Name: Wesley Dondoni Colombo
Type: PhD thesis
Publication date: 17/05/2021

Namesort descending Role
Cecília Waichert Monteiro Co-advisor *
Celso Oliveira Azevedo Advisor *

Examining board:

Namesort descending Role
Ana Carolina Loss Rodrigues Internal Alternate *
Cecília Waichert Monteiro Co advisor *
Celso Oliveira Azevedo Advisor *
Dalton de Souza Amorim External Examiner *
FERNANDA TONINI GOBBI External Alternate *
Orlando Tobias Silveira External Examiner *
Taissa Rodrigues Marques da Silva Internal Examiner *

Summary: Epyrinae are the second most diverse subfamily of Bethylidae, with approximately 950
described species and 13 valid genera, of which 12 are extant: Anisepyris Kieffer;
Aspidepyris Evans; Bakeriella Kieffer; Calyozina Enderlein; Chlorepyris Kieffer;
Disepyris Kieffer; Epyris Westwood; Formosiepyris Terayama; Holepyris Kieffer; Laelius
Ashmead; Trachepyris Kieffer and Xenepyris Kieffer; and one is extinct: †Elektroepyris.
Epyrinae have a complex taxonomic history and only in the last decade their monophyly
has recovered. Due to the difficulty in delimiting Epyrinae, many generic taxa were
consequently classified inappropriately. Studies revising the concept of the epyrine genera
are necessary and crucial, but they have been neglected due to the difficulty in interpreting
morphostructural diversity. As a solution, molecular data has been used to help elucidate
the morphostructural impasses. In this dissertation, the diversity of Epyrinae is explored,
both alphaxonomically and phylogenetically, through molecular and morphological data,
and by reconstructing phylogenetic trees, under different optimization criteria (Maximum
Parsimony, Bayesian Inference and Maximum Likelihood). Additionally, the subfamily's
paleodiversity was revised and, at the end, the Epyrinae were organized and redefined and
the evolution of their lineages discussed. This thesis consists of four chapters, the first
three of which aimed to explore and list the morphological and molecular patterns of the
Epyrinae, providing data and information for the last chapter, which aimed to test the
monophyly of the Epyrinae genera. In Chapter 1, an alpha-taxonomic review of Epyrinae
from Papua New Guinea is present, exploring and listing morphological patterns and
filling in gaps in Old World taxonomic knowledge. Finally, 39 species were recognized,
both morphologically and molecularly, of which 36 are new and described. In Chapter 2,
the extinct genus †Elektroepyris was revised and cladistically positioned. Morphological
characters were coded, summing up to 69. The caracteres were used to perform Maximum
Parsimony reconstruction, including representatives of all subfamilies of Bethylidae, extant
or extinct. †Elektroepyris was recovered as a lineage distinct from other subfamilies of
Bethylidae and for this reason, a new subfamily, †Elektroepyrinae, was described, based
on an autapomorphy of the anterior wing with the third abscissa of the Cu vein. In Chapter
3, the paleodiversity of Epyrinae was reviewed alpha-taxonomically. Thirty extinct species
of Epyrinae were recognized, distributed in six genera (Anisepyris, Chlorepyris, Epyris,
†Gloxinius, Holepyris and Laelius). A new genus, †Gloxinius, has been proposed for the
subfamily to allocate †G. bifossatus and two species were transferred from Epyris to Chlorepyris. Three species were transferred from Epyris to Pristocerinae: †Merascylla is
proposed to allocate the species †M. atavellus, and two other species are transferred to
Pseudisobrachium. One species is transferred from Epyris to Scleroderminae: †Mael is
proposed to allocate the species †M. longiceps. The species †Laelius nudipennis is not a
bethylid and has been transferred to Platygastroidea incertae sedis. In Chapter 4, a
Bayesian inference analysis was performed to reconstruct a total evidence topology from a
matrix with 195 terminal taxa and 3599 characters, containing 232 morphological
characters and the genes 16S, 18S, 28S, COI and Cytb. This is the first phylogenetic study
that examined all 42 generic names of Epyrinae, including junior synonyms and extinct
taxa. The results recovered Anisepyris, Bakeriella, Calyozina, Chlorepyris, Disepyris,
Laelius, and Trachepyris as monophyletic and Formosiepyris, Epyris, and Holepyris as
polyphyletic. Two new genera are proposed, GenusA to be proposed as gen. nov. and
GenusB to be proposed as gen. nov., and six genera were revalidated, five junior synonyms
of Epyris and one of Holepyris: Calyoza to be proposed as stat. rev., Dolus to be proposed
as stat. rev., Muellerella to be proposed as stat. rev., Psilepyris to be proposed as stat.
rev., Rysepyris to be proposed as stat. rev. and Trissepyris to be proposed as stat. rev. In
addition, all 962 species of Epyrinae were reviewed via direct observation of the holotype,
illustrations, or literature and were distributed over 17 genera: Anisepyris, Aspidepyris,
Bakeriella, Calyoza to be proposed as stat. rev., Calyozina, Chlorepyris, Dolus to be
proposed as stat. rev., Epyris, GenusA to be proposed as gen. nov., GenusB to be proposed
as gen. nov., †Gloxinius, Holepyris, Laelius, Muellerella to be proposed as stat. rev.,
Psilepyris to be proposed as stat. rev., Rysepyris to be proposed as stat. rev. and
Trissepyris to be proposed as stat. rev..

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