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Taxonomy, Conservation, and the Future of Native Aquatic Snails in the Hawaiian Islands

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Taxonomy, Conservation, and the Future of Native Aquatic Snails in the Hawaiian Islands diversity Perspective Taxonomy, Conservation, and the Future of Native Aquatic Snails in the Hawaiian Islands Carl C. Christensen 1,2, Kenneth A. Hayes 1,2,* and Norine W. Yeung 1,2 1 Bernice Pauahi Bishop Museum, Honolulu, HI 96817, USA; [email protected] (C.C.C.); [email protected] (N.W.Y.) 2 Pacific Biosciences Research Center, University of Hawaii, Honolulu, HI 96822, USA * Correspondence: [email protected] Abstract: Freshwater systems are among the most threatened habitats in the world and the biodi- versity inhabiting them is disappearing quickly. The Hawaiian Archipelago has a small but highly endemic and threatened group of freshwater snails, with eight species in three families (Neritidae, Lymnaeidae, and Cochliopidae). Anthropogenically mediated habitat modifications (i.e., changes in land and water use) and invasive species (e.g., Euglandina spp., non-native sciomyzids) are among the biggest threats to freshwater snails in Hawaii. Currently, only three species are protected either federally (U.S. Endangered Species Act; Erinna newcombi) or by Hawaii State legislation (Neritona granosa, and Neripteron vespertinum). Here, we review the taxonomic and conservation status of Hawaii’s freshwater snails and describe historical and contemporary impacts to their habitats. We conclude by recommending some basic actions that are needed immediately to conserve these species. Without a full understanding of these species’ identities, distributions, habitat requirements, and threats, many will not survive the next decade, and we will have irretrievably lost more of the unique Citation: Christensen, C.C.; Hayes, books from the evolutionary library of life on Earth. K.A.; Yeung, N.W. Taxonomy, Conservation, and the Future of Keywords: Pacific Islands; Gastropoda; endemic; Lymnaeidae; Neritidae; Cochliopidae Native Aquatic Snails in the Hawaiian Islands. Diversity 2021, 13, 215. https://doi.org/10.3390/ d13050215 1. Introduction Freshwater ecosystems are negatively impacted by anthropogenic threats and almost Academic Editors: Russell Minton one in three freshwater species is threatened with extinction [1]. This extinction plight and Luc Legal is well reflected in freshwater molluscan faunas [2]. Globally, the number of freshwater Received: 15 April 2021 molluscs threatened with extinction varies, with 44% in Europe [3], 29% in Africa [4] Accepted: 14 May 2021 and 17% in Indo-Burma [5]. More than a decade ago at least two-thirds of all species of Published: 18 May 2021 unionid mussels from Canada and the United States were listed as extinct, possibly extinct, critically imperiled, imperiled, or vulnerable [6], and nearly 41% of all freshwater mussel Publisher’s Note: MDPI stays neutral species assessed by the IUCN are considered Near Threatened, Vulnerable, Endangered, with regard to jurisdictional claims in or Critically Endangered [7]. The conservation status of nearly 100% of mammal and bird published maps and institutional affil- species is known, but only 7% of mollusc species have conservation status assessments by iations. IUCN and the extinction rate for poorly studied molluscs is probably orders of magnitude greater than currently estimated [8–11]. Many of these unrecorded extinctions are occurring on Pacific Islands such as Hawaii [12–14]. The Hawaiian archipelago is one of the most isolated landmasses in the world, Copyright: © 2021 by the authors. with California over 3700 km to the northeast and Japan nearly 6000 km to the west– Licensee MDPI, Basel, Switzerland. northwest [15]. The Hawaiian Islands have a small but highly specialized native stream This article is an open access article fauna (Figure1), and colonizers have reached these islands by dispersal through or over distributed under the terms and substantial expanses of ocean, either as planktonic veligers or as passive hitchhikers on conditions of the Creative Commons migrating birds or other dispersal vectors [15,16]. Attribution (CC BY) license (https:// creativecommons.org/licenses/by/ 4.0/). Diversity 2021, 13, 215. https://doi.org/10.3390/d13050215 https://www.mdpi.com/journal/diversity Diversity 2021, 13, 215 2 of 12 Diversity 2021, 13, x FOR PEER REVIEW 2 of 12 Figure 1. Live representatives of Hawaii’s endemic freshwater snail fauna. (A) Neripteron cariosum (Wood, 1828), (B) Figure 1. Live representatives of Hawaii’s endemic freshwater snail fauna. (A) Neripteron cariosum (Wood, 1828), (B) Neritona Neritona granosa (Sowerby, 1825), (C) and Neripteron vespertinum (Sowerby, 1849), (D) Erinna newcombi H. Adams and A. granosa Neripteron vespertinum Erinna newcombi in Adams(Sowerby, in A. Adams, 1825), 1855, (C) and (E,F) Pseudosidora c.f. producta(Sowerby, (Mighels, 1849), 1845) (D) in its natural seepH. habitat Adams (E and) and A. in Adams artificialA. Adams,habitat 1855,in the (Elaboratory,F) Pseudosidora (F). Photoc.f. productacredits: (A(Mighels,–C) Keoki 1845) Stender, in its (D natural) Gordon seep Smith, habitat U.S. (E )Fish and and in artificialWildlife habitatService, in and the laboratory(E,F) Kenneth (F). PhotoHayes. credits: (A–C) Keoki Stender, (D) Gordon Smith, U.S. Fish and Wildlife Service, and (E,F) Kenneth Hayes. Hawaiian streams are highly variable in flow, spatially and temporally, with sudden peaksHawaiian and long streams troughs are in highly flow rates. variable Peak in flow,flows spatially aid the andflushing temporally, of debris with from sudden the peaksstreambeds, and long which troughs in turn in signal flow rates. to aquati Peakc organisms flows aid theto migrate flushing and of spawn debris from[15–18]. the streambeds,Native Hawaiian which stream in turn species signal have to adapted aquatic to organisms these variable to migrate conditions, and and spawn many, [15 but–18 ]. Nativenot all, Hawaiiannon-native stream species species are unable have adapted to survive to these in these variable freshwater conditions, habitats and many,[16]. butAdaptations not all, non-nativein native species species include are unable freshwat to surviveer fish with in these fused freshwater pelvic fins habitats that allow [16 ]. Adaptationsthem to cling into nativesubstrate species during include torrential freshwater flows or to fish climb with waterfalls, fused pelvic and finsseveral that native allow themstream to fishes, cling tocrustaceans, substrate during and molluscs torrential are flows amphidromous or to climb [19–22]. waterfalls, Unfortunately, and several many native streamof the low-elevation fishes, crustaceans, freshwater and molluscs habitats are in amphidromousthe islands have [19 been–22 ].heavily Unfortunately, impacted many by ofdevelopment the low-elevation and are freshwaterdominated by habitats introduced in the species, islands leaving have beenthe remaining heavily impacted native taxa by developmentto seek refuge andin the are less dominated impacted upper by introduced elevations species, reaches leaving[23]. In turn, the remaininghigher-elevation native taxahabitats to seek are refugeimpacted in theby the less alteration impacted of upper the lower elevations reaches, reaches making [23 ].even In turn,the upper- higher- elevationelevation habitatsrefuges susceptible are impacted to invasion by the alteration and impacts of theof introduced lower reaches, species. making even the upper-elevationAlthough the refuges Hawaiian susceptible aquatic to malacofauna invasion and is impacts small, eight of introduced recognized species. species are presentAlthough in three the families: Hawaiian Neritidae, aquatic Lymnae malacofaunaidae, and is small,Cochliopidae. eight recognized The fauna species is highly are presentendemic, in threeand native families: species Neritidae, inhabit Lymnaeidae, various fresh- and Cochliopidae.and brackish-water The fauna habitats is highly from en- demic,estuaries and and native coastal species ponds inhabit to variousfreshwater fresh- springs and brackish-water and seeps. habitatsHere, we from provide estuaries a andtaxonomic coastal overview ponds to freshwaterof the Hawaiian springs freshwat and seeps.er molluscs, Here, we the provide impacts a taxonomic that have reduced overview oftheir the numbers, Hawaiian their freshwater conservation molluscs, status, the and impacts what thatsteps have must reduced be taken their to conserve numbers, them their conservationnow and in the status, future. and what steps must be taken to conserve them now and in the future. 2.2. TaxonomicTaxonomic OverviewOverview of the Fauna WithinWithin thethe Neritidae, Neritidae, there there are are three thre endemice endemic species; species; two two of these, of these,Neripteron Neripteron vesper- tinumvespertinum(Sowerby, (Sowerby, 1849) and1849)N. and cariosum N. cariosum(Wood, (Wood, 1828), 1828), are euryhaline are euryhaline and inhabit and inhabit coastal ponds,coastal anchialineponds, anchialine pools, stream pools, mouths,stream mouths and estuaries, and estuaries [24,25]. [24,25]. The third The species, third species,Neritona granosaNeritona(Sowerby, granosa (Sowerby, 1825), is 1825), anadromous is anadromous (Figure (Figure2). Adults 2). Adults of this of species this species live in live clear in steep-gradientclear steep-gradient streams streams from from sea level sea level to at to least at least 400 400 m in m elevation, in elevation, and and planktonic planktonic lar- vaelarvae develop develop in in the the sea sea before before returning returning to to freshwater freshwater environmentsenvironments [24,26,27]. [24,26,27]. Eggs Eggs are are attachedattached to boulders
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