The
Cambrian period saw the emergence of group of animals that
would conquer
the marine environments world wide and go on to make the first
pioneering steps onto land. Arthropods
comprise the creatures with jointed legs, from fleas to crabs
to the extinct trilobites to
the extant but primitive horseshoe crab. Since first appearing,
probably during the Precambrian,
their variability has been nothing short of astonishing, and
their
impact on the living
earth enormous. Arthropods (Greek for jointed feet) comprise
the largest phylum of animals and include the insects, arachnids,
crustaceans, and many others as given in the table below. Approximately
80% of extant animal species are arthropods, with over a million
modern species described and an extensive fossil record dating
back to the base on the Cambrian.
Arthropods
are most characterized by their jointed limbs and cuticles,
which are mainly made of a-chitin. The cuticles of
crustaceans
are also biomineralized with calcium carbonate, as was that of
the extinct trilobite. The cuticle is sufficiently inflexible
as to inhibit growth such that it must be periodically replaced
by molting, a characteristic that unites nine phyla in Superphylum
Ecdysozoa (Arthropoda, Onychophora, Tardigrada, Kinorhyncha,
Priapulida, Loricifera, Nematoda and Nematomorpha). The arthropod
body plan
consists of repeated segments, each with
a pair of appendages. The embryos of all arthropods are segmented,
built from a series of repeated modules. The last common ancestor
of living arthropods probably consisted of a series of undifferentiated
segments, each with a pair of appendages that functioned as
limbs. However all known living and fossil arthropods have
grouped segments into tagmata in which segments and their limbs
are specialized in various ways.
Parvancorina
and Spriggina are Ediacaran animals from around 555 Mya that
are among the earliest putative
arthropods in the fossil record. Bivalve-like fossil shells
have been found in China that date to some 541 to 539
million
years
ago. The earliest Cambrian trilobite fossils date to about 530
million years ago, but because they were already diverse and
dispersed
worldwide,
they certainly must have already
existed for a
long period. Many arthropods are described from the Burgess
Shale dating
to some 505 million years ago. A large number of arthropods are
also described from the older (525 to 520 million years) Chengjiang
Maotianshan Shales. The earliest fossil crustaceans date
to about 513 million years ago the earliest fossil
shrimp to about 500 million years ago. The eariest
identifiable body fossils of land animals are arachnids
and chilopods from the late Silurian (419 million year ago) of
England. Arthropod ichnofossils from the late Cambrian
have been intertidal sand dune deposits in Canada and Wisconsin.
Because
of their beauty and their extensive availablity in the fossil
record, the trilobite
is the unequivocal favorite among fossil collectors. Though
trilobites
dominated the Paleozoic marine environments, came back strong
after several mass extinctions, they faded out by the end
of the
Permian, their niches on various marine environments taken
over by their crustacean cousins. The spiders thrived on
land as did
the insects. Insects underwent an amazing adaptive radiation.
Excluding microbial organisms, modern times are dominated
by the
insects, with beetles alone making up some 25% of known organisms.
Subphylum
|
Class |
Common
Examples |
Trilobitamorpha
(note1) |
Trilobita |
Trilobites
and Relatives |
Aglaspidida
or Aglaspida |
Aglaspids |
Chelicerata |
Arachnida |
Spiders,
scorpions, harvestmen, ticks, and mites |
Merostomata |
Horseshoe
crabs and eurypterids. |
Pycnogonida |
Sea
spiders |
Myriapoda |
Archipolypoda
(note 2) |
early
myriapod |
Chilopoda |
Centipedes |
Diplopoda |
Millipedes |
Pauropoda |
|
Symphyla |
Garden
centipedes |
Hexapoda |
Diplura |
|
Collembola |
Springtails |
Protura |
|
Insecta |
|
Crustacea |
Branchiopoda |
Brine
shrimp |
Remipedia |
|
Cephalocarida |
Horseshoe
shrimps (no fossil record)) |
Maxillopoda |
Barnacles |
Ostracoda |
Seed
shrimp |
Malacostraca |
Crabs,
mole crabs, lobsters, isopods (woodlice and sowbugs), true
shrimps, and Phyllocarids (?) |
1
- Trilobitomorpha is a subphylum of the phylum Arthropoda
that includes the trilobites. Originally a variety of peculiar
forms, mostly from the lower Cambrian, were included as the
Class Trilobitoidea. However, the many species do not appear
to be closely related to the trilobites or, in many cases,
to each other either, and are now generally placed in separate
subphyla when classified at all.
2 - The Archipolypoda comprise the early Myriapods that are
also often classified under the Diplopoda (millipedes). Archipolypods
differed from millipedes, however, in having less coalesced
segments, larger head reletive
to the body, and large compound eyes. The fossil record indicates
appearance of Archipolypoda in the Silurian and extinction
at the end of the Carboniferous. Well-preserved specimens
are known from the Mazon Creek Formation.
References:
- Budd,
G. E., Butterfield, N. J., and Jensen, S., "Crustaceans
and the "Cambrian Explosion?",
Science 294 (5549): 2047 (2001).
- Yuan, X.; Xiao, S., Parsley, R.L., Zhou,
C., Chen, Z. and Hu, J., "Towering sponges in an Early Cambrian
Lagerstätte: Disparity between nonbilaterian and bilaterian
epifaunal tierers at the Neoproterozoic-Cambrian transition",
Geology 30 (4): 363–366 (2002).
- Lin,
J. P.; Gon, S.M.; Gehling, J.G.; Babcock, L.E.; Zhao,
Y.L.; Zhang, X.L.; Hu, S.X.; Yuan, J.L.; Yu, M.Y.; Peng,
J., "A Parvancorina-like arthropod from the Cambrian
of South China", Historical Biology 18 (1): 33–45,
(2006).
- Lieberman, B. S., "Testing the Darwinian
legacy of the Cambrian radiation using trilobite phylogeny
and biogeography", Journal of Paleontology 73 (2):
176 (1999).
- Jeram AJ, Selden PA, Edwards D. Land animals
in the Silurian: arachnids and myriapods from Shropshire,
England. Science.
;250:658–661. (1990).
- MacNaughton RB, Cole JM, Dalrymple RW, Braddy SJ, Briggs
DEG, Lukie TD. First steps on land: Arthropod trackways
in Cambrian-Ordovician eolian sandstone, southeastern Ontario,
Canada. Geology. (2002).
- Hoxie, C., Late Cambrian Arthropod Trackways
in Subaerially Exposed Environments (2005).
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