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PHYLOGENY AND CLASSIFICATION OF EUPELMIDAE, WITH A REVISION OF THE WORLD GENERA OF CALOSOTINAE AND METAPELMATINAE (HYMENOPTERA: CHALCIDOIDEA)

Published online by Cambridge University Press:  31 May 2012

Gary A. P. Gibson*
Affiliation:
Biosystematics Research Centre, Agriculture Canada Research Branch, K.W. Neatby Building, Ottawa, Ontario, Canada K1A 0C6
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Abstract

Three subfamilies are classified in Eupelmidae: Calosotinae Bouček, Eupelminae Walker, and Metapelmatinae Bouček. Diagnoses of these three subfamilies and of Tanaostigmatidae are given, together with a key to distinguish members from each other and from other Chalcidoidea. Genera of Calosotinae and Metapelmatinae are revised for the world, with a key to genera given for both subfamilies. For each genus the following is provided: synonymy, description of structural features of males and females, notes on distribution and hosts, available keys to species listed by biogeographic region, and a catalog of species. New generic combinations are made in the catalogs of species based on examination of type specimens of the species. Eight genera are included in Calosotinae [type species in brackets]: Archaeopelma gen.nov. [A. tropeotergum sp.nov.], Licrooides gen.nov. [L. umbilicatus sp.nov.], Paraeusandalum gen.nov. [P. chilense sp.nov.], Eusandalum Ratzeburg, Chirolophus Haliday, Calosota Curtis, Balcha Walker, and Tanythorax gen.nov. [T. spinosus sp.nov.]. Four genera are included in Metapelmatinae: Metapelma Westwood, Neanastatus Girault, Eopelma gen.nov. [E. mystax sp.nov.], and Lambdobregma gen.nov. [L. schwarzii (Ashmead) comb.nov.]. The following are proposed as new synonymies: Notosandalum De Santis and Exosandalum Bouček = Eusandalum Ratzeburg, and Metacalosoter Masi = Calosota Curtis. Eighteen structural features of adults of the three eupelmid subfamilies, and of other Chalcidoidea including Tanaostigmatidae, Encyrtidae, Aphelinidae, and Pteromalidae, are studied to delimit character states and determine their distribution among the higher taxa. Twenty-two additional features of adults of Calosotinae and Metapelmatinae are studied for supplemental evidence of relationships among the genera in these two subfamilies. Observed character-state distributions are used to postulate character polarity and homoplastic states, and hypotheses of monophyly and relationships among taxa are based on proposed synapomorphic states. Aphelinidae sensu lato (including the subfamily Eriaporinae) are indicated as either a paraphyletic or polyphyletic taxon if the Eriaporinae are included but the Signiphoridae are excluded. Tanaostigmatidae sensu lato (including the genus Cynipencyrtus Ishii) are indicated as the sister group of Encyrtidae based on a relatively long mesoscutal process for the muscle pl2–t2c, and structure of the articulation between the mesoscutum and scutellar-axillar complex. The genus Cynipencyrtus Ishii is indicated to be most closely related to Encyrtidae based on common possession of transverse axillae and mesotibial apical pegs. Possible relationships among Calosotinae, Metapelmatinae, and Eupelminae, and among these and Tanaostigmatidae + Encyrtidae remain unresolved. There are no derived character states unique to either Eupelmidae, or Eupelmidae + (Tanaostigmatidae + Encyrtidae), so that these taxa and relationships are not definitively supported as monophyletic. Eupelmidae may represent a grade-level taxon with membership determined by similar suites of apomorphic states that function to enhance jumping ability. Cladograms are used to illustrate alternate hypotheses of character-state evolution and relationships among the genera of Calosotinae and Metapelmatinae. Distribution of character states for the higher taxa and for each genus of Calosotinae and Metapelmatinae is summarized in two tables. Scanning electron photomicrographs are used to illustrate structural features.

Résumé

Trois sous-familles sont classées parmi les Eupelmidae: les Calosotinae Bouček, les Eupelminae Walker et les Metapelmatinae Bouček. Une analyse de ces trois sous-familles et des Tanaostigmatidae est présentée, ainsi qu'une clé permettant de distinguer les membres entre eux et de les différencier des autres Chalcidoidea. Les genres des Calosotinae et des Metapelmatinae sont révisés à l'échelle mondiale, et une clé d'identification des genres donnés pour les deux sous-familles est présentée. Pour chaque genre, on fournit les renseignements suivants: les synonymes, la description des caractères structuraux des mâles et des femelles, des notes sur la distribution et les hôtes, les clés disponibles pour l'identification des espèces énumérées par région biogéographique et un catalogue d'espèces. Les nouvelles combinaisons de genres qui figurent dans la catalogue sont basées sur l'examen de spécimens type de l'espèce. Huit genres sont inclus dans les Calosotinae [espèce type entre crochets]: Archaeopelma gen.nov. [A. tropeotergum sp.nov.], Licrooides gen.nov. [L. umbilicatus sp.nov.], Paraeusandalum gen.nov. [P. chilense sp.nov.], Eusandalum Ratzeburg, Chirolophus Haliday, Calosota Curtis, Balcha Walker et Tanythorax gen.nov. [T. spinosus sp.nov.]. Quatre genres sont inclus dans les Metapelmatinae: Metapelma Westwood, Neanastatus Girault, Eupelma gen.nov. [E. mystax sp.nov.], et Lambdobregma gen.nov. [L. schwarzii (Ashmead) comb.nov.]. Les nouveaux synonymes suivants sont proposés: Notosandalum De Santis et Exosandalum Bouček = Eusandalum Ratzeburg, et Metacalosoter Masi = Calosota Curtis. Dix-huit caractères structuraux des adultes des trois sous-familles d'eupelmidés et d'autres Chalcidoidea, y compris les Tanaostigmatidae, les Encyrtidae, les Aphelinidae et les Pteromalidae sont étudiés afin de délimiter les caractéristiques et d'établir leur distribution parmi les taxons supérieurs. Vingt-deux autres caractères additionnels des adultes des Calosotinae et des Metapelmatinae sont étudiés afin d'obtenir d'autres preuves de relations entre les genres dans ces deux sous-familles. La distribution des caractéristiques qui a été établie sert à postuler la polarité de transformation des caractères et les caractères homoplastiques, tandis que les hypothèses de monophylétisme et de relations entre les taxons sont basées sur les synamorphies proposées. Les Aphelinidae au sens large (y compris la sous-famille des Eriaporinae) forme un taxon paraphylétique ou polyphylétique si les Eriaporinae en font partie et qu'en sont exclus les Signiphoridae. Les Tanaostigmatidae au sens large (y compris le genre Cynipencyrtus Ishii) forment un taxon frère des Encyrtidae en raison de la présence d'une apophyse mésoscutale relativement longue pour le muscle pl2–t2c, et de la structure de l'articulation entre le mésoscutum et le complex scutellaire-axillaire. Le genre Cynipencyrtus Ishii est le plus étroitement relié aux Encyrtidae en raison de la possession commune d'axilles transverses et d'organes sensoriels en forme de cheville à l'apex du mésotibia. Les relations possibles entre les Calosotinae, les Metapelmatinae et les Eupelminae, et entre ces derniers et les Tanaostigmatidae + Encyrtidae ne sont pas encore élucidées. Aucune caractéristique dérivée n'est unique aux eupelmidae ni aux Eupelmidae + (Tanaostigmatidae + Encyrtidae), aussi n'y a-t-il pas de preuve définitive à l'appui du monophylétisme de ces taxons et de ces relations. Les Eupelmidae peuvent représenter un taxon informel dont les membres présentent des séries semblables de caractères dérivés favorisant une meilleure aptitude au saut. Des cladogrammes sont utilisés pour illustrer les autres hypothèses sur l'évolution des caractéristiques et les relations entre les genres des Calosotinae et des Metapelmatinae. La distribution des caractéristiques pour les taxons supérieurs et pour chaque genre des Calosotinae et des Metapelmatinae est résumée dans deux tableaux. Des photographies prises au microscope électronique à balayage servent à illustrer les caractères structuraux.

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Research Article
Copyright
Copyright © Entomological Society of Canada 1989

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