The
Echinodermata, (from the Greek meaning spiny skin), is a phylum
containing some 13,000 extinct and 7,000 extant species. Living
representatives are only found in marine environment, making the
phylum the largest lacking terrestrial and fresh water forms. Echinoderms
evolved from bilaterally symmetric animals exhibiting fivefold radial
symmetry in portions of their body at some stage of life. This pentameral
symmetry is readily apparent in familiar adult starfish and sand
dollars. Other echinoderms both extant and extinct lack the five
point morphology because it was lost somewhere during development
(i.e., during ontogeny).
Echinoderms have
a vascular system that carries water and which in some echinoderms
end in suckered feet enabling the creature to grip and move objects.
Reproduction in Echinoderms reproduction is usually via external
fertilization through eggs and sperm discharged into the water,
and the majority of echinoderms have several planktonic larval
stages before reverting to a sessile
existence on the seafloor.
Since
most echinoderms have some type of calcareous
support exoskeleton (actually often interlocking plates of calcium
carbonate), there exists an extensive fossil record tracing echinoderm
evolution. Yet, many aspects of their early evolutionary origins
are confounded, such that the classification table below is but
one of many interpretations to be found in the literature. Importantly,
the Echinodermata phylogenetic relationship to other phyla is
poorly understood because they were already well differentiated
by the Cambrian, and
their unique characteristics are not present in other groups.
While echinoderms are known from the Cambrian on, the Vendian
period has a few soft-bodied fossils that are putative echinoderms
or their ancestors. These include Arkarua and Tribrachidium from
the Ediacara Hills of Australia. Homalozoans, from which echinoderm
may have descended, and eocrinoids,
that are not directly ancestral to the true crinoids, are abundant
in the early Cambrian fossil record.
A
possible early crinoid is Echmatocrinus from the famous Burgess
Shale of the middle Cambrian, though many researchers doubt it
was a true crinoid. Cotyledion
from the much younger early Cambrian Chengjiang
Maotianshan Shale
is another potential primitive crinoid. Other Cambrian echinoderms
included the unusual helicoplacoids. Asterozoans (starfish and
brittle stars) appeared in the Ordovician, as did the earliest
echinozoans. The oldest asterozoans (the Somasteroidea) have morphological
similarities to both starfish and brittle stars, supporting the
theory that starfish and brittle stars probably diverged from
a common somasteroid ancestor. After the Ordovician, there is
an extensive echinoderm fossil record dominated by crinoids and
blastoids; such as this stunning Triacrinus
crinoid death assemblage from the Devonian Hunsruck
Slates near Bundenbach, Germany. Some Paleozoic limestone
formations are comprised of almost nothing other than crinoid
and blastoid pieces. All the blastoids and most of the crinoids
met extinction at the end of the Permian, leaving only the Asterozoans
and echinozoans that remain extant today. The Holothurians, or
sea cucumbers, are prevalent echinoderms but are extremely rarely
fossilized.
Complete
fossil starfish are also very rare, and often are but partial
plates or segments of arms. The poor fossilization
results because the skeleton is not ridged like echinoids (sea
urchins), but comprised of numerous small plates (or ossicles)
that quickly fall apart after decay of the soft parts of the animal.
Great
fields (so to speak) of crinoid gardens inhabited shallower waters
during the Paleozoic, essentially from the Ordovician on, and
particularly in the Carboniferous
(for example, see the famous Crawfordsville
crinoids). However, crinoids suffered a major crisis during
the Permian period (the P-T even) when most met extinction, with
but few survivors into the Triassic
period. The Mesozoic era realized another large crinoid radiation,
with more modern forms having flexible arms becoming widespread.
After another extinction event at the end of the Cenozoic they
again declined, with most remaining species constrained to deep
waters until present time.
Phylum
Echninodermata Taxonomy |
Subphylum |
Class |
Order |
Subphylum
Blastozoa |
Eocrinoidea
(Cambrian to Silurian) |
>
30 genera |
Parablastoidea
(Ordovician) |
3
genera |
Cystoidea
(Ordovician to Devonian) |
Rhombifera |
>
50 genera |
Diploporita
|
>
40 genera |
Blastoidea
(Cambrian to Permian) |
>
90 genera |
Subphylum
Asterozoa (starfish and kin, also called Stelleroidea) |
Asteroidea
(True starfish or sea stars - Lower Ordovician to recent) |
Forcipulatida |
Paxillosida |
Platyasterida |
Spinulosida |
Valvatida |
Ophiuroidea
(Brittle stars - Lower Ordovician to recent) |
Oegophiurida |
Ophiurida |
Phrynophiurida |
Crinozoa |
Paracrinoidea
(Ordovician to Silurian) |
>
10 genera |
Crinoidea
(Crinoids / sea lilies - Cambrian to recent) |
Articulata
(Triassic to recent, the only remaining crinoids) |
Cladida
(extinct) |
Flexibilia
(Lower Ordovician to Upper Permian) |
Camerata
(Middle Ordovician to Permian) |
Subphylum
Echinozoa |
Echinoidea
(Sea Urchins - Ordovician to recent) |
2
subclasses containing 13 orders and > 700 genera |
Class
Holothuroidea (Sea Cucumbers - Ordovician to recent) |
some
200 genera |
Class
Edrioasteroidea (Early Cambrian to Carboniferous) |
35
genera |
Class
Edrioblastoidea (Ordovician) |
single
genus |
Class
Helicoplacoidea (Cambrian) |
3
genera |
Class
Cyclocystoidea (Ordovician to Devonian) |
8
genera |
Additional
notes
- Parablastoidea
are small class of primitive blastozoan echinoderms containing
three genera found in the early Middle Ordovician
- The
Ophiuroids or brittle stars are probably the most common
extant echinoderms. They occur in large numbers in all
parts of the oceans, from the shallowest coastal waters
to the deep sea trenches.
- Blastoids
persisted until their extinction at the end of Permian,
some 250 million years ago. Although never as diverse
as the crinoids, blastoids are prodigious as fossils,
especially from the Carboniferous, and are particularly
abundant in some U.S. midwestern states.
- Crinoidea
has more than 1000 genera, with some 80 extant species.
|
|