the larval structures of Hetaerina duplex. We are also grateful to Juan David Palencia for creating the distribution maps of all Hetaerina species in Colombia, and to Professor Rodolfo Novelo for his valuable comments on the manuscript. References Bota-Sierra, C. A., Cordero-Rivera, A., Novelo-Gutiérrez, R., SánchezHerrera, M., Londoño, G. A. (2024). Can high temperatures affect body size in insects? The case of rubyspot damselflies in the Colombian Western Andes. Diversity, 16, 743. doi:10.3390/ d16120743 Bota-Sierra, C., Álvarez-Álvarez, K., Amaya, V., Carrillo-Camargo, B., Garzón-Salamanca, L., Hoyos, A., Mendoza-Penagos, C., Montes-Fontalvo, J., Palacino-Rodríguez, F., Pérez-Gutiérrez, L., Realpe, E., Sánchez Herrera, M., Sandoval-H, J., StandPérez, M., Torres-Pachón, M., Velásquez M. I. & Cano-Cobos, Y. (2024). Commented checklist of the Odonata from Colombia. International Journal of Odonatology, 27, 103–150. doi:10.48156/1388.2024.1917280 Bota-Sierra, C., Sandoval-H, J., Ayala-Sánchez, D. & Novelo-Gutiérrez, R. (2019). Libélulas de la Cordillera Occidental Colombiana, una mirada desde el Tatamá. Panamericana Formas e Impresos, Bogotá, 198 pp. Bota-Sierra, C. & Palacino, F. (2016). Hetaerina aurora. The IUCN Red List of Threatened Species 2016: e.T49254606A49254949. Retrieved Ocotber, 23, 2024, from doi:10.2305/IUCN.UK.2016-1. RLTS.T49254606A49254949.en Bota-Sierra, C. & Rache, L. (2016). Hetaerina duplex. The IUCN Red List of Threatened Species 2016: e.T49254256A49254969. Retrieved Ocotber, 23, 2024, from doi:10.2305/IUCN.UK.2016-1. RLTS.T49254256A49254969.en Montes-Fontalvo & Stand-Pérez The larva of Hetaerina duplex with a key to known Colombian larvae species 22 International Journal of Odonatology │ Volume 28 │ pp. 16–22 De Marmels, J. (1985). On the true Hetaerina capitalis Selys, 1873 and its sibling species Hetaerina smaragdalis spec. nov. (Zygoptera: Calopterygidae). Odonatologica, 14, 177–190. Garrison, R. W. (1990). A synopsis of the genus Hetaerina with descriptions of four new species (Odonata: Calopterygidae). Transactions of the American Entomological Society, 116, 175–259. Garrison, R. W & von Ellenrieder, N. (2024). Taxonomic revisions of some Neotropical Zygoptera (Odonata). Zootaxa, 5405(1), 001– 042. doi:10.11646/zootaxa.5405.1.1 Geijskes, D. C. (1943). Notes on Odonata of Surinam. IV. Nine new or little known zygopterous nymphs from the inland waters. Annals of the Entomological Society of America, 36, 165–183. doi:10.1093/aesa/36.2.165 Guillermo-Ferreira, R, & Bispo, P. (2012). Description of the larva of Mnesarete pudica (Hagen in Selys, 1853) (Odonata: Calopterygidae) and notes on known genera of South American Calopterygidae larvae. Zootaxa, 3482, 77–81. doi:10.11646/zootaxa.3482.1.5 Miguel, T. B., Oliveira-Junior, J. M. B., Ligeiro, R. & Juen, L. (2017). Odonata (Insecta) as a tool for the biomonitoring of environmental quality. Ecological Indicators, 81, 555–566. doi:10.1016/j. ecolind.2017.06.010 Oliveira Junior, J. M. B., Shimano, Y., Gardner, T. A., Hughes, R. M., Marco Júnior, P. & Juen, L. (2015). Neotropical dragonflies (Insecta: Odonata) as indicators of ecological condition of small streams in the eastern Amazon. Austral Ecology, 40(6), 733–744. doi:10.1111/aec.12242 Selys Longchamps, E. de (1853) Synopsis des Caloptérygines. Bulletin de l’Académie Royale des sciences, des lettres et des beauxarts de Belgique, 20 (Annexe), 1‒73 Standring, S., Sánchez-Herrera, M., Guillermo-Ferreira, R., Ware, J. L., Vega-Sánchez, Y. M., Clement, R., Drury, J. P., Grether, G. F., González-Rodríguez, A., Mendoza-Cuenca, L., Bota-Sierra, C. & Bybee, S. (2022). Evolution and biogeographic history of rubyspot damselflies (Hetaerininae: Calopterygidae: Odonata). Diversity, 14, 757. doi:10.3390/d14090757 Watson, M. C. (1956). The utilization of mandibular armature in taxonomic studies of anisopterous nymphs. Transactions of the American Entomological Society, 81, 155–205. Zloty, J., Pritchard, G. & Esquivel, C. (1993). Larvae of the Costa Rican Hetaerina (Odonata: Calopterygidae) with comments on distribution. Systematic Entomology, 18, 253–265. doi:10.1111/j.1365-3113.1993.tb00665.x