''Beddomeia fultoni'' ''(B. fultoni)'' is a

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

of small

freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...

snail belonging to the family

Tateidae Tateidae is a family of very small and minute aquatic snails with an operculum, gastropod mollusks in the superfamily Truncatelloidea.Bouchet, P. (2014). Tateidae Thiele, 1925. Accessed through: World Register of Marine Species at http://www.ma ...

. The species is

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...

to Australia. Commonly known as Farnhams Creek hydrobiid snail, the species used to belong to the

Hydrobiidae Hydrobiidae, commonly known as mud snails, is a large cosmopolitan family of very small freshwater and brackish water snails with an operculum; they are in the order Littorinimorpha. Distribution Hydrobiidae are found in much of the world, ...

family ''sensu lato (s.l.)'' and is listed as one of 37 threatened ''

Beddomeia ''Beddomeia'' is a genus of very small freshwater snails that have a gill and an operculum, aquatic operculate gastropod mollusks in the family Tateidae. Species Species within the genus ''Beddomeia'' include: *'' Beddomeia acheronensis'' * ...

'' species on the Tasmanian ''

Threatened Species Protection Act 1995 The ''Threatened Species Protection Act 1995'' (TSP Act), is an act of the Parliament of Tasmania that provides the statute relating to conservation of flora and fauna. Its long title is An Act to provide for the protection and management of th ...

''. Found within north-western

Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...

, the species has a somewhat small range. Previously, they have only been identified in sites within six different streams which are separated by a maximum of 4km.Richards, K. (2010) ''An ecological, morphological and molecular investigation of Beddomeia species (Gastropoda: Hydrobiidae) in Tasmania'' (PhD). University of Tasmania. A large portion of the Beddomeia species, including ''B. fultoni'' fall into the category of narrow-range

endemic species Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...

. These narrow-range

taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

are generally poorly reserved due to limitations caused by small distributions, size, mobility, biogeographical boundaries and highly specific habitat requirements.Richards, K. (2013). Listing Statement for Beddomeia fultoni (Hydrobiid Snail (Farnhams Creek)). Retrieved 26 April 2021, from https://www.naturalvaluesatlas.tas.gov.au/downloadattachment?id=13550 Habitat disturbance from

agricultural Agriculture or farming is the practice of cultivating Plant, plants and livestock. Agriculture was the key development in the rise of Sedentism, sedentary human civilization, whereby farming of Domestication, domesticated species created food ...

and domestic land use,

climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...

and

invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...

are currently the principal threats contributing to the vulnerability of the species.Ponder, W., & Colgan, D. (2002). What makes a narrow-range taxon? Insights from Australian freshwater snails. ''Invertebrate Systematics'', ''16''(4), 571−582.

Taxonomy

Freshwater Mollusca can be divided into two classes, the

bivalves Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...

(mussels, clams and pea shells and the

gastropods The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...

(snails). These two classes differ significantly in physical appearance, biology and ecology but are grouped due to similarities found between body plans.Ponder, W.F., Walker, K. (2003). From Mound Springs to Mighty Rivers: The conservation Status of Freshwater Molluscs in Australia. ''Aquatic Ecosystem Health & Management'', ''6''(1), 19-28. The Hydrobiidae family, in which used to belong the ''Beddomeia'' genus and ''Beddomeia fultoni'' species, is the most diverse group of the taxonomic subclass

Caenogastropoda Caenogastropoda is a taxonomic clade, a large diverse group which are mostly sea snails and other marine gastropod mollusks, but also includes some freshwater snails and some land snails. The clade is the most diverse and ecologically successfu ...

and has over 267 known species. The ''Beddomeia'' complex can be separated into found distinct genera; These genera include ''

'', ''

Phrantela ''Phrantela'' is a genus of minute freshwater snails with an operculum, aquatic gastropod molluscs or micromolluscs in the family Hydrobiidae Hydrobiidae, commonly known as mud snails, is a large cosmopolitan family of very small freshwa ...

'', '' Nanocochlea'' and ''

Victodrobia ''Victodrobia'' is a genus of very small freshwater snails with an operculum, aquatic gastropod mollusks in the family Hydrobiidae. Species Species within the genus ''Victodrobia'' include: * ''Victodrobia burni'' * ''Victodrobia elongata'' * ...

.'' Of the 67 species that belong within the ''Beddomeia'' complex, 47 of these species are of the ''Beddomeia'' genus. This genus is endemic and restricted to Tasmania and occupies the northern areas of the state. Within this clade the

Planorbidae Planorbidae, common name the ramshorn snails or ram's horn snails, is a family of air-breathing freshwater snails, aquatic pulmonate gastropod molluscs. Unlike most molluscs, the blood of ram's horn snails contains iron-based hemoglobin ins ...

family is the next largest with about 40 species. Approximately half of the known hydrobiids are located in Tasmania, however, speciose faunas occur in south-eastern Australia and in the

Great Artesian Basin The Great Artesian Basin (GAB), located in Australia, is the largest and deepest artesian basin in the world, stretching over , with measured water temperatures ranging from . The basin provides the only source of fresh water through much of ...

. While some Hydrobiids have widespread and abundant populations, others, like ''B. fultoni'' are restricted to singular streams where genetic divergences occur within the same habitat.

Description

Hydrobiid snails are often around 1.0 -7.0 mm in length and often have smooth shells that can be opaque to a darker brown in colour. ''B. fultoni'' has a shell that is 3.07-3.78 mm in length and 2.24 – 2.78 mm in width and have a

protoconch A protoconch (meaning first or earliest or original shell) is an embryonic or larval shell which occurs in some classes of molluscs, e.g., the initial chamber of an ammonite or the larval shell of a gastropod. In older texts it is also called ...

of approximately 1.5 whorls. The umbilicus of the shell is not

sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

and is either very small or closed and is characterized by 0.16-0.39mm wide chink.Ponder, W.F, Clark, G.A, Miller, A.C & Toluzzi, A (1993). On a major radiation of fredhwater snails in Tasmania and eastern Victoria: a preliminary overview of the ''Beddomeia'' group (Mollusca: Gastropoda: Hydrobiidae). ''Invertebrate Taxonomy'' 7 :501-750 Due to their small size and cryptic nature, information surrounding the

ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...

and

biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary i ...

of hydrobiid snails is relatively limited. At a species level, the identification of hydrobiids and other closely related families can be challenging. Distinguishing factors include a number of defined anatomical and shell characteristics, however, in more recent times a move from morphological taxonomy has seen a rise in the use of allozyme

electrophoresis Electrophoresis, from Ancient Greek ἤλεκτρον (ḗlektron, "amber") and φόρησις (phórēsis, "the act of bearing"), is the motion of dispersed particles relative to a fluid under the influence of a spatially uniform electric fie ...

and DNA-based techniques for species identification. A study conducted in 2005 used a variety of Tasmanian species, including ''Pseudotricula, Nanocochlea,

Austropyrgus ''Austropyrgus'' is a genus of minute freshwater snails with an operculum, aquatic gastropod molluscs or micromolluscs in the Hydrobiidae family. ''Austropyrgus'' species are endemic to Australia, where they are found in virtually all freshwat ...

'' and ''Beddomeia'' species in order to phylogenetically analyse the subterranean genera and is currently the only reported sequencing of ''Beddomeia'' species. Due to high variability within each species of the ''Beddomeia'' family, certain morphological traits are referred to as ‘plastic’. These traits and characteristics are also convergent as multiple similarities can be seen between two or more species and therefore, cannot reliably be used as a direct indication of species. ''B. fultoni'' co-exist with some species of ''

'' which may cause confusion due to similar appearances. However, ''B. fultoni'' can usually be identified by the absence of operculum peg, a characteristic not appearing on any ''Beddomeia'' species.

Distribution and Habitat

Patterns of high diversity are generally presented by gastropods with increasing

species richness Species richness is the number of different species represented in an ecological community, landscape or region. Species richness is simply a count of species, and it does not take into account the abundances of the species or their relative a ...

and increasing

endemicity In epidemiology, an infection is said to be endemic in a specific population or populated place when that infection is constantly maintained at a baseline level without extra infections being brought into the group as a result of travel or simi ...

at lower latitudes. However, an exception of this is Tasmania, which holds the most diverse freshwater fauna in Australia. Through the use of anatomical traits to investigate speciation within a hydrobiid complex located in south-eastern Australia, 67 hydrobiid species were classified by Ponder ''et al.'' and the ''Beddomeia'' complex was categorised into four genera; three of which (''Beddomeia,

'' and '' Nanocochlea'') are endemic to Tasmania, with Victoria being the sole habitat of the fourth genus, ''

'' ''.'' There is a minimal geographical separation between these four genera particularly between the morphologically similar ''Austropyrgus and Nanocochlea'' and the supposedly closely related ''Beddomeia'' and ''Phrantela'' genera. The ''B.fultoni'' species can be described as cryptic in habitat due to the tendency to be located within environments within small and large streams of low hydrological variation. ''B. fultoni'' is geographically restricted to the Montagu catchment in north-western Tasmania, where it has been located within the headwater

tributaries A tributary, or affluent, is a stream or river that flows into a larger stream or main stem (or parent) river or a lake. A tributary does not flow directly into a sea or ocean. Tributaries and the main stem river drain the surrounding drainage b ...

of Fixters and Farnhams creeks. Within Fixters Creek the species is generally found on roots, wood, leaves and stones at Farnhams Creek. ''B.fultoni'' have only been identified within only 6 different streams with a maximum of 4km separating these sites, revealing the extremely narrow range of the species. This can be seen in Table 1, a population summary for ''Beddomeia fultoni'' assembled by K. Richards. Currently, the length of stream that the species occurs in unknown; however, inhospitable environments downstream of these catchments, caused by factors such as intensive dairy production, limit further radiation. Inhospitable environments such as wide streams, areas with high levels of

agricultural development Agricultural expansion describes the growth of agricultural land (arable land, pastures, etc.) especially in the 20th and 21st centuries. The agricultural expansion is often explained as a direct consequence of the global increase in food and ene ...

, production forestry, and domestic residences further separate subpopulations occurring within these catchments. Species such as ''B.fultoni'' which exhibit small ranges of distribution do so as a result of limitation by size, dispersal capabilities and mobility. Historical biogeographical obstructions often determine the small ranges of species, however for those with minimal dispersal capabilities, migration between suitable habitats is significantly hindered. The presence of substances such as root mates, various substrates, CWD, and detritus have been associated with the occurrence of ''B. fultoni'' within its known habitats.DPIWE, 2003. ''Aquatic Ecology of the Montagu River Catchment''. Hobary: The Department of Primary Industries, Water and Environment. * NRM region = Natural Resource Management region

Conservation and Threats

In 1995 ''Beddomeia fultoni'' was listed as rare on the Tasmanian ''

Threatened Species protection Act 1995 The ''Threatened Species Protection Act 1995'' (TSP Act), is an act of the Parliament of Tasmania that provides the statute relating to conservation of flora and fauna. Its long title is An Act to provide for the protection and management of th ...

''. In 2009 this status was elevated to endangered due to the species meeting the listing criterion for in two sections of criterion B. This includes criterion B1, as the species distribution is extremely fragmented and has been identified at no more than 5 locations and B2, as the species has a shown that a continuous population decline could be observed, inclined or projected due to occurrence and quality of habitat. Species are regarded as endangered when factors that have contributed to its decline continue to operate in the same manner and minimise the long-term chances of the species' survival prospects. Key factors that have been identified as threats to ''B.fultoni'' and other ''Beddomeia'' species include habitat modification and destruction,

interspecific competition Interspecific competition, in ecology, is a form of competition in which individuals of ''different'' species compete for the same resources in an ecosystem (e.g. food or living space). This can be contrasted with mutualism, a type of symbiosis. ...

from

introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived there ...

and stochastic effects.

Habitat Disturbance

Areas within North-Western Tasmania have been targeted for

and intensive vegetable production due to the deep ferrosols, including deep basalt soils, present. As a result, ''B.fultoni'' are highly vulnerable to anthropogenic habitat destruction and modification. Habitat suitability for ''B.fultoni'' is reduced by increases to stream temperatures as a result of the permanent removal of riparian vegetation. Retention of this riparian vegetation within hydrobiid habitats is a crucial factor in the survival of these populations and in terms of conservation, forestry, agricultural and mining activities have shown to have negatively impacted these areas. These damages are yet to be quantified however, it is predicted that a range from short to long term effects will be observed, with some being permanent. Currently the effectiveness of various riparian buggers are being investigated and reviewed in order to limit the impacts on water quality and ecology in areas that are subject to anthropogenic disruption. Without intervention, it is predicted that by the year 2100, human land-use alteration will have the greatest global impact on biodiversity, followed by more popularised events such as climate change and nitrogen deposition.

Climate Change

Alongside invasive species,

is one of the greatest biodiversity threats and is expected to provoke or contribute to the

extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

of many species such as ''B. fultoni'' in the future. As a result of increased

global temperatures Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of Meteorology, meteorological variables over a time spanning from months to millions of years. Some of the M ...

, predictions of increased rainfall, along with

severe storm A thunderstorm, also known as an electrical storm or a lightning storm, is a storm characterized by the presence of lightning and its acoustic effect on the Earth's atmosphere, known as thunder. Relatively weak thunderstorms are someti ...

and

flooding A flood is an overflow of water ( or rarely other fluids) that submerges land that is usually dry. In the sense of "flowing water", the word may also be applied to the inflow of the tide. Floods are an area of study of the discipline hydrolog ...

events, create major concern for freshwater populations. These events decrease habitat availability and often contribute to the modification of

riparian vegetation A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the terrestrial biomes of the Earth. Plant habitats and communities along the river margins and banks ar ...

communities. Other factors caused by climate change, such as increased water temperature, sea level, human activity and habitat availability, further disrupt the ability of species, such as ''B. fultoni,'' to survive.

Management

As of 2020 there are over 180 rare, vulnerable, or endangered animal species that have been identified under the ''Threatened Species Protection Act 1995.'' Over half of these listed species are invertebrates and 37 species are from the genus ''Beddomeia''. Management of ''B. fultoni'' alongside other members of the ''Beddomeia'' genus is predominantly governed by a series of parliamentary acts such as Tasmania’s ''Threatened Species Protection Act 1995; Nature conservation act 2000'' and '' Environment Protection and Biodiversity Conservation Act, 1999.'' The conservation measures are implemented through careful planning, restrictions and rule-sets such as the Forest Practices Code. However, currently species management is hampered by the lack of understanding and knowledge of the species’ habitat preferences and responses to

disturbance Disturbance and its variants may refer to: Math and science * Disturbance (ecology), a temporary change in average environmental conditions that causes a pronounced change in an ecosystem * Disturbance (geology), linear zone of faults and folds ...

. For ''B. fultoni'' management, the main objective is to decrease extinction risk through the maintenance of the integrity of habitat at known sites. In addition to this, identifying new subpopulations of the species and increasing ecological understanding are also critical factors in the long-term management of the species.

Behaviour and Ecology

Reproduction and lifestyle

The ''Beddomeia'' species have shown no signs of seasonal reproductive peaks and are able to breed throughout the entire year. The species reproduces sexually and lays single eggs which are contained within a secreted sand-grain capsule. The size of the egg capsules is approximately one third of the adult body size and individual capsules are equipped with broad attachment bases. This design allows for eggs to be attached to the underside of submerged rocks. Incubation periods are unknown; however, prior to emergence, eggs develop into completely formed juvenile snail. It is predicted that most ''Beddomeia'' species live to approximately 5 years; however, these species develop slowly and are only expected to reach

sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural definitio ...

after 2-3 years. Due to low egg capsule to snail abundance ratios being recorded, it is believed that the fecundity of ''B. fultoni'' is low. These methods of reproduction prevent ''Beddomeia'' species from dispersing into new habitat, unlike most other molluscs which have a free-swimming

larval A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The l ...

stage.

Predators and competition

Molluscs in general have been found to be a large contributor to the diet of

platypus The platypus (''Ornithorhynchus anatinus''), sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal Endemic (ecology), endemic to Eastern states of Australia, eastern Australia, including Tasmania. The platypu ...

; however, in Tasmania, stonefly nymphs, introduced

trout Trout are species of freshwater fish belonging to the genera '' Oncorhynchus'', ''Salmo'' and ''Salvelinus'', all of the subfamily Salmoninae of the family Salmonidae. The word ''trout'' is also used as part of the name of some non-salmoni ...

and platypus have been outlined as key

predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...

of hydrobiids. With exception to shell morphology, physiological predation responses of Hydrobiids are currently poorly known.

Native Native may refer to: People * Jus soli, citizenship by right of birth * Indigenous peoples, peoples with a set of specific rights based on their historical ties to a particular territory ** Native Americans (disambiguation) In arts and entert ...

snail populations and species like ''B. fultoni,'' particularly in lowland rural and urban streams, have also been negatively affected through the introduction of species such a '' Potamopyrgus antipodarium.''

Feeding and Digestion

In most occurrences, snails consume food via their mouths as they crawl over surfaces. Food is obtained in processes such as "area restricted searching" and "giving-up-time", in which the snails travel rapidly until they encounter a source of food and remain in this area until the resources run low have been.Dimock, R. and Dillon, R., 2000. The Ecology of Freshwater Molluscs. ''The Journal of Parasitology'', 86(5), p.1046. A method called " tropotaxis" allows the snails to orient in still waters by sensing and measuring the concentration of attractants in nearby locations. Upon arrival of these food sources, similar chemicals which stimulate tropotaxis are used to stimulate a feeding response. A muscular organ which contains the trophic apparatus, called the

proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...

, is the first part of a snail to make contact with the substrate. It is common to find sand grains in the

stomachs The stomach is a muscular, hollow organ in the gastrointestinal tract of humans and many other animals, including several invertebrates. The stomach has a dilated structure and functions as a vital organ in the digestive system. The stomach is ...

of freshwater snails such as ''Beddomeia fultoni.'' It is likely that this material is gathered and actively ingested in order to be used in the

trituration Trituration (Latin, ''grinding'') is the name of several different methods used to process materials. In one sense, it is a form of comminution (reducing the particle size of a substance). In another sense, it is the production of a homogeneous pow ...

of food.CARRIKER, M., 1946. OBSERVATIONS ON THE FUNCTIONING OF THE ALIMENTARY SYSTEM OF THE SNAIL LYMNAEA STAGNALIS APPRESSA SAY. ''The Biological Bulletin'', 91(1), pp.88-111.

References

External links

* * {{Taxonbar, from=Q307387 Gastropods of Australia Tateidae Beddomeia Vulnerable fauna of Australia Endemic fauna of Australia Gastropods described in 1993 Taxonomy articles created by Polbot