the primary freshwater crabs of guatemala …

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 Rica2 Unidad de Investigación Pesquera y Acuicultura (UNIP), Centro de Investigación de Ciencias del

Mar y Limnología (CIMAR), Universidad de Costa Rica, 11501-2060 San José, Costa Rica3 Instituto Nacional de Pesquisas da Amazônia, Av. André Araújo 2936, 69067-375 Manaus, Amazonas, Brazil4 Centro de Estudios Atitlán, Universidad del Valle de Guatemala, campus Altiplano, Km 137 Caserío Xolbé,

Cantón El Tablón, Sololá, Guatemala

A B S T R A C T

The freshwater crabs of the family Pseudothelphusidae Ortmann, 1893 are restricted to the Neotropics and many species have an elevatedrisk of extinction. The majority of pseudothelphusids is considered to be data-deficient, which impedes the assessment of their vulnerabilitylevels. More than 50% of pseudothelphusids are endemic to Guatemala, and half of the species are categorized as data-deficient. In orderto 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 specimenscomprising 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 of57 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), Raddausmertensi (Bott, 1956), and two undescribed species (Phrygiopilus sp., Zilchia sp.). Almost half of the 17 freshwater crab species (includingthe two undescribed) are either data-deficient or not yet assessed, and 65% (11 species) are endemic to Guatemala. Conservation strategiesare urgently needed to preserve the integrity of the sites and habitats occupied by freshwater crabs, along with studies aiming to fill the gapof the data-deficient species.

KEY WORDS: biodiversity, Central America, conservation status, faunal surveys, Neotropical region

DOI: 10.1163/1937240X-00002478

INTRODUCTION

Freshwater habitats are among the most species-rich and en-dangered ecosystems in the world (Revenga and Kura, 2003;Dudgeon et al., 2006; Abell et al., 2008). Although freshwa-ter represents an insignificant portion of the world’s waterand just 0.8% of the earth’s surface, it harbors an extraordi-nary wealth of species, roughly 7% of all described speciesworldwide (Darwall et al., 2008). This species diversity isthreatened by numerous factors, mainly associated with an-thropogenic activities such as increasing demand for drink-ing water, dam construction, river flow modification, andpollution (Revenga and Kura, 2003; Dudgeon et al., 2006;Abell et al., 2008; Darwall et al., 2008). As a consequence,the extinction risk for freshwater species is higher than fortheir terrestrial counterparts (Collen et al., 2014).

Primary, or true, freshwater crabs are heterotreme bra-chyurans that live in freshwater, semi-terrestrial, or terres-trial habitats, and that reproduce by direct development,lacking free-living larval stages (Yeo et al., 2008; Cumber-lidge and Ng, 2009). Primary freshwater brachyuran crabsare represented by more than 1300 species and comprise

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

one-fifth of all the world’s brachyurans or true crabs (Cum-berlidge and Ng, 2009). The study by Cumberlidge et al.(2009) on the conservation status of freshwater crabs re-vealed that one-sixth of all assessed species have an ele-vated risk of extinction; however, given that almost half ofthe species lack information on vulnerability, that assess-ment is an underestimation of the true risk level. Accordingto Cumberlidge et al. (2009), Trichodactylidae and Pseu-dothelphusidae (both restricted to the Neotropics) have thehighest threat. Cumberlidge et al. (2014) assessed the con-servation status of the freshwater crabs in the Neotropics andconcluded that 34% of pseudothelphusids have an elevatedrisk of extinction, and 56% of the species of this family aredata-deficient.

The primary freshwater crab fauna of Central Americais fairly well studied, and is composed mainly by repre-sentatives of Pseudothelphusidae with only a few speciesof Trichodactylidae in Nicaragua and Panama (e.g., Smal-ley, 1970; Pretzmann, 1972, 1978, 1980; Rodriguez, 1982;Magalhães and Türkay, 1996, 2008; Rodriguez and Magal-hães, 2005; Lara et al., 2013; Magalhães et al., 2013). Costa

© The Crustacean Society, 2016. Published by Brill NV, Leiden DOI:10.1163/1937240X-00002478

Dow

nloaded from https://academ

ic.oup.com/jcb/article-abstract/36/6/776/2735695 by guest on 19 M

ay 2020

http://dx.doi.org/10.1163/1937240X-00002478

mailto:[email protected]

mailto:[email protected]

http://dx.doi.org/10.1163/1937240X-00002478

WEHRTMANN ET AL.: FRESHWATER CRABS OF GUATEMALA 777

Rica, with 15 species (Magalhães et al., 2015), and Panama,with 14 species (Cumberlidge et al., 2014), are the CentralAmerican countries with the highest diversity of pseudothel-phusids, followed by Guatemala with 12 species (Cumber-lidge et al., 2014). Endemic pseudothelphusid species arerelatively common in the Central American region, espe-cially in Panama (12), Guatemala (nine) (Cumberlidge et al.,2014), and Costa Rica (six) (Magalhães et al., 2015).

Guatemala is a fairly small country (108,889 km2) lo-cated in northern Central America, bordering with Mex-ico, Belize, Honduras, and El Salvador. The country har-bors a diverse freshwater crab fauna, but the taxonomicstatus of several species is uncertain because species andsubspecies have been described and/or synonymized by dif-ferent authors (Smalley, 1970; Pretzmann, 1972, 1978, 1980;Rodriguez, 1982, 1992). In a recent revision (Cumberlidgeet al., 2014), six genera and 12 species were reportedfor Guatemala, including three cave-dwelling pseudothel-phusid crabs. Rodriguez (1982) considered an additional fivespecies for Guatemala as incertae sedis. Studies of the fresh-water crabs of Guatemala were mostly undertaken between

the studies of H. Milne Edwards (1853) and Rodriguez(2001), and only recently Wehrtmann et al. (2014) clarifiedthe species identity of the freshwater crabs caught by localfishermen of Lake Atitlán. Practically all published infor-mation concerns taxonomic aspects, and virtually nothing isknown about the biology and ecology of these species.

There is an urgent need to update and broaden our knowl-edge of the freshwater crabs of Guatemala, especially con-sidering that more than 50% of the Guatemalan pseudothel-phusids are endemic to the country and half of which arecategorized as “data-deficient” (Cumberlidge et al., 2014).Here we report the findings of recent surveys carried outacross Guatemala as well as preliminary results of the ex-amination of collections that contain specimens of freshwa-ter crabs from this country, particularly from a relativelylarge collection deposited at the Universidad del Valle deGuatemala, Guatemala City.

MATERIALS AND METHODS

We visited numerous locations in Guatemala during 2014 (January andJuly) and 2015 (August) (Fig. 1). All licenses required for research,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 editionof this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.

Dow



ay 2020

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 theConsejo Nacional de Areas Protegidas de Guatemala (CONAP). Crabs werecollected during daytime by lifting rocks directly in or close to streams andother water bodies. In some cases, local people collaborated by collectingcrabs in larger streams, diving at depths of up to 2 m. The collectedspecimens were stored in individual plastic bags with 70% ethyl alcohol.At each sampling location, we recorded the altitude and coordinates witha GPS (Garmin Foretrex 401, Olathe, KS) and photographed (NikonCoolpix P7000) the sampling sites where the crabs were obtained. Voucherspecimens were deposited in the Museo de Zoología, Universidad deCosta Rica, San José (UCR-MZ) and Instituto Nacional de Pesquisas daAmazônia (INPA), Manaus, Brazil.

A considerable number of unidentified freshwater crabs from Guatemalawas studied in the zoological collection of the Universidad del Valle deGuatemala (UVG), Guatemala City. This collection contained 57 lots ofpseudothelphusid crabs collected throughout Guatemala between 1975 and2000 (Fig. 2), making it the largest assemblage of Guatemalan freshwatercrabs currently preserved for scientific studies (Table 1). Specimens weredirectly identified in the facilities of the UVG, but some individuals weretaken to INPA for further identification.

We also examined material from Guatemala deposited in 18 institutionsin North and Central America and Europe (Table 1). The largest holdingsare kept in the United States National Museum of Natural History, Smith-sonian Institution, Washington, DC (USNM), Muséum national d’Histoirenaturelle, Paris (MNHN), Museo de Zoología, Universidad de Costa Rica,

San José, Costa Rica (UCR-MZ), and Field Museum of Natural History,Chicago (FMNH). One of us (CM) examined most of the material in 12of these institutions, whereas information about material housed in sixother institutions were obtained online from the Global Biodiversity In-formation Facility (GBIF portal; http://www.gbif.org). Primary types of all24 taxa described or recorded from Guatemala are housed in MNHN (12taxa), USNM (five taxa), Forschungsinstitut und Naturmuseum Sencken-berg, Frankfurt a.M. (SMF) (three taxa), Naturhistorisches Museum Wien,Vienna (NHMW) (two taxa), and The Natural History Museum, London(NHM) and Naturhistorisches Museum Basel, Basel (NHMB) (one taxoneach).

We followed the classification scheme for pseudothelphusids as indi-cated in Villalobos Hiriart and Álvarez (2008), and specimens were identi-fied following the descriptions and keys in Pretzmann (1980) and Rodriguez(1982), complemented by notes on morphology and illustrations obtainedby one of us (CM) after examining the type specimens of most Guatemalantaxa.

RESULTS

Species Diversity

We collected a total of 55 freshwater crabs comprising sixpseudothelphusid species during the 2014 and 2015 sur-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.

Dow



ay 2020

http://www.gbif.org




Table 1. Number and identification status of the lots with primary fresh-water crabs from Guatemala (all belonging to Pseudothelphusidae) de-posited in collections. AMNH, American Museum of Natural History, NewYork; CNCR, Colección Nacional de Crustáceos, Universidad NacionalAutónoma de México, Mexico City; FLMNH, Florida Museum of Nat-ural History, University of Florida, Gainesville; FMNH, Field Museumof Natural History, Chicago; INPA, Instituto Nacional de Pesquisas daAmazônia, Manaus; MCZ, Museum of Comparative Zoology, Harvard Uni-versity, Cambridge; MNHN, Muséum national d’Histoire naturelle, Paris;NHM, The Natural History Museum, London; NHMB, NaturhistorischesMuseum Basel, Switzerland; NHMW, Naturhistorisches Museum Wien,Vienna; SMF, Forschungsinstitut und Naturmuseum Senckenberg, Frank-furt a.M.; SNSD, Senckenberg Naturhistorische Sammlungen Dresden;TMNH, Tulane Museum of Natural History, Tulane University, New Or-leans; UCR-MZ, Museo de Zoología, Escuela de Biología, Universidad deCosta Rica, San José; USNM, United States National Museum of NaturalHistory, Smithsonian Institution, Washington, DC; UVG, Universidad delValle de Guatemala, Guatemala City; YPM, Yale Peabody Museum of Nat-ural History, New Haven; ZMH, Zoologisches Museum, Universität Ham-burg. ∗Data from direct examination; ∗∗Data from the literature or GlobalBiodiversity Information Facility’s Portal (www.gbif.org).

Collection Numberof lots

Identifiedto species

level

Not identified(juveniles,females, ordamaged)

AMNH∗ 7 2 5CNCR∗∗ 1 1FLMNH∗ 1 1FMNH∗ 11 9 2INPA∗ 7 7MCZ∗∗ 2 2MNHN∗ 14 14NHM∗∗ 2 1 1NHMB∗∗ 1 1NHMW∗ 2 2SMF∗ 3 3SNSD∗∗ 2 2TMNH∗ 6 5 1UCR-MZ∗ 19 17 1USNM∗ 38 30 8UVG∗ 57 42 15YPM∗∗ 3 3ZMH∗ 1 1Total 177 139 37

carcinini (Table 2). The most common species were Potamo-carcinus armatus, Phrygyopilus ibarrai, Raddaus bocourti,and Potamocarcinus magnus representing 20.0%, 14.5%,14.5%, and 14.5% of the material identified to species level,respectively. The remaining species, Raddaus tuberculatus(11.0%) and Elsalvadoria zurstrasseni (7.3%), were less fre-quent. Nearly one fifth (18.2%) of the collection consistedof females and juveniles that could not be identified even togeneric level. The 57 lots of the UVG collection containeda total of 69 individuals of Pseudothelphusidae; 44 of theselots held specimens that could be identified at least to genus.This material was comprised by a total of nine pseudothel-phusid species, two of Hypolobocerini and seven species ofPotamocarcinini (Table 2). The predominant species were R.bocourti (61.7% of all identified lots) and Potamocarcinusmagnus (20.5%).

A revision of the literature (H. Milne Edwards, 1853;A. Milne-Edwards, 1866; Rathbun, 1896, 1897, 1905; Bott,1956, 1970; Pretzmann, 1965, 1971, 1978, 1980; Smalley,

1970; Delamare Deboutteville, 1976; Rodriguez, 1982; Ro-driguez and Hobbs, 1989; Rodriguez, 2001) and examina-tion of collections (Table 1) revealed that a total of 27 taxa offreshwater crabs, all belonging to Pseudothelphusidae, haveso far been described or recorded from Guatemala. Only 14species and one subspecies, however, are currently recog-nized as valid taxa, whereas five taxa are considered syn-onyms, five are incertae sedis, and two remain undescribed.

SYSTEMATICS

Pseudothelphusidae Ortmann, 1893Tribe Hypolobocerini Pretzmann, 1971

Elsalvadoria tomhaasi Bott, 1970

Distribution.—Guatemala: Guatemala (this study) andSacatepéquez (Bott, 1970) departments.

Elsalvadoria zurstrasseni (Bott, 1956)

Distribution.—Guatemala: Escuintla, Guatemala, and Su-chitepéquez (this study) departments; El Salvador (Bott,1956; Rodriguez, 1982).

Remarks.—New record for Guatemala (Fig. 3).

Tribe Potamocarcinini Ortmann, 1897Phrygiopilus acanthophallus Smalley, 1970

Distribution.—Guatemala: Alta Verapaz (Smalley, 1970;Rodriguez, 1982) and Zacapa (this study) departments.

Phrygiopilus chuacusensis Smalley, 1970

Distribution.—Guatemala: Baja Verapaz (Smalley, 1970)and Zacapa (this study) departments.

Phrygiopilus ibarrai (Pretzmann, 1978)

Distribution.—Guatemala: Baja Verapaz (this study) andGuatemala (Pretzmann, 1978; Rodriguez, 2001) depart-ments.

Phrygiopilus strengerae (Pretzmann, 1965)

Distribution.—Guatemala: Alta Verapaz department (Pretz-mann, 1965).

Remarks.—Pretzmann (1965) stated that the type localityis “Trace Aguas, Caco” in Guatemala department, butaccording to Reddel (1981: 317), Cacao, Trece Aguas isprobably in the Seamay-Sejul karstik system, Alta Verapazdepartment.

Phrygiopilus n. sp.

Distribution.—Guatemala: Escuintla (this study) depart-ment.

Potamocarcinus armatus H. Milne Edwards, 1853Potamocarcinus (Potamocarcinus) nicaraguensis aequispinosus Pretzmann,

1971: 20.

Distribution.—Mexico (Pretzmann, 1972); Guatemala: AltaVerapaz (this study) and Petén (Pretzmann, 1971, 1980; thisstudy) departments.

Remarks.—Pretzmann (1971) only mentioned the countryas the type locality of P. (P.) nicaraguensis aequispinosus.

Dow



ay 2020

http://www.gbif.org


Tabl

e2.

Upd

ated

listo

fth

ecu

rren

tlyva

lidsp

ecie

sof

prim

ary

fres

hwat

ercr

abs

(Pse

udot

help

husi

dae)

from

Gua

tem

ala

with

cons

erva

tion

stat

us(a

ccor

ding

toC

umbe

rlid

geet

al.,

2009

)an

dnu

mbe

rof

lots

with

iden

tified

spec

imen

sde

posi

ted

inco

llect

ions

exam

ined

(exc

eptw

here

mar

ked

byan

aste

risk

),in

clud

ing

hist

oric

alre

cord

sas

wel

las

spec

ies

colle

cted

duri

ngou

rre

cent

surv

eys.

AM

NH

,Am

eric

anM

useu

mof

Nat

ural

His

tory

,New

Yor

k;FM

NH

,Fie

ldM

useu

mof

Nat

ural

His

tory

,Chi

cago

;IN

PA,I

nstit

uto

Nac

iona

lde

Pesq

uisa

sda

Am

azôn

ia,M

anau

s;M

NH

N,M

uséu

mna

tiona

ld’

His

toir

ena

ture

lle,P

aris

;N

HM

W,

Nat

urhi

stor

isch

esM

useu

mW

ien,

Vie

nna;

SMF,

Fors

chun

gsin

stitu

tun

dN

atur

mus

eum

Senc

kenb

erg,

Fran

kfur

ta.

M.;

TM

NH

,T

ulan

eM

useu

mof

Nat

ural

His

tory

,T

ulan

eU

nive

rsity

,N

ewO

rlea

ns;

UC

R-M

Z,

Mus

eode

Zoo

logí

a,U

nive

rsid

adde

Cos

taR

ica,

San

José

;U

SNM

,U

nite

dSt

ates

Nat

iona

lM

useu

mof

Nat

ural

His

tory

,Sm

ithso

nian

Inst

itutio

n,W

ashi

ngto

n,D

C;

UV

G,

Uni

vers

idad

del

Val

lede

Gua

tem

ala,

Gua

tem

ala

City

.Con

serv

atio

nca

tego

ries

:DD

,Dat

a-D

efici

ent;

LC

,Lea

stC

once

rn;N

A,N

otA

sses

sed;

V,V

ulne

rabl

e.

Con

serv

atio

nst

atus

Spec

ies

Col

lect

ions

Obs

erva

tions

Ref

eren

ces

Pres

ent

sur-

veys

AMNH

FMNH

INPA

MNHN

NHMW

SMF

TMNH

UCR-MZ

USNM

UVG

DD

Els

alva

dori

ato

mha

asi

Bot

t,19

701

1B

ott(

1970

),th

isst

udy

VE

lsal

vado

ria

zurs

tras

seni

(Bot

t,19

56)

11

11

New

reco

rdfo

rG

uate

mal

aT

his

stud

y

VP

hryg

iopi

lus

acan

thop

hall

usSm

alle

y,19

70

11

3Sm

alle

y(1

970)

,Rod

rigu

ez(1

982)

,thi

sst

udy

NA

Phr

ygio

pilu

sch

uacu

sens

isSm

alle

y,19

701

21

Smal

ley

(197

0),t

his

stud

y

DD

Phr

ygio

pilu

sib

arra

i(P

retz

man

n,19

78)

21

11

1Pr

etzm

ann

(197

8),t

his

stud

y

DD

Phr

ygio

pilu

sst

reng

erae

(Pre

tzm

ann,

1965

)1

Pret

zman

n(1

965)

Phr

ygio

pilu

ssp

.1

Und

escr

ibed

spec

ies

Thi

sst

udy

LC

Pota

moc

arci

nus

arm

atus

H.M

ilne

Edw

ards

,185

32

32

Pret

zman

n(1

971,

1980

),th

isst

udy

LC

Pota

moc

arci

nus

mag

nus

Rat

hbun

,189

64

26

52

46

Rat

hbun

(190

5),P

retz

man

n(1

980)

,Rod

rigu

ez(1

982)

,thi

sst

udy

LC

Rad

daus

boco

urti

(A.M

ilne-

Edw

ards

,186

6)4

21

86

11

927

Rat

hbun

(190

5),P

retz

man

n(1

980)

,Rod

rigu

ez(1

982)

,thi

sst

udy

NA

Rad

daus

mer

tens

i(B

ott,

1956

)1

New

reco

rdfo

rG

uate

mal

aT

his

stud

y

DD

Rad

daus

tube

rcul

atus

(Rat

hbun

,189

7)2

22

Rat

hbun

(189

7),R

odri

guez

(198

2),t

his

stud

y

LC

Typh

lops

eudo

thel

phus

am

itch

elli

Dem

alar

eD

ebou

ttevi

lle,1

976

1∗R

epor

ted

from

cave

sD

emal

are

Deb

outte

ville

(197

6)

LC

Typh

lops

eudo

thel

phus

aju

bert

hiei

Dem

alar

eD

ebou

ttevi

lle,1

976

1∗R

epor

ted

from

cave

sD

emal

are

Deb

outte

ville

(197

6)

DD

Zil

chia

falc

ata

Rod

rigu

ezan

dH

obbs

,198

92

Rep

orte

dfr

omca

ves

Rod

rigu

ezan

dH

obbs

(198

9)

NA

Zil

chia

zilc

hige

rman

iPr

etzm

ann,

1978

12

Thi

sst

udy

Zil

chia

sp.

2U

ndes

crib

edsp

ecie

sT

his

stud

y

Dow



ay 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 onlineedition of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/journals/1937240x.

Potamocarcinus magnus (Rathbun, 1896)Potamocarcinus guatemalensis Rathbun, 1905: 313, text-fig. 101, pl. 14,

fig. 7.

Distribution.—Mexico (Rodriguez, 1982); Guatemala: AltaVerapaz (Rathbun, 1905; this study), Escuintla (Rodriguez,1982), Guatemala (Rathbun, 1905; Pretzmann, 1980; thisstudy), Izabal (Pretzmann, 1980; this study), Jalapa (thisstudy), Péten (this study), Sololá (Pretzmann, 1980; Wehrt-mann et al., 2014), Suchitepéquez (this study), and Zacapa(Pretzmann, 1980; this study) departments; Honduras (Pretz-mann, 1980); El Salvador (Rodriguez, 1982); Costa Rica(Magalhães et al., 2015).

Raddaus bocourti (A. Milne-Edwards, 1866)Pseudothelphusa similis Rathbun, 1905: 295, text-fig. 87, pl. 15, fig. 4.Pseudothelphusa cobanensis Rathbun, 1905: 296, text-fig. 88, pl. 15, fig. 5.Pseudothelphusa grallator Rathbun, 1905: 297, text-fig. 89, pl. 15, fig. 6.

Distribution.—Mexico (Rodriguez, 1982); Belize (Pretz-mann, 1980; Rodriguez, 1982); Guatemala: Alta Verapaz(Rathbun, 1905; Pretzmann, 1980; Rodriguez, 1982), Baja

Verapaz (this study), Chiquimula (Pretzmann, 1980), Chi-maltenango (this study), Escuintla (this study), Guatemala(Pretzmann, 1980; Rodriguez, 2001; this study), Izabal (thisstudy), Petén (Pretzmann, 1980; this study), Quiché (thisstudy), Santa Rosa (this study), Sololá (Rodriguez, 2001;Wehrtmann et al., 2014), Suchitepéquez (this study), To-tonicapán (Rathbun, 1905), and Zacapa (this study) de-partments; Honduras (Pretzmann, 1980); El Salvador (Ro-driguez, 1982).

Raddaus mertensi (Bott, 1956)

Distribution.—Guatemala: Chiquimula department (thisstudy); El Salvador (Bott, 1956).

Remark.—New record from Guatemala (Fig. 3).

Raddaus tuberculatus (Rathbun, 1897)

Distribution.—Mexico (Pretzmann, 1972); Guatemala: SanMarcos (this study), Sololá (Rathbun, 1897, 1905), andSuchitepéquez (this study) departments.

Dow



ay 2020



Typhlopseudothelphusa juberthiei Delamare Deboutteville,1976

Distribution.—Guatemala: Alta Verapaz department (Dela-mare Deboutteville, 1976).

Typhlopseudothelphusa mitchelli Delamare Deboutteville,1976

Distribution.—Guatemala: Alta Verapaz department (Dela-mare Deboutteville, 1976).

Zilchia falcata Rodriguez and Hobbs, 1989

Distribution.—Guatemala: Alta Verapaz department (Ro-driguez and Hobbs, 1989).

Zilchia zilchi germani Pretzmann, 1978

Distribution.—Guatemala: Guatemala department (thisstudy).

Remarks.—Pretzmann (1978: 3; 1980: 658) stated that thetype locality of this subspecies is “Rio Guyamas, NE ElProgresso, Honduras.” This is an error, and the actual typelocality is “Guatemala, Department of Guatemala, 14 kmsouth of Guatemala City, near San José Pinula.” Thisinformation is given in a letter dated 9 April 1976 by IvoPoglayen-Newall to G. Pretzmann, filed in the crustaceansection of NHMW (CM).

Zilchia n. sp.

Distribution.—Guatemala: Escuintla department (thisstudy).

Incertae SedisPseudothelphusa bisunturalis Rathbun, 1897

Distribution.—Guatemala: Suchitepéquez department, SanAgustín stream, near Atitlán volcano, Pacific slope (Rath-bun, 1897).

Boscia gracilipes A. Milne-Edwards, 1866

Distribution.—Guatemala: Alta Verapaz department, exactlocality unknown (A. Milne-Edwards, 1866).

Gordonia longipes Pretzmann, 1965

Distribution.—Guatemala: Alta Verapaz department, Lan-quín (Pretzmann, 1965).

Pseudothelphusa propinqua Rathbun, 1905

Distribution.—Guatemala: Alta Verapaz department, Cobán(Rathbun, 1905).

Pseudothelphusa proxima Rathbun, 1905

Distribution.—Guatemala: Alta Verapaz department, Cobán(Rathbun, 1905).

DISCUSSION

The current taxonomic knowledge on the freshwater crabfauna of Guatemala is based on relatively few specimensthat have been sporadically collected since the 19th cen-tury and deposited in collections of European and NorthAmerican institutions. Following the paradigm of the pe-riod, the first descriptions of species were based mainly

on somatic characters of the carapace, and several of thetaxa were described based on female specimens (H. MilneEdwards, 1853; A. Milne-Edwards, 1866; Rathbun, 1896,1897, 1905). Many of the types were preserved as driedspecimens. Almost all of this material described between1853 and 1905 is deposited at the MNHN (types of ninespecies), except for the specimens of two species found atUSNM and one at NHMB. From the mid-20th century on,the examination of characters of the male first gonopod be-came essential for a reliable identification of the taxa, andseveral taxonomists (Bott, 1956, 1970; Pretzmann, 1965,1978; Smalley, 1970; Delamare Deboutteville, 1976; Ro-driguez and Hobbs, 1989) relied on this structure to diagnosespecies. The primary types of the Guatemala species de-scribed after 1956 are housed in MNHN (three types), SMF(three types), USNM (three types), NHMW (two types),and NHMB (one type). In addition to these type specimens,few non-type material, usually consisting of single males orfew specimens, were available for study (Tables 1 and 2).This situation, as well as the uncertain taxonomic status ofsome taxa, have so far prevented a more accurate assess-ment of the diversity and distribution of the freshwater crabsof Guatemala. Wehrtmann et al. (2014), for instance, com-mented on the implications of managing the artisanal crabfishery in Lake Atitlán as a single-species fishery (Potamo-carcinus guatemalensis = P. magnus), when it actually in-volved two species: Raddaus bocourti and P. magnus. Ro-driguez (1982) listed five species as incertae sedis becausethey were described from female specimens, which makesit virtually impossible to ascertain their specific identity be-cause the current classification relies almost exclusively oncharacters of the male first gonopod.

Such limitations affected the recent evaluation for con-servation assessment purposes of the freshwater crabs ofGuatemala by Cumberlidge et al. (2014). They reported 12species, whereas here we recognized 17 species from sixgenera. In spite of the uncertainties, Guatemala has a di-verse freshwater crab fauna, comparable to that of CostaRica and Panama. Magalhães et al. (2013) mentioned sixgenera and 17 species for Panama and four genera and 15species for Costa Rica (Magalhães et al., 2015). Our knowl-edge of the freshwater crabs from El Salvador, Honduras,and Nicaragua has not been updated since the early 1980s(Pretzmann, 1980; Rodriguez, 1982). All the collected sam-ples are kept in European or North American institutions,and it is not known if institutions in these three countriesmaintain collections of freshwater crabs.

Our current knowledge of the conservation status of thefreshwater crabs of Guatemala is still scarce: 47% of the 17species (including two yet undescribed species) are consid-ered as data-deficient or have not been yet evaluated (Ta-ble 2). Two species (Elsalvadoria zurstrasseni and Phry-giopilus acanthophallus) are classified as vulnerable accord-ing to IUCN categories and criteria (IUCN, 2003) (Table 2),and 65% (11 species, including the two undescribed species)are endemic to Guatemala, a high level of endemism fora continental country (Cumberlidge et al., 2009). Further-more, three cave-dwelling species inhabit Guatemala, andthese species are especially vulnerable to disturbance andpollution (Ng and Yeo, 2007; Cumberlidge et al., 2009).

Dow



ay 2020


This setting, together with the alarmingly poor quality offresh water (Elmore et al., 2005; Padilla Cámbara et al.,2010), represents a serious threat to the crab fauna of thecountry. Other threats related to anthropogenic habitat dis-turbances are imminent as well: the increasing demographicpressure (Castillo et al., 1997; IARNA, 2003; MARN, 2010;Rejmánková et al., 2011), increasing and accelerating de-terioration of the environment, risk of pollution (includingintensive use of pesticides and chemical fertilizers), and theincreasing levels of waste, are some of the serious threats(CEMAT, 1999; Elmore et al., 2005; Padilla Cámbara etal., 2010). Considering that even widely distributed specieswith an apparent tolerance to land-use changes have declineddrastically in abundance as a consequence of changes in landuse, hydrology, or pesticide regimes (Cumberlidge et al.,2009), conservation managers of governmental as well asnon-governmental organizations need to recognize that thisfreshwater crab fauna is seriously threatened. Conservationstrategies are urgently needed to preserve the integrity ofthe habitats along with studies to fill gaps in data-deficientspecies (Cumberlidge et al., 2009, 2014).

ACKNOWLEDGEMENTS

We thank the Consejo Nacional de Ciencia y Tecnología, Costa Rica(CONICIT; project IQ-0001-11) and the Conselho Nacional de Desenvolvi-mento Científico e Tecnologico (CNPq; Brazil, projects 490314/2011-2) fortheir financial support to ISW and CM. CM also thanks CNPq for an on-going research grant (304736/2015-5). Additional support for ISW was re-ceived from Universidad de Costa Rica (project VI 808-B3-504). We areespecially grateful to Octavio Cajas (Consejo Nacional de Areas Protegi-das de Guatemala) for his support with the collection and export permits,and to Raquel Romero Chaves (Unidad de Investigación Pesquera y Acui-cultura, Universidad de Costa Rica) for the preparation of the distributionmaps. We would like to express our gratitude to Margarita Palmieri andGabriela Palomo (Biology Department, Universidad del Valle de Guatemala(UVG)), who facilitated the study of the UVG collection, and to StefaniaRodríguez and Christine LeBeau (AMNH), Jochen Gerber (FMNH), LaureCorbari, Paula Martin-Lefrève and Sébastien Soubzmaigne (MNHN), Pe-ter Dworschak (NHMW), the late Michael Türkay (SMF), Hank Bart Jr.,and Nelson Rios (TMNH), and Rafael Lemaitre and Karen Reed (USNM)for their assistance during the visits of CM to their respective institutions.Thanks are also due to Oscar Rolando Gonzáles (UVG) for his supportin checking the coordinates for the collection sites from the UVG collec-tion. Two anonymous reviewers provided constructive comments, whichare greatly appreciated. We are also grateful to Tayler Clarke (Universityof British Columbia, Vancouver, Canada) for revising the English.

REFERENCES

Abell, R., M. L. Thieme, C. Revenga, M. Bryer, M. Kottelat, N. Bogut-skaya, B. Coad, N. Mandrak, S. Contreras Balderas, W. Bussing, M. L. J.Stiassny, P. Skelton, G. R. Allen, P. Unmack, A. Naseka, R. Ng, N. Sin-dorf, J. Robertson, E. Armijo, J. V. Higgins, T. J. Heibel, E. Wikra-manayake, D. Olson, H. L. López, R. E. Reis, J. G. Lundberg, M. H.Sabaj Pérez, and P. Petry. 2008. Freshwater ecoregions of the world:a new map of biogeographic units for freshwater biodiversity conserva-tion. BioScience 58: 403-414.

Bott, R. 1956. Dekapoden (Crustacea) aus El Salvador. 3. Süsswasserkrab-ben (Pseudothelphusa). Senckenbergiana biologica 37: 229-242.

. 1970. Bemerkungen zu einigen Süsswasserkrabben (Crustacea:Decapoda). Senckenbergiana biologica 51: 355-461.

Castillo, L. E., E. de la Cruz, and R. Clemens. 1997. Ecotoxicologyand pesticides in tropical aquatic ecosystems of Central America.Environmental Toxicology and Chemistry 16: 41-51.

CEMAT (Centro Mesoamericano de Estudios sobre Tecnología Apropiada).1999. Diagnóstico de la Condición Ambiental del Agua y del Manejode Desechos en Guatemala. SIGA/SICA-CCAD-CEMAT, Ciudad deGuatemala.

Collen, B., F. Whitton, E. E. Dyer, J. E. M. Baillie, N. Cumberlidge, W. R. T.Darwall, C. Pollock, N. I. Richman, A.-M. Soulsby, and M. Böhm. 2014.Global patterns of freshwater species diversity, threat and endemism.Global Ecology and Biogeography 23: 40-51.

Cumberlidge, N., and P. K. L. Ng. 2009. Systematics, evolution, andbiogeography of freshwater crabs, pp. 491-508. In, J. W. Martin, K. A.Crandall, and D. L. Felder (eds.), Decapod Crustacean Phylogenetics.CRC Press, Boca Raton, FL.

, F. Alvarez, and J. L. Villalobos. 2014. Results of the global conser-vation assessment of the freshwater crabs (Brachyura, Pseudothelphusi-dae and Trichodactylidae): the Neotropical region, with an update on di-versity. ZooKeys 457: 133-157.

, P. K. L. Ng, D. C. J. Yeo, C. Magalhães, M. R. Campos, F. Alvarez,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 crabsand the biodiversity crisis: importance, threats, status, and conservationchallenges. Biological Conservation 142: 1665-1673.

Darwall, W., K. Smith, D. Allen, M. Seddon, G. McGregor Red, V. Claus-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.Stuart (eds.), The 2008 Review of the IUCN Red List of ThreatenedSpecies. IUCN, Gland.

Delamare Deboutteville, C. 1976. Intérêt biologique et écologique desCrabes cavernicoles du Guatemala et du Mexique appartenant augenre Typhlopseudothelphusa Rioja. Comptes rendus hebdomadaires desSéances de l’Académie des Sciences (Paris) (Série D) 283: 837-840.

Dudgeon, D., A. H. Arthington, M. O. Gessner, Z.-I. Kawabata, D. J.Knowler, C. Leveque, R. J. Naiman, A.-H. Prieur-Richard, D. Soto, M. L.Stiassny, and C. A. Sullivan. 2006. Freshwater biodiversity: importance,threats, status and conservation challenges. Biological Reviews 81: 163-182.

Elmore, A. C., G. R. Miller, and B. Parker. 2005. Water quality in Lemoa,Guatemala. Environmental Geology 48: 901-907.

IARNA (Instituto de Agricultura, Recursos Naturales y Agrícolas, Univer-sidad Rafael Landívar, Guatemala). 2003. Estado actual de los recursoshídricos de Guatemala. Informe Técnico No. 1, available online at http://biblio3.url.edu.gt/IARNA/serie_amb/4.PDF.

IUCN (International Union for Conservation of Nature). 2003. Guidelinefor Application of IUCN Red List Criteria at Regional Levels: Version3.0. IUCN Species Survival Commission. IUCN, Gland.

Lara, L. R., I. S. Wehrtmann, C. Magalhães, and F. L. Mantelatto.2013. Species diversity and distribution of freshwater crabs (Decapoda:Pseudothelphusidae) inhabiting the basin of the río Grande de Térraba,Pacific slope of Costa Rica. Latin American Journal of Aquatic Research41: 685-695.

Magalhães, C., and M. Türkay. 1996. Taxonomy of the Neotropical fresh-water crab family Trichodactylidae II. The genera Forsteria, Melocarci-nus, Sylviocarcinus, and Zilchiopsis (Crustacea: Decapoda: Brachyura).Senckenbergiana biologica 75: 97-130.

, and . 2008. Taxonomy of the Neotropical freshwater crabfamily Trichodactylidae, IV. The genera Dilocarcinus and Poppiana(Crustacea: Decapoda: Brachyura). Senckenbergiana biologica 88: 185-215.

, M. R. Campos, and M. Türkay. 2013. Freshwater crabs fromeastern Panamá: a new species of Potamocarcinus H. Milne Edwards,1853, and new records of two little-known species (Crustacea: Decapoda:Pseudothelphusidae, Trichodactylidae). Zootaxa 3702: 348-356.

, I. S. Wehrtmann, L. R. Lara, and F. L. Mantelatto. 2015. Freshwa-ter crabs from Costa Rica, with a taxonomic revision of the genus Pty-chophallus Smalley, 1964 (Crustacea: Decapoda: Pseudothelphusidae).Zootaxa 3905: 301-344.

MARN (Ministerio de Ambiente y Recursos Naturales de Guatemala).2010. Informe Ambiental del Estado de Guatemala. GEO, Guatemala,available online at http://biblio3.url.edu.gt/IARNA/MARN/GEO2009.pdf.

Milne-Edwards, A. 1866. Description de trois nouvelles espèces du genreBoscia, crustacés brachyures de la tribu des Thelphuseusiens. Annales dela Société entomologique de France (Série 4) 6: 203-205.

Milne Edwards, H. 1853. Mémoire sur la famille des Ocypodiens. Annalesdes Sciences naturelle (Zoologie) 20: 163-228.

Ng, P. K. L., and D. C. J. Yeo. 2007. Malaysian freshwater crabs:conservation prospects and challenges, pp. 95-120. In, L. Chua, L. G.Kirton, and L. G. Saw (eds.), Proceedings of the Seminar on the Status ofBiological Diversity in Malaysia and Threat Assessment of Plant Species

Dow



ay 2020

http://biblio3.url.edu.gt/IARNA/serie_amb/4.PDF

http://biblio3.url.edu.gt/IARNA/serie_amb/4.PDF

http://biblio3.url.edu.gt/IARNA/MARN/GEO2009.pdf

http://biblio3.url.edu.gt/IARNA/MARN/GEO2009.pdf


in Malaysia, 28-30 June 2005. Forest Research Institute of Malaysia,Kuala Lumpur.

Padilla Cámbara, T. A., N. García Álvarez, and W. Pérez Duarte. 2010.Caracterización físico-química y bacteriológica, en dos épocas del año,de la subcuenca del río Quiscab, Guatemala. Revista Ciencias TécnicasAgropecuarias (La Habana) 19: 43-46.

Pretzmann, G. 1965. Vorläufiger Bericht über die Familie Pseudothel-phusidae. Anzeiger der Österreichischen Akademie der WissenschaftenMathematische Naturwissenschaftliche Klasse: 1-10.

. 1971. Fortschritte in der Klassifizierung der Pseudothelphusidae.Sitzungsberichte der Österreichischen Akademie der Wissenschaften,Mathematisch Naturwissenschaftliche Klasse (Abteilung 1) 179: 15-24.

. 1972. Die Pseudothelphusidae (Crustacea Brachyura). Zoologica120: 1-182.

. 1978. Neue Potamocarcinini, Poglayen-Neuwall Leg. 1975 (vor-läufige Mitteilung). Sitzungsberichte der Österreichischen Akademieder Wissenschaften, Mathematisch Naturwissenchaftliche Klasse,(Abteilung 1) 187: 51-54.

. 1980. Von Dr. Ivo Poglayen-Neuwall 1975 in Mittelamerikagesammelte Krabben. Annalen des Naturhistorischen Museums in Wien83: 651-666, pls. 1-18.

Rathbun, M. J. 1896. Descriptions of two new species of fresh-water crabsfrom Costa Rica. Proceedings of the United States National Museum 18:377-379, pls. 19-30.

. 1897. Descriptions de nouvelles espèces de crabes d’eau douceappartenant aux collections du Muséum d’Histoire naturelle du Paris.Bulletin du Muséum nationale d’Histoire naturelle (Paris) 3: 58-61.

. 1905. Les crabes d’eau douce. Nouvelles Archives du Muséumd’Histoire naturelle (Paris) (4ème série) 7: 159-322.

Reddel, J. R. 1981. A review of the cavernicole fauna of Mexico,Guatemala, and Belize. The Bulletin of the Texas Memorial Museum 27:1-327.

Rejmánková, E., J. Komárek, M. Dix, J. Komárková, and N. Girón. 2011.Cyanobacterial blooms in lake Atitlan, Guatemala. Limnologica 41: 296-302.

Revenga, C., and Y. Kura. 2003. Status and trends of biodiversity of inlandwater ecosystems. Montreal. Technical Series no. 11. Secretariat of theConvention on Biological Diversity, Montreal, QC.

Rodriguez, G. 1982. Les Crabes d’Eau Douce d’Amérique. Familie desPseudothelphusidae. Faune Tropicale 22. ORSTOM, Paris.

. 1992. The Freshwater Crabs of America. Family Trichodactylidaeand Supplement to the Family Pseudothelphusidae. Faune Tropicale 31.ORSTOM, Paris.

. 2001. New species and records of pseudothelphusid crabs (Crus-tacea: Brachyura) from Central America in the Museum of Natural His-tory of Tulane University. Proceedings of the Biological Society of Wash-ington 114: 4-443.

, and H. H. Hobbs, Jr. 1989. A new cavernicolous crab, Zilchia fal-cata, from Guatemala, with notes on the genera of the Potamocarcini(Crustacea Decapoda, Pseudothelphusidae). Bulletin du Muséum na-tionale d’Histoire naturelle (Paris) (4ème série) 11(Section A): 183-192.

, and C. Magalhães. 2005. Recent advances in the biology of theNeotropical freshwater crab family of Pseudothelphusidae (Crustacea,Decapoda, Brachyura). Revista Brasileira de Zoologia 22: 354-365.

Smalley, A. E. 1970. A new genus of freshwater crabs from Guatemala,with a key to the Middle American genera (Crustacea, Decapoda,Pseudothelphusidae). American Midland Naturalist 83: 96-106.

Villalobos Hiriart, J. L., and F. Álvarez. 2008. Los cangrejos de la familiaPseudothelphusidae (Decapoda: Brachyura: Eubrachyura) de México,con un apéndice de las especies citadas para América hasta el 2006,pp. 239-300. In, F. Álvarez and G. Rodríguez-Almaraz (eds.), Crustáceosde México: Estado Actual de su Conocimiento. Universidad Autónomade Nuevo León, Monterrey.

Wehrtmann, I. S., C. Magalhães, and M. Orozco. 2014. Freshwater crabsin lake Atitlán, Guatemala: not a single-species fishery. Journal ofCrustacean Biology 34: 123-125.

Yeo, D. C. J., P. K. L. Ng, N. Cumberlidge, C. Magalhães, S. R. Daniels, andM. R. Campos. 2008. Global diversity of crabs (Crustacea: Decapoda:Brachyura) in freshwater. Hydrobiologia 595: 275-286.

RECEIVED: 1 June 2016.ACCEPTED: 18 August 2016.AVAILABLE ONLINE: 7 September 2016.

Dow



ay 2020

the primary freshwater crabs of guatemala …

Documents