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Making the most of phylogeny - OPUS - Universität Würzburg

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The phylum Tardigrada Part I. Introduction<br />

time <strong>of</strong> one year. Suzuki (2008) reported a low frequency <strong>of</strong> males in <strong>the</strong> culture <strong>of</strong><br />

M. tardigradum.<br />

The eggs are colourless or brown and <strong>the</strong> diameter ranges from 70–110 µm<br />

(Kinchin 1994). Suzuki (2003) found that <strong>the</strong> <strong>most</strong> rapidly developing animal laid<br />

eggs 12 days after hatching.<br />

Hypsibius dujardini<br />

Hypsibius dujardini is an obligatory par<strong>the</strong>nogenetic species (Ammermann 1967).<br />

H. dujardini can be cryopreserved and cultured continuously for long time. The<br />

genome size <strong>of</strong> about 78 Mbp (Gregory et al. 2007) is one <strong>of</strong> <strong>the</strong> smallest genomes<br />

<strong>of</strong> animals (Gabriel et al. 2007). This is about three-quarters <strong>of</strong> C. elegans and less<br />

than half <strong>the</strong> size <strong>of</strong> <strong>the</strong> D. melanogaster genome. The generation time is about<br />

14 days at room temperature, <strong>the</strong> embryos have a stereotyped cleavage pattern<br />

with asymmetric cell divisions, nuclear migrations, and cell migrations occurring in<br />

reproducible patterns.<br />

The rate <strong>of</strong> protein evolution in H. dujardini is similar to <strong>the</strong> rate in D. melanogaster<br />

and, thus similar to <strong>most</strong> o<strong>the</strong>r known genomes in metazoans (Gabriel et al. 2007).<br />

Only sparse molecular data are available, but Bavan et al. (2009) described a purinergic<br />

receptor expressed in H. dujardini and Gabriel and Goldstein (2007) examined<br />

<strong>the</strong> expression <strong>of</strong> Pax3/7 during embryogenesis.<br />

Richtersius coronifer<br />

Richtersius coronifer, formerly Adorybiotus or Macrobiotus, is a large tardigrade with<br />

body sizes up to 1 mm (Kinchin 1994). R. coronifer is a bryophilic (moss-living)<br />

tardigrade, which can be found all over <strong>the</strong> world mainly in alpine and arctic<br />

environments up to 5600 m (Ramløv and Westh 2001).<br />

R. coronifer can survive severe dehydration for years (Ramløv and Westh 2001)<br />

and is capable <strong>of</strong> tolerating temperatures down to −196 ◦ C independent from <strong>the</strong><br />

actual state (tun or normal living) (Ramløv and Westh 1992; Westh and Ramløv<br />

1991). The disaccharide trehalose is accumulated up to approximately 2.3 % dry<br />

weight during desiccation (Westh and Ramløv 1991).<br />

Genus Paramacrobiotus<br />

Recently <strong>the</strong> genus Paramacrobiotus was erected within <strong>the</strong> family <strong>of</strong> Macrobiotidae<br />

using morphological characters and gene sequences. Within <strong>the</strong> genus several<br />

cryptic species from different places around <strong>the</strong> world could be found, which can<br />

not be clearly separated by morphological or common molecular markers alone<br />

(Guidetti et al. 2009). This cryptic species have been already used for biochemical<br />

6

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