Eldoniid

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Eldonioidea
Temporal range: Cambrian–Devonian
Eldonia berbera 1.JPG
Fossils of Eldonia berbera , from the Late Ordovician of Morocco
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Clade: Cambroernida
Class: Eldonioidea
Dzik, 1991
Families

Eldoniids or eldonioids are an extinct clade of enigmatic disc-shaped animals which lived in the early to middle Paleozoic (Cambrian to Devonian). They are characterized by their "medusoid" (jellyfish-shaped) bodies, with the form of a shallow dome opening below to an offset mouth supplemented by filamentous tentacles. [1] Internally, they have a distinctive C-shaped cavity encompassing the gut, as well as hollow radial (radiating) structures arranged around a central ring canal. Most eldoniids are soft-bodied and can only be preserved in lagerstätten, but a few species may have hosted mineralized deposits. [2] [3] Historically, the affinities of eldoniids was enigmatic; recently, they been assessed as cambroernid deuterostomes. [4] [5] Their lifestyle is still an unresolved question; some authors reconstruct eldoniids as free-floating planktonic predators similar to jellyfish, [1] while others argue that they were passive detritivores, embedded within the seabed for much of their life. [3]

Contents

Classification

Eldoniids have a frequently fluctuating species composition and their relationship to other animals has been controversial. Some authors argued that they represented early holothurians (sea cucumbers), while others stressed their similarity to lophophorates (a subset of lophotrochozoans with ciliated tentacles known as lophophores). [3] [1] In 2010, eldoniids were allied with two other unusual Cambrian genera: Herpetogaster and Phlogites . Together, they comprise the cambroernids, an informal clade. Cambroernids are probably deuterostomes, specifically stem- or crown-group ambulacrarians related to echinoderms and hemichordates. [4] [5]

Eldoniidae was a family first established in reference to Eldonia , a disc-shaped fossil named in 1911. Eldonia was one of many mysterious soft-bodied animals discovered in the Cambrian-age Burgess Shale of Canada. [2] It is also a major component of the Late Ordovician Tafilalt biota of Morocco. Eldonia was initially mistaken for a scyphozoan (jellyfish), and was not the only member of the group originally misidentified as a cnidarian. Others include Vellumbrella (mistaken for a jellyfish), Discophyllum (mistaken for a coral), and Paropsonema (mistaken for a jellyfish-like porpitoid hydrozoan). [6] Eldoniids, in a broad definition, also include Rotadiscus , [3] Stellostomites , [1] Pararotadiscus , [1] and Seputus. [7] Apart from Eldonia, the most abundant and well-preserved eldoniid species are from the Cambrian of China: Stellostomites eumorphus and Rotadiscus grandis are from the Chengjiang biota of Yunnan, and Pararotadiscus guizhouensis is from the Kaili Formation of Guizhou. [8]

Dzik (1989) named the class Eldonioidea to encompass "velumbrellids" (i.e. eldoniids) and the goblet-shaped Cambrian animal Dinomischus . Eldonioidea was divided further into the orders Dinomischida (containing solely Dinomischus) and Vellumbrellida. The latter order was divided into Eldoniidae and Rotadiscidae. [9] Most subsequent authors doubt a close relationship between Dinomischus and eldoniids. As a result, "Eldonioid" and "Eldoniid" have been used interchangeably by many recent publications. Some authors classify the rotadiscids and post-Cambrian "paropsonemids" as informal groups within Eldoniidae, [1] [4] while others maintain rotadiscids and "paropsonemids" as separate eldonioid families. [8] [5]

Anatomy

Cross-sectional diagrams of Stellostomites eumorphus (= Eldonia eumorpha).
A: Vertical cross-section
B: Transverse section near the dorsal disc
C: Transverse section near the ventral disc
D: Underside of the ventral disc Stellostomites diagrams.png
Cross-sectional diagrams of Stellostomites eumorphus (= Eldonia eumorpha).
A: Vertical cross-section
B: Transverse section near the dorsal disc
C: Transverse section near the ventral disc
D: Underside of the ventral disc

The basic eldonioid body plan is oriented around two circular discs sealed together with the viscera in between. [1] The ventral (lower) disc is concave from below while the dorsal (upper) is convex from above, providing a dome-like profile to the body. The rim of the body is smooth, though the margin may furl slightly inwards under the ventral disc. Sediment infillings or differences in preservation allow for hollow internal structures to be differentiated from solid sheets of tissue within the body. [1]

Disc structures

Fossils of Eldonia ludwigii, showing the prominent coiled sac, radial lobes, and central ring canal Walcott Cambrian Geology and Paleontology II plate 09.jpg
Fossils of Eldonia ludwigii , showing the prominent coiled sac, radial lobes, and central ring canal

The ventral disc acts as a platform for numerous elongated internal structures arranged in a radiating pattern. [1] [4] [6] These narrow structures, known as “radial sacs” [2] [1] (also “radial lobes” [8] or "internal lobes" [8] ), were most likely hollow cavities. They are among the most visible features in most fossils. The sacs are divided from each other by thin ligamentous walls, described as “septa” or “mesenteries”. [1] In most eldoniid species, each radial sac bifurcates (divides into two parts) near the rim of the disc. [1] The radial sacs thin towards the center of the disc, but do not converge at a single point. Instead, they intersect with a small ring-shaped cavity or canal in the central part of the ventral disc. [1] The radial sacs and ring cavity were likely filled with fluids, akin to a hydrostatic skeleton [4] or (less likely) [1] [4] an echinoderm-like water vascular system. [2]

The dorsal disc is usually more rigid and resistant to distortion than the ventral disc. In several taxa, the dorsal disk is ornamented by one or more irregular concentric (ring-shaped) wrinkles, which may be growth lines or artefacts of compression. [2] [3] [1] [4] [7] [8] According to some interpretations, radial ridges, wrinkles, or grooves also ornament part of the dorsal disc in some rotadiscids. [3] [7] [8] “Paropsonemids” have even more complex dorsal discs, combining radial and concentric ornamentation at a fine level of detail. [7] [8] In Eldonia and Stellostomites , the dorsal disc appears to bear radial internal structures, mimicking the radial sacs above the ventral disc. The dorsal disc has been interpreted as almost completely solid, with the radiating structures identified as very thin tubular canals. This is opposite to what is seen in the ventral disc, which has proportionally larger radiating cavities and slender solid septa. [1]

Body cavity

In most eldonioids, the center of the body is solid, but in other species, there is a rigid central cavity which tapers towards the ventral or dorsal disc. [2] [3] [1] All eldonioids bear a thick, easily-recognizable coiled sac which rings around the middle of the body. The coiled sac is a horseshoe-shaped cavity that curves dextrally (clockwise) from the mouth to the anus. [2] [3] [1] [4] [6] It sheaths the alimentary canal (gut), which is thinner and more difficult to discern in most fossils. In the best-preserved eldoniids, the gut can be divided into three regions: the esophagus or pharynx (front), stomach (middle), and intestine (rear). [2] [1] [4] The stomach is usually the broadest part of the gut, and the portion of the coiled sac surrounding it is stained a dark color. [2] The mouth and anus are positioned close to each other and open through the ventral disc. [1]

Feeding tentacles

A small number of circumoral feeding tentacles project out from near the mouth. Their basic form is shrub-shaped, with a pair of main shafts that split away from each other and divide further into smaller filaments. [3] [1] [4] Some authors referred to the circumoral tentacles as a “lophophore”, a term used for the hollow filamentous tentacles of lophophorates such as brachiopods and bryozoans. [2] [3] [1] In Rotadiscus, a pair of tightly coiled spiral structures connect to the base of the tentacles. These structures covered a hole called the coelomopore, seen in living ambulacrarians. Rotadiscus helps to establish homology between the coelomopore and additional pharyngeal openings present in tunicates and cephalochordates, as well as part of the vertebrate pituitary gland. Moreover, it suggests that the tentacles develop from outgrowths of the coelom (body cavity), more akin to ambulacrarians rather than lophophorates. Further similarities include detachment of the tentacles from the mouth, and their tapering structure. [5]

Taxonomy

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Bryozoa are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about 0.5 millimetres long, they have a special feeding structure called a lophophore, a "crown" of tentacles used for filter feeding. Most marine bryozoans live in tropical waters, but a few are found in oceanic trenches and polar waters. The bryozoans are classified as the marine bryozoans (Stenolaemata), freshwater bryozoans (Phylactolaemata), and mostly-marine bryozoans (Gymnolaemata), a few members of which prefer brackish water. 5,869 living species are known. Originally all of the crown group Bryozoa were colonial, but as an adaptation to a mesopsammal life or to deep-sea habitats, secondarily solitary forms have since evolved. Solitary species has been described in four genera; Aethozooides, Aethozoon, Franzenella and Monobryozoon). The latter having a statocyst-like organ with a supposed excretory function.

<span class="mw-page-title-main">Sipuncula</span> Phylum of invertebrates, peanut worms

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<span class="mw-page-title-main">Hyolitha</span> Palaeozoic lophophorates with small conical shells

Hyoliths are animals with small conical shells, known from fossils from the Palaeozoic era. They are at least considered as being lophotrochozoan, and possibly being lophophorates, a group which includes the brachiopods, while others consider them as being basal lophotrochozoans, or even molluscs.

<span class="mw-page-title-main">Bilateria</span> Animals with embryonic bilateral symmetry

Bilateria is a large clade or infrakingdom of animals called bilaterians, characterized by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front and a rear end, as well as a left–right–symmetrical belly (ventral) and back (dorsal) surface. Nearly all bilaterians maintain a bilaterally symmetrical body as adults; the most notable exception is the echinoderms, which extend to pentaradial symmetry as adults, but are only bilaterally symmetrical as an embryo. Cephalization is also a characteristic feature among most bilaterians, where the special sense organs and central nerve ganglia become concentrated at the front/rostral end.

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<i>Yunnanozoon</i> Cambrian fossil chordate

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<i>Dinomischus</i> Genus of stalked filter-feeding animals

Dinomischusis an extinct genus of stalked filter-feeding animals within the Cambrian period, with specimens known from the Burgess Shale and the Maotianshan Shales. While long of uncertain affinities, recent studies have suggested it to be a stem-group ctenophore.

<span class="mw-page-title-main">Phoronid</span> Phylum of marine animals

Phoronids are a small phylum of marine animals that filter-feed with a lophophore, and build upright tubes of chitin to support and protect their soft bodies. They live in most of the oceans and seas, including the Arctic Ocean but excluding the Antarctic Ocean, and between the intertidal zone and about 400 meters down. Most adult phoronids are 2 cm long and about 1.5 mm wide, although the largest are 50 cm long.

<span class="mw-page-title-main">Brachiopod</span> Phylum of marine animals also known as lamp shells

Brachiopods, phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection. Two major categories are traditionally recognized, articulate and inarticulate brachiopods. The word "articulate" is used to describe the tooth-and-groove structures of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. This is the leading diagnostic skeletal feature, by which the two main groups can be readily distinguished as fossils. Articulate brachiopods have toothed hinges and simple, vertically oriented opening and closing muscles. Conversely, inarticulate brachiopods have weak, untoothed hinges and a more complex system of vertical and oblique (diagonal) muscles used to keep the two valves aligned. In many brachiopods, a stalk-like pedicle projects from an opening near the hinge of one of the valves, known as the pedicle or ventral valve. The pedicle, when present, keeps the animal anchored to the seabed but clear of sediment which would obstruct the opening.

<i>Eldonia</i> Extinct genus of soft-bodied animals

Eldonia is an extinct soft-bodied cambroernid animal of unknown affinity, best known from the Fossil Ridge outcrops of the Burgess Shale, particularly in the 'Great Eldonia layer' in the Walcott Quarry. In addition to the 550 collected by Walcott, 224 specimens of Eldonia are known from the Greater Phyllopod bed, where they comprise 0.43% of the community. Species also occur in the Chengjiang biota, Siberia, and in Upper Ordovician strata of Morocco.

<span class="mw-page-title-main">Asterozoa</span> Phylum of marine invertebrates

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<i>Herpetogaster</i> Extinct genus of Cambrian animals

Herpetogaster is an extinct cambroernid genus of animal from the Early Cambrian Chengjiang biota of China, Blang Formation of China, Pioche Formation of Nevada and Middle Cambrian Burgess Shale of Canada containing the species Herpetogaster collinsi and Herpetogaster haiyanensis.

<i>Stellostomites</i>

Stellostomites is a discoidal animal known from the Cambrian Chengjiang biota and classified with the eldoniids.

<i>Rotadiscus</i> Extinct genus of disc-shaped animal

Rotadiscus is a genus of discoidal animal known from the Cambrian Chengjiang biota and classified with the eldonioids.

<i>Phlogites</i> Genus of ambulacrarians

Phlogites, a senior synonym of Cheungkongella ancestralis, is a cambroernid, and thus a member of the deuterostome total group. It is known from the Lower Cambrian Haikou Chengjiang deposits of China, and was initially compared to the tunicates. It was originally described as a tunicate, though, this identification has been questioned, especially with the discovery of another Chengjiang tunicate, Shankouclava. It has most recently been interpreted as a cambroernid related to Eldonia, Rotadiscus and Herpetogaster.

<span class="mw-page-title-main">Cambroernid</span> Extinct clade of animals

The cambroernids are a clade of unusual Paleozoic animals with coiled bodies and filamentous tentacles. They include a number of early to middle Paleozoic genera noted as "bizarre" or "orphan" taxa, meaning that their affinities with other animals, living or extinct, have long been uncertain. While initially defined as an "informal stem group," later work with better-preserved fossils has strengthened the argument for Cambroernida as a monophyletic clade.

Phoronis psammophila is a species of marine horseshoe worm in the phylum Phoronida. It lives in a tube projecting from the sea floor in shallow seas around the world.

<span class="mw-page-title-main">Cincta</span> Extinct class of marine invertebrates

Cincta is an extinct class of echinoderms that lived only in the Middle Cambrian epoch. Homostelea is a junior synonym. The classification of cinctans is controversial, but they are probably part of the echinoderm stem group.

References

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