Headshield slugs | |
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A live individual of Chelidonura varians in the family Aglajidae, head end towards the lower left | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Mollusca |
Class: | Gastropoda |
Subclass: | Heterobranchia |
Cohort: | Tectipleura |
Order: | Cephalaspidea |
Superfamilies | |
See text |
The order Cephalaspidea, also known as the headshield slugs and bubble snails, is a major taxon of sea slugs and bubble snails, marine gastropod mollusks within the larger clade Euopisthobranchia. [1] Bubble shells is another common name for these families of marine gastropods, some of which have thin bubble-like shells. [2] This clade contains more than 600 species. [3]
Members of this worldwide clade used to be considered the most ancestral of the opisthobranchs, but now they are considered as derived and specialized members of the Euthyneura Spengel, 1881. [4]
Headshield slugs are the most morphologically diverse group of all the opisthobranchs.[ citation needed ]
The vast majority possess a shell, although it may be reduced or internal. They have a well-developed headshield, a characteristic broadening at the head, which is used to plow beneath the surface of the sand. This headshield prevents the sand entering the mantle cavity. There is a muscular foot with or without parapodia (fleshy wing-like flaps).
Headshield slugs often live just beneath the surface of the sand and can also be seen crawling on rocks. They have well-developed sensory structures to detect prey, which may be other opisthobranchs, polychaetes or bristleworms and foraminiferans. Several species are voracious carnivores.
Members of the brightly colored genus Chelidonura also have well-developed eyes on the anterior end of the head and bundles of sensory cilia around the mouth. With these cilia they are able to track their prey by following the victim's mucous trail.
The Hancock's organ is a chemosensory organ situated between the foot and the headshield. It plays a role in olfactory and sensory detection. It is visible as a dark brown pit at the base of the right rhinophore.
The taxonomy of the shelled cephalaspideans, the bubble snails, like that of many shelled mollusks, used to be based very simply on shell characteristics. But because there are some similarities in shell morphology throughout this group, more recently taxonomists have taken other anatomical characteristics into consideration, such as the radula, gizzard, penis, and Hancock's organ.
In 2015 a new study based on molecular phylogenetics has changed significantly the taxonomy of the Cephalaspidea. [3] The monophyly of the Cephalaspidea was confirmed, but the families Cylichnidae, Diaphanidae, Haminoeidae, Philinidae, and Retusidae were found non-monophyletic. This had led to the creation of new families ((Alacuppidae, Colinatydidae, Colpodaspididae, Mnestiidae, Philinorbidae) ) and one new genus ( Alacuppa ). Two family names (Acteocinidae, Laonidae) and two genera ( Laona , Philinorbis ) are reinstated as valid
In the taxonomy of Bouchet & Rocroi (2005), the clade Cephalaspidea is arranged as follows:
The superfamily Acteonoidea has been included into the new Informal Group "Lower Heterobranchia" and the superfamily Cylindrobulloidea becomes part of the Group Cylindrobullida.
Malaquias et al. (2009) [5] have rearranged taxonomy of Cephalaspidea sensu lato:
The taxonomy of Cephalaspidea sensu lato by Malaquias et al. (2009) [5] is arranged as follows (there are listed genera under molecular analysis; not analyzed families are under "incertae sedis"):
Architectibranchia Haszprunar, 1985
Runcinacea Burn, 1963
Cephalaspidea Fischer, 1887 – This means Cephalaspidea sensu stricto
Subsequently, Malaquias (2010) [6] moved Bullacta exarata (formerly the only member of Bullactidae) into the family Haminoeidae. [6]
Jörger et al. (2010) [1] moved Cephalaspidea sensu stricto and Runcinacea into the Euopisthobranchia and they confirmed the placement of Acteonoidea within the Lower Heterobranchia. [1] All families of Architectibranchia were already within the Lower Heterobranchia in the taxonomy of Bouchet & Rocroi, except for the Notodiaphanidae, which has been placed in the Lower Heterobranchia since 2010, in order that the Architectibranchia can be considered to be monophyletic.
The publication by Oskars T.R., Bouchet P. & Malaquias M.A. (2015). A new phylogeny of the Cephalaspidea (Gastropoda: Heterobranchia) based on expanded taxon sampling and gene markers. in the journal Molecular Phylogenetics and Evolution. 89 came to the following conclusion, with the creation of new families [3]
The superfamily Bulloidea was not supported in the Bayesian phylogenetic hypothesis and Diaphanoidea was found polyphyletic. The superfamilies Haminoeoidea and Philinoidea were accepted. The composition of each of the superfamilies was drastically rearranged.
Diaphanoidea is a taxonomic superfamily of small sea snails, marine opisthobranch gastropod mollusks or micromollusks in the Cephalaspidea, the headshield slugs and bubble snails.
Notodiaphana fragilis is a species of marine snail, gastropod mollusk in the clade Architectibranchia.
Haminoeoidea is a taxonomic superfamily of small sea snails or bubble shells, marine opisthobranch gastropod mollusks in the clade Cephalaspidea, the headshield slugs and bubble snails.
Haminoeidae, commonly known as the haminoeid bubble snail family, is a taxonomic family of sea snails, marine opisthobranch gastropod mollusks in the superfamily Haminoeoidea.
Philinoidea is a taxonomic superfamily of sea slugs, specifically headshield slugs, marine gastropod mollusks in the order Cephalaspidea.
Aglajidae is a family of often colorful, medium-sized, sea slugs, marine opisthobranch gastropod mollusks. These are not nudibranchs; instead they are headshield slugs, in the clade Cephalaspidea.
Bulloidea is a superfamily of sea snails, or bubble snails, marine gastropod mollusks.
Philinidae is a family of medium-sized sea slugs, marine opisthobranch gastropod mollusks. These are headshield slugs, in the order Cephalaspidea.
Cylichnidae, common name the "chalice bubble snails" or "canoe bubble snails" is a family of sea snails or bubble snails, marine gastropod mollusks in the superfamily Cylichnoidea.
Runcinoidea is a taxonomic superfamily or a clade Runcinaecea of sea slugs, marine gastropod mollusks in the order Runcinida
This overview lists proposed changes in the taxonomy of gastropods at the family level and above since 2005, when the taxonomy of the Gastropoda by Bouchet & Rocroi (2005) was published. In other words, these are recent updates in the way various groups of snails and slugs are classified.
Scaphandridae is a family of sea snails, marine gastropod mollusks in the superfamily Philinoidea.
Architectibranchia is a clade of marine snails, gastropod molluscs.
Rhizoridae is a family of very small sea snails, barrel-bubble snails, marine opisthobranch gastropod molluscs. These are headshield slugs, in the superfamily Bulloidea.
Tornatinidae is a family of very small sea snails, barrel-bubble snails, marine opisthobranch gastropod molluscs. These are headshield slugs, in the superfamily Bulloidea.
Smaragdinella is a genus of medium-sized sea snails or bubble snails, marine opisthobranch gastropod molluscs in the family Haminoeidae, the haminoea bubble snails, part of the clade Cephalaspidea, the headshield slugs and bubble snails.
Biuve fulvipunctata, the white-speckled headshield slug, is a species of sea slug or headshield slug, a marine opisthobranch gastropod mollusc in the family Aglajidae. This species is widespread in the Indian and Pacific Oceans but has invaded the Mediterranean Sea since 1961, despite apparently being absent from the Red Sea until recorded there in the 21st century. It is the only species in the monotypic genus Biuve.
Cylichnoidea is a superfamily of sea snails or bubble snails, marine gastropod molluscs in the order Cylichnidae, the "chalice bubble snails".
Laonidae is a family of small marine snails belonging to the superfamily Philinoidea, though Laonidae and Philinoidea are genetically distinct. They are monophyletic, meaning they are developed from a single ancestor. Laonidae also has the synonymised name Laoninae, which is largely unaccepted in the modern day scientific community. The name Laoninae is introduced by Alice Pruvot-Fol who proposes it as the subfamily to the genea Loana, A.Adams 1865.
Phanerophthalmus is a genus of medium-sized sea snails or bubble snails, marine opisthobranch gastropod molluscs in the family Haminoeidae, the haminoea bubble snails, part of the clade Cephalaspidea, the headshield slugs and bubble snails.