Beddomeia fultoni

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Beddomeia fultoni
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Caenogastropoda
Order: Littorinimorpha
Family: Tateidae
Genus: Beddomeia
Species:
B. fultoni
Binomial name
Beddomeia fultoni
Ponder & Clark, 1993
Scientific sketch of Beddomeia fultoni found in Tasmania Beddomeia fultoni sketch.png
Scientific sketch of Beddomeia fultoni found in Tasmania

Beddomeia fultoni(B. fultoni) is a species of small freshwater snail belonging to the family Tateidae. [2]

Contents

The species is endemic to Australia. Commonly known as Farnhams Creek hydrobiid snail, the species used to belong to the family Hydrobiidae sensu lato (s.l.)  and is listed as one of 37 threatened Beddomeia species on the Tasmanian Threatened Species Protection Act 1995 . [3]

Found within north-western Tasmania, 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 4 km. [4] A large portion of the Beddomeia species, including B. fultoni fall into the category of narrow-range endemic species. These narrow-range taxa are generally poorly reserved due to limitations caused by small distributions, size, mobility, biogeographical boundaries and highly specific habitat requirements. [5] Habitat disturbance from agricultural and domestic land use, climate change and invasive species are currently the principal threats contributing to the vulnerability of the species. [6]

Taxonomy

Freshwater Mollusca can be divided into two classes, the bivalves (mussels, clams and pea shells and the gastropods (snails). These two classes differ significantly in physical appearance, biology and ecology but are grouped due to similarities found between body plans. [7]

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 and has over 267 known species. [8] [7] The Beddomeia complex can be separated into found distinct genera; These genera include Beddomeia , Phrantela , Nanocochlea and Victodrobia. [4] 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. [7]

Within this clade the family Planorbidae is the next largest with about 40 species. [9] Approximately half of the known hydrobiids are located in Tasmania, however, speciose faunas occur in south-eastern Australia and in the Great Artesian Basin. 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. [7]

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 of approximately 1.5 whorls. The umbilicus of the shell is not sexually dimorphic and is either very small or closed and is characterized by 0.16-0.39mm wide chink. [2]

Due to their small size and cryptic nature, information surrounding the ecology and biology of hydrobiid snails is relatively limited. [4] 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 and DNA-based techniques for species identification. [4] A study conducted in 2005 used a variety of Tasmanian species, including Pseudotricula, Nanocochlea, Austropyrgus and Beddomeia species in order to phylogenetically analyse the subterranean genera and is currently the only reported sequencing of Beddomeia species. [10]  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. [11]

B. fultoni co-exist with some species of Austropyrgus 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. [5]

Distribution and habitat

Patterns of high diversity are generally presented by gastropods with increasing species richness and increasing endemicity at lower latitudes. However, an exception of this is Tasmania, which holds the most diverse freshwater fauna in Australia. [12] 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. [2] and the Beddomeia complex was categorised into four genera; three of which (Beddomeia, Phrantela and Nanocochlea ) are endemic to Tasmania, with Victoria being the sole habitat of the fourth genus, Victodrobia [5] . 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. [4] 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. [13]

B. fultoni is geographically restricted to the Montagu catchment in north-western Tasmania, where it has been located within the headwater tributaries of Fixters and Farnhams creeks. Within Fixters Creek the species is generally found on roots, wood, leaves and stones at Farnhams Creek. [4] B.fultoni have only been identified within only 6 different streams with a maximum of 4 km 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. [5] 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, production forestry, and domestic residences further separate subpopulations occurring within these catchments. [5]

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. [4]  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. [13]

Table 1. Population summary for Beddomeia fultoni [5]
LocationTenureNRM region *1:25 000 MapsheetYear Last (first) SeenExtent of Subpopulation (ha)Abundance
1Farnhams Creek, on the Bass

Highway, west of Christmas

Hills

Private PropertyCradle CoastTogari1989unknownLow
2Tributary of Fixters Creek,

north end of Brittons Swamp

State ForestCradle CoastTogari1989unknownLow
3Tributary of Fixters Creek,

southern end of Brittons

Swamp

State ForestCradle CoastTogari2006unknownLow
4Tributary of Fixters Creek,

central Brittons Swamp

(informal reserve)

State ForestCradle CoastTogari2010unknownModerate
5Tributary of Fixters Creek,

central Brittons Swamp

(informal reserve)

State ForestCradle CoastTogari2010unknownHigh
6Tributary of Farnhams Creek,

off Riseborough Road, Togari

State Forest

State ForestCradle CoastMella2010unknownLow

* 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 . [14]   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. [5] 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. [13]

Key factors that have been identified as threats to B.fultoni and other Beddomeia species include habitat modification and destruction, interspecific competition from introduced species, climate change and stochastic effects. [6]  

Habitat disturbance

Areas within North-Western Tasmania have been targeted for agricultural development 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. [5] Habitat suitability for B.fultoni is reduced by increases to stream temperatures as a result of the permanent removal of riparian vegetation. [15]  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. [16] 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. [2] 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. [17] 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. [18]

Climate change

Alongside invasive species, climate change is one of the greatest biodiversity threats and is expected to provoke or contribute to the extinction of many species such as B. fultoni in the future. [19] As a result of increased global temperatures, predictions of increased rainfall, along with severe storm and flooding events, create major concern for freshwater populations. These events decrease habitat availability and often contribute to the modification of riparian vegetation communities. [4] 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. [20]

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 [21] 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. [22] However, currently species management is hampered by the lack of understanding and knowledge of the species' habitat preferences and responses to disturbance. [7] 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. [4]

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. [5] 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. [4] 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. [23] 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 after 2–3 years. [4]  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 stage. [24]  

Predators and competition

Molluscs in general have been found to be a large contributor to the diet of platypus; however, in Tasmania, stonefly nymphs, introduced trout and platypus have been outlined as key predators of hydrobiids. [25] With exception to shell morphology, physiological predation responses of Hydrobiids are currently poorly known. [4] Native 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. [13]

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. [26] A method called "tropotaxis" allows the snails to orient in still waters by sensing and measuring the concentration of attractants in nearby locations. [27] Upon arrival of these food sources, similar chemicals which stimulate tropotaxis are used to stimulate a feeding response. [28] A muscular organ which contains the trophic apparatus, called the proboscis, is the first part of a snail to make contact with the substrate. [26] It is common to find sand grains in the stomachs 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 of food. [29]

See also

Related Research Articles

Beddomeia is a genus of very small freshwater snails that have a gill and an operculum, aquatic operculate gastropod mollusks in the family Tateidae.

Beddomeia averni is a species of very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk in the family Hydrobiidae. This species is endemic to Australia. The Hydrobiidae snail has been categorized into four distinct groups: Beddomeia being one of them. Very little is known about their ecology and habitat. Low funding in Australia for the conservation of threatened species is partly responsible for this; moreover, there are only a few good sources that contribute to its limited data required to have a better understanding. Studies have shown that it is hard to distinguish the species from a morphology point of view. A non-taxonomist may find it arduous to tell differences among different members of the species. Therefore the anatomical study is incorporated to understand and differentiate them. It is found that human-led actions such as agriculture, forestry, mining, etc have caused major degradation in their population. Other factors such as changes in water conditions due to sedimentation, varying temperature, mineral level, salinity can either positively/negatively impact their population.

Beddomeia camensis is a species of very small freshwater snail that is endemic to Australia. The invertebrate is 2-4 mm in size, it is a gastropod mollusk and belongs to the Hydrobiidae family, a large family of freshwater snails recognised by their small size and cosmopolitan distribution. Beddomeia camensis is one of the least populated species within its family distributed across only 5 small to medium-sized streams that flow into the Cam River catchment in North-West Tasmania.

Beddomeia capensis is an aquatic operculate gastropod mollusk, a species of very small freshwater snail that has a gill and an operculum, in the family Hydrobiidae. The species is classified as endangered and known to be endemic to five streams within a relatively small area on the island state of Tasmania, Australia. However, recent surveys have only been able to locate Beddomeia capensis in two small streams on Table Cape, with the total length of inhabited stream less than 80 metres (260 ft).

<i>Beddomeia hallae</i> Species of gastropod

Beddomeia hallae, also known as Buttons Rivulet hydrobiid snail, is a species of small freshwater snail that is endemic to Australia. The species is an aquatic operculate gastropod mollusk in the family Hydrobiidae. Beddomeia hallae belongs to the genus Beddomeia, which is the largest group in the family Hydrobiidae, consisting of 47 species. In the Threatened Species Protection Act 1995, this species is one of the 37 Beddomeia species listed as endangered, however, on the International Union for Conservation of Nature Red List, the species is listed as vulnerable. Found in Tasmania, in the streams of Buttons Rivulet and Castra Rivulet, Beddomeia hallae is sighted in its natural habitat amongst wood, leaves and under stones. Nonetheless, the Beddomeia species including Beddomeia hallae are geographically isolated, existing within restricted ranges.

Beddomeia krybetes is a species of very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk in the family Tateidae. It was first described in 1993 by Winston Ponder, G.A. Clark, Alison Miller and A Toluzzi.

Beddomeia launcestonensis is a species of very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk in the family Hydrobiidae. This species is endemic to Australia.

Beddomeia mesibovi is a species of very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk in the family Tateidae. It was first described in 1993.

Beddomeia minima is a population of freshwater snails that are endemic to Australia. It is commonly classified as a hydrobiid snail. This population was listed as Vulnerable on the IUCN Red List in 2011 due to its distribution being restricted to a single location and range of possible threats present in Tasmania. Beddomeia minima is one of a few fresh water snail species belonging to the same genus Beddomeia that survive in Tasmania, Australia, specifically a single location in the Scottsdale area.

Beddomeia petterdi is a species of snail in the family Tateidae. It is a very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk. It was first described in 1993.

Beddomeia phasianella is a species of snail in the family Tateidae. It is a very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk. This species is endemic to Tasmania.

Beddomeia salmonis is a species of very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk. This species is endemic to Australia, and was first described in 1993 by Winston Ponder and G.A. Clark.

Beddomeia topsiae is a species of snail in the family Tateidae first described in 1993.

Beddomeia waterhouseae, also known as Claytons Rivulet freshwater snail, is a species of freshwater snail in the family Tateidae. This species is endemic to northern Tasmania in Australia. The holotype specimen was found in a very small tributary of Little Clayton's Rivulet and is held at the Australian Museum. B. waterhouseae is small and as an adult has a shell measuring between 1.7 to 3.7 mm in length. The shell shape is ovate-conic to broadly conic and has a thin inner lip and no columellar bulge. This species feeds on algae and detritus on rocks. The female of the species lay single eggs in capsules made of sand grains and attached to the underside of rocks or wood. B. waterhouseae is considered vulnerable by the IUCN as it has a very small range and is sensitive to water quality and so may be threatened by disturbances of its habitat. Other threats include habitat loss. Conservation activities such as assessment of the aquatic ecosystem and vegetal surveys are being undertaken in an attempt to preserve this species.

Beddomeia zeehanensis is a species of very small freshwater snail that has a gill and an operculum, an aquatic operculate gastropod mollusk in the family Hydrobiidae. The species was first described in 1993 by Winston Ponder and G.A. Clark.

<i>Nanocochlea</i> Genus of gastropods

Nanocochlea is a genus of minute freshwater snails with an operculum, aquatic gastropod mollusks in the family Tateidae. The genus was first described in 1993 by Winston Ponder and others. The type species is Nanocochlea monticola.

Austropyrgus niger is a species of minute freshwater snail with an operculum, an aquatic gastropod mollusc or micromollusc in the family Hydrobiidae. This species is endemic to southeastern Tasmania, Australia. It is found in small coastal streams along the D'Entrecasteaux Channel.

Austropyrgus rectoides is a species of minute freshwater snail with an operculum, an aquatic gastropod mollusc or micromollusc in the Hydrobiidae family. This species is endemic to Tasmania, Australia. It is found in small coastal rivers and streams in northern and western Tasmania and also on Hunter Island.

Austropyrgus ronkershawi is a species of minute freshwater snail with an operculum, an aquatic gastropod mollusc or micromollusc in the Hydrobiidae family. This species is endemic to Tasmania, Australia. It is known from a number of small streams and rivers in central northern Tasmania.

Westrapyrgus is a genus of minute freshwater snails with an operculum, aquatic gastropod molluscs or micromolluscs in the family Hydrobiidae. The genus is endemic to south Western Australia. It is the first hydrobiid genus to be described from this state.

References

  1. ↑ Ponder, W.F. (1996). "Beddomeia fultoni". IUCN Red List of Threatened Species . 1996: e.T2692A9468393. doi: 10.2305/IUCN.UK.1996.RLTS.T2692A9468393.en . Retrieved 16 November 2021.
  2. 1 2 3 4 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
  3. ↑ Threatened Species Section (2017). Beddomeia fultoni (Hydrobiid Snail (Farnhams Creek)): Species Management Profile for Tasmania's Threatened Species Link. https://www.threatenedspecieslink.tas.gov.au/Pages/Hydrobiid-Snail-(Farnhams-Creek).aspx. Department of Primary Industries, Parks, Water and Environment, Tasmania. Accessed on 21/3/2021.
  4. 1 2 3 4 5 6 7 8 9 10 11 12 Richards, K. (2010) An ecological, morphological and molecular investigation of Beddomeia species (Gastropoda: Hydrobiidae) in Tasmania (PhD). University of Tasmania.
  5. 1 2 3 4 5 6 7 8 9 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
  6. 1 2 Ponder, W., & Colgan, D. (2002). What makes a narrow-range taxon? Insights from Australian freshwater snails. Invertebrate Systematics, 16(4), 571−582.
  7. 1 2 3 4 5 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.
  8. ↑ Smith, B.  J. (1992)  Non-Marine Mollusca.  In: W.  W. K. Houston (Ed.), Zoological Catalogue of Australia. Australian Government Publishing Service, Canberra
  9. ↑ Ponder, W.F. (1997). Conservation status, threats and habitat requirements of Australian terrestrial and freshwater molluscs. Memoirs Of The Museum Of Victoria, 56(2), 421-430.
  10. ↑ Perez, K., Ponder, W., Colgan, D., Clark, S., & Lydeard, C. (2005). Molecular phylogeny and biogeography of spring-associated hydrobiid snails of the Great Artesian Basin, Australia. Molecular Phylogenetics And Evolution, 34(3), 545-556.
  11. ↑ Hershler, R. & Ponder, W.F. (1998). A review of morphological characters of Hydrobioid snails. Smithsonian Contributions to Zoology, Number 600, 55.
  12. ↑ Strong, E., Gargominy, O., Ponder, W., & Bouchet, P. (2007). Global diversity of gastropods (Gastropoda; Mollusca) in freshwater. Hydrobiologia, 595(1), 149-166.
  13. 1 2 3 4 DPIWE, 2003. Aquatic Ecology of the Montagu River Catchment. Hobary: The Department of Primary Industries, Water and Environment.
  14. ↑ Tasmanian Legislation Online. (2020). Retrieved 26 April 2021, from https://www.legislation.tas.gov.au/view/html/inforce/current/act-1995-083
  15. ↑ Mitchell, S. (1999). A Simple Model for Estimating Mean Monthly Stream Temperatures After Riparian Canopy Removal. Environmental Management, 24(1), 77-83.
  16. ↑ Cornish, P. (2001). The effects of roading, harvesting and forest regeneration on streamwater turbidity levels in a moist eucalypt forest. Forest Ecology And Management, 152(1-3), 293-312.
  17. ↑ Borg, H., Hordacre, A., & Batini, F. (1988). Effects of logging in stream and river buffers on watercourses and water quality in the southern forest of Western Australia. Australian Forestry, 51(2), 98-105.
  18. ↑ Sala, O., (2000). Global Biodiversity Scenarios for the Year 2100 . Science, 287(5459), pp.1770-1774.
  19. ↑ Gallardo, B., & Aldridge, D. (2013). Evaluating the combined threat of climate change and biological invasions on endangered species. Biological Conservation, 160, 225-233.
  20. ↑ Hastie, L., Cosgrove, P., Ellis, N., & Gaywood, M. (2003). The Threat of Climate Change to Freshwater Pearl Mussel Populations. AMBIO: A Journal Of The Human Environment, 32(1), 40-46.
  21. ↑ State of Queensland. NATURE CONSERVATIONAND OTHER LEGISLATIONAMENDMENT ACT 2000 (2000). Queensland.
  22. ↑ Forest Practices Board (2000). Forest Practices Code. Forest Practices Board, Hobart, Tasmania
  23. ↑ FPA Biodiversity research and monitoring. Fpa.tas.gov.au. (2015). Retrieved 16 May 2021, from https://www.fpa.tas.gov.au/research_and_monitoring/biodiversity_program_research_and_monitoring.
  24. ↑ Bryant, S.  L.  and Jackson,  J.  (1999). Tasmania’s  Threatened  Fauna Handbook:  what,  where and  how  toprotect Tasmania’s threatened animals. Threatened Species Unit, Parks and Wildlife Service, Hobart.
  25. ↑ Faragher, R., Grant, T., & Carrick, F. (1979). Food of the platypus (Ornithorhynchus anatinus) with notes on the food of brown trout (Salmo trutta) in the Shoalhaven River, N.S.W. Austral Ecology, 4(2), 171-179.
  26. 1 2 Dimock, R. and Dillon, R., 2000. The Ecology of Freshwater Molluscs. The Journal of Parasitology, 86(5), p.1046.
  27. ↑ Townsend, C., 1973. The role of the osphradium in chemoreception by the snail Biomphalaria glabrata (Say). Animal Behaviour, 21(3), pp.549-556.
  28. ↑ DALDORPH, P. and THOMAS, J., 1988. The chemical ecology of some British freshwater gastropod molluses: behavioural responses to short chain carboxylic acids and maltose. Freshwater Biology, 19(2), pp.167-178.
  29. ↑ 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.
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