Linguamyrmex
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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 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 in the formicid subfamily
Haidomyrmecinae Haidomyrmecinae, occasionally called Hell ants, are an extinct subfamily of ants (Formicidae) known from Cretaceous fossils found in ambers of North America, Europe, and Asia, spanning the late Albian to Campanian, around 100 to 79 million years ...
, and is one of only nine genera placed in the subfamily
Haidomyrmecinae Haidomyrmecinae, occasionally called Hell ants, are an extinct subfamily of ants (Formicidae) known from Cretaceous fossils found in ambers of North America, Europe, and Asia, spanning the late Albian to Campanian, around 100 to 79 million years ...
. 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'', ...
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 are ...
.


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 seve ...
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 generat ...
adults which have been preserved as inclusions in transparent chunks of
Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. The ...
. 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 M ...
specimens were recovered from deposits in
Kachin State Kachin State ( my, ကချင်ပြည်နယ်; Jingpho language, Kachin: ), also known by the endonym Kachinland, is the northernmost administrative divisions of Myanmar, state of Myanmar. It is bordered by China to the north and east ...
, in Myanmar. Burmese amber has been radiometrically dated using U- Pb isotopes, yielding an age of approximately 98.79 ± 0.62  million years old, close to the
Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...
 –
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 ...
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 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" which means ant, and the
Latin Latin (, or , ) is a classical language belonging to the Italic languages, 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 ...
"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, bo ...
''vladi'' is a patronym honoring Vlad III, known as
Vlad the Impaler Vlad III, commonly known as Vlad the Impaler ( ro, Vlad Țepeș ) or Vlad Dracula (; ro, Vlad Drăculea ; 1428/311476/77), was Voivode of Wallachia three times between 1448 and his death in 1476/77. He is often considered one of the most im ...
, 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 s ...
. 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 busi ...
's
Count Dracula Count Dracula () is the title character of Bram Stoker's 1897 gothic horror novel ''Dracula''. He is considered to be both the prototypical and the archetypal vampire in subsequent works of fiction. Aspects of the character are believed by some ...
, 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. ...
'', '' 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 ''Haidomyrmex'' is an extinct genus of ants in the formicid subfamily Haidomyrmecinae, and is one of nine genera placed in the subfamily Haidomyrmecinae. The genus contains three described species ''Haidomyrmex cerberus'', ''Haidomyrmex scimit ...
'', '' 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, palae ...
''. ''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 La ...
''. 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 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 ...
placed just to the front of the vertex, and the bulging ovoid
compound eye 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 disti ...
s 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 agriculture, agricultural hand tool for mowing grass or Harvest, harvesting Crop, crops. It is historically used to cut down or reaping, reap edible grain, grains, before the process of threshing. The scythe has been largely ...
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 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 broad and has a coating of fine setae. To the rear of the mesosoma, the propodeum 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
apodeme An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
s 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
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species). It is one of the modern descendan ...
Hymenoptera of Asia Haidomyrmecinae Cretaceous insects of Asia Burmese amber Insects described in 2017