Liometopum Apiculatum
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''Liometopum apiculatum'' is a
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
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
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 botanical subfamily names with "-oideae", and zo ...
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 wor ...
. ''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 list of regions of the United States, region of the United States that includes Arizona and New Mexico, along with adjacen ...
and
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
to
Quintana Roo Quintana Roo, officially the Free and Sovereign State of Quintana Roo, is one of the 31 states which, along with Mexico City, constitute the 32 administrative divisions of Mexico, federal entities of Mexico. It is divided into municipalities of ...
.


Taxonomy

''Liometopum apiculatum'' was first described by
Mayr Mayr is a German surname. Notable people with the surname include: * Andrea Mayr (born 1979), Austrian female long-distance runner * Ernst Mayr (1904–2005), German American evolutionary biologist * Franz Xaver Mayr (1875–1965), Austrian gastro ...
(1870), who described the workers of this species. Emery (1895) later described the
queens Queens is the largest by area of the Boroughs of New York City, five boroughs of New York City, coextensive with Queens County, in the U.S. state of New York (state), New York. Located near the western end of Long Island, it is bordered by the ...
of this species; Wheeler (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 list of regions of the United States, region of the United States that includes Arizona and New Mexico, along with adjacen ...
and
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
to
Quintana Roo Quintana Roo, officially the Free and Sovereign State of Quintana Roo, is one of the 31 states which, along with Mexico City, constitute the 32 administrative divisions of Mexico, federal entities of Mexico. It is divided into municipalities of ...
. They extend from
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
through
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
,
New Mexico New Mexico is a state in the Southwestern United States, Southwestern region of the United States. It is one of the Mountain States of the southern Rocky Mountains, sharing the Four Corners region with Utah, Colorado, and Arizona. It also ...
, southeastern
Arizona Arizona is a U.S. state, state in the Southwestern United States, Southwestern region of the United States, sharing the Four Corners region of the western United States with Colorado, New Mexico, and Utah. It also borders Nevada to the nort ...
, 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 The pinyon or piñon pine group grows in southwestern North America, especially in New Mexico, Colorado, Arizona, and Utah, with the single-leaf pinyon pine just reaching into southern Idaho. The trees yield edible Pine nut, nuts, which are a sta ...
zones, up to the
ponderosa pine ''Pinus ponderosa'', commonly known as the ponderosa pine, bull pine, blackjack pine, western yellow-pine, or filipinus pine, is a very large pine tree species of variable habitat native to mountainous regions of western North America. It is t ...
and
riparian zone A riparian zone or riparian area is the interface between land and a river or stream. In some regions, the terms riparian woodland, riparian forest, riparian buffer zone, riparian corridor, and riparian strip are used to characterize a ripari ...
s; at lower elevations they inhabit
creosote bush ''Larrea tridentata'', called creosote bush, greasewood, and chaparral is a medicinal herb. In Sonora, it is more commonly called ''hediondilla''; Spanish ''hediondo'' = "smelly". It is a flowering plant in the family Zygophyllaceae. The specific ...
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 such as clover, and other herbs. Grasslands occur ...
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 cottonwood
willow Willows, also called sallows and osiers, of the genus ''Salix'', comprise around 350 species (plus numerous hybrids) of typically deciduous trees and shrubs, found primarily on moist soils in cold and temperate regions. Most species are known ...
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 th ...
''. 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 ''Yucca'' ( , YUCK-uh) is both the scientific name and common name for a genus native to North America from Panama to southern Canada. It contains 50 accepted species. In addition to yucca, they are also known as Adam's needle or Spanish-bayon ...
'' spp. They have also been collected in glass containers and rubber tires and among the roots of various perennial
xerophyte A xerophyte () is a species of plant that has adaptations to survive in an environment with little liquid water. Examples of xerophytes include cacti, pineapple and some gymnosperm plants. The morphology and physiology of xerophytes are adapted to ...
s such as ''
Agave ''Agave'' (; ; ) is a genus of monocots native to the arid regions of the Americas. The genus is primarily known for its succulent and xerophytic species that typically form large Rosette (botany), rosettes of strong, fleshy leaves. Many plan ...
'' spp., ''
Opuntia ''Opuntia'', commonly called the prickly pear cactus, is a genus of flowering plants in the cactus family Cactaceae, many known for their flavorful fruit and showy flowers. Cacti are native to the Americas, and are well adapted to arid clima ...
'' spp., ''
Myrtillocactus geometrizans ''Myrtillocactus geometrizans'' (bilberry cactus, whortleberry cactus, blue myrtle cactus, or blue candle) is a species of cactus in the genus '' Myrtillocactus'', native to central and northern Mexico.Germplasm Resources Information Network''Myr ...
'', ''
Yucca filifera ''Yucca filifera'' is a member of the subfamily Agavaceae, family Asparagaceae, native to central Mexico. History It was discovered in 1840 in northeastern Mexico between Saltillo and Parras () on 19 May 1847 by merchant and explorer Josi ...
'', '' Senecio praecox'', ''
Schinus molle ''Schinus'' is a genus of flowering trees and tall shrubs in the sumac family, Anacardiaceae. Members of the genus are commonly known as pepper trees. The Peruvian pepper tree ('' Schinus molle'') is the source of the spice known as pink pep ...
'' or ''
Prosopis juliflora ''Neltuma juliflora '' (, ''Cuji'' in Venezuela, ''Trupillo'' in Colombia, ''Aippia'' in the Wayuunaiki language and long-thorn kiawe in Hawaii), formerly ''Prosopis juliflora'', is a shrub or small tree in the family Fabaceae, a kind of mesquit ...
''. 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 This is a glossary of terms used in the descriptions of ants. A B D E F G H M N O P Q R S T U W See also ...
(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
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s,
annelid The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to vario ...
s,
mollusk Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000  extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The ...
s, dead
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s, 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 ''Yucca'' ( , YUCK-uh) is both the scientific name and common name for a genus native to North America from Panama to southern Canada. It contains 50 accepted species. In addition to yucca, they are also known as Adam's needle or Spanish-bayon ...
'' 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''), aphids, and other ants (''
Pogonomyrmex barbatus ''Pogonomyrmex barbatus'' is a species of harvester ant from the genus ''Pogonomyrmex''. Its common names include red ant and red harvester ant. These large (5– to 7-mm) ants prefer arid chaparral habitats and are native to the Southwestern Uni ...
'', '' Camponotus sayi'' and '' Solenopsis xyloni''). In some habitats the honeydew produced by
hemipteran Hemiptera (; ) is an order of insects, commonly called true bugs, comprising more than 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 ...
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 citrus ...
'' 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'') a ...
'', 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 California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
. 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 flight Nuptial flight is an important phase in the reproduction of most ant, termite, and some bee species. It is also observed in some fly species, such as '' Rhamphomyia longicauda''. During the flight, virgin queens mate with males and then land ...
s 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 typica ...
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