33

Bull. Br. arachnol. Soc. (1992) 9 (2), 33-37

A review of the ethology of jumping spiders (Araneae,
Salticidae)
David B. Richman
Department of Entomology, Plant Pathology and Weed Science,
New Mexico State University,
Las Cruces, New Mexico 88003, USAandRobert R. Jackson
Department of Zoology,
University of Canterbury,
Christchurch, New ZealandIntroduction

Ethology is the study of behaviour. Jumping spiders(Salticidae), because of their complex behaviour andunique, complex eyes, have been (among spiders) disproportionately well-represented in ethological studies. We
will review five areas of salticid ethology: predatory
strategies, courtship, maternal behaviour, silk utilisationand mimicry. First, however, it will be useful to present
some background information on the characteristic thatmakes salticids so unique among spiders.
Most spiders have simple eyes and poorly developed
vision, but salticids have acute vision (Drees, 1952). The
unusual and highly evolved visual system of salticids,
extensively studied by neurobiologists (Forster, 1982a;
Blest, 1985; Land, 1985), is probably the most importantcharacteristic of this family. Because no other spiderfamily has comparable eyes, the evolutionary relationshipof salticids to other spider families is unclear.
Our understanding of salticid eyes is derived, to a large
degree, from the ground-breaking work of Homann(1928, 1971) and Land (1969a,b), and major extensionsof this work from David Blest's laboratory (e.g. Blest &Carter, 1987). The salticid antero-median (AM), or principal, eyes are responsible for acute vision. Three pairs of
secondary eyes, the antero-lateral (AL), postero-median(PM) and postero-lateral (PL) eyes, are primarily movement detectors. In most salticids, the PM eyes are minuteand vestigial, but these eyes are large and functional in a
few primitive species. It is the AM eyes, however, that areespecially unusual.
All eight eyes have fixed cuticular lenses, but the AM
eye tubes are long and have six pairs of muscles attachedto them. The salticid uses these muscles to co-ordinate
precise and complex rotational and side-to-side movements of the eye tube, important in tracking moving preyand in recognising shapes. However, the AM eye tube is
not capable of accommodation; it cannot move forwards
and backwards to change focal length.
The structure of the AM eye retina also distinguishes
these eyes. The retina is tiered: light coming from the lenssystem passes successively through layers 4, 3, 2 then 1.
The tiered structure is crucial in enabling the salticid tocompensate for chromatic aberration. In essence, the AM
eye is a miniature telephoto system, much like a Galilean
telescope. First, because the retina is at the rear of along eye tube, the AM eye has a long focal length. Moreremarkably, there is a second (diffracting) lens at the rearof the eye tube, just in front of the retina, which increases
the magnifying power of the lens (Williams & Mclntyre,1980). The AM lens system and retina combine to givesalticid eyes extraordinary resolving power and allow thesalticid to perform phenomenal feats of visual discrimination (Jackson & Blest, 1982a). The resolving power ofsalticid eyes has no doubt been an important factor in
the evolution of complex visually mediated behaviour,
including prey capture, agonistic display and visual
courtship.
The AM eye retina also enables salticids to perceive
colour, with sensitivity ranging from red to ultraviolet(Peaslee & Wilson, 1989). The ability to see colour probably was important in the evolution of one of the mostdistinctive characteristics of salticids as a group: these
spiders are often outlandishly colourful. The salticid's
colourful markings are especially evident during courtship, suggesting that colour is important in intraspecificcommunication in this family.
Predatory strategies

Jumping spiders are, for the most part, diurnal hunters,
which is not surprising for a group with such a highly
developed visual system. Forster (1982b) analysed indetail how visually hunting salticids usually catch theirprey. Typically, the salticid first orients by swivelling its
cephalothorax around to bring the AM eyes to bear on the
prey. Next, the salticid aligns its abdomen with its cephalothorax and begins to stalk slowly, almost cat-like, towards
the prey. When close to the prey, the salticid pauses, lowers
its body and fastens a dragline to the substrate, then leapson the prey.
Salticids resemble web-building spiders in being able
to detect and precisely locate prey at a distance (Enders,
1975). Web-builders do this by accurately interpretingvibrations coming across the web from the ensnared prey.
Salticids, in contrast to the web-builders, do not depend
on a silk structure erected in the environment. Salticids,
instead, use sight to gain accurate information about
potential prey.
Remarkably, a salticid.does not have to take a straightline path to reach its prey. It can take a circuitous route(detour) to its prey, sometimes temporarily losing sight ofthe prey (Hill, 1979). Detouring, suggesting remarkableproblem-solving abilities for a spider, is currently a
topic of intensive research by Michael Tarsitano at the
University of Canterbury.
Some salticids are apparently active searchers (Enders,
1975), spending much of their time walking and stopping
periodically to look around. If they come near prey, they
actively pursue it. Other species spend more time visuallyscanning the environment from one place, then actively
stalking prey that comes near. Phaeacius, a Sri Lankan
salticid that sits motionless while facing downwards
on tree trunks, has taken sit-and-wait predation to an
extreme. This salticid rarely stalks its prey; instead, it
prefers simply to lunge down on prey that walks in front ofit (Jackson & Hallas, 1986a).

34

Ant-eating salticids also use unusual and specialised
prey-capture techniques. Most salticids exclude worker
ants from their diets, but Corythalia canosa (Walckenaer),
Habrocestum pulex (Hentz) and Pystira orbiculata(Keyserling) routinely eat ants [Edwards et al., 1974;
Cutler, 1980; Jackson & van Olphen (1991)]. Ant-eating
salticids have prey-specific predatory behaviour for catching ants. Corythalia canosa, for example, circles around in
front of the ant, then lunges over the insect to grab it
behind the head. Ant-eating salticids, however, do notappear to be limited to eating ants. For example, they arealso efficient predators of flies, which they catch using thestandard predatory behaviour outlined by Forster(1982b), and attack from any direction.
Web-invasion is another unusual predatory tactic ofcertain salticids. Some salticids are known to leap into,or walk across, webs to catch web-building spiders(araneophagy) or insects (kleptoparasitism) ensnared in
the webs (e.g. Robinson & Valerio, 1977). Phidippus
audax (Hentz) has been observed attacking Tgtragnatha
orb-weavers in this manner (Richman, unpubl. observation). Four genera, Brettus, Cyrba, Gelotia and Portia,
however, are more specialised web-invaders (Jackson &Blest, 1982b; Jackson & Hallas, 1986a,b; Jackson, 1990band in press). These salticids go slowly on to the alien web
and manipulate silk with their legs and palps, making
vibratory signals. The web-owner spider sometimes
responds to these signals by approaching as if the signal
came from a small ensnared insect. Instead of getting ameal, the web spider becomes the aggressive mimic's meal.
As with the ant-eating species, web-invading salticids do
not limit predation to one type of prey. The web-invading
salticids are also efficient at using other predatory tactics
against other types of prey. For example, Portia fimbriata(Doleschall) from Queensland, Australia, has a specialised way of stalking and catching other (ordinary) salticids, which involves a slow, mechanical, stalking motion(Jackson & Blest, 1982b). The behaviour of web-invaders,like that of ant-eaters, suggests, in the Salticidae, thenotion that the jack-of-all-trades is the master of none
does not apply (Jackson & Hallas, 1986c).
Courtship

Courtship is another context in which salticid behaviour
tends to be complex. Peckham & Peckham (1889, 1890)
were the first to make a systematic study of salticid courtship, emphasising the importance of intersexual selection
as a factor in the evolution of salticid behaviour. During
the Peckhams' time, and for most of this century as well,intersexual selection was not well understood; yet intersexual selection is now one of the most important topics in
ethology. Recent studies (e.g. Jackson, 1981) support the
Peckhams' contention that intersexual selection has
shaped the evolution of salticid courtship by forcing
males to overcome female resistance by performing
more complex dances and by being more strikingly
ornamented.
Researchers are also interested in using information
about courtship to understand salticid phytogeny andclassification. Crane (1949), in a well-known and exten-

Ethology of jumping spiders

sive comparative study of salticid display, examined
broad phylogenetic trends. She did not, however, proposea phylogenetic classification system based on behaviour.
She divided jumping spiders into three broad behaviouralgroups: runners, hoppers and intermediates. These groups
often crossed phylogenetic lines and were referred to by
her as stages. More recently, Richman (1982) showed thatsalticid displays, if used with care, are useful characters insystematics.
Individual salticid displays are often elaborate. For
example, Habronattus hallani (Richman) males run sideways, then jump at the end of the run, displaying theiriridescent femora; Habronattus coecatus (Hentz) males
creep slowly towards the female with front legs raised,suddenly raising then lowering their third pair of legs fromtime to time as they advance. Hentzia palmarum (Hentz)
males approach the female in a zigzag path while tilting
their abdomen to the side and spreading their chelicerae
apart (Richman, 1982). Each salticid species tends to haverepertoires of numerous discrete displays, this being
another source of complexity. For example, Corythalia
canosa has a repertoire of about 30 major displays thatcan be combined in diverse ways (Richman, 1982; Jackson& MacNab, 1989). There also appears to be considerable
variation between species, not only in the details of eachspecies' display repertoire, but also in repertoire size.
For example, Platycryptus undatus (DeGeer), an exceptionally cryptic North American salticid, appears to havea considerably smaller repertoire of displays than does
C. canosa (Richman, 1982).
All salticids use visual displays during courtship (i.e.
special behaviours that function to provide a special
visually perceived stimulus to a conspecific of the oppositesex). This is not surprising in a spider family with such
acute vision. However, acute vision has not limited salticids to visual communication. On the contrary, multichannel communication is common in the Salticidae, andthis is an important factor contributing to the complexityof salticid display.
Many salticids perform tactile displays when the maleand female come into contact with each other before
mating, the repertoire of these displays sometimes being
large and complex (e.g. Jackson & Harding, 1982).
Several salticids communicate by stridulating (Edwards,
1981; Gwynne & Dadour, 1985; Maddison & Stratton,
1988a,b). Also, pheromones are known to be importantfor many species (Crane, 1949; Richman, 1982; Pollard
et al., 1987; Jackson, 1987). Besides volatile pheromones
perceived by olfaction, salticids commonly use pheromones associated with silk and perceived by contact
chemoreception.
There is yet another common mode of communicationin the Salticidae: vibratory signals transmitted across silk(e.g. Jackson & MacNab, 1989). Although web-building
spiders from other families are well known for communicating with vibratory signals on silk (Robinson, 1982),
silk-borne signals are surprising in the salticids, hunting
spiders with acute vision. Yet salticids, like all spiders, use
silk, even if they do not normally build webs. In particular, most salticids spin a cocoon-like nest as a resting,moulting and oviposition site. The male of many salticid

D. B. Richman & R. R. Jackson

species performs vibratory courtship on the silk of thefemale's nest and, if successful, enters the nest to matewith the female. The male does not have to be able to see
to perform this type of courtship (Jackson, 1977). Indeed,nests are often built in places with little light, such as
under rocks.
The same individual that courts with vibratory displays
adult females he finds in nests will use visual displays if hefinds an adult female outside her nest. He uses yet anothertactic if the female he finds inside her nest is subadult: firsthe courts her using vibratory displays, then he spins asecond chamber on her nest and cohabits until he canmate with her in the nest after she matures (Jackson,1982).
There is an interesting parallel between predatory
and courtship behaviour in the Salticidae. In both cases,
salticids often have conditional strategies (i.e they use
discretely different tactics in different circumstances). In
the conditional mating strategies, different tactics areused depending on the location and maturity (or evenreceptivity) of the female; in the conditional predatory
strategies, different tactics are used depending on the typeof prey. The study of conditional strategies is an important current area of research in zoology (e.g. Dominey,1984), and salticids seem to be a group in which conditional strategies are especially well developed (Jackson&Wilcox, 1990).
Closely related to courtship displays are male-male
confrontations or agonistic displays. These visual displays are similar to courtship, but have added aggressive aspects that make them distinct (e.g. Crane, 1949;
Richman, 1982; Jackson & MacNab, 1989). We lackspace for a thorough discussion of male-male aggressive
displays, which do not appear to occur in all salticid
species. These are apparently lacking, for example, inmost of the species of Habronattus, but occur in suchvaried genera as Corythalia, Phidippus, Hentzia and
Menemerus (see Richman, 1981 for a review of theliterature to that date).
Maternal behaviour

Courtship and mating are followed by oviposition,and maternal care of eggs and recently hatched juveniles appears to be widespread, if not universal, in the
Salticidae. Salticids spin silken eggsacs and stay withthese, presumably guarding them. Exactly what salticid
females guard against and how they guard the eggs is
unclear (Eberhard, 1974), but the maternal female could
probably deter many egg predators and parasitoids.
The most common type of salticid eggsac is, more or
less, a spherical mass of eggs wrapped in dense silk andembedded in the interior of a cocoon-like nest. However,other designs are known. For example, lyssomanine salticids stand in the open on leaves (usually on the underside) over eggs that are only loosely covered with silk,and the eggs are usually widely spaced under the silk
(Richman & Whitcomb, 1981; Jackson, 1990a).
A European species, Marpissa rwnpfi (Scopoli) [ = M.
muscosa (Clerck)], stacks three to five flat eggsacs one on
top of the other inside a nest (Holm, 1940). Holcolaetis

35

stands in the open over a flat papery eggsac on tree trunks(Wanless, 1985). Euryattus chooses a dead, rolled-up leaf
as an oviposition site and hangs the leaf up on a tree trunkor a rock face by several heavy guylines (Jackson, 1985a).
Pellenes nigrociliatus (L. Koch) does something similar,
only this species chooses a snail shell instead of a leaf
(Mikulska, 1961). Thiania seals a pair of green leaves
together with strong silk rivets and oviposits in the tightspace between the leaves (Jackson, 1986).
Silk utilisation

Besides using silk to make nests and eggsacs, all salticids trail a dragline behind them when they walk. If thesalticid loses its footing in the vegetation, it can return toits starting point by climbing up the line. As an alternative
to a cocoon-like nest, some salticids rest at night hanging from draglines (Carroll, 1977) and some drop on
draglines when feedtng^Robinson & Valerio, 1977).
Salticids also use silk for dispersal by using a behaviour
known as ballooning. While standing at the edge of vegetation or hanging from a dragline (Eberhard, 1987), thesalticid is carried away on the wind when air currents
catch hold of strands of silk from the spinnerets.
Some salticids even build pre-catching webs. Pellenes
arcigerus (Walckenaer) (Lopez, 1986) and Spartaeus(Jackson & Pollard, 1990) build sheet webs. Spartaeus
primarily catches moths in its web, but the predatory
behaviour of Pellenes arcigerus has not been studied.
Portia, besides invading alien webs (as noted earlier), alsobuilds an unusual funnel-shaped aerial web, and uses thisweb to catch other spiders (Jackson & Hallas, 1986b).
Euryattus, besides hanging up a dead leaf as an oviposition site (see above), also spins a flimsy space web;primarily the juveniles build these webs (Jackson, 1985a).
Simaetha spins a dense space web, often embedding a
cocoon-like nest within the web, or it may just build a nest
(Jackson, 1985b). Simaetha has another unusual habit;
it often builds its web within the communal webs of
Badumna Candida (L. Koch), a social amaurobiid inAustralia, and gleans trapped insects off the sticky webof these social spiders.
Web-building in the Salticidae is unexpected. The salticids are a group renowned for their well-developed eyes
and generally thought of as a classic example of hunting
spiders. Perhaps web-building in salticids is a relic of
the evolution of this family from web-building ancestors(Jackson, 1985c).
Mimicry

We have reviewed one type of mimicry already —aggressive mimicry in which a predator tricks its prey bymaking deceptive signals on a web. Batesian mimics areprey species that deceive their predators, and this type of
mimicry may be common in salticids. Ant mimicry is thebest known (Edmunds, 1978; Reiskind, 1977).
Many salticids are remarkably similar to ants in morphology and in locomotory behaviour. Ants are avoided
by many predators of spiders, suggesting that the ant-likesalticids are Batesian mimics (Edmunds, 1974). Data are36

scarce to support this supposition, but Engelhardt (1971)
showed that birds fail to distinguish between ants and an
ant-like salticid, Synageles venator (Lucas).
Besides ant-like salticids, there are also wasp-like andbeetle-like species. Phidippus apacheanus Chamberlin &Gertsch, for example, is remarkably similar to velvet ants(mutillid wasps) in both general appearance and locomotory behaviour (Edwards, 1984; Richman, unpubl. obs.).
Agassa cyanea (Hentz) resembles flea beetles and has
been collected in the company of flea beetles (Richman,
unpubl. obs.). Numerous salticids, including species inat least four genera (Agassa, Sassacus, Cylistella andCoccorchestes) resemble chrysomelid beetles.
Batesian mimicry is a reasonable hypothesis to accountfor the wasp mimicry because wasps have potent stings.
The advantage of beetle mimicry is not as clear, but thehard exoskeleton and the prevalence of noxious tastes insome beetle species lends credence to the notion that
Batesian mimicry has been important in the evolution of
beetle-like Salticidae.
Orsima formica Peckham & Peckham from Borneo
is apparently a "generalized, reverse insect mimic"(Reiskind, 1976). This salticid waves its abdomen andspinnerets in a fashion to resemble the head of an ant or
beetle. That is, this salticid uses its rear end to mimic thefront end of an insect.
There is another way the tables can be turned; sometimes the insect tricks the salticid. Certain fruit flies havemarkings on their wings that resemble a salticid performing a threat display. A salticid that sees the fly's wing
displays back, as if it had seen a rival, giving itself away tothe fly (Whitman et al., 1988). An even more remarkablemimic of this type is a recently discovered fulgorid planthopper nymph that bears a close resemblance to a jumping spider when viewed from the rear (G. Zolnerowich,
pers. comm.). These are unusual instances in which preyspecies mimic a predator to gain protection from the same
predator mimicked.
Concluding remarks

This review has been, by necessity, selective both in
the topics we covered and the examples we used for each
topic, but the interested reader can find other examples
and topics in the references we have cited (see also
Richman, 1981).
Salticids have a combination of traits that make themunique subjects for ethological study. These traits include,
among other things, the specialised salticid visual system,
the complexity of salticid behaviour, a tendency of these
spiders to evolve conditional strategies and interspecificdiversity in behaviour. Although much research has been
done already, our impression is that the potential ofthese animals for ethological research is not as widely
appreciated as it should be.
Acknowledgements

This is Journal Article 1579, New Mexico Agricultural
Experiment Station, Las Cruces, New Mexico.

Ethology of jumping spiders

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