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20 EUROPEAN ARACHNOLOGY 2005 hypothesis that they are stem-lineage chelicerates is correct we would still expect to fi nd some ‘missing links’ bridging this morphological gap, whereby the more anterior limbs lose the exopod and become primarily adapted for walking while the posterior ones are either lost completely or modifi ed into plate-like, gill-bearing opercula. The enigmatic Silurian fossil Offacolous kingi ORR et al., 2000 might be such a missing link. These authors reconstructed this probable early chelicerate from computer images of serial sections through the nodules which encase them and discovered that most of its prosomal appendages are still biramous. This suggests a more basal grade of organisation than xiphosurans which have only one biramous limb pair (the 6th ) bearing the fl abellum (Fig. 12). Interestingly, the ‘great-appendage’ arthropods lack clear tagmosis into a prosoma and opisthosoma and express a head region probably bearing four pairs of appendages, including the raptorial pair. This ‘head’ in the chelicerate stem-lineage associated with four limb pairs is potentially segmentally homologous with the cephalosoma of pycnogonids (VILPOUX, WALOSZEK 2003, see also above), to the propeltidium of the carapace in some arachnids and the proterosoma region characteristic for many mites (see also DUNLOP, ARANGO 2005, fi g. 5). Thus the ‘great-appendage’ hypothesis might alter interpretations of polarity for a number of arachnid characters. A Final Word: Sanctacaris Finally, one of the most famous candidates for the oldest chelicerate is Sanctacaris uncata BRIGGS, COLLINS, 1988 from the Burgess Shale; a fossil which has entered the popular literature (e.g. GOULD 1989) as an arachnid ancestor. In the original description it was specifi cally referred to Chelicerata on account of: (1) a head with at least six pairs of appendages, (2) a cardiac lobe, i.e. a swelling on the head shield such as that seen in horseshoe crabs, (3) division of the body into a putative prosoma and opisthosoma and (4) an anus on the last trunk segment. A common criticism of this interpretation is the fact that it lacks chelicerae (or antennae for that matter), although BOXSHALL (2004) suggested that chelicerae might be present, but indistinct. BUDD (2002) has even proposed that the ‘six’ pairs of prosomal limbs are just outgrowths from the articles comprising a single pair of ‘great-appendages’ (see above). Arachnomorph cladograms (e.g. WILLS et al. 1998) generally did not resolve Sanctacaris as sister-group of Chelicerata, drawing its affi nities into question. Interestingly, what Wills et al. did recover was Sanctacaris close to Yohoia. Bearing in mind the new position proposed for Yohoia on the chelicerate stem-lineage, and the possibility that Sanctacaris has ‘great-appendages’ too, a new look at the affi nities of Sanctacaris is clearly warranted. Its interpretation as a stem-lineage chelicerate may yet prove to be correct, albeit for different reasons to those proposed in the original description. Acknowledgements: I thank Gerhard Scholtz, Dieter Waloszek and Gonzalo Giribet for helpful discussions and (in part) for access to manuscripts in press, Franz Krapp for pointing out the Bousfi eld paper and Simon Braddy for valuable suggestions on the manuscript during review. Received: 06.08.2005 Accepted: 10.04.2006 References ANDERSON L. I., P. A. SELDEN 1997. Opisthosomal fusion and phylogeny of Palaeozoic Xiphosura. - Lethaia, 30: 19-31. AX P. 1987. The Phylogenetic System. Chichester, New York, J. Wiley & Sons, 340 p. BERGSTRÖM J., X. G. HOU 2003. Arthropod origins. - Bulletin of Geosciences, 78: 323-334. BRIGGS D. E. G., D. COLLINS 1988. A Middle Cambrian chelicerate from Mount Stephen, British Columbia. - Palaeontology, 31: 779-798. BRIGGS D. E. G., R. A. FORTEY 1989. The early radiation and relationships of the major arthropod groups. - Science, 246: 241-243.
J. Dunlop: Euchelicerate origins BRIGGS D. E. G., D. L. BRUTON, H. B. WHITTINGTON 1979. Appendages of the arthropod Aglaspis spinifer (Upper Cambrian, Wisconsin) and their signifi cance. - Palaeontology, 22: 167-180. BROILI F. 1932. Ein neuer Crustacee aus dem rheinischen Unterdevon. - Sitzungsberichte der Bayerischen Akademie der Wissenschaften, 1932: 27-38. BOUDREAUX H. B. 1979. Arthropod Phylogeny - with Special Reference to Insects. New York, Wiley, 320 p. BOUSFIELD E. L. 1995. A contribution to the natural classifi cation of Lower and Middle Cambrian arthropods: food gathering and feeding mechanisms. - Amphipacifi ca, 2: 3-34. BOXSHALL G. A. 2004. The evolution of arthropod limbs. - Biological Reviews, 79: 253-300. BUDD G. E. 2002. A palaeontological solution to the arthropod head problem. - Nature, 417: 271-275. CHEN J.-Y., D. WALOSZEK, A. MAAS 2004. A new ‘great-appendage’ arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial antero-ventral appendages. - Lethaia, 37: 3-20. COLLINS D. 1996. The “evolution” of Anomalocaris and its classifi cation in the arthropod class Dinocarida (nov.) and order Radiodonta (nov.). - Journal of Paleontology, 70: 280-293. COTTON T., S. J. BRADDY 2004. The phylogeny of arachnomorph arthropods and the origins of Chelicerata. - Transactions of the Royal Society of Edinburgh: Earth Sciences, 94: 169-193. DAMEN W. G. M., M. HAUSDORF, E. A. SEYFARTH, D. TAUTZ 1998. A conserved mode of head segmentation in arthropods revealed by the expression of Hox genes in a spider. - Proceedings of the National Academy of Science, USA, 95: 10665-10670. DUNLOP J. A., C. P. ARANGO 2005. Pycnogonid affi nities: a review. - Journal of Zoological Systematics and Evolutionary Research, 43: 8-21. DUNLOP J. A., P. A. SELDEN 1998. The early history and phylogeny of the chelicerates. - In: FORTEY R. A., R. H. THOMAS (eds): Arthropod Relationships. London, Chapman & Hall, 221-235. DUNLOP J. A., M. WEBSTER 1999. Fossil evidence, terrestrialization and arachnid phylogeny. - Journal of Arachnology, 27: 86-93 EDGECOMBE G. D., G. D. F. WILSON, D. J. COLGAN, M. R. GRAY, G. CASSIS 2000. Arthropod cladistics: combined analysis of histone H3 and U2 snRNA sequences and morphology. - Cladistics, 16: 155-203. EMERSON M. J., F. R. SCHRAM 1997. Theories patterns, and reality: game plan for arthropod phylogeny. - In: FORTEY R. A., R. H. THOMAS (eds): Arthropod Relationships. London, Chapman & Hall, 69-86. FORTEY R. A., H. B. WHITTINGTON 1989. The Trilobita as a natural group. - Historical Biology, 2: 125-138. GIRIBET G., C. RIBERA 2000. A review of arthropod phylogeny: new data based on ribosomal DNA sequences and direct character optimization. - Cladistics, 16: 204-231. GIRIBET G., G. D. EDGECOMBE, W. C. WHEELER 2001. Arthropod phylogeny based on eight molecular loci and morphology. - Nature, 413: 157-161. GOULD S. J. 1989. Wonderful Life. New York, W. W. Norton, 347 p. GRASSHOFF M. 1978. A model of the evolution of the main chelicerate groups. - Symposia of the Zoological Society of London, 42: 273-284. GRÜNER H.-E. (ed.) 1993. Lehrbuch der Speziellen Zoologie. Band I, 4. Teil, Arthropoda (ohne Insecta). Jena, Gustav Fisher Verlag, 1267 p. HAHN G. 1989. Reconstruction of the phylogenetic relationships among the higher taxa of trilobites. - Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF), 28: 187-199. HEDGPETH J. 1947. On the evolutionary signifi cance of the Pycnogonida. - Smithsonian Miscellaneous Collections, 106(18): 1-53. HEIDER K. 1913. Entwicklungsgeschichte und Morphologie der Wirbellosen. - In: HINNEBERG P. (ed.): Der Kultur der Gegenwart, Teil 3, Abt. 4, Bd. 2, Leipzig & Berlin, B. G. Teubner, 176-332. HESSELBO S. P. 1992. Aglaspidida (Arthropoda) from the Upper Cambrian of Wisconsin. - Journal of Paleontology, 66: 885-923. HOU X.-G. 1987. Two new arthropods from the Lower Cambrian Chengjiang, Eastern Yunnan. - Acta Palaeontologica Sinica, 26: 236-256. HOU X.-G., J. BERGSTRÖM 1997. Arthropods of the Lower Cambrian Chengjiang fauna, southwest China. - Fossils & Strata, 45: 1-116. 21
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Lycosid abundance R. Jocqué & M. A
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