Erigone Atra
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''Erigone atra'' is a species of dwarf spider or money spider, in the family
Linyphiidae Linyphiidae, spiders commonly known as sheet weavers (from the shape of their webs), or money spiders (in the United Kingdom, Ireland, Australia, New Zealand, and in Portugal, from the superstition that if such a spider is seen running on you, ...
. It is commonly found in North America, Europe, parts of Russia (European to
Far East The ''Far East'' was a European term to refer to the geographical regions that includes East and Southeast Asia as well as the Russian Far East to a lesser extent. South Asia is sometimes also included for economic and cultural reasons. The ter ...
), Central Asia, China, Mongolia, Korea, and Japan. This spider is one of the most common ''Erigone'' spiders. ''E. atra'' is an important spider for agriculture, as it preys on pests such as
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s which are commonly found on crops. ''E. atra'' spiders are aeronautical spiders, as they travel via
ballooning Ballooning may refer to: * Hot air ballooning * Balloon (aeronautics) * Ballooning (spider) * Ballooning degeneration, a disease * Memory ballooning See also * Balloon (disambiguation) A balloon is a flexible container for (partially or fully) co ...
. This technique, sometimes referred to as kiting, allows ''E. atra'' spiders to traverse large distances and find new habitats when environmental or human stresses create unfit living environments. ''E. atra'' is difficult to differentiate from other congeneric species because of their similar sizes and coloring.


Description

The female ''E. atra'' ranges from 1.8 to 2.8 mm in length. Their
cephalothorax 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 ...
region is 0.8 to 1.0 mm in length, black or dark brown in color, and is hairless. The
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
of their
cephalothorax 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 ...
have smaller teeth or no teeth compared to males. Their
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
s are larger than their cephalothorax regions ranging from 0.8 to 2.0 mm and are black. Their legs are all attached to the cephalothorax and are uniformly brown. Members of the
Linyphiidae Linyphiidae, spiders commonly known as sheet weavers (from the shape of their webs), or money spiders (in the United Kingdom, Ireland, Australia, New Zealand, and in Portugal, from the superstition that if such a spider is seen running on you, ...
family are often identified by the spinal patterns on the tibia portion of their legs. ''E. atra'' and similar, congeneric species have a 2-2-2-1 tibial spinal pattern. ''E. atra'' are covered in hairs. The
pedipalp Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") and ...
s of the females have no observable claw. Female ''E. atra'' look very similar to congeneric species ''Erigone arctica''. The male ''E. atra'' ranges from 1.9 to 2.5 mm in length. Their cephalothorax region is 1.0 to 1.2 mm in length, hairless, and black or brown. The carapace of their cephalothorax have marginal teeth along the sides. The head region of their carapace is slightly raised compared to females. Their abdomens are smaller and more pointed than female abdomens, ranging from 0.7 to 1.5 mm are black. Their legs are all attached to the cephalothorax and are uniformly brown and have a 2-2-2-1 tibial spinal pattern similar to females. Male pedipalps are longer than females, are covered in hair and tooth-like apophyses. Male ''E. atra'' look very similar to congeneric species ''Erigone dentipalpis''. ''E. atra'' eyes are similar to other members of the
Linyphiidae Linyphiidae, spiders commonly known as sheet weavers (from the shape of their webs), or money spiders (in the United Kingdom, Ireland, Australia, New Zealand, and in Portugal, from the superstition that if such a spider is seen running on you, ...
family. They have two rows of four eyes. The back row of eyes are straight while the front row is slightly curved. The front middle eyes are significantly smaller than the side eyes.


Taxonomy

''Erigone atra'' was first described by arachnologist
John Blackwall John Blackwall (20 January 1790 – 11 May 1881) was an English naturalist with a particular interest in spiders. Life Blackwall was born in Manchester on 20 January 1790. He lived at Hendre House near Llanrwst in north Wales from 1833 until ...
in 1833. They belong to the
Linyphiidae Linyphiidae, spiders commonly known as sheet weavers (from the shape of their webs), or money spiders (in the United Kingdom, Ireland, Australia, New Zealand, and in Portugal, from the superstition that if such a spider is seen running on you, ...
family, which are also known as sheet weaver spiders. The
Erigoninae Erigoninae are the largest subfamily of sheet weavers (Linyphiidae), which is itself the second largest spider family. In the United States they are known as dwarf spiders, while they are called money spiders in England. The exact taxonomic lim ...
subfamily are known as dwarf spiders in the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
and money spiders in the
United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
. ''Erigone'' is one of the many genera in the Erigoninae subfamilies and was discovered by Jean Victor Audouin in 1826. As of October 2020 the genus contains 111 accepted species.


Distribution and habitat


Geographic range

''E. atra'' has a
holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical region ...
distribution, from North America to Japan. The spider is most abundant in Western Europe.


Habitat

''E. atra'' spiders live in a variety of
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s. This is because of their ability to travel long distances via
ballooning Ballooning may refer to: * Hot air ballooning * Balloon (aeronautics) * Ballooning (spider) * Ballooning degeneration, a disease * Memory ballooning See also * Balloon (disambiguation) A balloon is a flexible container for (partially or fully) co ...
. They are primarily found in
grassland 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 natur ...
s,
fallow Fallow is a farming technique in which arable land is left without sowing for one or more vegetative cycles. The goal of fallowing is to allow the land to recover and store organic matter while retaining moisture and disrupting pest life cycles ...
s, and crop fields. ''E. atra's'' colonial habitat is subject to variation based on season and other environmental factors. They prefer grasslands with high yields of vegetation and fewer populations of different species. ''E. atra'' prefer perennial grasslands over crops because perennial habitats are sustained over time. During the winters, ''E. atra'' prefer higher vegetation habitats as they provide more protection from predators and increased insulation from the cold. The increased canopy cover creates a dry shelter with ground vegetation which is required for
overwintering Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal activi ...
. Some exchange between grassland and crop habitat exists during a spider's lifetime. ''E. atra'' can be found in regions where both cows and sheep graze. Combined grazing of cows and sheep create a uniform expanse of short grass which provides an ideal high-density, low-species habitat.


Diet


Prey selection

''E. atra'' eat
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s,
springtail Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called Ento ...
s,
gnats A gnat () is any of many species of tiny flying insects in the dipterid suborder Nematocera, especially those in the families Mycetophilidae, Anisopodidae and Sciaridae. They can be both biting and non-biting. Most often they fly in large num ...
and other crop pests. Their diet consists primarily of species of aphids and isotomidae springtails, as both families of pest are selected for by ''E. atra''. Both insects are found in high density in crop fields, one of the primary habitats of ''E. atra''. Aphids are caught more frequently by the spiders, however, springtails are typically consumed more because they are more easily identifiable by the spiders. ''E. atra'' select prey based on size. Minor preferences for larger springtails where observed in ''E. atra'' females, and females ate a greater percentage of larger prey compared to smaller prey. Female ''E. atra'' have higher predation rates then males. Prey selection is dependent on the habitat the spiders are in. The variability in habitat due to the spider's large range from aeronautical travel leads to variability in prey selection.


Prey capture

''E. atra'' capture prey both by web capture and by hunting. Small webs are sometimes built a few millimeters off of the ground around crops or other feeding places for aphids, springtails, and other pests. These webs are used for prey capture by killing prey entangled in web threads. Webs are also built above bare soil and can be used as a base for actively attacking and killing prey rather than entangling prey. ''E. atra'' have also been observed searching for prey on plants and holding prey within their
chelicerae The chelicerae () are the mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as "jaws", chelicerae may be shaped as either articulated fangs, or similarly ...
without webs nearby. Adult males almost exclusively hunt without a web.


Behavior


Ballooning

Ballooning Ballooning may refer to: * Hot air ballooning * Balloon (aeronautics) * Ballooning (spider) * Ballooning degeneration, a disease * Memory ballooning See also * Balloon (disambiguation) A balloon is a flexible container for (partially or fully) co ...
is the behavioral trait where aeronautical insects shoot web threads into the air and causes them to become airborne. In ''E. atra'', ballooning is a form of aerial dispersal in which the spiders use thin threads of
spider silk Spider silk is a protein fibre spun by spiders. Spiders use their silk to make Spider web, webs or other structures, which function as sticky nets to catch other animals, or as nests or cocoons to protect their offspring, or to wrap up prey. ...
, often called gossamers, to catch electric field currents and
air current In meteorology, air currents are concentrated areas of winds. They are mainly due to differences in atmospheric pressure or temperature. They are divided into horizontal and vertical currents; both are present at mesoscale while horizontal ones d ...
s. ''E. atra'' first undergo a behavior called tip-toeing to become airborne. A spider will climb up to an elevated position, to avoid threads coming into contact with the ground or other objects, and raise their abdomens in the air. This position maximizes silk released from the
spinneret A spinneret is a silk-spinning organ of a spider or the larva of an insect. Some adult insects also have spinnerets, such as those borne on the forelegs of Embioptera. Spinnerets are usually on the underside of a spider's opisthosoma, and are ...
s and this initial silk release occurs via muscular release. The silk strands are pulled by drag, which both further pulls out released gossamers from the spinnerets while also pulling the spider into the air. Static electricity fields may also contribute to lift along with drag from the air currents. Wind turbulence is a major cue for spiders to begin tip-toe behavior. Food deprivation is another cue for spiders to disperse. Acute feeding stress, feeding history and age all influence how food deprivation influences tip-toe behavior. Increases in temperature are another cue for ''E. atra'' dispersal behavior. In late summer, there is a mass ballooning dispersal triggered by high temperatures. However, the spiders’ propensity of ballooning (long-distance dispersal) was the lowest at 30 degrees celsius. Variation in ''E. atra'' ballooning behavior is due to developmental variations. The habitat chosen by the mother can affect ballooning behavior in offspring because of temperature, food availability, and other environmental circumstances during rearing of young.


Rappelling

In addition to ballooning, there is also a second kind of dispersal, and that is the short-distance dispersal: rappelling. These spiders tend to disperse by utilizing silk as either a sail, which means that ballooning is being used, or rappelling. This means that this technique is being used to bridge the threads. There is a higher risk of mortality and cost associated with ballooning than rappelling, which is why spiders tend to select for the more controlled behavior or rappelling.


Reproduction and lifespan


Lifespan

Two generation of ''E. atra'' are observed per year. The first generation is laid by females which have overwintered as adults. These eggs hatch and mature in early summer and copulation takes place. Later in summer, the second generation of egg sacs arise. The second generation of ''E. atra'' hatch and mature in early fall. Some females are fertilized before winter but store sperm while overwintering and produce egg sacs in the spring. In comparing the two sexes' lifespan, the females live on average 18 days longer than the males.


Reproduction

Female ''E. atra'' create
egg sac Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species dive ...
s, which are sometimes referred to as a cocoon, to hold their eggs. A female will create egg sacs their whole adult life and they will typically die less than 10 days after they have made their last egg sac. Females will lay on average 4.4-11.8 eggs a day at optimal temperature, and clutch size is on average 12-14 eggs. Also, in comparison to temperature groups 15, 20, 25, and 30 degrees Celsius, the groups reared in 20 and 25 degrees Celsius had the highest level of fecundity. Hence, temperature does impact the amount of eggs in the first sac as well as the overall number of eggs produced in the lifetime. ''E. atra'' typically have smaller clutches but produce egg sacs at a rapid rate. As females age, the number of eggs per egg sac decreases. Egg sacs are often produced near the female's web. Increased prey availability corresponds with increased reproductive ability. Depending on prey availability, females will kill more prey than can be consumed because as prey availability increases, so does the amount of prey eaten per egg produced.


Development

''E. atra'' development is dependent on temperature and prey abundance. ''E. atra'' develop very quickly in warm weather. In a laboratory study by De Keer et al. they were observed to reach adulthood in 20 days when they were reared at 20 °C. When spiders where reared at 10 °C development lasted on average 150 days and when reared at 5 °C spiders only
moulted In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...
once. Juvenile mortality increases as temperature decreases. Prey abundance has a major effect on the duration of development. As availability of prey decreased, length of spider development increased and mortality rate of spiderlings increased. Development usually takes place during warm periods in early summer and fall, allowing for rapid development. Rapid development is needed because of population loss due to dispersal, egg sac parasites, and human agricultural practices.


Egg sac parasites

''Gelis festinans'' is a small insect specialized parasitoid that specifically targets ''E. atra'' egg sacs. They are wingless, on average 3 millimeters long, and found in open, grassy habitats. A female's parasitoid will deposit one or several eggs into an ''E. atra'' egg sac wall. A larva will hatch that eats the spider eggs and after on average two week, a single adult parasitoid emerges from the egg sac. ''G. festinans'' are attracted to plant
odor An odor (American English) or odour (English in the Commonwealth of Nations, Commonwealth English; American and British English spelling differences#-our, -or, see spelling differences) is caused by one or more volatilized chemical compounds ...
s from ''E. atra'' habitats, such as wheat or grass odors. ''G. festinans'' habitat preference is dependent on ''E. atra'' availability and population density. Female ''G. festinans'' spend a significantly more time on ''E. atra'' webbing zones rather than non-webbing zones. Once a ''G. festinans'' has encountered webbing they change from a random search pattern to a restricted search pattern for female ''E. atra.'' This change in search behavior could be cued by either chemicals in the webbing, ''E. atra''
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s, or because of the webbing structure. ''G. festinans'' can mark search areas. Marked regions are searched less by other females. Female ''G. festinans'' are also able to distinguish between parasitized and unparasitized egg sacs. No
superparasitism Superparasitism is a form of parasitism in which the host (typically an insect larva such as a caterpillar) is attacked more than once by a single species of parasitoid. Multiparasitism or coinfection, on the other hand, occurs when the host has be ...
has been observed in ''G. festinans.''


Webs

As members of the
Linyphiidae Linyphiidae, spiders commonly known as sheet weavers (from the shape of their webs), or money spiders (in the United Kingdom, Ireland, Australia, New Zealand, and in Portugal, from the superstition that if such a spider is seen running on you, ...
family, ''E. atra'' are sheet-weaving spiders. The webs are small webs typically anchored onto vegetation and are composed of a mesh sheet of silk threads. There is usually no order or pattern in creating webs unlike
orb-weaver spider Orb-weaver spiders are members of the spider family Araneidae. They are the most common group of builders of spiral wheel-shaped webs often found in gardens, fields, and forests. The English word "orb" can mean "circular", hence the English name ...
s. ''E. atra'' usually build their webs over depressions in the ground or over bare soil. In a study done by Alderweireldt et al. analyzing prey selection and capture techniques the median web size of ''E. atra'' was found to be 7.6 cm2 in crop fields. When taking into account individual variation, the 7.6 cm2 is in agreement with the average web size of 4 cm2 found for the
Erigoninae Erigoninae are the largest subfamily of sheet weavers (Linyphiidae), which is itself the second largest spider family. In the United States they are known as dwarf spiders, while they are called money spiders in England. The exact taxonomic lim ...
family. ''E. atra'' webs can be used to entangle prey but are not vital for prey capture or feeding. Webs can also be used as a base from which ''E. atra'' can actively attack and kill prey nearby.


References


External links

* {{Taxonbar, from=Q1058958 Linyphiidae Articles created by Qbugbot Spiders described in 1833