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This work is licensed under the Creative Commons Attribution-Noncommercial-Share Alike 3.0 United States License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-sa/3.0/us/ or send a letter to Creative Commons, 171 Second Street, Suite 300, San Francisco, California, 94105, USA. QUAESTIONES ENTOMOLOGICAE ISSN 0033-5037 A periodical record of entomological investigation published at the Department of Entomology, University of Alberta, Edmonton, Alberta. Volume 20 Number 3 1984 CONTENTS Ashe-Generic Revision of the Subtribe Gyrophaenina (Coleoptera: Staphylinidae: Aleocharinae) with Review of the Described Subgenera and Major Features of Evolution 129 GENERIC REVISION OF THE SUBTRIBE GYROPHAENINA (COLEOPTERA: STAPHYLINIDAE: ALEOCHARINAE) WITH A REVIEW OF THE DESCRIBED SUBGENERA AND MAJOR FEATURES OF EVOLUTION James S. Ashe Field Museum of Natural History Roosevelt Road at Lakeshore Drive Chicago, Illinois 60605 Quaestiones Entomologicae U. S. A. 20:129-349 1984 ABSTRACT The world genera of the subtribe Gyrophaenina are revised and described; subgenera are reviewed. Comparative morphological studies of adults reveal a great variety of characters available for taxonomic and phylogenetic study when gyrophaenines are examined in sufficient detail. Structures in the mouthparts, particularly the maxilla, proved especially useful. Illustrations of variation in structural features are provided. Gyrophaenines are inhabitants of polypore and gilled mushrooms, where both larvae and adults feed by scraping maturing spores, basidia, cystidea and hyphae from the hymenium surface. Known features of natural history of gyrophaenines are reviewed. Many of these features are related to unusual features of mushrooms as habitats. The subtribe is redefined, characterized, and larval characteristics are reviewed. The Gyrophaenina are shown to be monophyletic based on structure of the maxilla and spermatheca. Thirteen genera (11 previously described and two newly described) are recognized in the subtribe: Gyrophaena Mannerheim, Phanerota Casey, Eumicrota Casey, Encephalus Kirby, Probrachida n. gen. (type species Brachida modesta Sharp), Brachida Mulsant and Rey, Agaricochara Kraatz, Sternotropa Cameron, Pseudoligota Cameron, Neobrachida Cameron, Adelarthra Cameron, Brachychara Sharp, and Agaricomorpha new genus (type species Gyrophaena (Agaricochara) apacheana Seevers). Given for each genus are, as appropriate, synonymic list, diagnosis, description, discussion of nomenclatorial and taxonomic history, notes on natural history, general geographic distribution, and review of major literature. Based on analysis of transformation series of 47 characters, a cladistic analysis of the genera is provided. Gyrophaenina is hypothesized to be sister group to the subtribe Bolitocharina. Within the Gyrophaenina, three lineages can be recognized, arbitrarily and informally designated the "Brachida", 'Sternotropa" and "Gyrophaena" lineages. The "Brachida" lineage (Probrachida, BrachidaJ is hypothesized to be sister group to all other gyrophaenines, and the 'Sternotropa" lineage (Sternotropa, Pseudoligota, Adelarthra, Agaricomorpha, Brachychara, Neobrachida and probably Agaricochara,) and the "Gyrophaena" lineage (Eumicrota, Gyrophaena, Phanerotaj are hypothesized to be sister groups. Cladistic relationships of Encephalus cannot be determined at present. Analysis of distribution of gyrophaenines among major types of host mushrooms compared with structural features in mouthparts and overlaid on a cladistic analysis of genera and analysis of major patterns of host relationships suggest hypotheses about major 130 Ashe features of evolution ofgyrophaenines. At least two factors have had fundamental influence on evolution of relationships between gyrophaenines and mushrooms. First, evolution of mouthpart structures that allowed beetles to graze on the hymenium, rather than feed on fungal flesh, opened a relatively unused portion of the mushroom habitat. Second, general characteristics of the mushroom as a habitat require that members of each species evolutionarily optimize among conflicting requirements. These include: need to use every mushroom encountered, physiological limitations suggested by the great chemical and physical diversity of mushrooms, and physiological and competitive advantages expected from specialization. In resolving these conflicting requirements, gyrophaenines have evolved tolerance to a range of physical and chemical characteristics provided by mushrooms. This tolerance is reflected in an "acceptability spectrum" and allows members of a gyrophaenine species to respond to seasonal, yearly and geographic variation in the mushroom flora. Major habitat types found among mushrooms, from ephemeral gilled mushrooms to persistent polypores, can be considered to provide a series of adaptive zones for gyrophaenines. Increasing reliance on hymenium scraping as a feeding mode is reflected in changes in structure of the maxilla. Life cycle adaptations to the ephemeral nature of gilled mushrooms was probably involved in attainment of this adaptive zone. This has occurred only among members 0/Gyrophaena and Phanerota. Other gyrophaenines appear to be restricted to polypores or habits are not known. RESUME L'auteur presente une revision generique de lafaune mondiale de la sous-tribu des Gyrophaenina et passe en revue les sous-genres deja decrits. Une etude de morphologie comparee des adultes revilent un grand nombre de caracteres utiles pour la taxonomie et la phylogenie lorsque les Gyrophaenines sont examines suffisamment en detail. Les structures les plus utiles sont celles des pieces buccales, particulierement des maxilles. La variation des caracteres structuraux est illustree. Les Gyrophaenines habitent les polypores et les champignons a lamelles, dans lesquels larves et adultes se nourrissent en raclant les spores en maturation, les basides, les cystides et les hyphes se trouvant a la surface de Vhyminium. L'auteur revoit les aspects connus de I'histoire naturelle des Gyrophaenines. Plusieurs de ces aspects sont relies it des traits inusitis de I'habitat que representent les champignons. La sous-tribu est redefinie et caracterisee, et les caracteristiques des larves sont revues. La structure des maxilles et de la spermatheque indiquent que les Gyrophaenina forment un groupe monophyletique. L'auteur reconnait 13 genres dans la sous-tribu (11 decrits anterieurement et deux nouvellement decrits): Gyrophaena Mannerheim, Phanerota Casey, Eumicrota Casey, Encephalus Kirby, Probrachida n. gen. (genotype Brachida modesta Sharp), Brachida Mulsanl et Rey. Agaricochara Kraatz, Sternotropa Cameron, Pseudoligota Cameron, Neobrachida Cameron, Adelarthra Cameron, Brachychara Sharp, et Agaricomorpha n. gen. (genotype Gyrophaena (AgaricocharaJ apacheana Seevers). Les items suivants sont presente pour chaque genre, lorsqu'appropries: liste des synonymes, diagnose, description, discussion de I'histoire nomenclatoriale et taxonomique, notes sur I'histoire naturelle, grandes lignes de la repartition geographique et revue de la litterature principale. V'etude des series de transformations de 47 caracteres a servi de base a une analyse cladistique. L'hypothese est emise a I'effet que les Gyrophaenina forment le taxonfrere de la sous-tribu des Bolitocharina. Parmi les Gyrophaenina, trois lignees se distinguent et sone designees defacon arbitraire et informelle sous les noms de "Brachida", 'Sternotropa" et TJyrophaena". La lignee "Brachida" (comprenant les genres Probrachida et Brachida,) former ait le taxonfrere de tous les autres Gyrophaenines, et les lignees, 'Sternotropa" (incluant Sternotropa, Pseudoligota, Adelarthra, Agaricomorpha, Brachychara, Neobrachida et probablement AgaricocharaJ et "Gyrophaena" (comprenant Eumicrota, Gyrophaena et PhanerotaJ seraient taxonsfreres. II n'est presentement pas possible d'etablir les relations cladistiques rf'Encephalus. La distribution des Gyrophaenines parmi les principaux types de champignons-hotes est comparee avec les caracteristiques structurales des pieces buccales. Cette comparaison est superposee a une analyse cladistique des genres ainsi qu'a une analyse des principaux types de relations avec les botes, ce qui permet de formuler des hypotheses sur les principaux aspects de revolution des Gyrophaenines. Au moins deuxfacteurs ont eu une influence fondamentale sur revolution des relations entre les Gyrophaenines et les champignons. Premierement revolution de structures particulieres des pieces buccales, qui permit a ces Coleopteres de Generic Revision of the subtribe Gyrophaenina 131 brouter sur I'hymenium plutbt que de consommer la chair des champignons, a rendu possible Vexploitation d'une portion relativement inutilisZe de Vhabitat constitue par les champignons. Deuxiemement, les caracteristiques genkrales des champignons en tant qu'habitat requierent que les membres de chaque espece de Gyrophaenines soient adaptes pour satisfaire optimalement a des exigences incompatibles. Ces exigences comprennent: la necessity d'utiliser chaque champignon rencontre, les limitations physiologiques que suggere la grande diversite physique et chimique des champignons, et les avantages physiologiques et competitifs decoulant de la specialisation. Pour repondre a ces exigences incompatibles, les Gyrophaenines ont evolue une tolerance a une gamme de caracteristiques physiques et chimiques des champignons. Cette tolerance est refletee par la variete des champignons acceptables et permet aux membres des Gyrophaenines