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''Inchoatia'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of air-breathing
land snail A land snail is any of the numerous species of snail that live on land, as opposed to the sea snails and freshwater snails. ''Land snail'' is the common name for terrestrial molluscs, terrestrial gastropod mollusks that have gastropod shell, shel ...
s,
terrestrial Terrestrial refers to things related to land or the planet Earth, as opposed to extraterrestrial. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on o ...
pulmonate Pulmonata or pulmonates is an informal group (previously an order, and before that, a subclass) of snails and slugs characterized by the ability to breathe air, by virtue of having a pallial lung instead of a gill, or gills. The group inclu ...
gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...
mollusk Mollusca is a phylum of protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000  extant species of molluscs are recognized, making it the second-largest animal phylum after Arthropoda. The ...
s in the family
Clausiliidae Clausiliidae, also known by the common name door snails, is a Taxonomy (biology), taxonomic family (biology), family of small, very elongate, mostly Gastropod shell#Chirality in gastropods, left-handed, air-breathing land snails, sinistral terre ...
, the door snails.MolluscaBase eds. (2020). MolluscaBase. Inchoatia E. Gittenberger & Uit de Weerd, 2006. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=996065 on 2020-12-15


Description

The species classified within the genus ''Inchoatia'' are conchologically similar. They all have slender to very slender, small to medium-sized shells with more or less prominent papillae along the suture. The clausilial apparatus is of the so-called "N-type", with a lamella spiralis and a plica principalis. As in genus ''
Albinaria ''Albinaria'' is a genus of air-breathing land snails, terrestrial pulmonate gastropod mollusks in the family Clausiliidae, called the door snails. Ecology and Life Cycle These species of snails live on limestone rocks, where they feed on alga ...
'', the
genital tract The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are al ...
shows a dimorphism in penial structure, with either a papilla or a caecum.


Distribution

The species within this genus occur in
limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...
areas of the central and western part of mainland
Greece Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...
, where some taxa are restricted to high altitudes.


Taxonomy

Some ''Inchoatia'' taxa were insufficiently known, even conchologically (in terms of a shell description) and were classified without clear reference to the underlying facts. In the past (Nordsieck 1972, Nordsieck 1974, Zilch 1981) species of the Alopiinae were classified in the genus'' Carinigera'' or '' Sericata'', mainly on the anatomical basis of having either a penial papilla or a caecum. This dogmatic approach led to a grouping together of clausiliid species with very different shells, which was eventually unmasked as unnatural by DNA analyses. This position is however not embraced by Nordsieck (2007),Nordsieck H. (2007). ''Worldwide door snails (Clausiliidae), recent and fossil''. 1-214. ConchBooks, Hackenheim. who lumped together several genus-group taxa, which "can be characterized neither by shell nor by genital characters", in his view. The genus ''Inchoatia'' was introduced by Gittenberger & Uit de Weerd (2006)Gittenberger E. & Uit de Weerd D. R. (2006). "The genus ''Inchoatia'' validated and three mistakes corrected". ''Basteria'' 70: 131-132. on the basis of molecular phylogenetic analyses by Uit de Weerd et al. (2004).Uit de Weerd D.R., Piel W. H. & Gittenberger E. (2004). "Widespread polyphyly among Alopiinae snail genera: when phylogeny mirrors biogeography more closely than morphology". ''
Molecular Phylogenetics and Evolution ''Molecular Phylogenetics and Evolution'' is a peer-reviewed scientific journal of evolutionary biology and phylogenetics. The journal is edited by E.A. Zimmer. Indexing The journal is indexed in: * EMBiology *Journal Citation Reports *Scopus ...
'' 33(3): 533-548.
This molecular phylogenetic study by Uit de Weerd et al. (2004) was not designed to answer questions about the status of the species group taxa within ''Inchoatia''. The fact that the ''Inchoatia'' taxa are very disjunctly distributed means that attempting to discriminate species as against subspecies is even more of a subjective opinion than usual, because
reproductive isolation The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, ethology, behaviors and physiology, physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensu ...
under natural conditions cannot be observed. An objective or a generally accepted method to weigh morphological differences does not exist. DNA analyses may be extremely helpful, but cannot always be considered decisive with regard to species or subspecies status.
Molecular phylogeny Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
reconstructions made clear that reproductive isolation and lineage splitting do not always run entirely parallel, resulting in paraphyletic species. In ''Inchoatia'', with some island-like distributions of presumably high alpine taxa, this may also be the case. Besides
synapomorphic In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
nucleotide Nucleotides are Organic compound, organic molecules composed of a nitrogenous base, a pentose sugar and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both o ...
substitutions, these taxa share a similar shell shape, and have more or less continuous
ranges In the Hebrew Bible and in the Old Testament, the word ranges has two very different meanings. Leviticus In Leviticus 11:35, ranges (כירים) probably means a cooking furnace for two or more pots, as the Hebrew word here is in the dual numbe ...
. According to the molecular data, the major division within ''Inchoatia'' is between an ''Inchoatia inchoata'' + "''Inchoatia regina''"
subclade In genetics, a subclade is a subgroup of a haplogroup. Naming convention Although human mitochondrial DNA (mtDNA) and Y chromosome DNA (Y-DNA) haplogroups and subclades are named in a similar manner, their names belong to completely separate syst ...
, and an ''Inchoatia haussknechti'' + ''Inchoatia megdova'' subclade. This pattern is congruent with genital anatomical differences between the two subclades, previously used to place them in separate genera '' Sericata'' and '' Carinigera'' and with conchological similarities between the species within each subclade. The
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
relationships between nominal subspecies within genus ''Inchoatia'' were examined by Uit de Weerd et al. (2009). A
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
based on sequences of cytochrome-c oxidase I (COI) genes showing phylogenic relations of ''Inchoatia'' by Uit de Weerd et al. (2009) (simplified):


Species

According to Gittenberger & Uit de Weerd (2009) and Uit de Weerd et al. (2009) the genus ''Inchoatia'' includes three species, each of which has a number of subspecies: * '' Inchoatia haussknechti'' (O. Boettger, 1886) - synonym: ''Carinigera haussknechti'' ** ''Inchoatia haussknechti haussknechti'' (O. Boettger, 1886) ** ''Inchoatia haussknechti alticola'' (Nordsieck, 1974) ** ''Inchoatia haussknechti hiltrudae'' (Nordsieck, 1974) ** ''Inchoatia haussknechti orina'' (Westerlund, 1894) ** ''Inchoatia haussknechti refuga'' (Westerlund, 1894) ** ''Inchoatia haussknechti semilaevis'' (O. Boettger, 1889) * '' Inchoatia inchoata'' (O. Boettger, 1889) - type species, synonym: ''Sericata inchoata'' ** ''Inchoatia inchoata inchoata'' (O. Boettger, 1889) ** ''Inchoatia inchoata klemmi'' (Nordsieck, 1972) ** ''Inchoatia inchoata paramythica'' (Nordsieck, 1974) ** ''Inchoatia inchoata regina'' (Nordsieck, 1972) * '' Inchoatia megdova'' (Nordsieck, 1974) - synonym: ''Carinigera megdova'' ** ''Inchoatia megdova megdova'' (Nordsieck, 1974) ** ''Inchoatia megdova bruggeni'' Gittenberger & Uit de Weerd, 2009 ** ''Inchoatia megdova palatalifera'' (Hausdorf, 1987) ** ''Inchoatia megdova tavropodensis'' (Fauer, 1993)


References

This article incorporates CC-BY-3.0 text from references.Gittenberger E. & Uit de Weerd D. R. (2009). "Summarizing data on the ''Inchoatia'' taxa, including ''Inchoatia megdova bruggeni'' subspec. nov. (Gastropoda, Pulmonata, Clausiliidae)". ''
Zoologische Mededelingen ''Zoologische Mededelingen'' was a peer-reviewed open access scientific journal publishing papers and monographs on animal systematics. The publisher was the National Museum of Natural History Naturalis in the Netherlands. The first issue appeared ...
'' 83 http://www.zoologischemededelingen.nl/83/nr03/a08
Uit de Weerd D. R., Schneider D. & Gittenberger E. (2009). "Molecular phylogenetic relationships of ''Inchoatia'' taxa". ''
Zoologische Mededelingen ''Zoologische Mededelingen'' was a peer-reviewed open access scientific journal publishing papers and monographs on animal systematics. The publisher was the National Museum of Natural History Naturalis in the Netherlands. The first issue appeared ...
'' 83: 589-592. http://www.zoologischemededelingen.nl/83/nr03/a09
* Bank, R. A. (2017). Classification of the Recent terrestrial Gastropoda of the World. Last update: July 16, 2017. {{Taxonbar, from=Q6014504 Clausiliidae