Linguamyrmex Brevicornis
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''Linguamyrmex'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of ant in the formicid subfamily Haidomyrmecinae, and is one of only nine genera placed in the subfamily Haidomyrmecinae. The genus contains three described species, ''Linguamyrmex brevicornis'', ''Linguamyrmex rhinocerus'', and the type species ''Linguamyrmex vladi'' all known from
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'', the ...
fossils found in
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
.


History and classification

''Linguamyrmex'' is known from a total of four 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 several ...
is specimen number "AMNH BuPH-01" from the American Museum of Natural History; the other three specimens described are also in the same collection, but were not placed as members of ''L. vladi''. The described specimens are of
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 ...
adults which have been preserved as inclusions in transparent chunks of Burmese amber. The
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 Ma ...
specimens were recovered from deposits in
Kachin State Kachin State ( my, ကချင်ပြည်နယ်; Kachin: ), also known by the endonym Kachinland, is the northernmost state of Myanmar. It is bordered by China to the north and east (Tibet and Yunnan, specifically and respectively); Sh ...
, in Myanmar. Burmese amber has been
radiometrically dated Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares t ...
using U- Pb isotopes, yielding an age of approximately 98.79 ± 0.62  million years old, close to the Albian –
Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in the s ...
boundary, in the earliest Cenomanian. The fossils were first studied by paleoentomologists Phillip Barden and David Grimaldi with their 2017
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 being published in the journal ''Systematic Entomology''. The genus name ''Linguamyrmex'' was formed as a combination of the Greek "myrmex" which means ant, and 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 ...
"lingua", which means tongue, as a reference to the shape and appearance of the clypeal projection. The
specific epithet In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
''vladi'' is a patronym honoring Vlad III, known as Vlad the Impaler, who was the ruler of
Wallachia Wallachia or Walachia (; ro, Țara Românească, lit=The Romanian Land' or 'The Romanian Country, ; archaic: ', Romanian Cyrillic alphabet: ) is a historical and geographical region of Romania. It is situated north of the Lower Danube and so ...
. It was coined as a reference to both Vlad's preferred execution style and for his inspiration of
Bram Stoker Abraham Stoker (8 November 1847 – 20 April 1912) was an Irish author who is celebrated for his 1897 Gothic horror novel '' Dracula''. During his lifetime, he was better known as the personal assistant of actor Sir Henry Irving and busine ...
's Count Dracula, who drank blood. The three remaining workers studied, BuPH-02, BuPH-03, and BuPH-04 were assigned to ''Linguamyrmex'' due to several factors. Each of the specimens is incomplete, all missing portions of the antennae, making comparison of the scape length to the other segments impossible, and all specimens are missing the terminal segments of the gaster. Based on the worker size and on paddle composition Barden and Grimaldi suggest the three workers may belong to undescribed species. ''Linguamyrmex'' is one of several ant genera described from Burmese amber, the others being ''
Burmomyrma ''Burmomyrma'' is an extinct genus of aculeate hymenopteran, suggested to be an ant. The genus contains a single described species, ''Burmomyrma rossi''. ''Burmomyrma'' is known from a single Middle Cretaceous fossil which was found in Asia. Hi ...
'', '' Camelomecia'', '' Ceratomyrmex'', ''
Gerontoformica ''Gerontoformica'' is an extinct genus of stem-group ants. The genus contains thirteen described species known from Late Cretaceous fossils found in Asia and Europe. The species were described between 2004 and 2016, with a number of the specie ...
'', '' Haidomyrmex'', '' Myanmyrma'', and ''
Zigrasimecia ''Zigrasimecia'' is an extinct genus of ants which existed in the Cretaceous period approximately 98 million years ago. The first specimens were collected from Burmese amber in Kachin State, west of Myitkyina town in Myanmar. In 2013, palaeo ...
''. ''Linguamyrmex'' is one of five genera in Haidomyrmecini, the other four being '' Ceratomyrmex'', ''Haidomyrmex'', '' Haidomyrmodes'' and ''
Haidoterminus ''Haidoterminus'' is an extinct genus of ant in the Formicidae subfamily Haidomyrmecinae, and is one of only nine genera placed in this subfamily. The genus contains a single described species ''Haidoterminus cippus'' and is known from one Late ...
''. The genus is distinguished from them by the well developed gasteral constriction on the abdomen, a feature only seen, to a lesser degree, in ''Haidomyrmodes mammuthus''. Additionally ''Linguamyrmex'' has an enlarged and paddle shaped clypeus, similar to the extremely modified clypeus of ''Ceratomyrmex'', but which is smooth along the stalk instead of sporting setae, and with trigger hairs placed at the base of the setae pad on the paddle instead of at the base of the paddle stalk.


Behavior and ecology

As with all haidomyrmecine members, the head structure of ''Linguamyrmex'' is interpreted to facilitate trap-jaw behavior, as is seen in modern specialized genera of Formicinae, Myrmicinae and Ponerinae. In haidomyrmecines the clypeus is modified to span from near the oral opening up past the mandibles to near the vertex of the head, the only ant group with such a modification. Along with the clypeal structuring a pair of long setae are positioned in the path of the mandibles. In modern trap-jaw ants such setae are used as triggers that initiate the rapid closure of the mandibles, often to capture prey or sometimes in a defensive action. The upper areas of the clypeus are always coated in a brush of stout setae in the region where the mandibles came to rest when triggered to close. Based on the modification of the clypeus, it is probable that ''Linguamyrmex'' would have captured prey from the front, other strategies being made improbable due to the upward motion of the mandibles. In haidomyrmecines with highly modified cleypeal areas, such as ''Linguamyrmex'' and ''Ceratomyrmex'' the horn probably served as a pinning or trapping point for the mandibles with captured prey. This hypothesis for the purpose of the clypeus in ''Linguamyrmex'' is further supported by data obtained from X-ray micro-computed tomography scans of specimen "BuPH-03". Scanning of the specimen shows density variation in the paddle, with the outer edges less dense than the central core area on the paddles underside. This indicates the paddle was strengthened with thickened cuticle or with metals sequestering in the cuticle, the latter being more likely from the distinct contrast shown in the x-ray. The location of the highest density is near the center of the paddle at the point of impact for the mandibles when they closed, likely to help brace the paddle against misfires of the mandibles, or impact through soft bodied prey, such as beetle larvae. Metals such as calcium, iron, manganese, and zinc are used by insects in both mandibles and ovipositors to reduce wear and increase durability.


Description

The head of the ''Linguamyrmex'' holotype measures from the front edge of the clypeus to the rear of the head at the occipital foramen, which is placed high on the back side, near the vertex. There are three ocelli placed just to the front of the vertex, and the bulging ovoid compound eyes are positioned high on the head capsule. The antennae are not fully preserved, with only three full segments, the scape, pedicel, and flagellomere I. The antennae insertion is just in front of and below the eyes, positioned in depressions on each side of the frontal triangle. The frontal triangle extends down from the vertex and merges with the base of the clypeal horn, which is reinforced with clear membrane-like cuticle. The clypeus is enlarged and modified into a paddle shaped horn that has a pad of setae on the underside. The stalk of the horn is smooth, with no setae, and the trigger hairs seen in haidomyrmecines sprout from near the base of the setae pad on the paddle. The pad is composed of long fine setae around the edges changing to shorter thicker setae in the center. The mandibles
scythe A scythe ( ) is an agricultural hand tool for mowing grass or harvesting crops. It is historically used to cut down or reap edible grains, before the process of threshing. The scythe has been largely replaced by horse-drawn and then tractor m ...
shaped in general outline, curving out and upwards from the head towards the clypeal horn. There are expansions near the base of each mandible which angle in towards each other, and which have upper surfaces that are concave and covered with setae pointing towards the mouth. The
mesosoma 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 ...
has a small propleuron that is hidden except where the mesosoma touches the head capsule, while the pronotum is broad and has a coating of fine setae. To the rear of the mesosoma, the
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, not subdivided, and bears a pair of spiracles. It is strongl ...
is gently rounded upper front face that curves to become a shear rear face. Both the propodeal spiracle and metapleural gland are visible on the sides of the mesosoma, with the spiracle elongated into a slit shape, while metapleural gland is a horizontal opening placed rear of a well-developed bulla. The petiole has a gradually sloping front face and rounded upper surface. The rear is broadly attached to gasteral segment I and the distinctly showing helcium is striated. An incomplete fusion line runs along the side of the petiole and gaster, with a thin membranous section present below, while a constriction between segments I and II that rings the full gaster. There is a forward projecting horn on the underside of sternite III that also has distinct striations. The cuticle of the gaster is translucent, which shows that the apodemes of gastral sternites IV and V are expanded and heavily sclerotized.


References


External links

* {{Taxonbar, from=Q41548333 Monotypic fossil ant genera Fossil taxa described in 2017 Hymenoptera of Asia Haidomyrmecinae Cretaceous insects of Asia Burmese amber Insects described in 2017