Liometopum Masonium
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''Liometopum apiculatum'' is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of ant in the
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...
Dolichoderinae Dolichoderinae is a subfamily of ants, which includes species such as the Argentine ant (''Linepithema humile''), the erratic ant, the odorous house ant, and the cone ant. The subfamily presents a great diversity of species throughout the world ...
. ''Liometopum apiculatum'' ants are found in arid and semi-arid regions of
southwestern United States The Southwestern United States, also known as the American Southwest or simply the Southwest, is a geographic and cultural region of the United States that generally includes Arizona, New Mexico, and adjacent portions of California, Colorado, Ne ...
and
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
to Quintana Roo.


Taxonomy

''Liometopum apiculatum'' was first described by Mayr (1870), who described the workers of this species. Emery (1895) later described the
queens Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
of this species;
Wheeler Wheeler may refer to: Places United States * Wheeler, Alabama, an unincorporated community * Wheeler, Arkansas, an unincorporated community * Wheeler, California, an unincorporated community * Wheeler, Illinois, a village * Wheeler, Indiana, a ...
(1905) described the males and Wheeler & Wheeler (1951) described the larvae. Shattuck (1994) considered ''L. apiculatum'' a senior synonym of ''Liometopum masonium''. This species was also confirmed by Del Toro ''et al.'' (2009).


Habitat

''Liometopum apiculatum'' ants are found in arid and semi-arid regions of
southwestern United States The Southwestern United States, also known as the American Southwest or simply the Southwest, is a geographic and cultural region of the United States that generally includes Arizona, New Mexico, and adjacent portions of California, Colorado, Ne ...
and
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
to Quintana Roo. They extend from
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of t ...
through
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
,
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
, southeastern Arizona, and south into Mexico. They are usually found at elevations of , but their prime habitat is oak forests found around . At higher elevations they are found in pinyon pine zones, up to the ponderosa pine and riparian zones; at lower elevations they inhabit creosote bush scrub and
grasslands A grassland is an area where the vegetation is dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes, like clover, and other herbs. Grasslands occur natural ...
in microhabitats of clay, under rocks, boulders, and decaying logs. They have also been found in foothill meadows, deciduous canyon forests, pinyon-cedar woodlands, ponderosa pine-cedar-oak woodlands, and cottonwoodwillow forests. At high elevations, their abundance decreases and they are replaced by '' L. luctuosum''. Altitude may play an important role in the distribution of ''
Liometopum ''Liometopum'' is a genus of ants that belongs to the subfamily Dolichoderinae, found in North America, Europe and Asia. Caterpillars of certain butterfly species have a symbiotic relationship with ''Liometopum'' ants. They produce secretions t ...
''. In regions of Mexico explored by , ''L. luctuosum'' and ''L. apiculatum'' are only found between . Although conditions below looked favorable, they are absent. In the U.S., ''L. apiculatum'' is found from .


Nests

Nests of ''L. apiculatum'' are typically located underground and have a very distinctive structure. They fill hollowed-out chambers with a branched network of carton-like material made out of agglomerated soil and oral secretions until the entire nest resembles Swiss cheese. Within the nest as many as three or four large chambers containing this honeycombed carton-like material can be found. The carton-like material of this species is much finer than that of ''L. uctuosum''. ''L. apiculatum'' tend to nest at higher densities than do ''L. luctuosum''. These ants are usually found nesting in dead logs, under stones, or in decaying stalks of '' Yucca'' spp. They have also been collected in glass containers and rubber tires and among the roots of various perennial xerophytes such as '' Agave'' spp., ''
Opuntia ''Opuntia'', commonly called prickly pear or pear cactus, is a genus of flowering plants in the cactus family Cactaceae. Prickly pears are also known as ''tuna'' (fruit), ''sabra'', ''nopal'' (paddle, plural ''nopales'') from the Nahuatl word f ...
'' spp., '' Myrtillocactus geometrizans'', '' Yucca filifera'', ''
Senecio praecox ''Senecio'' is a genus of flowering plants in the daisy family (Asteraceae) that includes ragworts and groundsels. Variously circumscribed taxonomically, the genus ''Senecio'' is one of the largest genera of flowering plants. Description Morp ...
'', '' Schinus molle'' or '' Prosopis juliflora''. In some habitats, the nests are deep under heavy boulders or large trees. The queen is always well protected and is usually in a remote place about from the largest chamber where the brood is stored. The chambers are connected by various galleries.


Colonies

Colonies of ''L. apiculatum'' are polydomous with segments of nests (or satellite colonies) scattered over the landscape. Colonies are variable in size, ranging from a few hundred to hundreds of thousands of workers. Colonies exploited for their brood by humans contain between 65,000 and 85,000 individuals, while undisturbed colonies may contain as many as 250,000 individuals. Colonies remain useful for repeated brood collection by humans for 4 to 12 years. Colony foundation in ''L. apiculatum'' is by haplometrosis (non-cooperative), that is, a single fertile queen founds each colony. Colony foundation behavior is not uniform among founding queens. The time spent exploring, excavating, and removing excavated materials and waste are usually higher throughout the day, while oviposition, brood care, and inactivity increase at night. Founding queens prefer sites close to bodies of water; however, sites slightly further from water are more conducive to the establishment of a successful colony as persistently high humidity will result in the early death of a colony due to fungal invasion.


Feeding

''Liometopum apiculatum'' are opportunistic carnivores and granivores, and have also been observed foraging on dead insects, larger colonies being more predaceous. ''L. apiculatum'' also feeds on crustaceans,
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
s, mollusks, dead vertebrates, animal droppings, and extrafloral nectar. These ants also obtain nectar or pollen from bear grass and substances from the outside of the ovaries of the flowers of century plants (''
Agave scabra ''Agave scabra'' (synonym ''Manfreda scabra''), also known as rough leaved agave, is a member of the subfamily Agavoideae. Description ''Agave scabra'' has rosettes growing in height and in width. It is similar in form to many other agaves. Th ...
'' and '' Agave chisosensis'') and Spanish dagger ('' Yucca'' spp.). Workers have been attracted to various foods used as baits including apple sauce, sausage, vegetable soup, sugar water, and cookies. ''L. apiculatum'' have also been observed soliciting honeydew from insects including membracids (''
Vanduzea segmentata ''Vanduzea'' is a genus of treehoppers in the family Membracidae. There are about 12 described species in ''Vanduzea''. Species These 12 species belong to the genus ''Vanduzea'': * ''Vanduzea albifrons'' Fowler * '' Vanduzea arquata'' Say (bl ...
''), aphids, and other ants ('' Pogonomyrmex barbatus'', '' Camponotus sayi'' and ''
Solenopsis xyloni The southern fire ant (''Solenopsis xyloni''), also known as the Californian fire ant or cotton ant, is a stinging fire ant native to southern parts of the United States. Its behaviour is similar to the red imported fire ant ''(S. invicta)'', alt ...
''). In some habitats the honeydew produced by
hemipteran Hemiptera (; ) is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from to around ...
s, ''
Cinara ''Cinara'', the conifer aphids or giant conifer aphids, is a genus of aphids in the family Aphididae. They are widespread in the Northern Hemisphere. These aphids specialize on conifers in the pine and cypress families.Jousselin, E., et al ...
'' spp., ''
Dysmicoccus brevipes ''Dysmicoccus brevipes'' is a mealybug. The scientific name was published for the first time by Theodore Dru Alison Cockerell in 1893. The species is found primarily on pineapple and other species in the genus ''Ananas'', but also infests citru ...
'' and ''
Saissetia oleae ''Saissetia oleae'' ( syn. ''Coccus oleae'') is a scale insect in the family Coccidae. It is considered one of the three main phytophagous parasites of the olive tree (''Olea europaea''), together with the olive fruit fly (''Bactrocera oleae'') ...
'', are the main energy sources. In other words, hemipteran exudates make up the bulk of the diet of ''L. apiculatum''. Their role in disrupting biological control has not been determined.


Foraging activity

''Liometopum apiculatum'' forage from March to September. Workers forage almost exclusively on trails as wide as on the soil surface, and when the temperature rises sharply at midday, they cease foraging and seek shelter under stones. The movement of this species is less erratic than '' L. occidentale'' at higher temperatures. An increase in temperature by changes the speed 15-fold, increasing exponentially from per second. There also appears to be little difference in the speed whether ants are moving towards or away from the nest, or between large and small workers during the summer months. However, after prolonged periods (two months or more) of low temperatures, the larger workers are faster than the small workers. Also within a range of , there appears to be little effect of temperature on the number of ants on trails. Maximal activity occurs between 12 p.m. and 12 a.m during the summer months in southern alpine habitats such as Mount Wilson, California. In natural environments, ants of this species forage in areas of , with an average of ; however, they only use between 16% and 30% of this area at any given time. The spatial distribution of the foraging areas for these species seems to be strongly correlated with the location of shrubs and trees infested by hemipterans.


Reproduction

Immature stages of reproductives have been found in ''L. apiculatum'' nests from May to August, whereas the rest of the year the brood is of the worker caste. Males and gynes have been collected outside the nest from June to August and queens (likely founding queens) have been collected in July and August under stones and other landscape features. Nuptial flights of this species occur during the day after a heavy rain during the months of April or May. Before a nuptial flight there is a great agitation of the workers, which leave the nest and run rapidly in a "zig-zag" fashion. The male and female alates leave the nest, but are less active. After a while the workers begin to bite the legs and wings of the alates, forcing them to climb the nearest plant. The workers continue to excite the alates with bites until they begin to beat their wings, and subsequently initiate flight one by one, not as a swarm. Mating takes place in the air, and mated males and females fall to earth together, often still attached. The life span of ''L. apiculatum'' queens is shorter than that of ''L. luctuosum'' queens (exact time difference not specified); however, their productivity (oviposition) is greater. The annual productivity for an established colony (60,000 to 85,000 workers) of ''L. apiculatum'' is about of brood per year.
Oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
by founding queens is large, however, only a small percentage reaches the adult stage of the F1 generation, partly because the smaller, "trophic" eggs are consumed as food. After laying her first batch of eggs, the queen delays laying more until the first eggs have developed into pupae. Once the first workers emerge, the queen discontinues laying trophic eggs, which lowers the total number of eggs laid but increases the proportion of viable eggs. Eggs are laid all year round. Some virgin queens of ''L. apiculatum'' emerge from the nest, remove their wings, and dig a nest without mating. They will lay eggs, care for them, and eat them to survive. However, they only care for more recently laid eggs that have not turned yellow or dried out.


Life cycle

A study by recorded the longevity of each of the reproductive castes and colony foundation of ''L. apiculatum''. In this study, they studied the life cycle of this species under different conditions of humidity, temperature, and substrate. Ant queens were placed either in glass tubes with moist cotton or in jars with soil, and were held at varying temperatures and relative humidity. Observations of the longevity of the various reproductive castes of ''L. apiculatum'' are as follows: males lived 15 to 37 days, virgin queens lived 19 to 268 days and fertilized queens lived 17 to 316 days. However, the last estimate is incredibly short for a queen, so either these ants are highly polygynous or they live much longer in the wild.


References


Bibliography

* * * * Emery, C. (1895) Beiträge zur kenntniss der Nordamerikanischen Ameisenfauna. ''Zoologische Jahrbücher Abteilung für Systematik Ökologie und Geographie der Tiere'' 8:257-360. * * Mayr, G. (1870) Neue Formiciden. ''Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien'' 20: 960–961. *Shattuck, S.O. (1994) Taxonomic catalog of the ant subfamilies Aneuretinae and Dolichoderinae (Hymenoptera: Formicidae). ''University of California Publications in Entomology'' 112:1–241. * *Wheeler, G. C., & J. Wheeler (1951) The ant larvae of the subfamily Dolichoderinae (Hymenoptera: Formicidae). ''Proceedings of the Entomological Society of Washington'' 53:169–210. *


External links

* {{Taxonbar, from=Q3913022 Dolichoderinae Hymenoptera of North America Insects described in 1870 Edible insects