More From the "They Don't Write Papers Like They Used To" Files

But before that, a couple of other things:

Recent Carnivals - Berry Go Round is here, Circus of the Spineless is here.

Also recently, the Open Laboratory 2008 collection of some of the year's best online science writing is now available. In the interests of disclosing all interests (naturally), yours truly has a piece featured in this publication - on citrus fruit. Anybody who has heard me commenting on my poor knowledge and outright terror of botany will know that this is a Very Funny Thing.

On to the main point of the post, which will not be anything I'm writing at all...

Recently, I was reading a paper from 1866 by a Dr H. Dohrn with the unprepossessing title of "Synopsis of the birds of Ilha do Principe, with some remarks on their habits and descriptions of new species" (Proceedings of the Zoological Society of London 1866: 324-332). Principe, for those who don't know it, is a part of the country of Sao Thomé and Principe in west-central Africa. This is the second paragraph of that publiction:

It seems to me that the most remarkable feature in the fauna of Ilha do Principe is that not a single bird of prey exists on the island, whilst they are abundant on the two other islands and on the nearest part of the continent. I saw hundreds of Milvus parasitus in San Thomé; Gypohierax angolensis and some other species are not uncommon in Fernando Po; but the whole tribe avoids Principe. The inhabitants of the latter place and of San Thomé assert that there is a deadly hatred between the Grey Parrots (Psittacus erythacus) of Principe and the Kites of San Thomé, and that, if ever a Milvus visits the neighbouring island, hundreds of Parrots fall upon him and kill him, and that the Kites take revenge if perchance a Parrot should venture a trip to their kingdom. There must be some family reason for this strange degree of enmity, for they seem to live in tolerable peace together on the coast.

You will probably not be surprised to hear that I was somewhat taken aback by this - even allowing for the different time period, it seemed a rather credulous account. But as I read further, I realised that Dohrn was employing something that is a little alien to modern academic writing: irony.

To demonstrate my point, here are my other favourite passages from Dohrn:

On a species of drongo:

The native name is "Maria Palu, feiticeira" (translated, Maria Palu, the sorceress). The bird is black, with red eyes; seems very indolent in daytime, and shows a great ability in the imitation of some other birds' voices. Of course there must be some "feiticeira" to it: therefore, sitting on the roof of a house and singing in its melancholy manner, it prophecies the death of one of the inhabitants; and this, of course, takes place, but often a long time after this prophecy.

On the bald ibis:

Soon after my arrival on the island I was informed by some natives that there was a very remarkable bird in the island called "Corvão." One told me that it was a kind of raven with splendid metallic wings; another described the bird "with the head of an owl and the feet of a duck, climbing up and down trees;" and others gave me other extravagant descriptions of it; but all of them agreed that the bird lived in almost inaccessible rocky and wooded localities of the southern district, and that if ever a specimen passed over the town it was a bad omen for the white inhabitants, who in such case were exposed to heavy disease or death. Of course I was very curious to see this species, and settled for a fortnight in a negro's hut in those desert parts of the island*. Whoever has visited those large tropical forests knows the difficulty of proceeding there. I enjoyed the special favour of heaven in arriving there when the rains set in a month before they usually do, and it was very hard work to run after these birds. I saw them daily at great distances, and heard them crowing like a Raven; but as soon as I entered the forests the monkeys made so much noise, barking and howling, as to alarm all the animals in the neighbourhood. Thus I was finally very glad when one of my native hunters appeared with a female specimen of the Corvão, which turned out to be Geronticus olivaceus.

*I think in this case "desert" means "deserted", not "arid".

The Uglier Side of the Family (Taxon of the Week: Ceratomorpha)

Lowland tapir, Tapirus terrestris, in a blatant attempt to exploit the cute factor. Photo by Antonio Pinheiro.

The modern perissodactyls are, sadly, but a shadow of their former glory. Once among the planet's dominant herbivores, the odd-toed hoofed mammals have become reduced to less than twenty living species (the majority of which are critically endangered to boot). Nevertheless, their secure position as charismatic megafauna means that they are familiar animals to most people (at least conceptually). One species in particular, the horse Equus caballus, has developed a close association with humanity and holds a high position in the human psyche (or at least the Eurasian and American psyche). But this post won't be dealing with horses - this is for the other side of the perissodactyls. The Ceratomorpha may not have been blessed with the aesthetic appeal of the horses, but they're not without their charms.


Skull of Tapirus terrestris. Note the severely recessed nasal bones, positioned above the eyes, that indicate the presence of the muscular trunk in life. Tapirs are often thought of as more "primitive" than other perissodactyls, but no other perissodactyl has a skull like that. Photo by Matthew Colbert.

The Ceratomorpha contains two living families, the Tapiridae (tapirs) and Rhinocerotidae (rhinoceroses). Most of the people reading this will, I'm guessing, probably be familiar with the appearance of both, though rhinos do get given a little more press than tapirs*. Rhinos are also marginally more diverse in the modern environment, with five species to the tapirs' four - but considering that at least two of the rhino species are hovering on the brink of extinction, that may yet change. As regards fossil taxa, the limits of Ceratomorpha are a little more hazy, mostly because different authors have applied slightly different concepts for 'Ceratomorpha' vs. the related name 'Tapiromorpha'. In my opinion, the most sensible definitions for both are those proposed by Holbrook (1999), who used 'Ceratomorpha' for the crown clade of tapirs + rhinos, and 'Tapiromorpha' for the total group of anything more closely related to tapirs and rhinos than to horses. These definitions are better than the alternatives proposed by Froehlich (1999) in being agnostic as to whether the extinct chalicotheres are tapiromorphs (as supported by Froehlich, 1999, among others) or not (as indicated by, e.g., Hooker & Dashzeveg, 2004, who placed chalicotheres outside the perissodactyl crown group).

*Completely unrelated aside to everything else - the Japanese name for 'tapir' is 'baku'. Originally, a baku was a trunked mythical creature that was supposed to feed on people's dreams (particularly nightmares), but living tapirs have a somewhat more material diet. This isn't the only example in Japanese of a living exotic animal being granted the name of a pre-'existing' mythical creature: giraffes are known as 'kirin'.

Within the Ceratomorpha, then, the primary division is between the Tapiroidea and the Rhinocerotoidea, each including (obviously) the taxa more closely related to one of the living families than the other*. The Rhinocerotoidea are far better understood that the Tapiroidea, which have a more spotty fossil record. Four families are generally assigned to the Tapiroidea - Helaletidae, Lophialetidae, Deperetellidae and Tapiridae. The mostly Eocene 'Helaletidae', small tapiroids of North America and Eurasia, however, are probably paraphyletic with regard to other tapiroids. Holbrook (1999), for instance, excluded the North American genus Heptodon from the Helaletidae as he found it to be sister to all other tapiroids. At least one 'helaletid' genus, the North American Oligocene Colodon, is of interest because it possessed significantly retracted narial opening, indicating the presence of a trunk as in modern tapirs. However, Colbert (2005) included Colodon within the Tapiridae, closer to modern tapirs than previous authors.

*Be warned, though - older references tend to use the term 'tapiroid' as a grade concept for non-rhinocerotoid tapiromorphs, including a number of taxa that would be regarded as stem-Ceratomorpha in this post.


Hyrachyus, an Eocene rhinocerotoid. Hyrachyus was very similar to the tapiroid 'Helaletidae' of the same time period (which is pretty much what one would expect, really) and represents the general basal morphology for Ceratomorpha. Picture from here, though it has a definite Zdenek Burian look about it.

The Lophialetidae and Deperetellidae were two strictly Asian late Eocene families that suffer from a severe lack of study. Both have been mostly regarded as tapiroids for as long as they have been known, but Holbrook (2001) pointed out that the evidence for doing so is pretty slight. Members of both families (supported as forming a monophyletic clade by Holbrook, 1999) showed a reduction in the number of toes from four to three and a longer, more slender foot than other tapiroids, indicating that they were more cursorial (Radinsky, 1969). The Eocene also saw the appearance of the first Tapiridae, which (with the possible exception of Colodon, depending on its position) seem to have been the only tapiroids to survive into the Oligocene. The earliest tapirid genus, Protapirus, has been described from both North America and Eurasia, but its monophyly is uncertain (Colbert, 2005).


Assortment of Amynodontidae as drawn by Stanton Fink - front to back, they are Cadurcodon, Gigantamynodon and Metamynodon. This isn't the first time I've used one of Stanton's drawings - other ones are here and here. I must confess to having something of a love-hate reaction to Stanton's work - his style isn't entirely to my taste aesthetically, but one thing I do think is great is how often he reconstructs animals that other artists just ignore. Besides, I can't draw to save myself, so I have absolutely no right to criticise.

The other superfamily, Rhinocerotoidea, has had a much greater diversity described, both in terms of number of species and morphological range. Three major families have been recognised - the Amynodontidae, Hyracodontidae and Rhinocerotidae. The middle Eocene to Middle Oligocene Amynodontidae were the really unfortunate members of the superfamily appearance-wise - as the saying goes, they would have not only been hit with the ugly stick, they would have fallen out of the ugly tree hitting every ugly branch on the way down. Amynodontids have mostly been characterised as subaquatic, like modern hippos, but Wall (1998) points out that only a single derived subgroup, the Metamynodontini, shows adaptations for such a lifestyle. The remaining amynodontids would have been terrestrial. One group of amynodontids, the Cadurcodontini, appears to have convergently evolved a short trunk like that of the tapirids.


A rather disgruntled looking Hyracodon, reconstructed by Heinrich Harder.

The Eocene to Oligocene Hyracodontidae are the real taxonomic dog's breakfast of the Rhinocerotoidea. Radinsky's (1966) influential definition of which taxa should be included in Hyracodontidae essentially amounted to "anything which doesn't belong to Amynodontidae or Rhinocerotidae". Characters that have been suggested to support a monophyletic Hyracodontidae, such as an elongate foot and three toes, are also found in other perissodactyls (in fact, if you look upwards you'll notice that I mentioned the exact same characters in connection with Lophialetidae). In general, hyracodontids were more cursorial than other rhinocerotoids, and small genera such as Hyracodon would have been fairly pony-like. Also usually included in the 'Hyracodontidae' where the gigantic Indricotheriinae, of which the central Asian Oligocene genus Paraceratherium is famed as the largest known land mammal*. The phylogenetic analysis of Holbrook (2001) supported treating the smaller Hyracodontinae as a separate family from the larger Indricotheriinae, but failed to support the latter as monophyletic.

*Though sometimes in disguise. Paraceratherium has held a few different names over the years - 'Indricotherium' and 'Baluchitherium' are two of the most commonly used. It remains a rather fraught question as to whether all these represent the same animal, or a number of closely related animals.


A herd of Paraceratherium (would Paraceratherium have really travelled in herds?) passing by scavenging Hyaenodon. Painting by Mauricio Antón.

And finally, the Rhinocerotidae. But I think I've rabbited on and wasted enough of everyone's time for today, so I guess I'll have to leave the hippo-like Teleoceras, or the double-horned Menoceras (two horns side by side, that is), or the gigantic Elasmotherium (with what must have been one of the most terrifying pieces of headgear this side of Arsinoitherium) for another time.

REFERENCES

Colbert, M. W. 2005. The facial skeleton of the Early Oligocene Colodon (Perissodactyla, Tapiroidea). Palaeontologia Electronica 8 (1): 8.1.12A.

Froehlich, D. J. 1999. Phylogenetic systematics of basal perissodactyls. Journal of Vertebrate Paleontology 19 (1): 140-159.

Holbrook, L. T. 1999. The phylogeny and classification of tapiromorph perissodactyls (Mammalia). Cladistics 15 (3): 331-350.

Holbrook, L. T. 2001. Comparative osteology of early Tertiary tapiromorphs (Mammalia, Perissodactyla). Zoological Journal of the Linnean Society 132 (1): 1-54.

Hooker, J. J., & D. Dashzeveg. 2004. The origin of chalicotheres (Perissodactyla, Mammalia). Palaeontology 47 (6): 1363-1386.

Radinsky, L. B. 1966. The families of the Rhinocerotoidea (Mammalia, Perissodactyla). Journal of Mammalogy 47 (4): 631-639.

Radinsky, L. B. 1969. The early evolution of the Perissodactyla. Evolution 23 (2): 308-328.

Wall, W. P. 1998. Amynodontidae. In Evolution of Tertiary Mammals of North America: Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals (L. L. Jacobs & K. M. Scott, eds.) pp. 583-588. Cambridge University Press.

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.