Discussion of Phylogenetic Relationships
Scarabaeoid family and subfamily definitions were established by the beginning of this century as were some notions on the evolutionary progression of the taxa. However, these concepts were often intuitively derived (confirmation of many of these traditional ideas has subsequently occurred). The first tangible contributions towards a phylogeny of the Scarabaeoidea were provided by the numerous morphological studies of the superfamily. Apart from providing vital morphological data a rough evolutionary picture of the superfamily also emerged.
The first attempt to apply modern cladistic techniques to the analysis of relationships within the group as a whole was that of Howden (1982). This study, based on 39 characters from 17 taxa, focused on the phylogenetic position of Taurocerastinae (Geotrupidae) in relation to many other scarabaeoid taxa. The study did fulfill one of its major functions in stimulating further investigations of the relationships among the various higher taxa.
The next major study, that of Scholtz (1990), provided a comprehensive review of the available scarabeoid literature. Although the main purpose of this work was to make available a complete data set for future cladistic analysis, some discussion of evolutionary trends was also included.
Browne (1993) and Browne & Scholtz (1995) examined the evolution and morphology of the hind wing articulation, base and venation. Although this study examined all higher scarabaeoid taxa (13 families, most subfamilies, 250 genera) and all taxa identified by Scholtz (1990) as being of uncertain phylogenetic status for the first time, it was very limited in that the resulting phylogram was based on only three character complexes (73 characters). However, there is strong evidence that wing related characters are the most reliable of all structures in elucidating relationships among higher taxa (Kukalov?-Peck 1983).
To date the most comprehensive study is that of Browne & Scholtz (in preparation). They considered all major character suites, 134 characters, which supported most of the relationships presented by Browne (1993) and Browne & Scholtz (1995). The main conclusions of this study are given in the phylogram above.
Although relationships among many taxa are controversial, recent analyses, especially those of Howden (1982), Browne (1993), Scholtz et al. (1994), Browne & Scholtz (1995, in preparation) have resolved the following:
- Glaresidae is the sistergroup of the remaining scarabaeoids and likely reflects the ancestral scarabaeoid due to its large number of unusual plesiomorphies.
- Lucanidae and Diphyllostomatidae together form the sistergroup of the Passalidae.
- Geotrupidae (Geotrupinae, Taurocerastinae and Lethrinae), Hybosoridae, Ceratocanthidae and Ochodaeidae together form the sistergroup of Passalidae, Lucanidae, Diphyllostomatidae, Trogidae, Bolboceratidae, Pleocomidae and Glaphyridae. Together these taxa form the sistergroup of the Scarabaeidae.
However, the following are still heatedly debated:
- Monophyly of the Geotrupidae. Howden (1982) includes Bolboceratinae, Geotrupinae, Taurocerastinae and Lethrinae in this family. Scholtz & Browne (in press) present evidence that this family is polyphyletic and elevated Bolboceratinae to familial status.
- The position of Glaphyridae. Once accorded superfamily status (Hinton 1967) this family has more commonly been placed among the so-called "intermediate" scarabaeoids (Geotrupidae, Hybosoridae, Ceratocanthidae and Ochodaeidae)(Scholtz 1990). Recent evidence suggests that it is a member of the so-called "primitive" scarabaeoids (Passalidae, Lucanidae, Diphyllostomatidae, Trogidae, Bolboceratidae, Pleocomidae and Glaphyridae).
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D. Jonathan Browne
University of Cape Town, South Africa
Clarke H. Scholtz
University of Pretoria, South Africa
Page copyright © 1995 D. Jonathan Browne and Clarke H. Scholtz
Citing this page:
Browne, D. Jonathan and Clarke H. Scholtz. 1995. Scarabaeiformia. Scarabaeoidea. Scarabs, stag beetles, dung beetles, rain beetles, etc.. Version 01 January 1995 (under construction). http://tolweb.org/Scarabaeoidea/9077/1995.01.01in The Tree of Life Web Project, http://tolweb.org/
Три пальца. Дело было вовсе не и кольце, a в человеческой плоти. Танкадо не говорил, он показывал.