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Primitive New Ants in Cretaceous Amber from Myanmar, New Jersey, and Canada (Hymenoptera: Formicidae)

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Primitive New Ants in Cretaceous Amber from Myanmar, New Jersey, and Canada (Hymenoptera: Formicidae) PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3485, 23 pp., 11 ®gures, 5 tables July 25, 2005 Primitive New Ants in Cretaceous Amber from Myanmar, New Jersey, and Canada (Hymenoptera: Formicidae) MICHAEL S. ENGEL1 AND DAVID A. GRIMALDI2 CONTENTS Abstract ....................................................................... 2 Introduction .................................................................... 2 Systematic Paleontology ......................................................... 5 Family Formicidae Latreille .................................................... 5 Subfamily ²Sphecomyrminae Wilson and Brown ............................... 5 ²Sphecomyrmodes, new genus ............................................. 5 Genus ²Sphecomyrma Wilson and Brown ................................... 7 Genus ²Haidomyrmex Dlussky ............................................ 11 Subfamily ²Brownimeciinae Bolton ......................................... 12 Genus ²Brownimecia Grimaldi, Agosti, and Carpenter ....................... 12 Subfamily Incertae Sedis ................................................... 13 ²Myanmyrma, new genus ................................................ 14 Subfamily Aneuretinae? Emery ............................................. 17 ²Cananeuretus, new genus ............................................... 17 Acknowledgments ............................................................. 20 References .................................................................... 20 Appendix ..................................................................... 22 1 Division of Invertebrate Zoology, American Museum of Natural History; Division of Entomology, Natural History Museum, and Department of Ecology and Evolutionary Biology, Snow Hall, 1460 Jayhawk Boulevard, University of Kansas, Lawrence, Kansas 66045±7523 ([email protected]). 2 Division of Invertebrate Zoology (Entomology), American Museum of Natural History ([email protected]). Copyright q American Museum of Natural History 2005 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3485 ABSTRACT New information is provided on the oldest fossil ants (Formicidae), including the description of a new species of ²Sphecomyrma (²Sphecomyrminae), a new genus of sphecomyrmines, a new genus of apparent myrmeciines, and a new genus of apparent aneuretines. New material from New Jersey amber (Turonian) includes workers of ²Sphecomyrma freyi Wilson and Brown preserved together in the same piece of amber, a worker of an unidenti®able ²Sphe- comyrma species, and a worker of ²Brownimecia clavata Grimaldi, Agosti, and Carpenter (²Brownimeciinae). A new species of ²Sphecomyrma in New Jersey amber is described and ®gured from a worker as ²S. mesaki, new species. Two worker specimens in Campanian amber from Canada are described, one of which is described as ²Cananeuretus occidentalis, new genus and species, and is tentatively placed in Aneuretinae. From Burmese amber (Albian- Cenomanian) are the oldest, de®nitive ants, along with ones in amber from Charente-Maritime of France (approximately contemporaneous in age). A new genus and species, allied to ²Sphe- comyrma, is described from these deposits as ²Sphecomyrmodes orientalis, along with a remarkable new ``poneroid'', ²Myanmyrma gracilis, new genus and species (Myrmeciinae?). A key to the species of ²Sphecomyrma is provided, the classi®cation of ants summarized, and the Cretaceous records of Formicidae brie¯y outlined. INTRODUCTION myrma freyi Wilson and Brown, in New Jer- sey amber. Since 1985, ants have been re- Ants can truly be said to shape the terres- ported in Cretaceous amber from Siberia, trial world. Of the 11,833 species of Formi- France, Canada, Burma, and additional new cidae3, many have a profound impact on nat- specimens and taxa in New Jersey amber (re- ural and manmade ecosystems, which is viewed in Grimaldi et al., 1997, with addi- made possible by their eusociality, frequently tions by Dlussky, 1999; Grimaldi and Agosti, large colony sizes, and abundance. There is 2000a; Grimaldi et al., 2002; Nel et al., 2004: scarcely a place outside of the polar regions where one cannot ®nd these insects or their vide appendix 1). Here we report important effects. Ants are among the most common new specimens of described ant taxa recently and diverse kind of insect in various Ceno- discovered in New Jersey amber, new species zoic deposits, and are particularly well of ²sphecomyrmines in both New Jersey and known in the fossilized faunas of Dominican, Burmese amber, as well as three new genera Sicilian, and Baltic ambers (Rasnitsyn and in Burmese and Canadian ambers (e.g., ®gs. Kulicka, 1990; Skalski and Veggiani, 1990; 1±3). While it is well established that New Wilson, 1985a), as are the less spectacularly Jersey amber is of Turonian age (Grimaldi et preserved compressions from LagerstaÈtte al., 2000) and that Canadian amber is Cam- such as Florissant (Carpenter, 1930), Green panian (Borkent, 1995), the dating of Bur- River (Dlussky and Rasnitsyn, 2003), and mese amber has been contentious. Formerly other Tertiary localities throughout the believed to be of Tertiary age, recent work world. Although Formicidae came into their has demonstrated that the Burmese deposit own during the Cenozoic, in the Mesozoic dates from the mid-Cretaceous (e.g., Zheri- and Early Tertiary (Paleocene) formicids khin and Ross, 2000; Grimaldi et al., 2002; were rare, and their earliest evolution has Cruickshank and Ko, 2003). Thus, those taxa been gradually unveiled with each new fossil in Burmese amber (Albian-Cenomanian) are discovery (e.g., Wilson et al., 1967; Wilson, among the current oldest records of ants 1985b; Grimaldi et al., 1997; Grimaldi and along with ants in amber of approximately Agosti, 2000a; Dlussky and Rasnitsyn, 2003; contemporaneous age from Charente-Mari- Dlussky et al., 2004; Nel et al., 2004). time, France (Nel et al., 2004), being at least Until 1985, the only true formicid known 8±10 million years older than previous re- from the Cretaceous Period was ²Spheco- cords of the family. In keeping with myrmecological tradition 3 Species total accurate as of 17 June 2005 (vide and literature, we generally use terminology www.antbase.org). for morphological structures (e.g., antennal 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 3 Figs. 1±3. Three Cretaceous amber ants. 1. ²Myanmyrma gracilis, new genus and species (AMNH Bu-014) in Burmese amber. 2. ²Cananeuretus occidentalis, new genus and species (TMP 8.89.7) in Canadian amber. 3. ²Sphecomyrmodes orientalis, new genus and species (AMNH Bu-351) in Burmese amber. parts) as outlined by HoÈlldobler and Wilson classi®cation, herein, noting in various places (1990) and Bolton (1994); however, for body where we believe it might eventually be regions we have used head, mesosoma (5 modi®ed. Material considered herein is de- alitrunk), and metasoma (5 petiole 1 gaster), posited in the Amber Collection, Division of as is standard in apocritan Hymenoptera. Invertebrate Zoology, American Museum of Throughout, ``head length'' is measured from Natural History, New York (AMNH); the the apex of the vertex to the anterior border Department of Palaeontology of the Natural of the clypeus. The higher classi®cation of History Museum, London (NHML); and the ants has recently undergone a major re- Royal Tyrell Museum of Palaeontology, arrangement owing to the work of Bolton Drumheller, Canada (TMP). Specimens in (2003) (summarized with minor modi®ca- the AMNH collection were embedded in ep- tions in table 1). We have employed that oxy for preparation and study, using the pro- 4 AMERICAN MUSEUM NOVITATES NO. 3485 TABLE 1 Hierarchical Suprageneric Classi®cation of the Ants (Formicidae) and Antlike Wasps (Armaniidae) (modi®ed from Bolton, 2003)a 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 5 cedure outlined by Nascimbene and Silver- (perhaps paraphyletic?) to Formicidae (vide stein (2000). table 1 and appendix 1). It is interesting to note that many primitive SYSTEMATIC PALEONTOLOGY ants have clypeal spicules with rounded api- ces (e.g., ²Sphecomyrmodes,²Myanmyrma; FAMILY FORMICIDAE LATREILLE Amblyoponinae). Apomyrma has similar DIAGNOSIS: Head prognathous; dorsal rim spicules, but these are located on the labrum of torulus often tuberculate or concealed un- rather than along the anterior clypeal margin. der vertical lamella of frons; antenna genic- The signi®cance of this trait is as of yet un- ulate. Primitively with anterior margin of clear (e.g., a ground-plan feature with nu- clypeus spiculate (apomorphically lost in merous, apomorphic losses; or functional many modern lineages: vide infra). Infrabuc- convergence). cal sac present between labium and hypo- pharynx. Pronotum with posterodorsal mar- SUBFAMILY ²SPHECOMYRMINAE gin weakly concave; posterolateral apex trun- WILSON AND BROWN cate anterior to tegula. Metapleural gland present in females, opening above metacoxa ²Sphecomyrmodes, new genus (rarely absent); meso- and metacoxae contig- TYPE SPECIES:²Sphecomyrmodes oriental- uous; inner metatibial spur modi®ed as cal- is, new species. car. Hind wing typically without jugal lobe DIAGNOSIS: Distinguished from all other (presence of the lobe is plesiomorphic within species of the tribe Sphecomyrmini by the the family and likely part of the familial minute, peglike denticles running along the ground plan). Metasoma petiolate; ®rst me- entirety of the anterior margin of the clypeus tasomal segment forming true node (strongly and from ²Sphecomyrma by
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