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Phylogenetic Studies Ofthe Free-Livingandarthropod

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Phylogenetic Studies Ofthe Free-Livingandarthropod Gayana Zool. 57 (1): 21-46, 1993 ISSN 0016-531X PHYLOGENETIC STUDIES OFTHE FREE-LIVING ANDARTHROPOD ASSOCIATED LAELAPIDAE (ACARI: MESOSTIGMATA) ESTUDIO FILOGENETICO DE LAELAPIDAE DE VIDA UBRE YASOCIADOS CONARTRÓPODOS (ACARI: MESOSTIGMATA) María E. Casanueva* ABSTRACT The genus Urozercon is included for the first time in the family Laelapidae; and the new genus Gecarcinolaelaps The genera of free-living and arthropod-associated is established in Iphiopsididae. The prior ranking of Laelapidae were reviewed in an attempt to determine the Euvarroa and Varroa as the family Varroidae Delfinado internal relationships of the family. Using phylogenetic and Baker, 1974 is refuted. methods, with computations on the PAUP and MacClade programs, a comparative study of eighty-three characters Keywords: Acari, Mesostigmata, Laelapidae, Phylogeny. in fifty generic taxa resulted in the construction of nine consensus cladograms illustrating familial and subfami- lial relationships. The Leptolaelapidae was used as the RESUMEN sister group and Ologamasidae as the outgroup. The Laelapidae was redefined to include eight sub- Se estudiaron los géneros de la familia Laelapidae de families: Hypoaspidinae Vitzthum, 1940;Melittiphidinae vida libre y asociados con artrópodos en un intento para Evans and Till, 1966;Haemogamasinae Oudemans, 1926; determinar las relaciones internas de la familia. Utilizan- AlphalaelapinaeTipton, 1960;LaelapinaeBerlese, 1892; do métodos fdogenéticos, con los programas PAUP y Myonyssinae Bregetova, 1956; Hirstionyssinae Evans MacClade, se realizó un estudio comparativo de 83 and Tul, 1 966 and Mesolaelapinae Tenorio and Radovsky, caracteres en 50 géneros, obteniéndose nueve cladogra- 1974. The new arrangement of the free-living and arthro- mas de consenso que ilustran las relaciones a nivel de pod-associated genera is the following: Family Laelapidae, familia y subfamilias. Las familias Leptolaelapidae y with two subfamilies: Hypoaspidinae, which includes the Ologamasidae se utilizaron como el grupo hermano y free-living, predatory and phoretic mites on Coleóptera grupo externo respectivamente. and Blattaria, with Tribe Pseudoparasitini and Tribe Se redefíne la familia Laelapidae, la cual incluye Hypoaspidini, and Melittiphidinae which includes laela- ocho subfamilias: Hypoaspidinae Vitzhum, 1940; Melit- pid mites associates of Hymenoptera (Apidae and tiphidinae Evans y Till, 1966; Haemogamasinae Formicidae) and Isoptera, with Tribe Laelaspidini, new Oudemans, 1926; Alphalaelapinae Tipton, 1960; tribe, Tribe Melittiphidini and Tribe Varroini. LaelapinaeBerlese, 1892; Myonyssinae Bregetova, 1956; The dermanyssoid mites associated with Chilopoda, Hirstionyssinae Evans y Till, 1966 y, Mesolaelapinae Diplopoda, Araneae and crustácea are included in the Tenorio y Radovsky, 1 974. La posición de los géneros de famili Iphiopsididae, with two subfamilies: Scissura- la familia Laelapidae, de vida libre y asociados con laelapinae, new subfamily and Iphiopsidinae which is artrópodos es la siguiente: a, Subfamilia Hypoaspidinae divided into Tribe Iphiopsidini and Tribe Gecarci- que incluye los ácaros de vida libre, depredadores y nolaelapini, new tribe. foréticos en Coleóptera y Blattaria, con dos tribus: Pseudoparasitini y Hypoaspidini y, b, Subfamilia Melittiphidinae dividida en tres tribus: Laelaspidini, nue- va tribu, Melittiphidini y Varroini, las cuales incluyen los laelápidos asociados a Hymenoptera (Apidae y * Laboratorio de Acarología, Departamento de Zoología, Formicidae) e Isoptera. Universidad de Concepción. Casilla 2407- Apartado 10, Los ácaros dermanisoídeos asociados con Chilopoda, Concepción, Chile. Diplopoda, Araneae y Crustácea son incluidos en la 21 GayanaZool. 57(1), 1993 Familia Iphiopsididae, con dos subfamilias: Scissura- familia Laelapidae y, el género Gecarcinolaelaps en la laelapinae, nueva subfamilia y, Iphiopsidinae. Esta últi- familia Iphiopsididae. ma es dividida en dos tribus: lphiopsidini y Gecarcino- laelapini, nueva tribu. Palabras claves: Acari, Mesostigmata, Laelapidae, Se incluye por primera vez el género Urozercon en la Filogenia. INTRODUCTION be hypothesized) within a monophyletic super- family Dermanyssoidea? The present study empha- sizes the predaceous soil and arthropod-associated The dermanyssine family Laelapidae is com- genera to discover major patterns of morphological plex, species-rich, very diverse, and very poorly variation in the lesser derivative forms and thus understood. Hypotheses concerning the evolution- provide a comparative and phylogenetic basis for ary history of this family and its relatives are mini- study of the derivative groups. Finally, compari- mally developed and the classifícation of the group sons with the eviphidoid family Leptolaelapidae as is consequently inadequate. Early authors (e. g., the hypothesized sister group (nearest evolutionary Canestrini, Berlese) described many genera and relative) and with Ologamasidae as the outgroup of species especially from Europe. Subsequent work- Laelapidae have been done. Thus this study pro- ers have continued with the description of new vides an improved classifícation and better under- forms and, at the same time, have attempted to standing of the evolutionary relationships within formúlate comprehensive classifications. These this diverse group of Mesostigmatid mites. efforts have failed. These mites have been placed in a single family (Vitzthum, 1943; Baker and Strandtmann, 1948) or in a number of different MATERIAL AND METHODS families (Baker and Wharton, 1952; Bregetova, 1956; Rosario, 1981). The difficulty in classifíca- tion might be attributed to the wide range of ana- This study was based on specimens in the tomical forms as a result of diverse adaptation to Acarology Laboratory at The Ohio State Univer- parasitism and to the consequent concentration of sity, and in other museums. The total number of studies on parasites on vertebrates while neglecting genera in this family is ca. 100; of this, one half is the numerous free-living and arthropod-associated available in the collection at the Acarology La- species. The Laelapidae exhibits a fascinating range boratory. The other half was made available by the of habitats and associations, mostly not studied in following persons and their institutions: Dr. Torb- detail. jorn Kronestedt, Swedish Museum of Natural His- Among the Laelapidae are many genera that tory, Sweden; Dr. Evert E. Lindquist, Biosystematic developed ectoparasitic (nest-dwelling or host- Research Institute, Canadian National Collection, dwelling) associations with mammals (35 genera), Ottawa; Dr. David C. Lee, South Australian Mu- and also many free-living, soil-dwelling predatory seum; Dr. Marisa Castagnoli, Istituto Sperimentale forms (10 genera). Lesser derivative elements are per la Zoologia Agraria, Florence, Italy ; Dr. George soil inhabiting or nest-associated predators. The C. Eickwort, Dept. of Entomology, Cornell Uni- greatest anatomical and ecological diversity, how- versity; Dr. John Kethley, Field Museum of Natural ever, is seen in the arthropod-associated species (43 History; Dr. R. L. Smiley, Smithsonian Institution, genera). Here occur associates of Coleóptera (8 USA and Dr. Mercedes Delfinado-Baker, Benefi- mite genera), Blattaria (2 genera), Isoptera ( 1 genus), cial Insects Laboratory, USDA. Hymenoptera (21 genera), Araneae (1 genus), Efforts to borrow specimens from Russia and Myriapoda (7 genera) and terrestrial Crustácea (2 from the Rijksmuseum van natuurlijke Historie, genera). Leiden, Netherlands, British Museum of Natural From the viewpoint of phylogenetic systemat- History, England and from the Potchefstroom Uni- ics, this family and included taxa and its relatives versity Collection in South África were unsuccess- are not adequately characterized and understood. ful. As a result, I was unable to examine the follow- Can the Laelapidae be recognized as a monophy- ing type species of genera: Angosomaspis letic group (that is, can a unique evolutionary origin multisetosus Costa, 1971; Dynastaspis walhallae 22 Phylogenetic studies of the free-living and arthropod: Casanueva, M E. Costa, 1971; Holostaspis isotricha Kolenati, 1858; lapidae Karg, 1978 and Rhodacaridae Oudemans, Hypoaspis (Hypohasta) simplexans Womersley, 1902 (sensu Lee, 1970). 1956; Iphiolaelaps myriapoda Womersley, 1956; All the data obtained from the study were Iphis astronomicus Koch, 1839; Laelaspisella analyzed using the computational programs PAUP epyginalis Marais and Loots, 1969; Lucanaspis and MacClade. These programs are based in the 1977 brachypedes Costa, '; Myrmeciphis crawleianus parsimony principie. Hull, 1923 ; Promacrolaelaps hunteri Costa, 1971; Reticulolaelaps faini Costa, 1968; and Zontia meliponensis Turk, 1948. For all these, it was nec- RESULTS AND DISCUSSION essary to resort literature-based data. I. HlSTORICAL REMARKS In some taxa a complete account of morpho- logical characters could not be made because the The morphological heterogeneity exhibited specimens available were poorly mounted and I had by the group has encouraged taxonomic and sys- no permission to remount them, or broken and body tematic studies by several scientists who have at- parts were missing. Example of this are the follow- tempted to define it. Berlese ( 1 892) recognized two ing taxa: Angosomaspis, Cyclothorax, Euandro- families: Dermanyssidae Kolenati, 1859, which laelaps, Laelaspisella, Myrmeciphis, Myrmolaelaps, included the obligatory parasites, and Laelapidae Myrrnoleichus, Promacrolaelaps, Reticulolaelaps,
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