Brownimecia
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''Brownimecia'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s, the only genus in the
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...
Brownimeciini and subfamily Brownimeciinae of the Formicidae. Fossils of the single identified species, ''Brownimecia clavata'', are known from the
Middle Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of t ...
of North America. The genus is one of several ants described from Middle Cretaceous
amber Amber is fossilized tree resin that has been appreciated for its color and natural beauty since Neolithic times. Much valued from antiquity to the present as a gemstone, amber is made into a variety of decorative objects."Amber" (2004). In ...
s of
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...
. ''Brownimecia'' was initially placed in the subfamily
Ponerinae Ponerinae is a subfamily of ants in the Poneromorph subfamilies group, with about 1,600 species in 47 extant genera, including '' Dinoponera gigantea'' - one of the world's largest species of ant. Mated workers have replaced the queen as the ...
, until it was transferred to its own subfamily in 2003; it can be distinguished from other ants due to its unusual
sickle A sickle, bagging hook, reaping-hook or grasshook is a single-handed agricultural tool designed with variously curved blades and typically used for harvesting, or reaping, grain crops or cutting Succulent plant, succulent forage chiefly for feed ...
-like mandibles and other morphological features that makes this ant unique among the Formicidae. The ant is also small, measuring , and a stinger is present in almost all of the specimens collected. The morphology of the mandibles suggest a high level of feeding specialization.


History and classification

''Brownimecia'' is known from three adult fossils: the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
—specimen number AMNH NJ-667—collected by Yale Goldman; the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...
; and a third described in 2005. At the time of the genus description, the type specimens were residing in the American Museum of Natural History, in New York City. All the described specimens are
worker caste Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping genera ...
adult females which have been preserved as inclusions in transparent chunks of
New Jersey amber New Jersey Amber, sometimes called Raritan amber, is amber found in the Raritan and Magothy Formations of the Central Atlantic (Eastern) coast of the United States. It is dated to the Late Cretaceous, Turonian age, based on pollen analysis of t ...
. The amber specimens were recovered from deposits of the South Amboy Fire Clay, part of the
Raritan Formation The Raritan Formation is a Cretaceous (Turonian) geologic formation. Dinosaur remains are among the fossils that have been recovered from the formation, although none have yet been referred to a specific genus.Weishampel, et al. (2004). "Dinosaur ...
. New Jersey amber has been dated to approximately 90 to 94 mya, placing it in the
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded b ...
of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
. Analysis of the amber composition indicates it originated as cupressaceous resins which were deposited in lagoons and salt water marshes along the Cretaceous eastern seaboard. ''Brownimecia clavata'' is one of several ant species described from New Jersey amber, the others being '' Sphecomyrma freyi'', '' Sphecomyrma mesaki'', '' Baikuris casei'', and '' Kyromyrma neffi''. The type fossils were first studied by paleoentomologists David Grimaldi, Donat Agusti, and James Carpenter of the American Museum. The team's 1997
type description A species description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species that have be ...
of the new genus and species was published in the journal ''
American Museum Novitates ''American Museum Novitates'' is a peer-reviewed academic journal published by the American Museum of Natural History. It was established in 1921. According to the ''Journal Citation Reports'', the journal has a 2013 impact factor The impact f ...
''. The genus name was coined by them as a patronym honoring the ant systematist William L. Brown, Jr. who co-described the first Cretaceous ant genus and species ''Sphecomyrma freyi''. The specific epithet ''clavata'' was chosen as a reference to the distinctive clubbed antennae seen in the workers. Grimaldi, Agusti, and Carpenter placed the new genus into the extant ant subfamily
Ponerinae Ponerinae is a subfamily of ants in the Poneromorph subfamilies group, with about 1,600 species in 47 extant genera, including '' Dinoponera gigantea'' - one of the world's largest species of ant. Mated workers have replaced the queen as the ...
, without tribal assignment, based on a slight constriction of the gaster. They also noted several features that are shared with ants of the amblyoponine group, then a part of Ponerinae. The workers have genal spurs on the lower edge area under the small compound eyes. Workers also have narrow long mandibles, as do some amblyoponine genera. Placement in Ponerinae was unchanged until a revision of the subfamily was published in 2003 by myrmecologist
Barry Bolton Barry Bolton is an English myrmecologist, an expert on the classification, systematics, and taxonomy of ants, who long worked at the Natural History Museum, London. He is known especially for monographs on African and Asian ants, and for encyclo ...
. In that paper the subfamily was shown to be a paraphyletic grouping and many of the included genera were moved to separate subfamilies, including the amblyoponines and ''Brownimecia''. Due to the unique combination of morphological features found in ''Brownimecia'', Bolton erected the new subfamily Brownimeciinae for the genus. Bolton notes that its sickle-like mandibles which lack teeth are not usual for ants and are often associated with dulotic, or slave-making, behavior in extant genera. In contrast to stem group ants such as the sphecomyrmines, the scapes of ''Brownimecia'' workers are more elongated like those of crown group ants. In his 2007 phylogenetic examination of Formicidae, entomologist Philip Ward notes the possibility that ''Brownimecia'' is a crown group genus and placed the subfamily into the informal "poneroid" grouping in his phylogony.


Description

Both the ''Brownimecia'' holotype worker and the 2005 worker are complete and fully preserved in curled positions and have their stings extended. The paratype worker is very incomplete; the amber in which it is encased is highly likely to have been at least partly tumble polished in a rock causing the ant to be exposed and portions of the head, left antenna, mid legs, hind legs, and all of the right antennae and forelegs are gone. The exposure allowed the interior cavity of the fossil to be examined under scanning electron microscope after being coated in a layer of
gold Gold is a chemical element with the symbol Au (from la, aurum) and atomic number 79. This makes it one of the higher atomic number elements that occur naturally. It is a bright, slightly orange-yellow, dense, soft, malleable, and ductile me ...
. The holotype has an overall body length of approximately with finely scattered
setae In biology, setae (singular seta ; from the Latin word for " bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
on all parts except the
notum The notum (plural nota) is the dorsal portion of an insect's thoracic segment, or the dorsal surface of the body of nudibranch gastropods. The word "notum" is always applied to dorsal structures; in other words structures that are part of the back ...
and
propodeum The propodeum or propodium is the first abdominal segment in Apocrita Hymenoptera (wasps, bees and ants). It is fused with the thorax to form the mesosoma. It is a single large sclerite A sclerite (Greek , ', meaning " hard") is a hardened bod ...
. The head is large with small rounded
compound eyes A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
composed of about one hundred
ommatidia The compound eyes of arthropods like insects, crustaceans and millipedes are composed of units called ommatidia (singular: ommatidium). An ommatidium contains a cluster of photoreceptor cells surrounded by support cells and pigment cells. The ...
. The holotype does not clearly show whether the
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
are present or absent, but they are clearly absent in the 2005 specimen. The mandibles are
scimitar A scimitar ( or ) is a single-edged sword with a convex curved blade associated with Middle Eastern, South Asian, or North African cultures. A European term, ''scimitar'' does not refer to one specific sword type, but an assortment of different ...
-shaped with no teeth present on the interior sides and almost the length of the head. The mandibles cross each other near their mid-length and the oral surfaces bear approximately thirty short needle-like setae. The clypeus is short but wide, crossing almost the whole width of the head. Perpendicular crenelations run along the upper margin of the clypeus on the 2005 specimen. The
gena Gena (Amharic: ገና) or qarsa (ቃርሳ) is a traditional field hockey game popular in the Ethiopian highlands. It is a game played in the space between villages but with no defined boundaries. It is played among two teams who attempt to throw ...
have a distinct tooth-like structure on the lower sides. The eleven segmented antennae are distinct in having a definite clubbed tip, a feature not seen in any other described Cretaceous ants. The apical
flagellomere Antennae ( antenna), sometimes referred to as "feelers", are paired appendages used for sensing in arthropods. Antennae are connected to the first one or two segments of the arthropod head. They vary widely in form but are always made of one o ...
segment is nearly twice the width of the other segments, while the third to the sixth segments are the shortest and the second flagellomere is slightly swollen. The metathoracic spiracles are raised on small cones projecting from the
alitrunk The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings. In hymenopterans of t ...
, while the
metapleural gland Metapleural glands (also called metasternal or metathoracic glands) are secretory glands that are unique to ants and basal in the evolutionary history of ants. They are responsible for the production of an antibiotic fluid that then collects in a ...
orifices are not notably raised, having a ridge leading to them. All of the alitrunk gland and spiracle orifices are to some degree obscured on the holotype by froths of bubbles that escaped them soon after the worker was entombed in the sap. The petiole top forms a pointed apex instead of a more rounded dome. There is a covering of fine microtrichia hairs on the petiole and a pair of small setae are situated near the apex. At the front of the petiole there is a narrow attachment area to the propodeum, while the rear is flared into a large attachment to the gaster. The helcium, a small exoskeleton plate between the petiole and gaster, is developed and shows crenelations on the rear edge. There is a small but distinct constriction between the first and second segments of the gaster. Both the holotype and 2005 specimens have a fully preserved gaster tip showing the extended sting, while the paratype is missing the tip area. Due to the unusual morphology of the mandibles, scientists have suggested that ''Brownimecia'' may have had a high level of feeding specialization.


References


External links

* {{Good article Brownimeciinae Monotypic fossil ant genera Cretaceous insects Prehistoric insects of North America New Jersey amber Fossil taxa described in 1997 Fossil taxa described in 2003