^ ^AMERICAN MUSEUM
rsovitates
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
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 from
Myanmar, New Jersey, and Canada
(Hymenoptera: Formicidae)
MICHAEL S. ENGEL' AND DAVID A. GRIMALDF
CONTENTS
Abstract 2
Introduction 2
Systematic Paleontology 5
Family Formicidae Latreille 5
Subfamily fSphecomyrminae Wilson and Brown 5
'\Sphecomyrmodes, new genus 5
Genus '\Sphecomyrma Wilson and Brown 7
Genus '\Haidomyrmex Dlussky 11
Subfamily fBrownimeciinae Bolton 12
Genus fBrownimecia Grimaldi, Agosti, and Carpenter 12
Subfamily Incertae Sedis 13
'\Myanmyrma, new genus 14
Subfamily Aneuretinae? Emery 17
'fCananeuretus, new genus 17
Acknowledgments 20
References 20
Appendix 22
' 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 (msengel@ku.edu).
' Division of Invertebrate Zoology (Entomology), American Museum of Natural History (grimaldi@amnh.org).
Copyright © American Museum of Natural History 2005 ISSN 0003-0082
AMERICAN MUSEUM NOVITATES
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ABSTRACT
New information is provided on tlie oldest fossil ants (Formicidae), including the description
of a new species of '\Sphecomyrma (fSphecomyrniinae), a new genus of sphecomyrmines, a
new genus of apparent myrmeciines, and a new genus of apparent aneuretines. New material
from New Jersey amber (Turonian) includes workers of '\Sphecomyrma freyi Wilson and
Brown 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
(tBrownimeciinae). A new species of 'fSphecomynna in New Jersey amber is described and
figured from a worker as '\S. mesaki, new species. Two worker specimens in Campanian amber
from Canada are described, one of which is described as ^Cananeuretus occidentalis, new
genus 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-Maritime
of France (approximately contemporaneous in age). A new genus and species, allied to '\Sphe-
comyrma, is described from these deposits as ^Sphecomyrmodes orientalis, along with a
remarkable new "poneroid", ^Myanmyrma gracilis, new genus and species (Myrmeciinae?).
A key to the species of ifSphecomyrma is provided, the classification of ants summarized, and
the 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-
cidae^, many have a profound impact on nat-
ural and manmade ecosystems, which is
made possible by their eusociality, frequently
large colony sizes, and abundance. There is
scarcely a place outside of the polar regions
where one cannot find these insects or their
effects. Ants are among the most common
and diverse kind of insect in various Ceno-
zoic deposits, and are particularly well
known in the fossilized faunas of Dominican,
Sicilian, and Baltic ambers (Rasnitsyn and
KuHcka, 1990; Skalski and Veggiani, 1990;
Wilson, 1985a), as are the less spectacularly
preserved compressions from Lagerstatte
such as Florissant (Carpenter, 1930), Green
River (Dlussky and Rasnitsyn, 2003), and
other Tertiary localities throughout the
world. Although Formicidae came into their
own during the Cenozoic, in the Mesozoic
and Early Tertiary (Paleocene) formicids
were rare, and their earliest evolution has
been gradually unveiled with each new fossil
discovery (e.g., Wilson et al., 1967; Wilson,
1985b; Grimaldi et al., 1997; Grimaldi and
Agosti, 2000a; Dlussky and Rasnitsyn, 2003;
Dlussky et al., 2004; Nel et al., 2004).
Until 1985, the only true formicid known
from the Cretaceous Period was fSpheco-
' Species total accurate as of 17 June 2005 {vide
www.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 new
specimens 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 important
new specimens of described ant taxa recently
discovered in New Jersey amber, new species
of tsphecomyrmines in both New Jersey and
Burmese amber, as well as three new genera
in Burmese and Canadian ambers (e.g., figs.
1-3). While it is well established that New
Jersey amber is of Turonian age (Grimaldi et
al., 2000) and that Canadian amber is Cam-
panian (Borkent, 1995), the dating of Bur-
mese amber has been contentious. Formerly
believed to be of Tertiary age, recent work
has demonstrated that the Burmese deposit
dates from the mid-Cretaceous (e.g., Zheri-
khin and Ross, 2000; Grimaldi et al., 2002;
Cruickshank and Ko, 2003). Thus, those taxa
in Burmese amber (Albian-Cenomanian) are
among the current oldest records of ants
along with ants in amber of approximately
contemporaneous age from Charente-Mari-
time, France (Nel et al., 2004), being at least
8-10 million years older than previous re-
cords of the family.
In keeping with myrmecological tradition
and literature, we generally use terminology
for morphological structures (e.g., antennal
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
?.t,.v ''VAI
Figs. 1—3. Three Cretaceous amber ants. 1. '\Myanmymia gracilis, new genus and species (AMNH
Bu-014) in Burmese amber. 2. ^Cananeuretus occidentalis, new genus and species (TMP 8.89.7) in
Canadian amber. 3. 'fSphecoinynnodes orientalis, new genus and species (AMNH Bu-351) in Burmese
amber.
parts) as outlined by Holldobler and Wilson
(1990) and Bolton (1994); however, for body
regions we have used head, mesosoma ( =
alitrunk), and metasoma (= petiole + gaster),
as is standard in apocritan Hymenoptera.
Throughout, "head length" is measured from
the apex of the vertex to the anterior border
of the clypeus. The higher classification of
the 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 places
where we believe it might eventually be
modified. Material considered herein is de-
posited in the Amber Collection, Division of
Invertebrate Zoology, American Museum of
Natural History, New York (AMNH); the
Department of Palaeontology of the Natural
History Museum, London (NHML); and
Royal Tyrell Museum of Palaeontology,
Drumheller, Canada (TMP). Specimens in
the AMNH collection were embedded in ep-
oxy for preparation and study, using the pro-
AMERICAN MUSEUM NOVITATES
NO. 3485
TABLE 1
Hierarchical Suprageneric Classiflcation of the Ants (Formicidae) and Antlilie Wasps (Armaniidae)
(modified from Bolton, 2003)»
Family fARMANIIDAE Dlussky
Family FORMICIDAE Latreille
Subfamily tSphecomyrminae Wilson & Brown
myrmicomorph group, Subfamily Myrmicinae (continued)
Tribe tSphecomymiini Wilson & Brown
dacetite tribal group
Tribe tHaidomyrmecini Bolton
Tribe Basicerotini Brown
"poneromorph" group [paraphyletic]
Tribe Dacetini Forel'
Subfamily tBrownimeciinae Bolton
Tribe Phalacromyrmecini Dlussky & Fedoseeva
Subfamily Amblyoponinae Forel''
cataulacite tribal group
Subfamily Paraponerinae Emery
Tribe Cataulacini Emery
Subfamily Heteroponerinae Bolton
Tribe Cephalotini Smith
Subfamily Ponerinae Lepeletier de Saint Fargeau
attite tribal group
Tribe Ponerini Lepeletier de Saint Fargeau
Tribe Blepharidattini Wheeler & Wheeler
Tribe Thaumatomyrmecini Emery
Tribe Attini Smith
Tribe Platythyreini Emery
solenopsidite tribal group
Subfamily Proceratiinae Emery
Tribe Stenammini Ashmead
Tribe Proceratiini Emery
Tribe Solenopsidini Forel
Tribe Probolomyrmecini Perrault
myrmicite tribal group
Subfamily Ectatomminae Emery
Tribe Myrmicini Lepeletier de Saint Fargeau
Tribe Ectatommini Emery
Tribe Tetramoriini Emery
Tribe Typhlomyrmecini Emery
Tribe Pheidolini Emery
leptanillomorph group""
Tribe Lenomyrmecini Bolton
Subfamily Leptanillinae Emery
Tribe ParatopuUni Wheeler
Tribe Anomalomyrmini Taylor
formicoxenite tribal group
Tribe Leptanillini Emery
Tribe Crematogastrini Forel
dorylomorph group
Tribe Ankylomyrmini Bolton
Subfamily Cerapachyinae Forel
Tribe Liomyrmecini Mayr
Tribe Acanthostichini Emery
Tribe Meranoplini Emery
Tribe Cylindromyrmecini Emery
Tribe Myrmicariini Forel
Tribe Cerapachyini Forel
Tribe Formicoxenini Forel
Subfamily Leptanilloidinae Bolton
formicomorph group
Subfamily Aenictinae Emery
Subfamily tFormiciinae Lutz
Subfamily Aenictogitoninae Ashmead
Subfamily Formicinae Latreille
Subfamily Ecitoninae Forel
Tribe Dimorphomyrmecini Emery""
Tribe Cheliomyrmecini Wheeler
Tribe Myrmecorhynchini Wheeler
Tribe Ecitonini Forel
plagiolepidite tribal group
Subfamily Dorylinae Leach
Tribe Lasiini Ashmead
myrmeciomorph group
Tribe Plagiolepidini Forel
Subfamily Myrmeciinae Emery
Tribe Myrmoteratini Emery
Tribe Myrmeciini Emery
formicite tribal group
Tribe Prionomyrmecini Wheeler
Tribe Oecophyllini Emery
Subfamily Pseudomyrmecinae Smith
Tribe Gigantiopini Ashmead
myrmicomorph group
Tribe Camponotini Forel=
Subfamily Agroecomyrmecinae Carpenter
Tribe Notostigmatini Bolton
Subfamily Myrmicinae Lepeletier de Saint Fargeau
Tribe Formicini Latreille
Tribe Stegomyrmecini Wheeler
Tribe Melophorini Forel
Tribe Mytmecinini Ashmead
Subfamily Aneuretinae Emery
Tribe Metaponini Forel
Subfamily Dolichoderinae Forel
Tribe Melissotarsini Emery
'■Bolton's (2003) informal groups of subfamilies as well as informal tribal groups are employed herein. As continued phylogenetic work
refines the higher classification of the Myrmicinae and Formicinae, these tribal groups may warrant formalization as supertribes. If so,
then we recommend that the suffix -iti be employed for the supertribal rank as has been done for other aculeate lineages (e.g., Engel, 2005).
In advance of this, we have used informal names for these tribal groups based on this suffix as to avoid confusion when referring infor-
mally to a particular tribe (i.e., to avoid confusion caused by, for example, using "attine" for members of the attine tribal group [which
might also include Blepharidatta] or for actual members of the Attini).
""The subfamily Apomyrminae Dlussky and Fedoseeva (formerly in the leptanillomorph group, sensu Bolton, 2003) is herein considered
a synonym of Amblyoponinae as proposed by Saux et al. (2004).
' In Grimaldi and Engel (2005) this name appeared as "Dacetonini" based on a misinterpretation of the Greek root on the part of MSE.
The name is taken from the Greek word daketon, meaning "biting animal". MSE erroneously believed the word to terminate in Greek as
5aKeTo>v which would result in an augmented stem and require the retention of the terminal "-on" in the family-group name (apparently
Forel, 1892, himself, believed this as well since he originally proposed the name as "Dacetonini", and innumerable myrmecologists also
used this form, alongside the form "Dacetini", until recently). However, the original Greek terminates with omicron (rather than omega),
i.e., SukEtov, which necessitates the dropping of the "-on" and leaving the combining stem as "dacet-". Hence the family-group name is
correctly spelled as Dacetini (in this case with the tribal suffix). The name appears here in its correct form and will so in future editions of
Grimaldi and Engel (2005)
"■Bolton (2003) used the junior name Gesomyrmecini since the type genus of Dimorphomyrmecini (i.e., Diinorphomynnex) is a syn-
onym of Gejomynnex. However, ICZN (1999: Art. 40.1) states that a family-group name cannot be rejected even if its type genus is con-
sidered a junior synonym of another genus. Since Dimorphomyrmecini dates from Emery (1895: as Dimorphomyrmii) and Gesomyrmeci-
ni from Ashmead (1905: as Gesomyrmicinae), the former name should be employed for the tribe.
Tlie subfamily tPalaeosminthurinae has recently been synonymized with Camponotini (Snelling, in press).
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
cedure outlined by Nascimbene and Silver-
stein (2000).
SYSTEMATIC PALEONTOLOGY
Family Formicidae Latreille
Diagnosis: Head prognathous; dorsal rim
of torulus often tuberculate or concealed un-
der vertical lamella of frons; antenna genic-
ulate. Primitively with anterior margin of
clypeus spiculate (apomorphically lost in
many 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 gland
present 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 within
the family and likely part of the familial
ground plan). Metasoma petiolate; first me-
tasomal segment forming true node (strongly
constricted anteriorly and posteriorly); first
metasomal sternum separated from second
metasomal sternum by deep constriction.
Morphologically distinct, sterile worker caste
typically present* (i.e., advanced eusocial);
reproductives typically macropterous, work-
ers apterous. Workers with pronotum fused
to 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 including
11,833 species (Bolton, 1995; www.
antbase.org) have a cosmopolitan distribution
and are among the most recognizable of all
insects. Numerous species exist in Cenozoic
deposits around the world and are relatively
commonly encountered. Species of Creta-
ceous formicids, which are very rare, are
briefly outlined in appendix 1.
Bolton (2003) considered the antlike
wasps of the Cretaceous family Armaniidae
to represent the basalmost subfamily of the
ants. We have, however, retained armaniids
at the family rank and as the sister group
•* Some inquilines have apomoqjhically lost the work-
er caste.
(perhaps paraphyletic?) to Formicidae {vide
table 1 and appendix 1).
It is interesting to note that many primitive
ants have clypeal spicules with rounded api-
ces (e.g., '\Sphecomyrmodes, 'IMyanmyrma;
Amblyoponinae). Apomyrma has similar
spicules, but these are located on the labrum
rather 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 functional
convergence).
Subfamily fSPHECOMYRMiNAE
Wilson and Brown
fSphecomyrmodes, new genus
Type Species: '\Sphecomyrmodes oriental-
is, new species.
Diagnosis: Distinguished from all other
species of the tribe Sphecomyrmini by the
minute, peglike denticles running along the
entirety of the anterior margin of the clypeus
and from fSphecomyrma by the absence of
a medial extension or process on the clypeal
margin.
Etymology: The new genus-group name
is a combination of '\Sphecomyrma, type ge-
nus of the subfamily, and the suffix-ofi?e5,
meaning "with the form of". The name is
masculine.
'fSphecomyrmodes orientalis, new species
Figures 3-4
Diagnosis: As for the genus {vide supra).
Description: Head. Relatively large,
height of head slightly less than length of
alitrunk. Length of head 1.23 mm (with man-
dibles closed). No apparent microsculpture
on cuticle of head. Clypeus setose; setae of
moderate length and widely separated. Man-
dible simple, with only two teeth; outer sur-
face with numerous, widely scattered, fine
setae. Antenna of moderate length, with
scape short, funicular article I (pedicel)
shortest antennal article, funicular article II
the longest article of funiculus. Lengths of
antennal articles (in mm): scape 0.23, pedicel
(funicular article [fa] I) 0.13, fall 0.32, falll
0.15, falV 0.15, faV 0.15, faVI 0.15, faVII
0.17, faVIII 0.17, falX 0.17, faX 0.17, faXI
0.27. Mesosoma. Mesosomal length 1.33
AMERICAN MUSEUM NOVITATES
NO. 3485
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, with
scattered fine, short setae on all visible sur-
faces, those on propodeum about twice as
long as other setae. Coxae large, slightly in-
flated, ventrally setose, setae numerous and
fine. Legs moderate length. Foreleg with tar-
somere I distinctly longer than combined
lengths of more distal tarsomeres; tarsomere
I with "antennal cleaner" (strigil) a velvety
notch on ventral margin of proximal end; cal-
car present, length slightly longer than great-
est width of profemur, ventral margin with
row of fine teeth and (apically) setae. Patch
of dense, elongate setae opposite strigil on
profemur; inner posterior surface of protibial
apex with three stout, spinelike setae minute-
ly curved inward at their extreme apices (vis-
ible on left foreleg). Pairs of stiff setae on
ventral surface of protarsomeres: tarsomeres
I-III with three pairs, IV with two small
pairs. Pretarsal claw with minute subapical
tooth. Metasoma. Attachment of petiole to
propodeum not particularly thick; thickness
(measured in lateral view) of anterior end of
petiolar peduncle 0.3 X greatest depth of pro-
podeum. Petiole length 0.38 mm, height 0.37
mm. Preserved portion of gaster 1.73 mm in
length. Integument without apparent micros-
culpturing, with scattered, fine setae. Distal-
most metasomal segments torn apart, al-
though sting bulb and sting well preserved
beneath metasoma (fig. 4).
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
Type Material: Holotype. AMNH Bu-
351, an incompletely preserved worker in a
piece of reddish-orange amber, from Myan-
mar. Collected in Kachin state, Tanai village,
on Ledo Road, 105 km NW Myitkyna, via
Leeward Capital Corp., 1999.
Etymology: The specific epithet is the
Latin word orientalis, meaning "of the east"
and is a reference to this being the first spe-
cies of the tribe fSphecomyrmini (sensu Bol-
ton, 2003) recorded from Myanmar.
Genus '\Sphec.omyrma Wilson and Brown
'\Sphecomyrma Wilson and Brown, In Wilson et
al., 1967: 8. Type species: '\Sphecomyrma freyi
Wilson and Brown, 1967, monobasic and orig-
inal designation.
Diagnosis: Scape short; funiculus long
and filiform, about four times length of
scape; promesothoracic suture complete and
well developed; trochantellus absent; petiole
with distinct, domed node widely separated
from propodeum and remainder of metasoma
by deep constrictions; cuticle without sculp-
turing, superficial microscopic relief, with
scattered and sparse setae.
Comments: The genus, which is defined
largely by plesiomorphies, is doubtfully
monophyletic. It contains three species:
'fSphecomyrma canadensis Wilson in Cana-
dian amber, and t^. freyi Wilson and Brown
and a new species in New Jersey amber (vide
infra). In addition, we provide information
from newly identified material of fS. freyi.
Key to Species of 'fSphecomyrma
(based on the worker caste)
1. Anterior margin of clypeus with short, broad
extension, surface with two, long setae at
most; compound eye round 2
— Anterior margin of clypeus with long, medial
lobe (fig. 5), surface with numerous, long
setae; compound eye oval [New Jersey am-
ber]
.... t-^- mesaki Engel and Grimaldi, n.sp.
2. Third antennal article slightly more than
twice as long as second article [New Jersey
amber]
t-^- fi'syi Wilson and Brown
— Third antennal article about as long as second
article [Canadian amber]
t^*- canadensis Wilson
'fSphecomyrma mesaki, new species
Figure 5
Diagnosis: Distinguished from all other
species of the genus by the median portion
of the clypeus having a long ventral lobe,
length of the clypeus through the lobe is
0.46 X the greatest width of clypeus; clypeus
setose; and scape very short (1.2X the length
of longest funicular article). The species can
be further distinguished from 'fS. freyi (the
other 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 t5.
freyi); and a large head (length of head/
length of mesosoma = 0.83, vs. 0.65 in fS.
freyi).
Description: Petiole and gaster not pre-
served, so only head, mesosoma, and legs
preserved. Head. Large, length of head
slightly less than length of alitrunk. Length
of head 2.20 mm (with mandibles closed);
width of head 1.95 mm; length of eye 0.66
mm. No microsculpture on cuticle of head.
Vertex with fine, sparse pilosity, setae ca. 0.2
mm long. Ocelli present, median ocellus sit-
uated just above dorsal tangent of compound
eyes. Face bare. Bases of antennae situated
in shallow, broad scrobes; length of scrobe
about equal to length of scape and articulat-
ing base. Eyes well developed, bare, situated
well above bases of antennae; gena deep.
Lateral portions of clypeus quadrate; median
portion distended into long ventral lobe that
extends to ventral margin of closed right
mandible. Clypeus setose, except on middle
part. Mandibles simple, with only two teeth.
Antennae of moderate length, with scape
short, funicular article I (pedicel) shortest an-
tennal article, funicular article II the longest
article of funiculus. Lengths of antennal ar-
ticles (in mm): scape 0.53, pedicel (funicular
article [fa] I) 0.20, fall 0.43, falll 0.35, falV
0.30, faV 0.23, faVI 0.23, fa VII 0.30, faVIII
0.22, falX 0.26, faX 0.30, faXI 0.33. IVIeso-
soma. Mesosomal length 2.66 mm; without
microsculpturing, except at posterolateral
margin of promesonotal suture, where eight
fine grooves occur. Dome of promesonotum
setose; several fine setae on dorsal surface of
metanotum and propodeum. Coxae large, in-
flated, ventrally setose. Attachment of petiole
AMERICAN MUSEUM NOVITATES
NO. 3485
,».l'V-rrC
metapleural prapodeunrt
gland opening
Fig. 5. Holotype worker of '\Sphecomyrma tnesaki, new species (AMNH NJ-1023); lateral habitus
and facial view.
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
to propodeum not particularly thick; thick-
ness (measured in lateral view) of anterior
end of petiolar peduncle 0.3x greatest depth
of propodeum. Metapleural gland opening
(MGO) and MG bulla obvious, situated on
posterolateral part of propodeum just above
metacoxa. MGO small, with groove running
between it and extended almost to vental
margin of propodeum. Legs moderate length.
Foreleg with tarsomere I slightly longer than
combined 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 with
3 pairs. III with 3 small pairs, IV with 2
small pairs. Pretarsal claw with subapical
tooth. Metasoma not preserved.
Type Material: Holotype. AMNH NJ-
1023, an incompletely preserved worker in a
piece of amber barely larger than the ant,
from Sayreville, New Jersey (Middlesex
Co.), White Oaks outcrop, coll. Bob Mesak
(fig. 5). The amber is an irregularly shaped
drop, 7X5X4 mm, made of clear yellow
amber. Portions of some appendages are
breached at the surface, and the petiole and
gaster were similarly lost at the surface.
Since very little amber exists between the ant
and surface of the amber piece, no trimming
or polishing was possible. Still, details
through the rough surface are highly visible
by immersing the piece in glycerine.
Etymology: The specific epithet is a pat-
ronymic for Bob Mesak, who collected and
donated this valuable specimen to the
AMNH.
'\Sphecomyrm.a freyi Wilson and Brown
Figure 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 from
New 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 tol4Xl5X4 mm.
The piece contains two workers of t^. freyi.
Descriptive Notes: Both workers are
largely but not completely preserved. Spec-
imen A has the frontal half of the head miss-
ing; most of the antenna is present except for
bases of the scapes; the entirety of the re-
mainder of the body is preserved and the
sting appears largely or fully extruded. Spec-
imen B has the dorsal and apical part of the
gaster missing, as well as portions of the
right hind leg; the anterior third of the head
is 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 mesosoma
1.39 mm (B), 1. 40 mm (A); length of petiole
0.29 mm (B), 0.33 mm (A); length of gaster
(A) 1.72 mm; length of extruded portion of
sting (A), 0.33 mm. Measurements of anten-
nal articles (as measured for right antenna in
A, 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 of
amber is highly significant and addresses
questions of the social behavior of primitive
ants like '\Sphecomyrma. As reviewed in Gri-
maldi et al. (1997), Dlussky (1987, 1988),
and Poinar et al. (1999) questioned whether
'ISphecomyrma was a true ant since it had
such a short scape [but see response to Poin-
ar et al. (1999) by Grimaldi and Agosti
(2000b)]. With such a short scape, Dlussky
argued that it would be impossible for
'\Sphecomyrma to antennate, and therefore it
was considered highly unlikely for fSphe-
comyrma to have been social. Among the ap-
proximately 1700 fossiliferous pieces of New
Jersey amber in the AMNH collection thus
far, four pieces contain worker or male sphe-
comyrmine ants. These ants are (and proba-
bly originally were) exceedingly rare, and the
probability that two workers would be pre-
served in one piece by chance alone is ex-
tremely remote. It is most parsimonious to
explain the occurrence of two workers in the
same piece as a result of social behavior.
'ISphecomyrma sp.
Material: AMNH NJ-942, a male in am-
ber from New Jersey: Middlesex Co., Say-
10
AMERICAN MUSEUM NOVITATES
NO. 3485
Fig. 6. Two workers of fSphecomynna freyi Wilson and Brown preserved in a single piece of New
Jersey amber (AMNH NJ-943).
reville. White Oaks outcrop. Collected by the
late Steve Swolensky. Specimen is in a clear
yellow piece of amber 5X7 mm, embedded
in epoxy and trimmed to 1.5 mm thickness
for a full lateral view of the ant.
Comments: Venation and other details of
this specimen are indistinguishable from
AMNH NJ-242, described and figured by Gri-
maldi et al. (1997) as '\Sphecomyrmal sp.
Measurements: Length of hind wing (forewing
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
11
TABLE 2
Measurements of Antennal Articles for
fSphecomyrma freyi (AMNH NJ-943)
Specimen
Antennal article
A»
Bi'
Scape
—
0.33 mm
fal (pedicel)
0.16 mm
0.12 mm
fa2
0.28 mm
0.24 mm
fa3
0.16 mm
0.16 mm
fa4
0.16 mm
0.16 mm
feS
0.16 mm
0.16 mm
m
0.16 mm
0.17 mm
fa7
0.16 mm
0.18 mm
&8
0.16 mm
0.17 mm
fe9
0.17 mm
0.17 mm
falO
0.16 mm
0.17 mm
fall
0.23 mm
0.26 mm
^Measurements taken from right antenna.
''Measurements taken from left antenna,
fa = funicular article.
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.
Genus '\Haidomyrmex Dlussky
'\Haidomyrmex Dlussky, 1996: 84. Type species:
'\Haidomyrmex cerberus Dlussky, 1996, mono-
basic and original designation.
Diagnosis: Clypeus a small, hemispheric
lobe 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 a
fine point apically. Compound eyes small,
length ca. 0.2 X length of head capsule; ocelli
absent. Mandibles elongate, scimitar-shaped,
without serrations or teeth. Propodeum
rounded in profile. Petiole one-segmented,
nodiform, with distinct constriction at artic-
ulation with remainder of metasoma. Known
from the worker caste only.
Comments: The head of ■\Haidomyrmex is
enigmatic (fig. 7) and much of the detail is
obscured below the clypeus and where the
mandibles articulate. Indeed, it is likely that
there has been significant 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, fine, stiff trigger setae lie on the inside
surface of the mandibles and on the oral mar-
gin. '\Haidomyrmex has no such setae, so per-
haps the clypeal brush functioned analogous-
ly. How 'fHaidomyrmex 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.
'fHaidomyrmex cerberus Dlussky
Figures 7-8
fHaidomyrmex cerberus Dlussky, 1996: 85.
Holotype: NHML In. 20182, partial work-
er specimen in amber from Myanmar.
Comments: We have tentatively followed
past authors in placing fHaidomyrmex within
tSphecomyrminae (e.g., Bolton, 2003).
However, it is important to note the signifi-
cant similarities between this genus and
'\Brownimecia in New Jersey amber iyide in-
fra). Both genera have large, dome-shaped
heads, with relatively small compound eyes,
lack ocelli, have large mandibles devoid of
serrations or dentition, and an elongate an-
terior extension, or collar, of the pronotum.
TABLE 3
Leg Measurements (in mm) for fSphecomyrma freyi (AMNH NJ-943)
Femur
Tibia
Tarsomeres
1
2
3
4
5
Fore
Mid
Hind
0.82
0.97
1.20
0.73
0.83
1.10
0.49
0.58
0.74
0.13
0.20
0.29
0.12
0.17
0.19
0.08
0.13
0.14
0.13
0.19
0.20
12
AMERICAN MUSEUM NOVITATES
NO. 3485
£W0
1,0mm
Fig. 7. Facial view of holotype worker of '\Haidomyrmex cerberus Dlussky (NHML ln.20182).
Subfamily I-Brownimeciinae Bolton
Genus fBrownimecia Grimaldi, Agosti,
and Carpenter
'\Brownimecia Grimaldi, Agosti, and Carpenter,
1997: 20. Type species: '\Browmmecia clavata.
Grimaldi, Agosti, and Carpenter, 1997, mono-
basic and original designation.
Diagnosis: Antenna distinctly clubbed,
apical funicular article twice the width of
basal ones and pedicel. Ocelli absent. Man-
dibles long, thin, scimitar-shaped, strongly
cruciate, 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 third
segment (also known as abdominal segments
III and IV; gastral segments 1 and 2). Known
from the worker caste only.
'IBrownimecia clavata Grimaldi, Agosti,
and Carpenter
'\Brownimecia clavata Grimaldi, Agosti, and Car-
penter, 1997: 20.
Material: AMNH NJ-941, in amber from
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
13
procoxa
Fig. 8. Lateral view of mesosoma and petiole, as preserved, of holotype worker of '\Haidomyrmex
Cerberus Dlussky (NHML In.20182).
New Jersey: Middlesex Co., Sayreville,
White Oaks outcrop, collected by Steve
Swolensky.
Comments: A beautifully preserved spec-
imen in clear yellow amber, 5X8 mm,
which was embedded in epoxy and trimmed
to 2 mm thickness. Like the holotype which
was described in 1997, it is curled up and the
sting is extruded. The piece also contains
some wood fragments and a frass pellet. The
length of head (including closed mandibles)
0.88 mm, greatest width of head 0.74 (be-
tween outer margins of eyes), length of eye
0.24 mm, length of trunk 0.94 mm, length of
petiole 0.32 mm. The mandibles are tightly
closed, so details are less visible than in the
holotype. The front portion of the head, how-
ever, and especially the clypeus, are more
visible. 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 rare
in 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 of
the formicid ground plan.
Subfamily Incertae Sedis
'\Myanniyrma, new genus
Type Species: ■\Myanmyrma gracilis, new
species.
Diagnosis: Gracile "poneroid" ant with
extremely long legs; mandibles nearly equal
to 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, second
article longest; articles slightly shorter api-
cally; integument of head spinose; clypeus
deeply incised along anterior margin and
with distinct peglike setae; eyes and ocelli
not apparent (but may be present); pre tarsal
claws with minute tooth; petiole with distinct
nodus; gastral constriction distinct; sting well
developed. Easily distinguished from fHai-
domyrmex, another highly modified ant in
Burmese amber, by the mandibular and clyp-
eal structure, head microsculpturing, meta-
somal constriction, long genal processes, and
very long legs.
Etymology: Taken directly from Myan-
mar, the country of the amber's origin; and-
14
AMERICAN MUSEUM NOVITATES
NO. 3485
myrma, a common suffix for ant genera. The
name is feminine.
'fMyanmyrma gracilis, new species
Figures 1, 9
Diagnosis: As for the genus {vide supra).
Description: Specimen has been com-
pressed, but body form was clearly extremely
gracile, with very long legs. Head. Propor-
tions difficult to determine due to compres-
sion; length ca. 1.76 mm. Front of head with
pair of carinae diverging dorsally; blunt
spines 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 median
emargination; clypeus with two lobes, mar-
gin of two lobes with row of ca. 14 minute
denticles on each lobe, labrum similarly bi-
lobed and denticulate. Lateral portion of
clypeus lobed, without denticles. Gena with
spatulate process, exact length difficult to
discern, but ca. 0.3 X length of mandible.
Mandibles long, 1.61 mm. (right one), nearly
equal to length of head, sickle-shaped; apical
tooth long, sharp; blunt subapical tooth, no
other teeth. No fine setae apparent. Apex of
right mandible longer than left, especially
apical tooth. Eyes well developed (difficult
to observe since they are sunken in a cavity),
length ca. 0.25 X length of head. Antenna
very long (7.2 mm); scape fairly short, with
wide base; funicular article (fa) 1 shortest,
fa2 longest; lengths of antennal articles (in
mm): scape 0.63, funicular article (fa) [ped-
icel] I 0.44, fall 1.05, falll 0.62, falV 0.53,
faV 0.61-faVI 0.45, faVII 0.40, faVIII 0.41,
falX 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 with
dorsal, preapical brush of fine setae; one
large, two finer apical spurs; ventral surface
of probasitarsus with fine pilosity, six pairs
of stiff setae; all tarsomeres with four apical
spines; each pretarsal claw with median
tooth. Mesotibia with pair of ventroapical
spurs; ventral surface of mesobasitarsus with
three rows of 10 stiff setae each; metatibia
with pair of apical spurs, one larger and pec-
tinate; ventral surface of metabasitarsus with
fine 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 portion
of petiole narrow, tubular; deep constriction
between second and third metasomal seg-
ments. Metasoma overall quite small, ca.
0.3x length of body (excluding antennae).
Terga and sterna telescoped in specimen, as
shown in figure 9; gaster (i.e., metasoma ex-
cluding petiole) approximately 2.95 mm
long. Sting well developed, only partly ex-
truded but internally visible; pygidium with
long fine setae.
Holotype: AMNH Bu-014, a worker, in
amber from northern Myanmar (Burma)
(figs. 1, 9). Collected in Kachin state, Tanai
village, on Ledo Road, 105 km NW Myit-
kyna, via Leeward Capital Corp., 1999. The
ant is in a transparent yellow piece of amber,
which was originally more than 20 mm in
diameter and 30 mm in length. The piece was
occluded with fractures and debris, so it re-
quired epoxy embedding and then trimming
to better observe the ant. The piece is now
rhomboid-shaped, 17 X 22 X 4 mm, the
broad surfaces being parallel to the lateral
surface of the ant. A flat edge also permits a
frontal view of the head. Other inclusions in
the piece are numerous frass pellets and
wood fragments, further suggestive (besides
body form) of arboreal/wood nesting habits
of the ant. The ant specimen shows a great
deal of distortion due to compression. The
cuticle is transparent, facilitating observation
of some internal structures (such as unex-
truded portions of the sting), but some com-
pressed surfaces could easily be mistaken for
flanges and other structures.
Additional Material: Two additional,
fragmentary specimens are perhaps represen-
tative of tAf. gracilis; however, because of
their poor preservation, we hesitate to des-
ignate them as paratypes. Both are preserved
in amber from northern Myanmar and with
the same collection data as the holotype.
AMNH Bu-225, a poorly preserved worker
but very similar to holotype in observable
traits. AMNH Bu-1509, a badly compressed
worker preserved in yellow amber; head
mostly crushed and difficult to discern, me-
tasoma similarly compressed; somewhat
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
15
-^if^r
Fig. 9. Holotype worker of 'fMyanmyrma gracilis, new genus and species (AMNH Bu-014); facial
view and general habitus.
16
AMERICAN MUSEUM NOVITATES
NO. 3485
TABLE 4
Leg Measurements" of Holotype Worker of fMyaninyrina gracilis (in mm)
Femur
Tibia
Tarsomeres
1
2
3
4
5
Claw
Fore
Mid
Hind
2.33
3.10
3.67
1.50
2.22
3.23
1.47
2.16
2.49
0.52
0.82
0.42
0.72
0.31
0.47
0.38
0.55
0.28
0.22
"As measured on the left legs.
smaller than holotype and Bu-225 but still
exhibiting the same elongate, slender legs,
constriction in first gastral segments, etc.,
this individual may represent a minor worker.
Etymology: The specific epithet is the
Latin word gracilis, meaning "slender", as
a reference to elongate legs and structure of
this ant.
Comments: Constriction of the metasoma
indicates the specimen is a "poneroid",
which is the third and oldest Cretaceous re-
cord of this paraphyletic grade of primitive
ants (the poneroid grade includes the poner-
omorph and myrmeciomorph subfamilies of
Bolton, 2003). The other Cretaceous "pone-
roids" are '\Brownimecia clavata (in New
Jersey amber: Turonian), and fCanapone
dentata Dlussky (in Canadian amber: Cam-
panian). It is interesting that fMyanmyrma is
only the fourth record of ants in Burmese
amber, but two of these records are for highly
modified species. '\Burmomyrma rossi Dlus-
sky is based on a single, alate, incomplete
specimen, with head and portion of the ali-
trunk missing. The most distinctive feature
of '\Burmoniyrma is the wing with highly re-
duced venation; thus, the taxonomic concept
of this genus is not entirely comparable with
that of the other two Burmese amber genera.
'\Burmomyrma, as discussed again later (vide
infra), has been tentatively placed in the
Aneuretinae (Dlussky, 1996; Bolton, 2003).
Both '\Haidomyrmex and 'fMyanmyrma are
extremely gracile and relatively large ants,
with large, highly modified mouthparts and
genae. The extremely long legs and slender
body are analogous to Leptomyrmex (Doli-
choderinae) and Oecophylla (Formicinae),
the latter of which is entirely arboreal. fHai-
domyrmex and '\Myanmyrma presumably
had similar habits. 'fHaidomyrmex and
'\Myanmyrma are clearly not closely related,
the former placed in the fSphecomyrminae
by Dlussky. The type and only specimen of
'IHaidomyrmex, however, has only the sec-
ond metasomal (i.e., first gastral) segment
preserved, 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 antenna
of '\Myanmyrma are highly significant. Fu-
nicular article 2 is the longest article in the
funiculus, as is the case for most fSpheco-
myrminae (except t^. canadensis Wilson).
This condition does not exist in the other
Cretaceous "poneroids", '\Brownimecia and
fCanapone. A long funicular article 2 was
proposed by Grimaldi et al. (1997: 8) as one
of only two or three possible synapomor-
phies for the fSphecomyrminae. If truly apo-
morphic, the long funicular article 2 would
have been independently derived in fMyan-
myrma. If a long funicular article 2 was ple-
siomorphic, the monophyly of the fSpheco-
myrminae would be seriously doubtful. Out-
group evidence from closely related aculeate
families suggests that, indeed, this feature is
symplesiomorphic for basal ants, including
'fMyanmyrma.
Placement of fMyanmyrma in a subfamily
is 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 the
Myrmeciinae (myrmeciomorph). Neither
placement is entirely satisfactory, but we be-
lieve inclusion in the latter subfamily is more
likely (although we tentatively retain the ge-
nus as subfamily incertae sedis). Like myr-
meciines, fMyanmyrma has a metasomal
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
17
constriction, the antennal sockets inclined to
nearly a vertical position, 12-segmented an-
tennae, pectinate inner metatibial spur, and
elongate mandibles. Unlike typical myrme-
ciines, however, the the new genus has man-
dibles that are sickle-shaped, with dentition
only at the apices; has a deeply incised clyp-
eal margin, with distinct spicules on the
lobes; has a spinose integument on the head;
has gracile legs; and elongate genal process-
es. These traits are, however, autapomorphic
and do not preclude placement in Myrmeci-
inae. Within Myrmeciinae, '\Myanmyrma
would 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 and
weakly so in '\Prionomyrmex, perhaps being
apomorphically lost in Nothomyrmecia. The
presence of spicules along the clypeal margin
in '\Myanmyrma is an interesting and enig-
matic feature that should be further explored
for its systematic implications.
This is the species that is referred to by
Grimaldi and Engel (2005) as "undescribed
Myrmeciinae?"
Subfamily Aneuretinae? Emery
fCananeuretus, new genus
Type Species: '\Cananeuretus occidentalis,
new species.
Diagnosis: Compound eyes present, small;
ocelli absent. Antennal sockets slightly in-
clined; scape elongate. Mandible of primitive
construction, with four distinct teeth (fig. 10),
basalmost tooth largest. Preoccipital carina
absent; ocelli absent. Alitrunk elongate, slen-
der; pronotal neck elongate. Meso- and me-
tatibia with a single spur; pretarsal claws
simple. Propodeal lobes, denticles, and
spines absent; petiole one-segmented, ante-
rior peduncle elongate, with distinct tubular,
postnodal section (more elongate than in
Aneuretus) articulating high on anterior sur-
face of second metasomal segment, petiolar
tergum and sternum fused; nodus not very
high. Metasoma excluding petiole short and
globular; without U-shaped emargination on
anterior margin of second metasomal seg-
ment; sting present and well developed.
Known from the worker caste only.
Etymology: The new genus-group name
is a combination of Canada, the country from
which this amber originates, and Aneuretus,
type genus of the Aneuretinae. The name is
masculine.
Comments: ■fCananeuretus might at first
be confused for fEotapinoma, a dolichoder-
ine described from Canadian amber (Dlus-
sky, 1999). '\ Cananeuretus differs notably
from 'lEotapinoma in the presence of a well-
developed sting (apparently vestigial in
'\Eotapinoma, like all dolichoderines), the
more elongate petiole with a distinct nodus
(shorter petiole, which does not have an
elongate peduncle and without developed no-
dus in fEotapinoma), and the reduced com-
pound eyes (large in '\Eotapinoma).
This new genus is tentatively placed in
Aneuretinae, although the group shows most-
ly what are presumed plesiomorphies for the
subfamily. The presence of a strong, well-
developed sting and absence of an acidopore
excludes placement in Dolichoderinae or
Formicinae. However, the genus is best
placed among the formicomorph subfamilies
(sensu Bolton, 2003) where it most closely
approximates the Aneuretinae. As more ma-
terial is recovered of these ants and a cladis-
tic framework for fossil aneuretines
achieved, it is possible that '\ Cananeuretus
and others may prove to be stem-group
Aneuretinae or even represent stem groups to
Aneuretinae + Dolichoderinae.
Today the subfamily Aneuretinae consists
of a single species occurring in Sri Lanka.
During the Tertiary aneuretines were distrib-
uted widely in the Northern Hemisphere as
evidenced by their occurrence in Baltic am-
ber (Wheeler, 1914) and Florissant, Colo-
rado (Carpenter, 1930). The mid-Cretaceous
genus fBurmomyrma, in amber from Myan-
mar, has been tentatively placed in the
Aneuretinae (Dlussky, 1996; Bolton, 2003)
and, until now, represented the sole Meso-
zoic record for the subfamily. It is increas-
ingly apparent that aneuretines were widely
distributed during the Mesozoic and early
Tertiary, and that Aneuretus is a notable rel-
ict in Sri Lanka today. The significant ex-
tinction of northern aneuretines was perhaps
a result of the infamous Eocene-Oligocene
18
AMERICAN MUSEUM NOVITATES
NO. 3485
\^/f
H
50 mm
Fig. 10. Holotype worker of 'fCananeuretus occidentalis, new genus and species (TMP 8.89.7).
climatic shift, a cooling event that dramati-
cally altered the evolutionary history and
biogeography of numerous insect lineages
that are today austral relicts (Grimaldi and
Engel, 2005).
\Cananeuretus occidentalis, new species
Figures 2, 10-1 1
Diagnosis: As for the genus {vide supra).
Description: Body with sparsely scattered.
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
19
pedicel
troctiBntellus
Fig. 11. Partial worker of '\Cananeuretus occidentalism (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, basalmost
apical tooth largest. Eyes present, relatively
small and low on head (difficult to observe).
Scape fairly long (0.46 mm), narrowed
slightly at base for articulation with bulb (fig.
10); pedicel 0.65 X length of scape. Meso-
soma. Elongate, not deep, length ca. 3.10
mm; pronotum forming a distinct neck an-
teriorly. Legs slender and long; trochantellus
present (fig. 11). Inner surfaces of protibia
and 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 metatibia
with a single spur; metatibial spur distinctly
and minutely setose (fig. 10). Propodeum
without lobes or spines, broadly rounded and
gently sloping to articulation with petiole.
Metasoma. Petiole with elongate peduncle.
distinct nodus, and relatively long posterior,
tubular section, articulating high on second
metasomal segment; petiolar tergum and
sternum fused; metasoma without constric-
tion between second and third segments (i.e.,
gastral segments 1 and 2); metasoma overall
quite globular. Sting well developed, extrud-
ed.
Holotype: TMP 8.89.7, labeled, "Grassy
Lake, Alberta, Campanian, Foremost For-
mation, Formicidae" (figs. 2, 10). Worker in
amber that is embedded in a thin block of
epoxy 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 slide
mounted. The specimen is partial and owing
to some minor differences in the shape of the
petiole, we have considered it best to only
tentatively assign it to this species.
Etymology: The specific epithet is the
20
AMERICAN MUSEUM NOVITATES
NO. 3485
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 the
manuscript and for correcting several inac-
curacies; to J. D. Gardner (TMP) for the loan
of specimens; and to A. J. Ross (NHML) for
hosting us during our 2002 visit to the De-
partment of Palaeontology at which time the
holotype of '\Haidomyrmex was studied. This
work was supported by NSF DBI-9987372
(to DAG) and by NSF EPSCoR grant
KAN29503 and NSF EF-0341724 (to MSE).
This is contribution Nr. 3416 of the Division
of Entomology, Natural History Museum and
Biodiversity Research Center, University of
Kansas.
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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 classificatory structure generally follows that of Bolton (2003). The fossil
Cretacoformica explicata Jell and Duncan (1986) from the Lower Cretaceous (Aptian) 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).
Taxon
Deposit
Age
Family FORMICIDAE Latreille
Poneromorph Group
Subfamily tSphecomyrminae Wilson and Brown
Genus ^Sphecomyrmodes Engel and Grimaldi, n.gen.
tS. orientalis Engel and Grimaldi, n.sp.
Genus ^Sphecomyrina Wilson and Brown
t5. canadensis Wilson
■\S.freyi Wilson and Brown
tS. mesaki Engel and Grimaldi, n.sp.
Genus ■fBaikuris Dlussky
tB. casei Grimaldi, Agosti, and Carpenter
tB. inandibularis Dlussky
tB. mirabilis Dlussky
Genus ■\Cretomyrina Dlussky
tC. amoldii Dlussky
tC. unicornis Dlussky
Genus ■\Dlusskyidris Bolton
to. zhericliini (Dlussky)
Genus ^Haidomyrmex Dlussky
t//. cerberus Dlussky
Subfamily Ponerinae? Lepeletier de Saint Fargeau
Genus ■\Afropone Dlussky, Brothers, and Rasnitsyn
tA. oculata Dlussky, Brothers, and Rasnitsyn
tA. orapa Dlussky, Brothers, and Rasnitsyn
Genus iCanapone Dlussky
tC. dentata Dlussky
Subfamily tBrownimeciinae Bolton
Genus iBrownimecia Grimaldi, Agosti, and Carpenter
tB. clavata Grimaldi, Agosti, and Carpenter
Myriueciomorph Group?
Subfamily Incertae Sedis (perhaps Myrmeciinae Emery)
Genus ■\Myanmyrnia Engel and Grimaldi, n.gen.
tM. gracilis Engel and Grimaldi, n.sp.
Myrmicomorph Group
Subfamily Myrmicinae Lepeletier de Saint Fargeau
Genus fAfromyrma Dlussky, Brothers, and Rasnitsyn
■^A. petrosa Dlussky, Brothers, and Rasnitsyn
Formicomorpb Group
Subfamily Incertae Sedis
Genus ■\Gerontoformica Nel and Perrault
tG. cretacica Nel and Perrault
Subfamily Formicinae Latreille
Genus ^Kyromyrma Grimaldi and Agosti
t^. rieffi Grimaldi and Agosti
Subfamily DoUchoderinae Forel
Genus ^Eotapinoma Dlussky
t£. macalpini Dlussky
Burmese (A)
Alb
Canadian (A)
New Jersey (A)
New Jersey (A)
Cam
Tur
Tur
New Jersey (A)
Taymyr (A)
Taymyr (A)
Tur
San
San
Taymyr (A)
Taymyr (A)
San
San
Taymyr (A)
Burmese (A)
Burmese (A)
San
Alb
Alb
Botswana (C)
Botswana (C)
Tur
Tur
Canadian (A)
Cam
New Jersey (A)
Tur
Burmese (A)
Botswana (C)
France (A)
New Jersey (A)
Canadian (A)
Alb
Tur
Alb
Tur
Cam
2005
ENGEL AND GRIMALDI: CRETACEOUS ANTS
23
APPENDIX
(Continued)
Taxon
Deposit
Age
Formicomorpb Group (continued)
Subfamily Aneuretinae? Emery
Genus ■\Burmomyrma Dlussky
tS. rossi Dlussky
Burmese (A)
Alb
Genus ■\Cananeurelus Engel and Grimaldi, n.gen.
tC. occidentalis Engel and Grimaldi, n.sp.
Canadian (A)
Cam
Family tARMANIIDAE Dlussky
bfamily tArmaniinae Dlussky
Genus -fArchaeopone Dlussky
■fA. kyddiarica Dlussky
Kazakhstan (C)
Alb
t/1. taylori Dlussky
Magadan (C)
Cen
Genus -fAnnania Dlussky (= ■fArmaniella Dlussky)
■fA. robusta Dlussky
Magadan (C)
Cen
■fA. capitata Dlussky
Ulya (C)
Alb
tA. pristiiia Dlussky
Ulya (C)
Alb
■fA. curiosa (Dlussky)
Magadan (C)
Cen
Genus fDolicliomyrma Dlussky
■fD. longiceps Dlussky
Kazakhstan (C)
Alb
Genus fKhetania Dlussky
^K. matidibulata Dlussky
Ulya (C)
Alb
Genus ■fOrapia Dlussky, Brothers, and Rasnitsyn
to. minor Dlussky, Brothers, and Rasnitsyn
Botswana (C)
Tur
to. rayneri Dlussky, Brothers, and Rasnitsyn
Botswana (C)
Tur
Genus -fPoiieropterus Dlussky
■fP. spliecoides Dlussky
Magadan (C)
Cen
Genus fPseudarmania Dlussky
fP. rasnitsyni Dlussky
Magadan (C)
Cen
■fP. aberrans Dlussky
Magadan (C)
Cen
ACULEATA Family Incertae Sedis
(nee Formicidae)
Genus -fCariridris Brandao and Martins-Neto"
tC bipetiolata Brandao and Martins-Neto
Genus fCretopone Dlussky""
■fC. magna Dlussky
Genus fPetropone Dlussky""
■fP. petiolata Dlussky
Santana (C)
Apt
Kazakhstan (C)
Alb
Kazakhstan (C)
Alb
"Verhaagh (1996) indicated a placement in Sphecidae (Apoidea) for ■fCariridis, while Rasnitsyn (in Rasnitsyn and Quicke, 2002)
suggested Ampulicidae (Apoidea).
"Orimaldi et al. (1997) indicated fCretopone and ■fPetropone to be best considered as Aculeata incertae sedis. More recently
Bolton (2003) has suggested that they may be genera of poneromorph ants but not assignable as to subfamily.
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