Linguamyrmex
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''Linguamyrmex'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
in the formicid subfamily
Haidomyrmecinae Haidomyrmecinae, occasionally called hell ants, are an extinct subfamily of ants (Formicidae) known from Cretaceous fossils primarily found in amber from North America, South America, Europe, and Asia, spanning the late Aptian to Campanian, aroun ...
, 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 primarily found in amber from North America, South America, Europe, and Asia, spanning the late Aptian to Campanian, aroun ...
. 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 more recent 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 ''cre ...
fossils found in
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
.


History and classification

''Linguamyrmex'' is known from a total of four adult fossils. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
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 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. Th ...
. The
amber Amber is fossilized tree resin. Examples of it have been appreciated for its color and natural beauty since the Neolithic times, and worked as a gemstone since antiquity."Amber" (2004). In Maxine N. Lurie and Marc Mappen (eds.) ''Encyclopedia ...
specimens were recovered from deposits in
Kachin State Kachin State (; Jingpho language, Kachin: ) is the northernmost administrative divisions of Myanmar, state of Myanmar. It is bordered by China to the north and east (Tibet Autonomous Region, Tibet and Yunnan, respectively), Shan State to the sou ...
, 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 (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
 –
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
boundary, in the earliest Cenomanian. The fossils were first studied by paleoentomologists Phillip Barden and David Grimaldi with their 2017 type description 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: 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 of all kno ...
"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 spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
"lingua", which means tongue, as a reference to the shape and appearance of the clypeal projection. The
specific epithet In Taxonomy (biology), taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin gramm ...
''vladi'' is a patronym honoring Vlad III, known as
Vlad the Impaler Vlad III, commonly known as Vlad the Impaler ( ) or Vlad Dracula (; ; 1428/31 – 1476/77), was Voivode of Wallachia three times between 1448 and his death in 1476/77. He is often considered one of the most important rulers in Wallachian hi ...
, who was the ruler of
Wallachia Wallachia or Walachia (; ; : , : ) is a historical and geographical region of modern-day Romania. It is situated north of the Lower Danube and south of the Southern Carpathians. Wallachia was traditionally divided into two sections, Munteni ...
. 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), better known by his pen name Bram Stoker, was an Irish novelist who wrote the 1897 Gothic horror novel ''Dracula''. The book is widely considered a milestone in Vampire fiction, and one of t ...
's
Count Dracula Count Dracula () is the title character of Bram Stoker's 1897 gothic horror novel ''Dracula''. He is considered the prototypical and archetypal vampire in subsequent works of fiction. Aspects of the character are believed by some to have been i ...
, 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'', '' Camelomecia'', '' Ceratomyrmex'', '' Gerontoformica'', ''
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 scimita ...
'', '' Myanmyrma'', and ''
Zigrasimecia ''Zigrasimecia'' is an Extinction, 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 20 ...
''. ''Linguamyrmex'' is one of five genera in Haidomyrmecini, the other four being '' Ceratomyrmex'', ''Haidomyrmex'', '' Haidomyrmodes'' and '' Haidoterminus''. 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 or ocellus (sometimes called a pigment pit) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish the ...
placed just to the front of the vertex, and the bulging ovoid
compound eye A compound eye is a Eye, visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidium, ommatidia, which are tiny independent photoreception units that consist of a cornea, lens (anatomy), lens, and p ...
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 In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
scythe A scythe (, rhyming with ''writhe'') is an agriculture, agricultural hand-tool for mowing grass or Harvest, harvesting Crop, crops. It was historically used to cut down or reaping, reap edible grain, grains before they underwent the process of ...
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. Wasps, bees and a ...
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 Propodeum labelled within the alitrunk of an ant worker The propodeum is a term that can refer to unrelated structures in insects or in mollusks. Insects The first abdominal segment in Apocrita Hymenoptera (wasps, bees and ants) is known a ...
is gently rounded upper front face that curves to become a shear rear face. Both the propodeal spiracle and
metapleural gland Metapleural glands (also called metasternal or metathoracic glands) are secretory glands that were considered unique to ants and basal in the evolutionary history of ants. They are responsible for the production of an antibiotic fluid that then col ...
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 () . is a skeleton that is on the exterior of an animal in the form of hardened integument, which both supports the body's shape and protects the internal organs, in contrast to an internal endoskeleton (e.g. that of a human) wh ...
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 or languages). It is one of the mo ...
Hymenoptera of Asia Haidomyrmecinae Cretaceous insects of Asia Burmese amber Insects described in 2017