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Jump to: ,

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Deuterostomia topics: | | | | |


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Here at Palaeos, we point out right from the start that we are not authorities in the subjects covered, and anyone reading this site should approach its statements with all the caution of a serious burn victim re-potting cacti. Sometimes, though, a particularly abstruse topic is broached that requires us to remind you of this rule. This is one of those times. The is a repository for what are patently some of the most bizarre organisms ever to clutter the sea-bed. If, indeed, they are stem echinoderms at all. They may be derived crown echinoderms, or stem , or organisms with no real connection to the echinoderms at all. Interpretations of these fossils are so varied and so divergent that one could be forgiven for failing to realize that they referred to the same animals. In the case of the , researchers can't even agree as to which end is anterior and which is posterior.

As a result, this section is a pastiche. With so many competing theories available, I have picked and chosen as seems appropriate (or as my whim takes me), and in the process I have quite likely constructed something which no researcher has supported, and to which no-one who has actually examined the fossils in question would give the time of day. My advice is to stop reading. Go straight to the bibliography, and get your information direct from the primary literature. You'll be better off.

For those who have chosen to ignore my advice, a few notes will be given on terminology. In light of the highly debatable nature of the echinoderm stem, I have decided to restrict the name “Echinodermata” to the only, allowing me to be more explicit about the origins of characters leading towards the modern body plan. This differs from the definition (often not explicitly stated) used in many publications.

Probably the majority (including the Treatise on Invertebrate Paleontology) identify Echinodermata by the origin of the , the skeleton with an apparently unique to this clade. However, Jefferies (e.g. 1986) and his supporters, who place the Stylophora as the stem group of chordates, would see the stereom as plesiomorphic for a much larger clade. If defined by the stereom, Echinodermata would then include Chordata.

Another alternative is defining Echinodermata by the origin of . However, this character is often widely altered in both modern and fossil groups of echinoderms. Also, the 2-1-2 system of in demonstrates that and pentaradial symmetry did not necessarily arise simultaneously.

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As an initial matter, the echinoderms and chordate phyla have been arranged in almost every possible order. Classical taxonomy was of no help, since we lacked enough data on basal to make even educated guesses about interrelationships. It was like trying to do a connect-the-dots picture with most of the dots missing and half the others in the wrong place. This confusion was succeeded by a large number of molecular studies reported by folks who had a whole lot of dots, but were a little unclear on how to connect them. Finally -- really within the last five years -- both new fossils and great improvements in molecular techniques have allowed us to put the big pieces together with confidence. The overall picture looks like this:

Deuterostomia
|--Ambulacraria
|  |--
|  `--
`--
   |--
   `--+--
      `--

We are particularly encouraged by studies like Furlong & Holland (2002) who re-analyzed prior sequence work with sophisticated , and obtained the same result with a great variety of individual sequences. See also Winchell et al. (2002) (same results, concatenating different sequences). Sequence phylogenies are still hit-and-miss; but the new fossils make it much easier to assess which have hit and which have missed.

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Arkarua adami is a small disc-shaped fossil, about 3 to 10 mm across, known from the of the Flinders Ranges in Australia (UCMP). All known examples are external moulds with a five-lobed central region that led to its identification as the earliest echinoderm. However, Arkarua lacks a stereom or any sign of a water-vascular system.

Arkarua's position as ancestral echinoderm has been widely accepted. The problem is that a number of other echinoderm stem groups ( and ) that appear to predate the origin of pentaradiality, but possess a stereom. As pointed out by Mooi (2001), this paradox has persisted because the organisms in question have rarely been compared. Those analyses that included Arkarua generally omitted the non-pentaradial taxa, and vice versa. There are two possible options if Arkarua is a stem echinoderm: (1) Arkarua is more crownward than the non-pentaradial taxa, and either the stereom and water-vascular system evolved independently on a number of occasions (very unlikely, as it would require the repeated appearance of an ultrastructure and a complex organ system that have not been recorded from any other clade) or Arkarua secondarily lost the stereom and water-vascular system (also unlikely, as it would require the complete absence of preservation of any contemporary forms retaining the stereom) (2) The non-pentaradial forms are secondarily derived from pentaradial ancestors. This latter is not impossible. Mooi (2001) suggests that unusual taxa are often placed too far down in trees by the difficulty of recognizing synapomorphies with less derived taxa. Compare, for instance, the recent transfer of from their originally basal position in the eukaryote tree to a much higher one within the . And some recent cladistic analyses have placed homalozoans within the echinoderm crown as derived . Unfortunately any cladistic analysis is only as good as the characters it uses and the analyses in question were based on anatomical interpretations different from those favored here. The recently discovered , which lack a stereom but have a comparable organization to Stylophora, appear to support a basal position for at least some non-pentaradial taxa.

The more likely explanation is that Arkarua's pentaradial symmetry is not homologous to that of echinoderms. After all, this is the single character uniting the two. Arkarua is therefore not here accepted as a deuterostome, and is more likely a cnidarian-grade animal or possibly even an alga.

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The Echinoderm Stem CKT050827; The Monophyly of Ambulacraria ATW051025, Arkarua CT050827

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