Pholcus phalangioides, commonly known as the cosmopolitan cellar spider, long-bodied cellar spider, or one of various types called a daddy long-legs spider, is a spider of the family Pholcidae. It was first described in 1775 by the Swiss entomologist Johann Kaspar Füssli (his surname is also spelled Fuesslin) as Aranea phalangoides. Its common name of "daddy long-legs" should not be confused with a different arachnid group with the same common name, the harvestman (Opiliones), or the crane flies of the superfamily Tipuloidea.

Females have a body length of about 8 mm while males tend to be slightly smaller. The length of the spider's legs are on average 5 or 6 times the length of its body. Pholcus phalangioides has a habit of living on the ceilings of rooms, caves, garages or cellars.

This spider species is considered beneficial in parts of the world because it preys on other spiders, including species considered dangerous such as redback spiders. Pholcus phalangioides is known to be harmless to humans and a potential for the medicinal use of their silk has been reported.

Taxonomy and phylogeny

Pholcus phalangioides was first described in 1775 by the Swiss entomologist Johann Kaspar Füssli. A member of the genus Pholcus in the family Pholcidae, P. phalangioides shares ancestry with roughly 1,340 similar cellar-spiders. All of these spiders are known for their characteristic long legs, which can range from 5 to 6 times the size of their bodies. This is not to be confused with organisms with similar physical appearances, such as the crane fly – an insect – and harvestmen of the arachnid order Opiliones.

Genetic population structure The population sizes of P. phalangioides are influenced greatly by the presence of human-made buildings since these spiders prefer warmer habitats indoors. The large number of buildings in the world has favoured P. phalangioides, though populations tend to be relatively small, widely dispersed, and greatly isolated from one another. This small size combined with low mobility of populations results in an increased importance placed on the role of genetic drift, more specifically the founder effect, on population structure. Although some gene flow does exist between populations, its importance has been insignificant when compared to that of geographical isolation-driven genetic drift. As a result, most P. phalangioides individuals of the same population that live in the same geographical region will have a very low degree of genetic variation (intrapopulation differentiation). On the other hand, this genetic drift results in significant interpopulation differentiation.

Description

Pholcus phalangioides are sexually dimorphic, where females are slightly larger than the males of the species. The body length of this species varies between males and females. Males tend to be around in length with the average male being around . The average female ranges from in length. As indicated by their common name, "daddy long-legs" (in North America), these spiders boast eight very long and thin legs which are covered in thin, grey bristles. On average, their legs are roughly 5 to 6 times as long as the spider's body. The average length of an adult female's legs is roughly .

The bodies of P. phalangioides, as with all spiders, can be divided into two parts: the prosoma and the opisthosoma. The prosoma is commonly known as the cephalothorax, and the opisthosoma is commonly known as the abdomen. Most of the prosoma is occupied by the brain. The opisthosoma is considered the posterior part of the body which contains most of the spider's internal organs including the heart, respiratory system, mid-gland, reproductive system, genital tract and silk glands. The translucent bodies of P. phalangioides tend to be a grey-pale brown color with a dark spot on the back of the prosoma and some dark, blurred spots on the dorsal side of the opisthosoma.

Although some other members of the family Pholcidae have six eyes, Pholcus phalangioides is an eight-eyed spider. The eyes are arranged such that there is a pair of smaller, dark eyes at the front of the prosoma followed by three parallel rows of pairs of larger eyes.

Similar to other species of spider, a hard exoskeleton coats the bodies of P. phalangioides. Depending on the age of the spider, this exoskeleton must be shed at differing intervals; younger spiders tend to molt much more often. During molting, the spider will produce certain enzymes that release the rest of its body from the underlying tissue of its exoskeleton. The spider is then able to escape the exoskeleton. The remnant outer skin or exoskeleton is known as the exuviae.

It takes about one year for these spiders to mature after they are born, and their life span is up to two years or more post-maturity.

Distribution and habitat

Because of its spread with humans worldwide, there has been some uncertainty about its exact original, native range, although it has been recognized as being likely in the subtropical parts of the Afro-Eurasia, thus with a preference for warmer climates, and recent authorities regards it as only native to Asia. As a synanthropic species, Pholcus phalangioides has largely had its modern geographic distribution determined by the spread of humans around the world. Today, these spiders can be found in most of the temperate countries of every continent in the world.

P. phalangioides are not suited for survival in cold environments, which is why they prefer the warmth of the indoors when in these regions, specifically inside human dwellings. These spiders have a particular affinity for dimly lit, dark areas that are quiet and calm. They are commonly found in the corners of buildings and people's homes as well as in attics. Populations of Pholcus phalangioides living outdoors can be found in caves and in between rock crevices.

Diet

P. phalangioides are carnivorous predators that feed on insects, other spiders, and other small invertebrates. Unlike many other spiders, who simply feed on prey that have gotten stuck in their webs, these spiders frequently venture out from their own webs to hunt other spiders resting in their respective webs and feed on them or their eggs. In times of low prey availability, both the males and females of the species will turn to cannibalism to meet their nutritional needs.

General ethology Web patterns In general, the webs of P. phalangioides are loose and horizontal with many irregularities. These webs are often intertwined with webs of other spiders of the same population. They live peacefully unless resources are low at which point the spiders turn to cannibalism.

Communication The extent of the P. phalangioides communication is seen in times of mating. The primary form of communication for these spiders is through the use of touch and chemicals, specifically pheromones.

Predation behaviors

Predators This species is preyed upon by jumping spiders of the Salticidae family. Some of these spiders simply leap into the webs of their prey and attack them. Others employ a certain strategy known as mimicry in order to trick P. phalangioides and capture them.

A jumping spider whose aggressive mimicry behavior towards P. phalangioides has been well studied is the species Portia fimbriata. During mimicry, the jumping spider produces certain specialized vibrations near the edge of the webs of P. phalangioides. These vibrations cause the webs of P. phalangioides to oscillate in such a way that they mimic the oscillations that would be produced when a form of prey gets stuck in the web. The jumping spider will then continue on with these vibrations for very long durations of time, up to three days in some instances. P. phalangioides often assume that this is an indication that they have caught some sort of prey and will move toward the host of the vibrations. At this point, the jumping spider is in an optimal position to leap onto and attack P. phalangioides, thus subduing them in many instances. In addition to employing mimicry, these jumping spiders are also particularly good at preventing P. phalangioides from inducing their whirling defense mechanism, which tends to be an effective way for P. phalangioides spiders to defend themselves from predators.

Defensive behavior The primary defense strategy performed by P. phalangioides in moments of predation is whirling. Whirling, or a gyration of the body, consists of the spider swinging its body around in a circle repeatedly while its legs remain fixed on the web. This whirling strategy is induced as soon as the individual recognizes any sort of movement occurring in its web. The duration of this whirling is related to the specific kind of predator that the spider encounters. Short-duration whirling can be induced simply by a human touching the spider's web or occasionally by spider of a different species. Long-duration whirling, which can last several hours or even days, is performed specifically in response to the presence of the more threatening Salticid, or jumping spiders, much more often than for spiders of other families. The rapid gyrating associated with the whirling disturbs the vision of the Salticid spiders such that they can no longer rely on their acute eyesight to pinpoint the location of P. phalangioides. This disruption results in the safety of the spiders from an otherwise deadly predator.

Mimicry Much like the Salticidae family of spiders, P. phalangioides also use mimicry as a predatory tactic to subdue their prey; however, unlike jumping spiders, P. phalangioides do not rely on vision for predation. This mimicry consists of creating specialized vibrations to trick the prey into thinking that it has caught an insect or another spider. The prey then slowly approaches its supposed catch at which point the P. phalangioides spider raises up on its long legs. The spider patiently waits until the exact moment at which the prey touches one of its legs. Then, the P. phalangioides spider quickly immobilizes its prey by using its legs to wrap it up in layers of silk. Its long legs give it plenty of distance from the prey to avoid being bitten in retaliation. After immobilizing its prey, P. phalangioides can administer their venomous bite to the prey and consume it.

Even forms of prey that do not fully make it onto the web of P. phalangioides are not safe. Often, prey will trip over the edges of the web, thus providing P. phalangioides with an optimal time to attack. P. phalangioides is capable of clinging onto their web with two of their legs while the rest of their body leans out of the web and shoots silk in the direction of the prey to subdue it.

Bite It is a common misconception that P. phalangioides is incapable of biting humans due to an inability of their fangs to penetrate the human epidermis. These spiders can bite humans since their fangs are roughly long, while the thickness of the human epidermis is less, around thick; however, there are hardly any reports on bites.

Venom Although these spiders are capable of hunting and killing some of the most venomous spiders in the world such as the redback spider, they are not dangerous to humans. According to researchers Greta Binford and Pamela Zobel-Thropp, the effects of P. phalangioides venom on humans and other mammals are negligible. In humans, the P. phalangioides bite simply results in a mild stinging sensation that has no long-term health consequences.

Reproduction

Male genitalia

Overall genital system structure