Umbilia gazing - part II

We last left off our survey of Umbilia in the middle Miocene where we looked at U. eximia the most abundant and widespread species.
Two of the remaining four described middle Miocene species are some of the weirdest of all crown-group cowries (I say crown group because there were some truly bizarre looking stem-group cowries, e.g. Gisortia).

U. siphonata (above) is one of these. It is a very large cowry, attaining a length of almost 17 cm, which is not far behind the biggest specimens of Macrocypraea cervus, the largest extant cowry, which used to reach sizes of 19 cm in length. However U. siphonata is cheating a little since the anterior and posterior rostra of this species are produced into great upwardly curving ‘horns’. The rudimentary flanges that support the bases of each rostrum of U. eximia are much better developed in this species. Even stranger is U. gastroplax, the ‘flanged cowry’ (specimen on the left is from Darragh 2002). This species also has elongated horn like rostra, although they are not as long as in U. siphonata. However the flanges have expanded outwards, merging together and making a continuous brim that encircles the entire base of the shell. The result looks much like a snow-shoe. Indeed it has been suggested that this is exactly what its function was and that it was an adaptation to living on soft, ‘soupy’ bottoms. One wonders then if U. siphonata was adapted to the same conditions but simply didn’t bother to keep itself on top of the sediment surface, and used its long rostra to carry its siphons up into clear water.
The fourth middle Miocene Umbilia we will look at is little U. leptorhyncha (below). Although common and widespread, good specimens are rare due to the thin-shelled fragility of this species. This species is a departure from the other mid Miocene species in its small size, globose shape and poorly developed rostra. In these respects it most closely resembles U. prosila and may be closely related to it.

All of these species can be found in both South Australia and Victoria (U. gastroplax has not been officially recorded from South Australia but I have personally collected two specimens from the Cadell Formation on the banks of the River Murray)
Darragh recorded the extant U. hesitata as a fifth middle Miocene species, albeit one that only appears at the end of the stage, with little to no time overlap with the previously mentioned species. The taxonomy of these rare later middle Miocene Umbilia is a complicated issue. Two species have been named Umbilia tatei and Umbilia cera. Both are short , with weakly developed beaks and heavily calluses surrounding the basal margins and probably represent the same species whatever they are. Problematically if these really are small specimens of the extant ‘wonder cowry’ (as U. hesitata is sometimes called) then we have the problem that U. tatei would have priority over U. hesitata. I’m sure all the avid cowry collectors would object to replacing the entrenched U. hesitata with U. tatei. Fortunately I don’t think they have to. Although U. hesitata does display a range of adult sizes which overlaps with the small shells of U. tatei, small modern U. hesitata resemble typical large specimens more than they do U. tatei. In particular no modern U. hesitata has such thick marginal calluses as U. tatei, nor do they develop the elongate coarse dentition seen on the holotype of U. cera (these are not present in the types of U. tatei but the dentition of these specimens appears to be underdeveloped due to immaturity).

A late Miocene U. 'hesitata', probably belonging to U. tatei.

From the late Miocene and Pliocene (there is no Pleistocene record of Umbilia at all) there is just a single species, the extant U. hesitata (although some of these are a little different from modern U. hesitata while others probably belong to U. tatei). Then in our modern seas we find five species: U. hesitata, U. armeniaca, U. capricornica, U. orriettae and U. petillirostris.
However unlike the middle Miocene where you can find up to four species at the same locality almost all of the modern species have mutually exclusive ranges (only U. capricornica and U. petillirostris overlap in the deep Capricorn Channel of the Great Barrier Reef. Moving anticlockwise around the Australian coast we find U. armeniaca (Western Australia to Kangaroo Island, South Australia), U. hesitata (south eastern South Australia to Southern Queensland), U. orriettae (Moreton Bay, Queensland) and U. capricornica/U. petillirostris on the Great Barrier Reef. It is interesting to note that this pattern matches the phylogenetic pattern recovered in a comprehensive molecular phylogenetic analysis of modern cowries (Meyer 2004). In this analysis U. armeniaca was the sister group to all other living species and U. hesitata was the sister group to U. capricornica + U. petillirostris (U. orriettae was not included but morphologically it appears to be intermediate between U. hesitata and U. capricornica). Thus the modern forms appear to be the result of a radiation that proceeded from west to east. All of these living species are rather similar to one another and have a rather generalised shell structure compared to the excesses of the middle Miocene.

Umbilia hesitata, the most abundant extant species.

Most specimens display a moderately well-developed posterior rostrum and have highly reduced anterior tubercles that are separated by an oblique sulcus. These characters suggest that the modern taxa are more closely related to U. eximia and U. tatei than any of the other extinct species. However the type specimens of U. petillirostris stand out as something unusual. Unlike other living Umbilia the types of U. petillirostris are globose and thin-shelled with a very short posterior rostrum. In these respects U. petillirostris resembles the smaller middle Miocene species, U. leptorhyncha and the late Oligocene U. prosila. Darragh (2002) suggested that these three species represented a lineage that had been separate since the Oligocene. But using the molecular phylogeny this would suggest that all of the modern species have been separate since at least the late Oligocene, despite sharing a similar hesitata-like morphology that does not show up in the fossil record until the late Miocene. However I strongly doubt that U. petillirostris is closely related to U. prosila and U. leptorhyncha. Despite its globose shape and thin shell it displays characters typical of the modern clade such as large size, a moderately produced posterior beak, and weak anterior tubercles. A greater sample of specimens shows that U. petillirostris and U. capricornica are quite variable and that individuals of each species can be found that approach the other in morphology. Indeed some have suggested that the two species are not distinct at all (Wilson and Clarkson 2004). Nevertheless limited genetic sampling does indicate that U. petillirostris does maintain a distinct haplotype (Meyer 2004). Excellent photographs of all the living forms can be seen here.

Next week: the origin of Umbilia.

References

Darragh, TA (2002) A revision of the Australian genus Umbilia (Gastropoda: Cypraeidae). Memoirs of the Museum of Victoria 59: 355-392.

Meyer CP (2004) Toward comprehensiveness: increased molecular sampling within Cypraeidae and its phylogenetic implications. Malacologia 46: 127-156.

Wilson B, Clarkson P (2004) Australia's Spectacular Cowries: A Review and Field Study of Two Endemic Genera-Zoila and Umbilia. Odyssey: El Cajon, 369 pp.

Lookin' out my back door

Well out my back window actually. We've been watching this industrious little guy for almost three years now. He is a masked weaver (Ploceus velatus), and in order to attract a mate he has to construct a nest of sufficient quality to encourage a female to lay her eggs in it. Sadly he's a bit of a loser. Although several females have checked him out he has never been able to seal the deal. After every rejection he would demolish the nest and start again. Now however it appears he is so riddled with frustration and self-doubt that he just builds and destroys nest after nest without even getting it looked at first. I kind of empathize with him, my early years (15 through to 26) weren't too dissimilar.

More sauropod vertebrae/ ceratopsian frill convergence

I was reminded by Mike Taylor's recent post, noting that a Camarasaurus vertebrae seems to have a ceratopsian frill growing out of it, that I had had the exact same thought when I saw 'Max'. Max is a diplodocid (identified as Apatosaurus but I have my doubts) found by the crew at the Saurier Museum in Aathal. However this time the 'frilloid' process is composed of the two postzygapophyses and the perforate interpostzygapophyseal lamina. Incidentally the interpostzyg laminae of most of Max's cervicals are similarly perforate. It is a real feature, not caused by damage - weird huh?

The last dicynodont

With so much going on I've had little time for blogging. Recently there was some discussion of the supposed Australian Cretaceous dicynodont (maxillary fragment of the specimen is pictured on the left) over at Chinleana. I'll add my two cents here rather than commenting there just to keep something ticking over on my blog. Randy Irmis made a startling comment that the consensus was that it was indeed a dicynodont. This is news to me, I had always thought that the identity of the specimen was always the weak part of the claim. I have to add that I've never seen the specimen myself. Randy goes on to add that because it was surface float it is the provenance of the specimen that is suspect. Here I have to add my voice in support of Thulborn's original assesment, whatever it was there can be little doubt that it came from the Cretaceous. As has been noted Australia is flat and rather geologically quiescent. The nearest Triassic rocks are many hundreds of kilometres away. Nor do these Triassic rocks have much in the way of dicynodonts in them anyway - just one beat-up quadrate from more than 20 years of intensive collecting in the Arcadia formation (the main fossil-bearing Triassic formation of south-eastern Queensland). When you are out prospecting in most parts of the world you almost never find fossils more than a few tens of metres from the formation that bore them (unless there is a transport mechanism such as a river). Australia certainly never had post-Triassic glaciations that can randomly transport objects over large distances. So if the morphology is definately saying dicynodont then hey, I'm prepare to accept this extraordinary claim. Indeed recently another clade thought to have died out before the end of the Triassic has been found to have survived until the Cretaceous (I can say no more) so survival of the Dicynodonts may not be so weird after all.

The lizard biters

The answer to the puzzle is that both were named Saurodectes, meaning ‘lizard biter’. One is an insect from the Early Cretaceous of Siberia, while the other is an early Triassic procolophonoid parareptile found in South Africa. I was part of the team that found the procolophonoid although I can’t claim that I found the specimen. In fact I found precious little while Ross Damiani found 'Saurodectes' despite being severely hampered by a broken ankle. Some people are just gifted when it comes to field-work and I am not one of them. The insect has priority over the name and we had to rename our procolophonoid Saurodektes (Modesto et al. 2004). Not that there is much shame in proposing a name that is preoccupied by an arthropod. With so many arthropods it seems to happen all the time. I got my first look at the real Saurodectes when I purchased Grimaldi and Engel’s magnificent tome ‘Evolution of the Insects’. And what a fascinating insect it is.
Described as a kind of louse, Saurodectes vrsanskyi (Rasnitsyn and Zherikhin 1999) has an unusual set of characters. Some of these such as the single claw at the end of its legs, short, widely spaced legs and membranous distensible abdomen are typical of ectoparasitic insects but the very large eyes and lack of spiny setae are not. What those handle-bar like structures sticking out its head are is anybody's guess. A recent survey of fossil lice could not find any convincing characters that definately placed the fossil amongst the lice (Pthiraptera) but could not suggest any alternative relationships either (Dalgleish et al. 2006).


The head of Saurodectes, from Grimaldi and Engel (2005).

The original describers (Rasnitsyn and Zherikhin, 1999) suggested that it was a pterosaur parasite on the basis that it was too big at 17 mm to parasitise Mesozoic mammals but had single clawed feet like modern mammal lice. This is seen as an adaptation to gripping hair shafts, and that since pterosaurs were also hairy then it was supposed that Saurodectes plied its way through ptero-fuzz. However I don’t see a close correspondance between the claws of modern mammal lice and those of Saurodectes. Actually this is not the only weird Mesozoic insect that has been claimed to be a pterosaur parasite. Sauropthirus longipes, a scorpionfly relative from the same formation as Saurodectes has also been hypothesized to have found its living on pterosaurs. Actually the stiff, backwardly pointed spines and eyelessness of this insect seems to be more fitting with this kind of lifestyle. It is, of course, not impossible for both of these to be pterosaur parasites but it strikes me that some of the odd features of Saurodectes may be explicable if it lived on the scaly hide of a large non-feathered dinosaur. In modern lice there is a loose correlation between parasite size and host size indicating that Saurodectes had a large host. Furthermore backwardly pointing setae may be of little use on a host that lacks filamentous integument. Eyes may also be of use to a large, exposed ectoparasite, not sheltering under a dense pelt of hair. Whatever the habits and relationships of Saurodectes, there can be little doubt there must have been hordes of parasites making a living off of dinosaurs that we have yet to learn about.

References

Dalgleish RC, Palma RL, Price RD, Smith VS (2006) Fossil lice (Phthiraptera) reconsidered. Systematic Entomology 31: 648-651.

Grimaldi D, Engel MS (2005) The evolution of Insects. Cambridge University Press.

Modesto SP, Damiani R, Neveling J, Yates AM (2004) Saurodektes gen. nov., a new generic name for the owenettid parareptile Saurodectes Modesto et al., 2003. Journal of Vertebrate Palaeontology 24: 970.

Rasnitsyn AP, Zherikhin VV (1999) First fossil chewing louse from the Lower Cretaceous of Baissa, Transbaikalia (Insecta,Pediculida ¼ Phthiriaptera, Saurodectidae fam. n.). Russian Entomological Journal 8: 253–255.

Puzzle Time: What's the connection?

Yes, see if you can guess the connection between these two fossils.

The Drakensburg Lavas and the First Great Dinosaur Dying

What you are looking at is a thick pile of basalt, that was extruded onto the Earth’s surface some 183 million years ago, during the latter part of the Pliensbachian stage (or at the Pliensbachian-Toarcian boundary, depending on whose timescale you follow) of the Early Jurassic. They are part of a 2 km thick sheet that is centred on the mountainous nation of Lesotho in Southern Africa. They take their name, the Drakensburg Group, from the Drakensburg Range, a ragged row of peaks said to resemble the back of a dragon that runs along the border of Lesotho and the South African province of Kwazulu-Natal. This large pile of basalt is an erosional remnant of a truly enormous volcanic province. Other parts of what was once a continous sheet of lava extend north to Zimbabwe, Botswana and Zambia, and westward into Namibia. What is even more jaw dropping is that if Gondwana is reassembled, then these southern African lavas (generally called the Karoo flood basalts) are just part of one enormous province that extends into the Southern tip of South America (the Chon Aike Province) and across Antarctica (the Ferrar Province) and into southern Australia. Taken together the total volume of magma that was either extruded onto the surface, or emplaced as intrusions below it, would come to more than two and a half million cubic kilometres (Wignall 2001). This volume actually exceeds the estimated original volume of the famous Deccan Traps of India, which were extruded at the end of the Cretaceous (when most dinosaur lineages famously kicked the bucket). Given that these vast volcanic outpourings seem to be linked with mass extinction events with disturbing regularity it seems odd that the truly enormous Karoo-Ferrar province is not linked to a big extinction event – or is it?
An extinction event amongst marine molluscs (yay! see molluscs have much to teach us!) in the late Pliensbachian has been recognised in Europe and South America and this has been tied to the Karoo-Ferrar eruptions (Hallam 1961, Aberhan and Fürsich 1996). But the general consensus is that this was a weak mass extinction, well below the level of the ‘big five’ mass extinctions.
But how sure can we be? One thing that is clear to dinosaur aficionados is that the early Middle Jurassic has an abysmal record of terrestrial faunas and this may well be masking the effects of a terrestrial mass extinction. Indeed the first stage of the Middle Jurassic Epoch, the Aalenian, is the only Mesozoic stage that does not have its own valid, diagnostic dinosaur taxon (or at least it didn’t a few years ago, maybe there is one now). Another thing that the dinosaur record shows is that prior to the middle Jurassic, dinosaur faunas were rather uniform the world over with a community structure dominated by basal sauropodomorphs (usually a massospondylid) with small coelophysid and larger dilophosaurids representing the theropod contingent and much rarer small basal ornithischians. This type of fauna can be found in Southern Africa (Elliot, Clarens and Forest Sandstone Formations), North America (Kayenta, Navajo and Portland Formations), Antarctica (Hanson Formation) and China (Lower Lufeng Formation). It is interesting that the two dominant components of this faunal association, the basal sauropodomorphs and the coelophysids are basically holdovers from the Triassic. However once the record picks up again higher up in the middle Jurassic things have changed a great deal. Gone are the coelophysoids and basal sauropodomorphs*. In their place we find ceratosaurs and tetanurans filling the large predator niches while eusauropods and eurypods (that is ankylosaurs and stegosaurs) occupy the large herbivore niches. This combination of taxa remained dominant around the world to the end of the Jurassic. So was this turnover a gradual affair? Maybe not, and I have suggested that there was actually a terrestrial mass extinction event that cleared away the coelophysoids and basal sauropodomorphs in my so far unpublished chapter in the upcoming Complete Dinosaur II. If so, it would seem very likely that this event was the same one that killed those poor little clams in the late Pliensbachian. In other words the Drakensburg and associated lavas really were significant for dinosaur evolution. Perhaps without them we may never have got such majestic beasts as Apatosaurus and Brachiosaurus. Quite independently Ronan Allain and Najat Aquesbi came to the same conclusion in their monograph on Tazoudasaurus, which I featured here. Ronan and myself must think alike for this isn’t the first time we’ve come up with the same idea more or less simultaneously. Earlier we both published the connection between the dating of the Karoo-Ferrar volcanics and the age of Vulcanodon at more or less the same time (Allain et al.2004, Yates et al. 2004).
Nevertheless there exists an alternative explanation. A team of French geologists led by Fred Jourdan have suggested that the late Pliensbachian extinction event was really mild because the Karoo-Ferrar basalts were extruded over an extended 8 million year period (Jourdan et al. 2005). Other continental flood basalt provinces show a pattern where 90% or more of their volume is extruded in a brief spell of less than 600 000 years. Jourdan et al. clearly demonstrated that the lavas to the north of South Africa were extruded over a period extending from 182 to 177 million years ago. Does this spell the end of the late Pliensbachian dinosaur extinction hypothesis? Perhaps but I’m not ready to discard this idea just yet. Note that the long duration of eruptions is restricted to regions north of South Africa. The Drakensburg (an erosiaonal remnant of a truly vast area shown by the intrusions that riddle the rest of the Karoo Basin) still yields a tight cluster of dates, while palaeomag indicates the whole pile experienced just one magnetic reversal (Duncan et al. 1997). What we need is a comprehensive sampling of the Antarctic, South American and Australian lavas to see whether they also extruded rapidly at the same time the Drakensburg lavas were extruded.

*There is one recorded Middle Jurassic basal sauropodomorph, Yunnanosaurus youngi, but I would like to see a better stratigraphic control on its age.

references

Aberhan M, Fürsich FT (1997). Diversity analysis of Lower Jurassic bivalves of the Andean Basin and the Pliensbachian-Toarcian mass extinction. Lethaia 29: 181-195

Allain R,Aquesbi N, Dejax J, Meyer CA, Monbaron M, Montenat C, Rechir P, Rochdy M, Russell DA and Taquet P (2004). A basal sauropod dinosaur from the Early Jurassic of Morocco. Comptes Rendus Palevol 3(3):199-208

Duncan RA, Hooper PR, Rehacek J, Marsh JS, Duncan AR (1997) The timing and duration of the Karoo igneous event, southern Gondwana. Journal of Geophysical Research 102 (B8): 18127-18138.

Hallam A (1961). Cyclothems, transgressions and faunal change in the Lias of North West Europe, Transactions of the Edinburgh Geological Society 18: 132–174.

Jourdan F, Féraud G, Bertrand H, Kampunzu AB, Tshoso G, Watkeys MK, Le Gall B (2005). The Karoo Large Igneous Province: brevity, origin and relation with mass extinction questioned by new 40Ar/39Ar age data. Geology 33: 745-748.

Wignall PB (2001) Large Igneous provinces and mass extinctions. Earth Science Reviews 53: 1-33.

Yates AM, Hancox PJ, Rubidge BS (2004). First record of a sauropod dinosaur from the upper Elliot Formation (Early Jurassic) of South Africa. South African Journal of Science 100: 504-506.