Portia labiata: Difference between revisions

Portia labiata
Scientific classification
Kingdom:	Animalia
Subkingdom:	Eumetazoa
Phylum:	Arthropoda
Subphylum:	Chelicerata
Class:	Arachnida
Order:	Araneae
Infraorder:	Araneomorphae
Family:	Salticidae
Subfamily:	Spartaeinae
Genus:	Portia
Species:	P. labiata
Binomial name
	Portia labiata;

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Portia labiata is a jumping spider (family Salticidae) found in Sri Lanka, India, Burma (Myanmar), Malaysia, Singapore, Java, Sumatra and the Philippines. Adult females have bodies 7 to 10 millimetres long,^[7]^: 433 while those of adult males are 5 to 7.5 millimetres long.^[8]^: 103-105 The carapaces of females and males are orange-brown, slightly lighter around the eyes, but have slightly different markings in the two sexes. The abdomens of females are mottled brown and black, while those of males are brown with lighter markings, and the abdomens of the two sexes have different patterns of hairs. Females' chelicerae, which house the fangs, are dark orange-brown, and those of males are orange-brown. The legs of both sexes are dark brown, with light markings in the femora. All species of the genus Portia have elastic abdomens, so that those of both sexes become almost spherical when well fed, and females can stretch as much when producing but not yet laying eggs.

The vision of a jumping spiders' main eyes is more acute than a cat's during the day and 10 times more acute than a dragonfly's,^[9] so that in practice they can see up to about 75 centimetres.^[10]^: 53 Like all jumping spiders, P. labiata can take in only a small visual field at one time, and takes a relatively long time to see objects, probably because a lot of scanning is needed. This makes P. labiata and other jumping spiders vulnerable to much larger predators such as birds, frogs and mantises, which a jumping spider often cannot identify.

The genus Portia have been called "Eight-legged Cats", as their hunting tactics are as versatile and adaptable as a lion's. All members of Portia have instinctive hunting tactics for their most common prey, but often can improvise by trial and error against unfamiliar prey or in unfamiliar situations, and then remember the new approach.^[9] While most jumping spiders prey mainly on insects and by active hunting, females of Portia also build webs to catch prey directly and sometimes join their own webs on to those of web-based spiders. When hunting in another spider's web, a Portia′s slow, choppy movements and the flaps on its legs make it resemble leaf detritus caught in the web. P. labiata and some other Portias use breezes and other disturbances as "smokescreens" in which these predators can approach web spiders more quickly, and revert to a more cautious approach when the disturbance disappears. P. labiata more often pursues small jumping spiders and web spiders than larger prey. A female P. labiata is effective against insects up to twice P. labiata′s size when the insect is stuck in a non-salticid's web, and against insects up to P. labiata′s size when not in webs. Both females and males prefer web spiders as prey, followed by other jumping spiders, and finally insects. In all cases females are more effective predators than males.

Populations from Los Baños and from Sagada, both in the Philippines, have slightly different hunting tactics. In laboratory tests, Los Baños P. labiatas rely more on trial and error than Sagada P. labiatas in finding ways to vibrate the prey's web and thus lure or distract the prey. Around Los Baños the web-building Scytodes pallida, which preys on jumping spiders, is very abundant, and spits a sticky gum on prey and potential threats. A P. labiata from Los Baños instinctively detours round the back of S. pallida while with plucking the web in a way that makes S. pallida believe the threat is in front of it. In areas where S. pallida is absent, the local members of P. labiata do not use this combination of deception and detouring for a stab in the back.

In a test to explore P. labiata′s ability to solve a novel problem, a miniature lagoon was set up, and the spiders had to find the best way to cross it. Specimens from Sagada, in the mountains, almost always repeated the first option they tried, even when that was unsuccessful. When specimens from Los Baños, beside a lake, were unsuccessful the first time, about three quarters switched to another option.

Adult P. labiatas sometimes uses "propulsive displays", in which an individual threatens a rival of the same sex, and unreceptive females also threaten males in this way.^[c] P. labiata females are extremely aggressive to other females, trying to invade and take over each other's webs, which often results in cannibalism. A test showed that they minimise the risk of confrontations by using silk draglines as territory marks. Another test showed that females can recognise the draglines of the most powerful fighters and prefer to move near the draglines of less powerful ones. Females try to kill and eat their mates during or after copulation, while males use tactics to survive copulation, but sometimes females outwit them. Before being mature enough to mate, juvenile females mimic adult females to attract males as prey.

Body structure and appearance

Carapace over cephalothorax

Abdomen

Eyes

Legs

Palps

Chelicerae, housing fangs

Main features of a jumping spider

Spiders are chelicerates, which differ from other arthropods in that the usual body segments are fused into only two tagmata, the cephalothorax and abdomen.^[1] The top of the cephalothorax is covered by a carapace,^[3]^: 572 that of jumping spiders has a distinctive rectangular shape,^[10]^: 51 and that of the genus Portia slopes gently upward almost to the back, then steeply down.^[11]^: 236 Spiders' abdomens bear appendages that have been modified into spinnerets that extrude silk from up to six types of silk glands within their abdomen. The cephalothorax and abdomen are joined by a small, cylindrical pedicel, which allows the abdomen to move while spinning silk.^[3]^: 571-574 While most jumping spiders do not build webs to catch prey, they use silk for other purposes,^[12] including resting, moulting and laying eggs.^[13]^: 495^[14]

Jumping spiders have large forelegs and short, powerful back legs.^[15] Unlike most arthropods, spiders have no extensor muscles in their limbs and instead extend them by increasing their blood pressure. Jumping spiders can leap up to 50 times their own length by powerfully extending the third or fourth pairs of legs,^[3]^: 578 reaching up to 200 millimetres with the forelimbs extended to grasp the prey.^[12] Spiders maintain balance when walking, so that legs 1 and 3 on one side and 2 and 4 on the other side are moving, while the other four legs are on the surface. To run faster, spiders increase their stride frequency.^[16]^: 328

Spider's chelicera, showing the fang almost completely folded away

In spiders and other chelicerates, there is only one pair of appendages before the mouth, the chelicerae,^[1] and in spiders these house fangs that inject poison into the prey.^[3] Behind the mouth is a pair of pedipalps ("palps" for short),^[1] and those of male spiders are quite large and are used for displaying and mating.^[3]

As in most species of the genus, the bodies of female Portia labiatas are 7 to 10 millimetres long^[7]^: 433 and and their carapaces are 2.8 to 3.8 millimetres long.^[8]^: 103-105 Males' bodies are 5 to 7.5 millimetres long, ^[7]^: 433 with carapaces 2.4 to 3.3 millimetres long.^[8]^: 103-105 The carapaces of females are orange-brown, slightly lighter around the eyes, where there are sooty streaks and sometimes a violet to green sheen in certain lights. Females' chelicerae are dark orange-brown and decorated with sparse white hairs, which form bands near the carapaces. The abdomens of females are mottled brown and black, and bear hairs of gold, white and black, and there are tufts consisting of brown hairs tipped with white. The carapaces of males are orange-brown, slightly lighter around the eyes, and have brown-black hairs lying on the surface but with a white wedge-shape stripe from the highest point down to the back, and white bands just above the legs. Males' chelicerae are also orange-brown with brown-black markings. The abdomens of males are brown with lighter markings and with brown-black hairs lying on the surface, and a short band of white hairs. The legs of both sexes are dark brown, with light markings in the femora (the sections of the legs nearest the body). ^[8]^: 103-105 All species of the genus Portia have elastic abdomens, so that those of both sexes can become almost spherical when well fed, and females' can stretch as much when producing eggs.^[13]^: 495

Spiders groom themselves regularly, and more often if wet or dirty. They moisten their fangs, draw the legs one at a time through the fangs, and "comb" the legs with the fangs and palps. The first and fourth pairs of legs are then used to groom other parts of the body, and the only place they appear not to reach is the dorsal surface of the carapace.^[10]^: 53

Movement

When not hunting for prey or a mate, Portia species adopt a special posture, called the "cryptic rest posture", pulling their legs in close to the body and their palps back beside the chelicerae ("jaws"), which obscures the outlines of these appendages. When walking, all Portia species have a slow, "choppy" gait that preserves their concealment: pausing often and at irregular intervals; waving their legs continuously and their palps jerkily up and down; and moving each appendage out of time with the others^[17]^[18]^: 6 and continuously varying the speed and timing.^[19]^: 418 Portia′s walk is unlike that of any other spider, and this gait and the spider's fringes gives the appearance of light flickering through the forest canopy and reflecting from piece of detritus.^[17]^[18]^: 6

In most species of spider, an individual lays a continuous dragline of non-sticky silk as the spider moves, and from time anchors the dragline to a surface with a spot of sticky silk. This allows the spider to return to the surface if the spider is dislodged. A spider about to jump first lays a sticky silk anchor, and then pays out a dragline as it flies.^[3]^: 574

When disturbed, most Portias leap upwards about 100 to 150 millimetres, often from the cryptic rest pose, and often over a wide trajectory. Usually Portia then either freezes or runs about 100 millimetres and then freezes.^[7]^: 434

When isolated on little islands, Portias can enter the water by slowly placing their forelegs in the surface of the water, pushing off with the back legs, and adopting a spreadeagle position. A Portia then swims by moving one leg forward at a time.^[20]^: 285

Senses

Although other spiders can also jump, salticids including Portia labiata are the only spiders with good vision,^[21]^: 521^[22] and their main eyes are more acute in daylight than a cat's and 10 times more acute than a dragonfly's.^[9] Jumping spiders have eight eyes, the two large ones in the center-and-front position (the anterior-median eyes, also called "principal eyes"^[10]^: 51) housed in tubes in the head and providing acute vision. The other six are secondary eyes, positioned along the sides of the carapace and acting mainly as movement detectors.^[9]^[23]^: 16 In most jumping spiders, the middle pair of secondary eyes are very small and have no known function, but those of Portias are relatively large, and function as well as those of the other secondary eyes.^[7]^: 424^[11]^: 232 The main eyes focus accurately on an object at distances from approximately 2 centimetres to infinity,^[10]^: 51 and in practice can see up to about 75 centimetres.^[10]^: 53 Like all jumping spiders, P. labiata can take in only a small visual field at one time,^[24] as the most acute part of a main eye can see all of a circle up to 12 millimeters wide at 20 centimeters away, or up to 18 millimeters wide at 30 centimeters away.^[25]^[a] Jumping spider's main eyes can see from red to ultraviolet.^[14]

Generally the jumping spider sub- family Spartaeinae, which includes the genus Portia, cannot discriminate objects at such long distances as the members of subfamilies Salticinae or Lyssomaninae can. However, members of Portia have vision about as acute as the best of the jumping spiders, for example: the salticine Mogrus neglectus can distinguish prey and conspecifics up to 320 millimetres away (42 times its own body length), while P. fimbriata can distinguish these up to 280 millimetres (47 times its own body length).^[26] The main eyes of a Portia can also identify features of the scenery up to 85 times its own body length, which helps the spider to find detours.^[18]^: 21

However, a Portia takes a relatively long time to see objects, possibly because getting a good image out of such tiny eyes is a complex process and needs a lot of scanning.^[e] This makes a Portia vulnerable to much larger predators such as birds, frogs and mantises, which a Portia often cannot identify because of the other predator's size.^[9]

Spiders, like other arthropods, have sensors, often modified setae (bristles), for smell, taste, touch and vibration protruding through their cuticle ("skin").^[27]^: 532-533 Unlike insects, spiders and other chelicerates do not have antennae.^[1]^[2]

Hunting tactics

Tactics used by most jumping spiders and by most of genus Portia

Almost all jumping spiders are predators,^[b] mostly preying on insects, on other spiders, and on other arthropods.^[22] The most common procedure is sighting the prey, stalking, fastening a silk safety line to the surface, using the two pairs of back legs to jump on the victim, and finally biting the prey.^[28] Most jumping spiders walk through out the day, so that they maximize their chances of a catch.^[29]

Members of the genus Portia have hunting tactics as versatile and adaptable as a lion's.^[9] All members of Portia have instinctive tactics for their most common prey, but can improvise by trial and error against unfamiliar prey or in unfamiliar situations, and then remember the new approach. They can also make detours to find the best attack angle against dangerous prey, even when the best detour takes a Portia out of visual contact with the prey,^[9] and sometimes the planned route leads to abseiling down a silk thread and biting the prey from behind. Such detours may take up to an hour,^[30] and a Portia usually picks the best route even if it needs to walk past an incorrect route.^[19]^: 422 If a Portia makes a mistake while hunting another spider, it may itself be killed.^[30]

While most jumping spiders prey mainly on insects and by active hunting,^[31]^: 340 females of Portia also build webs to catch prey directly.^[9]^[30] These webs are suspended from boughs and rocks, and are funnel-shaped^[14]^[13]^: 513 and about 4,000 cubic centimetres in volume.^[7]^: 429-431 The web is initially built in about 2 hours, and then gradually made stronger.^[11]^: 239 Portias also can join their own webs on to those of web-based spiders.^[9]

A web spider's web is an extension of the web spider's senses, informing the spider of vibrations that signal the arrival of prey and predators. If the intruder is another web spider, these vibrations vary widely depending on the new web spider's species, sex and experience. A Portia can pluck another spider's web with a virtually unlimited range of signals, either to lure the prey out into the open or calming the prey by monotonously repeating the same signal while the Portia walks slowly close enough to bite it.^[31]^: 340-341 Such tactics enable Portias to take web spiders from 10% to 200% of a Portia′s size,^[9] and Portias hunt in all types of webs.^[13]^: 491 In contrast, other cursorial spiders generally have difficulty moving on webs, and web-building spiders find it difficult to move in webs unlike those they build: sticky webs adhere to cursorial spiders and to web-builders of non-sticky webs; builders of cribellate webs have difficulty with non-cribellate webs, and vice versa.^[7]^: 424 Where the web is sparse, a Portia will use "rotary probing", in which it moves a free leg around until it meets a thread.^[7]^: 433-434 When hunting in another spider's web, a Portia′s slow, choppy movements and the flaps on its legs make it resemble leaf detritus caught in the web and blown in a breeze.^[13]^: 514 P. labiata and some other Portias use breezes and other disturbances as "smokescreens" in which these predators can approach web spiders more quickly, and revert to a more cautious approach when the disturbance disappears.^[32]^: 313 A few web spiders run far away when they sense the un-rhythmical gait of a Portia entering the web - a reaction Wilcox and Jackson call "Portia panic".^[19]^: 418

If a large insect is struggling in a web, Portia usually does not usually take the insect, but waits for up to a day until the insect stops struggling, even if the prey is thoroughly stuck.^[7]^: 448 When an insect stuck in a web owned by P. labiata, P. schultzi or any regional variant of P. fimbriata, and next to a web spider's web, the web spider sometimes enters the Portia′s web, and the Portia pursues and catches the web spider.^[7]^{: 440-441, 444}

When catching an insect outside a web, a Portia sometimes lunges and sometimes uses a "pick up",^[7]^: 441 in which it moves its fangs slowly into contact with the prey. In some pick ups, Portia first slowly uses its forelegs to manipulate the prey before biting.^[7]^: 441 P. labiata and P. schultzi also occasionally jump on an insect.^[7]^: 448 However, Portias are not very good at catching moving insects^[13]^: 516 and often ignore them,^[25] while some other salticid genera, especially the quick, agile Brettus and Cyrba, perform well against small insects.^[13]^: 516

When a Portia stalks another jumping spider, the prey generally faces the Portia and then either runs away or displays as it does to another member of its own species.^[7]^: 444^[d]

The webs of spiders on which Portias prey sometimes contain dead insects and other arthropods which are uneaten or partly eaten. P. labiata and some other Portias such as P. fimbriata (in Queensland) and P. schultzi sometimes scavenge these corpses if the corpses are not obviously decayed.^[7]^: 448

A Portia typically takes 3 to 5 minutes to pursuit prey, but some pursuits can take must longer, and in extreme cases close to 10 hours when pursuing a web-based spider.^[7]^: 439

All Portias eat eggs of other spiders, including eggs of their own species and of other cursorial spiders, and can extract eggs from cases ranging from the flimsy ones of Pholcus to the tough papery ones of Philoponella. While only P. fimbriata (in Queensland) captures cursorial spiders in their nests, all Portias steal eggs from empty nests of cursorial spiders.^[7]^: 448

Portias' venom is unusually powerful against spiders.^[13]^: 491 When a Portia stabs a small to medium spider(up to the Portia′s weight^[7]^: 428), including another Portia, the prey usually rans away for about 100 to 200 millimetres, enters convulsions, becomes paralysed after 10 to 30 seconds, and continues convulsing for 10 seconds to 4 minutes. Portia slowly approaches the prey and and takes it.^[7]^: 441-443 Portia usually needs to inflict up to 15 stabbings to completely immobilise a larger spider(1.5 to 2 times to the Portia′s weight^[7]^: 428), and then Portia may wait about 20 to 200 millimetres away for 15 to 30 minutes from seizing the prey.^[7]^: 441-443 Insects are usually not immobilised so quickly but continue to struggle, sometimes for several minutes. If Portia cannot make further contact, all types of prey usually recover, making sluggish movements several minutes after the stabbing but often starting normal movement only after an hour.^[7]^: 441-443

Spiders have a narrow gut that can only cope with liquid food, and have two sets of filters to keep solids out. Some spiders pump digestive enzymes from the midgut into the prey and then suck the liquified tissues of the prey into the gut, eventually leaving behind the empty husk of the prey. Others grind the prey to pulp using the fangs and the bases of the pedipalps, while flooding it with enzymes; in these species the fangs and the bases of the pedipalps form a preoral cavity that holds the food they are processing.^[3]^: 576

Occasionally a Portia is killed or injured while pursuing prey up to twice Portia′s size. P. labiata is killed in 2.1% of pursuits and injured but not killed in 3.9%, P. schultzi is killed in 1.7% and injured but not killed in 5.3%, and P. fimbriata in Queensland is killed in 0.06% of its pursuits and injured but not killed in another 0.06%. A Portia′s especially tough skin often prevents injury, even when its body is caught in the other spider's fangs. When injured, Portia bleeds and may sometimes loses one or more legs. Spiders' palps and legs break off easily when attacked, Portia′s palps and legs break off exceptionally easily, which may be a defence mechanism, and Portias are often seen with missing legs or palps, while other salticids in the same habitat are not seen with missing legs or palps.^[7]^: 440-450

Tactics used by Portia labiata

All performance statistics summarise result of tests in a laboratory, using captive specimens.^[7]^: 429-430

The following table shows the hunting performance of adult females. In addition to P. labiata, the table shows for comparison the hunting performances of P. africana, P. schultzi and three regional variants of P. fimbriata.^[7]^{: 424, 432, 434}

Differences in hunting tactics of females^[7]^{: 434, 437, 439}
Prey	Performance	P. labiata	P. africana	P. schultzi	P. fimbriata (Q)	P. fimbriata (NT)	P. fimbriata (SL)
Salticid	Tendency to pursue prey	63%	77%	58%	87%	50%	94%
Salticid	Efficiency in capturing prey	40%	29%	36%	93%	10%	45%
Web-building spider	Tendency to pursue prey	83%	74%	84%	91%	94%	64%
Web-building spider	Efficiency in capturing prey	79%	65%	72%	92%	81%	83%
Insect	Tendency to pursue prey	35%	48%	52%	27%	30%	43%
Insect	Efficiency in capturing prey	71%	67%	69%	41%	83%	78%

Notes on this table:

"Tendency to pursue prey" is the percentage of tests in which the subject pursues the potential prey, and a pursuit starts when the Portia either approaches the prey or shakes the prey's web.^[7]^: 428-429
"Efficiency in capturing prey" is the percentage of pursuits in which the subject captures the prey.^[7]^: 428-429
P. labiata specimens from Sri Lanka were used in this analysis.^[7]^: 425
"(Q)", "(NT)" and "(SL)" identify P. fimbriatas from Queensland, Northern Territory and Sri Lanka.^[7]^: 425

P. labiata often hangs a capture web from pliant stems and leaves of shrubs and lower branches of trees, rather than from rocks and tree trunks.^[7]^: 432

A female P. labiata more often pursues small jumping spiders and web spiders than larger prey. While it more often catches small jumping spiders than larger ones, it is about equally effective with all sizes of web spiders up to twice P. labiata′s size.^[7]^: 437-439 A female P. labiata is effective against insects up to twice P. labiata′s size when the insect is stuck in a non-salticid's web, and against insects not in webs and up to P. labiata′s size, while P. labiata seldom pursues or catches a larger insect in the open. A female P. labiata very seldom pursues or catches a larger insect in her own web, and is slightly less effective against smaller insects in P. labiata′s web than in other situations.^[7]^: 439 Males are less efficient in all cases.^[7]^: 436

A test in 1997 showed that P. labiata from the Philippines and from Sri Lanka have similar preferences for different types of prey, and that the order of preference is: web spiders; jumping spiders; and insects.^[33]^: 337-339 These preferences apply to both live prey and motionless lures, and to P. labiata specimens without prey for 7 days ("well-fed"^[33]^: 335) and without prey for 14 days ("starved"^[33]^: 335). P. labiata specimens without prey for 21 days ("extra-starved") showed no preference for different types of prey.^[33]^: 339 The test included as prey several species of web spiders and jumping spiders, and the selection of the prey species showed no evidence of affecting the results.^[33]^: 337-339 Insects were represented by the house fly Musca domestica.^[33]^: 335

Unlike the Queensland variant of P. frimbriata, P. labiata has no special tactics when hunting other jumping spiders.^[33]^: 343

P. labiata does not prey on ants,^[34]^: 45-46 but is preyed on by the ants Oecophylla smaragdina and Odontomachus sp. (species uncertain).^[34]^: 47

P. labiata sometimes approaches a translucent nest contain a spider. Usually P. labiata waits faces the prey for up to several hours. Occasionally P. labiata leaps at the prey in the nest, but this is ineffective.^[7]^: 447

Populations from Los Baños and from Sagada, both in the Philippines, have slightly different hunting tactics, and Los Baños has some very dangerous prey spiders. In laboratory tests, Los Baños P. labiatas rely more on trial and error than Sagada P. labiatas in finding ways to vibrate the prey's web and thus lure or distract the prey.^[20]^: 283-284 Around Los Baños the web-building Scytodes pallida, which preys on jumping spiders, is very abundant. All members of the genus Scytodes spit a sticky gum on prey and potential threats, and this can immobilise a Portia long enough for the Scytodes to wrap the Portia in silk and then bite it. Around Los Baños, P. labiata instinctively detours round the back of S. pallida that is not carrying eggs while with plucking the web in a way that makes S. pallida believe the threat is in front of it. P. labiata prefers to stalk a female S. pallida carrying eggs, as then S. pallida is reluctant to drop the eggs in order to spit, and in this case P. labiata sometimes uses a direct attack. In areas where S. pallida is absent, the local members of P. labiata do not use this combination of plucking other spiders' webs to deceive the prey and detouring for a stab in the back.^[35]

A test in 2001 showed that four jumping species take nectar, either by sucking it from the surface of flowers or biting the flowers with their fangs. The spiders fed in cycles of two to four minutes, then groomed, especially their chelicerae, before another cycle. A more formal part of the test showed that 90 juvenile jumping spiders, including P. labiata, generally prefer to suck from blotting soaked with a 30% solution of sugar in water rather than paper soaked with pure water. The authors suggest that, in the wild, nectar may be a frequent, convenient way to get some nutrients, as it would avoid the work, risks and costs (such as making venom). Jumping spiders can benefit from amino acids, lipids, vitamins and minerals normally found in nectar.^[36]

The spiders were divided into four groups:

Rewarded only, if:
- Jumped and then swam
- Swam without jumping
Penalised only, if:
- Jumped and then swam
- Swam without jumping

A test in a deliberately artificial environment explored P. labiata′s ability to solve a novel problem by trial and error. A little island was set up in the middle of a miniature atoll, and the space between with them was filled with water. The gap was too wide for the spiders to jump all the way, and the spiders' options were to leap and then swim or to swim only. The testers encouraged some specimens by using a small scoop to make waves toward the atoll when the spiders chose the option the testers preferred (leap and then swim for some spiders, and swim only for others), and discouraged some specimens by making waves back toward the island when the spiders chose the option the testers did not want - in other words, the testers "rewarded" one group for "successful" behaviour and "penalised" the other group for "unwanted" behaviour.^[20]^: 284-286 Specimens from Sagada almost always repeated the first option they tried, even when that was unsuccessful. When specimens from Los Baños were unsuccessful the first time, about three quarters switched to the other option, irrespective of whether the first attempt was by leaping and then swimming or by swimming only.^[20]^: 287-288

Reproduction and lifecycle

Before courtship, a male Portia spins a small web between boughs or twigs, and he hangs under that and ejaculates on to it.^[7]^: 467 He then soaks the semen into reservoirs on his pedipalps, ^[3]^: 581-583 which are larger than those of females.^[3]^: 572-573

Females of many spider species, including P. labiata,^[37]^: 33^[13]^: 517 emit volatile pheromones into the air, and these generally attract males from a distance.^[37]^: 36 The silk draglines of female jumping spiders also contain pheromones, which stimulate males to court females and may give information about each female's status, for example whether the female is juvenile, subadult or mature.^[37]^: 43 Pheromones may help to find jumping spiders' nests, which are usually hidden under rocks or in rolled leaves, making them difficult to be seen.^[37]^: 36

Portias sometimes use "propulsive displays", with which a member threatens a rival of the same species and sex, and unreceptive females also threaten males in this way.^[31]^: 343 A propulsive display is a series of sudden, quick movements including striking, charging, ramming and leaps.^[7]^: 455

A laboratory test showed how males of P. labiata minimise the risk of meeting each other, by recognising fresh pieces with blotting paper, some containing their own silk draglines and some containing another male's. Males also were attracted by fresh blotting paper containing females' draglines, while females do not response to fresh blotting paper containing males' draglines. This suggested that the males usually search for females, rather than vice versa. Neither sex responded to one week-old blotting paper, irrespective of whether it contained males' or females' draglines. A similar series of tests showed that P. fimbriata from Queensland showed the same patterns of responses between the sexes.^[38]

Among P. labiata and some other Portias, when adults of the same species but opposite sexes recognise each other, they display at 10 to 30 centimetres. Males usually wait for 2 to 15 minutes before starting a display, but sometimes a female starts a display first.^[7]^: 461

A female P. labiata that sees a male may approach slowly or wait. The male then walks with erect and displaying by waving his legs and palps. If the female does not run away, she gives a propulsive display first. If the male stands his ground and she does not ran away or repeat the propulsive display, he approaches and, if she is mature, they copulate.^[7]^: 461-464 If the female is sub-adult (one moult from maturity), a male may cohabit in the female's capture web.^[7]^: 467 Portias usually mate on a web or on a dragline made by the female.^[13]^: 518 P. labiata typically copulates for about 100 seconds,^[7]^: 465 while other genera can take several minutes or even several hours.^[13]^: 518^[7]^: 465

Females of P. labiata and P. schultzi try to kill and eat their mates during or after copulation, by twisting and lunging. The males wait until the females have hunched their legs, making this attack less likely. Males also try to abseil from a silk thread to approach from above, but females may manoeuvre to get the higher position. If the female moves at all, the male leaps and runs away.^[31]^: 343

Before being mature enough to mate, females of P. labiata and also P. shultzi mimic adult females to attract males as prey.^[39]

P. labiata females are extremely aggressive to other females, trying to invade and take over each other's webs, which often results in cannibalism. A laboratory test showed how they minimise the risk of meeting each other, by recognising pieces with blotting paper containing their own silk draglines and pieces contain another P. labiata females' draglines. If obstacles make it impossible to see whether the other is physically present, she avoids blotting paper containing the other's draglines, but moves with no constraint if she can see that the other female is not around. Draglines seem to act as territory marks, much as many mammals identify conspecifics by scent marking.^[40] P. labiata females also avoid rival females of higher fighting ability and spend more time around less powerful fighters. A laboratory test collected samples of the draglines of equal-sized females and then pitted some of them in contests. Other females avoided the draglines of the victors, and spent the majority of their time on draglines of the losers. Similar tests showed that females of P. fimbriata from Australia and P. schultzi from Kenya do not avoid draglines of a powerful fighter.^[41]^: 753

In P. labiata and in some other species, contests between males usually last only 5 to 10 seconds, and only their legs make contact. ^[7]^: 466 Contests between Portia females are violent^[13]^: 518 and embraces in P. labiata typically take 20 to 60 seconds. These occasionally including grappling that sometimes breaks a leg, but more usually the final move is a lunge. Sometimes one knocks the other on her back and, the other may be killed and eaten if she does not right herself quickly and run way. If the loser has a nest, the winner takes over and eats any eggs there.^[7]^: 466-467

Portia females have never been seen eating their own eggs, but in nature females with eggs of their own have been seen eating eggs of other females of the same species. In a test, P. labiata females did not eat their eggs if the testers put them in other female's nests, showing that the test females could identify their own eggs, possibly by chemical means. When the test females and their eggs were restored to their own nests and another females' eggs were also placed in the same nest, the test females ate neither their own eggs nor the "foreign" ones. In nature a female is unlikely to find foreign eggs in her nest, and it might be safest for females to avoid any eggs in their own nests.^[42]

P. labiata usually lays eggs on dead, brown leaves about 20 millimetres long, suspended near the top of its capture web. If there is no dead leaf available, the female will make a small horizontal silk platform in the capture web, and lay eggs on the underside of the platform.^[7]^: 434-435

For moulting, all Portias spin a horizontal web whose diameter is about twice the spider's body length and is suspended only 1 to 4 millimetres below a leaf. The spider lies head down, and often slides down 20 to 30 millimetres during moulting.^[13]^: 496 Portias spin a similar temporary web for resting.^[13]^: 513

Ecology

P. labiata is found in Sri Lanka, India, Burma (Myanmar), Malaysia, Singapore, Java, Sumatra and the Philippines. ^[7]^: 425^[8]^: 103-105^[43]

The populations of P. labiata in Los Baños and in Sagada, both in the Philippines, have different environments: Los Baños is a low-lying tropical rainforest where there are many species of spiders, some of which are especially dangerous to P. labiata; and Sagada is at higher altitude, with pine-forest and fewer species of spiders, none of which are as dangerous to P. labiata. The Los Baños variant has a slightly wider repertoire of tactics.^[35]^[20]^: 283-284

In the Philippines, P. labiata does not prey on ants,^[34]^: 45-46 but is preyed on by the ants Oecophylla smaragdina and Odontomachus sp. (species uncertain),^[34]^: 47 and solitary Odontomachuss have been seen attacked a P. labiata.^[34]^: 50 In a test the ant Diacamma vagans usually killed single-handed a P. labiata.^[34]^: 52

Taxonomy

P. labiata is one of 17 species in the genus Portia as of May 2011.^[4] This species has been named Sinis fimbriatus (Hasselt, 1882; misidentification), Linus labiatus (Thorell, 1887), Linus dentipalpis (Thorell, 1890), Erasinus dentipalpis (Thorell, 1892), Erasinus labiatus (Simon, 1903) and Portia labiata (Wanless, 1978), and the last name has been used since then.^[6]

Portia is in the subfamily Spartaeinae,^[44] which is thought to be primitive.^[13]^: 491 Molecular phylogeny, a technique that compares the DNA of organisms to reconstruct the tree of life, indicates that Portia is a member of the clade Spartaeinae, that Spartaeinae is basal (quite similar to the ancestors of all jumping spiders), that Portia′s closest relative is the genus Spartaeus, and that the next closest are Phaeacius and Holcolaetis.^[5]^: 53

Notes

a: ^ Jackson and Blest (1982) say, "The resolution of the receptor mosaic of Layer I in the central retina was estimated to be a visual angle of 2.4 arc min, corresponding to 0-12 mm at 20 cm in front of the spider, or 0-18 mm at 30 cm."^[25]

b: ^ Several species of cursorial spiders drink nectar as an occasional supplement their diet, and juveniles of some orb-web spiders digest pollen while re-cycling their webs.^[45] One jumping spider (as of 2010), Bagheera kiplingi, is almost totally herbivorous.^[46]

c: ^ "Propulsive displays" are sudden, quick movements including striking, charging, ramming and leaps.^[7]^: 455

d: ^ Except that the Queensland variant of Portia fimbriata generally uses a "cryptic stalking" technique which makes most salticids unaware of this predator.^[7]^: 444

e: ^ The retina is at the end of a tube. The inner end of the tube moves from side to side in one to two cycles per second, and twists 50° in a cycle that takes 10 seconds.^[47]^: 180-181

References

^ ^a ^b ^c ^d ^e Ruppert, E.E., Fox, R.S. and Barnes, R.D. (2004). "Chelicerata: Form". Invertebrate Zoology (7 ed.). Brooks / Cole. p. 555. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Chelicerata: Arachnida". Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 559–564. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Chelicerata: Araneae". Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 571–584. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c Platnick, Norman I. (2011). "World Spider Catalog - Genus Portia". The American Museum of Natural History. Retrieved 4 May 2011. {{cite web}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b Maddison, Wayne P. (2008). "Salticid spider phylogeny revisited, with the discovery of a large Australasian clade (Araneae: Salticidae)" (PDF). Zootaxa. 1893. Magnolia Press: 49–64. ISSN 1175-5334. Retrieved 14 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b Platnick, Norman I. (2011). "World Spider Catalog - Portia labiata". The American Museum of Natural History. Retrieved 21 June 2011. {{cite web}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah ^ai ^aj ^ak ^al ^am ^an ^ao ^ap ^aq ^ar ^as ^at ^au ^av Jackson, Robert R. (1986). "Comparative biology of jumping spiders Portia africana, P. albimana, P. fimbriata, P. labiata and P. schultzi, areanophagic, web-building jumping spiders (Araneae: Salticidae) utilisation of webs, predatory versatility, and intraspecfic interactions". New Zealand Journal of Zoology. 13: 423–489. ISSN 0301-4223. Retrieved 17 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c ^d ^e Wanless, F.R. (1978). "A revision of the spider genus Portia (Araneae: Salticidae)" (PDF). Bulletin of the British Museum (Natural History) Zoology. 34 (3). London: British Museum: 83–124. Retrieved 14 June 2011.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Harland, D.P., and Jackson, R.R. (2000). ""Eight-legged cats" and how they see - a review of recent research on jumping spiders (Araneae: Salticidae)" (PDF). Cimbebasia. 16: 231–240. Retrieved 5 May 2011.{{cite journal}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c ^d ^e ^f Forster, Lyndsay M. (1977). "A qualitative analysis of hunting behaviour in jumping spiders (Araneae: Salticidae)" (PDF). New Zealand Journal of Zoology. 4: 51–62. Retrieved 27 April 2011.
^ ^a ^b ^c Jackson, Robert R. (1986). "Web Building, Predatory Versatility, and the Evolution of the Salticidae". In William A. Shear (ed.). Spiders - webs, behavior, and evolution. Stanford University Press. p. 492. ISBN 9780804712033. Retrieved 22 May 2011.
^ ^a ^b Hillyard, Paul D. (2007). "Jumping spiders". The private life of spiders. New Holland Publishers. pp. 33–35. ISBN 9781845376901. Retrieved 11 April 2011.
^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o Jackson, Robert R. (1986). "Predatory versatility and intraspecies interactions of spartaeine jumping spiders (Araneae, Salticidae): Brettus adonis, B. cingulata, Cyrba algerina, and Phaeacius sp. indet". New Zealand Journal of Zoology. 13: 491–520. ISSN 0301-4223. Retrieved 1 April 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c Richman, David B. (1992). "A review of the ethology of jumping spiders (Araneae, Salticidae)" (PDF). Bulletin of the British Arachnology Society. 9 (2): 33–37. Retrieved 12 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Hogue, Charles Leonard (1993). "Jumping spiders". Latin American insects and entomology. University of California Press. p. 117. ISBN 9780520078499. Retrieved 11 April 2011.
^ Anderson, D.T. (1998). "The Chelicerata". In D.T. Anderson (ed.). Invertebrate Zoology (1 ed.). Oxford University Press Australia. p. 467. ISBN 0915539419. {{cite book}}: Check |isbn= value: checksum (help)
^ ^a ^b Harland, D.P. (2000). "Cues by which Portia fimbriata, an araneophagic jumping spider, distinguishes jumping-spider prey from other prey". The Journal of Experimental Biology. 203. Company of Biologists: 3485–3494. Retrieved 4 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
^ ^a ^b ^c Harland, Duane P. (2004). "Portia Perceptions: The Umwelt of an Aranephagic Jumping Spider". In Frederick R. Prete (ed.). Complex worlds from simpler nervous systems. MIT Press. pp. 5–40. ISBN 9780262661744. Retrieved 12 April 2011. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c Wilcox, R. Stimson (1998). "Cognitive Abilities of Araneophagic Jumping Spiders". In Russell P. Balda, Irene Maxine Pepperberg, Alan C. Kamil (ed.). Animal cognition in nature: the convergence of psychology and biology in laboratory and field. Academic Press. ISBN 9780120770304. Retrieved 23 May 2011. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: multiple names: editors list (link)
^ ^a ^b ^c ^d ^e Jackson, Robert R. (2006). "Geographic Variation in a Spider's Ability to Solve a Confinement Problem by Trial and Error". International Journal of Comparative Psychology. 19: 282–296. Retrieved 8 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Jarman, Elizabeth A.R. (1986). "The biology of Taieria erebus (Araneae, Gnaphosidae), an araneophagic spider from New New Zealand: silk utilisation and predatory versatility". New Zealand Journal of Zoology. 13: 521–540. ISSN 0301-4223. Retrieved 1 April 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b Hoefler, Chad D. (2006). "The Potential of a Jumping Spider, Phidippus clarus, as a Biocontrol Agent" (PDF). J. Econ. Entomol. 99 (2): 432–436. ISSN 0022-0493. Retrieved 25 April 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Hill, David Edwin (2010). "Use of location (relative direction and distance) information by jumping spiders (Araneae, Salticidae, Phidippus) during movement toward prey and other sighted objectives" (PDF). Peckhamia. 83 (1): 1–103. ISSN 1944-8120. Retrieved 12 April 2011. {{cite journal}}: Unknown parameter |month= ignored (help)
^ Piper, Ross (2007). "The Quest for Food: Portia spider". Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals (PDF). Westport, CT 06881: Greenwood Press. pp. 98–100. ISBN 978–0–313–33922–6. Retrieved 31 March 2011. {{cite book}}: Check |isbn= value: invalid character (help)CS1 maint: location (link)
^ ^a ^b ^c Jackson, R.R. (1982). "The distances at which a primitive jumping spider, Portia fimbriata, makes visual discriminations" (PDF). Journal of Experimental Biolology. 97: 441–445. Retrieved 12 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Harland, Duane P. (1999). "Distances at which jumping spiders (Araneae: Salticidae) distinguish between prey and conspecific rivals". Journal of Zoology. 247 (3): 357–364. doi:10.1111/j.1469-7998.1999.tb00998.x. Retrieved 4 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)
^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Arthropoda: Sense organs". Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 532–537. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ Richman, David B. (2008). "Jumping Spiders (Class Arachnidae, Order Araneae, Family Salticidae)". In John L. Capinera (ed.). Encyclopedia of entomology. Springer. pp. 2066–2068. ISBN 9781402062421. Retrieved 11 April 2011.
^ Forster, Lyn M. (1977). "Some factors affecting feeding behaviour in young Trite auricoma spiderlings (Araneae: Salticidae)". New Zealand Journal of Zoology. 4. The Royal Society of New Zealand: 435–442. Retrieved 24 April 2011. {{cite journal}}: Unknown parameter |month= ignored (help)
^ ^a ^b ^c Wilcox, S. and Jackson, R. (2002). "Jumping Spider Tricksters". In Bekoff, M., Allen, C., and Burghardt, G.M. (ed.). The Cognitive Animal: Empirical and Theoretical Perspectives on Animal Cognition (PDF). MIT Press. pp. 27–34. ISBN 0262523221. Retrieved 12 May 2011.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c ^d Jackson, Robert R. (1997). "Jumping spiders mating strategies: sex among cannibals in and out of webs". In Jae C. Choe, Bernard J. Crespi (ed.). The Evolution of Mating Systems in Insects and Arachnids. Cambridge University Press. p. 340-351. ISBN 9780521589765. Retrieved 6 June 2011. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Wilcox, R. Stimson (1996). "Spiderweb smokescreens: spider trickster uses background noise to mask stalking movements" (PDF). Animal Behaviour. 51. The Association for the Study of Animal Behaviour: 313–326. ISSN 0003–3472. Retrieved 23 May 2011. {{cite journal}}: Check |issn= value (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c ^d ^e ^f ^g Li, Daiqin (1997). "Prey preferences of Portia labiata, P. africana, and P. schultzi, araneophagic jumping spiders (Araneae: Salticidae) from the Philippines, Sri Lanka, Kenya, and Uganda". New Zealand Journal of Zoology. 24 (4): 333–349. doi:10.1080/03014223.1997.9518129. Retrieved 21 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c ^d ^e ^f Nelson, Ximena J. (2004). "Predation by ants on jumping spiders (Araneae: Salticidae) in the Philippines" (PDF). New Zealand Journal of Zoology. 31. The Royal Society of New Zealand: 45–56. ISSN 0301–4223. Retrieved 13 June 2011. {{cite journal}}: Check |issn= value (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b Nelson, Ximena J. (2011). "Flexibility in the foraging strategies of spiders". In Marie Elisabeth Herberstein (ed.). Spider Behaviour: Flexibility and Versatility. Cambridge University Press. p. 45-47. ISBN 9780521749275. Retrieved 29 May 2011. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Jackson, Robert R. (2001). "Jumping spiders (Araneae: Salticidae) that feed on nectar" (PDF). Journal of Zoolology. 255. The Zoological Society of London: 25–29. Retrieved 18 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ ^a ^b ^c ^d Gaskett, A.C. (2007). "Spider sex pheromones: emission, reception, structures, and functions" (PDF). Biological Reviews. 82. Cambridge Philosophical Society: 27–48. doi:10.1111/j.1469-185X.2006.00002.x. Retrieved 23 June 2011.
^ Clark, R.J. (1995). "Dragline-mediated sex recognition in two species of jumping spiders (Araneae Salticidae), Portia labiata and P. fimbriata". Ethology Ecology & Evolution. 7: 73–77. Retrieved 13 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Roberts, J. Andrew (2003). "Kinship and food availability influence cannibalism tendency in early-instar wolf spiders (Araneae: Lycosidae)" (PDF). Behaviour, Ecology and Sociobiology. 54. Springer-Verlag: 416–422. doi:10.1007/s00265-003-0646-8. Retrieved 18 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Clark, R.J. (1994). "Self recognition in a jumping spider: Portia labiata females discriminate between their own draglines and those of conspecifics". Ethology Ecology & Evolution. 6: 371–375. Retrieved 13 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Clark, Robert J. (1999). "Draglines and Assessment of Fighting Ability in Cannibalistic Jumping Spiders". Journal of Insect Behavior. 12 (6). Springer: 753–766. doi:10.1023/A:1020900925863. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Clark, Robert J. (1994). "Portia labiata, a cannibalistic jumping spider, discriminates between own and foreign eggsacs". International Journal of Comparative Psychology. 7 (1): 38–43. ISSN 0889-3667. Retrieved 19 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Proszynski, Jerzy. "Global Species Database of Salticidae (Araneae): Portia labiata (Thorell, 1887)". Global Species Database of Salticidae (Araneae). Warsaw: Museum and Institute of Zoology. Retrieved 21 June 2011.
^ Barrion, A.T. (1995). Riceland spiders of South and Southeast Asia. International Rice Research Institute / C.A.B. International. p. 45. ISBN 9780851989679. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Chen, Xiaoqiong (2010). "A survey of nectar feeding by spiders in three different habitats" (PDF). Bulletin of Insectology. 63 (2): 203–208. ISSN 1721-8861. Retrieved 20 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Meehan, Christopher J. (13 October 2009). "Herbivory in a spider through exploitation of an ant–plant mutualism". Current Biology. 19 (19). Elsevier Ltd.: R892–R893. Retrieved 20 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
^ Land, Michael F. (2006). "General-purpose and special-purpose visual systems". In Eric Warrant, Dan-Eric Nilsson (ed.). Invertebrate vision (PDF). Cambridge University Press. p. 167-210. ISBN 0521830885. Retrieved 21 June 2011. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

External links

[RFB2004CheliGen-1] Ruppert, E.E., Fox, R.S. and Barnes, R.D. (2004). "Chelicerata: Form". Invertebrate Zoology (7 ed.). Brooks / Cole. p. 555. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)

[RFB2004ArachGen-2] Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Chelicerata: Arachnida". Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 559–564. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)

[RFB2004Spiders-3] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Chelicerata: Araneae". Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 571–584. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)

[PlatnickEtcCatalPortia-4] Platnick, Norman I. (2011). "World Spider Catalog - Genus Portia". The American Museum of Natural History. Retrieved 4 May 2011. {{cite web}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[MaddisonEtc2008SaltPhylRev-5] Maddison, Wayne P. (2008). "Salticid spider phylogeny revisited, with the discovery of a large Australasian clade (Araneae: Salticidae)" (PDF). Zootaxa. 1893. Magnolia Press: 49–64. ISSN 1175-5334. Retrieved 14 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[PlatnickEtcCatalPorLab-6] Platnick, Norman I. (2011). "World Spider Catalog - Portia labiata". The American Museum of Natural History. Retrieved 21 June 2011. {{cite web}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[JacksonHallas1986Comp-7] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o ^p ^q ^r ^s ^t ^u ^v ^w ^x ^y ^z ^aa ^ab ^ac ^ad ^ae ^af ^ag ^ah ^ai ^aj ^ak ^al ^am ^an ^ao ^ap ^aq ^ar ^as ^at ^au ^av Jackson, Robert R. (1986). "Comparative biology of jumping spiders Portia africana, P. albimana, P. fimbriata, P. labiata and P. schultzi, areanophagic, web-building jumping spiders (Araneae: Salticidae) utilisation of webs, predatory versatility, and intraspecfic interactions". New Zealand Journal of Zoology. 13: 423–489. ISSN 0301-4223. Retrieved 17 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[Wanless1978RevisPortia-8] Wanless, F.R. (1978). "A revision of the spider genus Portia (Araneae: Salticidae)" (PDF). Bulletin of the British Museum (Natural History) Zoology. 34 (3). London: British Museum: 83–124. Retrieved 14 June 2011.

[HarlandJackson2000Cats-9] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j Harland, D.P., and Jackson, R.R. (2000). ""Eight-legged cats" and how they see - a review of recent research on jumping spiders (Araneae: Salticidae)" (PDF). Cimbebasia. 16: 231–240. Retrieved 5 May 2011.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[Forster1977SaltHunting-10] ^ ^a ^b ^c ^d ^e ^f Forster, Lyndsay M. (1977). "A qualitative analysis of hunting behaviour in jumping spiders (Araneae: Salticidae)" (PDF). New Zealand Journal of Zoology. 4: 51–62. Retrieved 27 April 2011.

[Jackson1986VersEvol-11] Jackson, Robert R. (1986). "Web Building, Predatory Versatility, and the Evolution of the Salticidae". In William A. Shear (ed.). Spiders - webs, behavior, and evolution. Stanford University Press. p. 492. ISBN 9780804712033. Retrieved 22 May 2011.

[Hillyard2007Private-12] Hillyard, Paul D. (2007). "Jumping spiders". The private life of spiders. New Holland Publishers. pp. 33–35. ISBN 9781845376901. Retrieved 11 April 2011.

[JacksonHallas1986SpartVersa-13] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o Jackson, Robert R. (1986). "Predatory versatility and intraspecies interactions of spartaeine jumping spiders (Araneae, Salticidae): Brettus adonis, B. cingulata, Cyrba algerina, and Phaeacius sp. indet". New Zealand Journal of Zoology. 13: 491–520. ISSN 0301-4223. Retrieved 1 April 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[RichmanJackson1992Etho-14] Richman, David B. (1992). "A review of the ethology of jumping spiders (Araneae, Salticidae)" (PDF). Bulletin of the British Arachnology Society. 9 (2): 33–37. Retrieved 12 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[Hogue1993LaAm-15] Hogue, Charles Leonard (1993). "Jumping spiders". Latin American insects and entomology. University of California Press. p. 117. ISBN 9780520078499. Retrieved 11 April 2011.

[Anderson1998Chelicerata-16] Anderson, D.T. (1998). "The Chelicerata". In D.T. Anderson (ed.). Invertebrate Zoology (1 ed.). Oxford University Press Australia. p. 467. ISBN 0915539419. {{cite book}}: Check |isbn= value: checksum (help)

[HarlandJackson2000Cues-17] Harland, D.P. (2000). "Cues by which Portia fimbriata, an araneophagic jumping spider, distinguishes jumping-spider prey from other prey". The Journal of Experimental Biology. 203. Company of Biologists: 3485–3494. Retrieved 4 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)

[HarlandJackson2004Umwelt-18] Harland, Duane P. (2004). "Portia Perceptions: The Umwelt of an Aranephagic Jumping Spider". In Frederick R. Prete (ed.). Complex worlds from simpler nervous systems. MIT Press. pp. 5–40. ISBN 9780262661744. Retrieved 12 April 2011. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[WilcoxJackson1998Cogn-19] Wilcox, R. Stimson (1998). "Cognitive Abilities of Araneophagic Jumping Spiders". In Russell P. Balda, Irene Maxine Pepperberg, Alan C. Kamil (ed.). Animal cognition in nature: the convergence of psychology and biology in laboratory and field. Academic Press. ISBN 9780120770304. Retrieved 23 May 2011. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)CS1 maint: multiple names: editors list (link)

[JacksonEtc2006ConfineProb-20] Jackson, Robert R. (2006). "Geographic Variation in a Spider's Ability to Solve a Confinement Problem by Trial and Error". International Journal of Comparative Psychology. 19: 282–296. Retrieved 8 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[JarmanJackson1986Taieria-21] Jarman, Elizabeth A.R. (1986). "The biology of Taieria erebus (Araneae, Gnaphosidae), an araneophagic spider from New New Zealand: silk utilisation and predatory versatility". New Zealand Journal of Zoology. 13: 521–540. ISSN 0301-4223. Retrieved 1 April 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[HoeflerEtc2006Biocontrol-22] Hoefler, Chad D. (2006). "The Potential of a Jumping Spider, Phidippus clarus, as a Biocontrol Agent" (PDF). J. Econ. Entomol. 99 (2): 432–436. ISSN 0022-0493. Retrieved 25 April 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[Hill2010Loc-23] Hill, David Edwin (2010). "Use of location (relative direction and distance) information by jumping spiders (Araneae, Salticidae, Phidippus) during movement toward prey and other sighted objectives" (PDF). Peckhamia. 83 (1): 1–103. ISSN 1944-8120. Retrieved 12 April 2011. {{cite journal}}: Unknown parameter |month= ignored (help)

[Piper2007Portia-24] Piper, Ross (2007). "The Quest for Food: Portia spider". Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals (PDF). Westport, CT 06881: Greenwood Press. pp. 98–100. ISBN 978–0–313–33922–6. Retrieved 31 March 2011. {{cite book}}: Check |isbn= value: invalid character (help)CS1 maint: location (link)

[JacksonBlest1982Dist-25] Jackson, R.R. (1982). "The distances at which a primitive jumping spider, Portia fimbriata, makes visual discriminations" (PDF). Journal of Experimental Biolology. 97: 441–445. Retrieved 12 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[HarlandEtc1999VisDist-26] Harland, Duane P. (1999). "Distances at which jumping spiders (Araneae: Salticidae) distinguish between prey and conspecific rivals". Journal of Zoology. 247 (3): 357–364. doi:10.1111/j.1469-7998.1999.tb00998.x. Retrieved 4 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help); Unknown parameter |month= ignored (help)

[RFB2004ArthroSens-27] Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). "Arthropoda: Sense organs". Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 532–537. ISBN 0030259827.{{cite book}}: CS1 maint: multiple names: authors list (link)

[Richman2008Ency-28] Richman, David B. (2008). "Jumping Spiders (Class Arachnidae, Order Araneae, Family Salticidae)". In John L. Capinera (ed.). Encyclopedia of entomology. Springer. pp. 2066–2068. ISBN 9781402062421. Retrieved 11 April 2011.

[Forster1977Feed-29] Forster, Lyn M. (1977). "Some factors affecting feeding behaviour in young Trite auricoma spiderlings (Araneae: Salticidae)". New Zealand Journal of Zoology. 4. The Royal Society of New Zealand: 435–442. Retrieved 24 April 2011. {{cite journal}}: Unknown parameter |month= ignored (help)

[WilcoxJackson2000Tricks-30] Wilcox, S. and Jackson, R. (2002). "Jumping Spider Tricksters". In Bekoff, M., Allen, C., and Burghardt, G.M. (ed.). The Cognitive Animal: Empirical and Theoretical Perspectives on Animal Cognition (PDF). MIT Press. pp. 27–34. ISBN 0262523221. Retrieved 12 May 2011.{{cite book}}: CS1 maint: multiple names: authors list (link)

[JacksonPollard1997SexCann-31] Jackson, Robert R. (1997). "Jumping spiders mating strategies: sex among cannibals in and out of webs". In Jae C. Choe, Bernard J. Crespi (ed.). The Evolution of Mating Systems in Insects and Arachnids. Cambridge University Press. p. 340-351. ISBN 9780521589765. Retrieved 6 June 2011. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[WilcoxEtc1996Smokescreen-32] Wilcox, R. Stimson (1996). "Spiderweb smokescreens: spider trickster uses background noise to mask stalking movements" (PDF). Animal Behaviour. 51. The Association for the Study of Animal Behaviour: 313–326. ISSN 0003–3472. Retrieved 23 May 2011. {{cite journal}}: Check |issn= value (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[LiEtcPreyPref-33] ^ ^a ^b ^c ^d ^e ^f ^g Li, Daiqin (1997). "Prey preferences of Portia labiata, P. africana, and P. schultzi, araneophagic jumping spiders (Araneae: Salticidae) from the Philippines, Sri Lanka, Kenya, and Uganda". New Zealand Journal of Zoology. 24 (4): 333–349. doi:10.1080/03014223.1997.9518129. Retrieved 21 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[NelsonEtc2004AntsSalt-34] ^ ^a ^b ^c ^d ^e ^f Nelson, Ximena J. (2004). "Predation by ants on jumping spiders (Araneae: Salticidae) in the Philippines" (PDF). New Zealand Journal of Zoology. 31. The Royal Society of New Zealand: 45–56. ISSN 0301–4223. Retrieved 13 June 2011. {{cite journal}}: Check |issn= value (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[NelsonJackson2011Flex-35] Nelson, Ximena J. (2011). "Flexibility in the foraging strategies of spiders". In Marie Elisabeth Herberstein (ed.). Spider Behaviour: Flexibility and Versatility. Cambridge University Press. p. 45-47. ISBN 9780521749275. Retrieved 29 May 2011. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[JacksonEtc2001Nectar-36] Jackson, Robert R. (2001). "Jumping spiders (Araneae: Salticidae) that feed on nectar" (PDF). Journal of Zoolology. 255. The Zoological Society of London: 25–29. Retrieved 18 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[Gaskett2007SpidSexPher-37] Gaskett, A.C. (2007). "Spider sex pheromones: emission, reception, structures, and functions" (PDF). Biological Reviews. 82. Cambridge Philosophical Society: 27–48. doi:10.1111/j.1469-185X.2006.00002.x. Retrieved 23 June 2011.

[ClarkJackson1995DragPlPfMF-38] Clark, R.J. (1995). "Dragline-mediated sex recognition in two species of jumping spiders (Araneae Salticidae), Portia labiata and P. fimbriata". Ethology Ecology & Evolution. 7: 73–77. Retrieved 13 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[RobertsEtc2003EarlyWolf-39] Roberts, J. Andrew (2003). "Kinship and food availability influence cannibalism tendency in early-instar wolf spiders (Araneae: Lycosidae)" (PDF). Behaviour, Ecology and Sociobiology. 54. Springer-Verlag: 416–422. doi:10.1007/s00265-003-0646-8. Retrieved 18 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[ClarkJackson1994DragPlFF-40] Clark, R.J. (1994). "Self recognition in a jumping spider: Portia labiata females discriminate between their own draglines and those of conspecifics". Ethology Ecology & Evolution. 6: 371–375. Retrieved 13 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[ClarkEtc1999DragFightAbility-41] Clark, Robert J. (1999). "Draglines and Assessment of Fighting Ability in Cannibalistic Jumping Spiders". Journal of Insect Behavior. 12 (6). Springer: 753–766. doi:10.1023/A:1020900925863. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[ClarkJacksonLabEggSac-42] Clark, Robert J. (1994). "Portia labiata, a cannibalistic jumping spider, discriminates between own and foreign eggsacs". International Journal of Comparative Psychology. 7 (1): 38–43. ISSN 0889-3667. Retrieved 19 June 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[ProszynskiDBPorLab-43] Proszynski, Jerzy. "Global Species Database of Salticidae (Araneae): Portia labiata (Thorell, 1887)". Global Species Database of Salticidae (Araneae). Warsaw: Museum and Institute of Zoology. Retrieved 21 June 2011.

[BarrionLitsinger1995Riceland-44] Barrion, A.T. (1995). Riceland spiders of South and Southeast Asia. International Rice Research Institute / C.A.B. International. p. 45. ISBN 9780851989679. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[ChenEtc2010Nectar-45] Chen, Xiaoqiong (2010). "A survey of nectar feeding by spiders in three different habitats" (PDF). Bulletin of Insectology. 63 (2): 203–208. ISSN 1721-8861. Retrieved 20 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[MeehanEtc2009Bagh-46] Meehan, Christopher J. (13 October 2009). "Herbivory in a spider through exploitation of an ant–plant mutualism". Current Biology. 19 (19). Elsevier Ltd.: R892–R893. Retrieved 20 May 2011. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[LandNilsson2006Vision-47] Land, Michael F. (2006). "General-purpose and special-purpose visual systems". In Eric Warrant, Dan-Eric Nilsson (ed.). Invertebrate vision (PDF). Cambridge University Press. p. 167-210. ISBN 0521830885. Retrieved 21 June 2011. {{cite book}}: More than one of |pages= and |page= specified (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

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