The Enigmatic Lobopodians
Lobopodians
are small, segmented animals of uncertain taxonomic placement
that have apparent affinity to both arthropods
and annelids,
and that have a fossil record dating
to the early Cambrian, but examples are also known from Lagerstatten from
the Ordovician, Silurian and Carboniferous (e.g., Mazon
Creek).
The lobopods resemble Onychophorans,
or velvet worms, that are terrestrial worms with legs, and
indeed
may be Onychophoran ancestors. A distinguishing characteristic
of Lobopodians is dorsal-lateral plates or sclerites. Six
genera,
each with a single species, are described from the Chengjiang
Biota, making the Chengjiang
Maotianshan Shales the primary
source of Lobopod fossils. Aysheaia from the Burgess
Shale and
the strange Hallucigenia have some resemblance, as does Xenusion
from early Cambrian sandstones of eastern Europe that is
known
from only two specimens; these animals are now placed in the
onychophorans (Superphyum
Lophotrochozoa). Lobopodian morphologies in the Cambrian
exhibit wide diversity, as if nature were experimenting (and
maybe it was), spurring on classification debates
for decades. The most recent cladistical work by Smith
and Ortega-Hernández
(Nature, 2014) concludes that Lobopodia should be placed
in an unranked paraphyletic taxon, Panarthropoda, comprising
three phyla Arthropoda, Tardigrada, Onychophora, as well as
the genus Thelxiope (a stem group spiney arthropod occuring
in the Burgess Shale, with no example with legs preserved).
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References:
- Smith MR, Ortega-Hernández J. Hallucigenia's onychophoran-like
claws and the case for Tactopoda. Nature. 2014 Oct 16;514(7522):363-6.
- Whittington,
H. B. The lobopod animal Aysheaia pedunculata Walcott, Middle
Cambrian, Burgess Shale, British Columbia. Phil. Trans. R.
Soc. Lond. B 284,
165–197
(1978).
- Hou,
X.-G. & Bergström, J. Cambrian lobopodians–ancestors
of extant onychophorans? Zool. J. Linn. Soc. 114, 3–19
(1995)
- Ramsköld,
L. & Chen, J.-Y. in Arthropod
Fossils and Phylogeny (ed. Edgecombe, G. D.) 107–150
(Columbia Univ. Press, 1998)
- Bergström,
J. & Hou,
X. Cambrian Onychophora or xenusians. Zool. Anz. 240,
237–245 (2001)
- Budd,
G. E. Tardigrades as ‘stem-group
arthropods’: the evidence
from the Cambrian fauna. Zool. Anz. 240, 265–279
(2001)
- Wills,
M. A., Briggs, D. E. G., Fortey, R. A., Wilkinson, M. & Sneath,
P. H. A. in Arthropod Fossils and Phylogeny (ed. Edgecombe,
G. D.) 33–105
(Columbia Univ. Press, 1998)
- Campbell,
L. I. et al. MicroRNAs and phylogenomics resolve the relationships
of Tardigrada and suggest
that velvet worms are the sister group of Arthropoda.
Proc.
Natl Acad.
Sci. USA 108, 15920–15924
(2011)
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