Erigone atra

E. atra has a holarctic distribution, from North America to Japan. The spider is most abundant in Western Europe.[3]

E. atra spiders live in a variety of habitats. This is because of their ability to travel long distances via ballooning. They are primarily found in grasslands, fallows, 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.[9] The increased canopy cover creates a dry shelter with ground vegetation which is required for overwintering.[10] 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.[11]

E. atra eat aphids, springtails, gnats 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.[12]

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 without webs nearby. Adult males almost exclusively hunt without a web.[13]

Spider tip-toe and ballooning behavior

Ballooning 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, often called gossamers, to catch electric field currents and air currents. 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 spinnerets 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.[14]

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.[15] 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.[16]

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.[15]

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.[17] In comparing the two sexes' lifespan, the females live on average 18 days longer than the males.[15]

Erigone atra

Female E. atra create egg sacs, 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 then 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.[15]

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.[17][18] Egg sacs are often produced near the female's web.[19] 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.[17]

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 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.[17]

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 odors 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 pheromones, 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 has been observed in G. festinans.[19]