Avitomyrmex
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''Avitomyrmex'' 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 bulldog ants in the subfamily
Myrmeciinae Myrmeciinae is a subfamily of the Formicidae, ants once found worldwide but now restricted to Australia and New Caledonia. This subfamily is one of several ant subfamilies which possess gamergates, female worker ants which are able to mate a ...
which contains three described species. The genus was described in 2006 from
Ypresian stage In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresian ...
( Early Eocene) deposits of
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
, Canada. Almost all the specimens collected are queens, with an exception of a single fossilised worker. These ants are large, and the eyes are also large and well developed; a sting is present in one species. The behaviour of these ants may have been similar to extant Myrmeciinae ants, such as foraging solitarily for
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
prey and never leaving
pheromone trail Trail pheromones are semiochemicals secreted from the body of an individual to affect the behavior of another individual receiving it. Trail pheromones often serve as a multi purpose chemical secretion that leads members of its own species towards ...
s to food sources. ''Avitomyrmex'' has not been assigned to any
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 ...
, instead generally being regarded as ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
'' within Myrmeciinae. However, its identity as an ant has been challenged, although it is undoubtedly a hymenopteran insect.


History and classification

''Avitomyrmex'' is an extinct genus of ants with three described species. Fossils of ''Avitomyrmex'', along with other extinct Myrmeciinae ants were first studied and described by Bruce Archibald, Stefan Cover and Corrie Moreau of the
Museum of Comparative Zoology A museum ( ; plural museums or, rarely, musea) is a building or institution that cares for and displays a collection of artifacts and other objects of artistic, cultural, historical, or scientific importance. Many public museums make thes ...
in
Cambridge Cambridge ( ) is a College town, university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cam ...
,
Massachusetts Massachusetts (Massachusett: ''Muhsachuweesut Massachusett_writing_systems.html" ;"title="nowiki/> məhswatʃəwiːsət.html" ;"title="Massachusett writing systems">məhswatʃəwiːsət">Massachusett writing systems">məhswatʃəwiːsət'' En ...
. They published their 2006 description of the genus and species in an ''
Annals of the Entomological Society of America Annals ( la, annāles, from , "year") are a concise historical record in which events are arranged chronologically, year by year, although the term is also used loosely for any historical record. Scope The nature of the distinction between ann ...
'' journal article. The genus name is a combination of the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
"''avitus''" meaning "ancient" or "grandfatherly" and the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
''myrmex'', meaning "ant". Included with the genus description, the paper contained the description of ''Avitomyrmex mastax'', ''Avitomyrmex systenus'', and the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
''Avitomyrmex elongatus''. These fossil species date back to the Middle Ypresian. Archibald and colleagues originally classified ''Avitomyrmex'' as ''incertae sedis'' (Latin for "of uncertain placement") within the ant subfamily Myrmeciinae, as the specimens are unable to be properly identified. In 2008, however, Cesare Baroni Urbani of the
University of Basel The University of Basel (Latin: ''Universitas Basiliensis'', German: ''Universität Basel'') is a university in Basel, Switzerland. Founded on 4 April 1460, it is Switzerland's oldest university and among the world's oldest surviving universit ...
, Switzerland, noted that no specimen in this genus allows a proper examination of the apomorphy (key diagnostic traits) of the subfamilial or familial characters. While Baroni Urbani excludes ''Avitomyrmex'' from Myrmeciinae and classifies it as ''incertae sedis'' in Hymenoptera, the morphological characters and wings show the specimens are undoubtedly a hymenopteran insect. A 2012 report by Russian palaeoentomologist Gennady M. Dlussky of the
Moscow State University M. V. Lomonosov Moscow State University (MSU; russian: Московский государственный университет имени М. В. Ломоносова) is a public research university in Moscow, Russia and the most prestigious ...
describing new Myrmeciinae accepted the classification of Archibald and colleagues without mentioning the comments of Baroni Urbani. The following
cladograms A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
generated by Archibald and colleagues show two possible phylogenetic positions of ''Avitomyrmex'' among some ants of the subfamily Myrmeciinae; the cladogram on the right included three additional extinct genera compared to that on the left. It is suggested that ''Avitomyrmex'' may be closely related to other extinct Myrmeciinae ants such as ''Macabeemyrma'' and ''Ypresiomyrma'', as well as the extant ''Nothomyrmecia macrops''.


Description

There are several characters which separate ''Avitomyrmex'' from other ant genera. The most notable feature is the distinctly slender nature of the queens and workers morphology. This is shown clearly in the shape of the petiole connecting the
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
and the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
. While similar to the modern myrmeciine genus '' Nothomyrmecia'' of Southern Australia, the two genera are distinguishable by the structure of the petiole, with ''Avitomyrmex'' lacking the peduncle seen in ''Nothomyrmecia''. The eyes are large and well developed, the mandibles are subtriangular but poorly preserved, and a sting is present on examined ''A. systenus'' fossils. As for ''A. elongatus'' and ''A. mastax'', it is unknown if the two ants have a sting, due to either poor preservation or the sting has not been preserved at all.


''A. elongatus''

''A. elongatus'' was described from a single
side Side or Sides may refer to: Geometry * Edge (geometry) of a polygon (two-dimensional shape) * Face (geometry) of a polyhedron (three-dimensional shape) Places * Side (Ainis), a town of Ainis, ancient Thessaly, Greece * Side (Caria), a town of an ...
of a compression fossil found from the Middle Ypresian
McAbee Fossil Beds The McAbee Fossil Beds is a Heritage Site that protects an Eocene Epoch fossil locality east of Cache Creek, British Columbia, Canada, just north of and visible from Provincial Highway 97 / the Trans-Canada Highway ( Highway 1). The McAbee Fo ...
, Tranquille Formation, near Cache Creek, British Columbia. The incomplete specimen of a queen, numbered 2003.2.8CDM032, is currently preserved in the
paleontology Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
collections housed at the Courtenay and District Museum, Courtenay, British Columbia. Archibald, Cover, and Moreau coined the specific epithet from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
"''elongatus''" meaning "prolonged" in reference to the elongated morphology of the type specimen. The species is discernible from the other two species of ''Avitomyrmex'' by its notably larger size, the preserved portion of the ant being over 20 millimetres (0.8 inches). The forewings are almost as large as the specimen, measuring around 18 millimetres (0.7 inches) while the hindwings are too poorly preserved to be studied. 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 ...
is preserved with a partly disarticulated gaster and is missing her head.


''A. mastax''

The second species described from the McAbee Fossil Beds is ''A. mastax'' which, unlike ''A. elongatus'', is known from two specimens. The holotype and
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 ...
are both included in the
Thompson Rivers University Thompson Rivers University (commonly referred to as TRU) is a Public university, public teaching and research university offering Undergraduate education, undergraduate and Postgraduate education, graduate Academic degree, degrees and Vocational ...
, Kamloops collections as UCCIPRL-18 F-850 and UCCIPRL-18 F-929 respectively. The holotype specimen is a partial queen which is incomplete, with one forewing and the head fairly preserved, and the other isolated body portions indistinct. The paratype is a mostly complete queen missing parts of her gaster, legs and hind wings. Overall the species is estimated to have been 15 millimetres (0.6 inches) long and has a forewing length of 13 millimetres (0.5 inches). ''A. mastax'' is distinguishable from the other species in ''Avitomyrmex'' by its smaller mandible size, being less than half the length of the head with eight teeth, and additionally the shape of the head capsule. The specimen also has large compound eyes. The specific epithet ''mastax'' is from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
"''mastax''" meaning "jaw" or "mandible", a reference to the small size of the mandibles compared to the other species of ''Avitomyrmex''.


''A. systenus''

Of the three described species of ''Avitomyrmex'' found at the McAbee Fossil Beds, only ''A. systenus'' is known from worker caste specimens. The holotype is currently deposited in the Courtenay and District Museum paleontology collections as 2003.2.11 CDM 035 while the paratype, UCCIPR L-18 F-989, and an additional hypotype worker, UCCIPR L-18 F-825, which is tentatively assigned to the species are both deposited in Thompson Rivers University collections. Based on the mostly complete workers, mature specimens are estimated to have been 15 millimetres (0.6 inches). Due to the size of adult workers they cannot be attributed to ''A. elongatus'' while the overall petiole, head capsule and mandible structure distinguish it from ''A. mastax''. The eyes are large and one-third the length of the head, and the legs are indistinctly preserved but long. The
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum ( dorsal), the prosternum (ventral), and the propleuron (lateral) on e ...
is almost flat, and the gaster is narrow. The shape of the head was the basis for Archibald, Cover and Moreau choosing the specific epithet ''systenus'', which is from the Greek word ''systenos'' meaning "tapering to a point".


Ecology

Archibald and colleagues suggested that the life habits of ''Avitomyrmex'' species may have been similar to that of extant Myrmeciinae ants. These ants may have nested in the soil or in trees, possibly being an
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose nu ...
nesting genus. This may be the case as one ''Myrmecia'' species is known to inhabit trees exclusively. Workers most likely preyed on arthropods, killing them with their sting and fed on nectar; workers would have been found foraging onto trees or low vegetation without leaving any pheromone trails to food sources or recruit nestmates, as they were solitary foragers. ''Avitomyrmex'' ants most likely used their large eyes to locate prey and for navigational purposes.


References


Cited text

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External links

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''Avitomyrmex'' at the AntWiki – Bringing Ants to the World''Avitomyrmex''
at AntCat {{Taxonbar, from=Q4829007 Myrmeciinae Ypresian insects Fossil ant genera Fossil taxa described in 2006 Prehistoric insects of North America