The
fossil record provides scant evidence of the temporal scenario
of the transition of metazoans to life on land. The oldest fossil
of an animal that putatively lived on land is a euthycarcinoid
from the late Ordovician of Australia. The oldest terrestrial
faunal assemblages are arachnids and myriapods from the Silurian
of UK. Except for some late ichnofossils, the fossil record
provides no evidence to set aside the hypothesis that prokaryotes
ruled the land to the end of the Cambrian and many millions
of years beyond. Of particular importance are ichnofossils of
the genus
Protichnites,
which occur in several North American locations dating from
the late Cambrian into the Ordovician.
The
Protichnites ichnogenera generally refers to a trackway two
rows of imprints (footprints) inside which is a medial impression
(such as a telson drag mark).
They occur throughout the Paleozoic, throughout the world. Notable
Protichnites sites occur in Ontario, Quebec, New York, and Missouri,
and all in subtidal, intertidal and/or eolian (i.e., as in dunes
above the tide line) sandstones. Among these sites, the Protichnites
from the Krukowski Quarry are the oldest at some 510 million
years ago. Arthropods underwent enormous change and numerous
radiations throughout the periods of the Cambrian. It is likely
that the makers of the Protichnites (and Diplichnites) ichnogenus
also had many arthropod makers that changed across geologic
time. But what animals made the upper Cambrian tracks found
in the Krukowski quarry. The main suspects are eurypterids,
horseshoe crabs, Euthycarcinoids
and the Aglaspids, or some closely related arthropod or
stem group among these taxons.
It
was a time when evolutionary adaptation was producing predators,
and a time when adaptation was producing defense from predation,
both powerful selective pressures in part fueling what is known
as the Cambrian explosion. It is reasonable then to conjecture
that animals found ways to first emerge on marine shorelines
for different reasons -- some to graze in safety on abundant
bacterial colonies, and possibly others to hunt the grazers.
The
animal that made Protichnites tracks could have been one of
the Earth's first air-breathing animals. The Protichnites ichnogenus
is a Repichnia Ichnofossil associated
with vagile locomotion. Sir William Logan described the Protichnites
genus in 1863, the time of Darwin,
as fossil evidence that animals had evolved more rapidly and
much earlier than thought at the time of its discovery. Notable
in Protichnites is the prominent markings of a tail or other
body part being dragged; footprints, when present, are subtle
by comparison. The maker of these fossil trackways might have
been an animal
resembling the extant but ancient horseshoe crab, except of
a stem
group
that lacked a hard shell to be preserved. Others have posited
that Protichnites was a soft-bodied progenitor of the large,
now extinct group of early predators, the Eurypterids (Chelicerate
"meaning biting claws" Arthropods). The "Eurypterid
theory" is particularly intriguing since it would imply
that just as animals were first moving to land to feed, hungry
new predators were already on the hunt.
Some
of the mystery of Protichnites trackways is just being cleared
up as, after more than a decade, the very first body fossils
of a putative Protichnites track maker have at last been discovered
in the Krukowski quarry. The creatures that were discovered
in a single dessication zone during the summer of 2003 have
affinity to some of the most problematic groups of Arthropods,
the Euthycarcinoids and the Aglaspids.
The euthycarcinoids that are
known from 13 species from Upper Cambrian or Lower Silurian-Middle
Triassic from Argentina, Western Australia, Europe (e.g., the
Rhynie chert) and the Mazon Creek in Illinois (Vacarri, 2004).
But it is only in the Argentina and Wisconsin sites that the
Protichnites trackways are also found. The Krukowski arthropods
may emerge as the oldest land-based animal fossil in the entire
fossil record. Similarly, the aglaspids
are arthropods whose taxonomy among arthropods is uncertain.
Also
see: Ichnofossil Nomenclature
References