Physella acuta

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Physella acuta
Blasenschnecke 01.jpg
A live individual of Physella acuta
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Superorder: Hygrophila
Family: Physidae
Genus: Physella
Species:
P. acuta
Binomial name
Physella acuta
(Draparnaud, 1805) [2]
Synonyms [3]

Haitia acuta (Draparnaud, 1805)
Physa acuta Draparnaud, 1805
Physa cubensis L. Pfeiffer, 1839
Physa heterostropha (Say, 1817)
Physa integra Haldeman, 1841
Physa virgata A. Gould, 1855

Contents

Physella acuta, also known as European physa, tadpole snail, bladder snail, or acute bladder snail, is a species of small, air-breathing freshwater snail of the family Physidae. It originates from North America and was first described in 1805 by Jacques Philippe Raymond Draparnaud based on a specimen found in France. Its shell is sinistral (left-coiling), as is typical of physids. It reaches up to 12 millimetres (0.47 in) in length and forms a high spiral with a pointed apex. The mantle is blue to dark grey with golden spots and is often visible through the thin shell. Like other physids, P. acuta presents a unique set of muscles called the physid musculature that allows it to rapidly twist its shell as a defense mechanism.

P. acuta is invasive on all continents except Antarctica and is considered by Dillon and colleagues (2002) as "the world's most cosmopolitan freshwater gastropod". Its first introduction outside North America likely occurred through the 18th century cotton trade to Europe, while later spread mainly happened through the aquarium trade. The species can occupy various freshwater habitats, from lakes and ponds to sewage drains and irrigation systems, and tolerates polluted and oxygen-poor environments. As a simultaneous hermaphrodite (male and female reproductive organs are present and active at the same time), P. acuta can reproduce with other individuals and also self-fertilise. The species can coexist with other, often non-native snails such as Stenophysa marmorata and Radix auriculata. However, it frequently outcompetes native species due to its high reproductive rate and tolerance to habitat degradation. It is prey to a variety of animals such as water bugs, crayfish, and leeches, and protects itself by shaking its shell or escaping to or above the waterline. Despite its cosmopolitan distribution, prevalence of parasitic infections within invasive P. acuta populations is often low. Toledo and colleagues (1999) reported no successful infections by the human parasites Hypoderaeum conoideum and Trichobilharzia regenti , but one 2024 study detected Echinostoma (which causes echinostomiasis) in an individual from Rio de Janeiro, Brazil. P. acuta often arrives in aquariums through ornamental plants and can become a "pest snail" due to its rapid reproduction. However, a balanced population can help clean up organic leftovers in aquariums and avoid bacterial growth.

Taxonomy and nomenclature

Physella acuta was first described as Physa acuta in 1805 by Jacques Philippe Raymond Draparnaud, based on a specimen found in the Garonne catchment in France. [2] The specific epithet acuta is Latin for "sharp" and refers to the pointed shell. [4] In 1817 Thomas Say independently described the same species in Pennsylvania, naming it Physa heterostropha. [5] Until the beginning of the 21st century this and several other newly described species, such as Physella virgata and Physella integra (described in North America), as well as Physella cubensis (described in Central America), were considered distinct from each other until molecular and reproductive studies revealed them to be synonyms of P. acuta. [6] Since 2021 molecular phylogenetic studies have reported stronger evidence for moving the species from its original genus Physa to Physella. [7] [8] As of 2025 MolluscaBase, the mollusk-oriented branch of WoRMS (World Register of Marine Species), accepts the taxonomic classification proposed by Albrecht and colleagues (2025). [3] They place P. acuta in Physella propose relationships to other physids as shown in the cladogram below. [8]

Physidae

While the family Physidae is well established, the structure of its subfamily Physinae remains unclear as of 2025, leading to mixed acceptance of P. acuta's classification as Physella. [9] [7] Consequently, other authors and databases continue to use synonyms like Physa acuta or Haitia acuta. [10] [11] Debates include the taxonomic relationships between Physinae members as well as the definition and number of physinine genera and species, all of which also affect the identity of P. acuta. [7] This taxonomic uncertainty is partly due to vague descriptions in early works, which could apply to multiple taxa, [11] and to the generalist nature of physids, including P. acuta, which lead to morphological plasticity and rapid evolution. [7]

Description

External anatomy

Shell

Shell of Physella acuta Physella acuta 01.JPG
Shell of Physella acuta

Like other members of the family Physidae, the shell of P. acuta lacks an operculum, a "trapdoor" that closes the aperture (shell opening), and is sinistral. Sinistral shells are left-coiling, meaning that when held with the aperture facing the observer and the spire (coiled part of the shell) pointing upward, the aperture is on the left-hand side. [12] The shell is thin and small, reaching up to 12 millimetres (0.47 in) length. [13] It forms a high spiral of four to five whorls (complete revolutions) which take about two-thirds of the shell length and end in a pointed apex (tip). Sutures (grooves between the whorls) are impressed and clearly visible. The aperture is ear-shaped and takes about three-quarters of the total shell height. Both the columella (central pillar within the shell) and the apertural lip (the opening's margin) are white. The shell surface is smooth, thin to the point of being slightly transparent, and ranges in colour from pale horn to brownish yellow. [14] [15]

According to the U.S. Geological Survey (USGS), the species presents a high diversity of shell shapes which led to numerous false species descriptions before the onset of molecular phylogenetic studies. [16] Shells of P. acuta can be especially difficult to distinguish from those of Physella gyrina and Stenophysa marmorata if live specimens are not available, since body morphology provides key distinguishing features. Typically, P. gyrina has a shorter spire with shallower sutures and a larger shell which can exceed 13 millimetres (0.51 in) length. [14] [17] The shell of S. marmorata is thinner and less globous. [18]

Soft parts

Physella acuta on an aquarium glass. Blasenschnecke 03.jpg
Physella acuta on an aquarium glass.

The body of P. acuta is very variable in colour, which can range from blue to dark grey. The top mantle (tissue covering the visceral mass of the animal) has golden spots often visible under the thin shell. [16] Finger-like lobes extend from the mantle on both sides of the body, smaller than in other physids such as Physa fontinalis or Stenophysa marmorata. They can act as an accessory gill by extracting additional oxygen from the environment and help detect predators through the touch-sensitive tissue. [19] [20] The tentacles are grey and follow the build of other members within the family Physidae: cylindrical and slender, almost transparent, with small black eyes at the base. [14] These pit eyes only distinguish between light and dark. [21] The foot is narrow and ends in a pointed tail, as is also typical of the family. The mouth edge is large and flared. The body is an important point of distinction from P. gyrina and S. marmorata:P. gyrina has whitish spots over its whole dark grey body including the tentacles. [14] The mantle extensions of S. marmorata are not digitated and extend broadly over the sides of the shell and this species has a black stripe running through the middle of the tail. [18]

Shell-shaking by Physella acuta to ward off a mating attempt
Body and penial complex of Physella acuta.png
Physella acuta's body (b) and male reproductive organs (c) including the penis (d). p = penis, pg = preputial gland, pp = prepuce, ps = penis sheath

Internal anatomy

Like all members of the family Physidae, P. acuta has a pulmonary cavity within the mantle which enables it to take oxygen from the water or from the air. [22] It also has a specialised set of muscles called the physid musculature. These muscles are unique among gastropods and allow the snail to rapidly twist its shell in clockwise rotations of up to 120°. This shell-shaking movement is an important defense against predators. They provide the twisting force, serve as a base for the movement, anchor the shell to the body, and help return the shell to its original position afterwards. [23]

P. acuta is a simultaneous hermaphrodite, meaning it has both male and female reproductive organs functioning at the same time. In Physidae, the male organs are important for taxonomic identification and consist of prepuce, penis sheath, and penis. [24] The prepuce is a tube that releases or receives sperm from outside and, in the case of P. acuta, includes a small, lentil-shaped gland (the preputial gland). It is wider and twice as long as the muscular penis sheath. [25] [26] The penis is white and 160 - 180 μm in length. [27] It is elongated and narrow along most of its length but ends in a wider, rounded tip. [28] The penis musculature consists of circular muscles in both the outer and inner layers, while the intermediate layer has only longitudinal muscles. Together with molecular phylogenetic data, this musculature pattern can be used to distinguish major groups within the superorder Hygrophila. [27] The female organs are less frequently described in detail. They consist of a convoluted oviduct (which transports the eggs to the outside), a nidamental gland (which secretes the egg capsule), and a vagina connected to the spermathecal duct (which receives sperm) opening to the outside. [25]

Distribution

Physella acuta found near Akashi, Japan, far outside the species' native range Acute Bladder Snail (Physella acuta) in Japan.jpg
Physella acuta found near Akashi, Japan, far outside the species’ native range

P. acuta is widely dispersed across the globe, largely due to the aquarium trade. [29] It is an invasive species which can be found on all continents except Antarctica and is considered by Dillon and colleagues (2002) as "the world's most cosmopolitan freshwater gastropod". [30]

P. acuta was originally thought to be native to the Mediterranean region. However, reproductive isolation experiments [30] and molecular genetic studies [31] revealed it to be the same species as the North American Physella heterostropha and Physa integra. Comparative anatomy, fossil evidence, and phylogenetic data also support a North American origin, specifically within the United States. However, its exact native range within the country remains debated, with hypotheses for both eastern and western origins. [32] [6] The timing and pathway of P. acuta's first arrival in Europe are also uncertain. One hypothesis links it back to eastern U.S. populations via the 18th century cotton trade, [6] although earlier or natural introductions have also been proposed. [11] Once within Europe, its initial spread was likely facilitated by man-made canals. [11] [33]

Ecology and behaviour

Physella acuta can survive in eutrophic habitats such as lakes with excess algal growth. A eutrophic lake, with excess algal growth.jpg
Physella acuta can survive in eutrophic habitats such as lakes with excess algal growth.

P. acuta can occupy a variety of freshwater habitats as well as habitats varying widely in water availability. [16] As a pulmonate snail, it tolerates harsh environmental conditions such as polluted and eutrophic (oxygen-deprived) waters, since it is able to come to the surface to breathe air. [34] [15] P. acuta has been reported in lakes, reservoirs, ponds, streams, ditches, as well as artificial sites such as sewage drains and irrigation systems. [35] [36] It is a scraper feeder and uses its radula (a tongue-like structure covered in small chitinous teeth) to scrape green algae, diatoms, and aquatic plants from the surface. [37]

P. acuta is a simultaneous hermaphrodite which is self-compatible, meaning it is capable of both outcrossing (reproduction with another individual) and self-fertilisation. [38] In natural populations, reproduction occurs mainly by outcrossing, but self-fertilisation rates still remain between 10 - 30% and can increase as an adaptation strategy when mates are scarce. [39] [40] Adults lay 50 - 100 eggs per week for up to a year after reaching sexual maturity. [16] Eggs are deposited in elongate gelatinous sacs and hatch after 15 – 20 days. Individuals reach sexual maturity after 17 – 18 months. [41]

Ecological interactions

Coexistence and competition

P. acuta can coexist with other non-native snails such as Stenophysa marmorata, [42] Potamopyrgus antipodarum, [43] Lithoglyphus naticoides , and Radix auricularia . [44] In these cases, competition may not be strong enough to cause exclusion, and species can differ in their competition strategies or life-history traits. [42] More often, however, its presence leads to the decline of native gastropod populations. [45] [6] It outcompetes Glyptophysa gibbosa in Australia, [46] Physafontinalis in Italy, [47] as well as Racesina luteola and Filopaludina bengalensis in India. [48] [49] In Mozambique, it displaced Bulinus forskalii to become the dominant gastropod. [50] Like many invasive freshwater snails, the competitive success of P. acuta can be explained by its higher fecundity, shorter egg development time, and broader tolerance to habitat degradation. [47] It also shows a stronger growth response under rising temperatures than some native species, a trait that can favour it under climate change. [45] These characteristics make P. acuta more efficient in competing for food when diets overlap. [50] The presence of P. acuta can also inhibit the growth of other species, while its own growth is stimulated by them, although exact mechanisms are unknown. [45]

Anentome helena is often kept in aquariums to control populations of smaller snail species such as Physella acuta. Clea helena.jpg
Anentome helena is often kept in aquariums to control populations of smaller snail species such as Physella acuta.

Predation

P. acuta is predated by a variety of animals, including water bugs, [51] marsh fly larvae, [52] crayfish, [53] leeches, [20] and various fish species. [54] [55] To a lesser degree, it is also prey to the invasive carnivorous snail Anentome helena. [49] Anti-predator behaviours include shell-shaking (rapid shell movements) and crawling to or above the waterline, [20] [21] as well as burying into the sediment, [55] leaping (shell-shaking combined with detachment from the surface, causing the snail to jerk away), [20] [56] clamping to the substrate, and detaching to float up to the surface. [57]

Parasitism

P. acuta is a potential intermediate host for parasites of both native wildlife and humans. [6] Due to its cosmopolitan distribution and invasive capacity, it has the potential to significantly influence the distribution of parasites within freshwater ecosystems. [58] Reported infections include the turtle parasites Krefftascaris spp. and spirorchiids, [58] [59] the bird parasite Cotylurus cornutus [59] , and the rat parasite Euparyphium albuferensis, [60] although infection prevalence within invasive P. acuta populations is often low. [6] Experiments exposing P. acuta to the human parasites Hypoderaeum conoideum and Trichobilharzia regenti yielded no successful infections. [61] This is consistent with the enemy-release hypothesis, the observation that non-native species carry fewer parasites outside their native range. [62] [8] The species was directly linked to a human illness once in 2024, when Moreira and colleagues recorded Echinostoma (a parasitic flatworm which causes echinostomiasis) in P. acuta individuals from public parks in Rio de Janeiro, Brazil. [63]

Aquarium trade

Physella acuta is often called a "pest snail" in freshwater fishkeeping. It is usually introduced with ornamental plants or decoration. A single introduced individual can be enough to establish a population due to the snail's capacity to self-fertilise and to reproduce quickly. An excessive reproduction of P. acuta may result from an oversupply of food in the aquarium, for example when fish and shrimp are overfed or offered poorly digestible food (undigested parts are excreted and become available to the snails). Infrequent water changes can also lead to the accumulation of plant debris, which is eaten by the snails. However, a balanced population of P. acuta in the aquarium can help clean up organic leftovers and prevent bacterial growth. [64] They are peaceful snails which will not attack healthy plants and can feed other tank inhabitants like pufferfish, predatory snails, and crayfish. [65]

References

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