Spiders in the family Salticidae are commonly known as "jumping spiders", for the impressive springing movements often exhibited when they are alarmed. This is the largest family of spiders, including more than 5,000 known species. They are common on all continents except Antatctica, occurring in a range of microhabitats from beneath leaf litter up into the forest canopy. Tropical regions have especially high species diversity (in contrast, only around 300 species, for example, are known from North America north of Mexico).
Although most jumping spiders are easily recognized as salticids, the appearance of these highly visually oriented spiders is nevertheless extremely diverse in both body form and color pattern. Many salticids are apparent mimics of ants or beetles; others may be camouflaged to blend in against a background of bark, grass, or stones. Still other salticids are brightly colored and patterned and males may use these decorations in complex courtship displays. Some salticids (such as many Phiddipus and many, if not all, Habronattus) have stridulatory mechanisms for communication, which are often used in courtship. In many genera, males may engage in ritualized agonistic behavior that may sometimes resemble courtship.
Maternal care of eggs seems to be the rule among salticids, with females closely associated with both eggs and newly emerged spiderlings until the spiderlings molt to the first free-living instar and disperse. Most salticids lay their eggs in a distinct sac, but females of some species (e.g., Lyssomanes) lay only a thin sheet of silk over the eggs.
Predatory behavior has been described as surprisingly reminiscent of that of cats, with some species showing remarkable abilities to locate prey and develop efficient prey captive techniques while minimizing risk of injury to themselves. Richmond and Jackson (1992) provide a review of salticid behavior.
(Richman et al. 2005 and references therein)
An excellent resource for anyone interested in accessing more information about salticids is the web page of Dr. Wayne Maddison at the University of British Columbia.
Astieae Simon, 1901
The group was proposed by Simon (1901), revised by Wanless (1988) and subsequently studied by Gardzińska (1996), Żabka (1991b, 1995, 2001, 2002), Patoleta & Żabka (1999) and Żabka & Pollard (2002). Including the taxa treated here, it comprises 9 genera and 55 valid species: Adoxotoma Simon, 1909 (10 spp.), Arasia Simon, 1901 (3 spp.), Astia L. Koch, 1879 (2 spp.), Helpis Simon, 1901 (7 spp.), Jacksonoides Wanless, 1988 (7 spp.), Megaloastia Żabka, 1995 ( 1 sp.),Sondra Wanless, 1988 (15 spp.), Tauala Wanless, 1988 (8 spp.) and Parahelpisgen. nov. (2 spp.). The diagnoses for particular genera are given by Wanless (1988), two of these are updated here.
Astieae group is currently known from Australia, New Guinea, New Zealand and adjacent islands.
- Gardzińska, Joanna, Żabka, Marek (2010): A new genus and five new species of Astieae (Araneae: Salticidae) from Australia, with remarks on distribution. Zootaxa 2526, 37-53: 38-38, URL:http://www.mapress.com/zootaxa/2010/f/zt02526p053.pdf
Salticidae Blackwall, 1841
Salticidae Blackwall, 1841
The spider family Salticidae (jumping spiders) includes 5678 described species placed in 597 genera (Platnick 2014), far more known species than in any other spider family; at least 315 species are known from North America north of Mexico (Richman and Cutler 1978; Richman et al. 2005). Although Mexico in reality likely has far more species than the United States and Canada, the fauna is far less well studied and fewer than 300 species are known from Mexico (Richman and Cutler 1988; Metzner 2013).
Salticids are generally easy to recognize as such, but Richman et al. (2005) note that they can bear some resemblance to corinnids, oxyopids, and thomisids. Salticids are highly variable in color, including brown, gray, and black, often with patterns of brilliant scales of red, yellow, or metallic or iridescent green, blue, copper, gold, or silver. The general body shape ranges from longer than wide (strikingly so in ant-like species) to as wide as long (in some beetle-like species). The face is relatively flat. The salticid eye pattern is distinctive. The posterior median eyes are small and sometimes difficult to see on the side of the head; the anterior lateral eyes and the posterior lateral eyes, which face sideways or backward, are larger, and the forward-facing anterior median eyes are huge. The posterior eye row is so curved it may appear to be two distinct rows. The chelicerae of salticids range from small to long and can be very stout, projecting, and heavily toothed (especially in males). The spinnerets are short and sometimes not obvious; the anterior (front) and posterior (rear) spinnerets are about the same length. (Richman et al. 2005; Bradley 2013)
Salticids are found in abundance on every continent except Antarctica and occur in a broad range of microhabitats from beneath leaf litter to the forest canopy. The highest diversity of salticids is in the tropics. Some salticids have established populations outside their native range, e.g., Menemerus semilimbatus, a Mediterranean species established in Argentina and California (Manolis and Carmichael 2010 and references therein). Cutler (1990) compiled information on synanthropic salticids in the northeastern United States, some of which are introduced from other continents.
Apparent mimicry (e.g., of ants or beetles or mutillid wasps [Edwards 1984]) and crypsis (e.g., resemblance to tree bark, grass, or stones) is common among salticids in both body form and color patterns and often in behavior. Some salticids are gaudily colored and others have evolved decorations on various body parts. Males are especially ornate and their decorations are often involved in complex courtship behavior. Some species (e.g., some Phidippus and many, if not all, Habronattus) have stridulatory mechanisms for communication that are often used in courtship.
Maternal care of eggs and newly hatched spiderlings appears to be the general rule for salticids. Most salticids lay their eggs in a distinct egg sac, although some (e.g., Lyssomanes) simply cover their eggs with a thin sheet of silk.
From a human observer's perspective, salticid behavior does not seem like that of most other spiders; these spiders seem almost vertebrate-like as they turn to look at us and even follow our movements. Salticids are mainly diurnal, colorful, and active in open areas where they are easily observed. Some species exhibit complex and flexible behavior that they use in hunting prey. Richman and Jackson (1992) reviewed the behavior of salticids.
Most salticid species are probably univoltine (i.e., there is just a single generation per year), with adults present during just a few months of the year, as in Phiddipus johnsoni and Lyssomanes viridis. The number of instars in these species apparently varies from 5 to 11 (Jackson 1978; Richman and Whitcomb 1981).
The complex taxonomic history of the North American salticid fauna was reviewed by Richman et al. (2005). Richman (1978) wrote the first complete key to North American salticid genera north of Mexico after Peckham and Peckham (1909) and Richman and Cutler (1978) published a list of the salticid species then known from the region. Cutler (1979, 1982), Edwards (1980), and Richman (1979, 1980) published corrections to the key and list. Richman et al. (2005) includes numerous citations for generic revisions, species descriptions, and taxonomic and systematic studies of the Nearctic salticid fauna prior to 2005. Edwards and Wolff (1995) published a list of Caribbean salticids. Richman et al. (2012) published an updated list of salticids known from North America, including Mexico, together with maps showing known distributions for each (this paper includes additional citations for significant taxonomic literature and distribution information). Hill and Richman (2009) addressed the evolutionary origins of the salticids. Hill and Edwards (2013) reviewed evidence on the origins of North American salticid lineages. Bodner and Maddison (2012) investigated the historical biogeography of salticid radiations on a global scale. There is an extensive literature on the behavior, ecology, and taxonomy and systematics of salticids from many regions of the world. Much of this information is accessible through several excellent websites. Wayne Maddison has provided a wealth of online information about the Salticidae (Maddison 2011). Heiko Metzner (2013) and Jerzy Prószyński (2013) maintain extraordinarily rich online resources on the salticids of the world.
(Richman et al. 2005; Bradley 2013)
This is the largest family of spiders, with over 5,000 species around the world.
Biogeographic Regions: nearctic (Native ); palearctic (Native ); oriental (Native ); ethiopian (Native ); neotropical (Native ); australian (Native )
These spiders are very distinctive. They vary a lot in size (3-17mm), and color, but the arrangement of their eyes will always give them away. They have three or four rows of eyes. The front row look forward, and the middle two eyes in that row are much larger than the others.
All spiders have two body-segments, a cephalothorax in front and an abdomen behind. They have eight legs, all attached to the cephalothorax. On the front they have two small "mini-legs" called palps. These are used to grab prey, and in mating, and are much bigger in male spiders than in females. All spiders have fangs that they use to bite their prey with, like most spiders, jumping spiders have venom glands that produce toxic chemicals that help paralyze and digest their food.
Female spiders are often much bigger than males.
Other Physical Features: bilateral symmetry ; polymorphic
Sexual Dimorphism: female larger; male more colorful
These spiders roam in a wide variety of habitats. Pretty much anywhere they can find prey, they will live.
Habitat Regions: temperate ; tropical ; terrestrial
Terrestrial Biomes: taiga ; desert or dune ; chaparral ; forest ; rainforest ; scrub forest ; mountains
Wetlands: marsh ; swamp ; bog
These spiders don't use silk to catch prey, they stalk and pounce on Insecta and other invertebrates, using their excellent vision.
This species is very alert, and jumps or runs away if danger threatens. Species in this family will sometimes try to scare away other small predators, waving their front legs and moving their fangs.
- other Araneae
- praying mantids
- small Squamata
- small Amphibia
Known prey organisms
Based on studies in:
This list may not be complete but is based on published studies.
- L. W. Swan, The ecology of the high Himalayas, Sci. Am. 205:68-78, from pp. 76-77 (October 1961).
Life History and Behavior
Communication and Perception
Some species in this group have courtship dances. The males are brightly colored, and they wave their front legs and dance to communicate to a female. She often does a dance in response.
Like all spiders, Jumping Spiders hatch from eggs, and the hatchlings are very similar to the adults, only smaller. As they grow they have to shed their skin all at once, this is called molting. Most Jumping Spiders have to molt 5 or 6 times before they become adults.
Most Jumping Spiders don't live for more than about a year, but some species may live longer, especially in cold climates where they have to be dormant for many months each year.
After mating, female jumping spiders make hiding place of silk in a crevice or rolled-up leaf. There they lay up to several hundred eggs. They protect the eggs with a covering made of many layers of silk. Then they may leave, or stay with the eggs to guard them until they die in the cold of late Fall or Winter.
Key Reproductive Features: iteroparous ; seasonal breeding ; sexual ; fertilization (Internal ); oviparous
Female jumping spiders sometimes guard their eggs, but not always.
Parental Investment: female parental care
Evolution and Systematics
The eight eyes of a jumping spider provide it with excellent vision via two principal eyes used for stereoscopic vision and the other six for a panoramic view around the spider.
"Jumping spiders are said to have the best vision of all invertebrates. They catch their prey by ambush, and the large eyes placed close together in the front of the head give accurate stereoscopic vision for judging distance when pouncing. The remainder of its eight eyes give it almost all-round vision, so it can spot prey moving anywhere near it." (Foy and Oxford Scientific Films 1982:73)
Learn more about this functional adaptation.
- Foy, Sally; Oxford Scientific Films. 1982. The Grand Design: Form and Colour in Animals. Lingfield, Surrey, U.K.: BLA Publishing Limited for J.M.Dent & Sons Ltd, Aldine House, London. 238 p.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:4426
Specimens with Barcodes:4246
Species With Barcodes:635
Relevance to Humans and Ecosystems
Economic Importance for Humans: Positive
Jumping spiders may eat flies and other insects that are pests.
The jumping spider family (Salticidae) contains more than 500 described genera and about 5,000 described species, making it the largest family of spiders with about 13% of all species. Jumping spiders have some of the best vision among arthropods and use it in courtship, hunting, and navigation. Although they normally move unobtrusively and fairly slowly, most species are capable of very agile jumps, notably when hunting, but sometimes in response to sudden threats. Both their book lungs and the tracheal system are well-developed, and they use both systems (bimodal breathing). Jumping spiders are generally recognized by their eye pattern. All jumping spiders have four pairs of eyes with one pair being their particularly large anterior median eyes.
Jumping spiders are among the easiest to distinguish from similar spider families because of the shape of the cephalothorax and their eye patterns. The families closest to Salticidae in general appearance are the Corinnidae (distinguished also by prominent spines on the back four legs), the Oxyopidae (the lynx spiders, distinguished by very prominent spines on all legs), and the Thomisidae (the crab spiders, distinguished by their front four legs, which are very long and powerful). None of these families however, has eyes that resemble those of the Salticidae. Conversely, the legs of jumping spiders are not covered with any very prominent spines. Their front four legs generally are larger than the hind four, but not as dramatically so as those of the crab spiders, nor are they held in the outstretched-arms attitude characteristic of the Thomisidae. In spite of the length of their front legs, Salticidae depend on their rear legs for jumping. The generally larger front legs are used partly to assist in grasping prey, and in some species, the front legs and pedipalps are used in species-recognition signalling.
The jumping spiders, unlike the other families, have faces that are roughly rectangular surfaces perpendicular to their direction of motion. In effect this means that their forward-looking, anterior eyes are on "flat faces", as shown in the photographs. Their eye pattern is the clearest single identifying characteristic. They have eight eyes, as illustrated. Most diagnostic are the front row of four eyes, in which the anterior median pair are more dramatically prominent than any other spider eyes apart from the posterior median eyes of the Deinopidae. There is, however, a radical functional difference between the major (AME) eyes of Salticidae and the major (PME) eyes of the Deinopidae; the large posterior eyes of Deinopidae are adapted mainly to vision in dim light, whereas the large anterior eyes of Salticidae are adapted to detailed, three-dimensional vision for purposes of estimating the range, direction, and nature of potential prey, permitting the spider to direct its attacking leaps with great precision. The anterior lateral eyes, though large, are smaller than the AME and provide a wider forward field of vision.
The rear row of four eyes may be described as strongly bent, or as being rearranged into two rows, with two large posterior lateral eyes furthest back. They serve for lateral vision. The posterior median eyes also have been shifted out laterally, almost as far as the posterior lateral eyes. They are usually much smaller than the posterior lateral eyes and there is doubt about whether they are at all functional in many species.
Jumping spiders range in size from a body length of 1 to 22 mm.
In addition to using their silk for safety lines while jumping, they also build silken "pup tents", where they shelter from bad weather and sleep at night. They molt within these shelters, build and store egg cases within them, and also spend the winter in them.
Jumping spiders live in a variety of habitats. Tropical forests harbor the most species, but they are also found in temperate forests, scrub lands, deserts, intertidal zones, and mountainous regions. Euophrys omnisuperstes is the species reported to have been collected at the highest elevation, on the slopes of Mount Everest.
Jumping spiders have four pairs of eyes; three secondary pairs that are fixed and a principal pair that is movable.
The posterior lateral eyes (PLE) are wide-angle motion detectors which sense motions from the side and behind. Combined with the other eyes, it gives the spider a near 360-degree view of the world.
The anterior lateral eyes (ALE) have the best visual acuity and are the most complex of the secondary eyes. It has been shown that they are able to distinguish some details as well, and without them no "looming response" will be triggered by motion. Even with all the other pairs covered, jumping spiders in a study could still detect, stalk and attack flies, using the anterior lateral eyes only, which are also sufficiently widely spaced to provide stereoscopic vision.
The anterior median eyes (AME) have very good vision. This pair of eyes are built like a telescopic tube with a corneal lens in the front and a second lens in the back that focus images onto a 4-layered retina, a narrow boomerang-shaped strip oriented vertically. Physiological experiments have shown they may have up to four different kinds of receptor cells, with different absorption spectra, giving them the possibility of up to tetrachromatic color vision, with sensitivity extending into the ultraviolet range. As the eyes are too close together to allow depth perception, and the animals don't make use of motion parallax, they have evolved a method called image defocus instead. Of the four photoreceptor layers in the retina, the first two closest to the surface contain ultraviolet-sensitive pigments while the two deepest contain green-sensitive pigments. The incoming green light is only focused on the deepest layer, while the other one receives defocused or fuzzy images. By measuring the amount of defocus from the fuzzy layer, it's possible to calculate the distance to the objects in front of them. It seems that all salticids, regardless of whether they have two, three, or four kinds of color receptors, are highly sensitive to UV light. Some species (for example, Cosmophasis umbratica) are highly dimorphic in the UV spectrum, suggesting a role in sexual signaling (Lim & Li, 2005). Color discrimination has been demonstrated in behavioral experiments.
The principal, anterior median, eyes have high resolution (11 min visual angle), but the field of vision is narrow, from 2 to 5°. The central region of the retina, where acuity is highest, is no more than six or seven receptor rows wide. However, the eye can scan objects off the direct axis of vision. As the lens is attached to the carapace, the eye's scanning movements are restricted to its retina through a complicated pattern of translations and rotations. This dynamic adjustment is a means of compensation for the narrowness of the static field of vision. It is analogous to the way most primates move their eyes to focus images of interest onto the fovea centralis. Such movements within the jumping spider's eyes are visible from outside when the attention of the spider is directed to various targets.
Jumping spiders are generally diurnal, active hunters. Their well-developed internal hydraulic system extends their limbs by altering the pressure of body fluid (hemolymph) within them. This enables the spiders to jump without having large muscular legs like a grasshopper. Most jumping spiders can jump several times the length of their bodies. When a jumping spider is moving from place to place, and especially just before it jumps, it tethers a filament of silk (or 'dragline') to whatever it is standing on to protect itself if the jump should fail. Should it fall, for example if the prey shakes it off, it climbs back up the silk tether. Some species, such as Portia, will actually let themselves down to attack prey such as a web spider apparently secure in the middle of its web. Like many other spiders that leave practically continuous silk trails, jumping spiders impregnate the silk line with pheromones that play a role in social and reproductive communication, and possibly in navigation.
Certain species of jumping spiders have been shown by experiment to be capable of learning, recognizing, and remembering colors, and adapting their hunting behavior accordingly.
The hunting behaviour of the Salticidae is confusingly varied compared to that of most spiders in other families. Salticids hunt diurnally as a rule, which is consistent with their highly developed visual system. When it detects potential prey, a jumping spider typically begins orienting itself by swivelling its cephalothorax to bring the anterior median eyes to bear. It then moves its abdomen into line with its cephalothorax. After that, it might spend some time inspecting the object of its attention and determining whether a camouflaged or doubtful item of prey is promising, before it starts to stalk slowly forward. When close enough, the spider pauses to attach a dragline, then springs onto the prey.
There are, though, many variations on the theme and many surprising aspects. For one thing, salticids do not necessarily follow a straight path in approaching prey. They may follow a circuitous course, sometimes even a course that takes the hunter through regions from which the prey is not visible. Such complex adaptive behaviour is hard to reconcile with an organism that has such a tiny brain, but some jumping spiders, in particular some species of Portia, can negotiate long detours from one bush down to the ground, then up the stem of another bush to capture a prey item on a particular leaf. Such behaviour still is the subject of research.
Some salticid species are continually on the move, stopping periodically to look around for prey, which they then stalk immediately. Others spend more time scanning their surroundings from one position, actively stalking any prey they detect. Members of the genus Phaeacius take that strategy to extremes; they sit on a tree trunk, facing downwards and rarely do any stalking, but simply lunge down on any prey items that pass close before them.
Some Salticidae specialise in particular classes of prey. Ants comprise one such class. Most spiders, including most salticids, avoid worker ants, but several species not only eat them as a primary item in their diets, but also employ specialised attack techniques — Corythalia canosa for example, circles round to the front of the ant and grabs it over the back of its head. Such myrmecophagous species, however, will not necessarily refuse other prey items, and will routinely catch flies and similar prey in the usual salticid fashion, without the special precautions they apply in hunting dangerous prey such as ants. Ants offer the advantages of being plentiful prey items for which there is little competition from other predators, but it remains profitable to catch less hazardous prey when it presents itself.
Some of the most surprising hunting behaviour occurs among the araneophagous Salticidae, and it varies greatly in method. Many of the spider-hunting species quite commonly will attack other spiders, whether fellow salticids or not, in the same way as any other prey, but some kinds resort to web invasion; nonspecialists such as Phidippus audax sometimes attack prey ensnared in webs, basically in acts of kleptoparasitism — sometimes they leap onto and eat the web occupant itself, or simply walk over the web for that purpose.
Salticidae in the genera Brettus, Cyrba, Gelotia, and Portia display more advanced web-invasion behavior. They slowly advance onto the web and vibrate the silk with their pedipalps and legs. In this respect, their behaviour resembles that of the Mimetidae, probably the most specialised of the araneophagous spider families. If the web occupant approaches in the manner appropriate to dealing with ensnared prey, the predator attacks.
The foregoing examples present the Salticidae as textbook examples of active hunters; they would hardly seem likely to build webs other than those used in reproductive activities, and in fact, most species really do not build webs to catch prey. However, exceptions occur, though even those that do build capture webs generally also go hunting like other salticids. Some Portia species, for example, spin capture webs that are functional, though not as impressive as some orb webs of the Araneidae; Portia webs are of an unusual funnel shape and apparently adapted to the capture of other spiders. Spartaeus species, on the other hand, largely capture moths in their webs. In their review of the ethology of Salticidae, Richman and Jackson speculate on whether such web building is a relic of the evolution of this family from web-building ancestors.
In hunting, Salticidae also use their silk for a tether to enable them to reach prey that otherwise would be inaccessible. For example, by advancing towards the prey to less than the jumping distance, then retreating and leaping in an arc at the end of the tether line, many species can leap onto prey on vertical or even on inverted surfaces, which of course in a gravitational field would not be possible without such a tether.
Having made contact with the prey, hunting Salticidae administer a bite to inject rapidly acting venom that gives the victim little time to react. In this respect, they resemble the Mimetidae and Thomisidae, families that ambush prey that often are larger than the predator, and they do so without securing the victim with silk; they accordingly must immobilise it immediately and their venom is adapted accordingly.
Although jumping spiders are generally carnivorous, many species have been known to include nectar in their diets, and one species, Bagheera kiplingi, feeds primarily on plant matter. None is known to feed on seeds or fruit. Extrafloral nectaries on plants, such as the partridge pea, provide jumping spiders with nectar; the plant benefits accordingly when the spiders prey on whatever pests they find.
Jumping spiders use their vision in complex visual courtship displays. Males are often quite different in appearance from females, and may have plumose hairs, colored or iridescent hairs, front leg fringes, structures on other legs, and other, often bizarre, modifications. These are used in visual courtship in which the colored or iridescent parts of the body are displayed and complex sideling, vibrational, or zigzag movements are performed in a courtship "dance". If the female is receptive to the male, she will assume a passive, crouching position. In some species, the female may also vibrate her palps or abdomen. The male will then extend his front legs towards the female to touch her. If the female remains receptive, the male will climb on the female's back and inseminate her with his palps.
A 2008 study of the species Phintella vittatain in Current Biology suggests female spiders react to the males reflecting ultraviolet B light before mating, a finding that challenges the previously held assumption that animals did not register ultraviolet B light. It has recently been discovered that many jumping spiders seem to have auditory signals as well; amplified sounds produced by the males resemble buzzes or drum rolls.
Taxonomy and systematics
|Jumping spider classification|
The monophyly of the family Salticidae is well established through both phylogenetic and morphological analyses, but no consensus exists on what other group of spiders are most closely related to the jumping spiders. Suggested sister groups have included the oxyopids (lynx spiders), thomisids (crab spiders), clubionoids (sac spiders), and web-building spiders.
Jumping spiders can be divided into three major lineages: the lyssomanines (subfamily Lyssomaninae), the spartaeines (subfamily Spartaeinae), and the salticoids (unranked clade Salticoida). Of these, the Salticoida account for over 90% of all jumping spider species. Salticoida can be further divided into numerous groups, including Amycoida, Astioida, Aelurilloida, Euophryinae, Heliophaninae, Marpissoida, and Plexippoida.
Models for mimicry
Some small insects are thought to have evolved an appearance or behavioural traits that resemble those of jumping spiders and this is suspected to prevent their predation, specifically from jumping spiders. Some examples appear to be provided by patterns on the wings of some Tephritid flies, nymph of a Fulgorid and possibly some moths.
Very few jumping spider fossils have been found. Of those known, all are from Cenozoic era amber. The oldest fossils are from Baltic amber dating to the Eocene epoch, specifically, 54 to 42 million years ago. Other fossil jumping spiders have been found in Chiapan amber and Dominican amber.
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Among the subfamilies in this clade are the following:
The Amycoida form a large neotropical radiation from which only the related Sitticus and Attulus have reached the Old World. The Marpissoida are also mainly found in the New World. The Plexippoida are, except for Habronattus, an Old World group, as are the Heliophaninae. This suggests that much of the diversification of the salticids occurred after the separation of the continents of Old and New World.
Among the salticids not included in the Salticoida are:
- "Lapsiines": Lapsias, Galianora, Thrandina
- Maddison, W.P. & Hedin, M.C. (2003): Jumping spider phylogeny (Araneae: Salticidae). Invertebrate Systematics 17: 529–549. PDF doi:10.1071/IS02044
- Maddison, Wayne P. (2006): New lapsiine jumping spiders from Ecuador (Araneae: Salticidae). Zootaxa 1255: 17-28. PDF
- Maddison, Wayne P.; Bodner, Melissa R. & Needham, Karen M. (2008): Salticid spider phylogeny revisited, with the discovery of a large Australasias clade (Araneae: Salticidae). Zootaxa 1893: 49-64. Abstract
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