Evolution and systematics
The phylum Rhombozoa includes three families, eight genera, and approximately 150 species. The three families are Conocyemidae, Dicyemidae, and Kantharellidae.
Edouard van Beneden proposed the name Mesozoa for these organisms as an intermediate between Protozoa and Metazoa in body organization. Later zoologists considered rhombozoans degenerated from metazoans such as trematodes because of adaptation for the parasitic lifestyle. Results of phylogenetic studies with nucleotide sequences of 18S recombinant DNA and Hox gene have suggested rhombozoans are spiralians (protostomes) such as turbellarians, nemerteans, annelids, and mollusks. Many scientists now place this group within the phylum Dicyemida.
Rhombozoans have two phases of body organization—the vermiform stages of vermiform embryo and adult (nemato-gen, rhombogen) and the infusoriform embryo. The body of vermiform rhombozoans consists of a central cylindrical cell called the axial cell and a layer of 8-30 ciliated external cells called peripheral cells. The number of peripheral cells is fixed and species specific. At the anterior region, 4-10 peripheral cells form the calotte, the cilia of which are shorter and denser than in more posterior peripheral cells. The calotte comprises two tiers of cells—propolar cells and metapolar cells. Calotte shape varies with species.
Infusoriform embryos consist of 37 or 39 cells, which are more differentiated than those of vermiform organisms. Inside the embryo are four large cells called urn cells, each containing a germinal cell that may give rise to the next generation. At the anterior region of the embryo is a pair of unique cells called apical cells, each containing a refringent body composed of magnesium inositol hexaphosphate. At the posterior region, external cells are ciliated.
Temperate and subtropical continental waters. Okhotsk Sea; Sea of Japan; northern, eastern, and western Pacific Ocean; New Zealand; Australia. Mediterranean Sea; northern, eastern, western Atlantic Ocean; Gulf of Mexico; Antarctic Ocean.
Vermiform stages of rhombozoans are restricted to the renal sac of cephalopods. In decapods, rhombozoans are also found in the renopancreatic coelom. Some rhombozoans are found in the pericardium of decapods. On the surface of the renal appendage, vermiform rhombozoans insert their heads into renal tubules and folds.
Vermiforms and infusoriform embryos swim with their cilia. There appears to be positive thigmotaxis to renal appendages in the vermiform stages.
The surface of the rhombozoan body possesses numerous cilia and has a folded structure believed to contribute to absorption of nutrients from the urine of hosts.
The vermiform embryo develops asexually from an agamete (axoblast) and grows into an adult. A high population density in the host renal sac can cause a shift from an asexual mode to a sexual mode of reproduction. The functionally hermaphroditic gonad, the infusorigen, forms at high population density. Mature spermatozoa without tails fertilize the pri-
mary oocytes. A fertilized egg develops into an infusoriform embryo. It remains to be understood how infusoriform larvae infect the new host and develop into vermiforms.
No species is listed by the IUCN.
Significance to humans
Dicyema acuticephalum taken from its host Octopus vulgaris. (Specimens in McConnaughey collection, Santa Barbara Museum of Natural History. Photo by Hidetaka Furuya. Reproduced by permission.)
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