The Primary Freshwater Crabs of Guatemala (Decapoda: Brachyura: Pseudothelphusidae), with Comments on Their Conservation Status

JOURNAL OF CRUSTACEAN BIOLOGY, 36(6), 776-784, 2016

THE PRIMARY FRESHWATER CRABS OF GUATEMALA (DECAPODA: BRACHYURA: PSEUDOTHELPHUSIDAE), WITH COMMENTS ON THEIR CONSERVATION STATUS

Ingo S. Wehrtmann 1,2,∗, Célio Magalhães 3, and Mónica N. Orozco 4

1 Museo de Zoología, Escuela de Biología, Universidad de Costa Rica, 11501-2060 San José, Costa Rica 2 Unidad de Investigación Pesquera y Acuicultura (UNIP), Centro de Investigación de Ciencias del Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020 Mar y Limnología (CIMAR), Universidad de Costa Rica, 11501-2060 San José, Costa Rica 3 Instituto Nacional de Pesquisas da Amazônia, Av. André Araújo 2936, 69067-375 Manaus, Amazonas, Brazil 4 Centro de Estudios Atitlán, Universidad del Valle de Guatemala, campus Altiplano, Km 137 Caserío Xolbé, Cantón El Tablón, Sololá, Guatemala

ABSTRACT The freshwater crabs of the family Pseudothelphusidae Ortmann, 1893 are restricted to the Neotropics and many species have an elevated risk of extinction. The majority of pseudothelphusids is considered to be data-deficient, which impedes the assessment of their vulnerability levels. More than 50% of pseudothelphusids are endemic to Guatemala, and half of the species are categorized as data-deficient. In order to update and broaden our knowledge of the freshwater crab fauna of Guatemala, several surveys were carried out from 2014 to 2015, combined with the examination of collections of freshwater crabs of Guatemala deposited in several museums. A total of 55 specimens comprising six pseudothelphusid species were collected. The most common species were Potamocarcinus armatus H. Milne Edwards, 1853 (20% of the material identified to the species level), Phrygiopilus ibarrai (Pretzmann, 1978), Raddaus bocourti (A. Milne-Edwards, 1866), and Potamocarcinus magnus Rathbun, 1896 (14.5% each). The collection of the Universidad del Valle de Guatemala consisted of 57 lots with 69 individuals of nine pseudothelphusid species; predominant species were R. bocourti (61.7% of all identified lots) and P. magnus (20.5%). The following species are reported here as new records for Guatemala: Elsalvadoria zurstrasseni (Bott, 1956), Raddaus mertensi (Bott, 1956), and two undescribed species (Phrygiopilus sp., Zilchia sp.). Almost half of the 17 freshwater crab species (including the two undescribed) are either data-deficient or not yet assessed, and 65% (11 species) are endemic to Guatemala. Conservation strategies are urgently needed to preserve the integrity of the sites and habitats occupied by freshwater crabs, along with studies aiming to fill the gap of the data-deficient species. KEY WORDS: biodiversity, Central America, conservation status, faunal surveys, Neotropical region DOI: 10.1163/1937240X-00002478

INTRODUCTION one-fifth of all the world’s brachyurans or true crabs (Cum- Freshwater habitats are among the most species-rich and en- berlidge and Ng, 2009). The study by Cumberlidge et al. dangered ecosystems in the world (Revenga and Kura, 2003; (2009) on the conservation status of freshwater crabs re- Dudgeon et al., 2006; Abell et al., 2008). Although freshwa- vealed that one-sixth of all assessed species have an ele- ter represents an insignificant portion of the world’s water vated risk of extinction; however, given that almost half of and just 0.8% of the earth’s surface, it harbors an extraordi- the species lack information on vulnerability, that assess- nary wealth of species, roughly 7% of all described species ment is an underestimation of the true risk level. According worldwide (Darwall et al., 2008). This species diversity is to Cumberlidge et al. (2009), Trichodactylidae and Pseu- threatened by numerous factors, mainly associated with an- dothelphusidae (both restricted to the Neotropics) have the thropogenic activities such as increasing demand for drink- highest threat. Cumberlidge et al. (2014) assessed the con- ing water, dam construction, river flow modification, and servation status of the freshwater crabs in the Neotropics and pollution (Revenga and Kura, 2003; Dudgeon et al., 2006; concluded that 34% of pseudothelphusids have an elevated Abell et al., 2008; Darwall et al., 2008). As a consequence, risk of extinction, and 56% of the species of this family are the extinction risk for freshwater species is higher than for data-deficient. their terrestrial counterparts (Collen et al., 2014). The primary freshwater crab fauna of Central America Primary, or true, freshwater crabs are heterotreme bra- is fairly well studied, and is composed mainly by repre- chyurans that live in freshwater, semi-terrestrial, or terres- sentatives of Pseudothelphusidae with only a few species trial habitats, and that reproduce by direct development, of Trichodactylidae in Nicaragua and Panama (e.g., Smal- lacking free-living larval stages (Yeo et al., 2008; Cumber- ley, 1970; Pretzmann, 1972, 1978, 1980; Rodriguez, 1982; lidge and Ng, 2009). Primary freshwater brachyuran crabs Magalhães and Türkay, 1996, 2008; Rodriguez and Magal- are represented by more than 1300 species and comprise hães, 2005; Lara et al., 2013; Magalhães et al., 2013). Costa

∗ Corresponding author; e-mail: [email protected]; [email protected]

Rica, with 15 species (Magalhães et al., 2015), and Panama, the studies of H. Milne Edwards (1853) and Rodriguez with 14 species (Cumberlidge et al., 2014), are the Central (2001), and only recently Wehrtmann et al. (2014) clarified American countries with the highest diversity of pseudothel- the species identity of the freshwater crabs caught by local phusids, followed by Guatemala with 12 species (Cumber- fishermen of Lake Atitlán. Practically all published infor- lidge et al., 2014). Endemic pseudothelphusid species are mation concerns taxonomic aspects, and virtually nothing is relatively common in the Central American region, espe- known about the biology and ecology of these species. cially in Panama (12), Guatemala (nine) (Cumberlidge et al., There is an urgent need to update and broaden our knowl- 2014), and Costa Rica (six) (Magalhães et al., 2015). edge of the freshwater crabs of Guatemala, especially con- Guatemala is a fairly small country (108,889 km2)lo- sidering that more than 50% of the Guatemalan pseudothel- cated in northern Central America, bordering with Mex- phusids are endemic to the country and half of which are categorized as “data-deficient” (Cumberlidge et al., 2014). ico, Belize, Honduras, and El Salvador. The country har- Here we report the findings of recent surveys carried out bors a diverse freshwater crab fauna, but the taxonomic

across Guatemala as well as preliminary results of the ex- Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020 status of several species is uncertain because species and amination of collections that contain specimens of freshwa- subspecies have been described and/or synonymized by dif- ter crabs from this country, particularly from a relatively ferent authors (Smalley, 1970; Pretzmann, 1972, 1978, 1980; large collection deposited at the Universidad del Valle de Rodriguez, 1982, 1992). In a recent revision (Cumberlidge Guatemala, Guatemala City. et al., 2014), six genera and 12 species were reported for Guatemala, including three cave-dwelling pseudothel- MATERIALS AND METHODS phusid crabs. Rodriguez (1982) considered an additional ﬁve We visited numerous locations in Guatemala during 2014 (January and species for Guatemala as incertae sedis. Studies of the fresh- July) and 2015 (August) (Fig. 1). All licenses required for research, water crabs of Guatemala were mostly undertaken between collection, transportation, and export of specimens to the Museo de

Fig. 1. Location of sampling stations in Guatemala in 2014 (January and July) and 2015 (August). This ﬁgure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x. 778 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 36, NO. 6, 2016

Zoología at the Universidad de Costa Rica were obtained from the San José, Costa Rica (UCR-MZ), and Field Museum of Natural History, Consejo Nacional de Areas Protegidas de Guatemala (CONAP). Crabs were Chicago (FMNH). One of us (CM) examined most of the material in 12 collected during daytime by lifting rocks directly in or close to streams and of these institutions, whereas information about material housed in six other water bodies. In some cases, local people collaborated by collecting other institutions were obtained online from the Global Biodiversity In- crabs in larger streams, diving at depths of up to 2 m. The collected formation Facility (GBIF portal; http://www.gbif.org). Primary types of all specimens were stored in individual plastic bags with 70% ethyl alcohol. 24 taxa described or recorded from Guatemala are housed in MNHN (12 At each sampling location, we recorded the altitude and coordinates with taxa), USNM (five taxa), Forschungsinstitut und Naturmuseum Sencken- a GPS (Garmin Foretrex 401, Olathe, KS) and photographed (Nikon berg, Frankfurt a.M. (SMF) (three taxa), Naturhistorisches Museum Wien, Coolpix P7000) the sampling sites where the crabs were obtained. Voucher Vienna (NHMW) (two taxa), and The Natural History Museum, London specimens were deposited in the Museo de Zoología, Universidad de (NHM) and Naturhistorisches Museum Basel, Basel (NHMB) (one taxon Costa Rica, San José (UCR-MZ) and Instituto Nacional de Pesquisas da each). Amazônia (INPA), Manaus, Brazil. We followed the classification scheme for pseudothelphusids as indi- A considerable number of unidentified freshwater crabs from Guatemala cated in Villalobos Hiriart and Álvarez (2008), and specimens were identi- was studied in the zoological collection of the Universidad del Valle de fied following the descriptions and keys in Pretzmann (1980) and Rodriguez Guatemala (UVG), Guatemala City. This collection contained 57 lots of (1982), complemented by notes on morphology and illustrations obtained pseudothelphusid crabs collected throughout Guatemala between 1975 and by one of us (CM) after examining the type specimens of most Guatemalan Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020 2000 (Fig. 2), making it the largest assemblage of Guatemalan freshwater taxa. crabs currently preserved for scientific studies (Table 1). Specimens were directly identified in the facilities of the UVG, but some individuals were RESULTS taken to INPA for further identification. We also examined material from Guatemala deposited in 18 institutions Species Diversity in North and Central America and Europe (Table 1). The largest holdings are kept in the United States National Museum of Natural History, Smith- We collected a total of 55 freshwater crabs comprising six sonian Institution, Washington, DC (USNM), Muséum national d’Histoire pseudothelphusid species during the 2014 and 2015 sur- naturelle, Paris (MNHN), Museo de Zoología, Universidad de Costa Rica, veys: one in tribe Hypolobocerini and five in tribe Potamo-

Fig. 2. Sampling location of the 57 lots with pseudothelphusid crabs found deposited in the collections of the Universidad del Valle de Guatemala, Guatemala City. This ﬁgure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/ content/journals/1937240x. WEHRTMANN ET AL.: FRESHWATER CRABS OF GUATEMALA 779

Table 1. Number and identification status of the lots with primary fresh- 1970; Delamare Deboutteville, 1976; Rodriguez, 1982; Ro- water crabs from Guatemala (all belonging to Pseudothelphusidae) de- driguez and Hobbs, 1989; Rodriguez, 2001) and examina- posited in collections. AMNH, American Museum of Natural History, New tion of collections (Table 1) revealed that a total of 27 taxa of York; CNCR, Colección Nacional de Crustáceos, Universidad Nacional Autónoma de México, Mexico City; FLMNH, Florida Museum of Nat- freshwater crabs, all belonging to Pseudothelphusidae, have ural History, University of Florida, Gainesville; FMNH, Field Museum so far been described or recorded from Guatemala. Only 14 of Natural History, Chicago; INPA, Instituto Nacional de Pesquisas da species and one subspecies, however, are currently recog- Amazônia, Manaus; MCZ, Museum of Comparative Zoology, Harvard Uni- nized as valid taxa, whereas five taxa are considered syn- versity, Cambridge; MNHN, Muséum national d’Histoire naturelle, Paris; NHM, The Natural History Museum, London; NHMB, Naturhistorisches onyms, five are incertae sedis, and two remain undescribed. Museum Basel, Switzerland; NHMW, Naturhistorisches Museum Wien, Vienna; SMF, Forschungsinstitut und Naturmuseum Senckenberg, Frank- SYSTEMATICS furt a.M.; SNSD, Senckenberg Naturhistorische Sammlungen Dresden; TMNH, Tulane Museum of Natural History, Tulane University, New Or- Pseudothelphusidae Ortmann, 1893 leans; UCR-MZ, Museo de Zoología, Escuela de Biología, Universidad de Tribe Hypolobocerini Pretzmann, 1971 Costa Rica, San José; USNM, United States National Museum of Natural Elsalvadoria tomhaasi Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020 History, Smithsonian Institution, Washington, DC; UVG, Universidad del Bott, 1970 Valle de Guatemala, Guatemala City; YPM, Yale Peabody Museum of Nat- Distribution.—Guatemala: Guatemala (this study) and ural History, New Haven; ZMH, Zoologisches Museum, Universität Ham- Sacatepéquez (Bott, 1970) departments. burg. ∗Data from direct examination; ∗∗Data from the literature or Global Biodiversity Information Facility’s Portal (www.gbif.org). Elsalvadoria zurstrasseni (Bott, 1956) Collection Number Identified Not identified Distribution.—Guatemala: Escuintla, Guatemala, and Su- of lots to species (juveniles, chitepéquez (this study) departments; El Salvador (Bott, level females, or 1956; Rodriguez, 1982). damaged) Remarks.—New record for Guatemala (Fig. 3). AMNH∗ 72 5 CNCR∗∗ 11 ∗ Tribe Potamocarcinini Ortmann, 1897 FLMNH 11Phrygiopilus acanthophallus Smalley, 1970 FMNH∗ 11 9 2 INPA∗ 77 Distribution.—Guatemala: Alta Verapaz (Smalley, 1970; MCZ∗∗ 22 Rodriguez, 1982) and Zacapa (this study) departments. MNHN∗ 14 14 NHM∗∗ 21 1 Phrygiopilus chuacusensis Smalley, 1970 ∗∗ NHMB 11 Distribution.—Guatemala: Baja Verapaz (Smalley, 1970) NHMW∗ 22 ∗ and Zacapa (this study) departments. SMF 33 ∗∗ SNSD 22Phrygiopilus ibarrai (Pretzmann, 1978) TMNH∗ 65 1 UCR-MZ∗ 19 17 1 Distribution.—Guatemala: Baja Verapaz (this study) and USNM∗ 38 30 8 Guatemala (Pretzmann, 1978; Rodriguez, 2001) depart- ∗ UVG 57 42 15 ments. YPM∗∗ 33 ZMH∗ 11Phrygiopilus strengerae (Pretzmann, 1965) Total 177 139 37 Distribution.—Guatemala: Alta Verapaz department (Pretz- mann, 1965). carcinini (Table 2). The most common species were Potamo- Remarks.—Pretzmann (1965) stated that the type locality carcinus armatus, Phrygyopilus ibarrai, Raddaus bocourti, is “Trace Aguas, Caco” in Guatemala department, but and Potamocarcinus magnus representing 20.0%, 14.5%, according to Reddel (1981: 317), Cacao, Trece Aguas is 14.5%, and 14.5% of the material identified to species level, probably in the Seamay-Sejul karstik system, Alta Verapaz respectively. The remaining species, Raddaus tuberculatus department. (11.0%) and Elsalvadoria zurstrasseni (7.3%), were less fre- quent. Nearly one fifth (18.2%) of the collection consisted Phrygiopilus n. sp. of females and juveniles that could not be identified even to Distribution.—Guatemala: Escuintla (this study) depart- generic level. The 57 lots of the UVG collection contained ment. a total of 69 individuals of Pseudothelphusidae; 44 of these lots held specimens that could be identified at least to genus. Potamocarcinus armatus H. Milne Edwards, 1853 This material was comprised by a total of nine pseudothel- Potamocarcinus (Potamocarcinus) nicaraguensis aequispinosus Pretzmann, phusid species, two of Hypolobocerini and seven species of 1971: 20. Potamocarcinini (Table 2). The predominant species were R. Distribution.—Mexico (Pretzmann, 1972); Guatemala: Alta bocourti (61.7% of all identified lots) and Potamocarcinus Verapaz (this study) and Petén (Pretzmann, 1971, 1980; this magnus (20.5%). study) departments. A revision of the literature (H. Milne Edwards, 1853; A. Milne-Edwards, 1866; Rathbun, 1896, 1897, 1905; Bott, Remarks.—Pretzmann (1971) only mentioned the country 1956, 1970; Pretzmann, 1965, 1971, 1978, 1980; Smalley, as the type locality of P. (P. ) nicaraguensis aequispinosus. 780 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 36, NO. 6, 2016 ational d’Histoire naturelle, Paris; NHMW, Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020 ory, Tulane University, New Orleans; UCR-MZ, n, DC; UVG, Universidad del Valle de Guatemala, This study This study Demalare Deboutteville (1976) Demalare Deboutteville (1976) Rodriguez and Hobbs (1989) This study (1982), this study (1982), this study This study ording to Cumberlidge et al., 2009) and number of lots with ted during our recent surveys. AMNH, American Museum of

Guatemala species Reported from caves Reported from caves caves species Guatemala

UVG

1 Undescribed 2 Undescribed

USNM

1 Pretzmann (1965) 1 New record for

UCR-MZ

TMNH

1 2 1 Smalley (1970), this study

SMF

1 1 Bott (1970), this study NHMW

1 2 This study

MNHN ∗ ∗

1 2 Reported from 1

INPA

FMNH

1 1 3 Smalley (1970), Rodriguez (1982), this study AMNH 1 1 1 1 New record for 22 14 2 1 6 3 5 1 1 2 2 4 6 Pretzmann (1978), this study Pretzmann Rathbun (1971, (1905), 1980), Pretzmann this (1980), study Rodriguez 4 2 1 8 6 1 1 9 27 Rathbun (1905), Pretzmann (1980), Rodriguez 2 2 2 Rathbun (1897), Rodriguez (1982), this study surveys Present (Bott, Smalley, Rodriguez sp. Demalare Species Collections Observations References Demalare sp. Elsalvadoria zurstrasseni (Bott, 1956) Elsalvadoria tomhaasi Bott, 1970 1970 Phrygiopilus acanthophallus Phrygiopilus chuacusensis Smalley, 1970 Phrygiopilus ibarrai (Pretzmann, 1978) Phrygiopilus strengerae (Pretzmann, 1965) Phrygiopilus Potamocarcinus armatus H. Milne Edwards, 1853 Potamocarcinus magnus Rathbun, 1896 Typhlopseudothelphusa juberthiei Zilchia falcata and Hobbs, 1989 Zilchia zilchi germani Pretzmann, 1978 Zilchia Deboutteville, 1976 Raddaus bocourti (A. Milne-Edwards, 1866) Raddaus mertensi Raddaus tuberculatus (Rathbun, 1897) Typhlopseudothelphusa mitchelli Deboutteville, 1976 1956) V Table 2. Updated list of the currently valid species of primary freshwater crabs (Pseudothelphusidae) from Guatemala with conservation status (acc Conservation status DD identiﬁed specimens deposited in collectionsNatural examined History, (except New where York; FMNH, marked Field by an Museum asterisk), of including NaturalMuseo historical History, de records Chicago; Zoología, as INPA, Universidad well Instituto de as Nacional Costa species de collec Rica, Pesquisas San da José; Amazônia, USNM, Manaus; United MNHN, States Muséum n National Museum of Natural History, Smithsonian Institution, Washingto Naturhistorisches Museum Wien, Vienna; SMF, Forschungsinstitut und NaturmuseumGuatemala Senckenberg, City. Conservation Frankfurt categories: a.M.; DD, TMNH, Data-Deﬁcient; LC, Tulane Least Museum Concern; of NA, Natural Not Assessed; Hist V, Vulnerable. V NA DD DD LC LC LC DD NA LC NA DD LC WEHRTMANN ET AL.: FRESHWATER CRABS OF GUATEMALA 781 Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020

Fig. 3. Sampling locations in Guatemala of Elsalvadoria zurstrasseni (Bott, 1956), Raddaus mertensi (Bott, 1956), and the two yet undescribed species, Phrygiopilus sp. (collection site in the original label only indicates as “Department Escuintia”) and Zilchia sp. This ﬁgure is published in colour in the online edition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.

Potamocarcinus magnus (Rathbun, 1896) Verapaz (this study), Chiquimula (Pretzmann, 1980), Chi- Potamocarcinus guatemalensis Rathbun, 1905: 313, text-fig. 101, pl. 14, maltenango (this study), Escuintla (this study), Guatemala fig. 7. (Pretzmann, 1980; Rodriguez, 2001; this study), Izabal (this study), Petén (Pretzmann, 1980; this study), Quiché (this Distribution.—Mexico (Rodriguez, 1982); Guatemala: Alta study), Santa Rosa (this study), Sololá (Rodriguez, 2001; Verapaz (Rathbun, 1905; this study), Escuintla (Rodriguez, Wehrtmann et al., 2014), Suchitepéquez (this study), To- 1982), Guatemala (Rathbun, 1905; Pretzmann, 1980; this tonicapán (Rathbun, 1905), and Zacapa (this study) de- study), Izabal (Pretzmann, 1980; this study), Jalapa (this partments; Honduras (Pretzmann, 1980); El Salvador (Ro- study), Péten (this study), Sololá (Pretzmann, 1980; Wehrt- driguez, 1982). mann et al., 2014), Suchitepéquez (this study), and Zacapa (Pretzmann, 1980; this study) departments; Honduras (Pretz- Raddaus mertensi (Bott, 1956) mann, 1980); El Salvador (Rodriguez, 1982); Costa Rica (Magalhães et al., 2015). Distribution.—Guatemala: Chiquimula department (this study); El Salvador (Bott, 1956). Raddaus bocourti (A. Milne-Edwards, 1866) Pseudothelphusa similis Rathbun, 1905: 295, text-fig. 87, pl. 15, fig. 4. Remark.—New record from Guatemala (Fig. 3). Pseudothelphusa cobanensis Rathbun, 1905: 296, text-fig. 88, pl. 15, fig. 5. Pseudothelphusa grallator Rathbun, 1905: 297, text-fig. 89, pl. 15, fig. 6. Raddaus tuberculatus (Rathbun, 1897) Distribution.—Mexico (Rodriguez, 1982); Belize (Pretz- Distribution.—Mexico (Pretzmann, 1972); Guatemala: San mann, 1980; Rodriguez, 1982); Guatemala: Alta Verapaz Marcos (this study), Sololá (Rathbun, 1897, 1905), and (Rathbun, 1905; Pretzmann, 1980; Rodriguez, 1982), Baja Suchitepéquez (this study) departments. 782 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 36, NO. 6, 2016

Typhlopseudothelphusa juberthiei Delamare Deboutteville, on somatic characters of the carapace, and several of the 1976 taxa were described based on female specimens (H. Milne Distribution.—Guatemala: Alta Verapaz department (Dela- Edwards, 1853; A. Milne-Edwards, 1866; Rathbun, 1896, mare Deboutteville, 1976). 1897, 1905). Many of the types were preserved as dried specimens. Almost all of this material described between Typhlopseudothelphusa mitchelli Delamare Deboutteville, 1853 and 1905 is deposited at the MNHN (types of nine 1976 species), except for the specimens of two species found at th Distribution.—Guatemala: Alta Verapaz department (Dela- USNM and one at NHMB. From the mid-20 century on, mare Deboutteville, 1976). the examination of characters of the male first gonopod be- came essential for a reliable identification of the taxa, and Zilchia falcata Rodriguez and Hobbs, 1989 several taxonomists (Bott, 1956, 1970; Pretzmann, 1965, Distribution.—Guatemala: Alta Verapaz department (Ro- 1978; Smalley, 1970; Delamare Deboutteville, 1976; Ro- driguez and Hobbs, 1989). driguez and Hobbs, 1989) relied on this structure to diagnose Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020 species. The primary types of the Guatemala species de- Zilchia zilchi germani Pretzmann, 1978 scribed after 1956 are housed in MNHN (three types), SMF Distribution.—Guatemala: Guatemala department (this (three types), USNM (three types), NHMW (two types), study). and NHMB (one type). In addition to these type specimens, few non-type material, usually consisting of single males or Remarks.—Pretzmann (1978: 3; 1980: 658) stated that the few specimens, were available for study (Tables 1 and 2). type locality of this subspecies is “Rio Guyamas, NE El This situation, as well as the uncertain taxonomic status of Progresso, Honduras.” This is an error, and the actual type some taxa, have so far prevented a more accurate assess- locality is “Guatemala, Department of Guatemala, 14 km ment of the diversity and distribution of the freshwater crabs south of Guatemala City, near San José Pinula.” This of Guatemala. Wehrtmann et al. (2014), for instance, com- information is given in a letter dated 9 April 1976 by Ivo mented on the implications of managing the artisanal crab Poglayen-Newall to G. Pretzmann, filed in the crustacean fishery in Lake Atitlán as a single-species fishery (Potamo- section of NHMW (CM). carcinus guatemalensis = P. magnus), when it actually in- Zilchia n. sp. volved two species: Raddaus bocourti and P. magnus.Ro- driguez (1982) listed five species as incertae sedis because Distribution.—Guatemala: Escuintla department (this they were described from female specimens, which makes study). it virtually impossible to ascertain their specific identity be- Incertae Sedis cause the current classification relies almost exclusively on Pseudothelphusa bisunturalis Rathbun, 1897 characters of the male first gonopod. Such limitations affected the recent evaluation for con- Distribution.—Guatemala: Suchitepéquez department, San servation assessment purposes of the freshwater crabs of Agustín stream, near Atitlán volcano, Pacific slope (Rath- Guatemala by Cumberlidge et al. (2014). They reported 12 bun, 1897). species, whereas here we recognized 17 species from six Boscia gracilipes A. Milne-Edwards, 1866 genera. In spite of the uncertainties, Guatemala has a diverse freshwater crab fauna, comparable to that of Costa Distribution.—Guatemala: Alta Verapaz department, exact Rica and Panama. Magalhães et al. (2013) mentioned six locality unknown (A. Milne-Edwards, 1866). genera and 17 species for Panama and four genera and 15 Gordonia longipes Pretzmann, 1965 species for Costa Rica (Magalhães et al., 2015). Our knowledge of the freshwater crabs from El Salvador, Honduras, Distribution.—Guatemala: Alta Verapaz department, Lan- and Nicaragua has not been updated since the early 1980s quín (Pretzmann, 1965). (Pretzmann, 1980; Rodriguez, 1982). All the collected sam- Pseudothelphusa propinqua Rathbun, 1905 ples are kept in European or North American institutions, and it is not known if institutions in these three countries Distribution.—Guatemala: Alta Verapaz department, Cobán maintain collections of freshwater crabs. (Rathbun, 1905). Our current knowledge of the conservation status of the Pseudothelphusa proxima Rathbun, 1905 freshwater crabs of Guatemala is still scarce: 47% of the 17 species (including two yet undescribed species) are consid- Distribution.—Guatemala: Alta Verapaz department, Cobán ered as data-deficient or have not been yet evaluated (Ta- (Rathbun, 1905). ble 2). Two species (Elsalvadoria zurstrasseni and Phry- giopilus acanthophallus) are classified as vulnerable accord- DISCUSSION ing to IUCN categories and criteria (IUCN, 2003) (Table 2), The current taxonomic knowledge on the freshwater crab and 65% (11 species, including the two undescribed species) fauna of Guatemala is based on relatively few specimens are endemic to Guatemala, a high level of endemism for that have been sporadically collected since the 19th cen- a continental country (Cumberlidge et al., 2009). Further- tury and deposited in collections of European and North more, three cave-dwelling species inhabit Guatemala, and American institutions. Following the paradigm of the pe- these species are especially vulnerable to disturbance and riod, the first descriptions of species were based mainly pollution (Ng and Yeo, 2007; Cumberlidge et al., 2009). WEHRTMANN ET AL.: FRESHWATER CRABS OF GUATEMALA 783

This setting, together with the alarmingly poor quality of Collen, B., F. Whitton, E. E. Dyer, J. E. M. Baillie, N. Cumberlidge, W. R. T. fresh water (Elmore et al., 2005; Padilla Cámbara et al., Darwall, C. Pollock, N. I. Richman, A.-M. Soulsby, and M. Böhm. 2014. 2010), represents a serious threat to the crab fauna of the Global patterns of freshwater species diversity, threat and endemism. Global Ecology and Biogeography 23: 40-51. country. Other threats related to anthropogenic habitat dis- Cumberlidge, N., and P. K. L. Ng. 2009. Systematics, evolution, and turbances are imminent as well: the increasing demographic biogeography of freshwater crabs, pp. 491-508. In, J. W. Martin, K. A. pressure (Castillo et al., 1997; IARNA, 2003; MARN, 2010; Crandall, and D. L. Felder (eds.), Decapod Crustacean Phylogenetics. Rejmánková et al., 2011), increasing and accelerating de- CRC Press, Boca Raton, FL. , F. Alvarez, and J. L. Villalobos. 2014. Results of the global conser- terioration of the environment, risk of pollution (including vation assessment of the freshwater crabs (Brachyura, Pseudothelphusi- intensive use of pesticides and chemical fertilizers), and the dae and Trichodactylidae): the Neotropical region, with an update on di- increasing levels of waste, are some of the serious threats versity. ZooKeys 457: 133-157. (CEMAT, 1999; Elmore et al., 2005; Padilla Cámbara et , P. K. L. Ng, D. C. J. Yeo, C. Magalhães, M. R. Campos, F. Alvarez, al., 2010). Considering that even widely distributed species T. Naruse, S. R. Daniels, L. J. Esser, F. Y. K. Attipoe, F. L. Clotilde-Ba, W. Darwall, A. McIvor, M. Ram, and B. Collen. 2009. Freshwater crabs with an apparent tolerance to land-use changes have declined and the biodiversity crisis: importance, threats, status, and conservation Downloaded from https://academic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 May 2020 drastically in abundance as a consequence of changes in land challenges. Biological Conservation 142: 1665-1673. use, hydrology, or pesticide regimes (Cumberlidge et al., Darwall, W., K. Smith, D. Allen, M. Seddon, G. McGregor Red, V. Claus- 2009), conservation managers of governmental as well as nitzer, and V. Kalkman. 2008. Freshwater biodiversity – a hidden re- source under threat, pp. 1-11. In, J.-C. Vié, C. Hilton-Taylor, and S. N. non-governmental organizations need to recognize that this Stuart (eds.), The 2008 Review of the IUCN Red List of Threatened freshwater crab fauna is seriously threatened. Conservation Species. IUCN, Gland. strategies are urgently needed to preserve the integrity of Delamare Deboutteville, C. 1976. Intérêt biologique et écologique des the habitats along with studies to fill gaps in data-deficient Crabes cavernicoles du Guatemala et du Mexique appartenant au species (Cumberlidge et al., 2009, 2014). genre Typhlopseudothelphusa Rioja. Comptes rendus hebdomadaires des Séances de l’Académie des Sciences (Paris) (Série D) 283: 837-840. Dudgeon, D., A. H. Arthington, M. O. Gessner, Z.-I. Kawabata, D. J. ACKNOWLEDGEMENTS Knowler, C. Leveque, R. J. Naiman, A.-H. Prieur-Richard, D. Soto, M. L. Stiassny, and C. A. Sullivan. 2006. Freshwater biodiversity: importance, We thank the Consejo Nacional de Ciencia y Tecnología, Costa Rica threats, status and conservation challenges. Biological Reviews 81: 163- (CONICIT; project IQ-0001-11) and the Conselho Nacional de Desenvolvi- 182. mento Científico e Tecnologico (CNPq; Brazil, projects 490314/2011-2) for Elmore, A. C., G. R. Miller, and B. Parker. 2005. Water quality in Lemoa, their financial support to ISW and CM. CM also thanks CNPq for an on- Guatemala. Environmental Geology 48: 901-907. going research grant (304736/2015-5). Additional support for ISW was re- IARNA (Instituto de Agricultura, Recursos Naturales y Agrícolas, Univer- ceived from Universidad de Costa Rica (project VI 808-B3-504). We are sidad Rafael Landívar, Guatemala). 2003. Estado actual de los recursos especially grateful to Octavio Cajas (Consejo Nacional de Areas Protegi- hídricos de Guatemala. Informe Técnico No. 1, available online at http:// das de Guatemala) for his support with the collection and export permits, biblio3.url.edu.gt/IARNA/serie_amb/4.PDF. and to Raquel Romero Chaves (Unidad de Investigación Pesquera y Acui- IUCN (International Union for Conservation of Nature). 2003. Guideline cultura, Universidad de Costa Rica) for the preparation of the distribution for Application of IUCN Red List Criteria at Regional Levels: Version maps. We would like to express our gratitude to Margarita Palmieri and 3.0. IUCN Species Survival Commission. IUCN, Gland. Gabriela Palomo (Biology Department, Universidad del Valle de Guatemala Lara, L. R., I. S. Wehrtmann, C. Magalhães, and F. L. Mantelatto. (UVG)), who facilitated the study of the UVG collection, and to Stefania 2013. Species diversity and distribution of freshwater crabs (Decapoda: Rodríguez and Christine LeBeau (AMNH), Jochen Gerber (FMNH), Laure Pseudothelphusidae) inhabiting the basin of the río Grande de Térraba, Corbari, Paula Martin-Lefrève and Sébastien Soubzmaigne (MNHN), Pe- Pacific slope of Costa Rica. Latin American Journal of Aquatic Research ter Dworschak (NHMW), the late Michael Türkay (SMF), Hank Bart Jr., 41: 685-695. and Nelson Rios (TMNH), and Rafael Lemaitre and Karen Reed (USNM) Magalhães, C., and M. Türkay. 1996. Taxonomy of the Neotropical fresh- for their assistance during the visits of CM to their respective institutions. water crab family Trichodactylidae II. 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