Formica Exsecta
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''Formica exsecta'' (the narrow-headed ant or excised wood ant) 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 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 from Western Europe to Asia. A rare formicine ant with a deeply excised head, ''F. exsecta'' forms small mounds up to around a foot in height consisting of much finer material than that used by "true"
wood ant The ''Formica rufa'' group is a subgeneric group within the genus ''Formica'', first proposed by William Morton Wheeler. This group contains the mound-building species of ''Formica'' commonly termed "wood ants" or "thatch-mound ants", which build ...
s of the ''F. rufa'' group. An interesting feature of ''F. exsecta'' is that it occurs in two distinct social forms: either a monogyne form where the colony has a single egg-laying queen, or a polygyne form where many egg-laying queens are part of the same colony. ''F. exsecta'' is placed in the ''Coptoformica'' subgenus within the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
and is closely related to '' Formica exsectoides'', an
American American(s) may refer to: * American, something of, from, or related to the United States of America, commonly known as the "United States" or "America" ** Americans, citizens and nationals of the United States of America ** American ancestry, pe ...
species. Both species may form vast colony networks. The largest known polydomous system of ''F. exsecta'' consists of 3,350 nests dispersed over about 22 ha in Transylvania, Romania. In
Great Britain Great Britain is an island in the North Atlantic Ocean off the northwest coast of continental Europe. With an area of , it is the largest of the British Isles, the largest European island and the ninth-largest island in the world. It is ...
, ''F. exsecta'' can be found only in a few scattered
heathland A heath () is a shrubland habitat found mainly on free-draining infertile, acidic soils and characterised by open, low-growing woody vegetation. Moorland is generally related to high-ground heaths with—especially in Great Britain—a cooler ...
locations in
South West England South West England, or the South West of England, is one of nine official regions of England. It consists of the counties of Bristol, Cornwall (including the Isles of Scilly), Dorset, Devon, Gloucestershire, Somerset and Wiltshire. Cities and ...
— principally
Chudleigh Knighton Heath Chudleigh Knighton Heath, near Chudleigh Knighton, Devon is a Site of Special Scientific Interest, so designated because of its rare lowland heath vegetation. It is a habitat for many rare species including the ant, ''Formica exsecta''. This is on ...
and nearby
Bovey Heath Bovey Heath is a 32 hectare (50 acre) area of heathland between Bovey Tracey and Heathfield in south Devon, England. Although primarily heathland habitat, the site has some wet mature oak woodland, and is home to rare species such as ...
, which are both managed by the
Devon Wildlife Trust The Devon Wildlife Trust is a member of The Wildlife Trusts partnership covering the county of Devon, England. It is a registered charity, established in 1962 as the Devon Naturalists Trust, and its aim is to safeguard the future of the county's ...
, and in the central
Scottish Highlands The Highlands ( sco, the Hielands; gd, a’ Ghàidhealtachd , 'the place of the Gaels') is a historical region of Scotland. Culturally, the Highlands and the Lowlands diverged from the Late Middle Ages into the modern period, when Lowland Sco ...
(including
Rannoch Moor Rannoch Moor (, gd, Mòinteach Raineach/Raithneach) is an expanse of around of boggy moorland to the west of Loch Rannoch in Scotland, where it extends from and into westerly Perth and Kinross, northerly Lochaber (in Highland), and the area of ...
). A population centre previously existed in the
New Forest The New Forest is one of the largest remaining tracts of unenclosed pasture land, heathland and forest in Southern England, covering southwest Hampshire and southeast Wiltshire. It was proclaimed a royal forest by William the Conqueror, featu ...
, and such eminent
myrmecologist Myrmecology (; from Greek: wikt:μύρμηξ, μύρμηξ, ''myrmex'', "ant" and λόγος, ''logos'', "study") is a branch of entomology focusing on the science, scientific study of ants. Some early myrmecologists considered ant society as th ...
s as
Horace Donisthorpe Horace St. John Kelly Donisthorpe (17 March 1870 – 22 April 1951) was an eccentric United Kingdom, British myrmecologist and coleopterist, memorable in part for his enthusiastic championing of the renaming of the genus ''Lasius'' after him as ...
recorded this species there and in
Parkhurst Forest Parkhurst Forest is a woodland to the north-west of Newport, Isle of Wight, England. The site is partly a site of special scientific interest. It consists of ancient woodland, relict heathland and plantation woodland. The woodland is freehold ...
on the
Isle of Wight The Isle of Wight ( ) is a county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the largest and second-most populous island of England. Referred to as 'The Island' by residents, the Isle of ...
in the last century, but this seems to have declined considerably over the past few decades, and recent searches in such locations have failed to find any trace of colonies. The narrow-headed ant is currently one of the target species in the Back from the Brink project, which aims to extend its range in England. ''F. exsecta'' has also been found in forests in
Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic country located on ...
,
Finland Finland ( fi, Suomi ; sv, Finland ), officially the Republic of Finland (; ), is a Nordic country in Northern Europe. It shares land borders with Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of B ...
,
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,
Tibet Tibet (; ''Böd''; ) is a region in East Asia, covering much of the Tibetan Plateau and spanning about . It is the traditional homeland of the Tibetan people. Also resident on the plateau are some other ethnic groups such as Monpa people, ...
, and
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
.


Behavior


Sex allocation

''F. exsecta'' is a
eusocial 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 ...
species, displaying a
dominance hierarchy In biology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social animal , social groups interact, creating a ranking system. A dominant higher-ranking i ...
among its individual colony members. Amongst the narrow-headed ant species, the two different types of colony structure are:
monogyny Monogyny is a specialised mating system in which a male can only mate with one female throughout his lifetime but the female may mate with more than one male. In this system the males generally provide no paternal care. In many spider species that ...
, one queen per colony, and
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any o ...
, more than one queen per colony. Most commonly, one of these two different types is prevalent within a population. These two types of colonies differ not only in the queen’s mating system, but also in the organization of types of offspring and its hierarchy system. For a polygynous colony to thrive, it must adjust its sex allocation practices contingent on the abundance of resources. Colonies produce a greater percentage of male offspring when restraint on resource availability exists, as well as when the colony has a larger number of queens. The opposite scenario is also found to be true. More female offspring are produced when an abundance of resources exists, as well as when the colony has a smaller number of queens. On average, a colony’s sex ratio is estimated to be 5.8% female, or one female for every 17.2 male offspring. This heavily male offspring-based sex ratio displays an obvious deviation from Fisher’s theory of 1:1 sex ratio. In colonies in which the male offspring are favored, workers tend to execute most of the female
gynes The gyne (, from Greek γυνή, "woman") is the primary reproductive female caste of social insects (especially ants, wasps, and bees of order Hymenoptera, as well as termites). Gynes are those destined to become queens, whereas female workers a ...
. In contrast, colonies where an excess of female gynes is produced, which is more than necessary for the simple act of queen replacement, they are all accepted into the colony to eliminate the possibility of
parasitism Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
by unrelated queens from neighboring populations.


Inbreeding

In monogynous colonies, a significant amount of inbreeding is found. Inbreeding coefficients were found positive for the workers of these colonies. No inbreeding was found between mother queens. Procreation between related individuals of the colony can be further explained by the queen-male relatedness coefficient of 0.23, found by experiments from Liselotte Sundström. Male offspring that were reared in an inbred colony tended to be smaller in mass. Gynes reared in inbred colony display no difference in mass compared to those in noninbred colonies. This result reflects a trade-off between the quantity of offspring and their reproductive potential. Gynes’ reproductive success is more dependent on their mass than that of a male.


Fitness and homozygosity

In single-queen colonies, the level of queen
homozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
is negatively associated with colony age. Reduced colony survival appears to be due to reduced queen lifespan resulting from queen homozygosity. Worker homozygosity appeared to affect reproductive allocation, with higher homozygosity being associated with less resources being allocated to the sexual brood and more to worker production.


Haplodiploidy

''F. exsecta'', much like other insects in the order
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
, have a
haplodiploid Haplodiploidy is a sex-determination system in which males develop from unfertilized eggs and are haploid, and females develop from fertilized eggs and are diploid. Haplodiploidy is sometimes called arrhenotoky. Haplodiploidy determines the se ...
sex determination system. An unusual 0.75 relatedness coefficient between full haplodiploid sisters is one of the main contributors to the frequency of evolution of eusocial species. The queen’s eggs that are fertilized grow into
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
daughters, which contain two pairs of chromosomes, whereas unfertilized eggs produce
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
males, which only contain the queen's chromosomes. The voluntary fertilization of eggs is done by the egg-laying mother. Therefore, ideally, the queen’s best reproductive interest is to lay a larger quantity of eggs or increase the number of eggs that produce individuals that can reproduce themselves. However, female-rich colonies emphasized the production of workers rather than gynes.


References


External links

* {{DEFAULTSORT:Formica Exsecta exsecta Hymenoptera of Asia Hymenoptera of Europe Insects described in 1846 Taxa named by William Nylander (botanist)