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Anatomic glossary of mesopleural structures in Bethylidae (Hymenoptera: Chrysidoidea)

Abstract

The mesopleuron of Bethylidae has many structures that are used in taxonomic and phylogenetic studies. The lack of understanding of these structures has generated independent terminologies and a series of confusing terms, hampering effective scientific communication. A morphological study and literature review were made in order to solve these problems. Our study resulted in an anatomic glossary with 49 terms that presented a large number of synonyms and polysemies. The glossary standardizes the terms used in the Bethylidae mesopleuron and in other Hymenoptera groups, which will facilitate hypotheses of primary homology in comparative biology.

Keywords.
Terminological polysemy; Terminological synonyms; Mesopleuron; Morphology

INTRODUCTION

Most of the morphological studies of Hymenoptera have a generalized anatomical approach, in which all or large areas of the body are analyzed together (e.g.,Gibson, 1985Gibson, G.A.P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. The Canadian Entomologist, 117(11): 1395-1443., 1993Gibson, G.A.P. 1993. Groundplan structure and homology of the pleuron in Hymenoptera based on a comparison of the skeletomusculature of Xyelidae (Hymenoptera) and Raphidiidae (Neuroptera). Memoirs of the Entomological Society of Canada, 125: 165-187.; Matsuda, 1970Matsuda, R. 1970. Morphology and evolution of the insect thorax. Memoirs of the Entomological Society of Canada , 102 (Suppl. 76): 5-431. http://doi.org/10.4039/entm10276fv
http://doi.org/10.4039/entm10276fv...
; Snodgrass, 1910Snodgrass, R.E. 1910. The thorax of the Hymenoptera. Proceedings of the United States National Museum, 39: 37-21. http://doi.org/10.5479/si.00963801.39-1774.37
http://doi.org/10.5479/si.00963801.39-17...
; Vilhelmsen et al., 2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
; Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). In addition, many of these studies are restricted to only a few groups within the order (e.g., Ibaliidae - Ronquist & Nordlander, 1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.; Platygastridae - Masner, 1979Masner, L. 1979. Pleural morphology in scelionid wasps (Hymenoptera, Scelionidae) - an aid to higher classification. The Canadian Entomologist , 111(9): 1079-1087. and Mikó et al., 2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.; Vespidae - Duncan, 1939Duncan, C.D. 1939. A Contribution to The Biology of North American Vespine Wasps. Stanford University Publications, Biological Sciences, 8: 1-233. and Michener, 1944Michener, C.D. 1944. The Social Behavior of the Bees: A Comparative Study. Cambridge, Harvard University Press.; Braconidae - Karlsson & Ronquist, 2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
and Wharton, 2006Wharton, R.A. 2006. The species of Sternaulopius Fischer (Hymenoptera: Braconidae, Opiinae) and the braconid sternaulus. Journal of Hymenoptera Research , 15: 317-347.; and Eupelmidae - Gibson, 1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.).

Although these studies provided great contributions to the understanding of Hymenoptera and insect anatomy, anatomical studies restricted to more limited anatomical regions allow their characters to be discussed in more detail, and facilitates a more comprehensive comparative analysis. Examples are Shcherbakov (1980Shcherbakov, D.E. 1980. Morphology of the pterothoracic pleura of Hymenoptera. 1. Groundplan. Zoologicheskii Zhurnal, 59(11): 1644-1653., 1981) and Gibson (1993Gibson, G.A.P. 1993. Groundplan structure and homology of the pleuron in Hymenoptera based on a comparison of the skeletomusculature of Xyelidae (Hymenoptera) and Raphidiidae (Neuroptera). Memoirs of the Entomological Society of Canada, 125: 165-187.), who contributed considerably to the understanding of the thoracic pleura of Hymenoptera, accomplishing a detailed study of the external and internal morphology, comparing structures, standardizing terms and discussing the variations in different taxa.

Despite the various morphological studies, many cladistic and taxonomic studies explore the anatomy of taxa in a restricted and particular way, creating new terms for the analyzed characters or using previous terms without further exploring the homology between the characters. Such factors contribute to a progressive increase in the number of polysemy and synonyms in the terminology. This is evident in groups within Hymenoptera, in which evolutionary events have caused the loss or fusion of various anatomical structures (e.g., mesopleural and mesopleurosternal sutures). To try to minimize these problems in Bethylidae, Kawada et al. (2015Kawada, R.; Lanes, G.O. & Azevedo, C.O. 2015. Evolution of metapostnotum in flat wasps (Hymenoptera, Bethylidae): implications for homology assessments in Chrysidoidea. PLoS ONE , 10(10): e0140051. http://doi.org/10.1371/journal.pone.0140051
http://doi.org/10.1371/journal.pone.0140...
) carried out a study of the skeletomusculature of the metapostnotum to solve problems of delimiting the boundary between the metanotum and the propodeum. However, there are areas that still need a skeletomuscular study to understand their structures and propose homologies, such as the mesopleuron.

The Bethylidae mesopleuron presents a wide variety of structures of sulci, pits, foveae and impressions that are frequently used as taxonomic and cladistic characters (see: Evans, 1959Evans, H.E. 1959. The genus Anisepyris in America North of Mexico (Hymenoptera, Bethylidae). Proceedings of the Entomological Society of Washington, 61: 97-120.; Alencar & Azevedo, 2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12.; Azevedo et al., 2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.; Alencar & Azevedo, 2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.; Waichert & Azevedo, 2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.). Many of them are poorly understood, unexploited and there is no clarity about their homologies (e.g., mesepimeral sulcus in Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.), Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.), Barbosa et al. (2014Barbosa, D.N.; Kawada, R. & Azevedo, C.O. 2014. Order Hymenoptera, family Bethylidae. Genus Israelius, Richards, 1952. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Department of the President’s Affairs. v. 5, p. 463-470.); sub-anterior fovea in Alencar & Azevedo (2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12.); lower mesopleural fovea in Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.)). Moreover, the variety of structures has generated many terms, resulting in high quantities of synonyms and polysemies, hampering the taxonomic and cladistic analysis (e.g., synonymous structures: mesepimeral suture by Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.); mesopleural groove by Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.) and postpectal carina by Lanes & Azevedo (2008Lanes, G.O. & Azevedo, C.O. 2008. Phylogeny and taxonomy of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae). Insect Systematic and Evolution, 39: 55-86. http://doi.org/10.1163/187631208788784165
http://doi.org/10.1163/18763120878878416...
). Polysemous structures: sub-tegular groove by Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.); subalar pit by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
) and Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.)).

In an attempt to mitigate the overlapping of terms, many authors (e.g.,Gibson, 1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.; Karlsson & Ronquist, 2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
; Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.; Mikó et al., 2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.; Ronquist & Nordlander, 1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.) have comparatively analyzed the terminology applied to Hymenoptera characters (e.g., HAO, Antbase - Agosti & Johnson (2005Agosti, D. & Johnson, N.J. 2005. Antbase. World Wide Web electronic publication. Available: Available: http://antbase.org . Access: 08/02/2021.
http://antbase.org...
), Antkey - Sarnat & Suarez (28/04/2021)Sarnat, E.M. & Suarez, A.V. 2012. Antkey. University of Illinois and Identification Technology Program, CPHST, PPQ, APHIS, USDA. Available: Available: http://www.antkey.org/en . Access: 28/04/2021.
http://www.antkey.org/en...
). Nevertheless, the characters of the mesopleuron are still little understood by many authors, being necessary a more detailed organization of the concepts and terminology of these characters.

This study aims to propose a detailed glossary of the terms and concepts of mesopleural structures, as well as a list of synonyms, which will assist in future investigations of mesopleuron in Bethylidae and other groups of Hymenoptera, serving as a basis for comparison, understanding and character recognition as well as their use in phylogenies.

MATERIAL AND METHODS

Sample

Specimens of 33 genera of Bethylidae belonging to all five extant subfamilies were examined covering most of the diversity of the mesopleuron (Table 1). The homologies with other Hymenoptera families were made by comparisons with literature.

Table 1
Genera examined.

The examined material is from the following institutions: CASC: California Academy of Sciences, USA, San Francisco (R. Zuparko); CZMA: Coleção Zoológica do Maranhão, Brazil, Caxias (F. Limeira-de-Oliveira); ISAM: Iziko South African Museum, South Africa, Cape Town (S. van Noort); MNHN: Museum National d’Histoire Naturelle, France, Paris (Agnièle Touret-Alby, Bernardo Santos); NMKE: National Museum of Kenya, Kenya, Nairobi (R. Copeland); QMSB: Queensland Museum South Bank, Australia, Brisbane (Chris Burwell); QSBG: Queen Sirikit Botanic Garden, Thailand, Chiang Mai (W. Srisuka); UFES: Laboratory of Bethylidae at Universidade Federal do Espírito Santo, Brazil, Vitória (Celso O. Azevedo).

Dissection techniques and images

Critical-point-dried and air-dried specimens (Gibson, 1985Gibson, G.A.P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. The Canadian Entomologist, 117(11): 1395-1443.; Vilhelmsen et al., 2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
; Kawada et al., 2015Kawada, R.; Lanes, G.O. & Azevedo, C.O. 2015. Evolution of metapostnotum in flat wasps (Hymenoptera, Bethylidae): implications for homology assessments in Chrysidoidea. PLoS ONE , 10(10): e0140051. http://doi.org/10.1371/journal.pone.0140051
http://doi.org/10.1371/journal.pone.0140...
) were used for morphological analysis, dissected with insect pins and razor blades. The specimen’s parts were glued on a card with information on the sample’s provenance and stored as vouchers.

The external and internal part of the mesopleuron was photographed with a Leica Z16 APO stereomicroscope coupled to a Leica DFC 2 video camera by Leica Microsystems (Switzerland) following the lighting procedures of Kawada & Buffington (2016Kawada, R. & Buffington, M. 2016. A scalable and modular dome illumination system for scientific microphotography on a budget. PLoS ONE , 11(5): e0153426. http://doi.org/10.1371/journal.pone.0153426
http://doi.org/10.1371/journal.pone.0153...
). The image stacks were combined with Leica Application Suite V3.6.0 by Leica Microsystems (Switzerland) or Helicon Focus (HeliconSoft).

Terminology

The terms and descriptions of the morphological characters and muscles follow The Hymenoptera Anatomy Ontology project (Seltmann et al., 2012Seltmann, K.C.; Yoder, M.J.; Mikó, I.; Forshage, M.; Bertone M.A.; Agosti, D.; Balhoff, J.; Borowiec, M.; Brady, S.; Broad, G.; Brothers, D.; Burks, R.; Buffington, M.; Campbell, H.; Dew, K.; Ernst, A.; Fernandez-Triana, J.; Gates, M.; Gibson, G.; Jennings, J.; Johnson, N.; Karlsson, D.; Kawada, R.; Krogmann, L.; Kula, R.; Ohl, M.; Rasmussen, C.; Ronquist, F.; Schulmeister, S.; Sharkey, M.; Talamas, E.; Tucker, E.; Vilhelmsen, L.; Ward, P. & Wharton, R. 2012. A hymenopterists’ guide to the Hymenoptera Anatomy Ontology: utility, clarification, and future directions. Journal of Hymenoptera Research , 27: 67-88. http://doi.org/10.3897/jhr.27.2961
http://doi.org/10.3897/jhr.27.2961...
) available through The Hymenoptera Glossary (HAO http://glossary.hymao.org) and Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). The homologisation of structures (primary homology) between Bethylidae and other Hymenoptera was made by comparing the shape, location and muscles associated with the structures. When it was not possible to establish homology through association with the musculature, we applied the criterion of the topographical sameness of structures (see Seltmann et al., 2012Seltmann, K.C.; Yoder, M.J.; Mikó, I.; Forshage, M.; Bertone M.A.; Agosti, D.; Balhoff, J.; Borowiec, M.; Brady, S.; Broad, G.; Brothers, D.; Burks, R.; Buffington, M.; Campbell, H.; Dew, K.; Ernst, A.; Fernandez-Triana, J.; Gates, M.; Gibson, G.; Jennings, J.; Johnson, N.; Karlsson, D.; Kawada, R.; Krogmann, L.; Kula, R.; Ohl, M.; Rasmussen, C.; Ronquist, F.; Schulmeister, S.; Sharkey, M.; Talamas, E.; Tucker, E.; Vilhelmsen, L.; Ward, P. & Wharton, R. 2012. A hymenopterists’ guide to the Hymenoptera Anatomy Ontology: utility, clarification, and future directions. Journal of Hymenoptera Research , 27: 67-88. http://doi.org/10.3897/jhr.27.2961
http://doi.org/10.3897/jhr.27.2961...
).

Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.) and Seltmann et al. (2012Seltmann, K.C.; Yoder, M.J.; Mikó, I.; Forshage, M.; Bertone M.A.; Agosti, D.; Balhoff, J.; Borowiec, M.; Brady, S.; Broad, G.; Brothers, D.; Burks, R.; Buffington, M.; Campbell, H.; Dew, K.; Ernst, A.; Fernandez-Triana, J.; Gates, M.; Gibson, G.; Jennings, J.; Johnson, N.; Karlsson, D.; Kawada, R.; Krogmann, L.; Kula, R.; Ohl, M.; Rasmussen, C.; Ronquist, F.; Schulmeister, S.; Sharkey, M.; Talamas, E.; Tucker, E.; Vilhelmsen, L.; Ward, P. & Wharton, R. 2012. A hymenopterists’ guide to the Hymenoptera Anatomy Ontology: utility, clarification, and future directions. Journal of Hymenoptera Research , 27: 67-88. http://doi.org/10.3897/jhr.27.2961
http://doi.org/10.3897/jhr.27.2961...
) use the term homonymy when the same name is used for more than one structure. However, according to Martins (2015Martins, S.C. 2015. A sinonímia, a polissemia e a homonímia no vocabulário da fauna e da flora. Estudos Linguísticos, 44: 186-207.), the homonymy is related to cases in which two or more meanings are presented by words with the same spelling and/or sound, but have different semantic roots, while polysemy is characterized when a word with the same semantic root has two or more meanings. Furthermore, Alves (2000Alves, I.A. 2000. Polissemia e homonímia em uma perspectiva terminológica. Alfa, Revista de Linguistica, 44: 261-272. https://periodicos.fclar.unesp.br/alfa/issue/view/300.
https://periodicos.fclar.unesp.br/alfa/i...
) explains that polysemy and homonymy may represent cases in which the same term is used to name different concepts, but polysemy deals with concepts from the same area or conceptual network, while homonymy from different conceptual areas or networks.

From that, we will adopt polysemy rather than homonymy when one name is applied to two or more structures and synonymy when the same structure has more than one different name applied to itself. We establish three criteria according to the order of the importance to define the preferential term: (1) was preferred the term that best represents the skeletal structure (shape and location in the body) according to Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.), Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.) and HAO. (2) more widely used term in the scientific community; (3) oldest proposition of the term. The criteria adopted are informed in the description of the structure.

In polysemy cases or if the adopted terminology was not adequate (second criterion described in the previous paragraph), a new name was proposed. Some descriptions have been expanded because the previous definitions were insufficient to describe the characters. These definitions were structured following the genus differentia format, using the existing definition and from it we add the new part to cover the structural concept.

The positioning of characters in relation to the body follows the Biological Spatial Ontology - BSPO (Dahdul et al., 2014Dahdul, W.M.; Cui, H.; Mabee, P.M.; Mungall, C.J.; Osumi-Sutherland, D.; Walls, R.L. & Haendel, M.A. 2014. Nose to tail, roots to shoots: spatial descriptors for phenotypic diversity in the Biological Spatial Ontology. Journal of Biomedical Semantics, 5(34): 1-13.; BSPOBiological Spatial Ontology (BSPO). Biological Spatial Ontology: An ontology for representing spatial concepts, anatomical axes, gradients, regions, planes, sides, and surfaces. Available: Available: http://www.obofoundry.org/ontology/bspo.html . Access: 08/02/2021.
http://www.obofoundry.org/ontology/bspo....
) and the interpretation follows The Hymenoptera Anatomy Ontology (Yoder et al., 2010Yoder, M.; Mikó, I.; Seltmann, K.; Bertone, M. & Deans, A. 2010. A gross anatomy ontology for Hymenoptera. PLoS ONE , 5(12). http://doi.org/10.1371/journal.pone.0015991
http://doi.org/10.1371/journal.pone.0015...
; Seltmann et al., 2012Seltmann, K.C.; Yoder, M.J.; Mikó, I.; Forshage, M.; Bertone M.A.; Agosti, D.; Balhoff, J.; Borowiec, M.; Brady, S.; Broad, G.; Brothers, D.; Burks, R.; Buffington, M.; Campbell, H.; Dew, K.; Ernst, A.; Fernandez-Triana, J.; Gates, M.; Gibson, G.; Jennings, J.; Johnson, N.; Karlsson, D.; Kawada, R.; Krogmann, L.; Kula, R.; Ohl, M.; Rasmussen, C.; Ronquist, F.; Schulmeister, S.; Sharkey, M.; Talamas, E.; Tucker, E.; Vilhelmsen, L.; Ward, P. & Wharton, R. 2012. A hymenopterists’ guide to the Hymenoptera Anatomy Ontology: utility, clarification, and future directions. Journal of Hymenoptera Research , 27: 67-88. http://doi.org/10.3897/jhr.27.2961
http://doi.org/10.3897/jhr.27.2961...
; HAO).

From the terminological glossary by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.) and the bibliography consulted, which presents information about structures of the mesopleuron in Hymenoptera (mainly morphological terms and concepts) and Bethylidae specifically (including taxonomic, morphological and cladistic terms applied to structures in mesopleuron), a new glossary for the mesopleural structures was made, in which the detailed description of the morphological characters and the commonly used synonyms were included. Valid terms and abbreviations are in bold and new terms are marked with one asterisk (*). In case of articles written in a non-English language, we translated the terms and kept the term in the original language in parentheses and marked with a double asterisk (**). When the term represents only part of a structure, we include the term “in part”.

RESULTS

General description of the mesopleuron in Bethylidae

Mesopleuron occupies a large part of the mesopectus (mesopleuron + mesosternum), being divided into lateral and ventral surfaces. Located dorsally in the lateral surface, the mesepimeral sulcus (epm2s) separates the posterior mesepimeral area (epm2pa) from the rest of the mesopleuron, which is composed of the mesepisternum and part of the mesepimeron. The posterior mesepimeral area is projected anteriorly, forming the mesepimeral lobe (mlo). The subalar area (saa) is located anterodorsally in the mesopleuron and is ventrally limited by the subalar impression (sai), within which are the anterior subalar pit (asup) and posterior subalar pit (psup). Ventral to the subalar impression may have the anterior mesopleural fovea (pl2af). Posteriorly to this fovea there are a mesopleural callus (pl2c) or an upper mesopleural fovea (pl2uf) or a posterior oblique sulcus of the mesopleuron (pl2pos). Centrally in the mesopleuron, the mesopleural pit (pl2p) is present, which can be internal to the upper mesopleural fovea or to the posterior oblique sulcus of the mesopleuron. Ventrally in the lateral surface, the lower mesopleural fovea (pl2lf) is usually present and at the posterior extremity there is the mesopleural epicoxal lobe (pl2el), which may be limited by the mesopleural epicoxal sulcus (pl2es). In the ventral surface, subpleural signum (sps) and sternaulus (st) are usually present.

Integumentary structures

Acropleural apodeme (acra:Figs. 1A,2D) = The apodeme located anterodorsally on the mesopleuron and correspondent to the site of origin of the third mesopleuro-mesonotal muscle (HAO_0000083). Although the acropleural apodeme is present in Bethylidae, we were not able to see the third mesopleuro-mesonotal muscle.

acropleural apodeme by Gibson (1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Figure 1
Mesopleuron. (A) Chlorepyris. (B) Epyris. (C) Chlorepyris. (D) Anisepyris. (E) Chlorepyris. (F) Pseudisobrachium. (A) dorsal view. (B-F) lateral view. Abbreviations: acra = acropleural apodeme; pl2af = anterior mesopleural fovea; psup = posterior subalar pit; pl2c = mesopleural callus; asup = anterior mesopleural pit; pl2lf = lower mesopleural fovea; pl2uf = upper mesopleural fovea; epm2s = mesepimeral sulcus; pl2pos = posterior oblique sulcus of the mesopleuron.

Figure 2
Mesopleuron. (A) Odontepyris. (B) Pristepyris. (C-D) Pristocera. (E) Pseudisobrachium. (F) Cleistepyris. (A, B, D, E) internal view. (C, F) lateral view. Abbreviations: epm2r = mesepimeral ridge; pl2dp = mesopleural dentate process; epm2i = mesepimeral inflection; cx2-sa2 = mesocoxo-mesosubalar muscle; sai = subalar impression; epm2pa = posterior mesepimeral area; epm2s = mesepimeral sulcus; st = sternaulus; ste = sternaular elevation; acra = acropleural apodeme; pl2a-fu2 = anterior mesopleuro-mesofurcal muscle; pl2a = mesopleural apodeme; pl2-cx2 = mesopleuro-mesocoxal muscle; mlo = mesepimeral lobe; pl2es = mesopleural epicoxal sulcus; pl2el = mesopleural epicoxal lobe.

Anterior mesopleural fovea (pl2af:Figs. 1B, C) = The small depression located near the anterior margin of the mesopleuron and ventral to the anterior end of the subalar impression (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

anterior fovea by Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.); Azevedo & Alencar (2010aAzevedo, C.O. & Alencar, I.D.C.C. 2010a. Rediscovery of the Afrotropical genus Trissepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Epyris Westwood. Journal of Afrotropical Zoology, 6: 3-7., bAzevedo, C.O. & Alencar, I.D.C.C. 2010b. Rediscovery of Melanepyris (Hymenoptera: Bethylidae): a new synonym of Epyris. Revista Brasileira de Zoologia , 27(3): 403-407. http://doi.org/10.1590/S1984-46702010000300012
http://doi.org/10.1590/S1984-46702010000...
); Azevedo & Barbosa (2010Azevedo, C.O. & Barbosa, D.N. 2010. Two new species Pararhabdepyris Gorbatovskii (Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa, 2668: 55-62.); Azevedo et al. (2005Azevedo, C.O.; Moreira, A.R. & Kawada, R. 2005. Taxonomia de Bakeriella (Hymenoptera, Bethylidae) da Bolívia. Iheringia, Série Zoológica , 95(2): 165-172., 2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Barbosa & Azevedo (2009Barbosa, D.N. & Azevedo, C.O. 2009. Laelius Ashmead (Hymenoptera, Bethylidae) from Madagascar, with description of two new species. Zootaxa, 2170: 1-14., 2010Barbosa, D.N. & Azevedo, C.O. 2010. Order Hymenoptera, family Bethylidae, genus Laelius . In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing. v. 3, p. 412-421., 2011bBarbosa, D.N. & Azevedo, C.O. 2011b. Taxonomy of Allobethylus Kieffer (Hymenoptera, Bethylidae) from southeastern Asia. Journal of Asia-Pacific Entomology, 14: 89-94., 2014aBarbosa, D.N. & Azevedo, C.O. 2014a. Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species. Zoologia, 31: 285-311., bBarbosa, D.N. & Azevedo, C.O. 2014b. Moczariella, a bizarre new genus of Mesitiinae (Hymenoptera, Bethylidae) from Arab Peninsula. Zootaxa, 3860: 291-297., 2015Barbosa, D.N. & Azevedo, C.O. 2015. Synopsis of Bradepyris Kieffer, 1905 (Hymenoptera, Bethylidae, Mesitiinae). European Journal of Taxonomy, 151: 1-16. http://doi.org/10.5852/ejt.2015.151
http://doi.org/10.5852/ejt.2015.151...
); Barbosa et al. (2013Barbosa, D.N.; Perkovsky, E.E. & Azevedo, C.O. 2013. Two new species of Laelius Ashmead (Hymenoptera: Bethylidae) from Upper Eocene Rovno amber. Neues Jahrbuch fur Geologie und Palaontologie, 267: 67-73. http://doi.org/10.1127/0077-7749/2012/0297
http://doi.org/10.1127/0077-7749/2012/02...
); Lanes & Azevedo (2008Lanes, G.O. & Azevedo, C.O. 2008. Phylogeny and taxonomy of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae). Insect Systematic and Evolution, 39: 55-86. http://doi.org/10.1163/187631208788784165
http://doi.org/10.1163/18763120878878416...
); Stein et al. (2011Stein, P.R.W.; Alencar I.D.C.C.; Barbosa, D.N. & Azevedo, C.O. 2011. Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species. Zootaxa, 3032: 17-32.); Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

anterior mesopleural fovea by Azevedo (2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
); Barbosa & Azevedo (2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Anterior subalar pit (asup:Fig. 1C) = The pit located anterodorsally on the mesopleuron that corresponds internally to the acropleural apodeme*. Criterion 1.

acropleural sulcus by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

subalar pit by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

subtegular fovea by Azevedo & Lanes (2009Azevedo, C.O. & Lanes, G.O. 2009. Cladistic assessment and redescription of Galodoxa torquata Nagy (Hymenoptera, Bethylidae), a striking species with swallow tailed metasomal sternite. Zoologische Mededelingen Leiden, 83(4): 841-851.); Barbosa & Azevedo (2009Barbosa, D.N. & Azevedo, C.O. 2009. Laelius Ashmead (Hymenoptera, Bethylidae) from Madagascar, with description of two new species. Zootaxa, 2170: 1-14., 2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275.); Barbosa et al. (2013Barbosa, D.N.; Perkovsky, E.E. & Azevedo, C.O. 2013. Two new species of Laelius Ashmead (Hymenoptera: Bethylidae) from Upper Eocene Rovno amber. Neues Jahrbuch fur Geologie und Palaontologie, 267: 67-73. http://doi.org/10.1127/0077-7749/2012/0297
http://doi.org/10.1127/0077-7749/2012/02...
); Lim et al. (2010Lim, J.; Kwon, H.; Lee, J. & Lee, S. 2010. A new Laelius species (Hymenoptera: Bethylidae) from China with a key to the species in Eastern Palaearctic region. Journal of Asia-Pacific Entomology , 2010: 365-368. http://doi.org/10.1016/j.aspen.2010.07.001
http://doi.org/10.1016/j.aspen.2010.07.0...
).

Dorsal mesopleural inflection (pl2di: fig. 21B inLanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.) = The inflection extending along the dorsal margin of the mesopleuron anterior to the posterior subalar pit, which articulates with the second axillary sclerite (HAO_0000257).

dorsal mesopleural inflection by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Lateral coxal articular process of the mesopleuron (lcapp2:http://api.hymao.org/public/figure/show_zoom/12802in HAO) = The articular process located ventrolaterally on the mesopleuron, where bears the lateral coxal condyle of the mesopleuron (HAO_0001464).

lateral coxal articular process of the mesopleuron by Boudinot (2015Boudinot, B.E. 2015. Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. European Journal of Taxonomy , 120: 1-62.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Lower mesopleural fovea (pl2lf:Figs. 1C, D, E) = Wide depression located ventrally to the mesopleural pit (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 2.

curved carina (in part) by Krombein (1992Krombein, K.V. 1992. Systematics of the genera of Epyrinae with ramose male antennae (Hymenoptera: Bethylidae). Proceedings of the Entomological Society of Washington , 94(3): 345-360.).

inferior margin of lower fovea by Alencar & Azevedo (2011bAlencar, I.D.C.C. & Azevedo, C.O. 2011b. Revision of Malagasy Odontepyris Kieffer (Hymenoptera, Bethylidae). Zootaxa, 2935: 26-40.); Stein et al. (2011Stein, P.R.W.; Alencar I.D.C.C.; Barbosa, D.N. & Azevedo, C.O. 2011. Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species. Zootaxa, 3032: 17-32.).

lower fovea (or **fóvea inferior) by Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.); Azevedo (1992Azevedo, C.O. 1992. Sobre os Rhabdepyris Kieffer (Hymenoptera, Bethylidae) coletadas em Piçarras, SC, Brasil. Revista Brasileira de Entomologia, 36: 251-254., 1999aAzevedo, C.O. 1999a. Additions to the Neotropical Epyrinae (Hymenoptera, Bethylidae), with description of a new species of Lepidosternopsis from Brazil. Iheringia, Série Zoológica, 87: 11-18., cAzevedo, C.O. 1999c. On the Neotropical Rhabdepyris Kieffer (Hymenoptera Bethylidae) of the genus Chlorepyris. Revista Brasileira de Zoologia , 16(3): 887-897. http://doi.org/10.1590/S0101-81751999000300026
http://doi.org/10.1590/S0101-81751999000...
, 2011Azevedo, C.O. 2011. Synopsis of Aspidepyris Evans, 1964 (Hymenoptera, Bethylidae). Zootaxa, 3016: 63-68., 2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
); Azevedo & Alencar (2010aAzevedo, C.O. & Alencar, I.D.C.C. 2010a. Rediscovery of the Afrotropical genus Trissepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Epyris Westwood. Journal of Afrotropical Zoology, 6: 3-7., bAzevedo, C.O. & Alencar, I.D.C.C. 2010b. Rediscovery of Melanepyris (Hymenoptera: Bethylidae): a new synonym of Epyris. Revista Brasileira de Zoologia , 27(3): 403-407. http://doi.org/10.1590/S1984-46702010000300012
http://doi.org/10.1590/S1984-46702010000...
); Azevedo & Barbosa (2010Azevedo, C.O. & Barbosa, D.N. 2010. Two new species Pararhabdepyris Gorbatovskii (Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa, 2668: 55-62.); Azevedo & Mugrabi (2014Azevedo, C.O. & Mugrabi, D.F. 2014. Three new species of Tuberepyris Lanes et Azevedo (Hymenoptera, Bethylidae), with amended diagnosis of the genus. Zootaxa, 3794: 556-564.); Azevedo et al. (2005Azevedo, C.O.; Moreira, A.R. & Kawada, R. 2005. Taxonomia de Bakeriella (Hymenoptera, Bethylidae) da Bolívia. Iheringia, Série Zoológica , 95(2): 165-172.); Barbosa & Azevedo (2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275.); Barbosa et al. (2013Barbosa, D.N.; Perkovsky, E.E. & Azevedo, C.O. 2013. Two new species of Laelius Ashmead (Hymenoptera: Bethylidae) from Upper Eocene Rovno amber. Neues Jahrbuch fur Geologie und Palaontologie, 267: 67-73. http://doi.org/10.1127/0077-7749/2012/0297
http://doi.org/10.1127/0077-7749/2012/02...
); Corrêa & Azevedo (2002Corrêa, M.S. & Azevedo, C.O. 2002. Systematics of Epyris (Hymenoptera, Bethylidae) from Reserva Biológica de Duas Bocas, Espírito Santo, Brazil. Revista Brasileira de Entomologia , 46: 129-132. http://doi.org/10.1590/S0085-56262002000200004
http://doi.org/10.1590/S0085-56262002000...
); Evans (1964Evans, H.E. 1964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bulletin of the Museum of Comparative Zoology , 132: 1-222., 1965Evans, H.E. 1965. A revision of the genus Rhabdepyris in the Americas (Hymenoptera: Bethylidae). Bulletin of the Museum of Comparative Zoology , 133: 68-151., 1966aEvans, H.E. 1966a. Further studies on neotropical Pristocerinae (Hymenoptera, Bethylidae). Acta Hymenopterologica, 2: 99-117., 1967bEvans, H.E. 1967b. Notes on Mexican and Southwestern U.S. Bethylidae (Hymenoptera): Part II, Epyrinae. Entomological News , 78: 93-100., 1969cEvans, H.E. 1969c. A revision of the genus Epyris in the Americas (Hymenoptera, Bethylidae). Transactions of the American Entomological Society, 95: 181-352., 1978Evans, H.E. 1978. The Bethylidae of America North of Mexico. Memoirs of the American Entomological Institute, 27: 1-332.); Gobbi & Azevedo (2006Gobbi, F.T. & Azevedo, C.O. 2006. The Brazilian Holepyris (Hymenoptera, Bethylidae), with description of a new species from Amazonian forest. Zootaxa, 1357: 61-68.); Kawada & Azevedo (2003Kawada, R. & Azevedo, C.O. 2003. New species of Bakeriella (Hymenoptera, Bethylidae) from the New World and new data on geographic distribution. Revista Brasileira de Entomologia , 47(3): 461-468. http://doi.org/10.1590/S0085-56262003000300017
http://doi.org/10.1590/S0085-56262003000...
); Krombein (1992Krombein, K.V. 1992. Systematics of the genera of Epyrinae with ramose male antennae (Hymenoptera: Bethylidae). Proceedings of the Entomological Society of Washington , 94(3): 345-360.); Moreira & Azevedo (2003Moreira, A.R. & Azevedo, C.O. 2003. Taxonomy of Anisepyris (Hymenoptera, Bethylidae) from Estação Biológica de Santa Lúcia, Santa Teresa, ES, Brazil. Boletim do Museu Biológico Mello Leitão, Nova Série, 15: 35-39.); Rosmann & Azevedo (2005Rosmann, W.L. & Azevedo, C.O. 2005. Dez espécies novas e notas sobre Rhabdepyris Kieffer (Hymenoptera, Bethylidae) da Mata Atlântica. Revista Brasileira de Entomologia , 49(3): 397-408.); Santos H.S. & Azevedo (2000Santos, H.S. & Azevedo, C.O. 2000. Sistemática das espécies de Anisepyris Kieffer, 1905 (Hymenoptera, Bethylidae) do Brasil. Revista Brasileira de Entomologia , 44(3-4): 129-165.); Santos L.M. & Azevedo (2008Santos, L.M. dos & Azevedo, C.O. 2008. Taxonomia de Anisepyris Kieffer (Hymenoptera, Bethylidae) da Mata Atlântica Brasileira. Revista Brasileira de Entomologia , 52(3): 328-339. http://doi.org/10.1590/S0085-56262008000300004
http://doi.org/10.1590/S0085-56262008000...
); Terayama (2006Terayama, M. 2006. Insects of Japan, Bethylidae (Hymenoptera). Vol. 1. Fukuoka, Touka Shobo Co., 2008Terayama, M. 2008. Two new species of Epyris (Insecta, Hymenoptera, Bethylidae) from Sulawesi island of Indonesia. Japanese Journal of Systematic Entomology, 14: 145-149.); Vargas & Azevedo (2016aVargas, J.M. & Azevedo, C.O. 2016a. Three new species of Pararhabdepyris Gorbatovsky (Hymenoptera, Bethylidae) from Central Africa and the Arabian Peninsula. Zootaxa, 4117: 429-439. http://doi.org/10.11646/zootaxa.4117.3.10
http://doi.org/10.11646/zootaxa.4117.3.1...
, b); Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

lower margin (in part) by Azevedo (2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
); Azevedo & Alencar (2010aAzevedo, C.O. & Alencar, I.D.C.C. 2010a. Rediscovery of the Afrotropical genus Trissepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Epyris Westwood. Journal of Afrotropical Zoology, 6: 3-7., bAzevedo, C.O. & Alencar, I.D.C.C. 2010b. Rediscovery of Melanepyris (Hymenoptera: Bethylidae): a new synonym of Epyris. Revista Brasileira de Zoologia , 27(3): 403-407. http://doi.org/10.1590/S1984-46702010000300012
http://doi.org/10.1590/S1984-46702010000...
); Azevedo & Mugrabi (2014Azevedo, C.O. & Mugrabi, D.F. 2014. Three new species of Tuberepyris Lanes et Azevedo (Hymenoptera, Bethylidae), with amended diagnosis of the genus. Zootaxa, 3794: 556-564.); Santos L.M. & Azevedo (2008Santos, L.M. dos & Azevedo, C.O. 2008. Taxonomia de Anisepyris Kieffer (Hymenoptera, Bethylidae) da Mata Atlântica Brasileira. Revista Brasileira de Entomologia , 52(3): 328-339. http://doi.org/10.1590/S0085-56262008000300004
http://doi.org/10.1590/S0085-56262008000...
).

lower mesopleural fovea by Azevedo et al. (2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Evans (1959Evans, H.E. 1959. The genus Anisepyris in America North of Mexico (Hymenoptera, Bethylidae). Proceedings of the Entomological Society of Washington, 61: 97-120., 1966bEvans, H.E. 1966b. A revision of the genus Anisepyris Kieffer (Hymenoptera, Bethylidae). Studia Entomologica, 9: 1-120., 1970Evans, H.E. 1970. West Indian wasps of the subfamilies Epyrinae and Bethylinae (Hymenoptera: Bethylidae). Proceedings of the Entomological Society of Washington , 72(3): 340-356.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Lim & Azevedo, 2014Lim, J. & Azevedo, C.O. 2014. Five new species of Disepyris Kieffer, 1905 (Hymenoptera: Bethylidae), with emended generic diagnosis. Zootaxa, 3900: 526-540. http://doi.org/10.11646/zootaxa.3900.4.4
http://doi.org/10.11646/zootaxa.3900.4.4...
; Stein & Azevedo (2007Stein, P.R.W. & Azevedo, C.O. 2007. Synopsis of lobatifrons species-group (Hymenoptera: Bethylidae, Rhabdepyris) from Neotropical region, with redefinition of the group. Zootaxa, 1444: 23-34.).

lower part of lower fovea (in part) by Evans, 1978Evans, H.E. 1978. The Bethylidae of America North of Mexico. Memoirs of the American Entomological Institute, 27: 1-332..

posterior fovea by Azevedo (1999aAzevedo, C.O. 1999a. Additions to the Neotropical Epyrinae (Hymenoptera, Bethylidae), with description of a new species of Lepidosternopsis from Brazil. Iheringia, Série Zoológica, 87: 11-18.).

prepectal carina (in part) by Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

prepectal depression (in part) by Vargas & Azevedo (2016bVargas, J.M. & Azevedo, C.O. 2016b. Discovery of the male of Galodoxa torquata Nagy (Hymenoptera, Bethylidae), with emended generic diagnosis. In: Robillard, T.; Legendre, F.; Villemant, C. & Leponce, M. (Eds.). Insects of Mount Wilhelm, Papua New Guinea. Paris, Muséum National d’Histoire Naturelle. p. 451-564.).

transepisternal line (in part) by Azevedo et al. (2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

upper margin of lower fóvea (in part) by Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.); Azevedo (1994Azevedo, C.O. 1994. Descriptions of two new species and notes on the genus Bakeriella Kieffer from Brazil and Ecuador (Hymenoptera, Bethylidae). Journal of Hymenoptera Research, 3: 145-150.); Azevedo & Alencar (2010bAzevedo, C.O. & Alencar, I.D.C.C. 2010b. Rediscovery of Melanepyris (Hymenoptera: Bethylidae): a new synonym of Epyris. Revista Brasileira de Zoologia , 27(3): 403-407. http://doi.org/10.1590/S1984-46702010000300012
http://doi.org/10.1590/S1984-46702010000...
); Stein & Azevedo (2007Stein, P.R.W. & Azevedo, C.O. 2007. Synopsis of lobatifrons species-group (Hymenoptera: Bethylidae, Rhabdepyris) from Neotropical region, with redefinition of the group. Zootaxa, 1444: 23-34.).

Medial coxal articular process of the mesopleuron (mcapp2: figs. 20B and 21A inLanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.) = The articular process located on the ventral margin of the mesopectus medially of the mesocoxal foramen, where bears the medial coxal condyle of the mesopectus (HAOHymenoptera Anatomy Ontology (HAO). 2020. The Hymenoptera Glossary: Hymenoptera Anatomy Consortium. Available: Available: http://glossary.hymao.org . Access: 05/02/2021.
http://glossary.hymao.org...
/Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

medial coxal articular process of the mesopectus by (HAO).

medial coxal articular process of the mesopleuron by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

median mesocoxal articulation by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

mesocoxal condyle by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
).

Mesepimeral inflection (epm2i:Fig. 2B) = The inflection extending along the dorsal margin of the mesopectus (HAO_0000536). Criterion 1.

mesepimeral inflection by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.).

posterior mesepimeral inflection by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Mesepimeral lobe (mlo:Fig. 2F) = The posterodorsal mesepimeral lobe covering the posterior thoracic spiracle (Karlsson & Ronquist, 2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
). Criterion 1.

epimeral lobe by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

mesepimeral lobe by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
).

Mesepimeral ridge (epm2r:Figs. 2A, B) = The ridge extending along the posterior margin of the mesopectus (HAO_0000537). Criterion 1.

mesepimeral ridge by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

posterior mesopleural apodeme by Duncan (1939Duncan, C.D. 1939. A Contribution to The Biology of North American Vespine Wasps. Stanford University Publications, Biological Sciences, 8: 1-233.).

Mesepimeral sulcus (epm2s:Figs. 1E,2C) = The sulcus extending along the posterior margin of the mesopectus, which delimits the mesepimeral area and corresponds to the mesepimeral ridge (HAO_0000538). Criterion 1.

acropleural fovea (in part) by Barbosa & Azevedo (2014aBarbosa, D.N. & Azevedo, C.O. 2014a. Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species. Zoologia, 31: 285-311.).

acropleural sulcus (in part) by Barbosa & Azevedo (2014aBarbosa, D.N. & Azevedo, C.O. 2014a. Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species. Zoologia, 31: 285-311., bBarbosa, D.N. & Azevedo, C.O. 2014b. Moczariella, a bizarre new genus of Mesitiinae (Hymenoptera, Bethylidae) from Arab Peninsula. Zootaxa, 3860: 291-297., 2015Barbosa, D.N. & Azevedo, C.O. 2015. Synopsis of Bradepyris Kieffer, 1905 (Hymenoptera, Bethylidae, Mesitiinae). European Journal of Taxonomy, 151: 1-16. http://doi.org/10.5852/ejt.2015.151
http://doi.org/10.5852/ejt.2015.151...
, 2016Barbosa, D.N. & Azevedo, C.O. 2016. Australomesitius gen. nov. (Hymenoptera, Bethylidae: Mesitiinae), an unexpected new genus from Australia. Zootaxa, 4061: 447-450.); Barbosa et al. (2017Barbosa, D.N.; Kawada, R. & Azevedo, C.O. 2017. Order Hymenoptera, family Bethylidae. Genus Glenosema, Kiefer. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Department of the President’s Affairs . v. 6, p. 322-335.).

episternal furrow by Azevedo & Barbosa (2010Azevedo, C.O. & Barbosa, D.N. 2010. Two new species Pararhabdepyris Gorbatovskii (Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa, 2668: 55-62.); Azevedo & Mugrabi (2014Azevedo, C.O. & Mugrabi, D.F. 2014. Three new species of Tuberepyris Lanes et Azevedo (Hymenoptera, Bethylidae), with amended diagnosis of the genus. Zootaxa, 3794: 556-564.); Barbosa & Azevedo (2010Barbosa, D.N. & Azevedo, C.O. 2010. Order Hymenoptera, family Bethylidae, genus Laelius . In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing. v. 3, p. 412-421.; 2011bBarbosa, D.N. & Azevedo, C.O. 2011b. Taxonomy of Allobethylus Kieffer (Hymenoptera, Bethylidae) from southeastern Asia. Journal of Asia-Pacific Entomology, 14: 89-94., cBarbosa, D.N. & Azevedo, C.O. 2011c. Order Hymenoptera, family Bethylidae, subfamily Mesitiinae. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing . v. 4, p. 375-404., 2012Barbosa, D.N. & Azevedo, C.O. 2012. Revision of Mesitiinae (Hymenoptera, Bethylidae) from Madagascar, with description of eleven species. Zootaxa, 3417: 1-44.); Barbosa et al. (2013Barbosa, D.N.; Perkovsky, E.E. & Azevedo, C.O. 2013. Two new species of Laelius Ashmead (Hymenoptera: Bethylidae) from Upper Eocene Rovno amber. Neues Jahrbuch fur Geologie und Palaontologie, 267: 67-73. http://doi.org/10.1127/0077-7749/2012/0297
http://doi.org/10.1127/0077-7749/2012/02...
); Lim & Azevedo (2014Lim, J. & Azevedo, C.O. 2014. Five new species of Disepyris Kieffer, 1905 (Hymenoptera: Bethylidae), with emended generic diagnosis. Zootaxa, 3900: 526-540. http://doi.org/10.11646/zootaxa.3900.4.4
http://doi.org/10.11646/zootaxa.3900.4.4...
); Stein et al. (2011Stein, P.R.W.; Alencar I.D.C.C.; Barbosa, D.N. & Azevedo, C.O. 2011. Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species. Zootaxa, 3032: 17-32.).

episternal groove by Alencar & Azevedo (2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12.); Alencar et al. (2016Alencar, I.D.C.C.; Colombo, W.D. & Azevedo, C.O. 2016. Pristocera masii: rediscovery of the holotype and its transfer to Acrepyris (Hymenoptera: Bethylidae). Acta Entomologica Musei Nationalis Pragae, 56(2): 795-803.); Azevedo (2008aAzevedo, C.O. 2008a. Characterization of the types of the Neotropical Pseudisobrachium (Hymenoptera, Bethylidae), with a key to species. Revista Brasileira de Zoologia , 25(4): 737-801. http://doi.org/10.1590/S0101-81752008000400020
http://doi.org/10.1590/S0101-81752008000...
, 2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
); Azevedo et al. (2018aAzevedo, C.O.; Alencar, I.D.C.C. & Colombo, W.D. 2018a. Pairs in copulation of the highly dimorphic genus Pristocera Klug (Hymenoptera, Bethylidae) from Madagascar solve taxonomic problems of male-female associations. Zootaxa, 4433: 1-49.); Azevedo & Alencar (2009Azevedo, C.O. & Alencar, I.D.C.C. 2009. Rediscovery of Pristepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Acrepyris Kieffer. Zootaxa, 2287: 45-54.); Barbosa & Azevedo (2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275.); Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

mesepimeral sulcus by Azevedo et al. (2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

mesepimeral suture by Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.).

mesopleural groove by Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.).

mesopleural suture by Barbosa & Azevedo (2014aBarbosa, D.N. & Azevedo, C.O. 2014a. Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species. Zoologia, 31: 285-311., bBarbosa, D.N. & Azevedo, C.O. 2014b. Moczariella, a bizarre new genus of Mesitiinae (Hymenoptera, Bethylidae) from Arab Peninsula. Zootaxa, 3860: 291-297., 2015Barbosa, D.N. & Azevedo, C.O. 2015. Synopsis of Bradepyris Kieffer, 1905 (Hymenoptera, Bethylidae, Mesitiinae). European Journal of Taxonomy, 151: 1-16. http://doi.org/10.5852/ejt.2015.151
http://doi.org/10.5852/ejt.2015.151...
, 2016Barbosa, D.N. & Azevedo, C.O. 2016. Australomesitius gen. nov. (Hymenoptera, Bethylidae: Mesitiinae), an unexpected new genus from Australia. Zootaxa, 4061: 447-450.); Barbosa et al. (2014Barbosa, D.N.; Kawada, R. & Azevedo, C.O. 2014. Order Hymenoptera, family Bethylidae. Genus Israelius, Richards, 1952. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Department of the President’s Affairs. v. 5, p. 463-470., 2017Barbosa, D.N.; Kawada, R. & Azevedo, C.O. 2017. Order Hymenoptera, family Bethylidae. Genus Glenosema, Kiefer. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Department of the President’s Affairs . v. 6, p. 322-335.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

postpectal carina by Lanes & Azevedo (2008Lanes, G.O. & Azevedo, C.O. 2008. Phylogeny and taxonomy of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae). Insect Systematic and Evolution, 39: 55-86. http://doi.org/10.1163/187631208788784165
http://doi.org/10.1163/18763120878878416...
).

pleural sulcus by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

sub-alar fovea (in part) by Barbosa & Azevedo (2009Barbosa, D.N. & Azevedo, C.O. 2009. Laelius Ashmead (Hymenoptera, Bethylidae) from Madagascar, with description of two new species. Zootaxa, 2170: 1-14., 2010Barbosa, D.N. & Azevedo, C.O. 2010. Order Hymenoptera, family Bethylidae, genus Laelius . In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing. v. 3, p. 412-421., 2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275.); Lim et al. (2010Lim, J.; Kwon, H.; Lee, J. & Lee, S. 2010. A new Laelius species (Hymenoptera: Bethylidae) from China with a key to the species in Eastern Palaearctic region. Journal of Asia-Pacific Entomology , 2010: 365-368. http://doi.org/10.1016/j.aspen.2010.07.001
http://doi.org/10.1016/j.aspen.2010.07.0...
).

subtegular fovea (in part) by Azevedo (2011Azevedo, C.O. 2011. Synopsis of Aspidepyris Evans, 1964 (Hymenoptera, Bethylidae). Zootaxa, 3016: 63-68.); Azevedo & Alencar (2010bAzevedo, C.O. & Alencar, I.D.C.C. 2010b. Rediscovery of Melanepyris (Hymenoptera: Bethylidae): a new synonym of Epyris. Revista Brasileira de Zoologia , 27(3): 403-407. http://doi.org/10.1590/S1984-46702010000300012
http://doi.org/10.1590/S1984-46702010000...
); Azevedo & Barbosa (2010Azevedo, C.O. & Barbosa, D.N. 2010. Two new species Pararhabdepyris Gorbatovskii (Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa, 2668: 55-62.); Azevedo & Lanes (2009Azevedo, C.O. & Lanes, G.O. 2009. Cladistic assessment and redescription of Galodoxa torquata Nagy (Hymenoptera, Bethylidae), a striking species with swallow tailed metasomal sternite. Zoologische Mededelingen Leiden, 83(4): 841-851.); Barbosa & Azevedo (2011bBarbosa, D.N. & Azevedo, C.O. 2011b. Taxonomy of Allobethylus Kieffer (Hymenoptera, Bethylidae) from southeastern Asia. Journal of Asia-Pacific Entomology, 14: 89-94., cBarbosa, D.N. & Azevedo, C.O. 2011c. Order Hymenoptera, family Bethylidae, subfamily Mesitiinae. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing . v. 4, p. 375-404., 2012Barbosa, D.N. & Azevedo, C.O. 2012. Revision of Mesitiinae (Hymenoptera, Bethylidae) from Madagascar, with description of eleven species. Zootaxa, 3417: 1-44.).

subtegular furrow (in part) by Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.);

subtegular groove (in part) by Alencar & Azevedo (2009Alencar, I.D.C.C. & Azevedo, C.O. 2009. Rediscovery of Planepyris Kieffer: A new synomym of Trachepyris Kieffer, (Hymenoptera, Bethylidae). Doriana, 8: 1-12., 2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12., bAlencar, I.D.C.C. & Azevedo, C.O. 2011b. Revision of Malagasy Odontepyris Kieffer (Hymenoptera, Bethylidae). Zootaxa, 2935: 26-40.); Alencar et al. (2016Alencar, I.D.C.C.; Colombo, W.D. & Azevedo, C.O. 2016. Pristocera masii: rediscovery of the holotype and its transfer to Acrepyris (Hymenoptera: Bethylidae). Acta Entomologica Musei Nationalis Pragae, 56(2): 795-803.); Azevedo (2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
); Azevedo & Alencar (2009Azevedo, C.O. & Alencar, I.D.C.C. 2009. Rediscovery of Pristepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Acrepyris Kieffer. Zootaxa, 2287: 45-54., 2010aAzevedo, C.O. & Alencar, I.D.C.C. 2010a. Rediscovery of the Afrotropical genus Trissepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Epyris Westwood. Journal of Afrotropical Zoology, 6: 3-7.); Azevedo & Mugrabi (2014Azevedo, C.O. & Mugrabi, D.F. 2014. Three new species of Tuberepyris Lanes et Azevedo (Hymenoptera, Bethylidae), with amended diagnosis of the genus. Zootaxa, 3794: 556-564.); Azevedo et al. (2018aAzevedo, C.O.; Alencar, I.D.C.C. & Colombo, W.D. 2018a. Pairs in copulation of the highly dimorphic genus Pristocera Klug (Hymenoptera, Bethylidae) from Madagascar solve taxonomic problems of male-female associations. Zootaxa, 4433: 1-49., bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Lanes & Azevedo (2007Lanes, G.O. & Azevedo, C.O. 2007. Redescription and placement of the Oriental Scaphepyris rufus Kieffer (Hymenoptera: Bethylidae). Zootaxa, 1654: 55-60., 2008Lanes, G.O. & Azevedo, C.O. 2008. Phylogeny and taxonomy of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae). Insect Systematic and Evolution, 39: 55-86. http://doi.org/10.1163/187631208788784165
http://doi.org/10.1163/18763120878878416...
); Lim & Azevedo (2014Lim, J. & Azevedo, C.O. 2014. Five new species of Disepyris Kieffer, 1905 (Hymenoptera: Bethylidae), with emended generic diagnosis. Zootaxa, 3900: 526-540. http://doi.org/10.11646/zootaxa.3900.4.4
http://doi.org/10.11646/zootaxa.3900.4.4...
); Lim & Lee (2013Lim, J. & Lee, S. 2013. Taxonomy of the family Bethylidae (Hymenoptera: Chrysidoidea) from Cambodia and adjacent countries. I. genus Odontepyris Kieffer (Bethylidae: Bethylinae) with four new species and two new records. Journal of Natural History , 47(31-32): 1-22. http://doi.org/10.1080/00222933.2012.763057
http://doi.org/10.1080/00222933.2012.763...
).

sub-tegular groove (in part) by Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

subtegular groove of mesopleuron (in part) by Lanes & Azevedo (2008Lanes, G.O. & Azevedo, C.O. 2008. Phylogeny and taxonomy of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae). Insect Systematic and Evolution, 39: 55-86. http://doi.org/10.1163/187631208788784165
http://doi.org/10.1163/18763120878878416...
).

subtegular mesopleural fovea (in part) by Azevedo (2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
).

Mesepimeron (epm2:http://api.hymao.org/public/figure/show_zoom/10827in HAO) = The epimeron that is located on the mesopleuron (HAO_0000539). Criterion 1.

epimeron of mesopleuron by Snodgrass (1942Snodgrass, R.E. 1942. The skeleto-muscular mechanisms of the honeybee. Smithsonian Miscellaneous Collections, 103: 1-120.).

mesepimeron by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Michener (1944Michener, C.D. 1944. The Social Behavior of the Bees: A Comparative Study. Cambridge, Harvard University Press.).

Mesepisternum (eps2:http://api.hymao.org/public/figure/show_zoom/10826in HAO) = The episternum located on the mesopleuron (HAO_0000541). Criterion 1.

mesepisternum by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Snodgrass (1910Snodgrass, R.E. 1910. The thorax of the Hymenoptera. Proceedings of the United States National Museum, 39: 37-21. http://doi.org/10.5479/si.00963801.39-1774.37
http://doi.org/10.5479/si.00963801.39-17...
).

mesothoracic episternum by Snodgrass (1942Snodgrass, R.E. 1942. The skeleto-muscular mechanisms of the honeybee. Smithsonian Miscellaneous Collections, 103: 1-120.).

Mesocoxal foramen (cx2f:http://api.hymao.org/public/figure/show_zoom/29356in HAO) = The coxal foramen located on the mesopectus (HAO_0001785).

mesocoxal foramen by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Mesopectus (pec2:http://api.hymao.org/public/figure/show_zoom/28201in HAO) = The sclerite which is U-shaped in cross section, and connected anteriorly with the pronotum and the propectus, dorsally with the basalare, the mesonotum, the second axillary sclerite and the subalare, posteriorly with the metapectus and bears the mesodiscrimenal lamella and the mesofurca (HAO_0000557).

mesopectus by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Mesopleural apodeme (pl2a:Figs. 2E,3B) = The apodeme correspondent to the site of origin of the second mesopleuro-mesonotal and the mesopleuro-mesocoxal muscles (HAO_0001359). Criterion 1.

mesopleural apodeme by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

pleural apodeme by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Mesopleural callus (pl2c:Figs. 1B, F) = The area corresponding to the site of origin of the second mesopleuro-mesonotal muscle and part of the mesofurco-mesopleural muscle. Externally, it is a bulging area located on the anterodorsal region of the mesopleuron, that is generally smoother than the rest*. Criterion 1.

acropleural area by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

callus by Azevedo (2004Azevedo, C.O. 2004. A new species of Caloapenesia from Vietnam with discovery of the female of the genus. Spixiana, 27(2): 143-146., 2007Azevedo, C.O. 2007. The Provancher types of Bethylidae (Hymenoptera, Chrysidoidea). Boletim do Museu Biológico Mello Leitão, 22: 57-67., 2008aAzevedo, C.O. 2008a. Characterization of the types of the Neotropical Pseudisobrachium (Hymenoptera, Bethylidae), with a key to species. Revista Brasileira de Zoologia , 25(4): 737-801. http://doi.org/10.1590/S0101-81752008000400020
http://doi.org/10.1590/S0101-81752008000...
); Azevedo et al. (2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Corrêa & Azevedo (2001Corrêa, M.S. & Azevedo, C.O. 2001. Taxonomia de Apenesia (Hymenoptera, Bethylidae) na Reserva Biológica de Duas Bocas, Espírito Santo, Brasil. Iheringia, Série Zoológica , 90: 167-174. http://doi.org/10.1590/S0073-47212001000100017
http://doi.org/10.1590/S0073-47212001000...
, 2006Corrêa, M.S. & Azevedo, C.O. 2006. O gênero Apenesia (Hymenoptera, Bethylidae) na Mata Atlântica: notas e descrição de sete espécies novas. Revista Brasileira de Entomologia , 50(4): 439-449. http://doi.org/10.1590/S0085-56262006000400001
http://doi.org/10.1590/S0085-56262006000...
;) Evans (1961Evans, H.E. 1961. A revision of the genus Pseudisobrachium in the North and Central America (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology, 126: 211-318., 1963aEvans, H.E. 1963a. A revision of the genus Pristocera in the Americas (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology , 129: 241-290., bEvans, H.E. 1963b. A revision of the Apenesia in the Americas (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology , 130: 249-359., 1964Evans, H.E. 1964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bulletin of the Museum of Comparative Zoology , 132: 1-222.; 1966aEvans, H.E. 1966a. Further studies on neotropical Pristocerinae (Hymenoptera, Bethylidae). Acta Hymenopterologica, 2: 99-117., 1967aEvans, H.E. 1967a. Notes on Mexican and Southwestern U.S. Bethylidae (Hymenoptera): Part I, Pristocerinae. Entomological News, 78: 13-23., cEvans, H.E. 1967c. New generic records of Bethylidae from South America. Proceedings of the Entomological Society of Washington , 69(3): 269-272., 1969aEvans, H.E. 1969a. The genus Pseudisobrachium in Argentina and Chile. Acta Zoologica Liloana, 25: 45-65., bEvans, H.E. 1969b. The genera Apenesia and Dissomphalus in Argentina and Chile (Hymenoptera, Bethylidae). Breviora, 311: 1-23., 1978Evans, H.E. 1978. The Bethylidae of America North of Mexico. Memoirs of the American Entomological Institute, 27: 1-332.); Gobbi & Azevedo (2014Gobbi, F.T. & Azevedo, C.O. 2014. Revision of Caloapenesia (Hymenoptera, Bethylidae), with description of sixteen new species. Zootaxa, 3860: 501-535., 2016Gobbi, F.T. & Azevedo, C.O. 2016. Revision of Calobrachium Gobbi & Azevedo, gen. nov. (Hymenoptera, Bethylidae), with description of seven new Oriental species. Zootaxa, 4083: 221-238.); Leal & Azevedo (2001Leal, M.S. & Azevedo, C.O. 2001. Taxonomy of Apenesia Westwood (Hymenoptera, Bethylidae) from Paraná, Brazil. Revista Brasileira de Zoologia , 18(3): 673-679.); Ramos & Azevedo (2009Ramos, M.S. & Azevedo, C.O. 2009. Sinopse das espécies neotropicais do grupo brasiliensis do gênero Apenesia (Hymenoptera, Bethylidae). Iheringia, Série Zoológica , 99(4): 349-363.); Waichert & Azevedo (2004Waichert, C. & Azevedo, C.O. 2004. Fourteen new Species of Pseudisobrachium (Hymenoptera, Bethylidae) from Atlantic Rain Forest of Espírito Santo, Brazil. Zootaxa, 661: 1-22.).

mesopleural callus by Alencar & Azevedo (2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12.); Alencar et al. (2016Alencar, I.D.C.C.; Colombo, W.D. & Azevedo, C.O. 2016. Pristocera masii: rediscovery of the holotype and its transfer to Acrepyris (Hymenoptera: Bethylidae). Acta Entomologica Musei Nationalis Pragae, 56(2): 795-803.); Azevedo & Alencar (2009Azevedo, C.O. & Alencar, I.D.C.C. 2009. Rediscovery of Pristepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Acrepyris Kieffer. Zootaxa, 2287: 45-54.); Azevedo et al. (2018aAzevedo, C.O.; Alencar, I.D.C.C. & Colombo, W.D. 2018a. Pairs in copulation of the highly dimorphic genus Pristocera Klug (Hymenoptera, Bethylidae) from Madagascar solve taxonomic problems of male-female associations. Zootaxa, 4433: 1-49.); Evans (1961Evans, H.E. 1961. A revision of the genus Pseudisobrachium in the North and Central America (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology, 126: 211-318., 1963bEvans, H.E. 1963b. A revision of the Apenesia in the Americas (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology , 130: 249-359., 1973Evans, H.E. 1973. Further studies on South American Bethylidae (Hymenoptera). Proceedings of the Entomological Society of Washington , 75: 194-204., 1978Evans, H.E. 1978. The Bethylidae of America North of Mexico. Memoirs of the American Entomological Institute, 27: 1-332.); Gobbi & Azevedo (2010Gobbi, F.T. & Azevedo, C.O. 2010. Taxonomia de Pseudisobrachium (Hymenoptera, Bethylidae) da Mata Atlântica Brasileira. Revista Brasileira de Entomologia , 54(2): 173-224. http://doi.org/10.1590/S0085-56262010000200003
http://doi.org/10.1590/S0085-56262010000...
); Zamprogno & Azevedo (2014Zamprogno, L.N. & Azevedo, C.O. 2014. Phylogeny and reclassification of Pristocera Klug (Hymenoptera: Bethylidae). Insect Systematics & Evolution , 45: 1-49. http://doi.org/10.1163/1876312X-04402003
http://doi.org/10.1163/1876312X-04402003...
).

mesopleural elevation by Alencar & Azevedo (2009Alencar, I.D.C.C. & Azevedo, C.O. 2009. Rediscovery of Planepyris Kieffer: A new synomym of Trachepyris Kieffer, (Hymenoptera, Bethylidae). Doriana, 8: 1-12.).

mesopleural upper elevation by Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.).

supra-posterior callus by Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.).

Mesopleural dentate process (pl2dp:Fig. 2A) = Projection located posterodorsally in mesopleuron (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

blunt tooth by Krombein (1996Krombein, K.V. 1996. Biosystematic studies of Ceylonese wasps, XXI: A revision of the Bethylinae and Epyrinae Cephalonomiini and Sclerodermini (Hymenoptera: Bethylidae). Smithsonian Contributions to Zoology, 579: 1-29. http://doi.org/10.5479/si.00810282.579
http://doi.org/10.5479/si.00810282.579...
).

dentate process by Alencar & Azevedo (2011bAlencar, I.D.C.C. & Azevedo, C.O. 2011b. Revision of Malagasy Odontepyris Kieffer (Hymenoptera, Bethylidae). Zootaxa, 2935: 26-40.); Azevedo et al. (2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Lim & Lee (2013Lim, J. & Lee, S. 2013. Taxonomy of the family Bethylidae (Hymenoptera: Chrysidoidea) from Cambodia and adjacent countries. I. genus Odontepyris Kieffer (Bethylidae: Bethylinae) with four new species and two new records. Journal of Natural History , 47(31-32): 1-22. http://doi.org/10.1080/00222933.2012.763057
http://doi.org/10.1080/00222933.2012.763...
); Polaszek & Krombein (1994Polaszek, A. & Krombein, K.V. 1994. The genera of Bethylinae (Hymenoptera; Bethylidae). Journal of Hymenoptera Research , 3: 91-105.).

dentate prominence (**dent aigüi) by Polaszek & Krombein (1994Polaszek, A. & Krombein, K.V. 1994. The genera of Bethylinae (Hymenoptera; Bethylidae). Journal of Hymenoptera Research , 3: 91-105.); Kieffer & Marshall (1904-1906Kieffer, J.J. & Marshall, T.A. 1904-1906. Proctotrypides. In: André, E. (Ed.). Species des Hyménoptères d’Europe & d’Algerie. Tome IX. Paris, Librairie Scientifique A. Hermann. p. 1-55, 21 pls. [Publication dates of the different parts: pages 1-64 1st January 1904; 65-144 1st April 1905; 145-288 1st November 1905; 289-368 31st January 1906; 369-551 1st October 1906].).

mesopleural dentate process by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Mesopleural epicoxal lobe (pl2el:Fig. 2F) = Posterior region of the mesopleuron, anteriorly delimited by the mesopleural epicoxal sulcus. It usually overlaps the base of the mesocoxa*. Criterion 1.

mesopleural epicoxal lobe by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.).

trochantial lobe by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

Mesopleural epicoxal ridge (pl2er) = Internal ridge correspondent to the mesopleural epicoxal sulcus (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

mesopleural epicoxal ridge by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Mesopleural epicoxal sulcus (pl2es:Fig. 2F) = The epicoxal sulcus located on the mesopleuron (HAO_0000560). Criterion 1.

mesopleural epicoxal sulcus by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

postpectal carina by Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

Mesopleural pit (pl2p:Fig. 3C) = The pit correspondent to the mesopleural apodeme (HAO_0001358). Criterion 1.

Figure 3
Mesopleuron. (A-B) Dissomphalus. (C) Prosierola. (D) Cleistepyris. (E) Pseudisobrachium. (F) Epyris. (A, C, D) lateral view. (B, E, F) internal view. Abbreviations: pl2pos = posterior oblique sulcus of the mesopleuron; sps = subpleural signum; pl2a = mesopleural apodeme; pl2por = posterior oblique ridge of the mesopleuron; saa = subalar area; sai = subalar impression; pl2p = mesopleural pit; st = sternaulus; pl2-ba2 = mesopleuron-mesobasalar; pl2-3ax2b = second mesopleuro-third axillary sclerite of forewing muscle; pl2-3ax2c = third mesopleuro-third axillary sclerite of forewing muscle; pl2-t2b = second mesopleuro-mesonotal muscle; pl2-cx2 = mesopleuro-mesocoxal muscle; pl2a-fu2 = anterior mesopleuro-mesofurcal muscle.

antero-central pit by Azevedo (2007Azevedo, C.O. 2007. The Provancher types of Bethylidae (Hymenoptera, Chrysidoidea). Boletim do Museu Biológico Mello Leitão, 22: 57-67.).

central pit by Alencar & Azevedo (2009Alencar, I.D.C.C. & Azevedo, C.O. 2009. Rediscovery of Planepyris Kieffer: A new synomym of Trachepyris Kieffer, (Hymenoptera, Bethylidae). Doriana, 8: 1-12.); Azevedo (1999bAzevedo, C.O. 1999b. Additions to the systematic of the bethylid fauna (Hymenoptera, Chrysidoidea) from Chile. Revista Brasileira de Zoologia, 16(1): 233-243., 2005Azevedo, C.O. 2005. A new species of Allobethylus (Hymenoptera, Bethylidae) from Australia, with a key to world species. Zootaxa, 1064: 25-30., 2006Azevedo, C.O. 2006. Two new genera of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae) with large scolebythid-like prostenums. Zootaxa, 1191: 35-47., 2008bAzevedo, C.O. 2008b. Synopsis of Prosierola (Hymenoptera: Bethylidae). Zootaxa, 1912: 45-58., 2009aAzevedo, C.O. 2009a. A new species of Solepyris Azevedo (Hymenoptera, Bethylidae) from Brazil, with amended diagnosis of the genus. Revista Brasileira de Entomologia, 53(3): 327-330. http://doi.org/10.1590/S0085-56262009000300002
http://doi.org/10.1590/S0085-56262009000...
, bAzevedo, C.O. 2009b. Synopsis of Lytopsenella Kieffer (Hymenoptera: Bethylidae). Zootaxa, 2286: 58-64., 2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
); Azevedo & Alencar (2010aAzevedo, C.O. & Alencar, I.D.C.C. 2010a. Rediscovery of the Afrotropical genus Trissepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Epyris Westwood. Journal of Afrotropical Zoology, 6: 3-7.); Azevedo & Barbosa (2010Azevedo, C.O. & Barbosa, D.N. 2010. Two new species Pararhabdepyris Gorbatovskii (Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa, 2668: 55-62.); Azevedo & Lanes (2009Azevedo, C.O. & Lanes, G.O. 2009. Cladistic assessment and redescription of Galodoxa torquata Nagy (Hymenoptera, Bethylidae), a striking species with swallow tailed metasomal sternite. Zoologische Mededelingen Leiden, 83(4): 841-851.); Barbosa & Azevedo (2011bBarbosa, D.N. & Azevedo, C.O. 2011b. Taxonomy of Allobethylus Kieffer (Hymenoptera, Bethylidae) from southeastern Asia. Journal of Asia-Pacific Entomology, 14: 89-94.); Corrêa & Azevedo (2002Corrêa, M.S. & Azevedo, C.O. 2002. Systematics of Epyris (Hymenoptera, Bethylidae) from Reserva Biológica de Duas Bocas, Espírito Santo, Brazil. Revista Brasileira de Entomologia , 46: 129-132. http://doi.org/10.1590/S0085-56262002000200004
http://doi.org/10.1590/S0085-56262002000...
); Gobbi & Azevedo (2006Gobbi, F.T. & Azevedo, C.O. 2006. The Brazilian Holepyris (Hymenoptera, Bethylidae), with description of a new species from Amazonian forest. Zootaxa, 1357: 61-68., 2014Gobbi, F.T. & Azevedo, C.O. 2014. Revision of Caloapenesia (Hymenoptera, Bethylidae), with description of sixteen new species. Zootaxa, 3860: 501-535.); Gupta et al. (2017Gupta, A.; Rajeshwari, S.K. & Azevedo C.O. 2017. Biology and description of Megaprosternum cleonarovorum sp. nov. (Hymenoptera: Bethylidae) a gregarious larval ectoparasitoid of Cleonaria bicolor Thomson (Coleoptera: Cerambycidae) from India. Zootaxa, 4237: 78-90.); Kawada & Azevedo (2003Kawada, R. & Azevedo, C.O. 2003. New species of Bakeriella (Hymenoptera, Bethylidae) from the New World and new data on geographic distribution. Revista Brasileira de Entomologia , 47(3): 461-468. http://doi.org/10.1590/S0085-56262003000300017
http://doi.org/10.1590/S0085-56262003000...
); Santos L.M. & Azevedo (2003Santos, L.M. dos & Azevedo, C.O. 2003. Taxonomy of Epyris (Hymenoptera, Bethylidae) from Paraná, Brazil. Iheringia, Série Zoológica , 93(2): 189-195. http://doi.org/10.1590/S0073-47212003000200010
http://doi.org/10.1590/S0073-47212003000...
); Vargas & Azevedo (2008Vargas, J.M. & Azevedo, C.O. 2008. Revision of Alongatepyris (Hymenoptera: Bethylidae) with descripition of a new species from Colombia. Revista Brasileira de Zoologia , 25(4): 843-846., 2016a, b); Waichert & Azevedo (2012Waichert, C. & Azevedo, C.O. 2012. The genus Prorops Waterson, 1923 (Hymenoptera, Bethylidae) from Madagascar. European Journal of Taxonomy , 16: 1-11.).

episternal scrobe by Michener (1944Michener, C.D. 1944. The Social Behavior of the Bees: A Comparative Study. Cambridge, Harvard University Press.); Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

mesopleural pit (or **die zentrale mesopleurale Grube) by Alencar & Azevedo (2011bAlencar, I.D.C.C. & Azevedo, C.O. 2011b. Revision of Malagasy Odontepyris Kieffer (Hymenoptera, Bethylidae). Zootaxa, 2935: 26-40., 2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.); Alencar et al. (2016Alencar, I.D.C.C.; Colombo, W.D. & Azevedo, C.O. 2016. Pristocera masii: rediscovery of the holotype and its transfer to Acrepyris (Hymenoptera: Bethylidae). Acta Entomologica Musei Nationalis Pragae, 56(2): 795-803.); Argaman (2003Argaman, Q. 2003. Generic synopsis of Mesitinae Kieffer, 1914 (Hymenoptera: Bethylidae). Entomofauna, 24: 61-95.); Azevedo & Mugrabi (2014Azevedo, C.O. & Mugrabi, D.F. 2014. Three new species of Tuberepyris Lanes et Azevedo (Hymenoptera, Bethylidae), with amended diagnosis of the genus. Zootaxa, 3794: 556-564.); Azevedo et al. (2018bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294., cAzevedo, C.O.; van Noort, S. & Notton, D.G, 2018c. Rediscovery of Disepyris semiruber (Kieffer) (Hymenoptera: Bethylidae) in Southern Africa, with first description of its male. Journal of Natural History, 52: 23-24); Evans (1969cEvans, H.E. 1969c. A revision of the genus Epyris in the Americas (Hymenoptera, Bethylidae). Transactions of the American Entomological Society, 95: 181-352.); Lanes & Azevedo (2008Lanes, G.O. & Azevedo, C.O. 2008. Phylogeny and taxonomy of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae). Insect Systematic and Evolution, 39: 55-86. http://doi.org/10.1163/187631208788784165
http://doi.org/10.1163/18763120878878416...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.); Ramos et al. (2014Ramos, M.S.; Perkovsky, E.E.; Rasnitsyn, A.P. & Azevedo, C.O. 2014. Revision of Bethylinae fossils (Hymenoptera: Bethylidae) from Baltic, Rovno and Oise amber, with comments on the Tertiary fauna of the subfamily. Neues Jahrbuch fur Geologie und Palaontologie , 271: 203-228.); Sorg (1988Sorg, M. 1988. Zur Phylogenie und Systematik der Bethylidae (Insecta, Hymenoptera, Chrysidoidea). Sonderveröffentlichungen des Geologisches Institut der Universität Köln, 63: 1-146.); Stein et al. (2011Stein, P.R.W.; Alencar I.D.C.C.; Barbosa, D.N. & Azevedo, C.O. 2011. Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species. Zootaxa, 3032: 17-32.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
); Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29., 2012).

mesopleural scrobe by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
).

pleural aphophysis by Snodgrass (1942Snodgrass, R.E. 1942. The skeleto-muscular mechanisms of the honeybee. Smithsonian Miscellaneous Collections, 103: 1-120.).

pleural pit by Masner, 1979Masner, L. 1979. Pleural morphology in scelionid wasps (Hymenoptera, Scelionidae) - an aid to higher classification. The Canadian Entomologist , 111(9): 1079-1087.; Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

small pit by (or **fossette) by Benoit (1952Benoit, P.L.G. 1952. Bethylidae (Hym.) nord africains nouveaux ou peu connus du Museo Civico di Storia Naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova, 1: 1-7., 1957Benoit, P.L.G. 1957. Hymenoptera - Bethylidae. Exploration du Parc National Albert. Mission G.F. De Witte, 1933-1935. Fascicule 88. Tervuren, Institut des Parcs Nationaux de Congo Belge. 57p.); Kieffer (1904Kieffer, J.J. 1904. Description de nouveaux Dryininae et Bethylinae du Musée Civique de Gênes. Annali del Museo Civico di Storia Naturale di Genova , Serie 3, 41: 351-412., 1905Kieffer, J.J. 1905. Description de nouveaux Proctotrypides exotiques avec une planche et une figure dans le texte. Annales de la Société Scientifique de Bruxelles, 29: 95-142., 1906Kieffer, J.J. 1906. Description de nouveaux Hyménoptères. Annales de la Société Scientifique de Bruxelles , 30: 113-178., 1908Kieffer, J.J. 1908. Nouveaux proctotrypides et cynipides d’Amérique recueillis par M. Baker. Annales de la Société Scientifique de Bruxelles , 32: 7-64., 1910Kieffer, J.J. 1910. Description de nouveaux microhyménoptères du Brésil. Annales de la Société Entomologique de France, 78: 287-348., 1911Kieffer, J.J. 1911. Cynipides et Béthylides de l’Afrique du sud (Hymenopt.). Annales de la Société Entomologique de France , 80: 451-462.).

Mesopleural wing process (pl2wp: figs. 28, 50 inGibson, 1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.) = Projection of the anterodorsal corner of the mesopleuron (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

mesopleural wing process by Gibson (1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.).

pleural wing process by Duncan (1939Duncan, C.D. 1939. A Contribution to The Biology of North American Vespine Wasps. Stanford University Publications, Biological Sciences, 8: 1-233.); Gibson (1993Gibson, G.A.P. 1993. Groundplan structure and homology of the pleuron in Hymenoptera based on a comparison of the skeletomusculature of Xyelidae (Hymenoptera) and Raphidiidae (Neuroptera). Memoirs of the Entomological Society of Canada, 125: 165-187.); Snodgrass (1909Snodgrass, R.E. 1909. The thorax of insects and the articulation of the wings. Proceedings of the United States National Museum, 36(1687): 511-595., 1935Snodgrass, R.E. 1935. Principles of insect morphology. New York, McGraw-Hill. http://doi.org/10.1093/aesa/28.3.408
http://doi.org/10.1093/aesa/28.3.408...
).

Posterior mesepimeral area (epm2pa:Fig. 2C) = The area extending along the posterior margin of the mesopleuron, which is delimited anteriorly by the mesepimeral sulcus (HAO_0000751). Criterion 1.

mesepimeral flange by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
).

posterior mesepimeral area by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Posterior oblique ridge of the mesopleuron (pl2por:Fig. 3B) = The internal oblique ridge corresponding to the posterior oblique sulcus of the mesopleuron (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

oblique mesopleural ridge by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

posterior oblique ridge of the mesopleuron by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Posterior oblique sulcus of the mesopleuron (pl2pos:Figs. 1F,3A, D) = Oblique sulcus arising close to mesopleural pit, which follows to the anteroventral direction (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

longitudinal groove (or **sillon longitudinal) by Kieffer (1910Kieffer, J.J. 1910. Description de nouveaux microhyménoptères du Brésil. Annales de la Société Entomologique de France, 78: 287-348.); Muesebeck (1934Muesebeck, C.F.W. 1934. Seven new species of Indian Bethylidae (Hymenoptera). Records of the Indian Museum, 36(2): 223-232.).

mesopleural sulcus by Barbosa & Azevedo (2011cBarbosa, D.N. & Azevedo, C.O. 2011c. Order Hymenoptera, family Bethylidae, subfamily Mesitiinae. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing . v. 4, p. 375-404., 2012Evans, H.E. 1967a. Notes on Mexican and Southwestern U.S. Bethylidae (Hymenoptera): Part I, Pristocerinae. Entomological News, 78: 13-23.).

oblique espisternal sulcus by Argaman (2003Argaman, Q. 2003. Generic synopsis of Mesitinae Kieffer, 1914 (Hymenoptera: Bethylidae). Entomofauna, 24: 61-95.).

oblique groove (or **sillon oblique) by Benoit (1956Benoit, P.L.G. 1956. Contributions à l’étude de la faune entomologique du Ruanda-Urundi (Mission P. Basilewsky 1953). Hymenoptera: CXII. Bethylidae. Annales Musei Royal du Congo Belge. Tervure. Sciences Geologiques, 51: 560-564., 1957Benoit, P.L.G. 1957. Hymenoptera - Bethylidae. Exploration du Parc National Albert. Mission G.F. De Witte, 1933-1935. Fascicule 88. Tervuren, Institut des Parcs Nationaux de Congo Belge. 57p., 1968Benoit, P.L.G. 1968 Contribution à l’étude des espèces africaines du genre Mesitius Spinola (Hym. Bethylidae). Revue de Zoologie et de Botanique Africaines, 77: 83-94.); Evans (1964Evans, H.E. 1964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bulletin of the Museum of Comparative Zoology , 132: 1-222., 1966aEvans, H.E. 1966a. Further studies on neotropical Pristocerinae (Hymenoptera, Bethylidae). Acta Hymenopterologica, 2: 99-117.); Kieffer (1908Kieffer, J.J. 1908. Nouveaux proctotrypides et cynipides d’Amérique recueillis par M. Baker. Annales de la Société Scientifique de Bruxelles , 32: 7-64.).

oblique mesopleural sulcus by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

posterior oblique sulcus of the mesopleuron by Barbosa & Azevedo (2016Barbosa, D.N. & Azevedo, C.O. 2016. Australomesitius gen. nov. (Hymenoptera, Bethylidae: Mesitiinae), an unexpected new genus from Australia. Zootaxa, 4061: 447-450.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

posterior oblique suture by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
); Shcherbakov (1981Shcherbakov, D.E. 1981. Morphology the pterothoracic pleura in Hymenoptera. 2. Modifications of the groundplan. Zoologicheskii Zhurnal , 60(2): 205-213.).

transversal furrow by Gobbi & Azevedo (2016Gobbi, F.T. & Azevedo, C.O. 2016. Revision of Calobrachium Gobbi & Azevedo, gen. nov. (Hymenoptera, Bethylidae), with description of seven new Oriental species. Zootaxa, 4083: 221-238.).

transverse groove by Argaman (1990Argaman, Q. 1990. Generic synopsis of Sierolomorphidae (Hymenoptera). Israel Journal of Entomology, 24: 29-33.); Barbosa & Azevedo (2011cBarbosa, D.N. & Azevedo, C.O. 2011c. Order Hymenoptera, family Bethylidae, subfamily Mesitiinae. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing . v. 4, p. 375-404.).

Posterior subalar pit (psup:Fig. 1B) = The pit located posterodorsally on the mesopectus corresponding to the posterodorsal edge of the mesopleuron (HAO_0000961). Criterion 1.

posterior subalar pit by Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.).

subalar pitGibson (1985Gibson, G.A.P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. The Canadian Entomologist, 117(11): 1395-1443.); Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Krogmann & Vilhelmsen (2006Krogmann, L. & Vilhelmsen, L. 2006. Phylogenetic implications of the mesosomal skeleton in Chalcidoidea (Hymenoptera, Apocrita) - Tree searches in a jungle of homoplasy. Invertebrate Systematics, 20: 615-674. http://doi.org/10.1071/IS06012
http://doi.org/10.1071/IS06012...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Michener (1944Michener, C.D. 1944. The Social Behavior of the Bees: A Comparative Study. Cambridge, Harvard University Press.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Posterodorsal edge of the mesopleuron (pdem: figs. 12, 108 inMikó et al., 2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.) = The edge located at the intersection of the mesepimeral ridge and the inflected dorsal margin of the mesopleuron (HAO_0000771). Criterion 1.

posterodorsal edge of the mesopleuron by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

process on mesepimeral ridge by Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.).

Sternaular elevation (ste:Fig. 2D) = Internal elevation of the mesopleuron corresponding to the sternaulus externally (*).

Sternaulus (st:Figs. 2C,3D) = The line or fovea marking the site of origin of the mesopleuro-mesobasalar muscle* (HAO_0000953). Criterion 1.

sternaulus by Castro et al. (2011Castro, C.; Nunes, J. & Penteado-Dias, A. 2011. A new species of Heredius Marsh 2002 (Hymenoptera: Braconidae: Doryctinae) from Brazil. Brazilian Journal of Biology, 71: 175-177.); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Wharton (2006Wharton, R.A. 2006. The species of Sternaulopius Fischer (Hymenoptera: Braconidae, Opiinae) and the braconid sternaulus. Journal of Hymenoptera Research , 15: 317-347.).

sub-anterior fovea by Alencar & Azevedo (2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12.); Stein et al. (2011Stein, P.R.W.; Alencar I.D.C.C.; Barbosa, D.N. & Azevedo, C.O. 2011. Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species. Zootaxa, 3032: 17-32.).

Subalar area (saa:Fig. 3C) = The area delimited by the mesepimeral inflection, corresponding to the subalar ridge externally, which bears the mesopleural wing process (HAO_0000960). Criterion 1.

subalar area by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Michener (1944Michener, C.D. 1944. The Social Behavior of the Bees: A Comparative Study. Cambridge, Harvard University Press.); Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.).

subtegular plate by Linsley & Michener (1939Linsley, E.G. & Michener, C.D. 1939. A generic revision of the North American Nomadidae (Hymenoptera). Transactions of the American Entomological Society , 65: 265-305.).

Subalar impression (sai:Figs. 2C,3C) = Depression anterodorsal of the mesopleuron that generally is posteriorly connected to the mesepimeral sulcus (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

acropleural fovea by Barbosa & Azevedo (2014aBarbosa, D.N. & Azevedo, C.O. 2014a. Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species. Zoologia, 31: 285-311.).

acropleural sulcus by Barbosa & Azevedo (2014aBarbosa, D.N. & Azevedo, C.O. 2014a. Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species. Zoologia, 31: 285-311., bBarbosa, D.N. & Azevedo, C.O. 2014b. Moczariella, a bizarre new genus of Mesitiinae (Hymenoptera, Bethylidae) from Arab Peninsula. Zootaxa, 3860: 291-297., 2015Barbosa, D.N. & Azevedo, C.O. 2015. Synopsis of Bradepyris Kieffer, 1905 (Hymenoptera, Bethylidae, Mesitiinae). European Journal of Taxonomy, 151: 1-16. http://doi.org/10.5852/ejt.2015.151
http://doi.org/10.5852/ejt.2015.151...
, 2016Barbosa, D.N. & Azevedo, C.O. 2016. Australomesitius gen. nov. (Hymenoptera, Bethylidae: Mesitiinae), an unexpected new genus from Australia. Zootaxa, 4061: 447-450.); Barbosa et al. (2017Barbosa, D.N.; Kawada, R. & Azevedo, C.O. 2017. Order Hymenoptera, family Bethylidae. Genus Glenosema, Kiefer. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Department of the President’s Affairs . v. 6, p. 322-335.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

sub-alar fovea by Barbosa & Azevedo (2009Barbosa, D.N. & Azevedo, C.O. 2009. Laelius Ashmead (Hymenoptera, Bethylidae) from Madagascar, with description of two new species. Zootaxa, 2170: 1-14., 2010Barbosa, D.N. & Azevedo, C.O. 2010. Order Hymenoptera, family Bethylidae, genus Laelius . In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing. v. 3, p. 412-421., 2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275.); Lim et al. (2010Lim, J.; Kwon, H.; Lee, J. & Lee, S. 2010. A new Laelius species (Hymenoptera: Bethylidae) from China with a key to the species in Eastern Palaearctic region. Journal of Asia-Pacific Entomology , 2010: 365-368. http://doi.org/10.1016/j.aspen.2010.07.001
http://doi.org/10.1016/j.aspen.2010.07.0...
).

subalar impression by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

subtegular fovea by Azevedo (2011Azevedo, C.O. 2011. Synopsis of Aspidepyris Evans, 1964 (Hymenoptera, Bethylidae). Zootaxa, 3016: 63-68.); Azevedo & Alencar (2010bAzevedo, C.O. & Alencar, I.D.C.C. 2010b. Rediscovery of Melanepyris (Hymenoptera: Bethylidae): a new synonym of Epyris. Revista Brasileira de Zoologia , 27(3): 403-407. http://doi.org/10.1590/S1984-46702010000300012
http://doi.org/10.1590/S1984-46702010000...
); Azevedo & Barbosa (2010Azevedo, C.O. & Barbosa, D.N. 2010. Two new species Pararhabdepyris Gorbatovskii (Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa, 2668: 55-62.); Azevedo & Lanes (2009Azevedo, C.O. & Lanes, G.O. 2009. Cladistic assessment and redescription of Galodoxa torquata Nagy (Hymenoptera, Bethylidae), a striking species with swallow tailed metasomal sternite. Zoologische Mededelingen Leiden, 83(4): 841-851.); Barbosa & Azevedo (2011bBarbosa, D.N. & Azevedo, C.O. 2011b. Taxonomy of Allobethylus Kieffer (Hymenoptera, Bethylidae) from southeastern Asia. Journal of Asia-Pacific Entomology, 14: 89-94., cBarbosa, D.N. & Azevedo, C.O. 2011c. Order Hymenoptera, family Bethylidae, subfamily Mesitiinae. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing . v. 4, p. 375-404., 2012Barbosa, D.N. & Azevedo, C.O. 2012. Revision of Mesitiinae (Hymenoptera, Bethylidae) from Madagascar, with description of eleven species. Zootaxa, 3417: 1-44.).

subtegular furrow by Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.);

subtegular groove by Alencar & Azevedo (2009Alencar, I.D.C.C. & Azevedo, C.O. 2009. Rediscovery of Planepyris Kieffer: A new synomym of Trachepyris Kieffer, (Hymenoptera, Bethylidae). Doriana, 8: 1-12., 2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12., bAlencar, I.D.C.C. & Azevedo, C.O. 2011b. Revision of Malagasy Odontepyris Kieffer (Hymenoptera, Bethylidae). Zootaxa, 2935: 26-40.); Alencar et al. (2016Alencar, I.D.C.C.; Colombo, W.D. & Azevedo, C.O. 2016. Pristocera masii: rediscovery of the holotype and its transfer to Acrepyris (Hymenoptera: Bethylidae). Acta Entomologica Musei Nationalis Pragae, 56(2): 795-803.); Azevedo (2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
); Azevedo & Alencar (2009Azevedo, C.O. & Alencar, I.D.C.C. 2009. Rediscovery of Pristepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Acrepyris Kieffer. Zootaxa, 2287: 45-54., 2010aAzevedo, C.O. & Alencar, I.D.C.C. 2010a. Rediscovery of the Afrotropical genus Trissepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Epyris Westwood. Journal of Afrotropical Zoology, 6: 3-7.); Azevedo & Mugrabi (2014Azevedo, C.O. & Mugrabi, D.F. 2014. Three new species of Tuberepyris Lanes et Azevedo (Hymenoptera, Bethylidae), with amended diagnosis of the genus. Zootaxa, 3794: 556-564.); Azevedo et al. (2018aAzevedo, C.O.; Alencar, I.D.C.C. & Colombo, W.D. 2018a. Pairs in copulation of the highly dimorphic genus Pristocera Klug (Hymenoptera, Bethylidae) from Madagascar solve taxonomic problems of male-female associations. Zootaxa, 4433: 1-49., bAzevedo, C.O.; Alencar, I.D.C.C.; Ramos, S.M.; Barbosa, D.N.; Colombo, W.D.; Vargas-R., J.M. & Lim, J. 2018b. Global Guide of the flat wasps (Hymenoptera: Bethylidae). Zootaxa, 4489: 1-294.); Lanes & Azevedo (2007Lanes, G.O. & Azevedo, C.O. 2007. Redescription and placement of the Oriental Scaphepyris rufus Kieffer (Hymenoptera: Bethylidae). Zootaxa, 1654: 55-60., 2008); Lim & Azevedo (2014Lim, J. & Azevedo, C.O. 2014. Five new species of Disepyris Kieffer, 1905 (Hymenoptera: Bethylidae), with emended generic diagnosis. Zootaxa, 3900: 526-540. http://doi.org/10.11646/zootaxa.3900.4.4
http://doi.org/10.11646/zootaxa.3900.4.4...
); Lim & Lee (2013Lim, J. & Lee, S. 2013. Taxonomy of the family Bethylidae (Hymenoptera: Chrysidoidea) from Cambodia and adjacent countries. I. genus Odontepyris Kieffer (Bethylidae: Bethylinae) with four new species and two new records. Journal of Natural History , 47(31-32): 1-22. http://doi.org/10.1080/00222933.2012.763057
http://doi.org/10.1080/00222933.2012.763...
).

sub-tegular groove by Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

subtegular groove of mesopleuron by Lanes & Azevedo (2008Lanes, G.O. & Azevedo, C.O. 2008. Phylogeny and taxonomy of Sclerodermini (Hymenoptera, Bethylidae, Epyrinae). Insect Systematic and Evolution, 39: 55-86. http://doi.org/10.1163/187631208788784165
http://doi.org/10.1163/18763120878878416...
).

subtegular mesopleural fovea by Azevedo (2014Azevedo, C.O. 2014. Synopsis of Bakeriella Kieffer, 1910 (Hymenoptera, Bethylidae). Zootaxa, 3878: 501-535. http://doi.org/10.11646/zootaxa.3878.6.1
http://doi.org/10.11646/zootaxa.3878.6.1...
).

Subalar tubercle (sat: see fig. 5A inKarlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
)) = The projection located posteriorly on the subalar area, which accommodates the posterior end of the second axillary sclerite of the forewing at the beginning of the upstroke of the wing (HAO_0001475). Criterion 1.

cup-shaped socket on pleuron by (HAO).

subalar tubercle by Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
).

Subpleural signum (sps:Figs. 3A, C) = The signum located submedially on the ventral face of the mesopleuron, which corresponds to the site of origin of the first mesopleuro-mesonotal (pl2-t2a) muscle (HAO_0000975). Criterion 3.

precoxal sulcus by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

precoxal suture by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
).

subpleural signum by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Michener (1944Michener, C.D. 1944. The Social Behavior of the Bees: A Comparative Study. Cambridge, Harvard University Press.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Upper mesopleural fovea (pl2uf:Figs. 1D, E) = Fovea located centrally in the mesopleura around the mesopleural pit (Lanes et al., 2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). Criterion 1.

mesopleural fovea by Alencar & Azevedo (2009Alencar, I.D.C.C. & Azevedo, C.O. 2009. Rediscovery of Planepyris Kieffer: A new synomym of Trachepyris Kieffer, (Hymenoptera, Bethylidae). Doriana, 8: 1-12., 2011bAlencar, I.D.C.C. & Azevedo, C.O. 2011b. Revision of Malagasy Odontepyris Kieffer (Hymenoptera, Bethylidae). Zootaxa, 2935: 26-40.); Azevedo & Alencar (2010aAzevedo, C.O. & Alencar, I.D.C.C. 2010a. Rediscovery of the Afrotropical genus Trissepyris Kieffer (Hymenoptera, Bethylidae), a new synonym of Epyris Westwood. Journal of Afrotropical Zoology, 6: 3-7.); Azevedo & Barbosa (2010Azevedo, C.O. & Barbosa, D.N. 2010. Two new species Pararhabdepyris Gorbatovskii (Hymenoptera, Bethylidae) from Australia and Thailand. Zootaxa, 2668: 55-62.); Azevedo & Lanes (2009Azevedo, C.O. & Lanes, G.O. 2009. Cladistic assessment and redescription of Galodoxa torquata Nagy (Hymenoptera, Bethylidae), a striking species with swallow tailed metasomal sternite. Zoologische Mededelingen Leiden, 83(4): 841-851.); Azevedo & Mugrabi (2014Azevedo, C.O. & Mugrabi, D.F. 2014. Three new species of Tuberepyris Lanes et Azevedo (Hymenoptera, Bethylidae), with amended diagnosis of the genus. Zootaxa, 3794: 556-564.); Barbosa & Azevedo (2010Barbosa, D.N. & Azevedo, C.O. 2010. Order Hymenoptera, family Bethylidae, genus Laelius . In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing. v. 3, p. 412-421., 2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275., bBarbosa, D.N. & Azevedo, C.O. 2011b. Taxonomy of Allobethylus Kieffer (Hymenoptera, Bethylidae) from southeastern Asia. Journal of Asia-Pacific Entomology, 14: 89-94.); Evans (1966bEvans, H.E. 1966b. A revision of the genus Anisepyris Kieffer (Hymenoptera, Bethylidae). Studia Entomologica, 9: 1-120., 1969cEvans, H.E. 1969c. A revision of the genus Epyris in the Americas (Hymenoptera, Bethylidae). Transactions of the American Entomological Society, 95: 181-352.); Lim & Lee (2013Lim, J. & Lee, S. 2013. Taxonomy of the family Bethylidae (Hymenoptera: Chrysidoidea) from Cambodia and adjacent countries. I. genus Odontepyris Kieffer (Bethylidae: Bethylinae) with four new species and two new records. Journal of Natural History , 47(31-32): 1-22. http://doi.org/10.1080/00222933.2012.763057
http://doi.org/10.1080/00222933.2012.763...
); Lim et al. (2010Lim, J.; Kwon, H.; Lee, J. & Lee, S. 2010. A new Laelius species (Hymenoptera: Bethylidae) from China with a key to the species in Eastern Palaearctic region. Journal of Asia-Pacific Entomology , 2010: 365-368. http://doi.org/10.1016/j.aspen.2010.07.001
http://doi.org/10.1016/j.aspen.2010.07.0...
); Stein & Azevedo (2007Stein, P.R.W. & Azevedo, C.O. 2007. Synopsis of lobatifrons species-group (Hymenoptera: Bethylidae, Rhabdepyris) from Neotropical region, with redefinition of the group. Zootaxa, 1444: 23-34.); Stein et al. (2011Stein, P.R.W.; Alencar I.D.C.C.; Barbosa, D.N. & Azevedo, C.O. 2011. Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species. Zootaxa, 3032: 17-32.); Vargas & Azevedo (2016aVargas, J.M. & Azevedo, C.O. 2016a. Three new species of Pararhabdepyris Gorbatovsky (Hymenoptera, Bethylidae) from Central Africa and the Arabian Peninsula. Zootaxa, 4117: 429-439. http://doi.org/10.11646/zootaxa.4117.3.10
http://doi.org/10.11646/zootaxa.4117.3.1...
); Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.); Zamprogno & Azevedo (2014Zamprogno, L.N. & Azevedo, C.O. 2014. Phylogeny and reclassification of Pristocera Klug (Hymenoptera: Bethylidae). Insect Systematics & Evolution , 45: 1-49. http://doi.org/10.1163/1876312X-04402003
http://doi.org/10.1163/1876312X-04402003...
).

oval fovea by Cameron (1905Cameron, P. 1905. On the phytophagous and parasitic Hymenoptera collected 1995 by Mr. E. Ernest Green in Ceylon. Spolia Zeylanica, 3: 67-97.).

pleural fovea by Gordh & Witethom (1995Gordh, G. & Witethom, B. 1995. A new species of Goniozus from Thailand (Hymenoptera: Bethylidae). Proceedings of the Hawaiian Entomological Society, 32: 145.).

upper fovea (or **fóvea superior) by Rosmann & Azevedo (2005Rosmann, W.L. & Azevedo, C.O. 2005. Dez espécies novas e notas sobre Rhabdepyris Kieffer (Hymenoptera, Bethylidae) da Mata Atlântica. Revista Brasileira de Entomologia , 49(3): 397-408.); Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.).

upper mesopleural fovea by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Musculature

Anterior mesopleuro-mesofurcal muscle (pl2a-fu2:Figs. 2D,3F) = The mesofurcal muscle that originates at the mesopleuron and inserts distally on the mesofurca (HAO_0000118). Criterion 1.

anterior mesopleuro-mesofurcal muscle by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

mesopleuro-mesofurcal muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

First mesopleuro-mesonotal muscle (pl2-t2a:http://api.hymao.org/public/figure/show_zoom/8424in HAO) = The mesonotal muscle that arises ventrally from the mesopectus and inserts on the mesoscutum laterally of the notauli and medially parascutal carina (HAO_0000332).

first mesopleuro-mesonotal muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
); Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
).

Mesocoxo-mesosubalar muscle (cx2-sa2:Fig. 2B) = The mesosubalare muscle that arises from the mesocoxa and inserts on the mesosubalare (HAO_0000544). Criterion 2.

mesocoxo-mesosubalar muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
); Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
).

mesocoxo-mesosubalare muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Mesopleuro-mesobasalar muscle (pl2-ba2:Figs. 3E, F) = The mesobasalar muscle that originates anteroventrally at the mesopleuron and inserts on the mesobalare (HAO_0000563). Criterion 1.

mesopleural-basalare muscle by Wharton (2006Wharton, R.A. 2006. The species of Sternaulopius Fischer (Hymenoptera: Braconidae, Opiinae) and the braconid sternaulus. Journal of Hymenoptera Research , 15: 317-347.).

mesopleuro-mesobasalar by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

mesopleuro-mesobasalar muscle by Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
).

mesopleuro-mesobasalare muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Mesopleuro-mesocoxal muscle (pl2-cx2:Figs. 2E,3F) = The mesocoxal muscle that arises from the mesopleuron and inserts on the mesocoxa (HAO_0000564). Criterion 1.

basalar muscle of the mesothorax by Snodgrass (1942Snodgrass, R.E. 1942. The skeleto-muscular mechanisms of the honeybee. Smithsonian Miscellaneous Collections, 103: 1-120.).

mesopleuro-mesocoxal by Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
).

mesopleuro-mesocoxal muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Prepecto-mesobasalar muscle (pre-ba2:http://api.hymao.org/public/figure/show_zoom/8930in HAO) = The mesobasalar muscle that arises from prepectus (HAO_0001805).

prepectus mesobasalarVilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Second mesopleuro-mesonotal muscle (pl2-t2b:Figs. 3E, F) = The mesonotal muscle that arises medially from mesopleuron and inserts on the ventral surface of the lateral axillar area (HAO_0000924). Criterion 1.

second mesopleuro-mesonotal muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
).

second mesopleuro-mesonotal by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

Second mesopleuro-third axillary sclerite of forewing muscle (pl2-3ax2b:Figs. 3E, F) = The fore wing third axillary muscle that arises anteroventrally from the mesopleuron (HAO_0001742). Criterion 1.

anterior mesopleuro-third axillary sclerite of forewing by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.); Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

second mesopleuro-third axillary sclerite of forewing by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

second mesopleuro-third axillary sclerite of forewing muscle by Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
).

Third mesopleuro-third axillary sclerite of forewing muscle (pl2-3ax2c:Figs. 3E, F) = The fore wing third axillary muscle that originates posterolaterally from the mesopleuron (HAO_0001214). Criterion 1.

posterior mesopleuro-third axillary sclerite of forewing muscle by Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

third mesopleuro-third axillary sclerite muscle by Vilhelmsen et al. (2010Vilhelmsen, L.; Mikó, I. & Krogmann, L. 2010. Beyond the wasp-waist: structural diversity and phylogenetic significance of the mesosoma in apocritan wasps (Insecta: Hymenoptera). Zoological Journal of the Linnean Society, 59: 22-194. http://doi.org/10.1111/j.1096-3642.2009.00576.x
http://doi.org/10.1111/j.1096-3642.2009....
).

third mesopleuro-third axillary sclerite of forewing by Willsch et al. (2020Willsch, M.; Friedrich, F.; Baum, D.; Jurisch, I. & Ohl, M. 2020. A comparative description of the mesosomal musculature in Sphecidae and Ampulicidae (Hymenoptera, Apoidea) using 3D techniques. Deutsche Entomologische Zeitschrift, 67: 51-67. http://doi.org/10.3897/DEZ.67.49493
http://doi.org/10.3897/DEZ.67.49493...
).

Axillary sclerites

First axillary sclerite of the forewing (1ax2: fig. 44 inGibson, 1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.) = The axillary sclerite that articulates anteriorly and medially with the anterior notal wing process laterally with the second axillary sclerite, and posteriorly with the posterior notal wing process (HAO_0001772). Criterion 1.

first axillary sclerite by Gibson (1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.); Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.); Snodgrass (1909Snodgrass, R.E. 1909. The thorax of insects and the articulation of the wings. Proceedings of the United States National Museum, 36(1687): 511-595.).

first axillary sclerite of the forewing by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Mesobasalare (ba2: figs. 9, 10, 12-16 inGibson, 1985Gibson, G.A.P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. The Canadian Entomologist, 117(11): 1395-1443.) = The basalare located in the mesothorax (HAO_0000542).

mesobasalare by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Mesosubalare (sa2:http://api.hymao.org/public/figure/show_zoom/16875in HAO) = The subalare located in the mesothorax (HAO_0000581).

mesosubalare by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.); Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.).

Second axillary sclerite of the forewing (2ax2: fig. 44 inGibson, 1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.) = The second axillary sclerite that is located on the mesothorax (HAO_0001673). Criterion 1.

second axillary sclerite by Gibson (1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.); Snodgrass (1909Snodgrass, R.E. 1909. The thorax of insects and the articulation of the wings. Proceedings of the United States National Museum, 36(1687): 511-595.).

second axillary sclerite of the forewing by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Third axillary sclerite of the forewing (3ax2: fig. 44 inGibson, 1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.) = The third axillary sclerite that is located on the mesothorax (HAO_0001632). Criterion 1.

third axillary sclerite by Gibson (1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.).

third axillary sclerite of the forewing by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

DISCUSSION

As proposed by Freixa (2006Freixa, J. 2006. Causes of denominative variation in terminology. A typology proposal. Terminology, 12: 51-77.), denominative variation in terminology responds to linguistic needs of different natures, being that they occur in a natural way and, sometimes, they can result from the enhancement about the interpretation of the studied objects (evolution of concepts). However, contrary to what was defended by Alves (2000Alves, I.A. 2000. Polissemia e homonímia em uma perspectiva terminológica. Alfa, Revista de Linguistica, 44: 261-272. https://periodicos.fclar.unesp.br/alfa/issue/view/300.
https://periodicos.fclar.unesp.br/alfa/i...
) and Martins (2015Martins, S.C. 2015. A sinonímia, a polissemia e a homonímia no vocabulário da fauna e da flora. Estudos Linguísticos, 44: 186-207.), we believe that, in some cases, the presence of synonyms, homonyms and polysemies can be considered problematic in specialized communication, especially when different denominations for the same concept are a product of imprecise and/or ambiguous definitions of conceptual boundaries (i.e., denominative variation caused by conceptual imprecision; see cognitive causes in Freixa, 2006Freixa, J. 2006. Causes of denominative variation in terminology. A typology proposal. Terminology, 12: 51-77.).

Based on the morphological analysis of several Bethylidae taxa and their respective descriptions and terminologies adopted, we verify that over the years, a large number of imprecise terms have been proposed to name the same mesopleural characters applied to Bethylidae or even to Hymenoptera. We believe that the huge number of terminological synonyms presented in the anatomical glossary can be evidence of that.

Many terms are not identified due to the conceptual imprecision regarding the structure to which it was referring, usually caused by incomplete descriptions and lack of illustrations. Among the characters identified, many are difficult to interpret and the factors that contribute to this are: the structural equivalence between the structure is difficult to establish, the lack of knowledge about what it represents morphologically and the lack of a clear delimitation of the structures in some cases (e.g., subalar impression, mesepimeral sulcus, sternaulus). Moreover, many areas and characters are little explored in taxonomic and cladistic contexts (e.g., anterior and posterior subalar pit, sternaulus, mesopleural epicoxal lobe and subalar area).

Synonymous and polysemous terms

In this topic we will discuss the main cases of the synonymous and polysemous terms found in Bethylidae literature (mainly) and, in some cases, groups of Hymenoptera.

In Bethylidae, the posterior oblique sulcus of the mesopleuron is present in Pristocerinae and Mesitiinae, and despite showing little variation in size, position and format, it has been labelled differently in both subfamilies (see Argaman, 2003Argaman, Q. 2003. Generic synopsis of Mesitinae Kieffer, 1914 (Hymenoptera: Bethylidae). Entomofauna, 24: 61-95.; Barbosa & Azevedo, 2011aBarbosa, D.N. & Azevedo, C.O. 2011a. Phylogeny and reclassification of Laelius (Hymenoptera, Bethylidae) and description of four new species. Insect Systematics & Evolution, 42: 237-275., bBarbosa, D.N. & Azevedo, C.O. 2011b. Taxonomy of Allobethylus Kieffer (Hymenoptera, Bethylidae) from southeastern Asia. Journal of Asia-Pacific Entomology, 14: 89-94.; 2012Barbosa, D.N. & Azevedo, C.O. 2012. Revision of Mesitiinae (Hymenoptera, Bethylidae) from Madagascar, with description of eleven species. Zootaxa, 3417: 1-44.; Gobbi & Azevedo, 2016Gobbi, F.T. & Azevedo, C.O. 2016. Revision of Calobrachium Gobbi & Azevedo, gen. nov. (Hymenoptera, Bethylidae), with description of seven new Oriental species. Zootaxa, 4083: 221-238.).

The use of names such as mesopleural sulcus (see Barbosa & Azevedo, 2011cBarbosa, D.N. & Azevedo, C.O. 2011c. Order Hymenoptera, family Bethylidae, subfamily Mesitiinae. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing . v. 4, p. 375-404.; 2012Barbosa, D.N. & Azevedo, C.O. 2012. Revision of Mesitiinae (Hymenoptera, Bethylidae) from Madagascar, with description of eleven species. Zootaxa, 3417: 1-44.), for example, can lead to misidentification with another sulcus present in the mesopleuron e.g., sometimes, the lower margin of the mesopleural fovea form a structure like a sulcus externally. Names such as transversal furrow (see Gobbi & Azevedo, 2016Gobbi, F.T. & Azevedo, C.O. 2016. Revision of Calobrachium Gobbi & Azevedo, gen. nov. (Hymenoptera, Bethylidae), with description of seven new Oriental species. Zootaxa, 4083: 221-238.) and other terms present in Bethylidae literature can make identification by systematists difficult, because in the name there is no inference about the location of the character, making it difficult to correlate the name and the structure it represents through literature, especially when interpreted by non-specialists in Bethylidae. Furthermore, this sulcus is also found in other Hymenoptera groups, e.g., Chrysididae and Pompilidae, and was termed posterior oblique sulcus of the mesopleuron by Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
), the same term established in Bethylidae as described by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.).

Establishing the correspondence of the structure can be difficult, even when there is little structural variation, as the mesopleural callus. This area was named by Evans (1961Evans, H.E. 1961. A revision of the genus Pseudisobrachium in the North and Central America (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology, 126: 211-318.) as “a more or less well defined swelling just below the base of the hind wings” to the genus Pseudisobrachium that belongs to Pristocerinae. Because of this, the term was widely used in workers of this subfamily. However, in some genera of the Epyrinae, this area is more accentuated, which took Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.) to name it as upper mesopleural elevation. Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.) standardized the term acropleural area to this region, but our muscular analysis is in disagreement with the definition presented by them (see definition proposed by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.) and our definition in the glossary section). Because of this, we will keep the term mesopleural callus (as proposed by Evans (1961Evans, H.E. 1961. A revision of the genus Pseudisobrachium in the North and Central America (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology, 126: 211-318.), but with a new conception. This terminological unification based on an anatomic study will make the works in Bethylidae safer and more understandable and it can serve as an area to be explored and studied in other groups of Hymenoptera.

The subalar impression has recently been proposed for Bethylidae and it has received many terms. Externally, this structure is fused posteriorly with mesepimeral sulcus, being considered and named as a single structure in the publications of the Bethylidae. Therefore, the terms found in the literature review refers to the subalar impression and part of the mesepimeral sulcus. Of all citations of this term, only the publications by Barbosa & Azevedo (2014aBarbosa, D.N. & Azevedo, C.O. 2014a. Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species. Zoologia, 31: 285-311., bBarbosa, D.N. & Azevedo, C.O. 2014b. Moczariella, a bizarre new genus of Mesitiinae (Hymenoptera, Bethylidae) from Arab Peninsula. Zootaxa, 3860: 291-297.; 2015Barbosa, D.N. & Azevedo, C.O. 2015. Synopsis of Bradepyris Kieffer, 1905 (Hymenoptera, Bethylidae, Mesitiinae). European Journal of Taxonomy, 151: 1-16. http://doi.org/10.5852/ejt.2015.151
http://doi.org/10.5852/ejt.2015.151...
; 2016Barbosa, D.N. & Azevedo, C.O. 2016. Australomesitius gen. nov. (Hymenoptera, Bethylidae: Mesitiinae), an unexpected new genus from Australia. Zootaxa, 4061: 447-450.), and Barbosa et al. (2017Barbosa, D.N.; Kawada, R. & Azevedo, C.O. 2017. Order Hymenoptera, family Bethylidae. Genus Glenosema, Kiefer. In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Department of the President’s Affairs . v. 6, p. 322-335.) seem to have followed the original proposition of Gibson (1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.), which used the term acropleural sulcus to this area.

The problems related to the terminology of the mesepimeral sulcus arises mainly from the lack of the knowledge and the misinterpretation of the type of structure it represents, whether a groove (e.g., episternal groove and mesopleural groove), suture (e.g., mesepimeral suture) or carina (e.g., postpectal carina) that do not represent the structure, more complete definitions and names about the structures can be found in Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.), Ronquist & Nordlander (1989Ronquist, F. & Nordlander, G. 1989. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). Entomologica Scandinavica, Supplement 33: 1-60.) and HAO. Despite these terms, the names mostly used in Bethylidae are episternal furrow and episternal groove, which was cited for the first time by Azevedo (2008aAzevedo, C.O. 2008a. Characterization of the types of the Neotropical Pseudisobrachium (Hymenoptera, Bethylidae), with a key to species. Revista Brasileira de Zoologia , 25(4): 737-801. http://doi.org/10.1590/S0101-81752008000400020
http://doi.org/10.1590/S0101-81752008000...
) and Barbosa & Azevedo (2010Barbosa, D.N. & Azevedo, C.O. 2010. Order Hymenoptera, family Bethylidae, genus Laelius . In: van Harten, A. (Ed.). Arthropod Fauna of UAE. Abu Dhabi, Dar Al Ummah Printing. v. 3, p. 412-421.). However, they did not define the term.

The mesopleural pit is an example of the lack of uniformity in terminology not just in Bethylidae, but also in other families of Hymenoptera (e.g., episternal scrobe to bees - Michener (1944Michener, C.D. 1944. The Social Behavior of the Bees: A Comparative Study. Cambridge, Harvard University Press.); pleural apophysis to bees - Snodgrass (1942Snodgrass, R.E. 1942. The skeleto-muscular mechanisms of the honeybee. Smithsonian Miscellaneous Collections, 103: 1-120.); mesopleural scrobe to Braconidae - Karlsson & Ronquist (2012Karlsson, D. & Ronquist, F. 2012. Skeletal morphology of Opius dissitus and Biosteres carbonarius (Hymenoptera: Braconidae), with a discussion of terminology. PLoS ONE, 7(4): e32573. http://doi.org/10.1371/journal.pone.0032573
http://doi.org/10.1371/journal.pone.0032...
); pleural pit Platygastroidea - Mikó et al. (2007Mikó, I.; Vilhelmsen, L.; Johnson, N.F.; Masner, L. & Pénzes, Z. 2007. Skeletomusculature of Scelionidae (Hymenoptera: Platygastroidea): head and mesosoma. Zootaxa, 1571: 1-78.)). Although there are no doubts about the identity of the mesopleural pit, several terms have been applied to this structure, which shows the importance in using the uniform terminology in taxonomy, cladistic and morphologic works. To adopt this practice can make the use of data from the literature more reliable and less laborious.

The lack of consistent criteria for proposing primary homologies (sensuDe Pinna, 1991De Pinna, M.C.C. 1991. Concepts and tests of homology in the Cladistic paradigm. Cladistics, 7: 367-394.) may be one of the factors that promotes proliferation of terminological polysemy or synonyms. Even when there is concern with the proposal of homologies through comparative anatomy, the superficial analysis of the characters is not always sufficient, since some of them may be externally similar, but internally they represent totally different structures. An example of the lack of the consistent criteria is the case of the “transepisternal line”. Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.) used the term “transepisternal line” to name a ventral sulcus of the lateral surface of the mesopleuron, but for Gibson (1985Gibson, G.A.P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. The Canadian Entomologist, 117(11): 1395-1443.) the “transepisternal line” marks the origin of muscles that does not originate from this sulcus, which, in fact, appears to be only the inferior margin of the lower mesopleural fovea.

Many works, mainly taxonomic, prioritize the description of the specimens, with no concern for recognizing whether the analyzed characters have already been described in other taxa that do not belong to the group of the study, which leads to the creation of new terms for such characters. In addition, terminological polysemy or synonyms may be the result of a lack of extensive research on the terminology (e.g., databases such as HAO and Antkey) and of the different motivations by systematics and morphologists. Thus, the stability in terms requires agreement as to which terms should be used to name the structures without affecting the scientific communication (Gibson, 1993Gibson, G.A.P. 1993. Groundplan structure and homology of the pleuron in Hymenoptera based on a comparison of the skeletomusculature of Xyelidae (Hymenoptera) and Raphidiidae (Neuroptera). Memoirs of the Entomological Society of Canada, 125: 165-187.).

The “subalar pit” is an example of the importance in agreement between systematics and morphologists. The mesopleuron has two small pits in the subalar region, anterior and posterior, however, in some taxa only one of the pits is present, so that Richards (1956Richards, O.W. 1956. An interpretation of the ventral region of the hymenopterous thorax. Proceedings of the Royal Entomological Society of London, 31: 99-104. http://doi.org/10.1111/j.1365-3032.1956.tb00216.x
http://doi.org/10.1111/j.1365-3032.1956....
) used the term “subalar pit” to name the anterior subalar pit, whereas Gibson (1985Gibson, G.A.P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. The Canadian Entomologist, 117(11): 1395-1443.) used the same term to name the posterior subalar pit.

Poorly explored characters

The study of the mesopleuron showed not only the large number of synonyms existing for its structures, but also revealed many little explored characters that can be useful in systematics.

Some of them are certainly under-explored because of the location and size of the structure, e.g., the anterior subalar pit, posterior subalar pit, subalar area and mesepimeral lobe. All of them are located at the anterior extremity of the mesopleuron, and are often difficult to observe because the wing generally covers them. They are small compared to other structures. In these cases, the state of preparation of the specimen can greatly influence observability, because the position of the wing can not only make it difficult to observe, but to correctly identify and interpret the structures. Thus, this area of mesopleuron probably is neglected due to the uncertainty of their homology, or because of the risk of damage to the specimen while studying them.

However, sternaulus and epicoxal mesopleural lobe are easy to observe, but little investigated in Bethylidae. The sternaulus, for example, was mentioned only three times for Bethylidae, as sub-anterior fovea by Alencar & Azevedo (2011aAlencar, I.D.C.C. & Azevedo, C.O. 2011a. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new synonym of Pristocera Klug, 1808. European Journal of Taxonomy, 4: 1-12.) and Stein et al. (2011Stein, P.R.W.; Alencar I.D.C.C.; Barbosa, D.N. & Azevedo, C.O. 2011. Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species. Zootaxa, 3032: 17-32.) and sternaulus by Lanes et al. (2020Lanes, G.O.; Kawada, R.; Azevedo, C.O. & Brothers, D.J. 2020. Revisited morphology applied for Systematics of flat wasps (Hymenoptera, Bethylidae). Zootaxa, 4752: 1-127.). The mesopleural epicoxal lobe was mentioned only once by Ramos & Azevedo (2012Ramos, M.S. & Azevedo, C.O. 2012. Revision of Eupsenella Westwood, 1874 (Hymenoptera, Bethylidae). Zootaxa, 3539: 1-80.). In our study we observed that these structures vary considerably and can be useful for the taxonomy of the group, as well as Hymenoptera.

Although we did not used any specific methodology to explicitly evaluate character usage throughout the literature, we believe that the little use of them may be related to some factors: (i) researchers generally follow standardized descriptions previously adopted, without considering their inherited descriptive and/or explanatory limitations for particular groups; (ii) researchers explore only a fixed set of characters determined by their historical usage in a particular group; and (iii) specialized literature that provides explicit information and explanation of characters is virtually absent for a particular group.

Negative effects of the absence of uniform terminology

The lack of a specific literature and the wide range of morphological variability lead specialists of different taxonomic groups to develop their terminologies independently, without standardized vocabularies, which generates many synonyms (Gibson, 1985Gibson, G.A.P. 1985. Some pro- and mesothoracic structures important for phylogenetic analysis of Hymenoptera, with a review of terms used for the structures. The Canadian Entomologist, 117(11): 1395-1443.; Seltmann et al., 2013Seltmann, K.C.; Pénzes, Z.; Yoder, M.J.; Bertone, M.A. & Deans, A.R. 2013. Utilizing descriptive statements from the biodiversity heritage library to expand the Hymenoptera anatomy ontology. PLoS ONE , 8(2): e55674. http://doi.org/10.1371/journal.pone.0055674
http://doi.org/10.1371/journal.pone.0055...
). This may result in the accumulation of terms that often are confusing or that do not represent the structure and uncertain homology, making the taxonomic and cladistic work mostly ineffective and constitutes a barrier to clear and accurate scientific communication.

According to Komarek & Beutel (2006Komarek, A. & Beutel, R.G. 2006. Problems in taxonomy and suggestions for a standardized description of new insect taxa. Entomological Problems, 36: 55-70.) insufficient morphological knowledge of the characters can result in imprecise or inadequate observations, and important features may be overlooked or misinterpreted. This shows the importance of morphological knowledge and consequently terminological standardization in a taxonomic context, especially to new taxonomists. At the beginning of their careers, many of them are unable to have a clear understanding of some structures, may replicate any eventual error or find the work dull and difficult, especially for under-studied taxonomic groups.

For cladistic analysis, the effects on the lack of uniformity of terminology can result in a character matrix with many problems, because the incorrect identification of the structure may not correspond to the primary hypotheses of homology. Examples of this problem can be found at Waichert & Azevedo (2009Waichert, C. & Azevedo, C.O. 2009. Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters. Zootaxa, 2284: 1-29.) and Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.). In the first situation, the term subtegular groove is used in their matrix of the characters, but it corresponds to two distinct structures, the subalar impression and mesepimeral sulcus, which are not delimited externally, and, therefore, seem to form a single structure. In the second situation, the terms supra-posterior callus and posterior elevation on upper fovea in Alencar & Azevedo (2013Alencar, I.D.C.C. & Azevedo, C.O. 2013. Reclassification of Epyrini (Hymenoptera: Bethylidae): a tribal approach with commentary on their genera. Systematic Entomology, 38: 45-80.), are used to name the same structure, present in different groups that is the mesopleural callus.

Furthermore, in groups with sexual dimorphism, some structures are difficult to correlate. Thus, the standardization of terms and determination of homologous regions among species is important to infer phylogenetic relationships (Gibson, 1986Gibson, G.A.P. 1986. Mesothoracic skeletomusculature and mechanics of flight and jumping in Eupelminae (Hymenoptera, Chalcidoidea: Eupelmidae). The Canadian Entomologist , 118(7): 691-728.).

CONCLUSION

The study of the mesopleuron of bethylids showed how much a study of certain regions of the body is necessary and how much it can not only add new characters, but also understanding about certain structures. The standardization of the terminology of the mesopleural structures will facilitate the study of the mesopleuron of Bethylidae, generating uniformity in future studies, which can serve for comparison and understanding not only within the family, but in other groups.

Even understanding the importance of terminological variation, especially when it relates to the improvement of concepts, we believe that the standardization of terminologies should be managed with due care, respecting recommended standards and established by consensus, considering terminological variation as part of the process.

Thereby, the efforts of The Hymenoptera Anatomy Ontology (HAOHymenoptera Anatomy Ontology (HAO). 2020. The Hymenoptera Glossary: Hymenoptera Anatomy Consortium. Available: Available: http://glossary.hymao.org . Access: 05/02/2021.
http://glossary.hymao.org...
) and many other morphological studies have contributed to reducing the problem related to this subject. Gibson (1993Gibson, G.A.P. 1993. Groundplan structure and homology of the pleuron in Hymenoptera based on a comparison of the skeletomusculature of Xyelidae (Hymenoptera) and Raphidiidae (Neuroptera). Memoirs of the Entomological Society of Canada, 125: 165-187.) stated that systematics and morphology are inextricably related because each is dependent on the other for its own success, which is increasingly necessary.

ACKNOWLEDGMENTS

We thank the curators of the collections for authorization to use the analyzed specimens. We thank Ricardo Kawada for loaning specimens for study. We thank the Estação Biológica de Santa Lúcia for allowing us to collect specimens used in this study. We are also very grateful to the subject editor Helena Onody and reviewers Lars Vilhelmsen and anonymous for the revision and suggestions regarding the manuscript. The first author was supported with a Ph.D. bursary of Demanda Social by CAPES (Coordenação de Aperfeiçoamento de Pessoal de Nível Superior); third author was supported with a bursary by CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico), INCT-CNPq HymPar grant # 303748/2018-4.

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  • Published with the financial support of the "Programa de Apoio às Publicações Científicas Períodicas da USP"

Edited by

Edited by: Helena Carolina Onody

Publication Dates

  • Publication in this collection
    28 July 2021
  • Date of issue
    2021

History

  • Received
    10 Feb 2021
  • Accepted
    04 May 2021
  • Published
    04 June 2021
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