PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, , NY 10024 Number 3485, 23 pp., 11 ®gures, 5 tables July 25, 2005

Primitive New in from Myanmar, , and Canada (: Formicidae)

MICHAEL S. ENGEL1 AND DAVID A. GRIMALDI2

CONTENTS Abstract ...... 2 Introduction ...... 2 Systematic Paleontology ...... 5 Family Formicidae Latreille ...... 5 Subfamily ² Wilson and Brown ...... 5 ²Sphecomyrmodes, new ...... 5 Genus ² 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 ? 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 ᭧ American Museum of Natural History 2005 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3485

ABSTRACT New information is provided on the oldest ants (Formicidae), including the description of a new of ²Sphecomyrma (²Sphecomyrminae), a new genus of sphecomyrmines, a new genus of apparent myrmeciines, and a new genus of apparent aneuretines. New material from () 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 amber from Canada are described, one of which is described as ²Cananeuretus occidentalis, new genus and species, and is tentatively placed in Aneuretinae. From (Albian- ) 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 (?). 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 , 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 or their vide appendix 1). Here we report important effects. Ants are among the most common new specimens of described taxa recently and diverse kind of 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 (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 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 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, (ϭ modi®ed. Material considered herein is de- alitrunk), and (ϭ petiole ϩ 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 () (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 ). 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; 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- 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 the absence of constricted anteriorly and posteriorly); ®rst a medial extension or process on the clypeal metasomal sternum separated from second margin. metasomal sternum by deep constriction. ETYMOLOGY: The new genus-group name Morphologically distinct, sterile worker caste is a combination of ²Sphecomyrma, type ge- typically present4 (i.e., advanced eusocial); nus of the subfamily, and the suf®x±odes, reproductives typically macropterous, work- meaning ``with the form of''. The name is ers apterous. Workers with pronotum fused masculine. to mesothorax (a freely articulating prono- tum, present in some species, is plesio- ²Sphecomyrmodes orientalis, new species morphic and undoubtedly part of the formi- Figures 3±4 cid ground plan), remaining segments typi- cally fused. Species advanced eusocial. DIAGNOSIS: As for the genus (vide supra). COMMENTS: The ants, currently including DESCRIPTION: Head. Relatively large, 11,833 species (Bolton, 1995; www. height of head slightly less than length of antbase.org) have a cosmopolitan distribution alitrunk. Length of head 1.23 mm (with man- and are among the most recognizable of all dibles closed). No apparent microsculpture insects. Numerous species exist in Cenozoic on cuticle of head. Clypeus setose; setae of deposits around the world and are relatively moderate length and widely separated. Man- commonly encountered. Species of Creta- dible simple, with only two teeth; outer sur- ceous formicids, which are very rare, are face with numerous, widely scattered, ®ne brie¯y outlined in appendix 1. setae. Antenna of moderate length, with Bolton (2003) considered the antlike scape short, funicular article I (pedicel) wasps of the Cretaceous family Armaniidae shortest antennal article, funicular article II to represent the basalmost subfamily of the the longest article of funiculus. Lengths of ants. We have, however, retained armaniids antennal articles (in mm): scape 0.23, pedicel at the family rank and as the (funicular article [fa] I) 0.13, faII 0.32, faIII 0.15, faIV 0.15, faV 0.15, faVI 0.15, faVII 4 Some inquilines have apomorphically lost the work- 0.17, faVIII 0.17, faIX 0.17, faX 0.17, faXI er caste. 0.27. Mesosoma. Mesosomal length 1.33 6 AMERICAN MUSEUM NOVITATES NO. 3485

Fig. 4. Holotype worker of ²Sphecomyrmodes orientalis, new genus and species (AMNH Bu-351); general habitus, sting, clypeal margin and mandibles, and antenna. mm; without apparent microsculpturing, with ly curved inward at their extreme apices (vis- scattered ®ne, short setae on all visible sur- ible on left foreleg). Pairs of stiff setae on faces, those on about twice as ventral surface of protarsomeres: tarsomeres long as other setae. Coxae large, slightly in- I-III with three pairs, IV with two small ¯ated, ventrally setose, setae numerous and pairs. Pretarsal claw with minute subapical ®ne. Legs moderate length. Foreleg with tar- tooth. Metasoma. Attachment of petiole to somere I distinctly longer than combined propodeum not particularly thick; thickness lengths of more distal tarsomeres; tarsomere (measured in lateral view) of anterior end of I with ``antennal cleaner'' (strigil) a velvety petiolar peduncle 0.3ϫ greatest depth of pro- notch on ventral margin of proximal end; cal- podeum. Petiole length 0.38 mm, height 0.37 car present, length slightly longer than great- mm. Preserved portion of gaster 1.73 mm in est width of profemur, ventral margin with length. Integument without apparent micros- row of ®ne teeth and (apically) setae. Patch culpturing, with scattered, ®ne setae. Distal- of dense, elongate setae opposite strigil on most metasomal segments torn apart, al- profemur; inner posterior surface of protibial though sting bulb and sting well preserved apex with three stout, spinelike setae minute- beneath metasoma (®g. 4). 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 7

TYPE MATERIAL: Holotype. AMNH Bu- ²Sphecomyrma mesaki, new species 351, an incompletely preserved worker in a Figure 5 piece of reddish-orange amber, from Myan- mar. Collected in Kachin state, Tanai village, DIAGNOSIS: Distinguished from all other on Ledo Road, 105 km NW Myitkyna, via species of the genus by the median portion Leeward Capital Corp., 1999. of the clypeus having a long ventral lobe, ETYMOLOGY: The speci®c epithet is the length of the clypeus through the lobe is Latin word orientalis, meaning ``of the east'' 0.46ϫ the greatest width of clypeus; clypeus and is a reference to this being the ®rst spe- setose; and scape very short (1.2ϫ the length cies of the tribe ²Sphecomyrmini (sensu Bol- of longest funicular article). The species can ton, 2003) recorded from Myanmar. be further distinguished from ²S. freyi (the other species of the genus in New Jersey am- ber) by broad, shallow scrobes at base of an- Genus ²Sphecomyrma Wilson and Brown tennae; an that is approximately 1/3 larg- ²Sphecomyrma Wilson and Brown, In Wilson et er and oval (vs. almost perfectly round in ²S. al., 1967: 8. Type species: ²Sphecomyrma freyi freyi); and a large head (length of head/ Wilson and Brown, 1967, monobasic and orig- length of mesosoma ϭ 0.83, vs. 0.65 in ²S. inal designation. freyi). DESCRIPTION: Petiole and gaster not pre- DIAGNOSIS: Scape short; funiculus long served, so only head, mesosoma, and legs and ®liform, about four times length of preserved. Head. Large, length of head scape; promesothoracic suture complete and slightly less than length of alitrunk. Length well developed; trochantellus absent; petiole of head 2.20 mm (with mandibles closed); with distinct, domed node widely separated width of head 1.95 mm; length of eye 0.66 from propodeum and remainder of metasoma mm. No microsculpture on cuticle of head. by deep constrictions; cuticle without sculp- Vertex with ®ne, sparse pilosity, setae ca. 0.2 turing, super®cial microscopic relief, with mm long. Ocelli present, median ocellus sit- scattered and sparse setae. uated just above dorsal tangent of compound COMMENTS: The genus, which is de®ned . Face bare. Bases of antennae situated largely by plesiomorphies, is doubtfully in shallow, broad scrobes; length of scrobe monophyletic. It contains three species: ²Sphecomyrma canadensis Wilson in Cana- about equal to length of scape and articulat- dian amber, and ²S. freyi Wilson and Brown ing base. Eyes well developed, bare, situated and a new species in New Jersey amber (vide well above bases of antennae; gena deep. infra). In addition, we provide information Lateral portions of clypeus quadrate; median from newly identi®ed material of ²S. freyi. portion distended into long ventral lobe that extends to ventral margin of closed right mandible. Clypeus setose, except on middle Key to Species of ²Sphecomyrma part. Mandibles simple, with only two teeth. (based on the worker caste) Antennae of moderate length, with scape 1. Anterior margin of clypeus with short, broad short, funicular article I (pedicel) shortest an- extension, surface with two, long setae at tennal article, funicular article II the longest most; compound eye round ...... 2 article of funiculus. Lengths of antennal ar- Ð Anterior margin of clypeus with long, medial ticles (in mm): scape 0.53, pedicel (funicular lobe (®g. 5), surface with numerous, long article [fa] I) 0.20, faII 0.43, faIII 0.35, faIV setae; compound eye oval [New Jersey am- 0.30, faV 0.23, faVI 0.23, faVII 0.30, faVIII ber] ...... 0.22, faIX 0.26, faX 0.30, faXI 0.33. Meso- .... ²S. mesaki Engel and Grimaldi, n.sp. soma. Mesosomal length 2.66 mm; without 2. Third antennal article slightly more than microsculpturing, except at posterolateral twice as long as second article [New Jersey amber] ...... margin of promesonotal suture, where eight ...... ²S. freyi Wilson and Brown ®ne grooves occur. Dome of promesonotum Ð Third antennal article about as long as second setose; several ®ne setae on dorsal surface of article [Canadian amber] ...... metanotum and propodeum. Coxae large, in- ...... ²S. canadensis Wilson ¯ated, ventrally setose. Attachment of petiole 8 AMERICAN MUSEUM NOVITATES NO. 3485

Fig. 5. Holotype worker of ²Sphecomyrma mesaki, new species (AMNH NJ-1023); lateral habitus and facial view. 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 9 to propodeum not particularly thick; thick- bedded and trimmed to 14 ϫ 15 ϫ 4 mm. ness (measured in lateral view) of anterior The piece contains two workers of ²S. freyi. end of petiolar peduncle 0.3x greatest depth DESCRIPTIVE NOTES: Both workers are of propodeum. Metapleural gland opening largely but not completely preserved. Spec- (MGO) and MG bulla obvious, situated on imen A has the frontal half of the head miss- posterolateral part of propodeum just above ing; most of the antenna is present except for metacoxa. MGO small, with groove running bases of the scapes; the entirety of the re- between it and extended almost to vental mainder of the body is preserved and the margin of propodeum. Legs moderate length. sting appears largely or fully extruded. Spec- Foreleg with tarsomere I slightly longer than imen B has the dorsal and apical part of the combined length of more distal tarsomeres; gaster missing, as well as portions of the tarsomere I with ``antennal cleaner'' (strigil) right hind leg; the anterior third of the head a velvety notch on ventral margin of proxi- is obscured by Schimmel and a bubble. mal end; calcar present, length slightly lon- Measurements of body: Width of head ger than greatest width of femur, ventral mar- (specimen B), 1.03 mm; length of head (B, gin with row of ®ne teeth and (apically) approximate), 1.20 mm; length of mesosoma hairs. Stiff setae on ventral surface of pro- 1.39 mm (B), 1. 40 mm (A); length of petiole tarsomeres: tarsomere I with 7 pairs, II with 0.29 mm (B), 0.33 mm (A); length of gaster 3 pairs, III with 3 small pairs, IV with 2 (A) 1.72 mm; length of extruded portion of small pairs. Pretarsal claw with subapical sting (A), 0.33 mm. Measurements of anten- tooth. Metasoma not preserved. nal articles (as measured for right antenna in TYPE MATERIAL: Holotype. AMNH NJ- A, left antenna in B) presented in table 2. 1023, an incompletely preserved worker in a Measurements of leg segmentation (as mea- piece of amber barely larger than the ant, sured on specimen A) presented in table 3. from Sayreville, New Jersey (Middlesex COMMENTS: The discovery of this piece of Co.), White Oaks outcrop, coll. Bob Mesak amber is highly signi®cant and addresses (®g. 5). The amber is an irregularly shaped questions of the social behavior of primitive drop, 7 ϫ 5 ϫ 4 mm, made of clear yellow ants like ²Sphecomyrma. As reviewed in Gri- amber. Portions of some appendages are maldi et al. (1997), Dlussky (1987, 1988), breached at the surface, and the petiole and and Poinar et al. (1999) questioned whether gaster were similarly lost at the surface. ²Sphecomyrma was a true ant since it had Since very little amber exists between the ant such a short scape [but see response to Poin- and surface of the amber piece, no trimming ar et al. (1999) by Grimaldi and Agosti or polishing was possible. Still, details (2000b)]. With such a short scape, Dlussky through the rough surface are highly visible argued that it would be impossible for by immersing the piece in glycerine. ²Sphecomyrma to antennate, and therefore it ETYMOLOGY: The speci®c epithet is a pat- was considered highly unlikely for ²Sphe- ronymic for Bob Mesak, who collected and comyrma to have been social. Among the ap- donated this valuable specimen to the proximately 1700 fossiliferous pieces of New AMNH. Jersey amber in the AMNH collection thus far, four pieces contain worker or male sphe- ²Sphecomyrma freyi Wilson and Brown comyrmine ants. These ants are (and proba- Figure 6 bly originally were) exceedingly rare, and the probability that two workers would be pre- ²Sphecomyrma freyi Wilson and Brown, In Wil- served in one piece by chance alone is ex- son et al., 1967: 8. Grimaldi et al., 1997: 12 tremely remote. It is most parsimonious to (redescription of some features, new specimens, neotype). explain the occurrence of two workers in the same piece as a result of social behavior. MATERIAL: AMNH NJ-943, in amber from New Jersey: Middlesex Co., Sayreville, ²Sphecomyrma sp. White Oaks outcrop, collected by Keith Luz- zi (®g. 6). The piece is transparent yellow, MATERIAL: AMNH NJ-942, a male in am- originally much larger than now; was em- ber from New Jersey: Middlesex Co., Say- 10 AMERICAN MUSEUM NOVITATES NO. 3485

Fig. 6. Two workers of ²Sphecomyrma freyi Wilson and Brown preserved in a single piece of New Jersey amber (AMNH NJ-943). reville, White Oaks outcrop. Collected by the COMMENTS: Venation and other details of late Steve Swolensky. Specimen is in a clear this specimen are indistinguishable from yellow piece of amber 5 ϫ 7 mm, embedded AMNH NJ-242, described and ®gured by Gri- in epoxy and trimmed to 1.5 mm thickness maldi et al. (1997) as ²Sphecomyrma? sp. for a full lateral view of the ant. Measurements: Length of hind wing (forewing 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 11

TABLE 2 ulation with remainder of metasoma. Known Measurements of Antennal Articles for from the worker caste only. ²Sphecomyrma freyi (AMNH NJ-943) COMMENTS: The head of ²Haidomyrmex is enigmatic (®g. 7) and much of the detail is obscured below the clypeus and where the mandibles articulate. Indeed, it is likely that there has been signi®cant distortion at the manibular bases resulting in the rather ``deep'' appearance they have relative to the clypeus. The peculiar clypeal setae may have served a sensory function, perhaps as trigger hairs for the large mandibles, much the way gaff- shaped mandibles of various myrmecines and ``poneroids'' function. In those living ants long, ®ne, stiff trigger setae lie on the inside surface of the mandibles and on the oral mar- gin. ²Haidomyrmex has no such setae, so per- haps the clypeal brush functioned analogous- ly. How ²Haidomyrmex might have fed itself is an enigma. It is possible that this worker is similar to the major workers of Eciton, which cannot feed themselves. apices are lost), 1.97 mm; length of mesosoma 1.29 mm; length of petiole 0.36 mm; length of gaster 1.05 mm; length of antenna 2.40 mm. ²Haidomyrmex cerberus Dlussky Figures 7±8 Genus ²Haidomyrmex Dlussky ²Haidomyrmex cerberus Dlussky, 1996: 85. ²Haidomyrmex Dlussky, 1996: 84. Type species: ²Haidomyrmex cerberus Dlussky, 1996, mono- HOLOTYPE: NHML In.20182, partial work- basic and original designation. er specimen in amber from Myanmar. DIAGNOSIS: Clypeus a small, hemispheric COMMENTS: We have tentatively followed lobe lying just below antennal bases, pos- past authors in placing ²Haidomyrmex within sessing brush of ca. 60 ®ne, stiff, whitish se- ²Sphecomyrminae (e.g., Bolton, 2003). tae; setae are evenly arranged, those on ven- However, it is important to note the signi®- tral margin thickened at base but taper to a cant similarities between this genus and ®ne point apically. Compound eyes small, ²Brownimecia in New Jersey amber (vide in- length ca. 0.2ϫ length of head capsule; ocelli fra). Both genera have large, dome-shaped absent. Mandibles elongate, scimitar-shaped, heads, with relatively small compound eyes, without serrations or teeth. Propodeum lack ocelli, have large mandibles devoid of rounded in pro®le. Petiole one-segmented, serrations or dentition, and an elongate an- nodiform, with distinct constriction at artic- terior extension, or collar, of the pronotum.

TABLE 3 Leg Measurements (in mm) for ²Sphecomyrma freyi (AMNH NJ-943) 12 AMERICAN MUSEUM NOVITATES NO. 3485

Fig. 7. Facial view of holotype worker of ²Haidomyrmex cerberus Dlussky (NHML In.20182).

SUBFAMILY ²BROWNIMECIINAE BOLTON with oral surface bearing about 30 short, spi- culelike setae. Metasoma with slight but def- Genus ²Brownimecia Grimaldi, Agosti, inite constriction between second and third and Carpenter segment (also known as abdominal segments ²Brownimecia Grimaldi, Agosti, and Carpenter, III and IV; gastral segments 1 and 2). Known 1997: 20. Type species: ²Brownimecia clavata. from the worker caste only. Grimaldi, Agosti, and Carpenter, 1997, mono- basic and original designation. ²Brownimecia clavata Grimaldi, Agosti, DIAGNOSIS: Antenna distinctly clubbed, and Carpenter apical funicular article twice the width of ²Brownimecia clavata Grimaldi, Agosti, and Car- basal ones and pedicel. Ocelli absent. Man- penter, 1997: 20. dibles long, thin, scimitar-shaped, strongly cruciate, without teeth or crenulations, but MATERIAL: AMNH NJ-941, in amber from 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 13

Fig. 8. Lateral view of mesosoma and petiole, as preserved, of holotype worker of ²Haidomyrmex cerberus Dlussky (NHML In.20182).

New Jersey: Middlesex Co., Sayreville, clearly a plesiomorphic feature and part of White Oaks outcrop, collected by Steve the formicid ground plan. Swolensky. COMMENTS: A beautifully preserved spec- SUBFAMILY INCERTAE SEDIS imen in clear yellow amber, 5 ϫ 8 mm, which was embedded in epoxy and trimmed ²Myanmyrma, new genus to 2 mm thickness. Like the holotype which was described in 1997, it is curled up and the TYPE SPECIES:²Myanmyrma gracilis, new sting is extruded. The piece also contains species. some wood fragments and a frass pellet. The DIAGNOSIS: Gracile ``poneroid'' ant with length of head (including closed mandibles) extremely long legs; mandibles nearly equal 0.88 mm, greatest width of head 0.74 (be- to length of head, sickle-shaped, with a long, tween outer margins of eyes), length of eye apical sharp tooth and blunt subapical one; 0.24 mm, length of trunk 0.94 mm, length of clypeal margin bilobate, denticulate; long, petiole 0.32 mm. The mandibles are tightly spatulate genal process; antennal sockets in- closed, so details are less visible than in the clined, antennal funiculus very long, second holotype. The front portion of the head, how- article longest; articles slightly shorter api- ever, and especially the clypeus, are more cally; integument of head spinose; clypeus visible. The clypeus has ®ne, parallel cren- deeply incised along anterior margin and ulations along and perpendicular to the dor- with distinct peglike setae; eyes and ocelli sal margin of the clypeus. Most signi®cantly, not apparent (but may be present); pretarsal ocelli are de®nitively absent (rather than ves- claws with minute tooth; petiole with distinct tigial or minute). Three large ocelli are rare nodus; gastral constriction distinct; sting well in ants, with scattered occurrence (e.g., Cer- developed. Easily distinguished from ²Hai- apachyinae [Cylindromyrmex, Simopone], domyrmex, another highly modi®ed ant in [Aphophomyrmex, Notostigma, Burmese amber, by the mandibular and clyp- Alloformica, Cataglyphis], Myrmeciinae eal structure, head microsculpturing, meta- [Myrmecia]), otherwise the ocelli are repeat- somal constriction, long genal processes, and edly lost, reduced, or modi®ed secondarily very long legs. (as is true for ²Brownimecia). The large, ETYMOLOGY: Taken directly from Myan- well-developed ocelli of ²Sphecomyrma are mar, the country of the amber's origin; and± 14 AMERICAN MUSEUM NOVITATES NO. 3485 myrma, a common suf®x for ant genera. The tinate; ventral surface of metabasitarsus with name is feminine. ®ne pilosity, 10 pairs of setae. Metasoma. Petiole attachment to second metasomal seg- ²Myanmyrma gracilis, new species ment narrow; petiole with high, narrow node; Figures 1, 9 length of petiole 1.26 mm; posterior portion of petiole narrow, tubular; deep constriction DIAGNOSIS: As for the genus (vide supra). between second and third metasomal seg- DESCRIPTION: Specimen has been com- ments. Metasoma overall quite small, ca. pressed, but body form was clearly extremely 0.3x length of body (excluding antennae). gracile, with very long legs. Head. Propor- Terga and sterna telescoped in specimen, as tions dif®cult to determine due to compres- shown in ®gure 9; gaster (i.e., metasoma ex- sion; length ca. 1.76 mm. Front of head with cluding petiole) approximately 2.95 mm pair of carinae diverging dorsally; blunt long. Sting well developed, only partly ex- spines on ridges of carinae and on frons; truded but internally visible; pygidium with spines absent below ventral limit of carinae, long ®ne setae. with irregular cuticular foveae on clypeus. HOLOTYPE: AMNH Bu-014, a worker, in Ventral margin of clypeus with deep median amber from northern Myanmar (Burma) emargination; clypeus with two lobes, mar- (®gs. 1, 9). Collected in Kachin state, Tanai gin of two lobes with row of ca. 14 minute village, on Ledo Road, 105 km NW Myit- denticles on each lobe, labrum similarly bi- kyna, via Leeward Capital Corp., 1999. The lobed and denticulate. Lateral portion of ant is in a transparent yellow piece of amber, clypeus lobed, without denticles. Gena with which was originally more than 20 mm in spatulate process, exact length dif®cult to diameter and 30 mm in length. The piece was discern, but ca. 0.3ϫ length of mandible. occluded with fractures and debris, so it re- Mandibles long, 1.61 mm. (right one), nearly quired epoxy embedding and then trimming equal to length of head, sickle-shaped; apical to better observe the ant. The piece is now tooth long, sharp; blunt subapical tooth, no rhomboid-shaped, 17 ϫ 22 ϫ 4 mm, the other teeth. No ®ne setae apparent. Apex of broad surfaces being parallel to the lateral right mandible longer than left, especially surface of the ant. A ¯at edge also permits a apical tooth. Eyes well developed (dif®cult frontal view of the head. Other inclusions in to observe since they are sunken in a cavity), the piece are numerous frass pellets and length ca. 0.25ϫ length of head. Antenna wood fragments, further suggestive (besides very long (7.2 mm); scape fairly short, with body form) of arboreal/wood nesting habits wide base; funicular article (fa) 1 shortest, of the ant. The ant specimen shows a great fa2 longest; lengths of antennal articles (in deal of distortion due to compression. The mm): scape 0.63, funicular article (fa) [ped- cuticle is transparent, facilitating observation icel] I 0.44, faII 1.05, faIII 0.62, faIV 0.53, of some internal structures (such as unex- faV 0.61±faVI 0.45, faVII 0.40, faVIII 0.41, truded portions of the sting), but some com- faIX 0.48, faX 0.48, faXI 0.65. Mesosoma. pressed surfaces could easily be mistaken for Elongate, not deep, length ca. 3.52 mm; ¯anges and other structures. small opening to metapleural gland (MPG) ADDITIONAL MATERIAL: Two additional, apparent just above hind coxa. Legs extreme- fragmentary specimens are perhaps represen- ly long (vide measurements in table 4); tro- tative of ²M. gracilis; however, because of chantellus apparently absent. Protibia with their poor preservation, we hesitate to des- dorsal, preapical brush of ®ne setae; one ignate them as paratypes. Both are preserved large, two ®ner apical spurs; ventral surface in amber from northern Myanmar and with of probasitarsus with ®ne pilosity, six pairs the same collection data as the holotype. of stiff setae; all tarsomeres with four apical AMNH Bu-225, a poorly preserved worker spines; each pretarsal claw with median but very similar to holotype in observable tooth. Mesotibia with pair of ventroapical traits. AMNH Bu-1509, a badly compressed spurs; ventral surface of mesobasitarsus with worker preserved in yellow amber; head three rows of 10 stiff setae each; metatibia mostly crushed and dif®cult to discern, me- with pair of apical spurs, one larger and pec- tasoma similarly compressed; somewhat 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 15

Fig. 9. Holotype worker of ²Myanmyrma gracilis, new genus and species (AMNH Bu-014); facial view and general habitus. 16 AMERICAN MUSEUM NOVITATES NO. 3485

TABLE 4 Leg Measurementsa of Holotype Worker of ²Myanmyrma gracilis (in mm)

smaller than holotype and Bu-225 but still had similar habits. ²Haidomyrmex and exhibiting the same elongate, slender legs, ²Myanmyrma are clearly not closely related, constriction in ®rst gastral segments, etc., the former placed in the ²Sphecomyrminae this individual may represent a minor worker. by Dlussky. The type and only specimen of ETYMOLOGY: The speci®c epithet is the ²Haidomyrmex, however, has only the sec- Latin word gracilis, meaning ``slender'', as ond metasomal (i.e., ®rst gastral) segment a reference to elongate legs and structure of preserved, so the lack of a constriction pos- this ant. terior to this segment is suggested, not de®n- COMMENTS: Constriction of the metasoma itive (vide supra). indicates the specimen is a ``poneroid'', The lengths of basal articles of the antenna which is the third and oldest Cretaceous re- of ²Myanmyrma are highly signi®cant. Fu- cord of this paraphyletic grade of primitive nicular article 2 is the longest article in the ants (the poneroid grade includes the poner- funiculus, as is the case for most ²Spheco- omorph and myrmeciomorph subfamilies of myrminae (except ²S. canadensis Wilson). Bolton, 2003). The other Cretaceous ``pone- This condition does not exist in the other roids'' are ²Brownimecia clavata (in New Cretaceous ``poneroids'', ²Brownimecia and Jersey amber: Turonian), and ²Canapone ²Canapone. A long funicular article 2 was dentata Dlussky (in Canadian amber: Cam- proposed by Grimaldi et al. (1997: 8) as one panian). It is interesting that ²Myanmyrma is of only two or three possible synapomor- only the fourth record of ants in Burmese phies for the ²Sphecomyrminae. If truly apo- amber, but two of these records are for highly morphic, the long funicular article 2 would modi®ed species. ²Burmomyrma rossi Dlus- have been independently derived in ²Myan- sky is based on a single, alate, incomplete myrma. If a long funicular article 2 was ple- specimen, with head and portion of the ali- siomorphic, the monophyly of the ²Spheco- trunk missing. The most distinctive feature myrminae would be seriously doubtful. Out- of ²Burmomyrma is the wing with highly re- group evidence from closely related aculeate duced venation; thus, the taxonomic concept families suggests that, indeed, this feature is of this genus is not entirely comparable with symplesiomorphic for basal ants, including that of the other two Burmese amber genera. ²Myanmyrma. ²Burmomyrma, as discussed again later (vide Placement of ²Myanmyrma in a subfamily infra), has been tentatively placed in the is challenging. As noted, the metasomal con- Aneuretinae (Dlussky, 1996; Bolton, 2003). striction implies placement among the po- Both ²Haidomyrmex and ²Myanmyrma are neroid grade where it best approximates spe- extremely gracile and relatively large ants, cies of the (poneromorph) or the with large, highly modi®ed mouthparts and Myrmeciinae (myrmeciomorph). Neither genae. The extremely long legs and slender placement is entirely satisfactory, but we be- body are analogous to Leptomyrmex (Doli- lieve inclusion in the latter subfamily is more choderinae) and Oecophylla (Formicinae), likely (although we tentatively retain the ge- the latter of which is entirely arboreal. ²Hai- nus as subfamily incertae sedis). Like myr- domyrmex and ²Myanmyrma presumably meciines, ²Myanmyrma has a metasomal 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 17 constriction, the antennal sockets inclined to ment; sting present and well developed. nearly a vertical position, 12-segmented an- Known from the worker caste only. tennae, pectinate inner metatibial spur, and ETYMOLOGY: The new genus-group name elongate mandibles. Unlike typical myrme- is a combination of Canada, the country from ciines, however, the the new genus has man- which this amber originates, and Aneuretus, dibles that are sickle-shaped, with dentition type genus of the Aneuretinae. The name is only at the apices; has a deeply incised clyp- masculine. eal margin, with distinct spicules on the COMMENTS:²Cananeuretus might at ®rst lobes; has a spinose integument on the head; be confused for ²Eotapinoma, a dolichoder- has gracile legs; and elongate genal process- ine described from Canadian amber (Dlus- es. These traits are, however, autapomorphic sky, 1999). ²Cananeuretus differs notably and do not preclude placement in Myrmeci- from ²Eotapinoma in the presence of a well- inae. Within Myrmeciinae, ²Myanmyrma developed sting (apparently vestigial in would appear to be closest to the tribe Myr- ²Eotapinoma, like all dolichoderines), the meciini (sensu Bolton, 2003; Ward and Bra- more elongate petiole with a distinct nodus dy, 2003) as evidenced by the elongate sec- (shorter petiole, which does not have an ond funicular segment and distinctly two- elongate peduncle and without developed no- segmented waist. The latter trait is likely ple- dus in ²Eotapinoma), and the reduced com- siomorphic as it occurs in Myrmecia and pound eyes (large in ²Eotapinoma). weakly so in ²Prionomyrmex, perhaps being This new genus is tentatively placed in apomorphically lost in . The Aneuretinae, although the group shows most- presence of spicules along the clypeal margin ly what are presumed plesiomorphies for the in ²Myanmyrma is an interesting and enig- subfamily. The presence of a strong, well- matic feature that should be further explored developed sting and absence of an acidopore for its systematic implications. excludes placement in or This is the species that is referred to by Formicinae. However, the genus is best Grimaldi and Engel (2005) as ``undescribed placed among the formicomorph subfamilies Myrmeciinae?'' (sensu Bolton, 2003) where it most closely approximates the Aneuretinae. As more ma- SUBFAMILY ANEURETINAE?EMERY terial is recovered of these ants and a cladis- tic framework for fossil aneuretines ²Cananeuretus, new genus achieved, it is possible that ²Cananeuretus and others may prove to be stem-group TYPE SPECIES:²Cananeuretus occidentalis, Aneuretinae or even represent stem groups to new species. Aneuretinae ϩ Dolichoderinae. DIAGNOSIS: Compound eyes present, small; Today the subfamily Aneuretinae consists ocelli absent. Antennal sockets slightly in- of a single species occurring in Sri Lanka. clined; scape elongate. Mandible of primitive During the Tertiary aneuretines were distrib- construction, with four distinct teeth (®g. 10), uted widely in the Northern Hemisphere as basalmost tooth largest. Preoccipital carina evidenced by their occurrence in Baltic am- absent; ocelli absent. Alitrunk elongate, slen- ber (Wheeler, 1914) and Florissant, Colo- der; pronotal neck elongate. Meso- and me- rado (Carpenter, 1930). The mid-Cretaceous tatibia with a single spur; pretarsal claws genus ²Burmomyrma, in amber from Myan- simple. Propodeal lobes, denticles, and mar, has been tentatively placed in the spines absent; petiole one-segmented, ante- Aneuretinae (Dlussky, 1996; Bolton, 2003) rior peduncle elongate, with distinct tubular, and, until now, represented the sole Meso- postnodal section (more elongate than in zoic record for the subfamily. It is increas- Aneuretus) articulating high on anterior sur- ingly apparent that aneuretines were widely face of second metasomal segment, petiolar distributed during the Mesozoic and early tergum and sternum fused; nodus not very Tertiary, and that Aneuretus is a notable rel- high. Metasoma excluding petiole short and ict in Sri Lanka today. The signi®cant ex- globular; without U-shaped emargination on tinction of northern aneuretines was perhaps anterior margin of second metasomal seg- a result of the infamous - 18 AMERICAN MUSEUM NOVITATES NO. 3485

Fig. 10. Holotype worker of ²Cananeuretus occidentalis, new genus and species (TMP 8.89.7).

climatic shift, a cooling event that dramati- ²Cananeuretus occidentalis, new species cally altered the evolutionary history and Figures 2, 10±11 biogeography of numerous insect lineages that are today austral relicts (Grimaldi and DIAGNOSIS: As for the genus (vide supra). Engel, 2005). DESCRIPTION: Body with sparsely scattered, 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 19

Fig. 11. Partial worker of ²Cananeuretus occidentalis? (TMP 91.149.3).

®ne setae; integument ®nely imbricate. distinct nodus, and relatively long posterior, Head. Length ca. 0.52 mm; narrowed slight- tubular section, articulating high on second ly toward apex; vertex gently rounded. Man- metasomal segment; petiolar tergum and dibles with short, pointed teeth, basalmost sternum fused; metasoma without constric- apical tooth largest. Eyes present, relatively tion between second and third segments (i.e., small and low on head (dif®cult to observe). gastral segments 1 and 2); metasoma overall Scape fairly long (0.46 mm), narrowed quite globular. Sting well developed, extrud- slightly at base for articulation with bulb (®g. ed. 10); pedicel 0.65ϫ length of scape. Meso- HOLOTYPE: TMP 8.89.7, labeled, ``Grassy soma. Elongate, not deep, length ca. 3.10 Lake, Alberta, Campanian, Foremost For- mm; pronotum forming a distinct neck an- mation, Formicidae'' (®gs. 2, 10). Worker in teriorly. Legs slender and long; trochantellus amber that is embedded in a thin block of present (®g. 11). Inner surfaces of protibia epoxy and slide mounted. and probasitarsus with row of short, ®ne se- ADDITIONAL MATERIAL: TMP 91.149.3, la- tae forming weakly de®ned brushes (®g. 10); beled, ``Hymenoptera, Grassy Lake, Alberta, tarsomeres beyond basitarsus triangular, lat- Campanian, Foremost Formation, Formici- erally with short, distinct, stiff setae; pretar- dae'' (®g. 11). Worker in amber that is em- sal claws simple. Mesotibia and metatibia bedded in a thin block of epoxy and slide with a single spur; metatibial spur distinctly mounted. The specimen is partial and owing and minutely setose (®g. 10). Propodeum to some minor differences in the shape of the without lobes or spines, broadly rounded and petiole, we have considered it best to only gently sloping to articulation with petiole. tentatively assign it to this species. Metasoma. Petiole with elongate peduncle, ETYMOLOGY: The speci®c epithet is the 20 AMERICAN MUSEUM NOVITATES NO. 3485 term occidentalis, meaning ``of the west'', micidae) of Burmese amber. Paleontologiches- and is a reference to this being the ®rst aneu- kiy Zhurnal 3: 83±89. [In Russian] retine or aneuretine-like ant from the Meso- Dlussky, G.M. 1999. New ant taxa (Hymenoptera, zoic of the Western Hemisphere. Formicidae) from Canadian amber. Paleonto- logicheskiy Zhurnal 4: 73±76. [In Russian] Dlussky, G.M., D.J. Brothers, and A.P. Rasnitsyn. ACKNOWLEDGMENTS 2004. The ®rst ants (Hyme- noptera: Formicidae) from southern Africa, We are grateful to D. Agosti and P. S. with comments on the origin of the Myrmici- Ward for their invaluable reviews of the nae. Insect Systematics and Evolution 35: 1± manuscript and for correcting several inac- 13. curacies; to J. D. Gardner (TMP) for the loan Dlussky, G.M., and A.P. Rasnitsyn. 2003. Ants of specimens; and to A. J. Ross (NHML) for (Hymenoptera: Formicidae) of Formation hosting us during our 2002 visit to the De- Green River and some other middle Eocene de- partment of Palaeontology at which time the posits of North America. Russian Entomologi- holotype of ²Haidomyrmex was studied. This cal Journal 11: 411±436. work was supported by NSF DBI-9987372 Emery, C. 1895. Die Gattung Fab. und die systematische Eintheilung der Formiciden. (to DAG) and by NSF EPSCoR grant Zoologische JahrbuÈcher, Abtheilung fuÈr Syste- KAN29503 and NSF EF-0341724 (to MSE). matik, Geographie und Biologie der Thiere 8: This is contribution Nr. 3416 of the Division 685±778. of Entomology, Natural History Museum and Engel, M.S. 2005. Family-group names for Biodiversity Research Center, University of (Hymenoptera: ). American Museum Kansas. Novitates 3476: 1±33. Forel, A. 1892. Attini und Cryptocerini. Zwei REFERENCES neue Apterostigma-Arten. Mitteilungen der Schweizerischen Entomologischen Gesellschaft Ashmead, W.H. 1905. A skeleton of a new ar- 8: 344±349. rangement of the families, subfamilies, tribes Grimaldi, D., and D. Agosti. 2000a. A formicine and genera of the ants, or the superfamily For- in New Jersey Cretaceous amber (Hymenop- micoidea. Canadian Entomologist 37: 381±384. tera: Formicidae) and early evolution of the Bolton, B. 1994. Identi®cation guide to the ant ants. Proceedings of the National Academy of genera of the world. Cambridge: Harvard Uni- Sciences of the United States of America. 97: versity Press, 222 pp. 13678±13683. Bolton, B. 1995. A new general catalogue of the Grimaldi, D., and D. Agosti. 2000b. The oldest ants of the world. Cambridge: Harvard Univer- ants are Cretaceous, not Eocene: Comment. Ca- sity Press, 504 pp. nadian Entomologist 132: 691±693. Bolton, B. 2003. Synopsis and classi®cation of Grimaldi, D., D. Agosti, and J.M. Carpenter. Formicidae. Memoirs of the American Ento- 1997. New and rediscovered primitive ants mological Institute 71: 1±370. (Hymenoptera: Formicidae) in Cretaceous am- Borkent, A. 1995. Biting midges in the Creta- ber from New Jersey, and their phylogenetic ceous amber of North America (Diptera: Cer- relationships. American Museum Novitates atopogonidae). Leiden: Backhuys Publishers, 3208: 1±43. 237 pp. Grimaldi, D., and M.S. Engel. 2005. Evolution of Carpenter, F.M. 1930. The fossil ants of North the insects. Cambridge: Cambridge University America. Bulletin of the Museum of Compar- Press, xvϩ755 pp. ative Zoology 70: 1±66. Grimaldi, D., A. Shedrinsky, and T. Wampler. Cruickshank, R.D., and K. Ko. 2003. Geology of 2000. A remarkable deposit of fossiliferous am- an amber locality in the Hukawng Valley, ber from the Upper Cretaceous (Turonian) of northern Myanmar. Journal of Asian Earth Sci- New Jersey. In D. Grimaldi (editor), Studies on ences 21: 441±455. in amber, with particular reference to the Dlussky, G.M. 1987. New Formicoidea (Hyme- Cretaceous of New Jersey: 1±76. Leiden: Back- noptera) from Late Cretaceous. Paleontologi- huys Publishers, viii ϩ 498 pp. cheskiy Zhurnal 1987: 131±135. [In Russian] Grimaldi, D.A., M.S. Engel, and P.C. Nascimbe- Dlussky, G.M. 1988. Ants from the amber of Sa- ne. 2002. Fossiliferous Cretaceous amber from khalin (Paleocene?). Paleontologicheskiy Zhur- Myanmar (Burma): its rediscovery, biotic di- nal 1988: 50±61. [In Russian] versity, and paleontological signi®cance. Amer- Dlussky, G.M. 1996. Ants (Hymenoptera, For- ican Museum Novitates 3361: 1±72. 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 21

HoÈlldobler, B., and E.O. Wilson. 1990. The ants. overall history of the order. Prace Muzeum Zie- Cambridge: Harvard University Press, xiiϩ mi 41: 53±64. [1]ϩ732 pp. Rasnitsyn, A.P., and D.L.J. Quicke. 2002. History International Commission on Zoological Nomen- of insects. Dordecht: Kluwer, xii ϩ 517 pp. clature. 1999. International Code of Zoological Saux, C., B.L. Fisher, and G.S. Spicer. 2004. Dra- Nomenclature [4th ed.]. London: International cula ant phylogeny as inferred by nuclear 28S Trust for Zoological Nomenclature, xxixϩ rDNA sequences and implications for ant sys- [1]ϩ306 pp. tematics (Hymenoptera: Formicidae: Ambly- Jell, P.A. 2004. The fossil insects of Australia. oponinae). Molecular and Evo- Memoirs of the Queensland Museum 50(1): 1± lution 33: 457±468. 124. Skalski, A.W., and A. Veggiani. 1990. Fossil resin Jell, P.A., and P.M. Duncan. 1986. Invertebrates, in Sicily and the northern Apennines: geology mainly insects, from the freshwater, Lower Cre- and organic content. Prace Muzeum Ziemi 41: taceous, Koonwarra fossil bed (Korumberra 37±49. Group), South Gippsland, Victoria. Memoir of Snelling, R.R. In press. The identity of the fossil the Association of Australasian Paleontologists ant, ``Palaeosminthurus juliae'' Pierce and Gi- 3: 111±205. bron (Hymenoptera: Formicidae). Journal of Nascimbene, P., and H. Silverstein. 2000. The the Kansas Entomological Society. preparation of fragile Cretaceous ambers for Verhaagh, M. 1996. Warum die aÈlteste bekannte conservation and study of organismal inclu- Ameise keine ist. BeitraÈge der Hymenopterol- sions. In D. Grimaldi (editor), Studies on fossils ogen-Tagung in Stuttgart 1996: 11. Ward, P.S., and S.G. Brady. 2003. Phylogeny and in amber, with particular reference to the Cre- biogeography of the ant subfamily Myrmeci- taceous of New Jersey: 93±102. Leiden: Back- inae (Hymenoptera: Formicidae). Invertebrate huys Publishers, viii ϩ 498 pp. Systematics 17: 361±386. Naumann, I.D. 1993. The supposed Cretaceous Wheeler, W.M. 1914 [1915]. The ants of the Bal- ant Cretacoformica explicata Jell and Duncan tic amber. Schriften der Physikalisch-oÈkonom- (Hymenoptera). Journal of the Australian En- ischen Gesellschaft zu KoÈnigsberg 55: 1±142. tomological Society 32: 355±356. Wilson, E.O. 1985a. Invasion and in Nel, A., G. Perrault, V. Perrichot, and D. NeÂrau- the West Indian ant fauna: evidence from the deau. 2004. The oldest ant in the lower Creta- Dominican amber. Science 229: 265±267. ceous amber of Charente-Maritime (SW Wilson, E.O. 1985b. Ants from the Cretaceous France) (Insecta: Hymenoptera: Formicidae). and Eocene amber of North America. Psyche Geologica Acta 2(1): 23±29. 92: 205±216. Poinar, G., Jr., B. Archibald, and A. Brown. 1999. Wilson, E.O., F.M. Carpenter, and W.L. Brown, Jr. New amber deposit provides evidence of Early 1967. The ®rst Mesozoic ants, with the descrip- Paleogene , paleoclimates, and past tion of a new subfamily. Psyche 74: 1±19. distributions. Canadian Entomologist 131: 171± Zherikhin, V.V., and A.J. Ross. 2000. A review of 177. the history, geology and age of Burmese amber Rasnitsyn, A.P., and R. Kulicka. 1990. Hymenop- (Burmite). Bulletin of the Natural History Mu- teran insects in Baltic amber with respect to the seum London (Geology) 56: 3±10. 22 AMERICAN MUSEUM NOVITATES NO. 3485

APPENDIX Cretaceous Ants (Formicidae) and Related Taxa The following table summarizes the described species of ants and antlike wasps (i.e., Armaniidae) from the Cretaceous. The classi®catory structure generally follows that of Bolton (2003). The fossil Cretacoformica explicata Jell and Duncan (1986) from the Lower Cretaceous () Koonwara beds of Australia is excluded since it is not only not a formicid, but not even an aculeate (Naumann, 1993; Grimaldi et al., 1997: vide etiam Jell, 2004). 2005 ENGEL AND GRIMALDI: CRETACEOUS ANTS 23

APPENDIX (Continued) Complete lists of all issues of the Novitates and the Bulletin are available at World Wide Web site http://library.amnh.org/pubs. Inquire about ordering printed copies via e-mail from [email protected] or via standard mail from: American Museum of Natural History, LibraryÐ Scienti®c Publications, Central Park West at 79th St., New York, NY 10024. TEL: (212) 769- 5545. FAX: (212) 769-5009.

a This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).