Field of Science

Squirrels and Cedar-Mice (Taxon of the Week: Sciuridae)

The momonga or Japanese flying squirrel, Pteromys momonga: less than a foot in length and possibly the most revoltingly adorable animal in existence. Photo from here.

The Sciuridae are a family of about 280 species of rodent native to all continents except Australia (where they are currently represented only by a population of the Indian palm squirrel Funambulus pennanti living in and around Perth Zoo that was first established in 1898). The family is primarily arboreal though some lineages in the subfamily Xerinae have become terrestrial. According to the classification adopted by Thorington & Hoffmann (2005), living squirels are divided between five subfamilies. There are also two currently standing fossil subfamilies that have been recognised in the Sciuridae.

The Cedromurinae (a name that can be translated as "cedar-mice") are a small group from the Oligocene of the North American Great Plains (Korth & Emry 1991). Cedar-mice lack certain features of the dentition of living squirrels, placing them outside the sciurid crown group. Most notable of these is the lack of sciuromorphy, a derived mode of attachment of the jaw muscles. The muscle attachment sites for cedar-mice do not extend as far forward on the skull as in crown squirrels. Sciuromorphous squirrels are known from even earlier in the fossil record than cedar-mice and the two groups would have been contemporary.

The neotropical pygmy squirrel Sciurillus pusillus, a phylogenetically isolated squirrel species found in Amazonian South America. Photo from here.

Also quite distinct in its dentition was the Chinese Pleistocene Aepyosciurus orientalis, placed in its own subfamily when originally described (Wang & Qiu 2003) though this probably needs further investigation*. Aepyosciurus had the highest tooth-crowns of any squirrel, probably an adaptation to living on a much tougher diet on the arid Tibetan plateau.

*Chinese vertebrate palaeontology seems to be awash with monotypic suprageneric taxa established for 'distinctive' species without any attempt to actually place the species phylogenetically. Many of these are later sunk into previously established, more inclusive groups (such as happened with Yuesthonyx). Explanatory factors for this situation probably include lack of communication between Chinese and foreign researchers and/or lack of opportunities to study non-Chinese specimens and publications, and a perceived link between personal prestige and the numbers and significance of taxa described.

Indian giant squirrel, Ratufa indica. Including the tail, giant squirrels can reach close to a metre in length. Photo by Rajiv Lather.

The known distributions of the cedar-mice and many of the earliest squirrels indicates a North American origin for the Sciuridae. However, squirrels invaded Eurasia early in their history with numerous subsequent migrations back to North America; probably the great majority of living American squirrels are descended from Eurasian ancestors (Mercer & Roth 2003). Perhaps the only living squirrel that might have a claim to a lineage untainted by Eurasian influence is the South American Sciurillus pusillus which may be the sister group to all other living squirrels. Also very basal are the Ratufa giant squirrels of southern Asia. The remaining squirrels form a clade divided between the Callosciurinae (the Asian palm squirrels), Sciurinae (including the 'typical' and flying squirrels) and Xerinae (African squirrels, chipmunks and ground squirrels).


Korth, W. W., & R. J. Emry. 1991. The skull of Cedromus and a review of the Cedromurinae (Rodentia, Sciuridae). Journal of Paleontology 65 (6): 984-994.

Mercer, J. M., & V. L. Roth. 2003. The effects of Cenozoic global change on squirrel phylogeny. Science 299: 1568-1572.

Thorington, R. W., Jr & R. S. Hoffmann. 2005. Family Sciuridae. In: Wilson, D. E., & D. M. Reeder (eds). Mammal Species of the World: A Taxonomic and Geographic Reference 3rd ed., vol. 2 pp. 754-818. John Hopkins University Press: Baltimore.

Wang B. & Qiu Z. 2003. Aepyosciurinae——a new subfamily of Sciuridae (Rodentia, Mammalia) from basal loess deposits at the northeastern border of Tibetan Plateau. Chinese Science Bulletin 48 (7): 691-695. (A Chinese version of this paper appears to have been published two months earlier.)

Earthworms of the Amazon (Taxon of the Week: Urobenus buritis)

This week's Taxon of the Week post has been delayed a little: getting hold of some of the reference for it required me to enter a real library and locate an actual journal physically printed on paper. Always an experience.

Urobenus buritis (no pictures of this one, I'm afraid) is an earthworm only recorded from the region of Manaus in the state of Amazonas in northern Brazil. This may or may not be significant. Perhaps not surprisingly, collections of earthworms around the Amazon have been somewhat scattered; most of our knowledge of the area (including U. buritis itself in Righi et al., 1976) can be credited to the work of one researcher, Gliberto Righi. Nevertheless, what we do know suggests that many species in the area have very localised ranges (James & Brown, 2006). Earthworms are poor dispersers and many of them are very particular in their habitat preferences, making them highly vulnerable to disturbance.

The genus Urobenus possesses three pairs of calciferous glands in segments VII-IX (many other glossoscolecid genera possess only two*) and can be distinguished from the similar genus Rhinodrilus by the shape of the glands. In Rhinodrilus, all three pairs of glands are tubular; in Urobenus, the first two pairs are tubular and the third pair (in segment IX) is sac-shaped (Righi, 1985). Urobenus buritis has three pairs of spermathecae, delicate anterior septa and the male pores opening at segments 20-21 (U. brasiliensis, which U. buritis was originally described as a subspecies of, has them at segments 19-20).

*Offhand, this is something to consider when thinking about how vertebrate diversity compares to invertebrate diversity. Two externally quite similar taxa, that the vast majority of people would regard as both being 'just worms' and pretty much identical, may actually differ in something as seemingly basic as how many organs they have.

I haven't seen the original description of Urobenus buritis, so I can't say whether there is any information on its lifestyle, but the closely related U. brasiliensis is an epigeic (living above the ground) species inhabiting leaf litter (James & Brown, 2006). Many of the earthworms around the Amazon have cyclical life histories to deal with the contrast between wet and dry seasons. Tuiba dianae migrates towards drier forest as flood waters rise, maintaining a distance of at least five metres from the water's edge. Other species, such as Andiorrhinus tarumanis, climb up the nearest tree during the wet season and take up residence in patches of leaf litter trapped in the forest canopy while the ground is flooded. Whether Urobenus buritis indulges in such behaviour is currently unknown.


James, S. W., & G. G. Brown. 2006. Earthworm ecology and diversity in Brazil. In Soil Biodiversity in Amazonian and other Brazilian Ecosystems (F. M. S. Moreira, J. O. Siqueira & L. Brussaard, eds) pp. 56-116. CABI Publishing.

Righi, G. 1985. Sobre Rhinodrilus e Urobenus (Oligochaeta, Glossoscolecidae). Boletim de Zoologia 9: 231-257.

Righi, G., I. Ayres & E. C. R. Bittencourt. 1976. Glossoscolecidae (Oligochaeta) do Instituto Nacional des Pesquisas da Amazônia. Acta Amazônica 6 (3): 335-367.

The Worst of Girault

Though most researchers have accepted the legitimacy of A. A. Girault's private publications, there are at least two taxa described therein that will almost certainly never see formal use.

The conflict between Girault and his then-manager Johann Illingworth (Illingworth was yet another person accused by Girault of placing economic considerations ahead of pure research and of crushing Girault's attempts to do otherwise) lead to Girault's 'publication' of the taxon Shillingsworthia shillingsworthi. Girault's description of this unusual micro-wasp1 reads:


Like Polynema but petiole, head, abdomen, mandibles absent. S. shillingsworthi, blank, vacant, inaneness perfect. Nulliebiety remarkable, visible only from certain points of view. Shadowless. An airy species whose flight cannot be followed except by the winged mind. From a naked chasm on Jupiter, August 5th, 1919.

This so thin genus is consecrated to Doctor Johann Francis Illingworth, in these days remarkable for his selfless devotion to entomology, not only sacrificing all of the comforts of life, but as well his health and reputation to the uncompromising search for truth and for love of "those filmy people of the air." Honour him!

More sinister than the description of fictional insects was Girault's naming of a new species of human - "New or Business Woman"2. Sadly, the 'proper role' of women was a prominent topic in many of Girault's later publications (one of its more unusual incarnations was the reference to Elizabeth I as the "madman Queen"3, 4). This was Girault's formal description of Homo perniciosus:

Abnormal female (loveless, without offspring); heart functionless; mammae aborted; psychology novel (as supposed) but artificial; gay, high-coloured, feral, brass-cheeked, shape lovely like Woman but nature hard (selfless, thoughtless, proud, unsympathetic, irresponsible, aggressive, irritant, insensible, luxurious, pugnacious, over-active, inquisitive, mischievous, voracious and even carnivorous; antagonistic, ungentle, immodest, critical, competitive, poisonous); conduct unstable (even inclined to treachery), the lips compressed, body strong. Everywhere but rare in natural habitat.
From young adults, these commonest, 1923, Australia.

1Some insects never before seen by Mankind. 1920.
2Homo perniciosus and new Hymenoptera. 1924.
3Some Insecta and a new All Highness. (Notes compiled in fear and sorrow). 1928.
4Description of a case of lunacy in Homo and of new six-legged articulates. 1929.


Alexandre Girault: A Man Against the World

Alexandre Girault at age 20, as he appeared in Gordh et al. (1979).

Having recently discussed the availability of privately published taxonomic works, it is appropriate that I recently received a copy of Gordh et al.'s (1979) reprinting of the privately published pamphlets of Alexandre Arsene Girault. A. A. Girault (1884-1941) is one of entomology's most controversial figures, an incredibly prolific worker possessed of a character that could charitably be described as difficult, uncharitably as barking mad. Girault named more than 3000 taxa over the course of his career (possibly more than any other taxonomist*), often under extremely difficult circumstances. As well as his monographic publications through more traditional venues, about 1000 of these taxa were published in privately produced pamphlets between 1917 and 1937.

*Update: Gunnar has reminded me in the comments of an even more prolific taxonomist, the crane-fly researcher Charles P. Alexander who published more than 11000 taxa. That represents a little less than one new species for every day of his working life.

Born and raised in Maryland in the United States, Girault worked for the U. S. Department of Agriculture from 1904-1911 until a request from the Queensland government for a capable entomologist sent him to Australia with glowing recommendations. Girault worked on pests in sugar plantations until 1914 when he decided to return to the States and focus entirely on purely taxonomic entomology. However, he was frustrated in that design and returned to Australia in 1917. After his return, his employment was irregular, including extended periods of unemployment and periods when he held day jobs as shopkeeper or quarry worker. His behaviour on a personal level became increasingly difficult and he was eventually admitted to a mental asylum in 1939 suffering from paraphrenia two years before his death.

Girault was obsessed with the importance of pure research for its own sake without recourse to low economic considerations ("The fear was, these things would no longer be studied was it thought they helped man's agriculture no more than the sunshine does. But who can doubt its greater use when love is the inspiration; man's soul is helped so"1). In between the basic taxonomic descriptions, his privately published papers are littered with poetry and commentaries on this subject including numerous invections against those of his colleagues whom Girault felt had betrayed this ideal ("Are we coming to be a nation of lilliputs, the socalled entomologists reduced to farmer's clercks and others accordingly? Is there a scientist here who dares to speak out what he sees or ought or who can e'en think freely, without his thought be limited by mandate to some "problem"? Find him" 1). Some of these inserts can be a little jarring, to say the least; one list of synonymies reads, "Hemiptarsenoideus is Hemiptarsenus. Baocharis marlatti is Aphelinus subapterus. Liberty is Soul. Rhopalicus americanus is Spintherus pulchripennis"1. They are also often disproportionately vituperative; a particularly bitter paragraph, complaining of "An outrage, I should think, robbing the poet as Tarquineus raped Lucrece; worse insulting the heavens"2, was written in response to an editor publishing a paper of Girault's as "New chalcid parasites from Malaya" when Girault had intended it to be called "New chalcid-flies from Malaya".

Girault in 1924 with his wife Elizabeth and three of their children. Elizabeth Girault (whom Girault described as a "faithful friend and true-hearted wife"4) became seriously ill with tuberculosis in 1923 and remained so until her death in 1931. His wife's health and the strain of raising five children on his irregular income added to Girault's difficulties. Photo from here.

It was primarily Girault's insistence on including these discourses that required him to publish privately. His acerbic nature lead a number of editors to make blanket refusals to handle any of his manuscripts. However, it is also true that research without direct economic application was not viewed favourably in a country passing through the fiscally straitened aftermath of the Great War. While he was working for the Queensland Department of Agriculture in the early 1930s, successive directors attempted to curtail his taxonomic work, presumably because the time he was devoting to it was intefering with his work on economic entomology. Also, economic restrictions after 1915 meant that Girault was no longer able to publish extensive manuscripts in the Memoirs of the Queensland Museum as he had done previously.

Setting aside their eccentric presentation, the descriptions of new taxa themselves vary in their usefulness. Some are dreadful, others were (at least by the standards of their time) very good. Often more problematic for succeeding researchers has been the exceedingly poor condition of many of Girault's type specimens. With no access to supplies other than what he could beg from more fortunate associates, Girault resorted to measures such as mounting multiple specimens under fragments of coverslip on a single slide. He also often mounted body parts such as heads on a separate slide from the remainder of the specimen. Girault also expressed disdain for such niceties as labelling ("We do not desire specimens any more per se. It is the label we want now"1), orthography ("A crooked line, a blot causes no worry; they are bodies, not souls"1) and detailed descriptions ("Longer descriptions are not needed, are even confusing"4).

The legitimacy of Girault's private publications for nomenclatural purposes was long disputed but over time a consensus has developed that they are acceptable (even if only because the usage of names from those publications by later authors has made it potentially more problematic to dismiss them than accept them). If there is one object lesson to be derived from Girault's failings, it is the value of type specimens. While there is no denying that the sheer volume of Girault's publications has represented a significant hurdle for later researchers, the continued availability of Girault's types (mostly in the Queensland Museum) has allowed progress where it would have otherwise been impossible. For instance, of the hundreds of Giraultian taxa referred to by Bouček (1988), only one was regarded as completely unidentifiable. For many others, the currently available information is inadequate but may be enough that further investigation has a reasonable chance of resolving matters.

1Descriptiones hymenopterorum chalcidoidicarum variorum cum observationibus. V. 1917.
2New animals of Australia and old men of the earth. 1921.
3Some gem-like or marvellous inhabitants of the woodlands heretofore unknown and by most never seen nor dreamt of. 1925.
4New pests from Australia, VIII. 1930.

REFERENCES (Girault's publications in the footnotes above)

Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 1-400 (including biography of Girault by E. C. Dahms).

More than Just Sophophora (Taxon of the Week: Drosophilidae)

The variety of drosophilids, from here. In the classification used in this post, the species illustrated would be Sophophora simulans and Idiomyia conspicura.

Today's Taxon of the Week is a group of which one particular species has already occupied a reasonable chunk of screen time at this site. The recent taxonomic controversy over Sophophora melanogaster* has been covered here, here and here, and I'm sure that by now we're all well and truly sick of the horrid little beast. The Drosophilidae is a family of some 3500 species of which S. melanogaster is but one. Surely it's time some of the other drosophilids were given their dues?

*As noted in earlier posts, there is really no justification for including the Sophophora species in the genus Drosophila. Even if you subscribe to a classificatory theory that forgives paraphyly, there are simply too many non-Drosophila genera separating Sophophora from Drosophila sensu stricto to make including the latter in one genus reasonable (see Linde et al., 2010, and various other references mentioned therein). It's like the entomological version of West and East Pakistan. [facetious snark] So now the decision about appropriate naming has been officially made, can we please all get over it and deal with the consequences? [/facetious snark]

Drosophilids are treated as part of a large grouping of flies known as the acalyptrates. Acalyptrates possess short antennae with a horseshoe-shaped fissure between the antennae and the upper part of the face but lack a calypter, a lower lobe at the base of the wing found in houseflies and blowflies. Distinguishing the various acalyptrate families from each other can be an absolute nightmare (I speak from painful experience) but the features of Drosophilidae include an incomplete subcostal vein in the wing, three (occasionally two) pairs of bristles on the face with the front pair pointing forwards and the remainder pointing backwards, and usually no bristles on the anepisternum (one of the side plates on the thorax) (Grimaldi, 1990).

Pair of Drosophila endobranchia courting on the carapace of the land crab Gecarcinus ruricola whose mouthparts will become home to their commensal larvae. From Stensmyr et al., 2008.

Most drosophilids are small flies (the largest belong to the genus Idiomyia and have wingspans of up to two centimetres). The majority of species feed as larvae on yeasts and the like growing on decaying plant matter but a number are more outré in their dietary habits (Ashburner, 1981). Scaptomyza larvae mine leaves while larvae of Lissocephala powelli, Drosophila carcinophila and D. endobranchia live as commensals on the mouthparts of land crabs, feeding on the bacteria growing there. Several species of various genera are predatory, mainly on small plant-sucking bugs such as scales and whiteflies though other targets are known (including two unrelated species feeding on frog eggs and at least one record of larvae of a generalist species taken from a secondarily infected wound of a surgery patient). The Drosophila simulivora group includes four West African species whose larvae are aquatic predators of midge and sandfly larvae. Perhaps the most incredible carnivorous drosophilid is the European Cacoxenus indigator which lays its eggs in the brood cells of the mason bee Osmia coerulescens. The mason bee arranges its cells in single file in a long tube and the female Cacoxenus will insert a single egg into each cell except the last. The fly larva feeds on the food stores left for the bee larva and eventually on the bee larva itself (though it is unclear whether the fly actually attacks the bee while it is alive or simply scavenges it after it dies from strarvation, as it has been known for the bee and fly to both emerge alive from a single cell). The reason the female Cacoxenus does not lay in the terminal cell is that, once her own larvae pupate into adult flies, they are unable to break free of the brood cell themselves and are dependent on the one surviving bee to release them as it frees itself from the nest.

Two males of the hammer-head fly Idiomyia heteroneura performing competitive displays. This is an endangered species with a restricted range on the island of Hawai'i. Photo from here.

The highest concentration of drosophilid diversity is found in the Hawaiian islands which are home to perhaps more than 1000 species (O'Grady et al., 2003) divided between the genera Scaptomyza and Idiomyia (the latter referred to by many authors as "Hawaiian Drosophila"). Many of these species, it should be noted, are exceedingly rare and localised and many remain undescribed. Molecular analysis indicates that the Hawaiian drosophilids form a single clade, indicating a single colonisation of the island (O'Grady & DeSalle, 2008). However, Scaptomyza also includes a number of continental species that appear to be derived from a Hawaiian ancestor.


Ashburner, M. 1981. Entomophagous and other bizarre Drosophilidae. In The Genetics and Biology of Drosophila vol. 3A (M. Ashburner, H. L. Carson & J. N. Thompson, eds) pp. 395-429. Academic Press: London.

Grimaldi, D. A. 1990. A phylogenetic, revised classification of genera in the Drosophilidae (Diptera). Bulletin of the American Museum of Natural History 197: 1-139.

Linde, K. van der, D. Houle, G. S. Spicer & S. J. Steppan. 2010. A supermatrix-based molecular phylogeny of the family Drosophilidae. Genetics Research 92: 25-38.

O'Grady, P., J. Bonacum, R. DeSalle & F. Do Val. 2003. The placement of Engiscaptomyza, Grimshawomyia, and Titanochaeta, three clades of endemic Hawaiian Drosophilidae (Diptera). Zootaxa 159: 1-16.

O'Grady, P., & R. DeSalle. 2008. Out of Hawaii: the origin and biogeography of the genus Scaptomyza (Diptera: Drosophilidae). Biology Letters 4: 195-199.

The Problem of Publication Again

I haven't been a member of the DML for a while so I missed the controversy of the last few days over the publication (or not) of a book called Notes on Early Mesozoic Theropods. Mike Taylor has written about the situation at SV-POW! and I recommend reading his post. Basically, the question of whether or not this book (and the new theropod taxon described within) counts as 'published' circles around the fact that the book is distributed as "print on demand". In other words, copies of the book are printed as they are ordered rather than being printed in a single run of multiple copies and some people have questioned whether this satisfies the ICZN requirement of "simultaneously obtainable copies [produced] by a method that assures numerous identical and durable copies" (Art. 8.1.3)*. Print-on-demand has been rare in the past because the cost of preparing templates made printing single copies uneconomical; as printing has become cheaper, it may be expected to become more standard. Indeed, one could argue that it should be so, being potentially less wasteful than producing large runs of potentially unwanted copies.

*In the comments for the SV-POW article, I indicated that the electronic publication procedure proposals being considered by the ICZN (ICZN, 2008) included disqualification of print-on-demand publications from availability. I was wrong about this; the proposal merely includes a discussion of issues potentially connected with print-on-demand (specifically, the possibility of alterations between printings).

In previous posts, I've discussed the availability of electronic publication, rare publications and self-published works. One issue I haven't discussed in detail, but which is perhaps the most important of all, is that it is becoming increasingly difficult for the non-specialist to decide what is 'published' and what isn't. In the past, media such as private letters, theses and conference handouts have been disqualified as valid 'publications' because they were of ephemeral nature and/or limited distribution. Conversely, anything that was widely available could usually be safely assumed to be published because there really was little way for it to be otherwise. Electronic publication has changed all that; supposedly 'unpublished' works may have audiences running into the thousands.

Even most printed journals are now more widely read as electronic versions. The largest zoological taxonomic journal currently operating is Zootaxa which now publishes issues on more or less a daily basis (twenty issues in a single day on 14 May earlier this year). The vast majority of Zootaxa's articles are distributed electronically; printed copies are effectively only produced to confirm availability. But most of us have only the word of the Zootaxa editors that these printed copies exist; we're not tracking them down ourselves because the electronic version meets our needs. For all most of us know, Zootaxa may have halted printed production some time ago (or be printing on demand). The same goes for a great many other major taxonomic journals.

The minutiae of determining whether or not a taxonomic work has been validly 'published' can be daunting enough for experienced workers; for non-specialists, they may seem impossible to follow. Nor, in an ideal world, should a non-specialist ever have to make such a decision, no more than someone should be expected to assemble the motor themselves before taking their new car for a drive. Like it or not, I'm afraid there is no escaping the conclusion that electronic publication of nomenclatural acts is inevitable and will be accepted by the taxonomic community with or without the endorsement of the Code (Taylor, 2009). It's simply too confusing otherwise.

So what, practically speaking, should we actually be doing about it? To begin with, I agree with Mike Taylor's statement in the SV-POW post that we need to make registration of new nomenclatural acts (as currently beeing developed for ZooBank) compulsory. There are potential objections that could be made to compulsory registration (I've made some in the past myself) but I think that, overall, the advantages outweigh the disadvantages. Registration will not solve the problems surrounding marginal publications but it will make it easier to manage them (especially if specialists are able to upload comments on the availability of registered names). And at least non-specialists would be able to feel secure that names not properly registered can be safely ignored.


International Commission on Zoological Nomenclature. 2008. Proposed amendment of the International Code of Zoological Nomenclature to expand and refine methods of publication. Zootaxa 1908: 57-67.

Polaszek, A., M. Alonso-Zarazaga, P. Bouchet, D. J. Brothers, N. Evenhuis, F.-T. Krell, C. H. C. Lyal, A. Minelli, R. L. Pyle, N. J. Robinson, F. C. Thompson & J. van Tol. 2005. ZooBank: the open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62 (4).

Taylor, M. P. 2009. Electronic publication of nomenclatural acts is inevitable, and will be accepted by the taxonomic community with or without the endorsement of the Code. Bulletin of Zoological Nomenclature 66 (3): 205-214.

More Wolfies (Taxon of the Week: Artoriinae)

Artoria mckayi, an inhabitant of alpine gravel river banks in eastern Australia. Photo by D. Paul.

Wolf spiders, or "wolfies" as I tend to refer to them, were previously covered here in an earlier post. The subject of today's post is one of the specific subgroups of wolf spiders, the Artoriinae.

Though one of primary assemblages of wolf spiders in the Australian region, the Artoriinae were only formally established as a distinct group in 2007 (Framenau, 2007). The group had previously been identified in molecular phylogenies as an unnamed clade sister to the combined clade of the subfamilies Lycosinae and Pardosinae. At least one morphological character has also been identified supporting the clade, the presence of a basal apophysis (side-branch) on the embolus, the intromittent part of the male's copulatory pedipalp.

A Tetralycosa specimen captured in the Great Victoria Desert. Photo by June Hudson.

Artoriinae are primarily restricted to Australia and the Pacific islands except for Artoria parvula which is found from northern Australia to the Philippines. Inclusion of the Sumatran Lycosella tenera is contingent on whether or not it is truly congeneric with Hawaiian Lycosella species (Framenau, 2007) while three African species listed in Artoria by Platnick's World Spider Catalog owe their position to Carl-Friedrich Roewer, whose classification of wolf spiders is generally regarded as unmitigated bollocks, and require a second look. More recently, it was suggested that the South American genera Lobizon and Navira may warrant consideration as possible artoriines (Piacentini & Grismado, 2009). Framenau (2007) placed eight genera in the Artoriinae (including Lycosella) plus two probable undescribed genera; Framenau's (2007) "new genus 2" has since been dubbed Kangarosa* (Framenau, 2010).

*One of the new Kangarosa species was named by Volker Framenau after his recently born son, placing Yannick Framenau in an exclusive club of people to have a new species named after them before they've even completed toilet training.

Lobizon corondaensis, an Argentinian species that may or may not be related to the Artoriinae. From Piacentini & Grismado (2009). 'Lobizón' is apparently Spanish for 'werewolf'.

Like other wolf spiders, artoriines are mostly conservative in their overall appearance. They do exhibit a reasonable size range, from the 2.6 mm Artoria palustris to 25 mm Tetralycosa species. Males of the South-West Australian species Artoria schizocoides possess a unique brush of spatulate setae on the underside of the first tibia (Framenau & Hebets, 2007). The members of the genus Tetralycosa are burrowing species that are invariably found in high salinity environments such as around salt lakes.


Framenau, V. W. 2007. Revision of the new Australian genus Artoriopsis in a new subfamily of wolf spiders, Artoriinae (Araneae: Lycosidae). Zootaxa 1391: 1-34.

Framenau, V. W. 2010. Revision of the new Australian wolf spider genus Kangarosa (Araneae: Lycosidae: Artoriinae). Arthropod Systematics and Phylogeny 68 (1): 113-142.

Framenau, V. W., & E. A. Hebets. 2007. A review of leg ornamentation in male wolf spiders, with the description of a new species from Australia, Artoria schizocoides (Araneae, Lycosidae). Journal of Arachnology 35 (1): 89-101.

Piacentini, L. N., & C. J. Grismado. 2009. Lobizon and Navira, two new genera of wolf spiders from Argentina (Araneae: Lycosidae). Zootaxa 2195: 1-33.