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 : 8-9

4.1.5.3. ‘ Calopterygoidea ’ group 3

(BS = 100, PP = 1, QS = 0.48/0.59/ 0.96).

This group contains nine different families with relatively little in common morphologically. Most striking is the wide variety of shapes of the nymphal caudal gills: flat and fanlike in Mesopodagrionidae , balloon-shaped in Lestoideidae and Thaumatoneuridae , balloon-shaped with lateral abdominal gills on abdominal segments two to eight in Euphaeidae , balloon-shaped with filamentous gill tufts in Pseudolestidae , sturdy and pyramidal with the epiproct terminating in three points and filamentous gill tufts below them in Devadattidae , and roundish and gradually tapering to a single point in both paraprocts and epiproct with filamentous gills tufts below them in Amphipterygidae . This suggests that there has been strong selection on the nymphal respiratory system in these groups, although nothing is known of the relative advantages of the different shapes of gills in their lotic habitats.

The genera Amanipodagrion and Mesopodagrion form a clade that is the sister group to all taxa in Group 3 that were previously placed as incertae sedis ( Bybee et al., 2008; Davis et al., 2011; Dijkstra et al., 2014; Dumont et al., 2010; van Tol and Reijnen, 2009). Amanipodagrion is monotypic with the only known species, A. gilliesi , being from Tanzania. The species is confined to a single rocky forest stream in the East Usambara Mountains where the nymph is yet to be discovered. It is a relatively large species with uncertain taxonomic affinities due to its overall morphology, banded wings and a habit of resting in a hanging position that does not match other species. Mesopodagrion is known from two species found in China and the northern regions of Vietnam, Thailand and Myanmar. The two Mesopodagrion species possess a combination of characters that does not fit any other genus: an apomorphy is the distinct extension of the terminal rim of the 10th tergite between the cerci ( Yu and Bu, 2009). The nymphs have flat horizontal caudal gills ( Yu, 2016), which are otherwise only found in the unrelated Argiolestidae and Protolestidae ( Kalkman et al. 2010). The unique character set of both adults and nymphs, combined with the molecular results, lead us to establish a new family to accommodate this genus (see below Familylevel revisions to the Classification of Zygoptera ). In our analyses Amanipodagrion is the sister to Mesopodagrion but with low QS values (BS = 100, PP = 1, QS = 0.48/0.59/0.96). The two genera are clearly different in morphology and behaviour and cannot be considered members of the same family. We therefore place them in their own respective families: Amanipodagrionidae and Mesopodagrionidae (see below Family-level revisions to the Classification of Zygoptera ).

Bybee et al. (2008) first established the sister-group relationship between Euphaeidae and Lestoideidae (BS = 100, PP = 1, QS = 0.8/0/ 0.99), which was supported with more extensive taxon sampling by Dijkstra et al. (2014). Sister to these two families is Pseudolestidae (BS = 100, PP = 1, QS = 0.57/0.62/0.97) with a single known species Pseudolestes mirabilis . Being known only from Hainan, P. mirabilis , has a very distinct adult and nymphal morphology, for example having the hindwing much shorter than the forewing, and unique behavior (Cordero-Rivera and Zhang, 2018 a, 2018 b; Yu and Bu, 2011). The Oriental Devadattidae and Mesoamerican Amphipterygidae are sister taxa (BS = 100, PP = 1, QS = 0.81/0/1). Their nymphs share the filamentous gill tufts below the caudal gills, also a trait of Pseudolestidae , but other morphological differences in the adults and nymphs make them distinct enough to keep them in their respective families. The last two families of this group are the Mesoamerican Thaumatoneuridae (BS = 100, PP = 1, QS = 1/NA/1) and the Oriental Rhipidolestidae (BS = 100, PP = 1, QS = 1/NA/0.98). The latter includes four genera ( Agriomorpha , Burmargiolestes , Bornargiolestes and Rhipidolestes ), which Dijkstra et al. (2014) regarded as incertae sedis as they were found to be paraphyletic when Thaumatoneuridae was not included. Agriomorpha and Rhipidolestes form a monophyletic group in our analyses and as Burmargiolestes and Bornargiolestes are closely related to Agriomorpha , we assume the four to form a monophyletic group. The name Rhipidolestinae was first used by Silsby (2001) although it seems that she did so accidentally, using it for a group that included Pseudolestes for which the name Pseudolestidae was already available. Nonetheless we propose to consider Silsby (2001) as the author for this family as her description, while brief, complies with the code of zoological nomenclature, including a citation of the name of the type genus, Rhipidolestes . The name is here used for the group of four mentioned genera, although Rhipidolestes stands apart due to different venation and a sturdy dorsal spine on the male’ s ninth abdominal segment. Further work might therefore show that the family should better be divided into two separate subfamilies or even families.