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Deuterostomia


, the Green Tree Frog (, , , , )



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The deuterostomes make up on of the earliest divisions with the . Compared to the , the deuterostomes are far less diverse, including the and but having as their main claim to fame the fact that belong to this sub-division. The characters uniting these taxa are mostly developmental (Miller & Harley, 1996). The name Deuterostomia ("second mouth") refers to the development of the embryo into the anus, with the mouth developing as a secondary opening, in contrast to protostomes ("first mouth") with the process reversed. The initial in deuterostomes is , with cleavage planes either parallel or perpendicular to the vertical axis of the embryo, and indeterminate (each early embryonic cell retains the capacity to develop into a complete embryo if isolated from other cells), and development is from outpockets of the gut (). Deuterostomes are often . have supported of the Deuterostomia.

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Deuterostomia topics: | | | | |


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Deuterostomia is here defined as all animals more closely related to (Chordata) and (Echinodermata) than to () or (). It should be noted that "" (the anus developmentally proceeding the mouth) is not unique to Deuterostomia, but is also found in , and . On this page, "Deuterostomia" and "deuterostome" always refer to the , while "deuterostomy" and "deuterostomous" refer to the developmental pathway.

The evolutionary relationships of deuterostomes have gone through a fair amount of contention in recent years. Traditionally. mainly after the work of , deuterostomes have been placed quite high up in the animal tree, forming a clade with the other deuterostomous taxa nestled within the clade. Within the Deuterostomia, chordates and hemichordates were united by possession of and a stiffened -like structure (the notochord in chordates, the in hemichordates). The between hemichordates and echinoderms were regarded as ancestral for deuterostomes, and subsequently lost in chordates.

Humans being the chauvanistic animals we are, there's a lot of appeal in this tree, which appears to sit us at the top of ever-increasing complexity. Unfortunately, recent phylogenies, mostly molecular, rather put the kybosh on this view (Maddison, 2002). Deuterostomia is much lower down, probably the second branch of Bilateria to split off after the . Compared to its sister group, the omnipresent and spectacularly diverse Protostomia (containing the arthropods, molluscs and as its particular success stories), the deuterostomes are merely minor players. Interestingly, while protostomy has been generally regarded as ancestral to deuterostomy, modern phylogenies suggest the opposite, as shown by this phylogeny based on Giribet et al. (2000) where D = deuterostomous and P = protostomous:

0--Deuterostomia (D) 
`--+--Chaetognatha (D)
   `--+--Bryozoa (D) 
      `--+--Ecdysozoa (P) 
         `--+-- (P) 
            `--+--Brachiozoa (D) 
               `--other  (P) 

While deuterostomy probably represents a secondary in Brachiozoa (other phylogenies place Brachiozoa nested within other Trochozoa, reinforcing this probability), protosotomous taxa as a whole are unmistakably nested within deuterostomous taxa.

Within the Deuterostomia, molecular data have argued convincingly for a rearrangement of the main relationships (Cameron et al. 2000). The primary division of living members appears to be between the chordates on one hand and the echinoderms + hemichordates (forming a clade called ) on the other. This suggests that the characters shared between chordates and hemichordates may be ancestral for deuterostomes as a whole, and subsequently lost in echinoderms - a not-too-surprising state of affairs considering the greatly altered morphology of echinoderms. Heinzeller & Welsch (1999) regarded the of echinoderms as homologous with the notochord of chordates, as suggested by similarities in development and gene expression. The -type larva of Ambulacraria can no longer be assumed to be ancestral for Deuterostomia - it may represent an of Ambulacraria.

Recently, a molecular study has suggested that the obscure worm might also fall within the Deuterostomia, specifically as the sister taxon to Ambulacraria. We therefore give it a look here. Also covered are the , a group of fossils that may represent the stem-group of crown deuterostomes, as well as a number of fossil groups regarded as stem echinoderms.

Not covered for now are a few other fossil groups that have been less certainly referred to Deuterostomia at one time or another, such as , and . Also absent are the (arrow-worms), which have been considered the basalmost branch of the Deuterostomia in the past, but are currently regarded by many as basal protostomes (Helfenbein et al., 2004).



  

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