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Heterick, B. E., 2006 Reprinted from PCAS, ser. 4, vol. 57 (April 2006), with corrected Table of Contents PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series Volume 57, No. 3, pp. 69–202, 105 figs. April 18, 2006 A Revision of the Malagasy Ants Belonging to Genus Monomorium Mayr, 1855 (Hymenoptera: Formicidae) Brian Heterick1 Department of Entomology, California Academy of Sciences, 875 Howard Street, San Francisco CA 94103-3009 In this revision of the Malagasy Monomorium ant fauna, thirty-six species are recognized, nineteen of them here described as new. The new species are Monomorium adiastolon, sp. nov., M. aureorugosum, sp. nov., M. bifidoclypeatum, sp. nov., M. chnodes, sp. nov., M. clarinodis, sp. nov., M. denticulus, sp. nov., M. ferodens, sp. nov., M. fisheri, sp. nov., M. flavimembra, sp. nov., M. gongromos, sp. nov., M. infuscum, sp. nov., M. lepidum, sp. nov., M. micrommaton, sp. nov., M. nigricans, sp. nov., M. notorthotenes, sp. nov., M. platynodis, sp. nov., M. robertsoni, sp. nov., M. ver- sicolor, sp. nov., and M. xuthosoma, sp. nov. Five species (four described from African material) pass into synonymy: M. leopoldinum Forel 1905 with M. madecassum Forel 1892, M. binatu Bolton 1987, M. exchao Santschi 1926 and M. imerinense Forel 1892 with M. termitobium Forel 1892 and M. valtinum Bolton 1987 with M. hanneli Forel 1907. Seventeen species remain unchanged. Monomorium minutissimum Santschi 1937 is transferred from synonymy under Monomorium mictile Forel 1910 to become a junior synonym of Monomorium exiguum Forel 1894. The Malagasy species are dis- tributed among seven species groups. Previously recognized in the literature are the destructor-, hanneli-, latinode-, monomorium- and salomonis- groups. The hilde- brandti species group comprises six new species as well as those formerly assigned to the fossulatum species group. (Prior to this revision, fossulatum Emery 1895 was rel- egated to synonymy under sechellense Emery 1894, and hildebrandti Forel 1892 is the earliest described species in this group.) The shuckardi species group, here erected, appears to be unique to Madagascar, and comprises a group of four Malagasy Monomorium with several plesiomorphic characters. A key to workers of the 36 Malagasy Monomorium species is provided. TABLE OF CONTENTS Introduction . 70 Monomorium: A genus badly in need of reinterpretation . 70 Systematics Taxonomic background of Malagasy Monomorium . 71 Synonymic list of Malagasy species of Monomorium . 73 Materials and Methods Material examined. 75 Distribution data . 75 Sources of borrowed material . 75 Use of DELTA program for generating natural language descriptions. 75 1 Current address: Research Fellow, Department of Environmental Biology, Curtin University of Technology, GPO Box U 1987, Perth WA 6845, Australia 69 70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Fourth Series, Volume 57, No. 3 Genus Monomorium Mayr Diagnosis of worker of Malagasy species. 79 Diagnosis of queen of Malagasy species . 80 Diagnosis of male of Malagasy species . 81 Key to Malagasy Monomorium workers. 84 Malagasy species groups recognized as a result of this project: The shuckardi-group . 89 The destructor-group. 96 The salomonis-group. 100 The latinode-group . 108 The monomorium-group . 110 The hanneli-group . 153 The hildebrandti-group . 157 Acknowledgements . 181 Literature Cited . 181 Illustrations (Figs. 14–105): Distribution maps, Graphs, Automontage images. 187 INTRODUCTION Monomorium is one of the more significant genera of myrmicine ants. Not only does the genus have endemic representatives in all major land masses, but those of its species with tramp tenden- cies have been spread by human commerce to countries that do not have a native Monomorium fauna (McGlynn 1999) and in those countries can often be considered pests. In his Catalogue of the world’s ant species, Bolton (1995) listed a world total of 293 species for the genus. Since that time broad revisionary studies have added additional species: 43 have been described from Australia (Heterick 2001; Heterick 2003) and 30 from Arabia (Collingwood and Agosti 1996). Other research has also resulted in the description of new taxa, including that of two new Monomorium from Brazil (Fernandez, in press), two from China (Zhou 2001) and one from Japan (Terayama 1996) and southern Europe respectively (Radchenko 1997). Two species have also been transferred to genus Monomorium from Antichthonidris (synonymy Heterick 2001). Allowing for loss of other species- level names due to synonymy arising from the work of the abovementioned authors, the current world tally stands at 359 species. The classic text for information on the Monomorium species of tropical Africa has been Bolton’s (1987) monograph. In this work, Bolton alluded to one Malagasy Monomorium species- group with a very primitive palp formula (PF) of 5,3 and mentioned in his discussion of individual species several whose distribution included Madagascar. However, he made no attempt to provide a description of the indigenous Monomorium fauna of Madagascar, which lay outside the mono- graph’s terms of reference, in its own right. This work attempts to provide a comprehensive survey of the Monomorium species of Madagascar, and is part of an ongoing effort to inventory and describe the life forms of this fascinating and biodiverse large island. Keys are provided here to enable researchers to identify workers of endemic Malagasy Monomorium as well as those intro- duced to the island. MONOMORIUM: A GENUS BADLY IN NEED OF REINTERPRETATION.— Unlike the case with a num- ber of myrmicine genera that have a clearly defined diagnosis based on a few salient characters, the concept of the genus Monomorium in recent years has ballooned to the point where it has become unwieldy, and in great need of redefinition. The genus, as it stands at the moment, lacks clearly defined autapomorphies. A complete reappraisal of Monomorium is not possible in this paper, par- tially because this is intended as a monograph of a regional fauna rather than a review of the genus HETERICK: MONOMORIUM OF MADAGASCAR 71 on a world basis, and partially because such a redefinition will require examination of reproductive and worker castes of all the known species groups. Genitalic characters, particularly of the males, and generation of molecular sequences should be a mandatory part of the analysis. As it is at the moment, Monomorium has become almost a meta-genus that is continuing to swallow up previous- ly discrete genera. Since 1987 these have included Chelaner and Syllophopsis (Bolton 1987), Antichthonidris (Heterick 2001) and Epelysidris, Nothidris and Phacota (Fernandez, in press). I report in this work on a Malagasy species in which the worker has a PF of 3,2, a condition in Monomorium previously recorded only for males of two species formerly placed in Antichthonidris (Snelling 1975; Heterick 2001). Interestingly, this new species is not an isolated taxon, but evident- ly a member of a significant Afrotropical and Malagasy species group (the M. hildebrandti species group) in which all the other known members have a PF of 2,2. The change in the diagnosis of Monomorium brought about by this and other recent discoveries (e.g., see Heterick 2003, Fernandez, in press) means that the autonomy of the small solenopsidine genera Anillomyrma, Megalomyrmex and, possibly, Bondroitia is under threat. Genera linked with Solenopsis and with Allomerus, which have a broader suite of distinctive anatomical features than the abovementioned (see Bolton 1987 for details of shared characters), stand somewhat further apart from the current concept of Monomorium and are likely to remain so. SYSTEMATICS Taxonomic background of Malagasy Monomorium The first described Monomorium species, the tramp species now known as Monomorium pharaonis, was placed in the then portmanteau genus Formica as ‘Formica pharaonis’ (Linnaeus 1758). The genus Monomorium originated with Mayr in 1855. An account of the general history of the various generic and subgeneric names now synonymized under Monomorium is given in Bolton (1987) and so is not reproduced here. The later synonymization of Antichthonidris, Epelysidris, Nothidris and Phacota is mentioned above. Six taxa assigned to Monomorium have been described from material collected from Madagascar. Monomorium hildebrandti, Monomorium imerinense and Monomorium madecassum were described by Forel in the same publication as supposed races of Monomorium minutum Mayr (Forel 1892c). All three taxa were raised to full species by Dalla Torre (Dalla Torre 1893). Also in 1892, in a separate publication (Forel 1892b), Forel described Monomorium termitobium as a species in its own right. Monomorium shuckardi was published by this author three year later (Forel 1895). In 1922, Emery positioned this latter species under the subgenus Notomyrmex in which sim- ilar South American and Australian forms had already been placed. Notomyrmex was raised to genus by Kusnezov in 1957, but reduced to a synonym of Chelaner by Ettershank in 1966 (Chelaner becoming in turn a junior synonym of Monomorium with Bolton’s 1987 revision). In Ettershank’s work, non-Australasian taxa previously incorporated under Notomyrmex were separat- ed from the new genus of Chelaner, and the species shuckardi was returned to Monomorium. The final Monomorium species described from Madagascar was Monomorium sakalavum (Santschi 1928). In addition to the above taxa, four Monomorium species described from material collected elsewhere have been recorded from Madagascar,
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