Calopterygoidea, Selys-Longchamps, 1850

Bybee, Seth M., Kalkman, Vincent J., Erickson, Robert J., Frandsen, Paul B., Breinholt, Jesse W., Suvorov, Anton, Dijkstra, Klaas-Douwe B., Cordero-Rivera, Adolfo, Skevington, Jeffrey H., Abbott, John C., Herrera, Melissa Sanchez, Lemmon, Alan R., Lemmon, Emily Moriarty & Ware, Jessica L., 2021, Phylogeny and classification of Odonata using targeted genomics, Molecular Phylogenetics and Evolution 160, pp. 107115-107115 : 9

4.1.5.4. ‘ Calopterygoidea ’ group 4

(BS = 100, PP = 1, QS = 0.37/0.42/ 0.96).

The fourth group contained within the ‘Calopterygoidea’ consists of three pairs of families. Philosinidae (BS = 100, PP = 1, QS = 1/NA/1) includes the Asian genera Philosina and Rhinagrion , which resemble each other strongly in adult morphology and have a clear apomorphy in the nymphal stage in the tube-shaped caudal gills, i.e. the outer gills are folded around the median gill ( Kalkman et al., 2010). The Philogangidae with its single genus Philoganga is its sister group with strong support (BS = 100, PP = 1, QS = 0.92/0/1). Adult Philoganga resembles Philosinidae in general appearance, being relatively large and robust, and resting with wings outstretched. The main difference is in the denser venation, with two antenodal crossveins in Philosinidae , but 11 to 13 in Philogangidae . The nymphs of both families also resemble each other in general build, with long lateral outer caudal gills and a slightly shorter central caudal gill, although the lateral ones are not tube-shaped in Philoganga .

For about a century, Megapodagrionidae served as a ‘dustbin’ family for damselfly genera with unclear relationships. Based on present and recent work ( Dijkstra et al., 2014; Kalkman and Theischinger, 2013), these are now divided across no less than fifteen families. The true Megapodagrionidae are limited to the genera Megapodagrion , Allopodagrion and Teinopodagrion with a total of only 29 species limited to tropical America. The only genus included in our study ( Teinopodagrion ) was found to be sister to Argiolestidae (BS = 100, PP = 1, QS = 0.86/0/ 0.99), a group restricted to the Afrotropics and Australasia. Argiolestidae was until recently considered as a subfamily of Megapodagrionidae , but raised to family level based on the morphology of caudal gills of the nymphs that are distinctively flat and fan-shaped and held in a horizontal plane ( Kalkman and Theischinger 2013). True megapodagrionid caudal gills lie in a vertical plane with the lateral pair triquetral and the median foliaceous ( De Marmels, 1999).

Finally, the well supported monophyletic families Calopterygidae (BS = 100, PP = 1, QS = 1/NA/1) and Chlorocyphidae (BS = 100, PP = 1, QS = 1/NA/1) form the core of a group which is colloquially often addressed as Caloptera. However, Caloptera has poor QS values (BS = 100, PP = 1, QS = -0.25/0.23/1). Nonetheless, all species are restricted to running waters and the majority of males have brightly colored (both metallic and pigmented) bodies and often wings used in wonderfully elaborate courtship displays.