Colobopsis Anderseni
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''Colobopsis anderseni'', synonym ''Camponotus anderseni'', is a species of mangrove
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
found in northern
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
.


Taxonomy

The species was originally placed in the ant genus '' Camponotus'' when described in 2001 as ''Camponotus (Colobopsis) anderseni''. In a revision of the ant subfamily
Formicinae The Formicinae are a subfamily within the Formicidae containing ants of moderate evolutionary development. Formicines retain some primitive features, such as the presence of cocoons around pupae, the presence of ocelli in workers, and little ...
published in 2016 the subgenus ''C. (Colobopsis)'' found to be
phylogenetically In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
distinct from the other subgenera for ''Componotus''. As a result, the subgenus was raised to full genus status as ''
Colobopsis ''Colobopsis'' is a genus of ant in the subfamily Formicinae. This genus was first described in 1861 by Mayr and contains 95 species. The type species is '' Colobopsis truncata''. Description As part of Camponotini, ''Colobopsis'' workers have ...
'', with the species moved out of ''Camponotus'', resulting in the new binomial ''Colobopsis anderseni''.


Description

''Colobopsis anderseni'' is light brown and glossy. Like any insect, its body contains three sections. The first part is called the
prosoma The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
while the middle part is called 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 ...
, and the posterior part is called the
metasoma The metasoma is the posterior part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory system, and circula ...
. The dorsum, or posterior side, of the mesosoma on ''C. anderseni'' is flattened. It also has distinct
metanotum The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites ( exoskeletal plates) are the metanotum (dorsal), the metasternum (ventral), and the metapleuron (lateral) ...
on the posterior of the
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
. It is lacking erect
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...
under its head, and its body is slightly raised on the
tibiae The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
, or fourth joint of each leg. Its eyes elongate looking outward, nearer to the
mandibles In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
than the
vertex Vertex, vertices or vertexes may refer to: Science and technology Mathematics and computer science *Vertex (geometry), a point where two or more curves, lines, or edges meet *Vertex (computer graphics), a data structure that describes the position ...
. It is dimorphic, meaning there are two separate forms based on class. In major workers, the clypeus is depressed and finely punctate. The anterior margin is convex. In minor workers, the clypeus is wide, and the anterior margin is convex and projecting.


Geography

''Colobopsis anderseni'' has only been found in the
mangroves A mangrove is a shrub or tree that grows in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in several ...
of northern Australia. It is found exclusively in twigs of the mangrove tree ''
Sonneratia alba ''Sonneratia alba'' is a mangrove tree in the family Lythraceae. The specific epithet ' is from the Latin meaning "white", referring to the flowers. Description ''S. alba'' grows up to tall with a trunk diameter up to . The cracked to fissured ...
''. These trees can be found in the
Kimberly Kimberly or Kimberley may refer to: Places and historical events Australia * Kimberley (Western Australia) ** Roman Catholic Diocese of Kimberley * Kimberley Warm Springs, Tasmania * Kimberley, Tasmania a small town * County of Kimberley, a c ...
region in the
North Territory The Northern Territory (commonly abbreviated as NT; formally the Northern Territory of Australia) is an Australian territory in the central and central northern regions of Australia. The Northern Territory shares its borders with Western Aust ...
of
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
. According to recent studies, ''C. anderseni'' are the only species of ant known to live in the twigs of these trees, and recent studies showed that approximately 81% of the ant nests in the canopy of the mangrove tree ''Sonneratia alba'' were occupied by ''C. anderseni''. The remaining nests were inhabited by three species of ''
Tapinoma ''Tapinoma'' (from Greek ''ταπείνωμα'' low position) is a genus of ants that belongs to the subfamily Dolichoderinae. The genus currently comprises 74 described species distributed worldwide in tropical and temperate regions ...
'', two species of ''
Crematogaster ''Crematogaster'' is an ecologically diverse genus of ants found worldwide, which are characterised by a distinctive heart-shaped gaster (abdomen), which gives them one of their common names, the Saint Valentine ant. Members of this genus are ...
'', ''
Monomorium floricola The bicolored trailing ant or flower ant (''Monomorium floricola'') is a species of ant of the subfamily Myrmicinae Myrmicinae is a subfamily of ants, with about 140 extant genera; their distribution is cosmopolitan. The pupae lack cocoons ...
'', and ''
Tetraponera punctulata ''Tetraponera'' is a genus of ants in the subfamily Pseudomyrmecinae that are commonly known as slender ants and are characterized by their arboreal nature and slender bodies. The 96 described species of ''Tetraponera'' all of which live in hol ...
''.


Ecology


Flooding in the mangroves

Some select species of ants live in the Australian mangroves because the advantage of living in a mangrove is less competition from other ants. Unfortunately, this is because the mangrove is considered to be highly undesirable by most ant species due to the two daily inundations. However, all mangrove ants have a remarkable ability that allows them to survive these floods which would otherwise exterminate all ant colonies in a single wave. All of these ants use some method to create a sealed pocket of air to protect from any water gathering inside the nest. To avoid drowning during high tide, some ants escape to higher parts of the trees, whereas others hide in air-filled cavities in the mangrove trees or in air pockets in the mangrove mud. These retreats have limited air volumes, and they are therefore subject to extreme changes in the oxygen and carbon dioxide concentrations. This creates a problem when there are limited oxygen levels for large colonies.


Oxygen deprivation in nests

This problem was studied by a group of professors from the Department of Biological Science in the
University of Aarhus Aarhus University ( da, Aarhus Universitet, abbreviated AU) is a public research university with its main campus located in Aarhus, Denmark. It is the second largest and second oldest university in Denmark. The university is part of the Coimbra Gr ...
, Denmark and the School of Science at
Charles Darwin University Charles Darwin University (CDU) is an Australian public university with a main campus in Darwin and eight satellite campuses in some metropolitan and regional areas. It was established in 2003 after the merger of Northern Territory University, ...
in Darwin, Australia. They studied ''Colobopsis anderseni'' in a series of experiments to determine varying carbon dioxide and oxygen levels in nests. During inundation, a soldier sacrifices herself to block the entrance hole with her head which effectively prevents flooding inside the nest. Once this occurs, the area can be very crowded, with the ants and
coccid Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than the ...
s filling up to half the volume, and the conditions in the nests during a flood become hypercapnic and hypoxic. Each nest has only one entrance, and the opening is only about 1.56mm in diameter. The diameter of the galleries is only approximately 2.31 mm. During normal conditions with open nests, the oxygen depletion is substantial in the part of the nest most distant from the opening, and in a 120 mm long nest the oxygen concentration can be as low as 15.7%. During simulated inundation, in which the nest entrances were blocked, the oxygen concentration dropped to less than 0.5% after one hour. After opening the nest entrance, the oxygen concentration increased again, but for a 100 mm long nest it took nearly 20 minutes before the concentration was back to the normal depressed level. The dilemma faced by ''C. anderseni'' is to avoid drowning without suffering anoxia or hypercapnia, and they show a remarkable ability to adapt to the extreme conditions in the mangrove and exploit a niche where the density of other ants is insignificant. By adapting to these hostile situations, mangrove ants have developed the ability to switch to anaerobic respiration. This was proven by the observed oxygen and carbon dioxide levels. Oxygen uptake in the nests eventually stopped, but the carbon dioxide production continued. This indicates high levels of anaerobic respiration. A similar study was conducted by Nielsen and other associates with another Australian mangrove ant, ''
Polyrhachis sokolova '' Polyrhachis sokolova'' is a species of ant from Hong Kong that recently was discovered to be capable of surviving tidal inundations. Researchers at the James Cook University gained attention in 2006 after finding and reporting this behavior ...
''. This mangrove species has been known to use the front four legs as oars and the back two legs as rudders in order to swim back to the nest directly before inundation. At this point, the ants retreat into their nest in the mud, which creates an air pocket and blocks out the incoming tide. The question was again raised about the ant's source of air, and the study on carbon dioxide levels in the nests of ''Polyrhachis sokolova'' was conducted. The experiment concluded with results similar to the study of ''C. anderseni'' nests with evidence that the ants had achieved
anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing re ...
. ''C. anderseni'' and the study involving this species have made significant contributions to the scientific community. This study alongside the study on ''Polyrhachis sokolova'' has supplied the only published work explaining the unique ability for mangrove ants to breathe anaerobically. Before these studies, there was no notable explanation among scientists for many ant species surviving in mangrove inundations.


Queens and workers

In another experiment by Morgens Gissel Nielsen, colonies of ''Colobopsis aderseni'' were observed and behaviors were recorded. New
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 ''C. anderseni'' were regularly observed gnawing small cavities in the green terminal shoots of ''Sonneratia'' trees, and this species occupied all of the examined terminal nest-cavities. It was thereby concluded that all ant galleries in ''Sonneratia'' branches are founded by this species. However the densities of these ants in ''Sonneratia'' trees differs greatly even in close vicinities because of the remote characteristics of these trees. The base of the ''Sonneratia'' trees is usually surrounded by sea water. Therefore, each tree functions as an island. After the queens have landed and lost their wings, they have very limited capacity for dispersal to neighboring trees. One of the notable observations in this study is that queens and workers of ''C. anderseni'' were never found in the same nest cavity. No queens were found in any of the 225 dissected
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materia ...
chambers containing ''C. anderseni''. In an additional 400 nests collected from other areas, queens were only found in the soft green
shoot In botany, a plant shoot consists of any plant stem together with its appendages, leaves and lateral buds, flowering stems, and flower buds. The new growth from seed germination that grows upward is a shoot where leaves will develop. In the spri ...
s at the terminal ends of the branches of ''Sonneratia'' and were never together with workers. In the older nest chambers, workers were always present, regardless of the number of
brood Brood may refer to: Nature * Brood, a collective term for offspring * Brooding, the incubation of bird eggs by their parents * Bee brood, the young of a beehive * Individual broods of North American Periodical Cicadas: ** Brood X, the largest b ...
and
alate Alate (Latin ''ālātus'', from ''āla'' (“wing”)) is an adjective and noun used in entomology and botany to refer to something that has wings or winglike structures. In entomology In entomology, "alate" usually refers to the winged form o ...
s. Furthermore, it was not possible the find an egg-laying queen in the main nest chambers of nests. Many believe that the only plausible explanation for the observed demographic variation in the nests of ''Colobopsis anderseni'' is to assume that the species has some form of
thelytokous parthenogenesis Thelytoky (from the Greek ''thēlys'' "female" and ''tokos'' "birth") is a type of parthenogenesis in which females are produced from unfertilized eggs, as for example in aphids. Thelytokous parthenogenesis is rare among animals and reported in a ...
, which is known from only very few species. This hypothesis is currently being tested with DNA micro satellite markers.


Behavior

''Colobopsis anderseni'' workers of neighboring nests are known to be highly aggressive to each other. This was observed in a careful study from the University of Aarhus by watching neighboring ants in their environment as well as in a laboratory. When ants from different nests were placed together in a
Petri dish A Petri dish (alternatively known as a Petri plate or cell-culture dish) is a shallow transparent lidded dish that biologists use to hold growth medium in which cells can be cultured,R. C. Dubey (2014): ''A Textbook Of Biotechnology For Class- ...
, several began to fight. This indicates that nests represent separate
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
.


Colony size

Because each nest represents a separate colony, there are exceptionally small colonies, which are a clear adaptation to the high degree of environmental variability. A large colony requires either many small nest chambers or a few larger ones. The exchange between the many small chambers will increase the time the workers spend outside the nests and therefore also the degree of
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
. More important for this species is the possibility of being washed away by the tide, waves or heavy rainstorms. Large nest cavities require very long hollowed twigs, which break easily during strong winds, potentially resulting in the ants being washed away. Larger cavities in thicker branches require the ants to excavate hard wood, which did not seem to be the case for this species. A more serious problem is the coccids, which primarily require cavities with a thin wall. Finally, a large cavity with several entrances and a large volume of air is much more difficult to keep waterproof during flooding.


Diet

''Colobopsis anderseni'' are rarely seen
foraging Foraging is searching for wild food resources. It affects an animal's Fitness (biology), fitness because it plays an important role in an animal's ability to survive and reproduce. Optimal foraging theory, Foraging theory is a branch of behaviora ...
which has led scientists to believe that these ants rely on another food source. Due to the high numbers of coccids, their main food source is most likely honeydew. Very little is known about the dispersion of coccids to new chambers. It seems unlikely that the new queens transport coccids during nuptial flights, because no coccids are ever found in chambers less than 6 weeks old.


References

{{Taxonbar, from=Q5028508 Colobopsis Hymenoptera of Australia Insects described in 2001