echinoderms

Echinodermata
tagghudingar

general things

Echinodermata

Echinodermata Klein, 1734 ex Bruguire, 1789

Ethymology: Greek echinos (hedgehog, spine) and derma (integument, skin)

Main characteristics:
all echinoderms have a mesodermal skeleton of porous calcite plates but: reduced to absent among some holothuroids skeleton covered by a thin skin Endoskeleton radial symmetry, typically five-rayed / pentameral, as adults but: not developed in stem-group echinoderms, or carpoids lost or obscured by secondary adaptations water-vascular system (complex internal apparatus of tubes and bladders containing fluid) with extensions emerging through the skeleton to the outside as tube feet or podia
(podia serve for locomotion, respiration and feeding a.o.)

but: has not been confirmed in all fossil groups

Echinodermata general features

Porous calcite plates (ossicles)

intracellular
reducing the ossicle-producing stroma cell to a thin layer that lines the skeletal meshwork (stereom).

each ossicle is a monocrystal

high-magnesium calcite, which is a spathic mineral (crystalline with strong cleavage)

spathic fracture has been changed to conchoidal by inserting organic macromolecules into the crystal lattice each ossicle is composed of two interlocked networks, one composed of mineral matter (stereom) and the other, of organic matter of mesodermal origin (stroma). the trabecular structure of the stereom does not allow cracks to propagate farther than the next cavity in fossils, pores of the stereom have become closed by diagenetic calcite in optical continuity.

epidermis stroma

stereom Echinodermata general features ossicles

Ossicles ...
... can be fused into a test (sea urchins, crinoids) ... can spread apart (sea cucumbers) ... can be intermediate and variable (sea stars)

Echinodermata general features ossicles

Exceptions!

Primary polycrystalline calcite in ...

... the cortex of primary spines of Cidaridae ... tooth sceleton of Clypeaster ... the accessory calcareous structures filling the crevice fold in the chewing areas of Diadematoida teeth.

Cortex (primary polycrystalline)

disturbed crystal structure (secondary!)

Echinodermata general features ossicles

Water vascular system (= ambulacral system)

Hydraulic system of fluid-filled canals
internal transport locomotion gas exchange food capture excretion fluid similar to seawater and is moved through system with cilia

Podia tube feet

suction cups at the end each tube foot works independently moved by muscles and hydraulics

Madreporite
perforated platelike structure which acts as the inlet for the water vascular system (important for orientation, arms/ambulacralia)

Stone canal
lime-walled tube which connects ring canal and madreporite
[Latin ambulacrum, walk planted with trees, from ambulare, to walk.]

Echinodermata general features water vascular system

Echinodermata general features water vascular system

ambulacra

Echinodermata general features water vascular system

Other fearures:

non-segmented no head open blood system nervous system simple without nerval center light sensors/eyespots but no eyes reproduce sexually (produce sperm and eggs) and asexually (regenerating lost parts)

Echinodermata other features

Ecology
exclusively marine and stenohaline typically benthic (infaunal, epifaunal) but a few pelagic (planktonic and pseudo-planktonic) forms exist live free (vagile, floating, or active swimming) or attached (sessile) live on all kinds of substrates, littoral to abyssal, in all latitudes (today typical in shallow coastal waters and ocean trenches) carnivorous / herbivorous / detritus eaters

Echinodermata ecology

Echinodermata ecology

Echinodermata ecology

Luidia ciliaris (Asteroidea)

Heterometra sp.

Echinocardium cordatum (Echinoidea)

Amphiura brachiata (Ophiuroidea, brittle star)

Echinodermata ecology

systematics
1) Stem-group Echinodermata

Echinodermata

Echinoderm classification
Arkarua adami from the Ediacaran Hills of Australia oldest putative echinoderm

Reference: Gehling, J.G. 1987. Earliest known echinoderm - a new Ediacaran fossil from the Pound Subgroup of South Australia. Alcheringa, 11:337-345.

Echinodermata classification

Helicoplacoidea (Lower Cambrian)

spirals of overlapping ossicles "mouth" was a long groove that also spiralled around their body lived probably in burrows and extending their bodies outward to feed three ambulacra complete fossils only found in the White Mountains in California

Reference: Dornbos, S. Q. & Bottjer, D. J. 2000. Taphonomy and environmental distribution of helicoplacoid echinoderms. Palaios, 16: 197-204.

Echinodermata classification Helicoplacoidea

Mixed ossicles (Lower Cambrian)

Homlozoa systematics
2) Homalozoa

Echinodermata

Homalozoa [= Calcichordata, Carpoidea]

(Middle Cambrian Middle Devonian)

bilateral or asymmetical main body (theca) constructed of two types of ossicles: marginalia and centralia theca typically flattened and with appendages (stele, aulacophore) use of appendages unknown (locomotion, feeding, tail for swimming etc.) Four main groups can be distinguished Ctenocystoidea Homostelea (=Cincta) Homoiostelea (= Soluta) Stylophora
[incl. Mitrata, Cornuta, Ankyroida]

Echinodermata classification - Homalozoa

marginalia

centralia
proximesidististele

presumed mouth

presumed anus

stele, aulacophore (tripartite)

arm =? ambulacrum anterior?

posterior?

Echinodermata classification - Homalozoa

Homostelea Stylophora, Cornuta

Stylophora, Ankyroida Echinodermata classification - Homalozoa

Homoiostelea

Ctenocystoidea

Calcichordate theory (Jefferies 1986):

Chordates (ourselves) evolved from homalozoan echinoderms Stele=tail Calcite skeleton was replaced by an apatite one Theory more or less abandoned by now

systematics
3) Pelmatozoa

Echinodermata

Pelmatozoa (since Middle Cambrian)

attached living, stalked echinoderms (some secondarily freeliving) cup-shaped head (the calyx) attached to the calyx are arms (brachiols/brachia) the calyx is usually connected tegmen oral plates to the bottom via a stem pentamere or radial symmetry Three main groups: Cystoidea Blastoidea Crinoidea (sea lilies)
stem internodal nodal columnars theca calyx

arm brachium brachiol

Echinodermata classification Pelmatozoa

Cystoidea (Ordovician to Devonian)

[kristallpplen]

respiratory pore structures traversing the plates of the theca pore structures are basis for taxonomy theca often slightly irregular stem short or absent well defined anal pyramid surrounded by trigonal anal plates laterally to perioral plates hydropore (slit near the perisome, probably entrance of water vascular system) arms are non-branching, biserial brachiols two pore types typify the two main groups of cystoids Diploporita
diplopores

Rhombifera
dichopores rhomb-shaped contour

Echinodermata classification Pelmatozoa Cystoidea

[anal pyramid, brachiols, dichopores]

Echinodermata classification Pelmatozoa Cystoidea

Blastoidea (Early Cambrian/Ordovician to Permian)

typically with pentamere symmetry short stem (rarely preserved) crown of brachioles (rarely preserved) typical plating pattern: 3 basal plates (BB) 5 radial plates (RR) 5 deltoid plates () 5 lancet plates below ambulacra star shaped mouth surrounded by spiracles (outlet system)

anispiracle

Echinodermata classification Pelmatozoa Blastoidea

Blastoidea sensu stricto

Eocrinoidea (Early Cambrian to Silurian) Gogia palmeri

Echinodermata classification Pelmatozoa Blastoidea

Crinoidea (since Ordovician)

Most common palaeozoic echinoderm fossils Long stems Crinoid meadows in shallow water Disarticulated stems rockforming

Crinoidea (since Ordovician)

organized in stem, theka and brachia with ambulacra
brachials radials basals monocyclic calyx radials basals infrabasals dicyclic calyx radianal plate

brachia Echinodermata classification Pelmatozoa Crinoidea

Crinoidea (major groups and relationships)

Early Ordovician to Permian

= Aethocrinea

Ordovician since Triassic

Early Ordovician to Permian

Middle Ordovician to Permian Early Ordovician to Permian

1 development of true arms with extension of the ambulacra 2 loss of the basal circlet 3 loss of the lintel circlet 4 fixed brachials and fixed interradials incorporated into the calyx 5 with the mouth exposed on the tegmen and loose plate sutures 6 loss of the anal plate and an entoneural system enclosed within the calyx plates

Echinodermata classification Pelmatozoa Crinoidea

Camerata

Echinodermata classification Pelmatozoa Crinoidea Camerata

Articulata (Comatulida)
occur from intertidal to abyssal depths retain a stalk as postlarvae, but shed all but the topmost segment and take up a free existence as juveniles and adults with cirri (columns of ossicles) radiating from the margin of the centrodorsal plate distal ossicle is a small claw (holdfast)

Leptometra celtica

Antedon bifida

Echinodermata classification Pelmatozoa Crinoidea Articulata Comatulida

In deep water settings: Stalked crinoids survive to the present Crown of arms directed against current for effective filtering

systematics
4) Eleutherozoa

Echinodermata

Eleutherozoa (since Early Cambrian / Ordovician)

the non-stalked, vagile echinoderms (with exceptions!) = sea cucumbers, sea stars and brittle stars, sea urchins and sand dollars and some small fossil groups of uncertain affinity, as well as sea daisies Edrioasteroidea (Early Cambrian to Early Carboniferous) Holothuroidea (Ordovician?, since Early Devonian)
(sea cucumbers)

Concentricycloidea (Recent)
(sea daisies)

Asterozoa (since Early Ordovocian)

(sea stars and cushion stars, brittle stars and basket stars, Somasteroidea)

Echinozoa (since Early Ordovician)

(sea urchins and sand dollars, Cyclocystoidea?)

Echinodermata classification Eleutherozoa

Edrioasteroidea
typically with well developed pentamere symmetry endothecal ambulacral system no arms or brachioles with anal pyramid and peristomal field sessile combine characteristics of Eleutherozoa and Pelmatozoa Early Cambrian to Early Carboniferous

Edrioaster (Cambrian)

Stromatocystites (Cambrian)

Spiraclavus (Carboniferous) Carneyella (Ordovician)

Echinodermata classification Eleutherozoa Edrioasteroidea

Holothuroidea (sea cucumbers)

mouth and anus at opposite ends (secondary ossicles are imbedded within leathery skin ossicles often reduced to little sclerites (typify certain families) only in mouth region is a rigid series of plates (perioral ring)

modified tube feet for feeding

ambulacra arranged in two sets parallel with the long axis (3 ventral, 2 dorsal) few complete fossils: 2 Upper Jurassic (Solnhofen), 1 Lower Devonian Hunsrck Ordovician?, since Early Devonian

Palaeocucumaria hunsrueckiana

Echinodermata classification Eleutherozoa Holothuroidea

Concentricycloidea (sea daisies)

discovered in 1986 disk-shaped flat bodies and are less than 1 cm in diameter. The two species were located on wood found in deep waters off the coasts of New Zealand and the Bahamas recently found in the North Pacific (Voight, 2005) water-vascular system, with tube feet on the body surface around the edge of the disk no obvious arms or mouth; appear to absorb nutrients through their body wall possibly an aberrant asteroid

Reference: Voight, J. R. 2005. First report of the enigmatic echinoderm Xyloplax from the North Pacific. Biological Bulletin, 208: 77-80.

Echinodermata classification Eleutherozoa Concentricycloidea

Asterozoa
echinoderms with depressed star-shaped body with central disc bearing mouth on underside symmetrical radiating arms tube feet normaly confined to lower side of body since Early Ordovician 3 main groups: Asteroidea
(sea stars and cushion stars)

Somasteroidea

Ophiuroidea
(brittle stars and basket stars) Echinodermata classification Eleutherozoa Asterozoa

Asteroidea (sea stars and cushion stars)

Asterozoans with relatively broad arms considerable hollow space between ossicular frame arms not seperated from central disc oral side with open ambulacral grooves with rows of tube feed since Early Ordovician

gen. et sp. indet. (Ordovician, Morocco)

Crateraster (Cretaceous, England)

Echinodermata classification Eleutherozoa Asterozoa Asteroidea

Somasteroidea
Asterozoans with oral surface bearing shallow radial channels axial skeleton with ambulacral ossicles in double series each ambulacral giving rise to a transverse series of ossicles (metapinnules) Early Ordovician to Late Devonian appear prior to Asteroidea and Ophiuroidea

Villebrunaster thorali (Early Ordovician, France) oral, ventral side

Archegonaster pentagonus (Early Ordovician, Czechia) oral, ventral side

Echinodermata classification Eleutherozoa Asterozoa Somasteroidea

Ophiuroidea (brittle stars and basket stars)

5 thin, flexible snake-like arms arms clearly demarcated from central disks and made up of a single row of large calcite plates termed vertebrae
[early members had double rows of alternating vertebrae]
Ophioderma (Jurassic)

typically suspension feeders, few carnivores live today mainly in bathyal and abyssal depths havent chanched much since their first appearance since Ordovician
Furcaster (Devonian)

Loriolaster (Devonian) modern brittle star Encrinaster (Devonian)

Echinodermata classification Eleutherozoa Asterozoa Ophiuroidea

Echinozoa (sea urchins)

echinoderms with globose or discoidal test typically with spines typical with 5 ambulacral fields 2 main groups: 1) Echinoidea (sea urchins and sand dollars) 2) Cyclocystoidea since Early Ordovician

Phymosoma sp. (Cretaceous)

Echinodermata classification Eleutherozoa Echinozoa

Echinoidea (sea urchins and sand dollars)

test (= corona) constructed of 20 radial plate rows (= 10 segments) narrow segments = ambulacra, broad segments = interambulacra interambulacral plates are large and tubercular, without perforations ambulacral plates have pored plates through which the tube feet emerge mouth with five hard teeth arranged in a circlet (known as Aristotle's lantern) [reduced in derived forms; absent in stem group echinoids] (used for grazing) move with tube feet or walk on their spines traditionally devided in regular (Regularia) and irregular echinoids (Irregularia)

Echinodermata classification Eleutherozoa Echinozoa - Echinoidea

Aristotle's lantern

perignathic girdle

Echinodermata classification Eleutherozoa Echinozoa - Echinoidea

Regularia and Irregularia

represent no phylogentic groups, still used as a convenient, informal classification Regularia: strict pentamere symmetry upper surface (aboral) with central apical disc bearing periproct with anus long spines for protection epibenthic grazers paraphyletic group pentamere symmetry with superimposed bilateral symmetry mouth located more anterior anus at posterior end test covered with a mat of short spines infaunal depost feeders most probably monophyletic

Irregularia:

Echinodermata classification Eleutherozoa Echinozoa - Echinoidea

Echinoidea classification
==o Echinoidea |--o Perischoechinoidea | | | `--o Cidariida |-- Cidaridae `-- Psychocidaridae
= Regularia = a.o. sand dollars = a.o. heart urchins = advanced stem group = Paleozoic Regularia, stem group

`--+-- Euechinoidea [paraphyletic?] |-- Gnathostomata `-- Atelostomata

Perischoechinoidea - stem group Echinoidea - more than 20 radial plate rows - perignathic girdle rudimentary or absent

Echinodermata classification Eleutherozoa Echinozoa - Echinoidea

Classification: - Arrangement of plates

in apical disc - Morphology of lantern - Construction of perignathic girdle

adambulacral - perradial - interradial sutures

Echinodermata classification Eleutherozoa Echinozoa Echinoidea regular echinoids

20 plare columns 5 interambulacral 5 ambulacral

Classification - Apical disc - Shape of petals - Plating of plastron

Echinodermata classification Eleutherozoa Echinozoa Echinoidea heart urchins

5 interambulacral columns 5 ambulacral columns Adapical portion of ambulacra developed int petals Spines are short and stubby 5 food grooves radiating from peristome Often with elongated perforations (lunules) Have a lantern and perignathic Girdle, which, however, can be resorbed in adult stages

Echinodermata classification Eleutherozoa Echinozoa Echinoidea sand dollars

Cyclocystoidea
enigmatic Paleozoic group (Early Ordovocian to Early Carboniferous) mainly known by their ring of submarginal plates mode of life and systematic position controverse in situ finds confirm oral side down oral disc with complicated ambulacral system

Echinodermata classification Eleutherozoa Echinozoa Cyclocystoidea

systematics
5) Problematica

Echinodermata

Echmatocrinus from the Middle Cambrian Burgess Shale described as the oldest known crinoid

interpretation refused by several authors probably best considered as a cnidarian with octocoral affinity (Ausich & Babcock, 2000)

Echmatocrinus brachiatus surface pattern with scales/plates

Reference: Ausich W. I. & Babcock, L. E. 2000. Echmatocrinus, a Burgess Shale animal reconsidered. - Lethaia, 33 (2): 92-94.

Echinodermata classification Problematica

Eldonia, Portalia and Redoubtia, all known from the Burgess Shale have originally been described as holothourians

Eldonia is now considered to be a jelly fish Portalia and Redoubtia is of uncertain affinity (polychaete or sponge?)

Eldonia ludwigi Middle Cambrian

Echinodermata classification Problematica

systematics
summary

Echinodermata

<==o Echinodermata (Spiny-skinned animals) |?- Arkarua adami |-- Homalozoa [= Calcichordata] `--+--o Helicoplacoidea | `-- Helicoplacus `--o Echinodermata sensu stricto (pentametric echinodermates) |?- Camptostroma |--o Pelmatozoa | |?= Lepidocystoidea | |?- Astrocystites ottawaensis Whiteaves, 1897 | |?-o Parablastoidea Hudson, 1907 | | `--o Plastocystidae Jaekel, 1918 | | |-- Blastocystis rossica Jekel, 1918 | | `-- Blastoidocrinus carchariaedens Billings, 1859 | |-- Blastoidea [incl. Eocrinoidea] | |-- Cystoidea [incl. Diploporida, Rhombifera] | `--+?- Echmatocrinus [Echmatocrinea: Echmatocrinida] | `--o Crinozoa (sea lilies and feather stars) | |?- Coronata | |?- Paracrinoidea Regnll, 1945 [Eustelea Jaekel, 1900; Deviata Jaekel, 1918] | `-- Crinoidea (sea lilies and feather stars) `--o Eleutherozoa |?- Stromatocystides |-- Edrioasteroidea `--+?- Holothuroidea (sea cucumbers) `--+?-o Concentricycloidea (sea daisies) | `--o Xyloplax [possibly aberrant asteroid] |-- Asterozoa (sea stars and brittle stars) `?-+-- Sollasina woodwardi (Sollas) Fedotov, 1926 [Ophiocistioidea Sollas, 1899] `--o Echinozoa |?- Cyclocystoidea `-- Echinoidea (sea urchins and sand dollars)
modified after: http://www.fmnh.helsinki.fi/users/haaramo/Metazoa/Deuterostoma/Echinodermata/Echinodermata.htm#Edrioasteroidea

Echinodermata systematics

Classical paleontological classification (e.g., Clarkson)

Phylum Echinodermata
Subphylum Blastozoa * Class Eocrinoidea (Cambrian - Silurian, 30-32 genera) * Class Parablastoidea (Ordovician, 3 genera) * Class Rhombifera = Cystoidea in part (Ordovician - Devonian, 60 genera) * Class Diploporita = Cystoidea in part (Ordovician - Devonian, 42 genera) * Class Blastoidea (Silurian - Permian, 95 genera) Subphylum Crinozoa * Class Crinoidea - sea lilies (Cambrian - Recent, 1005 genera) * Class Paracrinoidea (Ordovician - Silurian, 13-15 genera) Subphylum Echinozoa * Class Echinoidea (Sea Urchins) (Ordovician - Recent, 765 genera) * Class Holithuriudea (Sea Cucumbers) (Ordovician - Recent, 200 genera) * Class Edrioasteroidea (early Cambrian - Carboniferous, 35 genera) * Class Edrioblastoidea (Ordovician, 1 genus) * Class Helicoplacoidea (Cambrian, 3 genera) * Class Cyclocystoidea (Ordovician - Devonian, 8 genera) Subphylum Asterozoa (Stelleroidea) * Class Asteroidea - starfish - (early Ordovician - Recent, 430 genera) * Class Ophiuroidea - Brittle Stars -(Ordovician - Recent, 325 genera) Subphylum Homalozoa * Class Stylophora (Cambrian - Devonian, 32 genera) * Class Homoiostelea (Cambrian - Devonian, 12-13 genera) * Class Homostelea (Cambrian, 3 genera) * Class Ctenocystoidea (Cambrian, 2 genera)

source: http://www.sidwell.edu/us/science/vlb5/Labs/Classification_Lab/Eukarya/Animalia/Echinodermata

Echinodermata systematics

Echinodermata systematics

Echinodermata systematics

Echinodermata systematics

Echinodermata systematics

Echinodermata evolution

slut

Echinodermata