Pseudoscorpions! Small, strange arachnids

Yes, this is a blog about spiders, and no, pseudoscorpions are not spiders. But they are members of the class Arachnida, like spiders, and fascinating, like spiders! I encountered them for the first time on a recent trip to the Okanagan with Sean, so here’s a post about some of their natural history.

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Two tiny pseudoscorpions Sean and I found while doing some rock flipping on a hillside near Vaseux Lake, BC. Photo: Sean McCann

Pseudoscorpions are really weird, and really awesome, creatures. Their name means “false scorpion” (which is also a common name for this order of arachnids), because superficially they look a lot like scorpions (members of another arachnid order), minus the “tail” with its stinger on the end.

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Drawings of a pseudoscorpion and a scorpion from JH Comstock’s book. The illustrations are most likely by Anna C. Stryke, but possibly by Mrs. Comstock. Note that the scale is different for each drawing.

Pseudoscorpions are tiny. The one below (photographed under a microscope) is only about 1 mm long! The largest pseudoscorpions can get as long as 10 mm. Their small size means they can live in tight spaces, like between floorboards, under tree bark, or even under the elytra (hardened forewings) of beetles – but more on this later.

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Pseudoscorpion found in Vero Beach, FL. Photo: Sean McCann

Pseudoscorpion morphology is strange. Like other arachnids, pseudoscorpions have two main body segments: a cephalothorax (the front part, a combined head-and-thorax bearing all of their appendages), and an abdomen. They may have one or two pairs of simple eyes on the sides of the cephalothorax (or none at all) and their vision is generally poor. As well as four pairs of legs, they have enormous chelate (pincer-like) pedipalps that they use for capturing prey and sensing their environment.

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A neobisiid pseudoscorpion nicely displaying its chelate jaws and pedipalps. Photo: Marshal Hedin, licensed under CC BY-NC 2.0.

Order Pseudoscorpionida

Pincer-like chelicera (singular of chelicerae) of a pseudoscorpion, bearing spinnerets on the movable finger. From Comstock’s book.

 

Their jaws (called chelicerae) are also like miniature pincers. Like spiders, pseudoscorpions can produce silk. Unlike spiders, who have abdominal silk glands and spinnerets, pseudoscorpions’ silk glands are in their cephalothoraxes, and their spinnerets are on the tips of their chelicerae! (The spitting spiders in the family Scytotidae are exceptional among spiders in also having silk glands in their cephalothoraxes, and “spitting” the silk out of their fangs along with venom.)

Pseudoscorpions use silk to build retreats* or cocoons for moulting, overwintering, and sometimes brooding their young.

Also like spiders, many pseudoscorpions use venom to subdue their prey, which includes mites and other tiny arthropods. The venomous pseudoscorpions are in suborder Iocheirata, which means “poison hands”. Their venom glands are in their modified pedipalps, with openings in the tips of one or both of the fingers of their claws.

 

To me, pseudoscorpion anatomy is all topsy-turvy (at least compared to spiders):        their silk comes out the wrong end and their venom comes out of clawed “hands” instead of fangs. 

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Pseudoscorpion – modified from the illustration in Comstock’s book.

Pseudoscorpions also have pretty strange reproduction. Males deposit a spermatophore (a package of sperm) on the ground, which a female must then pick up and insert into her reproductive opening. Males of different kinds of pseudoscorpions have various methods of ensuring that a female finds and uses their sperm. Some are carefree: the male deposits the spermatophore, walks away and (figuratively) crosses his fingers and hopes that a female will encounter it by chance. Other tactics are rather more direct and reliable: the male engages the female in an elaborate “mating dance” and eventually pulls her over his spermatophore to ensure that she picks it up. In one species, Serianus carolinensis, males only produce spermatophores when a female is nearby, then they spin special silk webs that direct her to the package.

Once a female’s eggs are fertilized, she keeps them in a brood pouch under her abdomen (or sometimes next to her in a silken brooding chamber). This brood-sac contains food for the developing pseudoscorpion embryos, which grow and moult into protonymphs (juveniles that look just like adults, but smaller) before emerging.

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A female Arctic pseudoscorpion, Wyochernes asiaticus, with her brood pouch. Photo: Crystal Ernst (used with permission)

Pseudoscorpions love books! Although I would like to think the little pseudoscorpion in the photograph below enjoys reading, what they really like about books is the booklice that sometimes live in them. Because pseudoscorpions can sometimes be found living between the pages of books and feeding on booklice, one common name for them is “book scorpions”.

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“Book scorpion” enjoying some literature while waiting for booklice. Photo: Sean McCann

Finally, pseudoscorpions are hitchhikers! Because they are so small and don’t fly, pseudoscorpions can’t get very far on their own. To overcome this obstacle, they hitch lifts on other organisms – usually larger arthropods like beetles and flies. This particular kind of symbiosis – in which the individual doing the carrying is apparently unharmed – is called phoresis.

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A longhorn beetle, Xylotrechus sagittatus, with phoretic pseudoscorpions hitching a ride on its leg. Photo: Sean McCann

Phoresis is a fantastic word that comes from Greek: phor means “carry, bear; movement” but it can also mean “thief”. Phoretic pseudoscorpions latch onto the bodies of their transportation with their pincers to steal a free ride. Some pseudoscorpions are stowaways under the elytra of comparatively gigantic harlequin beetles, and feed on phoretic mites and find mates while they travel. Piotr Naskreki has a wonderful blog post with pictures of these tiny ecosystems on a beetle’s back.

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A closer view of the cluster of tiny travelers. Photo: Sean McCann

Want to find out more about pseudoscorpions? Of course you do! Here are some references and further reading from around the web:

10 Facts about Pseudoscorpions. Fantastic blog post by arachnologist Chris Buddle, who also created this great photographic key to the pseudoscorpions of Canada.

Comstock JH (1912). THE SPIDER BOOK: a manual for the study of the spiders and their near relatives, the scorpions, pseudoscorpions, whip-scorpions, harvestmen, and other members of the class Arachnida, found in America north of Mexico, with analytical keys for their classification and popular accounts of their habits. Doubleday, Page & Company. (Available from the Biodiversity Heritage Library, with wonderful illustrations)

Harvey MS (2013). Pseudoscorpions of the World, version 3.0. Western Australian Museum, Perth.

Pseudoscorpions page on the Massey University Guide to New Zealand Soil Invertebrates website. *Includes photos of pseudoscorpions’ silk retreats!

Pseudoscorpions page on the Encyclopedia of Life.

The order of the Pseudoscorpiones. Nice summary of pseudoscorpion biology by F. Schramm, with lots of photos and scholarly references.

Zeh DW & Zeh JA (1992). On the function of harlequin beetle-riding in the pseudoscorpion, Cordylochernes scorpioides (Pseudoscorpionida: Chernetidae). Journal of Arachnology, 47-51.

A cunning crab spider

Sean McCann recently returned from an epic journey through the rainforest of Guyana in search of caracaras, and is busy blogging about his adventures over at Ibycter.com (so you should check it out!). To make up for not taking me along, he found and photographed lots of awesome spiders for me to blog about. Here’s the first one!

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Here we have a male crab spider (family Thomisidae) with its prey: an ant in the genus Dolichoderus*. At first glance, this may not seem particularly exciting. A small, hairy, black spider eats an insect. What’s so special about this scene?

Well, the spider appears to be Strophius nigricans, reported to be a specialist predator of ants. Most animals are not big on eating ants because they are generally distateful and well-defended by strong mandibles, stings, and defensive compounds. So specializing on ants is not particularly common, and tends to come along with some neat adaptations.

Strophius nigricans is not well studied, but I managed to find one paper about its predation behaviour. Oliveira and Sazima (1985) observed a male S. nigricans carrying an dead worker ant (Camponotus crassus) in the field in Brazil, and took it back to the laboratory to make some observations. The spider never let go of his ant carcass – this would come in handy later. In captivity, he was provided with some more C. crassus workers, and here his secrets were revealed.

The Strophius male used his ant corpse as a shield for protection against ants during predation. To eat ants, one must spend time near ants, but they don’t take kindly to intruders. The dead-ant-shield provides a clever disguise. Most ants don’t have especially good vision, and instead rely mainly on their senses of touch and smell. Whenever the Strophius male was approached by an ant, he would present his previous meal, which (obviously) feels just like another ant, and, if recently deceased, probably smells right too. As the spider pursued his prey, always attempting to sneak up from behind, he held the ant corpse aloft. Once within striking distance of an unsuspecting ant, he quickly dropped the shield and bit his new victim. Once dead, this new ant was used as a shield.

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Unlike a related crab spider species that preys on ants, Aphantochilus rogersi (a remarkable ant-mimic), Strophius doesn’t look all that much like its victims.  A. rogersi specializes on Cephalotes ants, which have relatively good vision, but its excellent mimicry is probably more important as a defense against visually hunting predators that avoid eating ants. However, a similar mode of defense is not necessarily out of the question for Strophius. The ants it preys on are black and have white hairs on the abdomen. From above, the spider carrying its ant-shield might look just enough like an ant carrying its comrade to fool a potential predator such as a bird.

As it turns out, the conclusion that Strophius nigricans are specialists on the ant species Camponotus crassus appears to be based on Oliveira and Sazima’s observations of the single male discussed above. Here we’ve seen that the spider also takes another species, and it seems that its prey capture technique should work just fine as long as the shield matches the subsequent target prey species. The visual mimicry would presumably be just as effective for any similar sized black ants. This certainly seems like a cool system that could use more investigation!

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*Thanks Alex Wild for the ID!

References:

Oliveira, P. S., & Sazima, I. (1984). The adaptive bases of ant‐mimicry in a neotropical aphantochilid spider (Araneae: Aphantochilidae). Biological Journal of the Linnean Society, 22(2), 145-155. (pdf)

Oliveira, P. S., & Sazima, I. (1985). Ant-hunting behaviour in spiders with emphasis on Strophius nigricans (Thomisidae). Bulletin of the British Arachnological Society.

Ghost spiders (Anyphaenidae)

Recently, Sean found a lovely pale spider in his mother’s garden in Victoria. Neither of us recognized it, and after a bit of research, I was able to identify it as a member of the family Anyphaenidae. Spiders in this family are called ghost spiders, and this is the first one I have ever encountered. She is now living in a petri dish on our bookshelf, and has made herself a cozy silk retreat inside a curled leaf.

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Anyphaena aperta female (missing her left front leg) photo: Sean McCann

Not much is known about the natural history of these secretive spiders. Apparently they are generally nocturnal hunters that spend their days in their silken retreats. Apart from building these “sleep sacs” under leaves or rocks (and of course spinning egg sacs), ghost spiders don’t have much use for silk.

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Anyphaena accentuata in her silken retreat under a leaf. Photo: Ferran Turmo Gort (licensed under CC BY-NC-SA 2.0)

Ghost spider body forms and lifestyles are similar to those of the sac spiders (family Clubionidae), and in fact they used to be considered clubionids. They also have some similarities to ground spiders in the family Gnaphosidae. Members of all three families are wanderers with prominent spinnerets. Luckily for me and others without regular access to a microscope, there are some fairly obvious characteristics that help distinguish Anyphaenidae from other spider families.

Less reliable, but easier to see, is the eye arrangement.

Ground spiders in the family Gnaphosidae often have some oval eyes, whereas anyphaenids’ eyes are all round.

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A ground spider in the genus Zelotes (Gnaphsosidae) with her lovely pink egg sac. Note the barrel-shaped spinnerets. I know you can’t really see her eyes – click the link above for a drawing. Photo: Sean McCann

And unlike many sac spiders in the family Clubionidae, the anterior median eyes (AME; the central pair closest to the front of the head) are usually smaller than the other eyes.

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Sac spider with relatively large anterior median eyes. Photo: Sean McCann

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Anyphaena eye arrangement. The anterior median eyes (central pair in the front row) are smallest. Photo: Dann Thombs (licensed under  CC BY-ND-NC 1.0)

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Dorsal view of eye arrangement of an Anyphaena male. Photo: Kyron Basu (licensed under CC BY-ND-NC 1.0)

The real giveaway characteristic of anyphaenids, however, is the tracheal spiracle (one of the openings of the respiratory system in spiders). To get a good look at this, it’s useful to have a Spi-Pot. In other similar-looking spiders the tracheal opening is located close to the spinnerets. In ghost spiders, it is closer to the centre of the abdomen.

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Ventral view of a male Anyphaena aperta. Photo: Ken Schnieder (licensed under CC BY-ND-NC 1.0)

The horizontal groove on the underside of this spider’s abdomen (across the darker patch in the middle) is the tracheal spiracle.

Although not much to look at from the outside, the tracheal system of the anyphaenids may provide us with some clues about their natural history. Internally, the tracheae of ghost spiders are much wider than those of sac spiders. This may be because they have a more active lifestyle than sac spiders; they are certainly very fast runners. Males have even larger tracheae than females, which Platnick (1974) concluded is likely related to their extremely vigorous courtship displays. During courtship,  Anyphaena accentuata males vibrate their abdomens up and down so quickly that it becomes a blur. In contrast, sac spider courtship has been described as “sluggish”.

 

The ghost spiders could certainly use more study, and I’ll be keeping an eye out for them from now on!

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Anyphaena aperta again. Photo: Sean McCann

 

References

Dondale, C. D., & Redner, J. H. (1982). The insects and arachnids of Canada. Part 9. The sac spiders of Canada and Alaska. Araneae: Clubionidae and Anyphaenidae (No. 1724). link

Platnick, N. (1974). The spider family Anyphaenidae in America north of Mexico. Bull Mus Comp Zool Harvard Univ. 146: 205-266. link

 

Bolas spiders: masters of deception

Bolas spiders (members of the genus Mastophora, in North America) are famous for their unusual prey capture technique: rather than a web, they produce a single silk line with a super-sticky ball of glue at the end, which they fling at their prey.

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Female Mastophora cornigera hunting with her ‘bolas’. (Photo: Matt Coors)

The common name ‘bolas spider’ is not particularly accurate, though. A real bolas – two or more weights connected by cord – is swung and thrown at an animal (like a horse, in the image below) in its entirety, and works by getting tangled around the legs of the target.

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By John Miers [Public domain], via Wikimedia Commons

The spider’s ‘bolas’ differs in that it never leaves its owner’s grip, and works by getting stuck to the target, which is invariably a moth. Eberhard (1980) observed that a more appropriate name would be “sticky yo-yo spiders”. The sticky yo-yo prey capture technique is impressive enough (inspired by their speed and accuracy with the bolas, Eberhard named one species dizzydeani for Jerome “Dizzy” Dean, one of the greatest baseball pitchers of all time), but to fully appreciate the wonders of bolas spider biology, we must delve into the secret lives of these aromatic and cryptic spiders. They are masters of deception, both olfactory and visual.

Seductive scents

Hunting with a sticky yo-yo is all very fierce and exciting, but what are the chances that a moth is ever going to fly close enough for the spider to swing at? Not very high. Unless, like the bolas spider, you have a trick or two up your sleeve… er, leg… coverings. Adult female bolas spiders have the incredible ability to produce a chemical cocktail that make them smell just like female moths advertising for mates (actually, no one knows yet which part of the spider’s body is responsible for this wonderful trick). Innocent male moths following what they perceive to be a pheromone trail (whose chemical message indicates that it leads to a sexually receptive female moth) are thus duped into coming in close enough for the spider to strike. This is called “aggressive chemical mimicry”, and it is awesome.

Moth sex pheromones are typically blends of two or more chemical compounds in very specific ratios. The particular chemicals and ratios allow male moths to discriminate between females of their own and other species. If a bolas spider produced just one moth pheromone, they probably wouldn’t do very well, because their diet would be restricted to only a single moth species. It turns out that each species of bolas spider attracts several kinds of male moths, and the best studied of these is Mastophora hutchinsoni.

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Bristly cutworm moth, Lacinopolia renigera. (Photo: Andy Reago & Chrissy McClarren; licensed under CC BY 2.0)

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Smoky tetanolita, Tetanotolita mynesalis. (Photo: kestrel 360; licensed under CC BY-NC-ND 2.0)

Mastophora hutchinsoni attracts four kinds of moths, but more than 90% of their prey consists of two species in the family Noctuidae: the smoky tetanolita (Tetanolita mynesalis) and the bristly cutworm (Lacinopolia renigera). These two moth species produce entirely different sex pheromones, and they are active at different times of night. The problem for the bolas spider is that the bristly cutworm pheromone interferes with the attractiveness of the smoky tetanolita’s pheromone.

 

Here’s where the bolas spiders start to get really fancy. Let’s join an M. hutchinsoni female for a night of hunting, and learn some of her secrets.

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M. cornigera female preparing for a night of moth hunting. (Photo: Matt Coors)

She begins by building her horizontal “trapeze” line, from which she then hangs motionless, with front legs extended in hunting position (but with no bolas, yet). She is already emitting the sex pheromones (well, analogs that are close enough!) of both prey species, but so far, only the early-flying bristly cutworm is active. They aren’t put off by the smell of smoky tetanolita females mixed in with the pheromone of a female of their own species, and soon one is winging its way toward the seductive scent coming from the female spider. It passes close by, but out of reach. This moth is lucky, for now. But no matter; his fate is not our immediate concern. The spider’s outstretched legs are covered with tiny vibration-sensitive hairs (called trichobothria) that allow her to detect the sound of the moth’s wing beats nearby. Now that she knows there is prey about, she springs into action and spends the next minute or two building her sticky bolas.

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Female M. cornigera hunting with her bolas. (Photo: Matt Coors)

 

Once her weapon is complete, she returns to her prey-capture position, with the bolas hanging from one of her outstretched front legs. For her next trick, she will again rely on her ability to detect the wingbeats of flying moths with her leg hairs. She waits patiently, silent and still.

 

 

 

Soon, another hapless male moth picks up the scent and starts winging towards the bolas spider. When her sensory hairs tell her the time is just right, she takes a swing at the approaching moth and connects. Although the moth struggles, shedding scales in its effort to escape, the wet stickiness of the bolas holds it fast. The spider reels the moth in and delivers a fatal venomous bite. She waits a few moments, then wraps her prize in swathes of silk and hangs it carefully from her trapeze line to eat later. The night is young, and the moths will continue flying for some hours yet. The bristly cutworms will remain active until 22:30. Our spider builds a fresh bolas, and settles in to wait. Gradually, the smell of bristly cutworm sex pheromone coming from the spider fades. The smell never disappears entirely, but is soon faint in comparison to the scent of a female smoky tetanolita. The smoky tetanolita males will come out after 23:00, and our spider will be ready for more deadly deception.

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Female M. phrysonoma with captured moths. (Photo: Keith Simmons; licensed under CC BY-NC-SA 2.0)

So far, we’ve discovered some of the adult female bolas spider’s secrets to success, but what about juveniles and males? They don’t use a bolas, but they are no less stealthy and deceitful than their counterparts. Young bolas spiders are also employ aggressive chemical mimicry to attract prey, but they specialize on male moth flies in the family Psychodidae. Each bolas spider species is especially attractive to a particular species of moth flies. It appears to be a pleasing coincidence that small bolas spiders prey on moth flies until they graduate to real moths. Whether or not the sex pheromones of the psychodids captured by each spider are similar to those of moths they specialize on is currently a mystery.

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Moth fly (Psychodidae). (Photo: Ted C. MacRae)

Optical illusions

Mastophora females are not only masters of chemical deception, but they are also visually cryptic, and hide in plain sight from their own potential predators. They do this by mimicking bird poop.

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Excellent bird-poop mimicry by Mastophora cornigera. (Photo: Matt Coors)

The female spiders spins herself a silken mat on the surface of a leaf, and clings to it with her legs drawn in tightly around her cephalothorax.

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Another bird poop mimic, female M. phrysonoma, with visitors! (Photo: Matt Coors)

But wait, what are those tiny red things? At first glance, they could easily be mistaken for mites, but no! These are tiny males, presumably interested in mating with the comparatively massive female. Bolas spider males are usually less than 2 mm long, while females are typically 10 – 15 mm long, and sometimes as large as 2 cm!

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Another shot of the incredible bird poop mimicry and extreme sexual dimorphism of M. phrysonoma. (Photo: Matt Coors)

Because the females are so cryptic and males are so tiny, almost nothing is known about the sexual behaviour of bolas spiders. As a researcher studying sexual communication and mating behaviour in spiders, I sure would love to know how the males in the photos above found the female and what happened next! Most likely, the female bolas spiders produce attractive sex pheromones just like the moths whose chemical communication they exploit. As far as I am aware, however, no one has investigated the sex pheromones of bolas spiders. One hypothesis that might explain the evolution of their mimicry of moth pheromones is that their own chemical signals have compounds in common with those of their prey. In fact, this is a hypothesis that Andy Warren is investigating for a different group of spiders that also mimic moth pheromones – orb weavers in the genus Argiope.

If you’re not familiar with spider systematics, it might seem odd that two groups of spiders that look so different and have such different prey-capture techniques share the amazing ability to lure male moths to their doom. In fact, bolas spiders are orb-weavers (at least, they are members of the orb-weaver family Araneidae), they just don’t build webs like most of their relatives. Like orb-weavers, bolas spiders regularly eat their silken traps and recycle the silk proteins to use another day.

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Argiope aurantia female on her orb-web. (Photo: Suzanne Cadwell; licenced under CC BY-NC 2.0)

See the family resemblance?

To see a female bolas spider in action, check out this video clip from David Attenborough’s “Life in the Undergrowth” series. (The bolas spider segment starts at 3:00, but the first few minutes about the redback spider are also worth a watch!)

 

Special thanks to Matt Coors for kindly letting me feature his fantastic photographs in this post!

References:

Eberhard, W. G. (1980). The natural history and behavior of the bolas spider Mastophora dizzydeani sp. n. (Araneidae). Psyche: A Journal of Entomology87(3-4), 143-169. http://dx.doi.org/10.1155/1980/81062

Haynes, K. F., Yeargan, K. V., & Gemeno, C. (2001). Detection of prey by a spider that aggressively mimics pheromone blends. Journal of insect behavior,14(4), 535-544. http://link.springer.com/article/10.1023/A:1011128223782

Haynes, K. F., Gemeno, C., Yeargan, K. V., Millar, J. G., & Johnson, K. M. (2002). Aggressive chemical mimicry of moth pheromones by a bolas spider: how does this specialist predator attract more than one species of prey? Chemoecology, 12(2), 99-105. http://link.springer.com/article/10.1007%2Fs00049-002-8332-2?LI=true

Yeargan, K. V. (1988). Ecology of a bolas spider, Mastophora hutchinsoni: phenology, hunting tactics, and evidence for aggressive chemical mimicry. Oecologia, 74(4), 524-530. http://www.jstor.org/stable/4218505

Yeargan, K. V. (1994). Biology of bolas spiders. Annual review of entomology39 (1), 81-99. DOI: 10.1146/annurev.en.39.010194.000501 

Yeargan, K. V., & Quate, L. W. (1996). Juvenile bolas spiders attract psychodid flies. Oecologia, 106(2), 266-271. http://link.springer.com/article/10.1007/BF00328607

Yeargan, K. V., & Quate, L. W. (1997). Adult male bolas spiders retain juvenile hunting tactics. Oecologia, 112(4), 572-576. http://link.springer.com/article/10.1007/s004420050347

 

 

Spiders of Lost Lagoon

This weekend, I was lucky to spend a rare rain-free November morning in Vancouver’s famous Stanley Park with Sean McCann, who kindly provided all the photos for this post. Despite the late season and cool weather, it turned out to be a very spidrous (a term coined by Sean) day indeed! We must remember that the end of Arachtober does not necessarily mean the end of spider season, as Eric Eaton pointed out in a recent Spider Sunday post.

Our first spider-observation stop was outside the Lost Lagoon Nature House. What at first glance looked like a dirty and unremarkable garage door turned out, upon closer inspection, to be a veritable spider heaven!

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Here I am making notes on an impressive aggregation of spiders.

Although not quite on the massive scale of the aggregation of thousands of orbweavers inside a wastewater treatment plant that Gwen Pearson wrote about recently, I estimate that the area around this single garage door is home to several hundred spiders all living in extremely close proximity. In this case, the aggregated webs are just a couple metres away from the edge of Lost Lagoon, a freshwater habitat that produces large numbers of chironomid midges as well as larger aquatic insects. Lights attract these insects, making this the perfect hunting ground at night, when the spiders are most active.

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It’s hard to see individual spiders at this scale, but looking closer, they are everywhere!

Here’s a closer look  at one small area of the door (zooming in on the upper right corner of the photo above). Two large females rest in the open while a third hides away in a silken retreat next to a cluster of 3 egg sacs.

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Almost all the spiders in this aggregation were members of a single species, the bridge orbweaver Larinioides sclopetarius (family Araneidae). Here are some closer shots of a male and female:

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Male Lariniodes sclopetarius against the blue backdrop of the building.

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Female Lariniodes sclopetarius posing on some nearby vegetation.

We did find a few spiders other than bridge orbweavers on the building. A female L. sclopetarius and what’s most likely a juvenile Eratigena (formerly Tegenaria) appeared to be sharing the same retreat in a cement pillar.

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Moments before this photo was taken, these two were hunkered down close together in the pit in the cement, apparently  oblivious to one another.

Right on the other side of the same pillar, this male longjawed orbweaver Tetragnatha elongata was hanging out.

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It turns out that Lariniodes and Tetrgnatha are the usual suspects when it comes to communal living and megawebs. Spiders in both genera like to live near water as was the case here next to the lagoon. Lariniodes sclopetarius often build their webs on human-made structures, and especially prefer to be next to lights (Greene et al. 2010 and references therein). This preference was very evident at the Nature House – of the four identical  garage door sections of the side of the building facing the lagoon, the spiders were almost exclusively living on the only one with a light.

Although our first stop provided plenty of arachnological excitement for one day, we eventually moved on and explored the rest of the lagoon.

We found this male Pimoa altioculata and several others of the same species on webbing underneath the railing of a small footbridge.

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Spiders in the family Pimoidae are related to the linyphiids, and also build sheet-webs.

When Sean removed the spider from his web to try to get some better shots, he immediately went into “play dead” mode, tucking all his legs up tightly around his body for defense.

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Several more longjawed orbweavers were hanging out under the railings of the same bridge.

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Another Tetragnatha sp. These spiders are typically found near water.

We also found this beautiful example of a hammock-shaped sheet web built by a small linyphiid.

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In this shot you can see the silhouette of the spider at the upper left hand side of the “hammock.”

We admired the web for some minutes before noticing the resident spider, who eventually came out onto the web for the lovely shot below:

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You can tell from the bulbous pedipalps that this is a subadult male. I am pretty sure the species is Neriene digna.

All in all it was a wonderful day at the park. I hope your November is just as spidrous as ours has been so far here on the wet coast!

Reference:

Greene, A., Coddington, J. A., Breisch, N. L., De Roche, D. M., & Pagac, B. B. (2010). An Immense Concentration of Orb-Weaving Spiders With Communal Webbing in a Man-Made Structural Habitat (Arachnida: Araneae: Tetragnathidae, Araneidae). American Entomologist, 56(3), 146-156.

Spiders Unraveled! Outreach at Iona Beach

This weekend was, to my knowledge, the first ever spider day hosted by Metro Vancouver Regional Parks. Along with a fantastic crew of volunteer spider enthusiasts and park staff, I had the great pleasure of sharing some of the incredible biology of spiders and the joys of being a naturalist with kids and adults alike.

Here are some of the highlights of the day in photographs, kindly provided by Mike Boers and Sean McCann.

Gwylim Blackburn greeted visitors at the become a naturalist station. Here kids could make or borrow all the supplies they might need to be arachnologists for the day! These included ‘pooters’ (aspirators) made out of straws for collecting small spiders, personalized field notebooks, pencils, magnifying glasses, collecting containers, and a customized field guide to the spiders of Iona Beach.

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Gwylim with his table of supplies for creating the tools to become a naturalist! photo: Sean McCann

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Building pooters and field notebooks. Photo: Sean McCann

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A triumphant young naturalist holds her field notebook aloft, prepared for the adventures ahead! Photo: Sean McCann

Tanya Stemberger served up cricket smoothies and crunchy beetle larvae at the eat like a spider station. Here visitors learned about the health and environmental benefits of eating insects like spiders do! Read more about this part of the event here.

Tanya serves up a cricket smoothie for some kids who are brave enough to try entomophagy! Photo: Mike Boers

Some were a bit sceptical about the idea of eating insects at first.

But most were enthusiastic!

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Adding the cricket protein powder to the smoothie. Photo: Sean McCann

In the end, almost everyone accepted the challenge and won a special entomophagy achievement award for their bravery!

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Tanya shows off the highly prized entomophagy award certificate. Photo: Sean McCann

Next, kids got to participate in the spider olympics! The first event was to avoid predation by a spider.  The challenge was to to climb through a web without creating vibrations that would ring the dinner bells and alert the resident spider to the presence of potential prey!

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The spider web challenge! Photo: Sean McCann

Other spider olympic events included trying to jump as far as a jumping spider (they can jump distances up to 25 times their body length!) and run as fast as a house spider (330 body lengths in 10 seconds!).

The highlight of the day for most visitors was the spider tour of the beach with Sean McCann, where kids got to learn how to find spiders and put their new field notebooks and pooters to use!

Every hour Sean set out with a new group of naturalists to find the hidden treasures of Iona Beach. Photo: Mike Boers

Sean took his crew of naturalists to several habitats in search of spiders, starting with his favourite: under the driftwood.

Sean demonstrates his log-flipping technique. Photo: Mike Boers

Collecting a specimen! Photo: Mike Boers

There are all kinds of wonders to be found under the logs if you look closely! Photo: Mike Boers

The tour then moved on to using beating sheets to collect spiders from vegetation, and also included sampling the rich spider fauna on the walls of the washroom building.

Samantha Vibert and I introduced interested arachnophiles (and a lot of unsuspecting cyclists stopping for water/washroom breaks!) to basic spider anatomy and diversity. We had live specimens of several local species representing both wandering hunters and web-builders.

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Basic spider anatomy poster featuring a hobo spider, one of the commonest spiders found under the logs at Iona Beach. Created by Sean McCann

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I greet my first visitor of the day, with much excitement and waving of hands. Photo: Sean McCann

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Sam and one of our most enthusiastic young arachnologists, who stayed most of the day and went on 3 of the 4 spider walks! Photo: Sean McCann

All in all, I think the day was a great success! We had over 120 visitors, and a ton of fun. I hope it happens again next year!

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A wolf spider, Arctosa perita, on the sand of Iona Beach. Photo: Sean McCann

 

Here’s a small sample of the diverse spiders that we found on the day, or brought from nearby sites (photos by Sean McCann).

A comb-tailed spider

Last weekend, I joined a group of fellow arachnophiles for a day at Burns Bog. We did not achieve our goal of finding the rare ground spider Gnaphosa snohomish (a bog specialist), but instead we met a very common spider that is nonetheless not well known: a comb-tailed spider in the family Hahniidae.

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Neoantistea magna, a common yet mysterious forest-dweller (photo Sean McCann).

A distinguishing feature of spiders in the subfamily Haniinae is the arrangement of the spinnerets in a single row like the teeth of a comb – thus the common name.

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The arrangement of the spinnerets of ‘comb-tailed’ spiders in the subfamily Hahniinae.         (Photo by Tom Murray, licensed under CC BY-ND-NC 1.0)

I generally think of spiders as being one of two basic types: wanderers or web builders. The wanderers include visually hunting ground dwellers like wolf spiders, whereas web building spiders are sit-and-wait predators that rarely leave their silken snares. This is overly simplistic, of course, but asking “web or not?” is often a useful first step in  classifying spiders. The genus Neoantistea, however, gave me a first encounter with members of an intermediate group known as vagrant web builders.

The sheet webs of Neoantistea spiders are tiny – typically less than 5 cm across. They are built in moss or across shallow depressions such as those formed by the tracks left by animals walking on soft ground. The diminutive spiders (their total body length is less than 5 mm) live under their webs, retreating into crevices in the litter or moss when disturbed.

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Small sheet-web of Neoantistea magna (photo Sean McCann).

What makes these spiders unusual for web builders is that although the web can be a useful aid for catching prey, it is not necessary. Neoantistea magna have reasonably large eyes and can recognize and hunt prey just as easily off of their webs as on them (Engers & Bultman 2006).

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A portrait of a male Neoantistea magna, showing the arrangement of the relatively large eyes (photo Sean McCann).

Although it was easy to identify the spiders we found to genus – the distinctive spinnerets leave no doubt as to the family, and of the North American members of the Hahniinae, Neoantistea is the only genus of web builders – determining the species was another matter entirely. Usually spider identification relies on close examination of the genitalia.

To ID this handsome fellow, two of the key features were the tibial apophysis and the patellar spur, tiny protrusions of the pedipalps which are very difficult to see without a microscope (here’s a diagram of the segments of the pedipalps).

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The key to identifying spiders often lies in the features of their elaborate genitalia. Here the large curved outgrowth on the tibia and the hooked spur on the patella of the pedipalp are circled (photo Sean McCann).

Speaking of genitalia, although very little is known about the biology of Neoantistea magna, there is one report of mating behaviour (Gardner & Bultman 2006). During copulation, the male clasps the female with his first two pairs of legs. The robust femur and tibia (see leg segment diagram) on each of these legs are studded with a double row of tubercles, giving them a serrated look. 

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Male N. magna. Note the burly front legs, presumably modified for grasping the female (photo Sean McCann).

Although the female may attempt to disengage from her partner, he is able to maintain a firm hold with his rather spectacularly modified legs and continue copulation.

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Female N. magna, with slender front legs (photo Sean McCann).

Fun with etymology:                                                                                                             The genus name Neoantistea means “new Antistea”. Antistea comes from the Latin word antistes, which means “one who stands in front of a temple, overseer, high priest”. Why were these tiny spiders given such a grandiose name? It’s a mystery.

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References

Engers, W., & Bultman, T. (2006). Foraging Habits of Neoantistea magna (Araneae: Hahniidae).

Gardner, D., & Bultman, T. (2006). Natural History and Reproductive Biology of a Hahniid Spider in Southwestern Michigan.

Opell, B. D., & Beatty, J. A. (1976). Nearctic Hahniidae (Arachnida: Araneae)Bull Mus Comp Zool Harvard Univ.

Tetragnatha revisited: dinner and romance at sunset

This post features photographs by Sean McCann. For more beautiful photography and natural history of arthropods and other wildlife, check out his blog, Ibycter.com

As a sequel to our recent encounter with some long jawed orb-weavers in the genus Tetragnatha (the tiny and cryptic Tetragnatha caudata), this week on an evening walk at Iona Beach, Sean and I observed some neat predation and mating behaviour in another species, most likely Tetragnatha laboriosa.

We made our first observation just as the sun was beginning to set, the beginning of the most active hunting hours for Tetragnatha laboriosa. This female had just captured her first meal of the evening, a bug in the family Miridae.IMG_1953

After biting it, she began wrapping it with silk, which she pulled out of her spinnerets with her last pair of legs (you can see her caught in the act below). IMG_1956

After wrapping the bug lightly with silk, she carried it back to the hub of her orb web and settled down to dine.IMG_1962

Unfortunately for the spider, dinner was interrupted by Sean’s efforts to get a good photograph. The disturbance prompted her to drop her meal and retreat to the vegetation at the edge of her web. Isn’t she just gorgeous?!
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After a minute or so, she went back for her abandoned prey.
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She then carried it off the web to resume her meal in peace. You can see from this image how the lovely coloration of these spiders allows them to blend in with plant stems when they adopt their cryptic stick-like posture.
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Later, when the sun had all but set and we were just about to head home, Sean spotted a pair of spiders (probably the same species, T. laboriosa) mating in a female’s web.
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Mating involves a fair bit of contortion for long jawed orb-weavers. Below you can see the male’s extremely long pedipalp (one of a pair of appendages modified for transferring sperm) engaged with the female’s epigyne (genital opening). The male’s short third pair of legs is used to position his partner’s abdomen. Throughout copulation he maintains a firm grip on the female’s jaws with his own.  IMG_2106

Here is a closer look at the mating position, where if you look closely you can see one of the female’s fangs interlocking with the special tooth on the male’s corresponding chelicera.jaws_clasping

Here is a drawing by B. J. Kaston of what the cheliceral embrace looks like close-up. The male, with larger jaws, is below, and the female above.

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Fig. 876 from Kaston 1948. Interlocking jaws of Tetragnatha pallescens (which looks very similar to T. laboriosa) during mating.

The female’s fangs get locked in underneath the special large tooth that protrudes from each of the male’s chelicerae.  tooth_landscape

As if we hadn’t had enough excitement already with the chance to closely witness such an intimate encounter, moments later we spotted two additional males waiting in the periphery of the female’s web. We were in for quite a show!

Here is one of the males that was waiting in the wings, posing elegantly and displaying his long jaws and even longer pedipalps. We’ll call him bachelor #2. IMG_2120

Not long after we spotted them, one of the lurking males made his move, lunging at the mating pair with his jaws held wide.  IMG_2108

A bit of a tussle ensued, after which the mating spiders disengaged. The attacking male pursued the mated male off the web and all the way to the substrate below. The female, apparently rather perturbed by this rude interruption, also left the web. One of the two rival males, apparently dominant, soon ascended back toward the web via his dragline. IMG_2112

Just as the winner of the first brief battle returned to the web, the third male entered the ring, and a second chase ensued. This cycle repeated a couple of times, until at last only one male returned victorious to the periphery of the web.IMG_2129

Bachelor # 2 (or was it #3?) settled down to wait at the edge of the web, while the female made her way back to the hub.     IMG_2140

It turns out that female T. laboriosa only mate once as a rule, and if copulation is interrupted as we observed, it’s a toss-up whether or not she will be willing to pick up where she left off (LeSar & Unzicker 1978). We couldn’t stay to see if our champion was able to successfully mate, but we wished him the best of luck!IMG_2138

Long, cryptic spiders

Iona beach, near the Vancouver International Airport in Richmond, BC, is fantastic place to go looking for spiders and other arthropods, and there always seems to be something new to discover on the dunes. So it was an unusual situation when last Friday evening, just before the park gates were about to close, Sean was lamenting not having found anything very interesting to photograph. We decided to spend our last minutes on the beach in an effort to turn up some sleeping hymenopterans in the vegetation bordering a walking path. After closely inspecting a couple of plants, instead of a sleeping bee or wasp, I found what at first glance looked like a tiny twig hanging in the middle of a spider’s orb web. Meet Tetragnatha caudata, the longest, most cryptic orb-weaver I have ever met!

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This slender female was not easy to spot among the grasses, even though she was hanging right in the middle of her web. Her body is only about 1cm long from tip to tail.

Tetragnathidae is the family of the longjawed orb-weavers. You can see where the name comes from in the photo of a male below. During mating, the male and female interlock their large chelicerae in toothy embrace. This is where the male’s exceptionally long pedipalps come in handy, allowing him to reach the female’s epigynum while maintaining his hold on his partner’s fangs.

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You can see this male’s large, powerful jaws sticking out to the sides underneath his very long, slender pedipalps.

Spiders in the genus Tetragnatha are sometimes called ‘stretch spiders’ because of their elongated bodies. They often rest with their first two pairs of legs stretched out in front. They can be very cryptic in this posture, especially if they cling to twigs or stems that are similar in colour to their bodies. Although they normally use their webs for hunting, sometime Tetragnatha will also snap up unsuspecting prey when lying low like this on vegetation.

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A male Tetragnatha in camouflage mode. This works better on dry grasses or twigs, I would think. But still pretty stealthy here.

Most species in the genus Tetragnatha are difficult to tell apart, but I got lucky with this one (it keys out in the second couplet of the key to the Canadian species in this genus). Tetragnatha caudata is so named because of its strange looking abdomen (caudata means ‘tailed’ in Latin). While I would expect the spinnerets to sit at the tip of a spider’s abdomen, in this species there is a slight kink where the spinnerets sit on the underside of the abdomen, and a pointy tail sticks out behind them.

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If you look closely you can see a strand of silk coming out of this female’s spinnerets, located on the underside of her abdomen just before it bends into a pointy tail.

The orb webs of Tetragnatha are usually horizontal or angled (unlike the vertical webs of spiders in the more familiar orbweaver family Araneidae) and have a hole in the centre where the spider sits waiting for prey.

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We found several of these tiny Tetragnatha caudata among tall grasses, but larger members of the genus typically build their horizontal orbwebs over water, allowing them to capture insects such as mosquitoes as they emerge.

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So let this be a lesson: next time you’re out looking for wildlife, stop and take a careful look in some seemingly boring vegetation! You never know what wonders you might turn up.

Thanks to Sean, as usual, for the great photos!

References:
Adams, R.J. (2014) Field Guide to the Spiders of California and the Pacific Coast States. University of California Press, Berkeley and Los Angeles.

Bradley, R. (2013) Common Spiders of North America. University of California Press, University of California Press, Berkeley and Los Angeles.

Dondale, C. D., Redner, J. H., Paquin, P., & Levi, H. W. (2003). The Insects and Arachnids of Canada. Part 23. The Orb-weaving Spiders of Canada and Alaska (Araneae: Uloboridae, Tetragnathidae, Araneidae, Theridiosomatidae)Ottawa, NRC Research Press.

The mystery of the burrow-dwelling sand dune spider

On a walk at Iona Beach a couple of weeks ago with Sean (who kindly provided all the photos that follow) I came upon a small hole in the sand, and after poking at it a bit, realized it was a silk-lined burrow with what looked like a trap door. We tried to figure out who lived there, but didn’t find anyone inside.

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The mystery burrow where this natural history adventure all began!

I went home puzzling about the mysterious burrow-dweller, and emailed our local spider expert Robb Bennett to see if he had any idea what kind of spider the owner might be. His response that it was probably a wolf spider burrow came as a surprise to me. I don’t know a lot about wolf spiders (family Lycosidae), and although I have occasionally seen them hunkered down in shallow depressions under stones and logs, I wasn’t aware that many species build quite elaborate silk-lined burrows. As it turns out, some lycosids overwinter or oviposit in burrows, or hunt by waiting just inside the burrow entrance for prey. One genus, Geolycosa, spends almost its entire life underground, in burrows up to 17cm deep (Dondale & Redner 1990)!

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Before this mystery began, I had only seen lycosids associated with shallow burrows, like this one we found on Mt. Tolmie.

A few days later, we returned to the beach to look for more spiders and other arthropods. We encountered several jumping spiders, including this one.

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 A Sitticus male?

After a while, Sean spotted this beautiful and well-camouflaged male wolf spider.

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After being disturbed, this spider alternated between lightning-fast sprints and freezing with its legs splayed out against the sand. It takes full advantage of its banded legs and mottled body coloration, which allow it to all but disappear against the background!

Next we encountered two females of the same species carrying their spiderlings on their abdomens. Lycosid females have special abdominal hairs onto which the inner layer of spiderlings cling (Rovner et al. 1973).

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The second female we found had a smaller brood of larger spiderlings hanging on mainly to the underside of her abdomen.

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Here’s a portrait of the same female as the previous photo, this time showing the characteristic lycosid eye arrangement.

A little while later, much to my excitement, I came across another silk-lined burrow!

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Another mystery burrow, very similar to the original!

Using a bit of dry grass to scratch at the sand-covered silk surrounding the burrow entrance, I was able to entice the resident spider towards the opening. And just like that, the mystery was solved! The burrow belonged to a female of the same wolf spider species we had been running into all evening as we explored the beach.

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The spider soon approached the entrance of her burrow to investigate the source of the disturbance.

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After some more serious disturbance resulting in the opening of the burrow now looking a lot more like the (also disturbed) entrance of the original mystery burrow, the spider was persuaded to come out entirely.

After a successful evening of sleuthing, we went home and identified our cryptic burrow-dwelling spider as Arctosa perita. This species is typically found on sand dunes or sandy heathland, and only the females construct silk-lined burrows (Dondale & Redner 1990). It is introduced to North America, and is apparently only present in certain areas of southern British Columbia. The similarly coloured beach wolf spider Arctosa littoralis (beautifully photographed by Ted MacRae here) is native to North America.

The burrow entrance can be cinched up and made effectively invisible to humans, but this does not prevent some predators from detecting the spiders inside. In Britain, Arctosa perita is the preferred prey of the spider wasp Pompilus plumbeus, which uses a combination of smell and touch to locate the spiders within their burrows (Bristowe 1948). This spider (and other lycosids) can detect polarized light, and this species is somewhat famous for being able to navigate using the sun or the moon (references in Dondale & Redner 1983).

Arctosa perita doesn’t seem to have a well established common name, but I found it referred to as the ‘sand bear-spider’ by Steven Falk on Flickr (check out his lovely photo set), which I quite like. The genus name Arctosa is based on the Greek word for bear, ἄρκτος (arctos). The species name ‘perita‘ means ‘mountain dweller’ in Greek which doesn’t make a whole lot of sense unless you consider the sand dunes that are apparently the preferred habitat of this species to be very small mountains. I will now always think of it as the mysterious burrow-dwelling tiny sandy mountain bear-spider! Catchy, right?

Bonus fun fact about wolf spider names:

The type genus is Lycosa, which of course means ‘wolf’. Several other genera were subsequently named to rhyme with Lycosa, with the names based on other carnivorous animals: Alopecosa (fox), Crocodilosa, Dingosa, Hyaenosa, Lynxosa, Mustelicosa (weasel), and Pardosa (leopard).

References:

Bristowe, W. S. (1948). NOTES ON THE HABITS AND PREY OF TWENTY SPECIES OF BRITISH HUNTING WASPS. Proceedings of the Linnean Society of London, 160: 12–37. doi: 10.1111/j.1095-8312.1948.tb00502.x

Dondale, C. D., & Redner, J. H. (1983). Revision of the wolf spiders of the genus Arctosa CL Koch in North and Central America (Araneae: Lycosidae)Journal of Arachnology, 11: 130.

Dondale, C. D., & Redner, J. H. (1990). The insects and arachnids of Canada. Part 17. The wolf spiders, nurseryweb spiders, and lynx spiders of Canada and Alaska. Araneae: Lycosidae, Pisauridae, and Oxyopidae. Publication-Agriculture Canada (English ; 1856).

Rovner, J. S., Higashi, G. A., & Foelix, R. F. (1973). Maternal behavior in wolf spiders: the role of abdominal hairs. Science, 182: 11531155. doi: 10.1126/science.182.4117.1153