Hristov, K.K. AvailableInd. J. Pure online App. Biosci. at www.ijpab.com (2020) 8(5), 1-10 ISSN: 2582 – 2845

DOI: http://dx.doi.org/10.18782/2582-2845.8319 ISSN: 2582 – 2845 Ind. J. Pure App. Biosci. (2020) 8(5), 1-10

Research Article

Peer-Reviewed, Refereed, Open Access Journal

Observations on turrita (Gmelin 1791) Breeding Behaviour in Laboratory Conditions

Kroum K. Hristov* Department of Chemistry and Biochemistry, Medical University - Sofia, Sofia - 1431, Bulgaria *Corresponding Author E-mail: [email protected] Received: 15.08.2020 | Revised: 22.09.2020 | Accepted: 24.09.2020

ABSTRACT Neritina turrita (Gmelin 1791) along with other Neritina, Clithon, Septaria, and other fresh- water are popular in ornamental aquarium trade. The need for laboratory-bred animals, eliminating the potential biohazard risks, for the ornamental aquarium trade and the growing demand for model systems for biomedical research reasons the work for optimising a successful breading protocol. The initial results demonstrate N. turrita as tough animals, surviving fluctuations in pH from 5 to 9, and shifts from a fresh-water environment to brackish (2 - 20 ppt), to sea-water (35 ppt) salinities. The females laid over 630 (at salinities 0, 2, 10 ppt and temperatures of 25 - 28oC) white oval 1 by 0.5 mm egg capsules continuously within 2 months after collecting semen from several males. Depositions of egg capsules are set apart 6 +/-3 days, and consist on average of 53 (range 3 to 192) egg capsules. Production of viable veligers was recorded under laboratory conditions.

Keywords: Neritina turrita, Sea-water, Temperatures, Environment

INTRODUCTION supposably different genera forming hybrids Neritininae are found in the coastal swamps of with each other, suggesting their close relation. most tropical regions. The existing of Describing them all as different species is Neritina were divided into 4 subgenera in the likely invalid, let alone allocating them to 20’s by Baker based on differences in shell different subgenera or, even worse, to shape, and in shape of the teeth of the radula. independent genera (Bandel, 2001). Later, Thiele suggests 5 subgenera, and in the Neritina turrita (Gmelin, 1791) snails 80’s Cunningham Vaught - 11 subgenera. have shells that are oblong-conical, lightly Nowadays, Haynes describes the subgeneric striated, shiny, spire elevated, pointed, yellow system as confusing, not reflecting the true with oblique, curved or rippled black stripes. relations among the existing species. Studying The shells are 25-32 mm in length. The the richest neritinid fauna around the Fiji aperture is white and the columellar area is Islands, she describes species belonging to yellow tinted.

Cite this article: Hristov, K.K. (2020). Observations on Neritina turrita (Gmelin 1791) Breeding Behaviour in Laboratory Conditions, Ind. J. Pure App. Biosci. 8(5), 1-10. doi:

http://dx.doi.org/10.18782/2582-2845.8319

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Hristov, K.K. Ind. J. Pure App. Biosci. (2020) 8(5), 1-10 ISSN: 2582 – 2845 It is found throughout the Indo-Pacific region: dense aggregations. Such migrations have including Madagascar, Japan to Indonesia, and been reported in Hawaii (Hau, 2007), Costa the western Pacific Ocean islands. It is not Rica, Japan (Kano, 2009), French Polynesia found in India and Australia. In the Mariana (Liu & Resh, 1997), and (Pyron & Islands, mainly Guam (Micronesia), N. turrita Covich, 2003; Gorbach et al., 2012). The is present, along with a total of 25 other controlling factors for such migrations are species of gastropods known to inhabit fresh unknown, although seasonal factors (like rain and brackish waters, including 18 native and flash-floods), and increased predation in species: 13 , 4 Thiaridae, and 1 the estuary (by fish and birds) have been Lymnaeidae; and 7 introduced species - 2 suggested (Schneider & Lyons, 1993). Ampullariidae, 2 Planorbidae, as well as 1 Neritina, and the related genera each from the Physidae, Thiaridae and Clithon and Septaria usually settle on tropical Viviparidae (Kerr, 2013). coasts, arriving after a planktonic larval stage Neritina turrita inhabits tidally- at sea. Veligers metamorphose in the estuaries, influenced brackish waters near the mouths of where the benthic young are exposed to the rivers and streams. It is often found near the influence of fresh-water. From the estuary, shore in mud, or on stones. Members of the many species crawl upstream and spend the gastropod families Neritidae and Thiaridae are rest of their adult lives in the fresh-water good travellers, having larvae that disperse on environment (Bandel, 2001). Female Neritidae ocean currents, so that they have a broad store sperm capsules (spermatophores) geographical distributions, and the adults received from the males, which permits move several kilometres up-stream estuaries - continuous spawning. Neritids lay flat, a habitat shift with growth. Migrating adults elliptical egg capsules on hard substrata, have higher densities in riffles than in pool including rocks and shells of other snails. habitats. Neritids prefer large substrates (Liu Newly laid capsules are white, becoming & Resh, 1997). In Moorea (French Polynesia) cream to brown in a few days. Each capsule and Mariana Islands - Guam, Palau and Yap contains up to 300 eggs that hatch several (Micronesia) they are found along with other weeks later. Morphological studies of larvae species introduced by man - either as potential and their shells are published since the 70’s by food, or due to accidental release from Robertson, Scheltema, Lauren, and by Bandel aquaculture or the aquarium trade. On at the present time. There is a limited time- Iriomote Island in the Ryukyu Islands, window for neritid larvae to reach salt-water: southern Japan, N. turrita is found together veligers held in fresh-water die within 6 days. with N. violacea and achieves high densities In Hawaii, larvae of are and is widely distributed in the mangrove found year-round in drift collections, in pools swamps and the adjacent seashore (Okuda & and run-off areas during low-flow conditions. Nishihira, 2002). The molluscan community is Floods cause a decrease in water temperature a key component of the mangrove food chain, and an increase in dissolved oxygen, providing preyed by many fish and waterbird species. optimum conditions for egg capsules to be Anthropogenic disturbances resulting in laid, and hatched larvae to be transported to habitat degradation impact the populations of the ocean. Genetic studies (Hodges & such amphidromous gastropods. Allendorf, 1998; Myers et al., 2000; Crandall Neritina turrita snails are herbivorous et al., 2010) confirm that larvae from in the adult stage, developing from planktonic amphidromous lineages are capable of pelagic veliger larvae. Its growth is faster in the dispersal across the open ocean that lasts from warmer season (April-October), and slower in weeks to months. The growth rate is higher the cooler season (Takada, 2016). Fresh-water and the life-span is longer in females than nerites are observed in massive up-stream males. The populations of amphidromous migrations, consisting of both adults and species are genetically structured at scales juveniles moving together in long lines and/or similar to fully marine species. As the Copyright © Sept.-Oct., 2020; IJPAB 2

Hristov, K.K. Ind. J. Pure App. Biosci. (2020) 8(5), 1-10 ISSN: 2582 – 2845 planktotrophic larvae of amphidromous ppt for 4, 4, 1, 4, 1, 4, and 1 week, species must settle in a rare, unstable habitat, respectively. No changes in their behaviour they could be selected for the ability to delay (grazing (Fig. 1B), and sleep/wake cycles) metamorphosis, and extend their planktonic were observed. Similarly, the changes of water life indefinitely (Abdou et al., 2015). pH using SeaChem Laboratories buffering Although popular animals in the salts did not influenced their behaviour. The ornamental aquarium trade, Neritina turrita animals were kept in 2 ppt brackish-water, and (Gmelin 1791) are not successfully bred in pH altered from 5 to 9 for a week at each pH captivity, and the available animals are wild- point. Neritina turrita were observed to caught and present a potential biohazard as copulate several times (Fig. 1C), regardless of well as depleting natural populations. There the salinity (2, and 15 ppt). They laid over 630 are no reports in the available literature for egg capsules (Fig. 1D) continuously over a their successful breeding in laboratory period of 2 months in both fresh (0 ppt) and condition. The aim of the undergoing study in brackish water (2 and 10 ppt) at temperatures my laboratory is the optimisation of a breeding of 25 - 28oC. Each deposition was set apart by protocol for Neritina turrita for the ornamental 6 +/-3 days, and comprised a mean of 53 (with aquarium trade, and the growing demand of a range of 3 to 192) egg capsules. Light animal model systems for the biomedical microscopy evaluation of N. turrita egg research. capsules and veligers used a Nikon research microscope. The typical for N. turrita egg MATERIALS AND METHODS capsule structure (Fig. 2A) was observed, and Neritina turrita adult snails were kept in 2, 10, the release of early veligers from the eggs and 60 L glass vessels. Tap-water was used to when placed in tap-water (Fig.2C). achieve a salinity range of 0 (fresh-water), 2, The family Neritidae is one of the 5, 10, 15, 20 ppt (brackish water), 35 ppt (sea- most primitive families in the , and water) in conjunction with ‘Salinity for reefs’, includes members that are diadromous species Aquavitro, SeaChem Laboratories Inc., capable of escaping the instability of fresh- Madison, USA. The brackish water (2 ppt) pH water habitats (due to droughts or cyclonic was adjusted using Discus Buffer (pH 5.8-6.8); flood events), to colonise new environments Neutral Regulator (pH 7.0); Malawi Victoria through oceanic dispersal. Amphidromy is one Buffer (pH 7.8-8.4); Tanganyika Buffer (pH of the modalities of diadromy: while the adults 9.0-9.4), SeaChem Laboratories Inc., Madison, grow, feed, and reproduce in rivers and low- USA. salinity areas of estuaries, larvae drift The snails were fed with spring and downstream after hatching to reach the marine autumn oak (Quercus robur) and plum (Prunus cerasifera) leaves, sliced salinity necessary for their development, and courgettes/zucchini (Cucurbita pepo), and to complete the dispersive, planktotrophic cucumber (Cucumis sativus) peel strips. marine larval stage. Later, they recruit to Light microscope images were estuaries, colonising the adult freshwater acquired with the help of Prof. N. E. Lazarov, habitats (Quintero-Galvis & Castro, 2013). and Mrs D. Brazicova with a Nikon research Neritina turrita (Gmelin 1791) inhabits microscope, equipped with a DXM1200c intertidal and supra-tidal rocks and mangroves, digital camera in the Department of Anatomy, brackish and fresh waters on temperate to Histology and Embryology, Medical tropical coasts. University - Sofia, Sofia - 1431, Bulgaria. The roots and stems of 5 species of mangrove - Avicennia marina, Bruguiera RESULTS AND DISCUSSION gymnorrhiza, Rhizophora mucronata, Ceriops Neritina turrita (Fig. 1A) tolerated salinities tagal, and Lumnitzera racemosa, and the from 0 (fresh-water) to 35 ppt (sea-water). The surrounding mud provide a substratum for a snails were kept at 0, 2, 5, 10, 15, 20, and 35 maroon algal felt termed the bostrychietum. Copyright © Sept.-Oct., 2020; IJPAB 3

Hristov, K.K. Ind. J. Pure App. Biosci. (2020) 8(5), 1-10 ISSN: 2582 – 2845 The bostrychietum is present throughout the strips, and successfully grew and reproduce on tropical world. The pneumatophores of the that diet. white mangrove A. marina, support the richest Other species of Neritina and related algal flora. Microcoleus chthonoplastes has the genera occupy similar habitats. For example, widest biogeographical range (Lambert et al., (Reeve 1855) is a tropical 1989). These algae are common in mangroves found in estuarine habitats in South around the world. While some are Africa: from Umgeni river, Natal; as far south opportunistic epiphytes (Colaconema sp. and as Umzimkulu river and Umlalazi river to Acrochaetium globosum), or topical tropical , and northwards to the Pangani shallow water algae (Canistocarpus river, Tanganyika, , at the head of cervicornis) not specific to mangroves, others estuaries (not in rivers). In the Mngazana are associated with mangroves (Bostrychia and estuary, Transkei, N. natalensis Caloglossa). While some are widespread, have been observed in salinity of 2ppt (Prof. others are specifically limited to the Indo- G. Branch, University of Cape Town, South Pacific ocean, and other occurs in - Africa, personal communication). In Kosrae USA, Malaysia, Micronesia, New Caledonia, (Micronesia), the mangrove forest of Yela and Saudi Arabia. Low-molecular weight River is dominated by Sonneratia alba and carbohydrates are contained differentially Bruguiera gymnorrhiza at its estuary, mature within the species with different geographical fresh-water swamp forest (dominated by distribution: Bostrychia moritziana and B. Terminalia carolinensis and Pandanus spp.) radicans specimens from Australia, , up to 40m asl, and undisturbed upland forest Georgia-USA, and Florida-USA, , (dominated by Campnosperma brevipetiolata) Micronesia, Peru, and have sorbitol above 40m asl (above sea level). Zonation in and dulcitol, but populations from North the distribution of snails with elevation, or Caroline to Connecticut, USA lack dulcitol distance from the river mouth has been (West et al., 2013). observed: Neritina canalis and Melanoides The Rhodophycean algae (Bostrychia, tuberculata are found only at the highest Caloglossa, Catenella, and Murrayella) stations (temperature ~25oC and conductivity comprise the bostrychietum along with the ~106 µS/cm). Neritina pulligera is found at other classes: Cyanophyceae, Chlorophyceae, the lower stations (temperature ~27oC and Phaeophyceae, and Bacillariophyceae. Fresh- conductivity ~83 µS/cm), along with Clithon water algae are also an associate of the castanea. Migrations of other neritids similar bostrychietum. Bostrychia tenella (supratidal to those in Puerto Rico and other places has localities) and B. scorpiodes (a salt marsh not been observed (Benstead et al., 2009). species) are cosmopolitan species, recorded as During the current study adults of a mangrove epiphyte in Brazil, New Zealand, Neritina turrita were found to tolerate Sumatra and Australia. B. tangatensis is a increases in salinity from 0 (fresh-water) to 35 mangrove epiphyte in Tanzania, Mozambique, ppt (sea-water) for extended periods of time. southern Africa, Australia and Japan, No changes in their behaviour (grazing and occurring on the pneumatophores of Avicennia sleep/wake cycles) were observed. Similarly, species. There is an algal zonation on roots of the changes of water pH from 5 to 9 at 2 ppt Rhizophora in Puerto Rico, where C. ripens brackish-water did not influenced their establishes on the drier sector of the behaviour. pneumatophore (Lambert et al., 1987). Neritidae have internal fertilisation, Although herbivorous algae-eaters, Neritina and encapsulate their eggs after fertilisation. A turrita were fed with spring and autumn oak unique organ, located close to the anus and (Quercus robur) and plum (Prunus cerasifera) oviduct opening, strengthens the egg capsule. leaves, sliced courgettes/zucchini (Cucurbita The aragonitic or calcitic spherulites, 5 - 500 pepo), and cucumber (Cucumis sativus) peel µm in diameter, are secreted in the digestive Copyright © Sept.-Oct., 2020; IJPAB 4

Hristov, K.K. Ind. J. Pure App. Biosci. (2020) 8(5), 1-10 ISSN: 2582 – 2845 tract of females. When secreted from the anus, Neritids in Hawaii (Neritina granosa) and a portion is carried by cilia to the crystal sac, Puerto Rico (N. punctulata and N. virginea) where the mineral particles are sorted and are observed to participate in up-stream stored. The base half of the egg capsule is migrations in coastal and insular streams and flattened and fixed to the substrate, and the rivers in massive aggregations (>3000 ind/m2), upper convex half lifts off when the young moving up to 10 km inland with speeds of 1 - escape. The walls are made of tough 50 m/day (Pyron & Covich, 2003; Blanco & conchiolin, lined internally by a membrane Scatena, 2005; Blanco & Scatena, 2007). enclosing an albuminous fluid in which eggs Successive flash floods draw them back to the float. Neritina pulligera collected from river mouth. Up-stream migrations are Kyushu Island, Japan, and maintained in fresh- frequent - once every 15 days. Migrations water at 27oC deposited clusters of 4 - 39 large typically occur within 5 days after a flood, and egg capsules mostly on shells, while Clithon end after nearly a week of continuous corona deposited their egg capsules on the upstream movement. During non-migratory aquarium devices and stones (Kano & periods, snails are dispersed and rarely form Fukumori, 2010). Similarly, Neritina zebra, a groups of more than 5 individuals. Neritids common species of muddy bottoms in brackish prefer large objects for attachment, more water environments found in , and turbulent and erosive habitats, and reahc along the cost of Brazil to Cabo Frio, Rio de greater densities in riffles and fast-flowing Janeiro, produces oval 1 - 1.5 mm-long egg areas. undergoes significant capsules containing ~68 eggs (32 - 106) that growth during upstream migrations. The become darker as development progresses upstream migration is probably a response to (Barroso & Mattews-Cascon, 2009). higher predation downstream, higher Regardless of the salinity (2 and 15 ppt), N. productivity upstream, and the tolerances of turrita were observed to copulate several times physical parameters (salinity and temperature) (Fig. 1C). Likewise, they laid over 630 egg by adults and juveniles (Pyron & Covich, capsules (Fig. 1D) continuously over a period 2003; Blanco & Scatena, 2005; Blanco & of 2 months in both fresh and brackish water Scatena, 2007). Interestingly, a number of (2 and 10 ppt). Each deposition was set 6 (2 - rivers in dry southern Puerto Rico lack 11) days apart, consisting on average of 53 (3 - populations of N. virginea. They are absent 192) egg capsules. from streams draining limestone that have high

Salinity is critical for hatching: egg concentrations of CaCO3, greater total capsules of N. zebra release veligers after 21 suspended solids and SiO2, elevated days at salinity 5 ppt (26-28oC). Despite the conductivity and high concentrations of presence of well-formed veligers, eggs do not dissolved ions (Na, Cl, K, Mg, and SO4) hatch at higher salinity (15 ppt), and (Blanco & Scatena, 2006). development is slower (25 days). Decreasing The settlement, metamorphosis, the salinity to 10 ppt results in the release of development and growth of veliger larvae of the veligers (Barroso & Mattews-Cascon, many marine gastropods have been 2009). In this study, N. zebra veligers could be extensively studied. Water-borne chemical and kept alive only for 2 days after leaving the egg substrate-associated cues (algal-surface capsules (Dr. C. X. Barroso, Universidade ligands, barnacle-associated substances, and Federal do Ceara, Instituto de Ciencias do bacterial-biofilm substances) induce settlement Mar, Brazil, personal communication). Unlike and metamorphosis (Pires & Hadfield, 1993; N. pulligera (Kano & Fukumori, 2010), and N. Plaut et al., 1995; Hadfield et al., 2000; zebra (Barroso & Mattews-Cascon, 2009), Nedved & Hadfield, 2009; Ruiz-Jones & which prey on their own egg capsules, N. Hadfield, 2011). The morphogenic effects are turrita was not observed to harm its own egg mediated through the central nervous system capsules. (Pires & Hadfield, 1993). K+ and Cs+ have Copyright © Sept.-Oct., 2020; IJPAB 5

Hristov, K.K. Ind. J. Pure App. Biosci. (2020) 8(5), 1-10 ISSN: 2582 – 2845 been shown as a potent metamorphosis with zebra (Dreissena polymorpha Pallas inducers, acting by depolarising the sensory 1769) and quagga mussels (Dreissena cells responding to the natural metamorphosis rostiformis bugensis Andrusov 1897). inducer, and also on the entire nervous system Interestingly, sodium concentration affects (Hadfield et al., 2000). It should be noted that veliger survival, and supplementation with potassium is toxic to fresh-water molluscs, and NaCl increases their survival (Moffitt et al., KCl, as potash has been suggested as a 2016). potential pesticide for treatment of infestation

Figure 1: Neritina turrita (Gmelin, 1791) were maintained in fresh, brackish (salinity from 2 to 20 ppt) and sea water (35 ppt) at 25 - 28oC (A). The grazing pattern on algae-covered glass surface (B). Copulating pair: the male is on the left, and the protruding penis is visible (C). N. turrita laid over 630 egg capsules, continuously within 2 months. Each deposition was set apart 6 +/-3 days, consisting on average of 53 (from 3 to 192) egg capsules (D).

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Hristov, K.K. Ind. J. Pure App. Biosci. (2020) 8(5), 1-10 ISSN: 2582 – 2845

Figure 2: Light microscopy evaluation of Neritina turrita (Gmelin, 1791) egg capsules. The distinct for N. turrita egg capsule structure (magnification 10x in A, and 20x in B) was observed. Egg mass after its release from the egg capsule (D), and the release of early veligers from the eggs when placed in tap-water (C, magnification 20x).

CONCLUSIONS metamorphosis-inducing cues present in the Neritina turrita snails are a hardy species, natural estuary habitat that regulate veliger’s surviving increase in salinity to levels of sea- faith. ii) Similarly to other amphidromous water, and change in acidity from pH 5 to 9. species (like Caridina multidentata for which Currently, I am working on two hypotheses there is an established breeding protocol), aimed at producing a reliable breeding veligers spend a short time in fresh-water after protocol: i) Everything happens in the estuary, hatching and are carried down-river to the where veligers stay in the mud after hatching, estuary, where they face the increased salinity, but those that don’t stay there have the ability necessary for the induction of metamorphosis to survive at sea, and disperse to new and the maturation of benthic juveniles. Is the locations. The adult snails are maintained at increased salinity acting as a cue that grounds the same below-sea-water salinity. I am the veligers in the mud at the estuary? As looking for developmental and noted (Abdou et al., 2015), considering the Copyright © Sept.-Oct., 2020; IJPAB 7

Hristov, K.K. Ind. J. Pure App. Biosci. (2020) 8(5), 1-10 ISSN: 2582 – 2845 high levels of larval mortality, and the effects Laboratory, University of Hawaii, USA); Dr. of diffusion, it seems unlikely that significant K. Stockton-Fiti (KASF Consulting LLC, numbers of larvae that drift away from their Henderson, USA); and Prof. J. R. Pawlik natal archipelago would be able to find (Department of Biology and Marine Biology, suitable fresh-water habitat for settlement. UNCW Center fro Marine Science, USA) for Therefore, local selection for traits that favour the valuable suggestions. self-recruitment is particularly strong for amphidromous species. Further, massive REFERENCES aggregations (>3000 ind/m2) of adults as well Abdou, A., Keith, P., & Galzin, R. (2015). as juveniles are observed to migrate up-stream Freshwater Neritids (: in coastal and insular streams and rivers Gastropoda) of tropical islands: (Pyron & Covich, 2003; Blanco & Scatena, Amphidromy as a life cycle, a review. 2005; Blanco & Scatena, 2007). They come Revue d’Ecologie (Terre et Vie) 70(4), from the estuary, where they breed, lay egg 387-397. capsules, and grow in their hundreds and Bandel, K. (2001). The history of thousands. Even the new arrivals, brought into and Neritina connected with the estuary habitats continuously by the ocean description and systematic evaluation currents have to grow and breed in this of related (Gastropoda). environment to achieve the numbers of snails Mitteilungen aus dem Geologisch- migrating up-river. It will be interesting to Palaontologischen Institut (GPI) der determine the reasons why some veligers stay Universitat Hamburg 85, 65-164. in the estuary and grow to move up-river later, Barroso, C. X., & Mattews-Cascon, H. (2009). and why some survive at sea and disperse. Spawning and intra-capsular development of Neritina zebra Acknowledgments (Bruguiere, 1792) (Mollusca: The author would like to express his gratitudes Gastropoda: Neritidae) under to Prof. N. E. Lazarov, and Mrs D. Brazicova laboratory conditions. Invertebrate for their help during the acquisition of the Reproduction and Development 53(3), results in the Department of Anatomy, 137-143. Histology and Embryology, Medical Barroso, C. X., Mattews-Cascon, H., & University - Sofia, Bulgaria; and also to Dr. Simone, L. R. L. (2012). Anatomy of M. Wilstermann-Hildebrand (Heimbiotop - Neritina zebra from Guyana and Aquaristik, Wirbellose und Garten, Brazil (Mollusca: Gastropoda: Warendorf, Germany); Dr. Y. Kano Neritidae). Journal of Conchology. (Department of Marine Ecosystems Dynamics 41(1), 49-64. Atmosphere and Ocean Research Institute, the Blanco, J. F., & Scatena, F. N. (2005). Floods, University of Tokyo, Japan); Emeritus Prof. G. habitat hydraulics and upstream Branch (Department of Biological Sciences, migration of Neritina virginea University of Cape Town, South Africa); (Gastropoda: Neritidae) in Emeritus Prof. C. Griffiths (Department of Northeastern Puerto Rico. Caribbean Biological Sciences, University of Cape Town, Journal of Science 41(1), 55-74. South Africa); Prof. Tony Pires (Dickinson Blanco, J. F., & Scatena, F. N. (2006). College, USA); Dr. C. X. Barroso Hierarchical contribution of river- (Universidade Federal do Ceara, Instituto de ocean connectivity, water chemistry, Ciencias do Mar, Brazil); Dr. M. Pyron hydraulics, and substrate to the (Department of Biology, Ball State University, distribution of diadromous snails in USA); Prof. J. F. Blanco-Libreros (Instituto de Puerto Rican streams. Journal of Biologia, University of Antioquia, ); North American Benthological Society Prof. M. G. Hadfield (Kewalo Marine 25(1), 82-98. Copyright © Sept.-Oct., 2020; IJPAB 8

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