Phylogeny and classification of Odonata using targeted genomics
Author
Bybee, Seth M.
Author
Kalkman, Vincent J.
Author
Erickson, Robert J.
Author
Frandsen, Paul B.
Author
Breinholt, Jesse W.
Author
Suvorov, Anton
Author
Dijkstra, Klaas-Douwe B.
Author
Cordero-Rivera, Adolfo
Author
Skevington, Jeffrey H.
Author
Abbott, John C.
Author
Herrera, Melissa Sanchez
Author
Lemmon, Alan R.
Author
Lemmon, Emily Moriarty
Author
Ware, Jessica L.
text
Molecular Phylogenetics and Evolution
2021
2021-02-18
160
107115
107115
journal article
10.1016/j.ympev.2021.107115
6643504
4.1.12.
Libelluloidea
(BS
=
100, PP
=
1, QS
=
0.81/0/1)
Libelluloidea
traditionally includes
Synthemistidae
,
Macromiidae
,
Corduliidae
and
Libellulidae (
Fraser, 1957
)
, and is extremely species rich (~1,500 species). We recover this as a well-supported monophyletic group.
Tillyard described
Synthemistidae
, which were united by synapomorphies both in the nymphal stage (e.g., premental shape, antennal segment number, tibial morphology) and adult stage (e.g., wing venation, male secondary genitalic morphology). However, the family was considered to be variable. Molecular phylogenies (
Letsch et al., 2016
ab;
Letsch, 2007
;
Ware
et al., 2007
) have suggested that the family should be split, with
Archaeophya
,
Gomphomacromia
,
Synthemis
,
Synthemiopsis
,
Eusynthemis
often recovered as a clade, with
Austrocordulia
,
Micromidia
,
Lathrocordulia
,
Macromidia
,
Cordulephya
recovered either as a paraphyletic group or a separate clade. Here, we recover a fully supported grouping of
Gomphomacromia
,
Eusynthemis
,
Choristhemis
and
Parasynthemis
, as clade also recovered by (
Letsch, 2007
) and (
Letsch et al., 2016a
,b).
Cordulephya
,
Micromidia
and
Austrocordulia
are also recovered as a monophyletic clade (BS = 100, PP = 1, QS = 1/NA/1), sister to the remaining libelluloids. Our findings strongly support the idea of
Synthemistidae
as a complex group deserving of extensive data collection, but are unclear regarding the number and arrangement of taxa in this group; hence we await further taxon sampling before naming additional clades in this complex.
We recover
Epophthalmia
,
Macromia
and
Phyllomacromia
in a fully supported group (BS = 100, PP = 1, QS = 1/NA/1).
Epophthalmia
,
Didymops
,
Macromia
and
Phyllomacromia
have routinely been recovered as a monophyletic
Macromiidae
, first by
Gloyd (1959)
, and again by several molecular studies (
Bybee et al., 2008
;
Carle et al., 2015
;
Dumont et al., 2010
;
Letsch et al., 2016
ab;
Letsch, 2007
;
Ware
et al., 2007
, in prep.). Nymphal synapomorphies include the shape and dentition of the prementum, the relative length of the hind legs to body size (giving them a “spider-like” appearance), and a frontal horn between their eyes; adult synapomorphies include the shape and size of the anal loop in the hindwing, eyes with a small protuberance on the lateral edge, and secondary penile characters.
Macromiidae
adults are fast fliers known to patrol long stretches of mainly lotic habitats.
Phyllomacromia
and
Epophthalmia
are recovered as sister taxa; this relationship was suggested by
May (1997)
based on male penile morphology.
Even with a limited sampling of corduliids we demonstrate the complexity of the phylogenetic relationships of this group. Nymphs of the family
Corduliidae
are difficult to distinguish from
Libellulidae
, save for a small number of premental characters (
Tennessen, 2019
;
Theischinger and Fleck, 2003
). (
Ware
et al., 2007
) recovered
Aeschnosoma
+
Pentathemis
as sister to the remaining
Corduliidae
; these genera have remarkably similar looking nymphs, with long spines on segment nine, despite having ranges in South America and eastern
Australia
, respectively.
Fleck and Legrand (2013)
hypothesize that
Libellulosoma
forms a clade with
Aeschnosoma
and
Pentathemis
. Here, we recover
Pentathemis
as sister to the
Libellulidae
, but with low bootstrap support (96%) and among the poorest of QC values (-0.76), suggesting that this relationship needs further investigation before confirmation of this grouping. In the absence of
Neocordulia
,
Lauromacromia
,
Idomacromia
,
Nesocordulia
and other taxa of
incertae sedis
, we cannot yet address with confidence the broader composition of
Corduliidae
.
Libellulidae
is recovered with full support. However, there is a great amount of discord within the group demonstrated by both the highest concentration and lowest nodal supports across the topology. We recover three extremely poorly supported clades suggesting instead a polytomy among the taxa included in our analyses. However, there are clades within the libellulids that are well supported. The Libellulinae (BS = 100, PP = 0.95, QS = 1/NA/1), a large subfamily is recovered in our analyses and has been consistently recovered in past molecular work.
Nannophlebia
and
Zygonyx
were recovered in a clade previously, and here we find support for their sister relationship (BS = 100, PP = 0.96, QS = 0.92/0/1).
Ware
et al. (2007)
found
Pantala
to be closely related to
Zygonyx
and
Nannophlebia
; here a clade comprising these three taxa is recovered with low QS values (BS = 100, PP = 1, QS = -0.46/0/0.97). Past studies have recovered
Rhyothemis
in a clade with
Sympetrum
(e.g.,
Ware
et al., 2007
). Herein we also recover this relationship but with the inclusion of never before sequenced
Austrothemis
(BS = 56, PP = 0.75, QS = 0.17/0.33/0.86). The family
Libellulidae
is extremely species rich, comprising well over 1,000 species. Many of the members of this family have an elongated, bisected anal loop in their hind wing, and an oblique vein immediately following the nodus. Their secondary penile characters, and general adult and nymphal morphology strongly support the monophyly of this family. Afurther look at this family with a much expanded taxon sampling is necessary to better understand the complex evolutionary history that certainly represents one of the most rapidly radiating lineages in
Odonata
.