Bdelloidea (Greek βδέλλα, bdella, "leech") is a class of rotifers found in freshwater habitats all over the world. There are over 450 described species of bdelloid rotifers (or 'bdelloids'), distinguished from each other mainly on the basis of morphology. The main characteristics that distinguish bdelloids from related groups of rotifers are exclusively parthenogenetic reproduction and the ability to survive in dry, harsh environments by entering a state of desiccation-induced dormancy (anhydrobiosis) at any life stage. They are often referred to as "ancient asexuals" due to their unique asexual history that spans back to over 25 million years ago through fossil evidence. Bdelloid rotifers are microscopic organisms, typically between 150 and 700 µm in length. Most are slightly too small to be seen with the naked eye, but appear as tiny white dots through even a weak hand lens, especially in bright light. In June 2021, biologists reported the restoration of bdelloid rotifers after being frozen for 24,000 years in the Siberian permafrost.
Evolutionary relationships
The phylum Rotifera traditionally included three classes: Bdelloidea, Monogononta and Seisonidea. Prior to 1990, phylogenetic studies based on morphology seemed to indicate that the sister group to bdelloid rotifers was Monogononta, with seisonid rotifers as an early-diverging outgroup.
Modern molecular phylogenetic studies demonstrate that this classic understanding of 'Rotifera' is incomplete (paraphyletic), because it omits a fourth clade of closely related organisms: the Acanthocephala, or thorny-headed worms. Originally classified as a separate phylum, molecular and morphological evidence accumulated between 1994 and 2014 to indicate that Acanthocephala forms a monophyletic group with Bdelloidea, Monogononta and Seisonidea. To accommodate this finding, some authors extend the term 'Rotifera' to include the highly modified, parasitic 'acanthocephalan rotifers' alongside bdelloid, monogonont and seisonid rotifers. Others refer to the grouping of the four taxa as Syndermata, a term derived from their shared syncytial epidermis.
The position of Bdelloidea within Syndermata (or Rotifera) is not entirely clear. Alternative possible phylogenetic relationships within the clade are illustrated by the accompanying cladograms. As of 2014, the "most comprehensive phylogenomic analysis of syndermatan relationships" to date was based on transcriptome data from all four groups, and provided "strong support" for the hypothesis illustrated in the bottom left of the figure, in which Seisonidea and Acanthocephala are sister taxa. The study further indicated that the sister group to this taxon is Bdelloidea, whereas Monogononta is the outgroup to all three. This would mean that the closest living relatives of bdelloid rotifers are not monogonont rotifers, as previously believed, but seisonid rotifers and acanthocephalans, despite their highly modified morphology.
Classification and identification
Bdelloidea is a class of the phylum Rotifera, consisting of three orders: Philodinavida, Philodinida and Adinetida. These orders are divided into four families and about 450 species. Since these organisms are asexual the usual definition of a species as a group of organisms capable of creating fertile offspring is inapplicable, therefore the species concept in these organisms is based on a mixture of morphological and molecular data instead. DNA studies suggest that the diversity is much greater than the original morphological classifications suggest.
Bdelloids can only be identified by eye while they are alive because many of the characteristics significant to classification are related to feeding and crawling; however, genetic identification of bdelloids is possible on dead individuals. Once preserved, the individuals contract into "blobs" which restricts analysis. There are currently three morphological identification methodologies, two of which are considered dated: Bartoš (1951) and Donner (1965). The third method is a diagnostic key developed in 1995 by Shiel.
Morphology
There are three main regions of the body of bdelloids: head, trunk and foot. The adjacent image depicts each area to show how body parts can be very different although they are named the same depending on the species involved. Bdelloids typically have a well-developed corona, divided into two parts, on a retractable head.