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http://dx.doi.org/10.18268/BSGM2019v71n3a2 P. 609‒629 VOL. 71NO. 3 BOL. SOC.GEOL.MEX.2019 2202, USA. Georgia, Athens, GA 30602- University of Ecology, Ecology, OdumSchool of of Institute Alan P. Covich Palacio, Durango, Mexico. Juárez del Estado de Durango, 35010 Gómez Facultad de Ciencias Biológicas, Universidad [email protected] José GamalielCastañeda-Gaytán David Ramiro Aguillón-Gutiérrez Jorge LuisBecerra-López Verónica Ávila-Rodríguez Iris GabrielaMeza-Sánchez Jorge Saenz-Mata Ulises Romero-Méndez José LuisEstrada-Rodríguez Alexander Czaja David Ramiro Iris Gabriela A “silent” localextirpation,withthedescriptionofanewspecies Fossil freshwatergastropodsfromnorthernMexico–Acaseofa Manuscript October 2,2018. accepted: Corrected manuscript received: 25,2018. September Manuscript received: August 29,2018. lexander Czaja Meza-Sánchez Aguillón-Gutiérrez , AlanP. Covich , Verónica , José Luis , José Gamaliel gastropods, habitat. freshwater, lossof Keywords: Paleobiology, extirpation, the Cuatrociénegas valley of still threatens the freshwater snailcommunities study that inthe area led to localextirpation of Cuatrociénegas same imperilment Basin.The diversity inNorth America remained inthe gastropods the greatest recent hotspots of of two hundred farther north, one kilometers (unrecorded) while at the same time some freshwater snailshappened silently pation of events.extirpation Inthis study area the extir are highlysensitive andvulnerable to such ranges (localendemics) with smallgeographic results show that especially hydrobiid loss caused by anthropogenic activities. The the area due to habitat inthe 20th century them disappeared from At least 24(75.0%)of Holocene onlyfour (12.5%)are still extant. study through the present in the area of freshwater 32species of the 20th century. Of of ently increased rapidlyinthe second half species appar Middle Holocene. lossof This Mexico that inthe pation began innorthern a“silent” localextir herein anexample of paleomalacological studies, weformer present recent times. Together with data from our them disappeared invery Mexico andallof genera) have not been known asfossil from these species (andsome gastropods. Most of cochliopid, hydrobid, planorbidandneritid subfossil and report several new records of and Durango. Here we describe anew species the LagunaDistrict, area asmall inCoahuila contains subfossil (Late of Holocene) deposits recent analogue ecosystems. study The site of conservation important perspectives infavor of called Conservation Paleobiology, canprovide events. new paleoecological approach, This distributions and local extinction or extirpation and are often the onlyway to document past cases, paleontological data canbe very helpful Mexican freshwater gastropods. Insuch of on the distribution but nodata onextinctions Mexico.Northern There are fewhistorical data Coahuila and Durango, Chihuahuan Desert of pronounced inaridregions such asinthe a worldwide phenomenon but it isespecially freshwater species diversity is decline of The ABSTRACT Ávila-Rodríguez Estrada-Rodríguez Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Castañeda-Gaytán , Jorge Luis . , Ulises Becerra-López - - - Romero-Méndez distribución y ninguna información sobre extinciones distribución yninguna información de México. Existen pocos datos históricos sobre la Chihuahuense de Coahuila yDurango, norte-centro áridas comoenelDesierto nunciado enlas regiones es unfenómeno mundial, pero esespecialmente pro El declive de la diversidad de especies dulceacuícolas RESUMEN hábitat. gasterópodos, dulceacuícolas, P dulceacuícolas del valle de Cuatrociénegas. estudio aún amenazan a las comunidades de caracoles local ennuestraque llevaron a la extirpación área de la cuenca de Cuatrociénegas. Las mismas amenazas en de gasterópodos enAmérica del Norte permaneció más al norte, unode los mayores focos de diversidad tras que al mismo tiempo unos doscientos kilómetros mien silenciosamente (sin registros), estudio ocurrió de los caracolesextirpación dulceacuícolas en el área de vulnerables La a tales acontecimientos de extirpación. (especies endémicas locales) son altamente sensibles y pequeños geográficos los hidróbidos conintervalos nicas. Nuestros resultados muestran que especialmente pérdida de hábitat causada por actividades antropogé ellos desaparecieron del área enelsiglo XXdebido a la Al menos 24(75.0%) de 4 (12.5%)aún existen. presentes enelárea de estudio durante elHoloceno, solo XX. Delas 32 especies de caracoles dulceacuícolas mitad del siglo cies aumentó rápidamente enla segunda comenzó enelHolocenoMedio. Esta pérdida de espe pación local “silenciosa” enelnorte deMéxicola cual anteriores, presentamos aquí unejemplo de una extir con datos de nuestros estudios paleomalacológicos desaparecieron entiempos muyrecientes. Enconjunto ros no se conocencomofósiles de Méxicoytodos ellos neritidos. La mayoría de estas especies yalgunos géne gasterópodos cochliópidos, hidróbidos, planórbidos y de nuevos varios registros especie yreportamos nueva Durango. fósiles describimos una Deestos depósitos unárea pequeña enCoahuila y del Distrito Laguna, fósiles (HolocenoTardío)de estudio contienedepósitos de ecosistemas análogos recientes.conservación Elsitio cionar perspectivas importantes para los esfuerzos de llamado Paleobiología puede propor de Conservación, enfoque paleoecológico, Este nuevo ción uextirpación. locales de extin distribuciones pasadas ylos eventos y, de documentar las a menudo, son la únicaforma casos, los datos paleontológicos pueden ser muyútiles de gasterópodos dulceacuícolas mexicanos. Entales alabras clave: Paleobiología, extirpación, , , Jorge /2019 Saenz-Mata pérdida de 609 , ------

Freshwater snails – a case of “silent” local extirpation from Mexico ABSTRACT INTRODUCTION / MATERIAL AND Freshwater snails – a case of “silent” local extirpation from Mexico METHODS 610 610 1. Introduction water-fed springs, andother habitats are threat where landscapes those inkarst as lakes, ground endemic specieswithrestricted distributions, such wideinterest especially among and occurs is of loss freshwater due tohabitatsnails extinctionThe of rmti eidsr n as eino south From region thissemi-desert and karst of study.inland lakesand springsinthearea of inated the habitats provided by the remaining elim finally (2013), Wolfe by detail in described pumping.groundwater aquifer depletion, This springs, andespecially by excessive alteration of the two rivers, as dammingandchannelization of accompanied bywas habitat perturbations such inMexico.and industry economicprocess This agriculture region became an important center of al. the early 1900s(Ortega-Guerrero, 2003; Butzer the Holocene, large lakes were still present until during smaller significantly become had lake this (Butzer America duringLate Pleistocene/Early Holocene the largest paleo-lakes in Latin basin, wasoneof Mexico (Figure region, 1). The alarge endorheic Durangostates, Coahuila and eastern northern western it comprises the adjoining portionsof regionthe LagunaDistrict,asmallkarst and studyThe area is located in the central part of impossible. shell convergences,or, almost as insomecasesof difficult, more much be would material subfossil of the apart of research systematicaffiliation our 2011a, 2011b, 2014a,2014b). Without these data (Hershler fromonly shellmorphology and hydrobiidor speciespreviously snails known newlycollected extant cochliopid biogeographyof advances have focused mainlyonsystematics and anatomically orusingmolecular methods. Recent livingspecieshave been studied ecologically,of diversionsa portion Only and damconstruction. water habitats are rapidly being altered by water Mexico ispoorly northern known andfresh of extant freshwater gastropods distributionThe of ened ( / , 2008). In the first half of the 20th century the century 20th the of half first the In 2008). , Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín e.g et al., ., Strong 2008;Czaja et al . 2008,Régnier et al ., 2014b). Although et al. et al. 2009). , 2002, et - - - - - /2019 2. Materialandmethods Guerrero-Arenas andJiménez-Hidalgo(2015). potential alreadyhas been pointed out indetail by far only once (Figueroa-Rangel this new interdisciplinary approach wasapplied so the present-dayunderstand conditions. InMexico, (2011) which utilizes the fossiland Flessa record to we follow the“near-time” approach their habitats inrecent times. Toof achieve that, distributionoriginal biogeographic priortothe loss a new species, report new records andshow their the present study isto describe aimof era. The planorbid andneritid freshwater gastropod gen subfossil shellsbelonging to cochliopid, hydrobiid, west Coahuila and northeast Durango, we report neighboring site was dated by Butzer but the same horizon withabundant snails from a sediments with These Paleo-Lake (Figure the former Irritila 1, site 1). of silt horizongray with abundant planorbid snails Vitta Viesca shell of (Figuresingle 1,sites5and4).A the paleo-lake in lacustrine sub-recentof deposits and River Aguanaval the of deposits fluviatile in and riograndensis bothspeciesof Shells of these springs (Figure 1, site 3). wetland area of Viesca a former springs in superficial sediments of nensis 1959; Wolfe, 2013).Shells of and dried out in the following years (Blásquez, ure 1, site 2). All springs were still active in 1959 Viesca, Coahuila (Fig near springs to the townof dried-out of sediments superficial present-day in Landye, 2011), lor, 1966), Shells of the bottom. at BP years ̴7130 than older not and section the brated Holocene playaas with radiometric cali deposits cf. Dilatata dilatata (Pfeiffer, 1839) were collected at west of west at collected were 1839) (Pfeiffer, 14 cf. gsbten32 0 ttetpo C ages between 3320 ± 100 at the top of usnea seemani gracilis gracilis Juturnia Pseudotryonia pasajae Pseudotryonia (Röding, 1798) was collected in a Promenetus exacuous (Frauenfeld, 1863) were collected (Gould, 1841) were collected Vitta (Pilsbry andFerriss,(Pilsbry 1906) Probythinella n. sp.,n. were not dated directly (Hershler, Liuand et al. hava J. coahuilae (Say, 1821)and Thiele, 1929, , 2012) but its et al. sensu (2008) (Tay Dietl - - - - - Area ofstudy. Lacustrine sub-recent of deposits thePaleo-Lake of Viesca.5=Sub-recentfluviatiledeposits theAguanavalRiver.Dottedsquares= of thePaleo-LakeIrritilaplaya deposits (Laguna Mayrán).2=Springs ofViesca.3=Former wetland area of thesprings of Viesca.4= Figure 1 criminant analysis was carried out. This was done was out. This carried was criminant analysis differences shell between subfossiland livingpopulations, adis morphological visualize To through sieves. vegetation, 4kgsediment per site was screened were hand collected from stones, sandandaquatic sites were snails last sampled in summer2017.The monitored by three 2009.These since the authors the localities that are the contains coordinates of study. water in the area Table of permanent 1 riversLa Peñaand thesmallspring stillcontain the Nazas and Aguanaval the dams of outflows of thisspring. Onlythe directly from sedimentsof Cuatrociénegaswere collected2017 summer in the extant the typelocalityforis species. this of shells Empty spring inSan Pedro de Ocuila, Durango, which 2016 fromcollected in September anunnamed hlso theextant species of shells the subfossilmaterial, empty For a comparison of Mapof thestudyareawithlocalizationof thesampled sitesof theLagunaof Coahuilaand District Durango. 1=Holocene Juturnia coahuilae Juturnia Pseudotryonia pasajae Pseudotryonia from Poza Churincein were were Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - the same microscope. Shell whorls were counted the shells were measured using ric variables of Stemi a Zeiss 2000-C microscope. Morphomet with a Zeiss AxioCamERc5s-camera attached to remove sediments.were Allshells photographed shells) were cleaned hydrogenusing peroxide to the (especially theaperturesSome specimens of two sieves with0.5mmand0.3meshsize. all localities were screened through Sediments of measurements). deviation andsamplesize are given intext (shell body whorl. mean, standard The and widthof bodywhorl ture height,aperture width,height of whorls, shell height, shell width, aper number of dataset (excludingmorphometric ratios): Total pods (Czaja other hydrobiid andcochliopid gastro shells of frequently been used to differentiate closely similar v 3.14(Hammer the softwareusing PAST (PAleontogical STatistics) et al ., 2017b).We used following shell et al., 2001). This technique 2001). This has /2019 611 611 - - -

Freshwater snails – a case of “silent” local extirpation from Mexico MATERIAL AND METHODS MATERIAL AND METHODS / Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS 612 612 3.1. SYSTEMATICS 3. Results Aguanaval riversand thesmallspring LaPeña). still containpermanent of waterinthestudyarea(outflowsofdamsNazas threelocalitiesthat Table 1.Thecoordinates and Site 3 Site 2 Site 1 supportive inthesedeterminations. neo- andpaleobiological approach ismutually of and population genetic aspects. Acombination considerationsspecies that includes biogeographic living of species concepts on insightfrom modern ever, are based definitions also morphospecies our paleobiological and paleontological studies. How widely used for species concept the morphological We emphasize that in the present paper we apply Malacological Collection. standard deviation; UJMC=UniversityJuárez body whorl; WBW, body whorl; sd, of width of aperture height;AW, aperture width;HBW, height whorls; SH,shellheight; SW,of shellwidth;AH, Abbreviations usedinthe text: WN, total number Mexico (CNMO). University of Mollusks, National Autonomous Collection of the new specieswere inthe National deposited of Durango(UJED).Paratypes State University of the Juarez Biological Science of the Faculty of of material is housedinthe Malacological Collection (1939). The Pilsbry according to the method of / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Type species: Superfamily NeritaceaLamarck, 1809 Subfamily NeritininaePoey, 1852 Family NeritidaeLamarck, 1809 Vitta 53′0 ,132′1 25°01′36″N,103°16′09″W 25°39′30″ N,103°22′51″W 25°13′14″N,103°26′58″W 25°27′55″ N,103°42′12″W 52′6 ,134′3 50′9 ,132′2 25°27′02″N, 25°07′49″N,103°22′22″W 25°27′56″ N,103°42′13″W cf. (Figure 2ato2e) Nerita virginea Nerita virginea Nazas River usnea Vitta (Röding, 1798) Mörch, 1852 Linnaeus, 1758 - /2019 Aguanaval River stitutes most of shell, surface smooth and porcel shell,surfacesmooth stitutes most of whorls rapidlyincreasing in size, bodywhorl con height 7.82 mm,width 6.79 mm,with 3.00 whorls, Description: the genus follow Eichhorst (2016) and placed this speciesin (Say, the recent literature, but we 1822) in much of visible.reported speciesis as The normally is than teeth the of more exposing off, leads uptothese teeth probablybeen brokenhas that outer shell layerThe on the parietal shelf on modern than defined more much the Further, specimen showsthe teeth on the columellar tentatively. identified be only can the shell,ourmaterialon out colorandpattern Remarks: coast), West Indies (Eichhorst, 2016). Distribution: Material Examined: = 3.00. 6.79 mm, AH = 5.49 mm, AW = 3.67 mm,WN Shell Measurements: visible only nearthesutures (Figure 2d). original color pattering (thin, parallel axial lines) the allel, brownto black remainsaxial linesas of the inner lip; numerous thin,par the bottom of teeth (Figure 2e), teeth absent from the top and straight with 12(or 13?) irregular parietal shelf ovate, the outer lip is thickened and without teeth, closely spacedonawhitish background; aperture dark, wavylines, which are suturesof consisting the sculpturing, visible in the area color pattern of microscopicaxial growth linesandnospiral sutureslanous; faintlyimpressed, withnumerous Vitta Because it is a single specimen with Because itisasingle Shell dextral, ovoid, with flat ,flat ovoid,dextral,with Shell because it lacks the characteristic Florida to Texas,Florida Mexico (gulf 1 specimen(UJMC360). La Peñaspring SH = 7.82 mm, SW = SH=7.82mm, 102°35′10″ W reclivata Vitta usnea s - - - - . UJMC361a. g)SpecimenUJMC361bwithcords onthebodywhorl. h)SpecimenUJMC361c,protoconchview. axial linesasremainsof theoriginal colorpattering. e)Parietalshelf withirregular teeth. f-h) Figure 2

Vitta cf. usnea (Röding, 1798)and Cochliopina riograndensis Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (Pilsbry&Ferriss, 1906).a-e)V. cf. Cochliopina riograndensis. usnea , UJMC360.d)Black /2019 f)Specimen 613 613

Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS 614 614 species hasa wide ecological range, occurrences 1000 km from the coast is notable. Although the Late Holocene presence in the study area almost (Eichhorst, 2016). However,inland inFlorida its foundfaces andwas freshwaterin living 160 km water, onrocks, plant stems, orother hard sur of hlso the genus shells of C. description of andFerrissPilsbry (1906) noticed in the original details to thosefromTexas.the typelocalityin inall and Coahuila correspond morphologically Remarks: Potosí, Tamaulipas) 2011). (Thompson, Texas andNorthwest Mexico (Coahuila, San Luis Distribution: 361a-j). Material Examined: (0.08) mm,WN=3.40(0.21). AWAH =1.15(0.08)mm, (0.31) mm, =1.02 SW theses; N=10)SH 1.65 (0.28)mm, = 2.41 Shell Measurements: fused tothepenultimate whorl. adapically; peristome thin, innerlippartly angled sutures;deep the aperture is roundly ovate and nent (Figure 2g); whorls moderately convex with one thread atmore theshoulderusually promi spiral threads or(two) cords onthe bodywhorls, ture; teleoconch nearly smoothor frequently with openly umbilicate; apical whorl withpitted struc height 1.39–2.02 mm,width 1.91–3.01 mm, Description: Type species: 100 and̴7130years BP, Butzer (radiometric calibrated Last recordthe area: in Middle/Late Holocene itats are not known initspresent-day distribution. in freshwater at long distances from the coast hab / Cochliopina riograndensis Cochliopina riograndensis Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Neritina. V. usnea Superfamily Gray, 1840 riss, 1906(by originaldesignation) Family Tryon, 1866 Genus The subfossilfromshells The Durango Shell small, broadly heliciform, Shell small, broadly heliciform, riograndensis Cochliopina Streams inVal Verde County, Fgr fto2h) (Figure 2f riograndensis , as it occurs infresh/brackish, asit occurs Valvata (Pilsbry andFerriss,(Pilsbry 1906) 14 10specimens(UJMC (mean, with sd in parenin (mean,with sd Morrison, 1946 Morrison, C agesbetween 3320± (Müller, 1773), but they agreat similarityto et al Pilsbry & Fer Pilsbry ., 2008). ------/2019 Juturnia gracilis Juturnia Type species: the shellsintogenus tures are not present in ourmaterial weso place which are radially interconnected. Both shell fea numerous spirallirae valvatid of sculpture in form the protoconchsof rounded apertures. According to Riedel (1993), thelastgenushave emphasized thatof species ( having larger shells with considerably more whorls resembles thegenus of species Diagnosis: Differential mm, AH= 1.07mm, AW = 1.01mm,WN= 10¼. SW = 1.47mm,HBW= 1.97mm,WBW = 1.30 Holotype: Measurements of (0.12) mm,WN=10.07(0.62). WBW =1.29 HBW =2.05 (0.21)mm, (0.08) mm, AWAH =1.11(0.11)mm, (0.13) mm, =0.95 SW theses; N=15)SH 5.75 (0.61)mm, = 1.49 Shell Measurements: plete; umbilicusrimate tobroadly open. thin orslightlythickened,theperistome com is to completely separate from parietal wall, outer lip inner lipthintoslightlythickened,broadly adnate 0.7–1.0 whorl, smooth(Figure 3g); aperture ovate, to brown periostracum preserved; protoconch of or transparent, in somespecimensfrom light tan spaced growth lines, from light tan to beige-white closely smooth whorls; teleoconch sculpture of wide, with8.00to11½strongly convex and 4.82–7.32 mmshell length and 1.29–1.78 shell Description: thin; umbilicusrimate tobroadly open. growth lines;aperture ovate, innerand outer lip spaced closely and fine very with only sculpture, 11½ stronglyconvex whorls; teleoconchwithout 8.00to ile, lighttanortransparent, teleoconchof Diagnosis: Genus J. coahuilae Juturnia and Romero-Méndez sp. nov. J. coahuilae is possibly is the described freshwater Shell medium-sized, turriform, frag Shell medium-sized, turriform, Durangonella coahuilae Shell turriform, medium-sized, Shell turriform, Czaja,Covich, Estrada-Rodríguez original designation) Hershler, LiuandStockwell, 2002 (Figure 3ato3g) but differs clearly from this by Valvata Juturnia Cochliopina. (mean, with sd in parenin (mean,with sd O the four extant Of exhibit aswell typical , the new species only , thenewspecies SH = 5.83 mm, SH = 5.83mm, Taylor, 1966(by - - - - s) Specimen UJMC-368b. t) Protoconchview,UJMC-368a. s) SpecimenUJMC-368b.t) Protoconch view.n-p) k-m) Protoconch view. j) UJMC-366a. f) Paratype2,UJMC-364.h-j) and ProbythinellaproteraPilsbry.a,b,g) Figure 3

Juturnia gracilis Probythinella emarginata, new species, new Juturnia coahuilae J. coahuilae Juturnia coahuilae J.gracilis (Taylor, 1966)Hershler, Liuand Stockwellfrom spring Tunel7,Viesca,Coahuila,Mexico, UJMC-367a. p) Protoconch view. q-t) UJMC-367a.p)Protoconchview. q-t) from Poza Churince, Cuatrociénegas, Coahuila, Mexico, a present-day specimen. m) specimen. present-day a Mexico, Churince,Cuatrociénegas,Coahuila, Poza from new species,Holotype,UJMC-362. g)Protoconch view. c,d)Paratype 1,UJMC-363.e, (Taylor, 1966) Hershler,Liu andStockwell, 1966) (Taylor, Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Probythinella protera. Probythinella emarginata q,r)SpecimenUJMC-368a. /2019 (Küster) 615 615

Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS 616 616 Coahuila. population from Spring Tunel7,Viesca, Coahuila.Triangles = gracilis Figure 4 coahuilae gracilis in that0.21–0.29 in is are as well the whorl height/whorl width ratio betweendiagnostic distinctions thesetwo species with mostwhorlssnail NorthAmerica). Reliablein dríguez, 25/vi/2016).Allfrom typelocality. (leg. Alexander CzajaandJoséEstrada-Ro Luis 363, Paratype 2(Figures 3e, 3f),UJMC-364. UJMC-362. Paratype 1 (Figures 3c,3d),UJMC- Type Material: recent andsubfossil species. analyses were efficient in distinguishing among the based discriminant description and morphometric combination between the detailed morphological endemic species from the Cuatrociénegas Basin.A the extant phologically clearly distinctto shells of shells of subfossil the that confirms data, morphometric (Figureanalysis seven 4),basedon standard shell / new species, subfossil population from Spring Tunel 4, Viesca, Coahuila. Square = Square Coahuila. Viesca, 4, subfossilTunel Springpopulation fromspecies, new Graphic depiction ofdepiction thediscriminantanalysisof Graphic variables of morphometric three populations of Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín J. coahuilae. n.sp. hasanarrower than general form (Figure 3ato3f).Also, thediscriminant J. gracilis As a result of this different ratio, different this of result a As n. sp. described herein are mor Holotype (Figures 3a, 3b, 3g), J. gracilis n.sp. and0.30–0.40 J. coahuila J. J. J. J. - - /2019 , apresent-daypopulation from PozaChurince, Cuatrociénegas, hlso shells of Remarks: Túnel 4springat Viesca, Coahuila,Mexico. Distribution: slender (lat. Etymology: the samelocality(UJMC-365a-l). Other MaterialExamined: cene, sub-recent. belowcm thepresent-daysurface). Latest Holo Stratum Typicum: m a.s.l.) Tunel 4 Spring (25° 20′ 20″ N, 102° 52′ 32″ W, 1099 Type Locality: and are reported fromonly extantof members withnumerous shells form whorls (more than7) onia America such as North southwestern cochliopinesepigean of several other and sculptural features with thoseof and Tryonia. Tryonia. Juturnia Hershler Hershler Both genera are difficult to separate to difficult are genera Both gracilis The species wasnamed after the The

J. coahuila J. gracilis However, suchnarrow and turri Mexico, Coahuila state, Viesca, Eremopyrgus, Littoridinops, Pseudotry broadly overlap insize, shape, eea hp fitsshells. ) generalshapeof Springsediments(layer5 c. et al. n.sp. isendemic to the (Taylor, 1966), a subfossil a 1966), (Taylor, (2002)mentioned that Juturnia. 12 specimens from12 specimens Circles= Juturnia Juturnia Juturnia - - -

eeso salinity(Ortega-Guerrero, 2003). levels of once fed the springs withwater, had(andhas)high springs at Viesca sincethe groundwater, which ronments. for can alsobeassumed the former This species of According to Hershler the spring deposits. the vertical section of studyof further requires hypothesis this of confirmation a separatedin (allopatric spring evolution) but a fossil between both species. Wethat suppose the sub undoubtedly anarrowphylogenic relationship structure of the shell proximitygraphical andthe similarity of coahuilae of shells subfossil in Viesca spring-outflows and the Cuatrociénegas Basin.In neighboring springs (Taylor, 1966),anextant endemic speciesfrom genus, the the three living speciesof both genera. Of is the unique useful differentiating feature between the well-developed growth lines on by shells and it seems that conchiologically on corresponds in itsconchologicalcorresponds features inalmost Remarks: Coahuila. the Cuatrociénegas basin, to thespringcomplex of Distribution: 366a-o). Material Examined: WN =6.12(0.21). WBW =1.19(0.09)mm, HBW =1.80(0.13)mm, AWAH =0.95(0.13)mm, mm, =0.76(0.04)mm, theses; N=15)SH3.55(0.34),SW = 1.27(0.10) Shell Measurements: species. this (1985)fromand Hershler the type locality of are identical to thosedescribedbyTaylor (1966) Description: Durangonella coahuilae J. gracilis Juturnia coahuilae Juturnia J. gracilis are very(see common below). geo This Durangonella coahuilae Juturnia J. coahuilae The subfossil The Morphologically, the subfossilshells The present-day The speciesisendemic n. sp.evolved has from n. sp. closelyresembles (Figure 3hto3j) are associated with saline envi Hershler, 1985,81,fig. 32 et al. and (D. W. Taylor, 1966) 15specimens(UJMC (mean, with sd in parenin (mean,with sd J. coahuilae (2002), all four extant J. gracilis Taylor, 1966 Juturnia n. sp. reflect sp. n. from Viesca J. coahuilae J. coahuilae shells Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín ly J. J. - - - - the south. the genus the (fossil) range of from Cuatrociénegas. present The record extends the extant species them within the variation of both populations, andwe consider not surprising the temporal and spatial distance of are, inviewof differences morphological small these However, 4). (Figure analysis discriminant the in reflected the subfossilspecies, which isalso rower of form the subfossiland extant the slightlynar is species (Figures 3h to 3m). The unique difference between all details to the recent species from Cuatrociénegas (Hershler, 1996). Hudson Bay, and Mackenzie River to the north Great Lakes, Riversissippi drainages of basin; and lentic inland habitats throughout the Mis Distribution: 367a-j). Material Examined: (0.30) mm,WN=4.78(0.25). WBW =2.43 HBW =3.05 (0.34)mm, (0.20) mm, AWAH =1.87(0.18)mm, (0.41) mm, =1.45 SW theses; N=10)SH 4.11 (0.63)mm, = 2.75 Shell Measurements: transparent. inner lips thin;shell clear-white, some specimens narrowlyadnate, orslightlyseparated, outer and sometimes weakly above, angled peristome usually sometimes planorboid, depressed; aperture ovate, ing from the suture to the whorl; base firstof whorl strong, regularlyspaced collabral striationsextend growth; teleoconch sculptured with toward endof often tilted; bodywhorl rarely abruptlydescending suture, suture impressed; apex blunt or depressed, well rounded, sometimes slightly shouldered below whorls; convexwhorls4¾–5¼ slightly inflated, to from 3.24–4.69mm,width2.24–3.18mm;with row-conic or sub-cylindrical, height ranging Description: Type species: Probythinella emarginata Genus Shell small, sub-globose tonar Shell small, sub-globose Eastern NorthAmericainlotic Eastern Probythinella Paludina emarginata (Figure 3nto3p) (mean, with sd in parenin (mean,with sd 10 specimens(UJMC Thiele, 1928 (Küster, 1852) Juturnia /2019 Küster, 1852 c. 300 km to c.300km 617 617 - - - - -

Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS 618 618 American (Hershler, 1996).Our North thissmallgroup of 3t), of a synapomorphy their shells (Figures 3p, extremely blunt apex of ica. Bothspeciesare well characterized bythe North Amer part of distributed along the eastern two extant species: Remarks: Type species: the Holocene, alsoabundant ininlandsites. during was, species this that confirm findings our inpresent-dayoccurs onlyincoastal regions but and lentic habitats assympatric species. species occurredduringthe Holocene inlotic That does notapplyto our area, where both appear to becontiguous, but non-overlapping. both recent distributions of Remarks: (Hershler, 1996). Mexico coast of Gulf watersish along the eastern Distribution: 368a-j). Material Examined: (0.10) mm,WN=5.50(0.12). WBW =1.64 HBW =2.13(0.12)mm, (0.07) mm, AWAH =1.16(0.04)mm, (0.10) mm, =0.99 SW theses; N=10)SH 2.69 (0.13)mm, = 1.69 Shell Measurements: the parietalwall,outerlipstrongly thickened. the coilingaxes, innerlipthickenedandadnate to ovate and inclinedapproximately 40° awayfrom apex blunt, depressed; sometimes aperture small, first whorl), ¼ whorlabsent; umbilicus almostplanorbiform; a to (accuracy whorls rounded mm, width1.51–1.82mm;with4¼–4½slightly height rangingfrom 2.54–2.92 to pupiform, Description: c. 300kmtothesouth. recordsspread outthe(fossil)range of / Type species: Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Genus Probythinella protera Hershler (1996) mentioned that Hershler the The genusThe Shell medium insize, ovate-conic original designation) Pseudotryonia Pseudotryonia Endemic to United States, brack Tryonia alamosae Amnicola emarginata (Figures 3qto3t) P. emarginata Probythinella (mean, with sd in parenin (mean,with sd 10 specimens(UJMC Pilsbry, 1953 Hershler, 2001 Probythinella and Taylor, 1987(by Küster, 1852 includesonly P. protera P. emarginata P. protera species , both - - - /2019

(0.08) mm, HBW = 1.64 (0.26) mm, WBW =1.23 HBW =1.64(0.26)mm, (0.08) mm, AWAH =0.98(0.14)mm, (0.22) mm, =0.77 SW theses; N=11)SH 2.21 (0.50)mm, = 1.38 Shell Measurements: in Durango. thisspecies (2011b, p. 16)from the type locality of identical to those described by Hershler Description: pasajae Pseudotryonia pasajae Pseudotryonia this speciesin Durango. Coahuila, NE fromc. 100km the type locality of shells were found inspringElMolino, Viesca, the species. subfossil The of sexual dimorphism the reflect may they that similar,conclude arewe both forms Since all other shell characteristics of differences, which is also observed in our material. ajae (Figure same author mentioned that 6). The they are alsovisible in the discriminant analysis Hershler among the two knownpopulations describedby differences are differences than eventhese smaller the shells, but ulations remain onlyinthe size of pop extant and subfossil the between differences the recent species. Small notypic variation of charactersare the phe within all morphometric Durango, nearthe type locality (Figure 6).Almost from anunnamedspringat San Pedro de Ocuila, theendemicextant species and shellsof from Viesca shells subfossil the among differences morphological significant reveal not did analysis Remarks: 2014a). Nazas basin,Durango, Mexico (Hershler CuencamédeCeniceros, Rio northwestsprings of Distribution: 369a-k). Material Examined: (0.19) mm,WN=4.23(0.54). Pseudotryonia pasajae Pseudotryonia sossxa iopimi omo shellssize of inform showssexual dimorphism figures. 2E–F, I–J, M–O, 3E–H,4D–E. et al. h eut fthe discriminant resultsThe of Morphologically, the shells are

(2014a) from the type locality and P. pasajae (Figure 5ato5d) Hershler Hershler Hershler, LiuandLandye, Hershler Hershler 2011 occurs living only in twoliving only occurs (mean, with sd in parenin (mean,with sd 11 specimens(UJMC et al. et al ., 2014:59–60. , 2011:14,16, P. pasajae P. pas et al., et al. - - - - UJMC-370. k) Protoconch view. l) T. seemani from Durango, syntype (BMNH 20001099) illustrated by Hershler by illustrated 20001099) (BMNH Durango,fromsyntype seemani T. l) Protoconch view. k) UJMC-370. de Ocuila,Durango, Mexico.h) Protoconchview.i-m) Male(?) specimen, UJMC-369b.d)Protoconch view,UJMC-369a.e-h) UJMC-369a. c) Figure 5 Fig. 12c). T. seemani umbilicus absent;periostracumtan. lines; aperture pyriform, parietal lip complete; stronggrowthfine impressed sutures. Sculpture of conch whorls highlyconvex, evenly rounded with height 5.99 mm, with5½ rounded whorls. Teleo Description: Durangonella seemani Hydrobia seemani Type species:

Pseudotryonia pasajae , apresent-day specimen fromanunnamed spring onwest sideofNombre deDios,Durango, Mexico(Hershler Tryonia Genus cf. Tryonia clathrata feld, 1865:525,pl.8. 1945, 19,pl.3,Fig. 1 original designation) Frauenfeld, 1863: 1025.Frauen Shell turriform, medium-sized, Shell turriform, Tryonia (Figure 5ito5k) seemani (Morrison, 1945): Morrison, 1945): Morrison, (Morrison, Hershler, Liuand Landye, 2011and Stimpson,1865 (Frauenfeld, 1863) Stimpson,1865(by T. seemani Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - . i-k) T. Tryonia cf. seemani, spired andstrongly convex-whorled specimen is to thegenus rehydrated material and transferred thisspecies (1945). Hershler the new erected genus later redescribed and placed as type species in seemani from Durango describedoriginallyas Remarks: 2011a). Durango City, DurangoState (Hershler Distribution: Material Examined: = 3.64mm,WBW2.25WN5½. 2.54 mm,AH= 2.09mm,AW = 1.56 mm,HBW Shell Measurements: cf. seemani P. pasajae, subfossil specimen from Viesca,Coahuila, Mexico, by Frauenfeldspecies was (1863). This (Frauenfeld, 1863).a-d) Our shellcloselyresemblesshells Tryonia Endemicto a few springs near apresent-dayspecimenfrom SanPedro et al. (Figure high 5m).Oursingle (2002) finally examined the examined finally (2002) 1 specimen(UJMC-370). Durangonella SH = 5.99 mm, SW = SH=5.99mm, et al. P. pasajae /2019 (2002. Fig. 1b). m) 1b). Fig. (2002. by Morrison et al. , specimen Hydrobia , 2011a, et al., 619 619

Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS 620 620 from Hershler spring atSanPedro deOcuila,Durango. Triangles = pasajae Figure 6 single single specimen was found in superficial sediments the subfossilshell from Viesca onlytentatively. The however, specimen we identify a single sinceitis unlikely, is convergence A conspecific. as sidered et al. (2002)(Figuretherefore 5l)andcan becon the species from Durango illustrated by Hershler of nearidentical tothesyntypes morphologically ¾ ‒ 5 ¾ rounded whorls that increase moderately 4 with discoidal; flattened, mm), 13.25 diameter Description: Biomphalaria “temascalensis” Biomphalaria obstructa theJuan Spring, Guerra Viesca. of / , a subfossil population from Spring Tunel 4, Viesca, Coahuila. Circles = Circles Coahuila. Viesca, 4, Tunel Springsubfossil population from a , Type species: Graphic depiction ofthediscriminant Graphicdepiction analysisofmorphometric variablesof threepopulationsof Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Family PlanorbidaeH.andA.Adams, 1855 et al. Genus , 2011b). Shells medium to large (maximum Biomphalaria Biomphalaria smithii (Figure 7ato7c) 25–34; pl.1. (Morelet 1849): Test. Noviss. I: 16. Preston, 1910 Rangel-Ruiz1987: (Pfeiffer, 1839) Preston, 1910 P. pasajae, - apresent-day population from elTanque, Durango, typelocality(data /2019 in areas alongtheAtlanticandPacific coasts. the genus inMexico andinhabits mainly species of jo-García (2003), 2011). According to Naran mined (Thompson, South American distribution remains to be deter USA, West Indies, Mexico andCentral America. Distribution: 371a-h). Material Examined: diameter) =12.10(0.83)mm,WN5.19(0.35). theses; N= 8) SH= 3.41 (0.20) mm, SW (= shell Shell Measurements: peristome isthin,withoutaperturallamellae. separated by sutures;deep aperture ovate, the growth pronouncedthe bodywhorl, on whorls of lines fine of sculpture flattened; spire diameter; in States andMexi extant species reported frequently from United subfossil shellsfrom Viesca resem are(DeJong longandconfusing the genus the recentof members of Remarks: P. pasajae The taxonomic The andsystematics history , a present-day population from an unnamedpopulation from present-day a , The species occurs in southeastern insoutheastern species occurs The co: B. havanensis B. havanensis 8specimens(UJMC (mean, with sd in parenin (mean,with sd Pseudotryonia isthe most common et al. l hlso two ble shells of and 2001). The 2001).The Biomphalaria B. obstructa. B. obstructa. . Square= P. - - -

havanensis Figure 7 Viesca, Coahuila, Mexico.g,h,j)SpecimenUJMC-373a. i)SpecimenUJMC-373b. UJMC-372a from fluviatile ofthe Aguanaval River,Durango, deposits Mexico. g-j) ke) lcdjs bv h etro the body; (keel), placed just above the center of whorls; body whorl withlesswell-developed carina enlarging few rapidly with spiral), (flat discoidal Description: 2016). havanensis from Oaxaca, Mexico) isajuniorsynonymof obstructa 2011) but recent molecular studies show that (DeJong species different Both species were considered for alongtime as

Type species: Biomphalaria havanensis Biomphalaria , fossil specimen UJMC-371a from a former wetland area of Viesca, Coahuila, Mexico. d-f) Mexico. Coahuila, Viesca, of wetlandarea former a from specimen UJMC-371a fossil , (aswell as (Aguiar-Silva Dilatata dilatata Genus Shell small, ultradextral, flattened, ultradextral, small, Shell (Figure 7dto7f) dilatatus Dilatata B. tamascalensis et al. (Gould,1841) Clessin,1885 (Pfeiffer, 1839), et al., , 2014;Rosser 2001; Thompson, 2001;Thompson, Gould,1841 Rangel-Ruiz Dilatata dilatata et al., Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín B. B. B. B. (Gould,1841)and Distribution: 372a-b). Material Examined: AH =1.11mm,AW =1.31mm,WN2.75. SW (= SH =1.22mm, shell diameter) = 2.68mm, 1.28 mm,AWWN =3.Specimen =1.59mm,2: mm, SW (= shell diameter) = 3.23 mm,AH= Shell Measurements: lips slightlythickened. growth; aperture umbilicusdeep; large, expanded; finerapidly linesincreasing of in size, sculpture of with 3 whorls (accuracyto a ¼ whorl), whorls (Albrecht Burch, 1989). In Mexico inZacatecas and Puebla andTexasFlorida to Canada(Baker, 1945; ent-daydistribution acrossUnited States from netus dilatatus P. exacuous et al., ). subfossil specimensfrom spring elMolino, Promenetus exacuous 2007; Thompson, 2011 as 2007; Thompson,

Dilatata dilat D. dilatata Specimen 1: SH = 1.63 2 specimens(UJMC ata has a broad has pres , subfossilspecimen , (Say, 1821). a-c) B (Say, 1821).a-c) /2019 Microme 621 621 - - .

Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS Freshwater snails – a case of “silent” local extirpation from Mexico RESULTS / DISCUSSION 622 622 eodo thisgenus from Mexico. record of fossil first the is It species. this of material recent inall details to the spond intheir shell morphology Remarks: r ii f curneo living occurrence of limit of ern size alone are insufficient for a distinction. The south shell in differences small that considered we because et al. megas. the Keeled species of Prescott andCurteanu (2004)distinguish two sub ferences, such some authors asClarke (1981)and Although there are noanatomical orradulardif this species. the shell of is the maincharacteristic of Remarks: (including Alaska)but doesnotreach Mexico. distribution acrossCanada and United States Distribution: 373a-j). Material Examined: diameter) = 6.96(0.59)mm,WN= 2.60(0.36). theses; N= 10)SH = 1.91(0.07)mm,SW (= shell Shell Measurements: shell repair scars. lip thin to slightly thickened: some specimens with umbilicus lines; aperturedeep; triangular orovate, outer andinner spiral fine and growth of lines increasing in size; spire fine flattened; sculpture of rapidly whorlswhorls, 2¼‒3½ with whorl; body the keeled); keel close toorabove the center of spira ultradextral, with carinate (sharply periphery (flat discoidal flattened, mm), 8.43 eter Description: Promenetus megas exacuous Promenetus exacuous exacuous sin Geol. Nat. Survey History Bull. 70, pt. 1, p. exacuous / Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (1998) and do not recognize these subspecies (1998)anddonot recognize these subspecies Type species: However, we follow Burch (1989)andTurgeon Molluscan familyPlanorbidae, p. 188. Molluscan familyPlanorbidae, p. 182. Genus Promenetus exacuous The prominentThe keel onthe body whorl The twoThe subfossilcorre specimens Shells medium (maximum diam 361, pi.23,fig. 1-5. P. exacuous (Say, 1821). Baker, 1928,Wiscon (Figure 7gto7j) Promenetus Planorbis exacuous Planorbis exacuous Promenetus: P. e. exacuous (W.H. Dall, 1905): Baker, 10 specimens (UJMC 10specimens (UJMC

(mean, with sd in paren has a broad present-day (Say, 1821): Baker, 1945, Baker, 1935 (Say, 1821) P. exacuous Say, 1821 is Tarrant and . . e P. l), ------/2019 4. Discussion Alabama, United States, where within 50 years 33 in NorthAmericatookplace in CoosaRiver,snails freshwater extinction event modern est single of occurred in Mexico larg and . The thatsuggested similarorgreater extinctionslikely ter gastropods inMexico but Johnson freshwa There are fewdata about extinctionsof from Mexico. genus this of record fossil first the is It south. the to the genus c. 1000 km spread out the (fossil) range of County, Texas (Pratt, 1983), and our present record endemic species in thisareaendemic species apparently depended all (43%) were endemics (Table of 2). Extirpation disappeared from the area during the Holocene, 14 the 28 speciesthat with a wider distribution. Of vulnerable to such extinction events than species rangeswere geographic Snails withsmall more contain onlyfour gastropod species(Table 2). sites are monitored by2009 and since the authors spring LaPeña waters. These containpermanent the Nazas andAguanaval rivers andthe small of dams the of outflows the Mexico.Only in regions regionvery andbelongsto dry the most populated study (Figure area 8). The is now a the area of freshwater gastropodin species the number of of 88% of the loss Statesmeans andCanada.This and threeico United in only (9%) occur species them(50%)stillliveMex in of totally extinct, half (12.5%) are stilllivinghere. Aquarter (25%) is throughfourLaguna Districtonly theHolocenein the 32 speciespresent 2017a, 2017b, 2017c).Of the same area (Czaja eral other subfossilreportedspecies recently from species described in the present study aswell sev inthe areapod extirpation (Table 2) are based on local gastro 20th century. estimated The times of 2014a, 2014b), but then increased rapidly in the intheearlybegan to middleHolocene(Czaja diversity mollusks States.general decline of This local freshwater speciesasinUnited loss of results Mexico from show asimilarrapid northern species went extinct (Johnson et al. , 2014a,2014b, 2015, et al., 2013).Our et al. (2013) et al. - - - - - , described inpresentpaper. Species = boldEndemic species, = * area. the same frompreviouslyreported subfossilspecies well as study present in the described Table 2.List ofgastropodspecies of thestudyareaand estimatedtimepriorof thelocalextirpationorextinctionbased onspecies Assimineidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Cochliopidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Hydrobiidae Lymnaeidae Planorbidae Planorbidae Planorbidae Planorbidae Planorbidae Planorbidae Planorbidae Physidae Family Pyrgulopsis paleoacarinatus* Pyrgulopsis paleominckleyi* Tryonia paleocircumstriata* Cochliopina riograndensisCochliopina Probythinella emarginata Probythinella Biomphalaria havenensis Biomphalaria excentricus Hebetancylus viescaensis* Mexipyrgus Pyrgulopsis manantiali* Pseudotryonia pasajae* Paludiscala thompsoni*Paludiscala Assiminea cienegensis* Assiminea parrasense* Promenetus exacuous Promenetus Probythinella proteraProbythinella californica Tryonia Juturnia coahuilae* Juturnia Phreatoceras taylori Cincinnatia integra Coahuilix landyei* Tryonia hershleri* Juturnia gracilis* Coahuilix hubbsi* duryiPlanorbella anceps pomilia Dilatata dilatata parvus Vitta Galba obrussa Species

cf. cf. usnea seemani Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín Holocene Records Holocene X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 20th Century Century 20th Records X X X X X X X X X X X X X X X X X X X X X X X X ? Occurrence in area in Occurrence /2019 Present-day Present-day X X X X 623 623

Freshwater snails – a case of “silent” local extirpation from Mexico DISCUSSION Freshwater snails – a case of “silent” local extirpation from Mexico DISCUSSION 624 624 the areaof studyduring theHolocene. Figure 8 inflows. Almost all endemic species of Cuatro alsobelongtothese hydrobiidciénegas families. species endemic all Almost inflows. North America,andthey ongroundwater depend most vulnerable freshwater gastropod families in inthe area. families belong to the bers These both currently have mem species) and none of were the Cochliopidae (12) and Hydrobiidae (7 families The most affected by the local extirpation activities impactedtheregion. study before human disappeared from the area of ilia Czaja andEstrada-Rodriguez, 2015, the28disappearedonly species thatIt seems of ranges.very tolerant species with wide geographic them are known as vived. Not surprisingly, all of (Menke, 1830) and Mexico. Only bythinella survive in thisarea. genera The Biomphalaria distribution such as However, alsotaxa with relatively wide recent reported from other sites (Hershler were endemics also extinctionsof of lar patterns cies of ability, which isespecially true for the phreatic spe factors, on, amongothers their limited dispersal / Declineinspeciesrichnessof freshwatergastropodsin Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín (Conrad,1834)and Coahuilix, Paludiscala apparently disappeared completely from n ohseiso and bothspeciesof Physella acuta (Wetherby, 1879), (Rowell, 1863), Promenetus, Dilatata, Cincinnatia, Vitta usnea (Draparnaud, 1805), (Draparnaud, and Probythinella Promenetus Phreatoceras. (Say, 1817) sur (Röding, 1798) Helisoma anceps Tryonia hershleri et al., Physella pom 2014b). and didnot Simi Pro ------/2019 were isestablished still living inthe 20th century the freshwater snails. That most snailsfrom Viesca of extirpation Turtle and this could also be time of the Viesca Mud approximate extinction date of 2013). Rhodin alum the Viesca MudTurtle ( existed around Viesca because of type specimens 1959). In1961,somefreshwater habitat must have but in 1959all were15 springs still active (Blásquez, determine to difficult is drying the of time exact the last century. The of pletely in the second half the Laguna Districtthat driedupcom part of Viesca inthe central 15springsof the complex of environmentalThe especially severe impact was in not onlyendemics but alsorelicts. had awider distribution; hence, they are currently springs. subfossilsingle records The show that they now endemic to very restricted areas, including study living butnear the all areafive of species are and Assiminea cienegensis, J. coahuilae, T. seemani, P. pasajae these aridregions. in affected snails hydrobid especially were there environment changeLaguna District.Also, in Northin time Americacoincide with the western by Hershler ure 9). Similar local extinction events documented atand alsoall15springs Viesca haddriedout(Fig 100 meters. Afewyearslater the lakesdisappeared a result, level the groundwater dropped by almost increasedpumps from12 toalmost3000and,as water the 1960s. Inthisperiod, the number of cially at Viesca) were from depleted to the 1920s resourcesgroundwater inLagunaDistrict(espe water hydrology. Wolfe described how(2013) has two largefurther dams altered the ground tion of production.agricultural In addition, the construc pumping principallyfor groundwater irrigated extensive overexploitation of after decades of aquatic habitatsViesca in occurred of loss The North Americainthe1930’s (Rader the springs.exotic This specieswasintroduced to culata byoccurrence with their common Iverson) were and Vogt, collected (Legler Pyrgulopsis manantiali Pyrgulopsis (Müller, 1774) in the superficial sediments of et al. et al. (2014b) from various sites inarid (2011) reported 1970 as the Kinosternon hirtipes megaceph Kinosternon (Hershler, 1985)are still Melanoides tuber et al., 2003). ------Figure 9 Almeida displacing native2004; populations (Thompson, long been suspected of species has District. This gastropod diversity inthe Laguna for the loss of Melanoides tuberculata It cannot be ruled out that such as Johnson CanadaandtheUnited States published by of endangered freshwater species appear in the list of included inthe IUCNRed Listanddoesalsonot more widespread States (Hershler, 1996).However, to the contrary currently a very limited distribution inUnited study,cene Rivers and lakesinthe area shows of few sites. United States in onlyavery in semi-arid region of (Johnson Canada and their populations are currently stable spread United States distribution and ineastern ginata Although Springdriedup Tunel1(Bilbao),Viesca,Coahuila,Mexico,that inthesecondcentury half (May2018). ofthelast are in Mexico, extirpated they have a wide et al. et al., P. protera, et al., Promenetus exacuous (2013). 2013).However, bothspeciesoccur 2018). Although extirpation and 2018).Although extirpation ainlatespecies common Holo P. emarginata, are at least partly responsible and this species is not species is this Probythinella emar Boletín de la Sociedad Geológica Mexicana Geológica Sociedad la de Boletín - - - 5. Conclusions never exceeds 0.1%. Today, however, specimens of few a only contain sediments superficial the that is unclear. current sampling demonstrates The parasitism or increased competition interspecific study,the area acausalrelationship of such as invasion temporallycoincide and spatially in asdls faquatic habitat inthe studyarea of caused loss activities, pumping, especially groundwater that trociénegas Basin.Unfortunately, the same human gastropodsdiversity in NorthAmerica, the Cua the currently greatest hotspots of from one of ognized only sometwo hundred away kilometers the Laguna District happened silently and unrec event local extirpation The described here from ties that stillcontainwater inthestudy area. now by far the most abundant speciesinthe locali Melanoides and its relative abundance /2019 Melanoides 625 625 is - - -

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Freshwater snails – a case of “silent” local extirpation from Mexico REFERENCES