PROTORICHTHOFENIOID BRACHIOPOD FROM URALS 
which were attached and retained spines. While some 
stratigraphically younger examples of this aspinose stock, in 
addition to those from northern Spain, may yet be discovered, it 
seems it did not persist after the Kazanian, when the Caucasian 
species of Likharev died out. The Permian genus Col//umatus 
would appear not to have been derived from this stock, but to be 
a true richthofenioid which had also lost its spines. A feature of 
taxonomical importance in the specimens described by 
Sutherland (1989) from Oklahoma, but not seen in this new 
material, is the juvenile ventral interarea. A small ventral 
interarea is also reported in the Teguliferininae (Muir-Wood & 
Cooper, 1960), which belongs in the Richthofenioidea. It is the 
presence of interareas in these Carboniferous species that places 
the Richthofenoidea in the Strophalosiidina, the structure being 
lost in the more common Permian richthofenioids. In Zalvera it 
seems the earliest growth of the ventral valve was normal for 
productids, but whether it involved the growth of a juvenile 
interarea is unknown. If present it might have become 
incorporated into the later subconical valve growth. In any 
event, once growth changed, after the initial 4-S5mm, from being 
a small finely spinose subtriangular valve to a rapidly expanding 
rounded cone with a spineless exterior, shell growth was 
holoperipheral. In the North American species there was a short 
period when the ventral interarea grew before the posterolateral 
mantle margins grew posteromedianly to continue the style of 
shell secretion seen for the rest of the valve. This left the juvenile 
interarea preserved, but with a short suture line posteromedianly 
where the two mantle margins had grown together and fused, 
allowing the typical teguliferinid cone to grow. In the Urals 
material this posterior fusion of the mantles occurred earlier in 
growth so that no interarea has been preserved. This raises the 
question as to whether Zalvera should be assigned to the 
Strophalosiidina. The alternative is to consider Za/vera within 
the Productidina as a unique aberrant offshoot showing 
tendencies towards the morphology of the Richthofenioidea. 
This seems less likely than placing Zalvera in the 
Richthofenioidea and accepting that the interarea never 
developed in these unusually-shaped brachiopods. It is hoped 
that more of this material will become available so as to allow 
more complete preparation and a better insight into the way in 
which this strange brachiopod grew. 
There is no clear evidence for the origin of the Zalveridae, but 
it is possible to suggest that the ancestral stock was within the 
Strophalostidina. The cone development seems to have been 
from exaggerated, and posteriorly fused, growth of the ventral 
59 
trail. Within strophalosiidines the Aulostegoidea includes many 
groups with elaborate trails, and as they lack a toothed 
articulation and have variably developed interareas, they provide 
possible ancestors for Zalvera. 
ACKNOWLEDGEMENTS. I thank Dr S. Lazarev, Palaeontological 
Institute, Moscow, for information about this material and other 
specimens in Russian collections; Dr Cor Winkler Prins, National 
Museum of Natural History, Leiden, for the loan of comparable 
material from Spain and Austria; Dr Patrick Sutherland, University of 
Oklahoma, for information about his specimens, and Dr R. Cocks for 
reading and commenting constructively on a draft version of this paper. 
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