Primitive New Ants in Cretaceous Amber from Myanmar, New ... · discovered in New Jersey amber, new species of †sphecomyrmines in both New Jersey and Burmese amber, as well as three

Copyright q American Museum of Natural History 2005 ISSN 0003-0082

P U B L I S H E D B Y T H E A M E R I C A N M U S E U M O F N AT U R A L H I S T O RY

CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024

Number 3485, 23 pp., 11 figures, 5 tables July 25, 2005

Primitive New Ants in Cretaceous Amber fromMyanmar, New Jersey, and Canada

(Hymenoptera: Formicidae)

MICHAEL S. ENGEL1 AND DAVID A. GRIMALDI2

CONTENTS

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Systematic Paleontology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

Family Formicidae Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Subfamily †Sphecomyrminae Wilson and Brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

†Sphecomyrmodes, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Genus †Sphecomyrma Wilson and Brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Genus †Haidomyrmex Dlussky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Subfamily †Brownimeciinae Bolton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12Genus †Brownimecia Grimaldi, Agosti, and Carpenter . . . . . . . . . . . . . . . . . . . . . . . 12

Subfamily Incertae Sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13†Myanmyrma, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Subfamily Aneuretinae? Emery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17†Cananeuretus, new genus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

1 Division of Invertebrate Zoology, American Museum of Natural History; Division of Entomology, Natural HistoryMuseum, and Department of Ecology and Evolutionary Biology, Snow Hall, 1460 Jayhawk Boulevard, Universityof Kansas, Lawrence, Kansas 66045–7523 ([email protected]).

2 Division of Invertebrate Zoology (Entomology), American Museum of Natural History ([email protected]).

2 NO. 3485AMERICAN MUSEUM NOVITATES

ABSTRACT

New information is provided on the oldest fossil ants (Formicidae), including the descriptionof a new species of †Sphecomyrma (†Sphecomyrminae), a new genus of sphecomyrmines, anew genus of apparent myrmeciines, and a new genus of apparent aneuretines. New materialfrom New Jersey amber (Turonian) includes workers of †Sphecomyrma freyi Wilson andBrown preserved together in the same piece of amber, a worker of an unidentifiable †Sphe-comyrma species, and a worker of †Brownimecia clavata Grimaldi, Agosti, and Carpenter(†Brownimeciinae). A new species of †Sphecomyrma in New Jersey amber is described andfigured from a worker as †S. mesaki, new species. Two worker specimens in Campanian amberfrom Canada are described, one of which is described as †Cananeuretus occidentalis, newgenus and species, and is tentatively placed in Aneuretinae. From Burmese amber (Albian-Cenomanian) are the oldest, definitive ants, along with ones in amber from Charente-Maritimeof France (approximately contemporaneous in age). A new genus and species, allied to †Sphe-comyrma, is described from these deposits as †Sphecomyrmodes orientalis, along with aremarkable new ‘‘poneroid’’, †Myanmyrma gracilis, new genus and species (Myrmeciinae?).A key to the species of †Sphecomyrma is provided, the classification of ants summarized, andthe Cretaceous records of Formicidae briefly outlined.

INTRODUCTION

Ants can truly be said to shape the terres-trial world. Of the 11,833 species of Formi-cidae3, many have a profound impact on nat-ural and manmade ecosystems, which ismade possible by their eusociality, frequentlylarge colony sizes, and abundance. There isscarcely a place outside of the polar regionswhere one cannot find these insects or theireffects. Ants are among the most commonand diverse kind of insect in various Ceno-zoic deposits, and are particularly wellknown in the fossilized faunas of Dominican,Sicilian, and Baltic ambers (Rasnitsyn andKulicka, 1990; Skalski and Veggiani, 1990;Wilson, 1985a), as are the less spectacularlypreserved compressions from Lagerstattesuch as Florissant (Carpenter, 1930), GreenRiver (Dlussky and Rasnitsyn, 2003), andother Tertiary localities throughout theworld. Although Formicidae came into theirown during the Cenozoic, in the Mesozoicand Early Tertiary (Paleocene) formicidswere rare, and their earliest evolution hasbeen gradually unveiled with each new fossildiscovery (e.g., Wilson et al., 1967; Wilson,1985b; Grimaldi et al., 1997; Grimaldi andAgosti, 2000a; Dlussky and Rasnitsyn, 2003;Dlussky et al., 2004; Nel et al., 2004).

Until 1985, the only true formicid knownfrom the Cretaceous Period was †Spheco-

3 Species total accurate as of 17 June 2005 (videwww.antbase.org).

myrma freyi Wilson and Brown, in New Jer-sey amber. Since 1985, ants have been re-ported in Cretaceous amber from Siberia,France, Canada, Burma, and additional newspecimens and taxa in New Jersey amber (re-viewed in Grimaldi et al., 1997, with addi-tions by Dlussky, 1999; Grimaldi and Agosti,2000a; Grimaldi et al., 2002; Nel et al., 2004:vide appendix 1). Here we report importantnew specimens of described ant taxa recentlydiscovered in New Jersey amber, new speciesof †sphecomyrmines in both New Jersey andBurmese amber, as well as three new generain Burmese and Canadian ambers (e.g., figs.1–3). While it is well established that NewJersey amber is of Turonian age (Grimaldi etal., 2000) and that Canadian amber is Cam-panian (Borkent, 1995), the dating of Bur-mese amber has been contentious. Formerlybelieved to be of Tertiary age, recent workhas demonstrated that the Burmese depositdates from the mid-Cretaceous (e.g., Zheri-khin and Ross, 2000; Grimaldi et al., 2002;Cruickshank and Ko, 2003). Thus, those taxain Burmese amber (Albian-Cenomanian) areamong the current oldest records of antsalong with ants in amber of approximatelycontemporaneous age from Charente-Mari-time, France (Nel et al., 2004), being at least8–10 million years older than previous re-cords of the family.

In keeping with myrmecological traditionand literature, we generally use terminologyfor morphological structures (e.g., antennal

2005 3ENGEL AND GRIMALDI: CRETACEOUS ANTS

Figs. 1–3. Three Cretaceous amber ants. 1. †Myanmyrma gracilis, new genus and species (AMNHBu-014) in Burmese amber. 2. †Cananeuretus occidentalis, new genus and species (TMP 8.89.7) inCanadian amber. 3. †Sphecomyrmodes orientalis, new genus and species (AMNH Bu-351) in Burmeseamber.

parts) as outlined by Holldobler and Wilson(1990) and Bolton (1994); however, for bodyregions we have used head, mesosoma (5alitrunk), and metasoma (5 petiole 1 gaster),as is standard in apocritan Hymenoptera.Throughout, ‘‘head length’’ is measured fromthe apex of the vertex to the anterior borderof the clypeus. The higher classification ofthe ants has recently undergone a major re-arrangement owing to the work of Bolton(2003) (summarized with minor modifica-tions in table 1). We have employed that

classification, herein, noting in various placeswhere we believe it might eventually bemodified. Material considered herein is de-posited in the Amber Collection, Division ofInvertebrate Zoology, American Museum ofNatural History, New York (AMNH); theDepartment of Palaeontology of the NaturalHistory Museum, London (NHML); andRoyal Tyrell Museum of Palaeontology,Drumheller, Canada (TMP). Specimens inthe AMNH collection were embedded in ep-oxy for preparation and study, using the pro-


TABLE 1Hierarchical Suprageneric Classification of the Ants (Formicidae) and Antlike Wasps (Armaniidae)

(modified from Bolton, 2003)a


cedure outlined by Nascimbene and Silver-stein (2000).

SYSTEMATIC PALEONTOLOGY

FAMILY FORMICIDAE LATREILLE

DIAGNOSIS: Head prognathous; dorsal rimof torulus often tuberculate or concealed un-der vertical lamella of frons; antenna genic-ulate. Primitively with anterior margin ofclypeus spiculate (apomorphically lost inmany modern lineages: vide infra). Infrabuc-cal sac present between labium and hypo-pharynx. Pronotum with posterodorsal mar-gin weakly concave; posterolateral apex trun-cate anterior to tegula. Metapleural glandpresent in females, opening above metacoxa(rarely absent); meso- and metacoxae contig-uous; inner metatibial spur modified as cal-car. Hind wing typically without jugal lobe(presence of the lobe is plesiomorphic withinthe family and likely part of the familialground plan). Metasoma petiolate; first me-tasomal segment forming true node (stronglyconstricted anteriorly and posteriorly); firstmetasomal sternum separated from secondmetasomal sternum by deep constriction.Morphologically distinct, sterile worker castetypically present4 (i.e., advanced eusocial);reproductives typically macropterous, work-ers apterous. Workers with pronotum fusedto mesothorax (a freely articulating prono-tum, present in some species, is plesio-morphic and undoubtedly part of the formi-cid ground plan), remaining segments typi-cally fused. Species advanced eusocial.

COMMENTS: The ants, currently including11,833 species (Bolton, 1995; www.antbase.org) have a cosmopolitan distributionand are among the most recognizable of allinsects. Numerous species exist in Cenozoicdeposits around the world and are relativelycommonly encountered. Species of Creta-ceous formicids, which are very rare, arebriefly outlined in appendix 1.

Bolton (2003) considered the antlikewasps of the Cretaceous family Armaniidaeto represent the basalmost subfamily of theants. We have, however, retained armaniidsat the family rank and as the sister group

4 Some inquilines have apomorphically lost the work-er caste.

(perhaps paraphyletic?) to Formicidae (videtable 1 and appendix 1).

It is interesting to note that many primitiveants have clypeal spicules with rounded api-ces (e.g., †Sphecomyrmodes, †Myanmyrma;Amblyoponinae). Apomyrma has similarspicules, but these are located on the labrumrather than along the anterior clypeal margin.The significance of this trait is as of yet un-clear (e.g., a ground-plan feature with nu-merous, apomorphic losses; or functionalconvergence).

SUBFAMILY †SPHECOMYRMINAE

WILSON AND BROWN

†Sphecomyrmodes, new genus

TYPE SPECIES: †Sphecomyrmodes oriental-is, new species.

DIAGNOSIS: Distinguished from all otherspecies of the tribe Sphecomyrmini by theminute, peglike denticles running along theentirety of the anterior margin of the clypeusand from †Sphecomyrma by the absence ofa medial extension or process on the clypealmargin.

ETYMOLOGY: The new genus-group nameis a combination of †Sphecomyrma, type ge-nus of the subfamily, and the suffix–odes,meaning ‘‘with the form of’’. The name ismasculine.

†Sphecomyrmodes orientalis, new speciesFigures 3–4

DIAGNOSIS: As for the genus (vide supra).DESCRIPTION: Head. Relatively large,

height of head slightly less than length ofalitrunk. Length of head 1.23 mm (with man-dibles closed). No apparent microsculptureon cuticle of head. Clypeus setose; setae ofmoderate length and widely separated. Man-dible simple, with only two teeth; outer sur-face with numerous, widely scattered, finesetae. Antenna of moderate length, withscape short, funicular article I (pedicel)shortest antennal article, funicular article IIthe longest article of funiculus. Lengths ofantennal articles (in mm): scape 0.23, pedicel(funicular article [fa] I) 0.13, faII 0.32, faIII0.15, faIV 0.15, faV 0.15, faVI 0.15, faVII0.17, faVIII 0.17, faIX 0.17, faX 0.17, faXI0.27. Mesosoma. Mesosomal length 1.33


Fig. 4. Holotype worker of †Sphecomyrmodes orientalis, new genus and species (AMNH Bu-351);general habitus, sting, clypeal margin and mandibles, and antenna.

mm; without apparent microsculpturing, withscattered fine, short setae on all visible sur-faces, those on propodeum about twice aslong as other setae. Coxae large, slightly in-flated, ventrally setose, setae numerous andfine. Legs moderate length. Foreleg with tar-somere I distinctly longer than combinedlengths of more distal tarsomeres; tarsomereI with ‘‘antennal cleaner’’ (strigil) a velvetynotch on ventral margin of proximal end; cal-car present, length slightly longer than great-est width of profemur, ventral margin withrow of fine teeth and (apically) setae. Patchof dense, elongate setae opposite strigil onprofemur; inner posterior surface of protibialapex with three stout, spinelike setae minute-

ly curved inward at their extreme apices (vis-ible on left foreleg). Pairs of stiff setae onventral surface of protarsomeres: tarsomeresI-III with three pairs, IV with two smallpairs. Pretarsal claw with minute subapicaltooth. Metasoma. Attachment of petiole topropodeum not particularly thick; thickness(measured in lateral view) of anterior end ofpetiolar peduncle 0.33 greatest depth of pro-podeum. Petiole length 0.38 mm, height 0.37mm. Preserved portion of gaster 1.73 mm inlength. Integument without apparent micros-culpturing, with scattered, fine setae. Distal-most metasomal segments torn apart, al-though sting bulb and sting well preservedbeneath metasoma (fig. 4).


TYPE MATERIAL: Holotype. AMNH Bu-351, an incompletely preserved worker in apiece of reddish-orange amber, from Myan-mar. Collected in Kachin state, Tanai village,on Ledo Road, 105 km NW Myitkyna, viaLeeward Capital Corp., 1999.

ETYMOLOGY: The specific epithet is theLatin word orientalis, meaning ‘‘of the east’’and is a reference to this being the first spe-cies of the tribe †Sphecomyrmini (sensu Bol-ton, 2003) recorded from Myanmar.

Genus †Sphecomyrma Wilson and Brown

†Sphecomyrma Wilson and Brown, In Wilson etal., 1967: 8. Type species: †Sphecomyrma freyiWilson and Brown, 1967, monobasic and orig-inal designation.

DIAGNOSIS: Scape short; funiculus longand filiform, about four times length ofscape; promesothoracic suture complete andwell developed; trochantellus absent; petiolewith distinct, domed node widely separatedfrom propodeum and remainder of metasomaby deep constrictions; cuticle without sculp-turing, superficial microscopic relief, withscattered and sparse setae.

COMMENTS: The genus, which is definedlargely by plesiomorphies, is doubtfullymonophyletic. It contains three species:†Sphecomyrma canadensis Wilson in Cana-dian amber, and †S. freyi Wilson and Brownand a new species in New Jersey amber (videinfra). In addition, we provide informationfrom newly identified material of †S. freyi.

Key to Species of †Sphecomyrma(based on the worker caste)

1. Anterior margin of clypeus with short, broadextension, surface with two, long setae atmost; compound eye round . . . . . . . . . . 2

— Anterior margin of clypeus with long, mediallobe (fig. 5), surface with numerous, longsetae; compound eye oval [New Jersey am-ber] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . †S. mesaki Engel and Grimaldi, n.sp.

2. Third antennal article slightly more thantwice as long as second article [New Jerseyamber] . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . †S. freyi Wilson and Brown

— Third antennal article about as long as secondarticle [Canadian amber] . . . . . . . . . . . . .. . . . . . . . . . . . . . . . †S. canadensis Wilson

†Sphecomyrma mesaki, new speciesFigure 5

DIAGNOSIS: Distinguished from all otherspecies of the genus by the median portionof the clypeus having a long ventral lobe,length of the clypeus through the lobe is0.463 the greatest width of clypeus; clypeussetose; and scape very short (1.23 the lengthof longest funicular article). The species canbe further distinguished from †S. freyi (theother species of the genus in New Jersey am-ber) by broad, shallow scrobes at base of an-tennae; an eye that is approximately 1/3 larg-er and oval (vs. almost perfectly round in †S.freyi); and a large head (length of head/length of mesosoma 5 0.83, vs. 0.65 in †S.freyi).

DESCRIPTION: Petiole and gaster not pre-served, so only head, mesosoma, and legspreserved. Head. Large, length of headslightly less than length of alitrunk. Lengthof head 2.20 mm (with mandibles closed);width of head 1.95 mm; length of eye 0.66mm. No microsculpture on cuticle of head.Vertex with fine, sparse pilosity, setae ca. 0.2mm long. Ocelli present, median ocellus sit-uated just above dorsal tangent of compoundeyes. Face bare. Bases of antennae situatedin shallow, broad scrobes; length of scrobeabout equal to length of scape and articulat-ing base. Eyes well developed, bare, situatedwell above bases of antennae; gena deep.Lateral portions of clypeus quadrate; medianportion distended into long ventral lobe thatextends to ventral margin of closed rightmandible. Clypeus setose, except on middlepart. Mandibles simple, with only two teeth.Antennae of moderate length, with scapeshort, funicular article I (pedicel) shortest an-tennal article, funicular article II the longestarticle of funiculus. Lengths of antennal ar-ticles (in mm): scape 0.53, pedicel (funiculararticle [fa] I) 0.20, faII 0.43, faIII 0.35, faIV0.30, faV 0.23, faVI 0.23, faVII 0.30, faVIII0.22, faIX 0.26, faX 0.30, faXI 0.33. Meso-soma. Mesosomal length 2.66 mm; withoutmicrosculpturing, except at posterolateralmargin of promesonotal suture, where eightfine grooves occur. Dome of promesonotumsetose; several fine setae on dorsal surface ofmetanotum and propodeum. Coxae large, in-flated, ventrally setose. Attachment of petiole


Fig. 5. Holotype worker of †Sphecomyrma mesaki, new species (AMNH NJ-1023); lateral habitusand facial view.


to propodeum not particularly thick; thick-ness (measured in lateral view) of anteriorend of petiolar peduncle 0.3x greatest depthof propodeum. Metapleural gland opening(MGO) and MG bulla obvious, situated onposterolateral part of propodeum just abovemetacoxa. MGO small, with groove runningbetween it and extended almost to ventalmargin of propodeum. Legs moderate length.Foreleg with tarsomere I slightly longer thancombined length of more distal tarsomeres;tarsomere I with ‘‘antennal cleaner’’ (strigil)a velvety notch on ventral margin of proxi-mal end; calcar present, length slightly lon-ger than greatest width of femur, ventral mar-gin with row of fine teeth and (apically)hairs. Stiff setae on ventral surface of pro-tarsomeres: tarsomere I with 7 pairs, II with3 pairs, III with 3 small pairs, IV with 2small pairs. Pretarsal claw with subapicaltooth. Metasoma not preserved.

TYPE MATERIAL: Holotype. AMNH NJ-1023, an incompletely preserved worker in apiece of amber barely larger than the ant,from Sayreville, New Jersey (MiddlesexCo.), White Oaks outcrop, coll. Bob Mesak(fig. 5). The amber is an irregularly shapeddrop, 7 3 5 3 4 mm, made of clear yellowamber. Portions of some appendages arebreached at the surface, and the petiole andgaster were similarly lost at the surface.Since very little amber exists between the antand surface of the amber piece, no trimmingor polishing was possible. Still, detailsthrough the rough surface are highly visibleby immersing the piece in glycerine.

ETYMOLOGY: The specific epithet is a pat-ronymic for Bob Mesak, who collected anddonated this valuable specimen to theAMNH.

†Sphecomyrma freyi Wilson and BrownFigure 6

†Sphecomyrma freyi Wilson and Brown, In Wil-son et al., 1967: 8. Grimaldi et al., 1997: 12(redescription of some features, new specimens,neotype).

MATERIAL: AMNH NJ-943, in amber fromNew Jersey: Middlesex Co., Sayreville,White Oaks outcrop, collected by Keith Luz-zi (fig. 6). The piece is transparent yellow,originally much larger than now; was em-

bedded and trimmed to 14 3 15 3 4 mm.The piece contains two workers of †S. freyi.

DESCRIPTIVE NOTES: Both workers arelargely but not completely preserved. Spec-imen A has the frontal half of the head miss-ing; most of the antenna is present except forbases of the scapes; the entirety of the re-mainder of the body is preserved and thesting appears largely or fully extruded. Spec-imen B has the dorsal and apical part of thegaster missing, as well as portions of theright hind leg; the anterior third of the headis obscured by Schimmel and a bubble.

Measurements of body: Width of head(specimen B), 1.03 mm; length of head (B,approximate), 1.20 mm; length of mesosoma1.39 mm (B), 1. 40 mm (A); length of petiole0.29 mm (B), 0.33 mm (A); length of gaster(A) 1.72 mm; length of extruded portion ofsting (A), 0.33 mm. Measurements of anten-nal articles (as measured for right antenna inA, left antenna in B) presented in table 2.Measurements of leg segmentation (as mea-sured on specimen A) presented in table 3.

COMMENTS: The discovery of this piece ofamber is highly significant and addressesquestions of the social behavior of primitiveants like †Sphecomyrma. As reviewed in Gri-maldi et al. (1997), Dlussky (1987, 1988),and Poinar et al. (1999) questioned whether†Sphecomyrma was a true ant since it hadsuch a short scape [but see response to Poin-ar et al. (1999) by Grimaldi and Agosti(2000b)]. With such a short scape, Dlusskyargued that it would be impossible for†Sphecomyrma to antennate, and therefore itwas considered highly unlikely for †Sphe-comyrma to have been social. Among the ap-proximately 1700 fossiliferous pieces of NewJersey amber in the AMNH collection thusfar, four pieces contain worker or male sphe-comyrmine ants. These ants are (and proba-bly originally were) exceedingly rare, and theprobability that two workers would be pre-served in one piece by chance alone is ex-tremely remote. It is most parsimonious toexplain the occurrence of two workers in thesame piece as a result of social behavior.

†Sphecomyrma sp.

MATERIAL: AMNH NJ-942, a male in am-ber from New Jersey: Middlesex Co., Say-


Fig. 6. Two workers of †Sphecomyrma freyi Wilson and Brown preserved in a single piece of NewJersey amber (AMNH NJ-943).

reville, White Oaks outcrop. Collected by thelate Steve Swolensky. Specimen is in a clearyellow piece of amber 5 3 7 mm, embeddedin epoxy and trimmed to 1.5 mm thicknessfor a full lateral view of the ant.

COMMENTS: Venation and other details ofthis specimen are indistinguishable fromAMNH NJ-242, described and figured by Gri-maldi et al. (1997) as †Sphecomyrma? sp.Measurements: Length of hind wing (forewing


TABLE 2Measurements of Antennal Articles for†Sphecomyrma freyi (AMNH NJ-943)

TABLE 3Leg Measurements (in mm) for †Sphecomyrma freyi (AMNH NJ-943)

apices are lost), 1.97 mm; length of mesosoma1.29 mm; length of petiole 0.36 mm; length ofgaster 1.05 mm; length of antenna 2.40 mm.

Genus †Haidomyrmex Dlussky

†Haidomyrmex Dlussky, 1996: 84. Type species:†Haidomyrmex cerberus Dlussky, 1996, mono-basic and original designation.

DIAGNOSIS: Clypeus a small, hemisphericlobe lying just below antennal bases, pos-sessing brush of ca. 60 fine, stiff, whitish se-tae; setae are evenly arranged, those on ven-tral margin thickened at base but taper to afine point apically. Compound eyes small,length ca. 0.23 length of head capsule; ocelliabsent. Mandibles elongate, scimitar-shaped,without serrations or teeth. Propodeumrounded in profile. Petiole one-segmented,nodiform, with distinct constriction at artic-

ulation with remainder of metasoma. Knownfrom the worker caste only.

COMMENTS: The head of †Haidomyrmex isenigmatic (fig. 7) and much of the detail isobscured below the clypeus and where themandibles articulate. Indeed, it is likely thatthere has been significant distortion at themanibular bases resulting in the rather ‘‘deep’’appearance they have relative to the clypeus.

The peculiar clypeal setae may have serveda sensory function, perhaps as trigger hairs forthe large mandibles, much the way gaff-shaped mandibles of various myrmecines and‘‘poneroids’’ function. In those living antslong, fine, stiff trigger setae lie on the insidesurface of the mandibles and on the oral mar-gin. †Haidomyrmex has no such setae, so per-haps the clypeal brush functioned analogous-ly. How †Haidomyrmex might have fed itselfis an enigma. It is possible that this worker issimilar to the major workers of Eciton, whichcannot feed themselves.

†Haidomyrmex cerberus DlusskyFigures 7–8

†Haidomyrmex cerberus Dlussky, 1996: 85.

HOLOTYPE: NHML In.20182, partial work-er specimen in amber from Myanmar.

COMMENTS: We have tentatively followedpast authors in placing †Haidomyrmex within†Sphecomyrminae (e.g., Bolton, 2003).However, it is important to note the signifi-cant similarities between this genus and†Brownimecia in New Jersey amber (vide in-fra). Both genera have large, dome-shapedheads, with relatively small compound eyes,lack ocelli, have large mandibles devoid ofserrations or dentition, and an elongate an-terior extension, or collar, of the pronotum.


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

SUBFAMILY †BROWNIMECIINAE BOLTON

Genus †Brownimecia Grimaldi, Agosti,and Carpenter

†Brownimecia Grimaldi, Agosti, and Carpenter,1997: 20. Type species: †Brownimecia clavata.Grimaldi, Agosti, and Carpenter, 1997, mono-basic and original designation.

DIAGNOSIS: Antenna distinctly clubbed,apical funicular article twice the width ofbasal ones and pedicel. Ocelli absent. Man-dibles long, thin, scimitar-shaped, stronglycruciate, without teeth or crenulations, but

with oral surface bearing about 30 short, spi-culelike setae. Metasoma with slight but def-inite constriction between second and thirdsegment (also known as abdominal segmentsIII and IV; gastral segments 1 and 2). Knownfrom the worker caste only.

†Brownimecia clavata Grimaldi, Agosti,and Carpenter

†Brownimecia clavata Grimaldi, Agosti, and Car-penter, 1997: 20.

MATERIAL: AMNH NJ-941, in amber from


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

New Jersey: Middlesex Co., Sayreville,White Oaks outcrop, collected by SteveSwolensky.

COMMENTS: A beautifully preserved spec-imen in clear yellow amber, 5 3 8 mm,which was embedded in epoxy and trimmedto 2 mm thickness. Like the holotype whichwas described in 1997, it is curled up and thesting is extruded. The piece also containssome wood fragments and a frass pellet. Thelength of head (including closed mandibles)0.88 mm, greatest width of head 0.74 (be-tween outer margins of eyes), length of eye0.24 mm, length of trunk 0.94 mm, length ofpetiole 0.32 mm. The mandibles are tightlyclosed, so details are less visible than in theholotype. The front portion of the head, how-ever, and especially the clypeus, are morevisible. The clypeus has fine, parallel cren-ulations along and perpendicular to the dor-sal margin of the clypeus. Most significantly,ocelli are definitively absent (rather than ves-tigial or minute). Three large ocelli are rarein ants, with scattered occurrence (e.g., Cer-apachyinae [Cylindromyrmex, Simopone],Formicinae [Aphophomyrmex, Notostigma,Alloformica, Cataglyphis], Myrmeciinae[Myrmecia]), otherwise the ocelli are repeat-edly lost, reduced, or modified secondarily(as is true for †Brownimecia). The large,well-developed ocelli of †Sphecomyrma are

clearly a plesiomorphic feature and part ofthe formicid ground plan.

SUBFAMILY INCERTAE SEDIS

†Myanmyrma, new genus

TYPE SPECIES: †Myanmyrma gracilis, newspecies.

DIAGNOSIS: Gracile ‘‘poneroid’’ ant withextremely long legs; mandibles nearly equalto length of head, sickle-shaped, with a long,apical sharp tooth and blunt subapical one;clypeal margin bilobate, denticulate; long,spatulate genal process; antennal sockets in-clined, antennal funiculus very long, secondarticle longest; articles slightly shorter api-cally; integument of head spinose; clypeusdeeply incised along anterior margin andwith distinct peglike setae; eyes and ocellinot apparent (but may be present); pretarsalclaws with minute tooth; petiole with distinctnodus; gastral constriction distinct; sting welldeveloped. Easily distinguished from †Hai-domyrmex, another highly modified ant inBurmese amber, by the mandibular and clyp-eal structure, head microsculpturing, meta-somal constriction, long genal processes, andvery long legs.

ETYMOLOGY: Taken directly from Myan-mar, the country of the amber’s origin; and–


myrma, a common suffix for ant genera. Thename is feminine.

†Myanmyrma gracilis, new speciesFigures 1, 9

DIAGNOSIS: As for the genus (vide supra).DESCRIPTION: Specimen has been com-

pressed, but body form was clearly extremelygracile, with very long legs. Head. Propor-tions difficult to determine due to compres-sion; length ca. 1.76 mm. Front of head withpair of carinae diverging dorsally; bluntspines on ridges of carinae and on frons;spines absent below ventral limit of carinae,with irregular cuticular foveae on clypeus.Ventral margin of clypeus with deep medianemargination; clypeus with two lobes, mar-gin of two lobes with row of ca. 14 minutedenticles on each lobe, labrum similarly bi-lobed and denticulate. Lateral portion ofclypeus lobed, without denticles. Gena withspatulate process, exact length difficult todiscern, but ca. 0.33 length of mandible.Mandibles long, 1.61 mm. (right one), nearlyequal to length of head, sickle-shaped; apicaltooth long, sharp; blunt subapical tooth, noother teeth. No fine setae apparent. Apex ofright mandible longer than left, especiallyapical tooth. Eyes well developed (difficultto observe since they are sunken in a cavity),length ca. 0.253 length of head. Antennavery long (7.2 mm); scape fairly short, withwide base; funicular article (fa) 1 shortest,fa2 longest; lengths of antennal articles (inmm): scape 0.63, funicular article (fa) [ped-icel] I 0.44, faII 1.05, faIII 0.62, faIV 0.53,faV 0.61–faVI 0.45, faVII 0.40, faVIII 0.41,faIX 0.48, faX 0.48, faXI 0.65. Mesosoma.Elongate, not deep, length ca. 3.52 mm;small opening to metapleural gland (MPG)apparent just above hind coxa. Legs extreme-ly long (vide measurements in table 4); tro-chantellus apparently absent. Protibia withdorsal, preapical brush of fine setae; onelarge, two finer apical spurs; ventral surfaceof probasitarsus with fine pilosity, six pairsof stiff setae; all tarsomeres with four apicalspines; each pretarsal claw with mediantooth. Mesotibia with pair of ventroapicalspurs; ventral surface of mesobasitarsus withthree rows of 10 stiff setae each; metatibiawith pair of apical spurs, one larger and pec-

tinate; ventral surface of metabasitarsus withfine pilosity, 10 pairs of setae. Metasoma.Petiole attachment to second metasomal seg-ment narrow; petiole with high, narrow node;length of petiole 1.26 mm; posterior portionof petiole narrow, tubular; deep constrictionbetween second and third metasomal seg-ments. Metasoma overall quite small, ca.0.3x length of body (excluding antennae).Terga and sterna telescoped in specimen, asshown in figure 9; gaster (i.e., metasoma ex-cluding petiole) approximately 2.95 mmlong. Sting well developed, only partly ex-truded but internally visible; pygidium withlong fine setae.

HOLOTYPE: AMNH Bu-014, a worker, inamber from northern Myanmar (Burma)(figs. 1, 9). Collected in Kachin state, Tanaivillage, on Ledo Road, 105 km NW Myit-kyna, via Leeward Capital Corp., 1999. Theant is in a transparent yellow piece of amber,which was originally more than 20 mm indiameter and 30 mm in length. The piece wasoccluded with fractures and debris, so it re-quired epoxy embedding and then trimmingto better observe the ant. The piece is nowrhomboid-shaped, 17 3 22 3 4 mm, thebroad surfaces being parallel to the lateralsurface of the ant. A flat edge also permits afrontal view of the head. Other inclusions inthe piece are numerous frass pellets andwood fragments, further suggestive (besidesbody form) of arboreal/wood nesting habitsof the ant. The ant specimen shows a greatdeal of distortion due to compression. Thecuticle is transparent, facilitating observationof some internal structures (such as unex-truded portions of the sting), but some com-pressed surfaces could easily be mistaken forflanges and other structures.

ADDITIONAL MATERIAL: Two additional,fragmentary specimens are perhaps represen-tative of †M. gracilis; however, because oftheir poor preservation, we hesitate to des-ignate them as paratypes. Both are preservedin amber from northern Myanmar and withthe same collection data as the holotype.AMNH Bu-225, a poorly preserved workerbut very similar to holotype in observabletraits. AMNH Bu-1509, a badly compressedworker preserved in yellow amber; headmostly crushed and difficult to discern, me-tasoma similarly compressed; somewhat


Fig. 9. Holotype worker of †Myanmyrma gracilis, new genus and species (AMNH Bu-014); facialview and general habitus.


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

smaller than holotype and Bu-225 but stillexhibiting the same elongate, slender legs,constriction in first gastral segments, etc.,this individual may represent a minor worker.

ETYMOLOGY: The specific epithet is theLatin word gracilis, meaning ‘‘slender’’, asa reference to elongate legs and structure ofthis ant.

COMMENTS: Constriction of the metasomaindicates the specimen is a ‘‘poneroid’’,which is the third and oldest Cretaceous re-cord of this paraphyletic grade of primitiveants (the poneroid grade includes the poner-omorph and myrmeciomorph subfamilies ofBolton, 2003). The other Cretaceous ‘‘pone-roids’’ are †Brownimecia clavata (in NewJersey amber: Turonian), and †Canaponedentata Dlussky (in Canadian amber: Cam-panian). It is interesting that †Myanmyrma isonly the fourth record of ants in Burmeseamber, but two of these records are for highlymodified species. †Burmomyrma rossi Dlus-sky is based on a single, alate, incompletespecimen, with head and portion of the ali-trunk missing. The most distinctive featureof †Burmomyrma is the wing with highly re-duced venation; thus, the taxonomic conceptof this genus is not entirely comparable withthat of the other two Burmese amber genera.†Burmomyrma, as discussed again later (videinfra), has been tentatively placed in theAneuretinae (Dlussky, 1996; Bolton, 2003).Both †Haidomyrmex and †Myanmyrma areextremely gracile and relatively large ants,with large, highly modified mouthparts andgenae. The extremely long legs and slenderbody are analogous to Leptomyrmex (Doli-choderinae) and Oecophylla (Formicinae),the latter of which is entirely arboreal. †Hai-domyrmex and †Myanmyrma presumably

had similar habits. †Haidomyrmex and†Myanmyrma are clearly not closely related,the former placed in the †Sphecomyrminaeby Dlussky. The type and only specimen of†Haidomyrmex, however, has only the sec-ond metasomal (i.e., first gastral) segmentpreserved, so the lack of a constriction pos-terior to this segment is suggested, not defin-itive (vide supra).

The lengths of basal articles of the antennaof †Myanmyrma are highly significant. Fu-nicular article 2 is the longest article in thefuniculus, as is the case for most †Spheco-myrminae (except †S. canadensis Wilson).This condition does not exist in the otherCretaceous ‘‘poneroids’’, †Brownimecia and†Canapone. A long funicular article 2 wasproposed by Grimaldi et al. (1997: 8) as oneof only two or three possible synapomor-phies for the †Sphecomyrminae. If truly apo-morphic, the long funicular article 2 wouldhave been independently derived in †Myan-myrma. If a long funicular article 2 was ple-siomorphic, the monophyly of the †Spheco-myrminae would be seriously doubtful. Out-group evidence from closely related aculeatefamilies suggests that, indeed, this feature issymplesiomorphic for basal ants, including†Myanmyrma.

Placement of †Myanmyrma in a subfamilyis challenging. As noted, the metasomal con-striction implies placement among the po-neroid grade where it best approximates spe-cies of the Ponerinae (poneromorph) or theMyrmeciinae (myrmeciomorph). Neitherplacement is entirely satisfactory, but we be-lieve inclusion in the latter subfamily is morelikely (although we tentatively retain the ge-nus as subfamily incertae sedis). Like myr-meciines, †Myanmyrma has a metasomal


constriction, the antennal sockets inclined tonearly a vertical position, 12-segmented an-tennae, pectinate inner metatibial spur, andelongate mandibles. Unlike typical myrme-ciines, however, the the new genus has man-dibles that are sickle-shaped, with dentitiononly at the apices; has a deeply incised clyp-eal margin, with distinct spicules on thelobes; has a spinose integument on the head;has gracile legs; and elongate genal process-es. These traits are, however, autapomorphicand do not preclude placement in Myrmeci-inae. Within Myrmeciinae, †Myanmyrmawould appear to be closest to the tribe Myr-meciini (sensu Bolton, 2003; Ward and Bra-dy, 2003) as evidenced by the elongate sec-ond funicular segment and distinctly two-segmented waist. The latter trait is likely ple-siomorphic as it occurs in Myrmecia andweakly so in †Prionomyrmex, perhaps beingapomorphically lost in Nothomyrmecia. Thepresence of spicules along the clypeal marginin †Myanmyrma is an interesting and enig-matic feature that should be further exploredfor its systematic implications.

This is the species that is referred to byGrimaldi and Engel (2005) as ‘‘undescribedMyrmeciinae?’’

SUBFAMILY ANEURETINAE? EMERY

†Cananeuretus, new genus

TYPE SPECIES: †Cananeuretus occidentalis,new species.

DIAGNOSIS: Compound eyes present, small;ocelli absent. Antennal sockets slightly in-clined; scape elongate. Mandible of primitiveconstruction, with four distinct teeth (fig. 10),basalmost tooth largest. Preoccipital carinaabsent; ocelli absent. Alitrunk elongate, slen-der; pronotal neck elongate. Meso- and me-tatibia with a single spur; pretarsal clawssimple. Propodeal lobes, denticles, andspines absent; petiole one-segmented, ante-rior peduncle elongate, with distinct tubular,postnodal section (more elongate than inAneuretus) articulating high on anterior sur-face of second metasomal segment, petiolartergum and sternum fused; nodus not veryhigh. Metasoma excluding petiole short andglobular; without U-shaped emargination onanterior margin of second metasomal seg-

ment; sting present and well developed.Known from the worker caste only.

ETYMOLOGY: The new genus-group nameis a combination of Canada, the country fromwhich this amber originates, and Aneuretus,type genus of the Aneuretinae. The name ismasculine.

COMMENTS: †Cananeuretus might at firstbe confused for †Eotapinoma, a dolichoder-ine described from Canadian amber (Dlus-sky, 1999). †Cananeuretus differs notablyfrom †Eotapinoma in the presence of a well-developed sting (apparently vestigial in†Eotapinoma, like all dolichoderines), themore elongate petiole with a distinct nodus(shorter petiole, which does not have anelongate peduncle and without developed no-dus in †Eotapinoma), and the reduced com-pound eyes (large in †Eotapinoma).

This new genus is tentatively placed inAneuretinae, although the group shows most-ly what are presumed plesiomorphies for thesubfamily. The presence of a strong, well-developed sting and absence of an acidoporeexcludes placement in Dolichoderinae orFormicinae. However, the genus is bestplaced among the formicomorph subfamilies(sensu Bolton, 2003) where it most closelyapproximates the Aneuretinae. As more ma-terial is recovered of these ants and a cladis-tic framework for fossil aneuretinesachieved, it is possible that †Cananeuretusand others may prove to be stem-groupAneuretinae or even represent stem groups toAneuretinae 1 Dolichoderinae.

Today the subfamily Aneuretinae consistsof a single species occurring in Sri Lanka.During the Tertiary aneuretines were distrib-uted widely in the Northern Hemisphere asevidenced by their occurrence in Baltic am-ber (Wheeler, 1914) and Florissant, Colo-rado (Carpenter, 1930). The mid-Cretaceousgenus †Burmomyrma, in amber from Myan-mar, has been tentatively placed in theAneuretinae (Dlussky, 1996; Bolton, 2003)and, until now, represented the sole Meso-zoic record for the subfamily. It is increas-ingly apparent that aneuretines were widelydistributed during the Mesozoic and earlyTertiary, and that Aneuretus is a notable rel-ict in Sri Lanka today. The significant ex-tinction of northern aneuretines was perhapsa result of the infamous Eocene-Oligocene


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

climatic shift, a cooling event that dramati-cally altered the evolutionary history andbiogeography of numerous insect lineagesthat are today austral relicts (Grimaldi andEngel, 2005).

†Cananeuretus occidentalis, new speciesFigures 2, 10–11

DIAGNOSIS: As for the genus (vide supra).DESCRIPTION: Body with sparsely scattered,


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

fine setae; integument finely imbricate.Head. Length ca. 0.52 mm; narrowed slight-ly toward apex; vertex gently rounded. Man-dibles with short, pointed teeth, basalmostapical tooth largest. Eyes present, relativelysmall and low on head (difficult to observe).Scape fairly long (0.46 mm), narrowedslightly at base for articulation with bulb (fig.10); pedicel 0.653 length of scape. Meso-soma. Elongate, not deep, length ca. 3.10mm; pronotum forming a distinct neck an-teriorly. Legs slender and long; trochantelluspresent (fig. 11). Inner surfaces of protibiaand probasitarsus with row of short, fine se-tae forming weakly defined brushes (fig. 10);tarsomeres beyond basitarsus triangular, lat-erally with short, distinct, stiff setae; pretar-sal claws simple. Mesotibia and metatibiawith a single spur; metatibial spur distinctlyand minutely setose (fig. 10). Propodeumwithout lobes or spines, broadly rounded andgently sloping to articulation with petiole.Metasoma. Petiole with elongate peduncle,

distinct nodus, and relatively long posterior,tubular section, articulating high on secondmetasomal segment; petiolar tergum andsternum fused; metasoma without constric-tion between second and third segments (i.e.,gastral segments 1 and 2); metasoma overallquite globular. Sting well developed, extrud-ed.

HOLOTYPE: TMP 8.89.7, labeled, ‘‘GrassyLake, Alberta, Campanian, Foremost For-mation, Formicidae’’ (figs. 2, 10). Worker inamber that is embedded in a thin block ofepoxy and slide mounted.

ADDITIONAL MATERIAL: TMP 91.149.3, la-beled, ‘‘Hymenoptera, Grassy Lake, Alberta,Campanian, Foremost Formation, Formici-dae’’ (fig. 11). Worker in amber that is em-bedded in a thin block of epoxy and slidemounted. The specimen is partial and owingto some minor differences in the shape of thepetiole, we have considered it best to onlytentatively assign it to this species.

ETYMOLOGY: The specific epithet is the


term occidentalis, meaning ‘‘of the west’’,and is a reference to this being the first aneu-retine or aneuretine-like ant from the Meso-zoic of the Western Hemisphere.

ACKNOWLEDGMENTS

We are grateful to D. Agosti and P. S.Ward for their invaluable reviews of themanuscript and for correcting several inac-curacies; to J. D. Gardner (TMP) for the loanof specimens; and to A. J. Ross (NHML) forhosting us during our 2002 visit to the De-partment of Palaeontology at which time theholotype of †Haidomyrmex was studied. Thiswork was supported by NSF DBI-9987372(to DAG) and by NSF EPSCoR grantKAN29503 and NSF EF-0341724 (to MSE).This is contribution Nr. 3416 of the Divisionof Entomology, Natural History Museum andBiodiversity Research Center, University ofKansas.

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APPENDIXCretaceous Ants (Formicidae) and Related Taxa

The following table summarizes the described species of ants and antlike wasps (i.e., Armaniidae) fromthe Cretaceous. The classificatory structure generally follows that of Bolton (2003). The fossilCretacoformica explicata Jell and Duncan (1986) from the Lower Cretaceous (Aptian) Koonwara bedsof 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).


APPENDIX(Continued)

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide Website http://library.amnh.org/pubs. Inquire about ordering printed copies via e-mail [email protected] or via standard mail from: American Museum of Natural History, Library—Scientific 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).

Primitive New Ants in Cretaceous Amber from Myanmar, New ... · discovered in New Jersey amber, new species of †sphecomyrmines in both New Jersey and Burmese amber, as well as three

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