Philodromus cespitum
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''Philodromus cespitum'' is a species of running crab spider in the family ''
Philodromidae Philodromidae, also known as philodromid crab spiders and running crab spiders, is a family of araneomorph spiders first described by Tord Tamerlan Teodor Thorell in 1870 (then known as subfamily Philodrominae within Thomisidae). It contains over ...
.'' It is found in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
,
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
, and parts of the
Middle East The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
and
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
. ''P. cespitum'' is a foliage-dweller, and is the most abundant species found in European fruit orchards. It acts as a
biological control Biological control or biocontrol is a method of controlling pests, such as insects, mites, weeds, and plant diseases, using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also invo ...
by preying on orchard pests. ''P. cespitum'' is a diurnal ambush hunter and preys on
aphids 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 ...
,
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
, and occasionally competitor spider species. Males court females by tapping on the females’ bodies with their forelegs. They then insert a genital plug into the female during copulation. Unlike in many other spider species, subsequent males can mate with plugged females by removing part of the plug prior to copulation. Males discriminate among females based on virginity and plug size, and can determine these factors using the females’ draglines and plug samples.


Description

Males of this species are approximately 3.5-5.0 mm long, and females are about 5.3 mm long. Body color is variable, mostly in shades of brown or yellow. Spots are also a common body pattern. Males tend to have more spots than females. Legs are usually yellowish brown or grey. Some members of the species also have spots on their legs. Males of the species have a noticeably asymmetric
sperm duct The vas deferens or ductus deferens is part of the male reproductive system of many vertebrates. The ducts transport sperm from the epididymis to the ejaculatory ducts in anticipation of ejaculation. The vas deferens is a partially coiled tube w ...
. Male
pedipalps 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 ...
contain nerve tissue and a sensory organ.


Phylogeny

''Philodromus cespitum'' belongs to the '' Philodromus aureolus'' species complex. ''P. cespitum'' is the only species in this group that is
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 ...
. Since different species in this group are distinguished by copulatory organs that are only fully visible when mature, spiderlings of different species in the group are difficult to tell apart. '' Philodromus lividus'' and '' Philodromus longipalpis'' were originally thought to be subspecies of ''P.'' cespitum. Since no evidence was found to place them in the subspecific category, they were made their own species. Males of ''P. lividus'' and ''P. cespitum'' have differently shaped sperm ducts''.'' Female ''P. longipalpis'' have a larger median plate and atrium of the epigyne than ''P. cespitum.'' ''Philodromus fuscolimbatus'', another species in the '' Philodromus aureolus'' species group, was also originally thought to be a subspecies of ''P. cespitum''.  However, the two species can be differentiated by the shapes of their sperm ducts.
Sympatry In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...
occurs between ''P. cespitum and P. fuscolimbatus'' in the northern
Alps The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, Sw ...
, a region where the two species overlap. ''Philodromus cespitum'' can also coexist with related species '' Philodromus albidus'' and '' Philodromus aureolus'' in the same geographic area. They accomplish this by occupying different
trophic Trophic, from Ancient Greek τροφικός (''trophikos'') "pertaining to food or nourishment", may refer to: * Trophic cascade * Trophic coherence * Trophic egg * Trophic function * Trophic hormone * Trophic level index * Trophic level * Trop ...
and habitat niches. Females of the species ''
Philodromus buchari ''Philodromus buchari'' is a spider species found in Europe. See also * List of Philodromidae species This page lists all described species of the spider family Philodromidae accepted by the World Spider Catalog : A ''Apollophanes'' ''Apo ...
'' very closely resemble females of ''P. cespitum,'' although ''P. buchari'' are slightly larger. Neural arrangement in the pedipalps of male cave spiders '' Hickmana troglodytes'' is almost identical to the neural arrangement in the pedipalps of male ''P. cespitum.''


Habitat and distribution

''Philodromus cespitum'' is a foliage-dweller, and is the dominant species found in Central European fruit orchards. It is more highly concentrated in orchard centers than edges. Larger individuals are found in the center and smaller individuals are found around the edges. This species is also found in cotton fields. ''P. cespitum'' is distributed across North America, Europe, North Africa, Turkey, Russia, Kazakhstan, China, Korea, and Japan. Adult females are most commonly observed in June and July. Spiderlings are present year-round but are most abundant from August to October.


Activity

Males are more active during the day than females, and females are more active at dawn and dusk. Both sexes show locomotor activity indicative of a twenty-four hour
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., Endogeny (biology), endogeno ...
cycle. However, the exact effect of these circadian rhythms on locomotion differs between males and females. Females also possess a secondary twelve hour cycle.


Diet

''Philodromus cespitum'' are diurnal hunters. They capture their prey mainly through
ambush An ambush is a long-established military tactics, military tactic in which a combatant uses an advantage of concealment or the element of surprise to attack unsuspecting enemy combatants from concealed positions, such as among dense underbru ...
. They feed on insects and spiders found in fruit orchards, such as
aphids 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 ...
, the pest ''
Cacopsylla pyri ''Psylla pyri'', commonly known as the pear psylla or pear psyllid, is an insect in the family Psyllidae. Originating in Europe and Asia, it has spread to North America. It is a pest of pear trees, sucking the sap, damaging the foliage, flowers ...
'', and ''
Theridion ''Theridion'' is a genus of tangle-web spiders with a worldwide distribution. Notable species are the Hawaiian happy face spider (''T. grallator''), named for the iconic symbol on its abdomen, and '' T. nigroannulatum'', one of few spi ...
'' spiders. They sometimes engage in
intraguild predation Intraguild predation, or IGP, is the killing and sometimes eating of a potential competitor of a different species. This interaction represents a combination of predation and competition (biology), competition, because both species rely on the sa ...
, in which they feed on competitor species as well as smaller prey. A diet of both competitor spiders and aphids increases the overall fitness of ''P. cespitum.'' However, since the costs are high due to retaliation and risk of injury from the competitor spider, intraguild predation does not often occur. Foraging aggressiveness in ''P. cespitum'' can sometimes lead to overkilling, where the organism kills more prey than it can consume. Aggressiveness in this species is positively correlated with size and capture success. Aggressiveness also significantly increases the probability of intraguild predation.


Reproduction and life cycle

Females lay around 250 eggs and construct five to seven egg cocoons. Females build egg cocoons during
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
. The number of eggs laid is not affected by changes in temperature. The number of larvae that hatch decreases after the third egg cocoon built. The time from oviposition to hatching is 20 days at 15°C and 10 days at 24°C. The developmental time of nymph-stages is about 430 days. Developmental time is longer for females than for males regardless of temperature. Females of this species tend to live longer than males. The lifespan of both sexes in a laboratory setting is shorter when housed together than when individually housed.


Mating


Courtship

A male will initiate courtship by tapping the female’s body with his forelegs. Males who have a proportionally larger
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
length in comparison to the females they are courting perform more taps than mating pairs who have a more similar ratio. The female may first run away and push away the male’s forelegs with her own legs. The male pursues the female until she stops moving and then taps continuously as he climbs over her body. The male then enters the mating position, in which he faces towards the tip of the female’s
opisthosoma The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma (cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects to a ...
. He inserts one of his pedipalps into one of the female’s copulatory organs and transfers sperm and genital plug material via the pedipalp. The female ends the copulation by shaking her body to dislodge the male. The male then attempts to restart the courtship process and mount the female again. The female will either allow him to mount again or will shake him off until he ceases the behavior. Females normally allow the male to engage in two to three rounds of courtship and copulation.


Genital plugs

Male spiders plug female genitalia when transferring sperm during copulation. The plugs are stored in the males’ genital bulb and they are
amorphous In condensed matter physics and materials science, an amorphous solid (or non-crystalline solid, glassy solid) is a solid that lacks the long-range order that is characteristic of a crystal. Etymology The term comes from the Greek ''a'' ("wi ...
. Since they do not have a complicated shape or structure, genital plugs are low cost for the male to produce. Plug size and quality are positively correlated with the number of times that the male taps the female’s body during courtship. Genital plugs are made up of a mixture of compressed
vesicles Vesicle may refer to: ; In cellular biology or chemistry * Vesicle (biology and chemistry), a supramolecular assembly of lipid molecules, like a cell membrane * Synaptic vesicle ; In human embryology * Vesicle (embryology), bulge-like features o ...
and
spermatozoa A spermatozoon (; also spelled spermatozoön; ; ) is a motile sperm cell, or moving form of the haploid cell that is the male gamete. A spermatozoon joins an ovum to form a zygote. (A zygote is a single cell, with a complete set of chromosomes, ...
. They extend halfway into the copulatory duct, with the other half being filled with only spermatozoa. In many species, genital plugging prevents copulation with future males. In ''P. cespitum'', however, males can mate even with fully plugged females by removing some of the plugs prior to mating. Females may determine the amount of plug material that a male deposits by choosing when to end the copulation. The amount of material deposited increases with increasing duration of female haematodocha expansion. When males are mated with females who already possess plugs rather than virgin females, the females tend to be more aggressive. They will often bite the males’ legs during courtship, and sperm transfer occurs less frequently. However, males are usually able to remove some of the plug material and mate with the female. Since the male must expend energy removing the plug and is not able to transfer as much sperm, this benefits the male who originally plugged the female. This may explain why plugs are still produced even though they can be removed by a future male.


Male mate choice

Males discriminate and prefer silk draglines from virgin females over mated females. They also prefer virgin females and mated females with small plugs to mated females with large plugs because they do not need to expend as much energy on removing the plug and will be able to transfer more sperm when copulating.


Enemies

Parasitism by organisms such as the small-headed fly ''Ogcodes fumatus'' can sometimes occur. The observed rate of parasitism in ''P. cespitum'' is 2.5%. ''P. cespitum'' exhibit abnormal web-building behaviors prior to the emergence of the larva parasite. ''Chrysoperla carnea'' and ''Coccinella septempunctata'' are natural enemies of ''P. cespitum''. “Philodromus cespitum.” CABI- Invasive Species Compendium, Nov. 2019, www.cabi.org/isc/datasheet/40100.


Interactions with humans and livestock

''Philodromus cespitum'' acts as a biological control agent by feeding on pests in fruit orchards. Because fruit orchards are one of the main habitats of ''P. cespitum,'' they are affected by the insecticides sprayed against the pests that they consume. ''P. cespitum'' exhibit different prey preferences when in the presence of insecticides. This may be due to an impaired sensory system, taste, or mobility as a result of the insecticides. Insecticides also increase the foraging aggressiveness of ''P. cespitum''. This may be due to reduced prey availability or behavioral changes as a result of altered internal states or gene expression. Insecticides can also be harmful to ''P. cespitum'', with different types causing different mortality rates, ranging from 0-80%. Since these spiders are beneficial to farmers in that they consume fruit orchard pests, insecticides that cause a lower mortality rate can work in conjunction with the spider to eliminate pests. ''Philodromus cespitum'' also acts as an important biological control in Chinese and Iranian cotton fields by preying on cotton pests.


Neonicotinoids

Neonicotinoids Neonicotinoids (sometimes shortened to neonics ) are a class of neuro-active insecticides chemically similar to nicotine, developed by scientists at Shell and Bayer in the 1980s. The neonicotinoid family includes acetamiprid, clothianidin, dinot ...
are typically effective in pest control, yet spiders are less sensitive to them given the altered structure of their
acetylcholine receptors An acetylcholine receptor (abbreviated AChR) is an integral membrane protein that responds to the binding of acetylcholine, a neurotransmitter. Classification Like other transmembrane receptors, acetylcholine receptors are classified according ...
where the neonicotinoids bind selectively.
Imidacloprid Imidacloprid is a systemic insecticide belonging to a class of chemicals called the neonicotinoids which act on the central nervous system of insects. The chemical works by interfering with the transmission of stimuli in the insect nervous system. ...
has quite severe effects with partial lethality after an hour.
Acetamiprid Acetamiprid is an organic compound with the chemical formula C10H11ClN4. It is an odorless neonicotinoid insecticide produced under the trade names Assail, and Chipco by Aventis CropSciences. It is systemic and intended to control sucking insects ...
has sublethal effects which are higher when the chemical is applied on the dorsal side. More males than females either are paralyzed or die with the dorsal application of neonicotinoids. This finding is consistent not only with ''P. cespitum'' but also with other prevalent spider families. Řezáč, Milan, et al. “Contact Application of Neonicotinoids Suppresses the Predation Rate in Different Densities of Prey and Induces Paralysis of Common Farmland Spiders.” Sci Rep, vol. 5, no. 9, ser. 1, Apr. 2019. 1.


References


External links

* {{Taxonbar, from=Q290886 cespitum Articles created by Qbugbot Spiders described in 1802