Diversity and Distribution of Cyatholaimidae (Chromadorida: Nematoda): A Taxonomic and Systematic Review of the World Records

Abstract

Nematoda is a very species-rich phylum that has successfully adapted to almost all types of ecosystems. Despite their abundance and ecological importance, the taxonomic knowledge of nematodes is still limited and the identification of species is not trivial. In Cyatholaimidae, a relatively common and abundant family of free-living nematodes, the identification of organisms is challenging due to the overlap of some generic diagnoses and the absence of updated systematic reviews. Here we systematically reviewed the knowledge about the family diversity, providing a list of valid species, the diagnostic characters to genus level, and the geographical and habitat distribution of species. The review systematized a total of 619 records. The occurrences were classified into biogeographic ecoregions and habitats. Cyatholaimidae includes 211 valid species, classified in 20 genera. The genera can be differentiated based on six diagnostics characters, namely: pattern of cuticle ornamentation; number of longitudinal rows of pore-complex in cuticle; structures of the buccal cavity; presence/absence of pharyngeal bulb; pre-cloacal supplements aspect; and the shape of gubernaculum. Cyatholaimidae includes mainly marine species, mostly occurring in the Coastal Zone. Four and three species were registered in freshwater and terrestrial habitats, respectively, all classified in the genus Paracyatholaimus. About 38% of the valid species occur in more than one type of habitat, under very different environmental conditions, suggesting a broad niche. The occurrence of congeneric species in different habitats types indicates that, throughout the evolutionary history of the family, multiple ecological shift events have occurred. The family occurs worldwide in 74 ecoregions, and the majority of the records and species are in the North Sea and Western Mediterranean. Most species are endemic to one ecoregion, and examples of broadly distributed ones may be a result of misidentifications or cases of long-distance dispersal, especially for those associated with biological substrates.

Two new species of Cyatholaimidae (Nematoda: Chromadorida) from the Southeastern Brazilian coast with emphasis on the pore complex and lateral pore-like structures

“…23–24 + 16 µm) and by the presence of lateral differentiation in the cuticle. The strong cuticularized vagina of females and the presence of eight longitudinal rows of pore complex in the cuticle (there are four rows in the other species) were observed only in the new species and in B. hopperi ( Sharma & Vincx, 1982 ) Cunha, Fonseca & Amaral, 2022 among all species of the genus. It differs from the new species by the absence of lateral differentiation in the cuticle and by the shape of the copulatory apparatus.…”

“…The cuticle ornamentation of enlarged hexagonal punctations in the anterior end is a unique characteristic of the genus. Based on that, the species Marylynnia dayi ( Inglis, 1963 ) Hopper, 1972 , M. hopperi Sharma & Vincx, 1982 , and M. punctata Jensen, 1985 were transferred to Biarmifer by Cunha, Fonseca & Amaral (2022) . Besides cuticle ornamentation, both genera may be differentiated by the number of longitudinal rows of pore complexes.…”

“…Based on morphology, it has been subdivided into four subfamilies (Cyatholaiminae Filipjev, 1918 , Paracanthonchinae De Coninck, 1965 , Pomponematinae Gerlach & Riemann, 1973 and Xenocyatholaiminae Gerlach & Riemann, 1973 ), however, none are supported by any synapomorphy ( Lorenzen, 1981 ). The same is true for many genera of the family, which are delimited by characters that may not be phylogenetically relevant ( Cunha, Fonseca & Amaral, 2022 ). Therefore, taxonomic studies of Cyatholaimidae are marked by changes in classifications, synonyms and transfers of species between genera ( e.g ., Micoletzky, 1924 ; Wieser, 1954 ; Miljutina & Miljutin, 2015 ).…”

“…The family Cyatholaimidae Filipjev, 1918 is a diverse group of mainly marine nematodes with 20 genera and 211 valid species distributed worldwide in different types of habitats ( Cunha, Fonseca & Amaral, 2022 ). Based on morphology, it has been subdivided into four subfamilies (Cyatholaiminae Filipjev, 1918 , Paracanthonchinae De Coninck, 1965 , Pomponematinae Gerlach & Riemann, 1973 and Xenocyatholaiminae Gerlach & Riemann, 1973 ), however, none are supported by any synapomorphy ( Lorenzen, 1981 ).…”

“…The genus was only found again over 40 years later when Pastor de Ward (2001) described Biarmifer madrynensis . Based on the cuticle ornamentation, which is unique to this group, it was recently proposed the reclassification of three species belonging to Marylynnia ( Hopper, 1972 ) Hopper, 1977 to Biarmifer ( Cunha, Fonseca & Amaral, 2022 ).…”