Maison Verreaux - Animal Specimens of All Varieties

Jules Verreaux. Photo from here.

Last week, I mentioned that I was attempting to research details of the lives of the 19th Century naturalists Jules and Édouard Verreaux, and indicated that I would share what I had been able to find. This is that post. I warn you, it's going to be a trifle incoherent. I'm no Brian Switek, and the problem is that despite the significant presence of Maison Verreaux in mid-1800s European zoology, none of the Verreauxs (Verreauxes?) seem to have left much in the way of personal accounts of their activities. As a result, one is forced to try and cobble together details from secondary sources and other writer's allusions, and when one does find more than one description of a certain event, the descriptions often conflict. (There are also a few significant references that I don't have access to.)

Jules Pierre* (born 1807), Jean-Baptiste Édouard (1810) and Joseph Alexis (the youngest) were the three sons of Pierre Jacques Verreaux, the founder in 1800 (or shortly afterwards) of the firm Maison E. Verreaux in Paris. Their mother Joséphine was the sister of Pierre Antoine Delalande, another well-known French naturalist. Maison Verreaux was a commercial taxidermy and natural history firm, bringing in specimens from all other the world and reselling them to interested parties, including a number of museums (the Muséum national d'Histoire naturelle seems to have been a big customer). In 1818, at the age of eleven, Jules accompanied his uncle Delalande to South Africa, and was there until 1820. He was to return to the Cape in 1825, after his uncle's death, and became a curator of the South African Museum in 1829.

*Not even his name is unequivocable - a couple of sources give it as "Pierre Jules".

In 1828, Jules Verreaux was sued for marriage by the family of Elisabeth Greef, a young woman who had borne an illegitimate child to Jules earlier that year (McKenzie, 2005). Jules had previously proposed marriage to Elisabeth, only to revoke the proposal later. Jules defended himself in the court with the claims that he was ignorant of having made such a proposal due to his not speaking Dutch, that he was a minor and so not empowered to make such a proposal without parental consent, and that it was all Elisabeth's fault anyway for being a slut (he even went to the extent of engineering for Elisabeth to mistakenly embrace a old boyfriend of hers while being watched by hidden witnesses). The court was not impressed by Jules' "artful, malicious and diabolical" behaviour - his claim of ignorance was obviously an outright lie, and Elisabeth was no more improprietous than any other young woman of the time. Unfortunately, Jules' minor status was an impassable legal hurdle, and the marriage suit had to be turned down. The judge did suggest to Elisabeth that she renew the suit once Jules reached his majority, but this never happened - as McKenzie (2005) suggests, perhaps the emotional trauma of being publicly scrutinised for slatternry was more than Elisabeth was willing to put herself through again.


The controversial "El Negro" while still on display in Spain in 2000. The exaggerated blackness of the skin was due to its being overly blackened with boot polish (chemicals such as arsenic probably used in preparing the mount would have bleached his skin). Widespread protest lead to the BaTswana man's repatriation and proper burial in Botswana shortly after this photo was taken. Photo from Univiersity of Botswana History Department, via South Pacific Taxidermy.

In 1829 or 1830, Édouard joined his brother in the Cape, taking custody of a large collection of South African specimens collected by Jules and returning with them to Paris in 1830 or 1831. One of these specimens, though it attracted relatively little attention at the time, was to become the subject of much controversy in the 1990s - the stuffed body of a young BaTswana man. See Parsons (2002) and Molina (2002) for the history of "El Negre", as this figure became known. Jules claimed in a letter to Georges Cuvier to have stolen the body from a graveyard the night after its burial, at no small risk to himself from its guards (though to be honest, I can't help wondering if he had jazzed up the story a little in order to increase the apparent value of the specimen - if so, the pitch failed. Cuvier didn't buy the body). The collection of human remains was not entirely uncommon at the time - the specimens from Delalande's earlier expedition to the Cape included nearly two dozen skeletons and a number of skulls, while the remains of Tasmanian aborigines were to be among material later collected by Jules in Australia. Only a few years earlier, in 1816, Cuvier himself had conducted his dissection of Sara Baartman, who had been patronisingly eroticised as the "Hottentot Venus" (despite not being a Hottentot). While Maison Verreaux never handled a huge number of human specimens (the unfortunate BaTswana notwithstanding), this was probably a matter of commercial interests rather than due to any ethical considerations (Molina, 2002).

During the period of 1832-1838, things get confusing (which is annoying, because this is one of the periods I most need to find out about). What is certain is that Édouard returned to the Cape in 1832 in the company of the youngest brother, Alexis, who was to remain in South Africa until his death in 1868 (Gunn & Codd, 1981). Édouard then took ship to east Asia in 1832 or 1833. In Jules' 1874 obituary in the Ibis, it states that Jules also travelled to east Asia, and implies that the two brothers both remained there until 1837. However, other sources (Stresemann, 1975; Gunn & Codd, 1981) state that Édouard returned to Paris in 1833 or 1834 to take over the running of Maison Verreaux's home office. Meanwhile, Stresemann (1975) implies and Dubow (2006) states that Jules remained as curator of the South African Museum until 1838! I have little idea how to decide between these accounts*, but it is certain that Jules returned to Paris in 1837 or 1838 and became established at Maison Verreaux. Also certain is that the wreck of the ship "Lucullus" in 1838 resulted in the destruction of a large number of specimens collected by Jules Verreaux on their way back to Paris from abroad (despite one source referring to Jules himself only narrowly surviving the wreck, in this regard at least I am inclined to trust the obituary, which states that Jules had returned on a separate ship from his specimens).

*Normally, I would trust the obituary as having been written closer to the time, but the obituary has a slipshod composition and 'boy's own story' tone that, to be honest, undermines its credibility for me. In cases where it disagrees with other sources, it also tends to form the minority.


Jules Verreaux's "Arab Courier attacked by Lions". This mount remains on display at the Carnegie Institute. Photo, again, from here.

For the next few years, Jules remained in Paris, but in 1842 (we're back on firmer dates now) he travelled to Australia with the backing of the Muséum national, where he was to remain until 1847. No detailed account of Jules' movements in Australia exists, but he seems to have mostly travelled in New South Wales and Tasmania. He may have also travelled to New Zealand about this time. After his return to Paris, he concentrated most of his attention on bird specimens, though continuing to handle a number of other taxa. His taxidermic diorama 'Arab Courier attacked by Lions', featuring a man mounted on a camel being (as the label said) attacked by two lions (the lions and the camel were all stuffed specimens; the man in this case was a mannequin*) was highly awarded in 1867 (dioramas were a great source of public entertainment before the invention of film). Offhand, the lions featured in the diorama were the Barbary subspecies, Panthera leo leo, which was to become extinct not long afterwards.

*Molina (2002) implies that it could have originally been a real human and later replaced by a mannequin, but other than the previous case of the BaTswana there seems little reason to entertain this suggestion.

Édouard Verreaux died in 1868. Jules Verreaux fled Paris at the beginning of the Franco-Prussian war in 1870, and was received enthusiastically in England by the young ornithologist Richard Bowdler Sharpe. He remained in England until his death in 1873.

While Jules Verreaux was highly praised by his associates during his lifetime, the affair with Elisabeth Greef was symptomatic of a certain, shall we say, economical attitude to the truth. This attitude has, unfortunately, clouded Jules Verreaux's legacy in zoology. Maison Verreaux specimens became notorious for poor locality data (usually with little more detail than 'Madagascar' or 'Australia'), and what little data there was has often proven unreliable. In at least a few cases, specimens were attributed to completely inaccurate localities (Olson et al., 2005). These misattributions may even have been deliberate, in order to give a specimen a more valuable provenance (Olson et al. mention, among others, a Virginia rail originally labelled 'Martinique' for which the original locality had been crossed out and 'Nlle Zelande' [New Zealand] written instead). As a result, many Verreaux specimens simply cannot be trusted as far as you can throw them. So the answer to my original question that sparked this search, whether a supposedly Australian Verreaux specimen held at Paris actually came from Australia, or whether it could perhaps have come from south-east Asia instead, can't be pinned down to anything more definite than "Who knows?"

REFERENCES

Dubow, S. 2006. A Commonwealth of Knowledge. Oxford University Press.

Gunn, M. & L. E. W. Codd. 1981. Botanical Exploration of Southern Africa: An Illustrated History of Early Botanical Literature on the Cape Flora. CRC Press.

McKenzie, K. 2005. Scandal in the Colonies. Melbourne University Publishing.

Molina, M. 2002. More notes on the Verreaux brothers. Pula: Botswana Journal of African Studies 16 (1): 30-36.

Olson, S. L., R. C. Fleischer, C. T. Fisher & E. Bermingham. 2005. Expunging the ‘Mascarene starling’ Necropsar leguati: archives, morphology and molecules topple a myth. Bulletin of the British Ornithologists’ Club 125 (1): 31-42.

Parsons, N. 2002. One body playing many parts- le Betjouana, el Negro, and il Bosquimano. Pula: Botswana Journal of African Studies 16 (1): 19-29.

Stresemann, E. 1975. Ornithology from Aristotle to the Present. Harvard University Press.

Taxon of the Week: Clausilioidea

The clausiliid Agathylla biloba. Photo by V. Wiese, via Palaeos.com.

Just a quick post today, as I've not much time. In fact, as I approach D-Day for the thesis, as well as tutoring labs for the next few months, I'm probably going to be putting a bit less up here. I'll try and keep the Taxon of the Week series going, at least.

The Clausilioidea are a superfamily of Stylommatophora (land snails). In the system of Bouchet et al. (2005), four families are assigned to this group - Palaeostoidae, Filholiidae, Anadromidae and Clausiliidae. The first three of these are all fossil families, so the Clausiliidae include all living clausilioids. The relationship of the fossil taxa with the Clausiliidae is up for debate - Schileyko (1979) pointed out that the Filholiidae weren't definite clausilioids (while he also thought that Urocoptidae, since excluded from the superfamily, were), while Wade et al. (2006) noted that the "Anadromidae" might be a polyphyletic group, with some related to the Clausilioidea and others to the Acavoidea.


Another clausiliid, Cochlodina laminata. Photo by D. Tymanov.

Their fossil associates notwithstanding, the Clausiliidae themselves are a clade of undoubted integrity (Wade et al., 2006), containing nearly 1300 living species. Most clausiliids are fairly small snails, though the largest, Megalophaedusa martensi, is a little under five centimetres long. All Clausilioidea are high-spired, with long narrow shells, and the Clausiliidae possess a unique feature called the clausilium, a calcareous plate attached to the opening of the shell that functions like an operculum, closing over the opening when the snail retreats inside (hence their common name of "door snails"). The foot of clausiliids appears surprisingly short (at least to me, but then I'm no expert on snails) compared to the length of the shell. The distribution of clausiliids seems somewhat spotty - main centres are Europe, eastern Asia and South America, with the South American taxa having probably invaded from Laurasia during the Cretaceous (Wade et al., 2006).


Mating clausiliid snails. Photo by T. Asami.

In the earlier post linked to above, I referred briefly to the division in land snails (which are always hermaphrodites) between "face-to-face" and "shell-mounting" copulation. Face-to-face mating is the more familiar behaviour, where the two mating snails (or slugs) lie alongside each other facing in opposite directions, extrude their genitalia and often both fertilise each other simultaneously. Among shell-mounters, in contrast, one snail climbs on top of the other, both face in the same direction, and generally insemination is unilateral with one individual donating and the other receiving sperm (however, after the iniatial sperm-donator has finished inseminating its partner, they may repeat the process with reversed roles). The two mating behaviours are closely (though not exactly) correlated with different shell shapes - face-to-face copulators are generally low-spired and rounded (like the common garden snail), while shell-mounters are usually high-spired (Davison et al., 2005). Clausiliidae, to match their high-spired shells, are invariably shell-mounters, though at least one clausiliid genus, Albinaria, differs from the others in that both individuals fertilise each other simultaneously.

In another earlier post where I discussed snail chirality (whether their shells were left-handed or right-handed), I mentioned that face-to-face copulators cannot generally mate if they have differing chiralities, because they can't readily bring their genital openings alongside each other. For shell-mounters, on the other hand, inter-chiral matings are far simpler and much more likely to be successful. In light of this, one might expect to see far more cases of reversed chirality among shell-mounters than face-to-face copulators, because the disadvantages of reversed chirality would not be as significant. And according to Asami et al. (1998), this is exactly what does happen - of those snail genera they surveyed including species with reversed chirality, over 90% were high-spired forms and probably shell-mounters.

REFERENCES

Asami, T., R. H. Cowie & K. Ohbayashi. 1998. Evolution of mirror images by sexually asymmetric mating behavior in hermaphroditic snails. American Naturalist 152; 225-236.

Davison, A., C. M. Wade, P. B. Mordan & S. Chiba. 2005. Sex and darts in slugs and snails (Mollusca: Gastropoda: Stylommatophora). Journal of Zoology 267 (4): 329-338.

Schileyko, A. A. 1979. The sytem of the order Geophila (=Helicida) (Gastropoda Pulmonata). Transactions of the Zoological Institute,
Academy of Sciences, USSR 80: 1-69.

Wade, C. M., P. B. Mordan & F. Naggs. 2006. Evolutionary relationships among the pulmonate land snails and slugs (Pulmonata, Stylommatophora). Biological Journal of the Linnean Society 87 (4): 593-610.

Request for Info

I'm hoping someone out there knows of a good summary of (and can potentially send me a copy of) the lives of the brother naturalists Jules and Édouard Verreaux. Specifically, I need to know when and where they were in Asia and Australia. Different sources have been giving me conflicting accounts.

I may share what I've been finding with you all at some point. There's no denying that the boys of Maison Verreaux led interesting (if somewhat reprehensible) lives...

The Sphinxes that aren't like the Others (Taxon of the Week: Smerinthini)

Female of the smerinthin moth Marumba quercus. Photo by Tony Pittaway.

Quick question - should the plural of "sphinx" be "sphinxes" or "sphinges"?

The sphinx moths or hawkmoths (Sphingidae) are one of the easiest lepidopteran families to recognise. Sphinxes tend to be fairly large (but not inordinately so), and are generally the speedsters of the moth world. Their rapid mobility is reflected in their wings, which are narrower, more streamlined and more pointed than those of other moth families. Sphinxes are not usually brightly-coloured, but they are none the less very handsome animals, with dapper patterns of earthy colours such as browns and soft pinks. The name "sphinx" is derived from their caterpillars - sphinx caterpillars have a way of sitting with the front of the body raised that is supposed to be reminiscent of the famed Egyptian statue.

Adult sphinxes are most famed as nectar feeders - with their long proboscides and ability to hover in front of the flowers they feed on, Old World sphinx moths have been described as the ecological counterparts of New World hummingbirds (I won't repeat the Xanthopan praedicta story here, but look it up if you're interested). One group of sphinxes, however, has decided to buck the trend. Sphinxes are divided into three subfamilies - Sphinginae, Macroglossinae and Smerinthinae. Smerinthinae are distinguished from the other two subfamilies because they lack the super-long proboscides. In at least one of the smerinthine tribes, Smerinthini, the adults are completely non-feeding (the subfamily as a whole is sometimes described as such, but the presence of pollen on proboscides of Ambulycini indicates that members of that tribe are still flower feeders - Beck et al., 2006). Adult Smerinthini are weaker fliers than other sphinx moths, and have less streamlined wings to match. Also, while caterpillars of other subfamilies tend to be fussy eaters, Smerinthini are relative gourmands, feeding on a wide range of host plants.


Paonias astylis, one of the few North American Smerinthini. The small-eyed and blinded sphinxes of the genus Paonias (so-called, I'm guessing, because the unusual relative position of the fore- and hindwings means that the eyespots on the latter are hidden) are perhaps some of the most distinctive sphinx moths. Photo by Jim McCormac.

Sphingidae as a whole are regarded as good dispersers, and European sphingid species tend to have wider ranges than moths of other families. However, a study of sphingid distributions in Indonesia found that, in line with their lower dispersal capabilities, smerinthine species showed a higher turnover between islands than members of the other two subfamilies (Beck et al., 2006). Smerinthini are mostly found in the Old World - only two species are found east of "Lydekker's line" between the Moluccas and New Guinea, and a handful of species are found in Nearctic North America.

It is easy to imagine a connection between the distinctive smerinthine life cycle and their poor dispersive abilities - with their short-lived adults and polyphagous larvae, female Smerinthini have neither the freedom nor the need to invest a lot of time in seeking out suitable host plants for their eggs. What is more uncertain is whether the smerinthine life cycle is derived from a more typically sphingid ancestor. The Smerinthinae have been suggested to be the basalmost subfamily of Sphingidae, and in some features they are more like members of closely-related families than other sphingids - their short proboscides (like Brahmaeidae) and preference for tannin-bearing trees over tannin-free shrubs (like Saturniidae). On the other hand, a genetic analysis by Regier et al. (2001) found support for a Sphinginae-Smerinthini clade excluding Macroglossinae. Mind you, Regier et al.'s analysis did not include representatives from the other smerinthine tribes.

Oh yes, and at least some Smerinthini have stridulatory apparatuses on their genital valves (Conner, 1999). These are singing moths.

REFERENCES

Beck, J., I. J. Kitching & K. E. Linsenmair. 2006. Wallace's line revisited: has vicariance or dispersal shaped the distribution of Malesian hawkmoths (Lepidoptera: Sphingidae)? Biological Journal of the Linnean Society 89 (3): 455-468.

Conner, W. E. 1999. 'Un chant d'appel amoureux': acoustic communication in moths. Journal of Experimental Biology 202: 1711-1723.

Regier, J. C., C. Mitter, T. P. Friedlander & R. S. Peigler. 2001. Re: Phylogenetic relationships in Sphingidae (Insecta: Lepidoptera): initial evidence from two nuclear genes. Molecular Phylogenetics and Evolution 20 (2): 311-316.

Linnaeus' Legacy #16 - Creationist Quote-Mine edition

The latest edition of Linnaeus' Legacy is up at Seeds Aside. Laurent has taken pity on the poor over-worked creationist labouring to prepare their quote-mines for the Darwin Day festivities, and so has presented them with a whole batch of quote-mines ready made and fresh off the press.

This month's keywords: fruity, non-existent, sex, bastard, sociability, highly elongate, impossible, sexual organs, appendages, genitalia, suckers, naked, control.

Most Unbelievable Organisms Evah!

Last week I asked for nominations for the title of Most Incredible Organism Ever. Thank you very much to those of you who responded with your selections. Some of them were organisms I'd already selected myself, some of you reminded me of amazing organisms that were even better than the ones that I'd considered*. Certainly, getting the list down to ten top nominations was not easy, and I'm sure anyone else would have chosen differently from myself. Allen Hazen pointed out that, strictly speaking, "incredible" means "inspires disbelief", and certainly some of the things I have lined up do exactly that.

*As an aside, something that never fails to amuse is looking up what Google search terms have brought people to Catalogue of Organisms. Trust me, "amazing organism" is bound to bring in the punters.

Honorable mentions should be given to those organisms that people nominated that I didn't end up using, because they're certainly all incredible. Allen Hazen suggested the platypus, while Alan nominated the aye-aye. Dave Coulter was all for the Osage orange, while Amie Roman asked me to "pick an onychophoran, any onychophoran".

But I'm afraid I ended up passing over these wonders. In no particular order, here are my nominations for "Most Incredible Organism" (click on the pictures to be taken to their source):



Homo sapiens Linnaeus, 1758: Both myself and Mike Keesey agreed on this one. As much as I hate to stoke this species' notoriously smug satisfaction, it has to be admitted that humans are pretty amazing. Douglas Adams once explained that "The History of every major Galactic Civilization tends to pass through three distinct and recognizable phases, those of Survival, Inquiry and Sophistication, otherwise known as the How, Why and Where phases. For instance, the first phase is characterized by the question How can we eat? the second by the question Why do we eat? and the third by the question Where shall we have lunch?" As far as we know, Homo sapiens is the only species on this planet to have reached Adam's second stage, let alone the third.



Polyascus polygenea (Lützen and Takahashi, 1997): Polyascus polygenea is a member of the Rhizocephala, notorious crustacean parasites of crabs. The larval rhizocephalan looks very similar to the larva of a barnacle (to which it is closely related), but when it finds a decapod host it burrows in and transforms into an almost fungus-like mass spreading through the hosts body. The only externally visible part of the parasite is its large egg-sac (the orange tube in the picture above, which does not show a Polyascus but another rhizocephalan species, Peltogaster paguri). The rhizocephalan egg-sac grows at the base of the crab's tail, where it would normally hold its own eggs. In order to make sure this spot is free, the rhizocephalan chemically castrates its host, preventing it from ever reproducing. It also affects its host's behaviour so that the crab lovingly tends the parasite's egg-sac as if it were its own. So powerful is the parasite's mental ju-ju that even male hosts that would not naturally produce eggs will tend the parasite just as a female would.

Vasha nominated the best-known rhizocephalan, Sacculina carcini, but I've decided to go with Polyascus polygenea because this species adds a further twist to the tale. A single Sacculina larva will give rise to a single egg-sac. But Polyascus reproduces within the host asexually by budding, so that one larva will give rise to multiple egg-sacs (Glenner et al., 2003).

Polyascus is also acting as the stand-in for all mind-controlling parasites. As we learn more about the natural history of parasitic organisms, it turns out that behavioral control of parasites over their hosts is not uncommon. Parasitic wasps make caterpillars guard the wasp's cocoons. Horsehair worms make crickets drown themselves so the aquatic adult worm can emerge. Tanya reminded me about Cordyceps unilateralis, a fungal parasite of ants that, when it's ready to produce spores, makes its host climb to the highest available point so that the spores will spread as far as possible. The ways of parasites are disturbing. And speaking of disturbing...



Acarophenax tribolii Newstead & Duvall, 1918: It is not uncommon for pregnant women to express delight at feeling their baby kick inside them. But what if it was doing more than just kicking? Mites of the genus Acarophenax are parasites of beetles that can claim to have perhaps the just-plain-ickiest life history of any animal. The sex ratio of this genus is highly skewed - depending on the species, a brood may contain up to thirty females, but usually only a single male. These offspring reach sexual maturity before they are even born, and the male proceeds to fertilise all of his sisters while still within their mother. In fact, the male doesn't even survive to become free-living - by the time the already-fertilised females emerge from their parent, the male has reached the end of his short (but extremely busy) lifespan. The advantage to the mite in this twisted incestuous life cycle? An exceedingly short generation time, of course - Acarophenax mahunkai, for instance, has a generation time of only three to five days (Steinkraus & Cross, 1993).



Mites of the closely related family Pyemotidae have a similar life cycle - the offspring reach full sexual maturity while in their mother, and begin copulating the instant that they emerge from their proud parent. Females of Pyemotes herfsi (shown in the picture above), known as "itch mites" and facultative biters of humans, can produce more than 250 fully mature offspring.



Welwitschia mirabilis Hook.f.: I also have to thank Tanya for reminding me of the wonder that is Welwitschia. Welwitschia mirabilis is unique to the Namib Desert in Angola and Namibia, and is a member of the gymnosperm order Gnetales along with the genera Ephedra and Gnetum. The Gnetales have received a lot of attention due to their much-debated phylogeny (morphological characters suggest they are the living sister group to angiosperms, while molecular analyses place them closer to conifers), but that's not what's so amazing about Welwitschia. It's not even the bright pink, insect-pollinated cones. What makes this plant so incredible is the way it grows. Welwitschia mirabilis only ever produces two adult leaves, followed by the death of the plant's apical meristem (growing tip). The two strap-like leaves, however, continue to grow indefinitely, and can reach lengths of over eight metres (most individuals look like they have more than two leaves, but this is only because of the leaves splitting as the ends get frayed). Welwitschia is very slow-growing, and individual specimens can live to be hundreds, if not thousands of years old.



Argentinosaurus huinculensis Bonaparte & Coria, 1993: There's no other way to say it - sauropods were just stupidly huge. And Argentinosaurus was one of the most ridiculous of all, being the largest well-characterised sauropod (potentially outdone only by such almost-apocryphal taxa as Amphicoelias fragillimus and Bruhathkayosaurus matleyi). With an estimated total length of nearly thirty metres, and potential weight of up to 80 tonnes... well, there's nothing much that can be said in response except "Whoa".

Sauropods are so huge that when a popular blog was set up dedicated to them, the site authors couldn't fit in the entire animal and were forced to dedicate themselves to a single section. I refer, of course, to the famed Sauropod Vertebra Picture Of the Week - SV-POW!. Rumour has it, however, that a second site is in the works devoted to sauropod crania, to be called "Sauropod Heads - Anatomy, Zoology And Morphology".



Rhizanthella slateri (Rupp) M. A. Clem. & P. J. Cribb, 1984: Rhizanthella is a small genus of three orchid species unique to Australia. What makes Rhizanthella so amazing is that its entire life cycle is spent underground. The plant is saprophytic, dependent on an associated fungus for nutrition, and its stems are entirely subterrean. Even the flowers do not have to break the surface - they are pollinated by minute gnats that can reach them through tiny cracks in the covering litter. The first known Rhizanthella specimens were discovered in 1928 when they were brought up by a farmer's plough, and only intermittent finds were made for a long time afterwards. Even today, their obscure habits mean that Rhizanthella species are poorly known. Sad to say, they are also all regarded as endangered. They are only known from restricted, scattered ranges, limited by the presence of their associated fungus and the tree of which it is in turn connected to mycorrhizally (in Rhizanthella gardneri, the tree is Melaleuca uncinata, but the associations of Rhizanthella slateri are still unknown).

Vasha reminded me of Rhizanthella by telling me of the American saprophytic plant Thismia americana, which also spends most of its life underground with only the minute flowers emerging above the surface. Thismia americana has not been recorded since 1916, and is feared to be extinct, though it is hard to know for certain. As described at the link, an intensive search in the early 1990s failed to find any specimens, but a concurrent dummy run using scattered white beads about the same size as T. americana flowers was also a failure.



Puccinia monoica Arthur, 1912: The object of the photo above is not a flower. It grew from a flowering plant, but it's not a flower. Puccinia monoica is a fungus parasitic on Brassicaceae (mustard) species. Like rhizocephalans on their crabs, Puccinia monoica changes the reproductive biology of its host, preventing it from growing its own flowers. Instead, it makes the host plant grow a tight whorl of leaves, which are covered by the bright yellow sporangia of the fungus. Not only does the fungus-induced 'false flower' look like a real flower, it even produces nectar and scent like a real flower, attracting insect pollinators just like a real flower would (Raguso & Roy, 1998). And just like pollen from a real flower, these pollinators carry spores from fungus to fungus, cross-fertilising the fungi as they do so.



Deinococcus radiodurans (ex Raj et al. 1960) Brooks and Murray 1981: A dose of radiation of 10 joules per kilogram will kill a human being. Sixty joules per kilogram will kill Escherichia coli. Deinococcus radiodurans may look like a fairly unremarkable bacterium at first glance, but it can withstand a radiactive dose of 5000 joules per kilogram and not even blink (that is, if it had eyes they wouldn't blink). It can withstand radiation so strong that its genome is simply blasted to pieces, stoically knitting the fragments back together again afterwards. Deinococcus can withstand extreme heat, extreme cold, and strong acidity. In a pun so bad that it demands to be repeated, this organism has been dubbed Conan the Bacterium. Pavlov et al. (2006) went so far as to suggest that Deinococcus' incredible resilience to radiation indicated an extraterrestrial origin, carried from Mars on an asteroid, but it seems more likely to be a by-product of resilience to other stressors such as desiccation (Cox & Battista, 2005). Still, one can't help wondering if, even if it didn't come from Mars in the first place, it has managed to make it over there on one of Earth's probes.

So resistant is Deinococcus to everything possibly imaginable, in fact, that we still have no idea where it lives naturally. It was first isolated from cans of irradiated beef, and has not yet been found to be abundant in any particular environment. Phylogenetically, Deinococcus forms a clade with the thermophilic bacterium Thermus (one species of which, Thermus aquaticus, is of enormous significance to molecular biology as the source of the Taq enzyme used in PCR). This clade is most commonly referred to (rather unimaginatively) as the Deinococcus-Thermus group, but I personally prefer the name given to them by Cavalier-Smith (2002) - Hadobacteria, the bacteria of Hades.



Proteus anguinus anguinus Laurenti, 1768: The white olm, the only truly cave-dwelling tetrapod (the closely related black olm, Proteus anguinus parkelj, is a surface-dweller). [Update: Much to my chagrinn, Nick Sly has reminded me that there are other cave-dwelling salamanders out there.] I've included the olm not only for its own sake, but as a representative of the entire world of troglobitic and stygobitic fauna (troglobitic animals are those that live in actual caves while stygobitic taxa live buried in the ground, usually in aquifers). In this strange, silent world, animals are almost entirely dependent on food particles washing down from the surface, so life underground is slow, and patient. Troglobites can go for incredible amounts of time without eating - Darren Naish informs us of an olm that was supposedly kept at the Faculty of Biotechnology in Ljubljana without food for twelve years! If that is what a large, complex vertebrate is capable of, imagine what is possible for the smaller invertebrates with their lower metabolic requirements.

And last, but certainly not least:



Wasmannia auropunctata (Roger, 1863): Commonly known as the little fire ant or electric ant (the latter name has been promoted in recent years to dissuade confusion with the larger, not closely related fire ants of the genus Solenopsis), Wasmannia auropunctata is regarded as one of the world's worst invasive organisms. It has been linked with decreases in biodiversity in locations to which it has been introduced, and has a painful sting to boot. It also has one of the world's most remarkable reproductive systems (Fournier et al., 2005). Like other ants, Wasmannia has both haploid males and diploid females, with the females divided between reproductive queens and non-reproductive workers. Genetically, though, Wasmannia is a little different from other ants. While males appear to mate with queens the normal way, only workers are produced by male fertilisation. Any new queens that are produced are genetically identical to their mothers. Still, the male lineage doesn't disappear - somehow, the male genes are able to eliminate the female genes from some of the eggs, and the resulting male Wasmannia are genetically identical to their fathers.

Wasmannia is one of very few organisms that exhibit androgenesis - clonally reproducing males. The only other known natural habitual cases are a cypress species, Cupressus dupreziana, and freshwater bivalves in the genus Corbicula, though odd cases of androgenesis have been recorded in laboratory and cultivated organisms (Hedtke et al., 2008). Effectively, the male and female Wasmannia are reproductively isolated from each other - they are separate species.

REFERENCES

Cox, M. M., & J. R. Battista. 2005. Deinococcus radiodurans — the consummate survivor. Nature Reviews: Microbiology 3 (11): 882–892.

Fournier, D., A. Estoup, J. Orivel, J. Foucaud, H. Jourdan, J. Le Breton & L. Keller. 2005. Clonal reproduction by males and females in the little fire ant. Nature 435: 1230-1234.

Glenner, H., J. Lützen & T. Takahashi. 2003. Molecular and morphological evidence for a monophyletic clade of asexually reproducing Rhizocephala: Polyascus, new genus (Cirripedia). Journal of Crustacean Biology 23: 548-557.

Hedtke, S. M., K. Stanger-Hall, R. J. Baker & D. M. Hillis. 2008. All-male asexuality: origin and maintenance of androgenesis in the Asian clam Corbicula. Evolution 62 (5): 1119-1136.

Pavlov, A. K., V. L. Kalinin, A. N. Konstantinov, V. N. Shelegedin & A. A. Pavlov. 2006. Was Earth ever infected by martian biota? Clues from radioresistant bacteria. Astrobiology 6 (6): 911-918.

Raguso, R. A., & B. A. Roy. 1998. 'Floral' scent production by Puccinia rust fungi that mimic flowers. Molecular Ecology 7 (9): 1127-1136.

Steinkraus, D. C, & E. A. Cross. 1993. Description and life history of Acarophenax mahunkai, n. sp. (Acari, Tarsonemina: Acarophenacidae), an egg parasite of the lesser mealworm (Coleoptera: Tenebrionidae). Annals of the Entomological Society of America 86 (3): 239-249.

Taxon of the Week: Stygnoplus

Cluster of an unidentified Stygnidae species. Photo from here.

South America is the current centre of described harvestmen diversity, with the bulk of this diversity represented by Laniatores - the shorter-legged, heavily armoured and often quite spiky suborder of Opiliones. Of the twenty-six families of Laniatores recognised in Pinto-da-Rocha & Giribet (2007), sixteen are found in the Neotropics (excluding the Podoctidae, whose sole "Neotropical" representative, Ibantila cubana from Cuba, was described from a specimen collected in a botanical garden and probably represents an introduction from parts unknown). The majority of research on South American harvestmen has focused on the largest family, the Gonyleptidae, but the remaining families are all waiting their turn.

The genus Stygnoplus belongs to one of these families, the Stygnidae. A number of features support the Stygnidae as a monophyletic group, perhaps the most apparent being their disassociated eyemound. Unlike most other harvestmen that have the two eyes located on a central eyemound, stygnids have the eyes placed some distance apart without a central mound - still fairly close in the subfamily Nomoclastinae, further apart in the subfamilies Stygninae and Heterostygninae (to the latter of which Stygnoplus belongs). About eighty species of Stygnidae have been described to date, but the existence of a number of undescribed species is known (Kury & Pinto-da-Rocha, 2002). Like most South American Opiliones, the stygnids have mostly only been studied from a taxonomic point of view, with very little known about their natural history. However, the Stygnidae are ahead of the game in that, unlike most other South American Opiliones (or, for that matter, Opiliones in general), they have been the subject of an actual phylogenetic analysis (Pinto-da-Rocha, 1997).


Another unidentified stygnid. Photo by artour_a.

The known centre of diversity for the Heterostygninae (including Stygnoplus) is the Lesser Antilles, though Stygnoplus was recorded from the mainland of South America by Villarreal-Manzanilla & Rodríguez (2004). (The type species, Stygnoplus forcipatus, had originally been described from the mainland, though with the completely uninformative and decidedly untrustworthy locality citation of "Colombia".)



In the absence of any other images of Stygnoplus online, here's Villarreal-Manzanilla & Rodríguez's (2004) figures of the Venezuelan species Stygnoplus lomion. Feel free to print them off, cut them out and see if you can assemble your own model of a South American arachnid. The appendages shown in the lower part of the plate are the pedipalps - stygnids, like many Laniatores, have absolutely terrifying raptorial pedipalps. If you look closely at the second photo on this post, you'll see that in this family the femora (the first large segment) of the pedipalps are quite long, giving these animals a quite impressive reach, perfectly designed to strike terror into the hearts (or other significant circulatory organs) of small invertebrates everywhere.

REFERENCES

Kury, A. B., & R. Pinto-da-Rocha. 2002. Opiliones. in Amazonian Arachnida and Myriapoda (J. Adis, ed.) pp. 345-362. Pensoft: Sofia.

Pinto-da-Rocha, R. 1997. Systematic review of the neotropical family Stygnidae (Opiliones, Laniatores, Gonyleptoidea). Arquivos de Zoologia 33(4): 163-342.

Pinto-da-Rocha, R., & G. Giribet. 2007. Taxonomy. In Harvestmen: The Biology of Opiliones (R. Pinto-da-Rocha, G. Machado & G. Giribet, eds.) pp. 88-246. Harvard University Press: Cambridge (Massachusetts).

Villarreal-Manzanilla, O., & C. J. Rodríguez. 2004. Descripción de una nueva especie y dos nuevos registros del género Stygnoplus (Opiliones, Stygnidae) para Venezuela. Revista Ibérica de Aracnología 10: 179-184.

Blinding Me with Science

Today's edition of Science, it turns out, is packed so chock-full of goodies that I hardly know where to turn. The discussion of how to distinguish species of bacteria? The beetles with male trimorphism? Blue butterfly larvae mimicking the sounds made by queen ants in order to be tended by the deluded worker ants? All of them very cool, and well worth discussion. But lets look at option four.



This little beastie (just under ten centimetres long) is called Schinderhannes bartelsi*, and its fossil remains are described in a paper by Kühl et al. (2009) (from whence comes the above reconstruction). Some of you may immediately recognise the similarity to the famed larger animals Anomalocaris and Laggania of the Cambrian Burgess Shale. However, Schinderhannes bears a few significant differences from those taxa: (1) it has that bizarre pair of 'wings' attached to the back of the head; (2) certain details of its anatomy suggest that it is more closely related to living arthropods than is Anomalocaris, showing that arthropods are descended from an 'anomalocarid' grade; and (3) it doesn't come from the Burgess Shale, but the German Hunsrück Slate, which is from the Lower Devonian, and shows that 'anomalocarid'-type animals were around for some 100 million years longer than we previously knew. I hate to repeat the old cliché about it being like discovering a Tyrannosaurus alive today, and in fact it's not like that, because the amount of time separating Tyrannosaurus from the present is considerably less than 100 million years.

*The name Schinderhannes is apparently derived from that of an 18th century bandit in the area from which it was found. Neat name, but it hints frustratingly at a back story that we are sadly denied in the paper.

Schinderhannes resembles anomalocarids in its radial mouth, and the large pair of spiny pre-oral appendages. However, certain of its features are more like true arthropods - it has a dorsum divided into distinct, sclerotised tergite plates, and it has biramous (two-branched) appendages like crustaceans. The combination of the large 'wings' and 'flukes' on either side of the tail spine suggest that it was an active swimmer.



Large raptorial pre-oral appendages (dubbed 'great appendages') have also been found in a number of Cambrian arthropods such as Leanchoilia and Yohoia. The phylogenetic position of such 'great-appendage' arthropods has been hotly debated. Budd (2002) suggested that they were a stem grade to the arthropod crown clade, but Cotton & Brady (2004) placed them within the crown clade, in the stem group for chelicerates. Researchers have also debated whether the great appendages of these arthropods are homologous to those of anomalocarids, and whether the great appendages are homologous to the chelicerae of modern chelicerates. The (admittedly pretty rudimentary) phylogenetic analysis of Schinderhannes by Kühl et al. (2009), the results of which are shown above, supports a position of great-appendage arthropods as stem chelicerates (despite the great appendages of these arthropods being a priori coded as homologous to those of anomalocarid-grade animals), which supports the comparison between great appendages and chelicerae. It also suggests that trilobites are closer to crustaceans than chelicerates, contrary to the idea of a trilobite + chelicerate "Arachnomorpha" clade. In some regards, this would make sense - trilobites, like crustaceans and insects, have lost the plesiomorphic state of grasping pre-oral appendages as found in chelicerates and have filamentous antennae instead. However, the position of trilobites in the tree above seems to be primarily due to the presence of antennae, so I don't know if it can be considered well-supported.

REFERENCES

Budd, G. E. 2002. A palaeontological solution to the arthropod head problem. Nature 417: 271-275.

Cotton, T. J., & S. J. Braddy. 2004. The phylogeny of arachnomorph arthropods and the origin of the Chelicerata. Transactions of the Royal Society of Edinburgh: Earth Sciences 94: 169-193.

Kühl, G., D. E. G. Briggs & J. Rust. 2009. A great-appendage arthropod with a radial mouth from the Lower Devonian Hunsrück Slate, Germany. Science 323: 771-773.

How to Kill Tropical Fish

A short while ago I was contacted by Ava of The Reef Tank, a site for keepers of tropical marine aquaria, asking if I would contribute something for their in-house blog. I did once keep a tropical (freshwater) aquarium, but sadly the main thing I learnt from the experience was that putting too much fish in the tank early on leads to their rapid demise due to the buildup of toxic nitrogen compounds (eventually, I discovered the attractive yet indestructable White Cloud mountain minnow). So I presented Ava with a piece on the bacteria responsible for maintaining the nitrogen cycle in fish tanks, and you can read it here.

More Evidence of Taxonomy's Importance

Another piece of news that shows the importance of proper taxonomy - Liberia is suffering from an abundance of crop-eating caterpillars, bad enough that the President has declared a state of emergency, and then they discover that they've been spraying for the wrong species of caterpillar.

Hat-tip to Agricultural Biodiversity Weblog for passing on this piece of news.

Oddments

First and foremost, Circus of the Spineless is back! Get your invertebraty goodness at The Other 95%.



Secondly, at some point in the not-too-distant future, I will be composing a post with a line-up of the the most incredible organisms of all time. As part of my preparation for that post, I'd like to hear what your nominations for the title would be. They can be living or fossil, abundant or exotic, animal, plant or otherwise. I only have one little challenge to the question - I'd like individual species. If you think the most incredible organisms are, say, ants or asteroids*, then try to think about what specifically is the coolest ant or asteroid out there. If you could also tell me why your nomination(s) are so cool, that'd be great.

*I'm being deliberately vague about whether I'm referring to asteroids the animals, or asteroids the plants.

Of Gregarines

An assortment of lecudinid eugregarines. The front end of the cell is towards the lower left in each individual. Image from Leander (2007) via Brian S. Leander.

The Sporozoa are perhaps the most famous group of protozoan parasites. As well as being one of the few protozoan groups distinctive enough to be well-characterised prior to the advent of electron microscopy and molecular analysis (albeit with a few hangers-on such as microsporidians and myxozoans that have since been cast aside from the sporozoan community), distinguished by their lack of motile organelles and intracellular invasion of their hosts, sporozoans include the causative agents of such maladies as malaria and toxoplasmosis. In many references, you may find the Sporozoa referred to as Apicomplexa, a name that refers to the apical complex, an organelle at the front end of the cell that is used to invade the cells of the sporozoan's host. However, as the apical complex is also found in some flagellates that are closely related to sporozoans (such as Colpodella), the name Apicomplexa is better used for the larger clade including these taxa while the name Sporozoa is restricted to the nested aflagellate clade (Cavalier-Smith & Chao, 2004).

Sporozoans have been divided into three classes, the invertebrate parasites Gregarinae (or Gregarinea), Coccidia (intestinal parasites of vertebrates) and Hematozoa (parasites of vertebrate blood cells). Phylogenetic analyses have indicated that the basal division in Sporozoa is between the vertebrate-parasitic Coccidia and Hematozoa on one branch (the Coccidiomorpha), and the Gregarinae on the other, though the Gregarinae is less well supported as monophyletic than the Coccidiomorpha and may yet be paraphyletic (Leander & Ramey, 2006; Leander et al., 2006). One notable exception is that the vertebrate parasite Cryptosporidium, previously regarded as a coccidian, may in fact be related to the gregarines or even derived from within them. Not surprisingly, their choice of hosts means that the Coccidiomorpha are by far the better studied of the two clades, while the Gregarinae have kind of been the poor relation. Nevertheless, it is the gregarines that are my focus today.

Gregarines have been divided into three groups, the archigregarines, eugregarines and neogregarines (Leander, 2007), but it seems more than likely that these represent a series of grades, with eugregarines paraphyletic to neogregarines and archigregarines paraphyletic to the eugregarine + neogregarine clade. In contrast to the complex life-cycles of some coccidiomorphs, gregarines have fairly simple life histories with only a single host. Transmission from one host to another is usually via oocysts released with the host faecal matter, but some gregarine oocysts hitch a ride with their host's gametes during copulation as protozoan STDs. Once inside the host, the oocysts hatch out into infective sporozoites that attach to or invade host cells and develop into feeding trophozoites. Some gregarines can reproduce asexually; the majority cannot. Sexual reproduction occurs by two trophozoites joining together as reproductive gamonts and becoming enclosed within a gametocyst; within the gametocyst they will each divide into hundreds of gametes that will fuse to form oocysts, ready for release.


Selenidium sp., showing the wriggling movement. From Leander (2007).

As I said before, the "archigregarines" probably represent the basal grade of gregarines. They are all intestinal parasites of marine invertebrates, and as such have been unfairly condemned as of little interest to anyone. Archigregarines have very similar sporozoites and trophozoites that are vermiform (worm-shaped) and generally move by wriggling (go here for videos of gregarine movement). Some archigregarines have cells with numerous longitudinal folds, others are smooth. Archigregarines also retain the ancestral characteristic of feeding on their host by using their apical complex to pierce the host cell and sucking out its contents.

The 'eugregarines' include the majority of gregarines (at least, the majority of described gregarines), and include parasites of freshwater and terrestrial as well as marine invertebrates. Again, their study has been biased towards those species that are parasites of insects, with the remainder being generally snubbed. Most marine eugregarines have been lumped together as the genus 'Lecudina', with little to unite them other than that they are marine eugregarines (Leander, 2007). Eugregarines differ from archigregarines in having morphologically quite distinct sporozoites and trophozoites. Their cell walls also became very rigid, and they lost the wriggling ability of archigregarines. Instead, eugregarines developed a system of gliding motility, with an actinomyosin skeleton running along the edge of the numerous surface folds. Ancestrally, eugregarines are intestinal parasites like archigregarines, but instead of the active feeding process of eugregarines, eugregarines absorb nutrients from the host through micropores on the cell surface (Leander et al., 2006).


Two conjoined gamonts of the polychaete coelom parasite Pterospora floridiensis. From Leander (2007).

Neogregarines are a derived subgroup of eugregarines with reduced trophozoites that specialise on terrestrial hosts (mostly insects) and mostly live in non-intestinal tissues. Another group of eugregarines, the urosporidians, became parasites of their host's coelom. Urosporidians lost their direct attachment to host cells, and became free-floating within the tissue as united gamont pairs. The gliding motility and longitudinal folds of other eugregarines were lost, and instead urosporidians move by pulsations of the cell walls. Cells became branched - many are V-shaped with two primary branches that each divide distally into a number of smaller "fingers".

Archigregarines in particular retain a number of features that are believed to be ancestral for sporozoans as a whole (such as sucking feeding), and Leander et al. (2006) suggest that they may constitute the ancestral group not just for eugregarines, but also for coccidiomorphs and hence sporozoans as a whole. Interestingly, gregarines (including archigregarines), so far as is known, lack the residual plastid found in coccidiomorphs. It is currently a subject of some debate as to whether the sporozoan (really coccidiomorph) plastid is homologous with that found in the related dinoflagellates or not (see here for an earlier take of mine on the issue), and the position of the plastid-less 'archigregarines' could have significant implications for this debate.

REFERENCES

Cavalier-Smith, T., & E. E. Chao. 2004. Protalveolate phylogeny and systematics and the origins of Sporozoa and dinoflagellates (phylum Myzozoa nom. nov.) European Journal of Protistology 40: 185-212.

Leander, B. S. 2007. Marine gregarines: evolutionary prelude to the apicomplexan radiation? Trends in Parasitology 24 (2): 60-67.

Leander, B. S., S. A. J. Lloyd, W. Marshall & S. C. Landers. 2006. Phylogeny of marine gregarines (Apicomplexa) — Pterospora, Lithocystis and Lankesteria — and the origin(s) of coelomic parasitism. Protist 157: 45-60.

Leander, B. S., & P. A. Ramey. 2006. Cellular identity of a novel small subunit rDNA sequence clade of apicomplexans: description of the marine parasite Rhytidocystis polygordiae n. sp. (host: Polygordius sp., Polychaeta). Journal of Eukaryotic Microbiology 53 (4): 280-291.