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The Crayfish of Middle America

Chapter · August 2015

DOI: 10.1201/b18723-22

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 18
The Crayfish of Middle America
Fernando Alvarez* and José Luis Villalobos

Introduction
Middle America, herein taken as Mexico, Guatemala, Belize, Honduras and Cuba, has a
crayfish fauna composed of 61 species of the family Cambaridae, grouped in three genera
(Cambarellus, Orconectes, and Procambarus) and one introduced species (Cherax
quadricarinatus) that belongs to the family Parastacidae (Table 18.1). The distribution
of species by country shows that 56 species occur across Mexico, two in Guatemala,
one in Belize, one in Honduras and four in Cuba; one of these species (Procambarus
pilosimanus) occurs in Mexico, Guatemala, Belize, while another one (Procambarus
williamsoni) is in Guatemala and Honduras. The three cambarid genera recorded
for the region occur in Mexico, whereas only species of Procambarus are present in
the other four countries. Their regional diversification is seen in the distribution of
number of species, as it decreases from north to south, a pattern that is consistent with
the hypothesized origin of the Cambaridae in the southeastern United States and the
subsequent colonization of areas to the south in Mexico, and Central America (Hobbs
1984, Hobbs 2001).
Recent studies have outlined how and when the diversification of Middle American
stocks could have occurred after they separated from populations distributed in the
southern United States (Pedraza-Lara et al. 2012). Two main routes could have operated,
one through the paleolakes of the southern United States and north-central Mexico,
that can explain the current distribution of the western species of Cambarellus along
the Pacific slope and the presence of the primitive Procambarus bouvieri and P. digueti
in the State of Michoacan; and a second one along the Gulf of Mexico coast which
experimented many expansions and contractions due to sea level changes, where several
lineages evolved creating groups of related species that we now recognize as subgenera,
such as Villalobosus and Austrocambarus (Villalobos 1982, 1983, Hobbs 1984).

Colección Nacional de Crustáceos, Instituto de Biología, Universidad Nacional Autónoma de México,

Apartado Postal 70-153, México 04510, D.F., México.
* Corresponding author: falvarez@unam.mx

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Table 18.1 List of crayfish species from Mexico, Cuba and Central America. The species are listed alphabetically by genus, subgenus and species, for each country. The
conservation status is taken from the IUCN Red List of Endangered Species (IUCN: Ex, extinct; CE, critically endangered; E, endangered; NT, near threatened; V, vulnerable;
LC, least concern; DD, data deficient) and the Mexican Red List of Endangered and Protected Species Nom-059-Semarnat-2010 (Nom-059).

Species Type locality Distribution Conservation status

IUCN, Nom-059
MEXICO
Cambarellus (Cambarellus) alvarezi Villalobos, El Potosi (24°48’ N, 100°16’ W), Nuevo León, Known only for the type locality. Ex, -
1952 23 km NW of Galeana, Mexico.
Cambarellus (Cambarellus) areolatus Faxon, Near Parras, Coahuila, Mexico. Known from the type locality and a few CE, -
1885 sites nearby the type locality.
Cambarellus (Cambarellus) chapalanus Faxon, Lake Chapala, Jalisco, Mexico. Jalisco and Michoacan. NT, -
1898
Cambarellus (Cambarellus) chihuahuae Hobbs, Ojo de Carbonera, 57.6 km W of Villa Ahumada Known only for the type locality. Ex, -
1980 (30°35’ N, 106°51’ W), Chihuahua, Mexico.
Cambarellus (Cambarellus) lermensis Villalobos, Swamps in the town of Lerma, State of Mexico, Swamps, lakes and channels of the Lerma -, -
1943 Mexico. River, around the town of Lerma, in the
state of Mexico, Mexico.
Cambarellus (Cambarellus) montezumae de Ponds in Chapultepec Park, Distrito Federal, Widely distributed along the central LC, -

The Crayfish of Middle America 449

Saussure, 1857 Mexico. portion of Mexico, from Jalisco to Puebla.
Cambarellus (Cambarellus) occidentalis Faxon, Mazatlán, Sinaloa, Mexico. Pacific slope of Nayarit and Sinaloa, LC, -
1898 Mexico.
Cambarellus (Cambarellus) patzcuarensis Lake Patzcuaro, Michoacan, Mexico. Known only from Lake Patzcuaro, E, -
Villalobos, 1943 Michoacan.
Cambarellus (Cambarellus) prolixus Villalobos Lake Chapala, Jalisco, Mexico. Known only from Lake Chapala, Jalisco. CE, -
and Hobbs, 1981
Cambarellus (Cambarellus) zempoalensis Zempoala Lagoons, Morelos, Mexico. Known only from the type locality. LC, -
Villalobos, 1943
Orconectes (Gremicambarus) virilis (Hagen, Lake Superior, USA. From southwestern and Southcentral Introduced
1870) Canada, throughout the central United
States to northern Mexico in Chihuahua.
In Europe in The Netherlands.

Table 18.1 contd....

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Table 18.1 contd.
Species
Procambarus (Austrocambarus) acanthophorus
Villalobos, 1948
Procambarus (Austrocambarus) catemacoensis
Rojas, Álvarez and Villalobos, 2000
Procambarus (Austrocambarus) cavernicola
Mejía-Ortiz, Hartnoll and Viccon-Pale, 2003
Procambarus (Austrocambarus) citlaltepetl
Rojas, Álvarez and Villalobos, 1999
Procambarus (Austrocambarus) llamasi
Villalobos, 1954
Procambarus (Austrocambarus) maya Álvarez,
López-Mejía and Villalobos, 2007
Procambarus (Austrocambarus) mexicanus
(Erichson, 1846)
Procambarus (Austrocambarus) mirandai
Villalobos, 1954
Procambarus (Austrocambarus) oaxacae oaxacae Guano Cave, 10 km NE of Valle Nacional,
Hobbs, 1973
Procambarus (Austrocambarus) oaxacae reddelli El Nacimiento Cave, San Antonio River, 10 km
Hobbs, 1973
Procambarus (Austrocambarus) olmecorum
Hobbs, 1987
Procambarus (Austrocambarus) pilosimanus
(Ortmann, 1906)
Type locality
El Castillo, 4 km E of Tuxtepec, Oaxaca,
Mexico.
Espagoya, Lake Catemaco (18°48’27” N,
95°06’51” W; 335 m), Veracruz, Mexico.
Gabriel Cave, Acatlán de Pérez Figueroa,
Oaxaca, Mexico.
Rincón de la Doncella Park (18°48’12” N,
97°10’51” W; 1400 m), Ciudad Mendoza,
Veracruz, Mexico.
Santa Rita, 47 km E of Escarcega, Campeche,
Mexico.
Salt marsh (19°49’38” N, 87°30’39” W), Sian
Ka’an Nature Reserve, Felipe Carrillo Puerto,
Quintana Roo, Mexico.
El Mirador de Zacuapan, 8 km NE of Huatusco,
Veracruz, Mexico.
Cerro Hueco, 4 km SE of Tuxtla Gutiérrez,
Chiapas, Mexico.
Oaxaca, Mexico.
SSW of Acatlan, Oaxaca, Mexico.
Metlac River, Fortín de las Flores, Veracruz,
Mexico.
See type locality in Guatemala.
450 Freshwater Crayfish
Distribution Conservation status
IUCN, Nom-059
Swamps, lagoons and rivers of the coastal LC, -
plain in Veracruz, Tabasco and Campeche.
Known only from the type locality. CE, -
Known only from the type locality. V, -
Ciudad Mendoza and Orizaba, and a few V, -
sites in the Zongolica region, Veracruz.
Swamps, lagoons and rivers of the coastal LC, -
plain in Campeche, Yucatan and Quintana
Roo.
Known only from the type locality. DD, -
Known from the type locality and a few LC, -
sites nearby the type locality.
In Chiapas, from Tuxtla Gutiérrez LC, -
southwards along the Central Depression.
Known only from the type locality. LC, -
Known from the type locality and a few -, -
sites nearby the type locality.
Known from the type locality and a few LC, -
sites nearby the type locality.
Chiapas and Quintana Roo, close to the LC, -
Guatemalan border.
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Procambarus (Austrocambarus) rodriguezi Ojo de Agua Cave, WNW of Hacienda Potrero Known only from the type locality. DD, -
Hobbs, 1943 Viejo, Paraje Nuevo, Córdoba, Veracruz,
Mexico.
Procambarus (Austrocambarus) ruthveni (Pearse, Cuautotolapan, Acayucan, Veracruz, Mexico. Known from the type locality and a few V, -
1911) sites nearby the type locality.
Procambarus (Austrocambarus) sbordonii Hobbs, El Nacimiento Cave, Santo Domingo River, Known only from the type locality. DD, -
1977 Bochil, Chiapas, Mexico.
Procambarus (Austrocambarus) vazquezae Playa Norte, Lake Catemaco, Veracruz, Mexico. Known only from Lake Catemaco, NT, -
Villalobos, 1954 Veracruz
Procambarus (Austrocambarus) veracruzanus Presidio, 30 km SE of Córdoba, Veracruz, Known from the type locality and a few DD, -
Villalobos, 1954 Mexico. sites nearby the type locality.
Procambarus (Austrocambarus) zapoapensis Zapoapan de Cabañas, 11 km SE of Catemaco, Known only from the type locality. NT, -
Villalobos, 1954 Veracruz, Mexico.
Procambarus (Girardiella) regiomontanus 5 km N of Monterrey, Nuevo León, Mexico. Known from the type locality and a few CE, CE
(Villalobos, 1954) sites nearby the type locality.
Procambarus (Mexicambarus) bouvieri Uruapan, Michoacán, Mexico. Known from the type locality and a few E, -
(Ortmann, 1909) sites nearby the type locality.
Procambarus (Ortmannicus) caballeroi Stream S of Villa Juárez, Xicotepec de Juárez, Known from the type locality and a few LC, -
Villalobos, 1944 Puebla, Mexico. sites nearby the type locality.

The Crayfish of Middle America 451

Procambarus (Ortmannicus) cuevachicae Cueva Chica Cave, 1.6 km NE of Pujal, Ciudad Known from the type locality and a few LC, -
(Hobbs, 1941) Valles, San Luis Potosi, Mexico. sites nearby the type locality.
Procambarus (Ortmannicus) gonopodocristatus Unión Paso Largo, 16 km NE of Martinez de la Known from the type locality and a few LC, -
Villalobos, 1958 Torre, Veracruz, Mexico. sites nearby the type locality.
Procambarus (Ortmannicus) hidalgoensis López- Stream in the road to Olotla, 2 km E of Known only from the type locality. LC, -
Mejía, Álvarez and Mejía Ortiz, 2005 Tlanchinol, Hidalgo, Mexico.
Procambarus (Ortmannicus) toltecae Hobbs, 1943 Puente de Palictla, small stream 8 km N of Known from the type locality and a few LC, -
Tamazunchale, San Luis Potosí, Mexico. sites nearby the type locality.
Procambarus (Ortmannicus) villalobosi Hobbs, Del Agua Cave, 75 km E of Ciudad Valles, Known only from the type locality. DD, -
1967 Aquismón, San Luis Potosí, Mexico.
Procambarus (Ortmannicus) xilitlae Hobbs and Hoya de las Guaguas, 10 km SSW of Aquismón, Known only from the type locality. DD, -
Grubbs, 1982 San Luis Potosi, Mexico.

Table 18.1 contd....

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Table 18.1 contd.
Species
Procambarus (Procambarus) digueti (Bouvier,
1897)
Procambarus (Paracambarus) ortmannii
(Villalobos, 1949)
Procambarus (Paracambarus) paradoxus
(Ortmann, 1906)
Procambarus (Pennides) roberti Villalobos and
Hobbs, 1974
Procambarus (Scapulicambarus) clarkii (Girard,
1852)
Procambarus (Scapulicambarus) strenthi Hobbs,
1977
Procambarus (Villalobosus) achilli López, Mejía
and Álvarez, 2003
Procambarus (Villalobosus) chacalli López-
Mejía, Álvarez and Mejía-Ortiz, 2004
Procambarus (Villalobosus) contrerasi (Creaser,
1931)
Procambarus (Villalobosus) cuetzalanae Hobbs,
1982
Procambarus (Villalobosus) erichsoni Villalobos,
1950
Procambarus (Villalobosus) hoffmanni
(Villalobos, 1944)
Procambarus (Villalobosus) hortonhobbsi
Villalobos, 1950
Type locality
Tributaries of the Santiago River, Jalisco,
Mexico
Los Estajos, 6 km NE of Zihuateutla, Puebla,
Mexico.
Tetela de Ocampo y La Cañada, 35 km NE of
Zacapoaxtla, Tetela de Ocampo, Puebla, Mexico.
Ditch in La Media Luna Lagoon, 7.7 km S of
Rio Verde San Luis Potosi, Mexico.
Between San Antonio and El Paso del Norte,
Texas, USA.
Small stream 24 km W of Ciudad Valles, San
Luis Potosi, Mexico.
Atezca Stream, 5 km NW of Molango, Hidalgo,
Mexico.
Dejigui Spring, 4 km E of Huayacocotla,
Veracruz, Mexico.
Tributary of the Cazones River, near Agua Fria,
19.2 km S of Miahuapan, Puebla, Mexico.
Tasalolpan Cave, 5 km SW of Cuetzalan,
Puebla, Mexico.
Puendo, Mamay and Bojoy Streams, Tenango de
Doria, Hidalgo, Mexico.
Presa de Necaxa Dam spillway, Juan Galindo,
Puebla, Mexico.
El Coyular, 7 km NE of La Unión, Zihuateutla,
Puebla, Mexico.
452 Freshwater Crayfish
Distribution Conservation status
IUCN, Nom-059
Jalisco and Michoacan. E, -
Known only from the type locality. CE, -
Known only from the type locality. CE, -
Known from the type locality and a few E, -
sites nearby the type locality.
The native range in Mexico comprises LC, -
northern Chihuahua, Coahuila and Nuevo
Leon. The introduced range in Mexico is
in Chiapas.
Known only from the type locality. DD, -
Known only from the type locality. LC, -
Known only from the type locality. DD, -
Known from the type locality and a few E, -
sites nearby the type locality.
Known from the type locality and a few NT, -
sites nearby the type locality.
Known from the type locality and a few DD, -
sites nearby the type locality.
Northern Puebla and central Veracruz. LC, -
Known from the type locality and a few E, -
sites nearby the type locality.
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Procambarus (Villalobosus) riojai (Villalobos,
1944)
Procambarus (Villalobosus) teziutlanensis
(Villalobos, 1947)
Procambarus (Villalobosus) tlapacoyanensis
(Villalobos, 1947)
Procambarus (Villalobosus) xochitlanae Hobbs,
1975
Procambarus (Villalobosus) zihuateutlensis
Villalobos, 1950
CUBA
Procambarus (A.) atkinsoni (Ortmann, 1913)
Procambarus (A.) cubensis cubensis (Erichson,
1846)
Procambarus (A.) cubensis rivalis (Faxon, 1912)
Procambarus (A.) niveus Hobbs and Villalobos,
1964
GUATEMALA
Procambarus (A.) pilosimanus (Ortmann, 1906)
Procambarus (A.) williamsoni (Ortmann, 1905)
BELIZE
Procambarus (A.) pilosimanus (Ortmann, 1906)
HONDURAS
Procambarus (A.) williamsoni (Ortmann, 1905)
INTRODUCED
Cherax quadricarinatus
Tributaries of the Necaxa River, Huauchinango,
Puebla, Mexico.
Chignautla, 5 km E of Teziutlan, Chignautla,
Puebla, Mexico.
Cañada de Tomata, Tlapacoyan, Veracruz,
Mexico.
Los Camarones Cave, 3 km NW of Xochitlán,
Puebla, Mexico.
Tlatentiloyan Stream, Los Estajos, Zihuateutla,
Puebla, Mexico.
Tributaries of Los Indios River, Pinos Island,
Cuba.
Cuba.
San Diego de los Baños, Pinar del Rio, Cuba.
Santo Tomas Cave, near Ponce, Pinar del Rio,
Cuba.
Cahabón River, Coban, Alta Verapaz,
Guatemala.
Los Amates, 6-8 km S of Motagua River, Izabal,
Guatemala.
See type locality in Guatemala.
See type locality in Guatemala.
Known from the type locality and a few LC, -
sites nearby the type locality.
Known from the type locality and a few DD, -
sites nearby the type locality.
Known from the type locality and a few DD, -
sites nearby the type locality.
Known from the type locality and a few DD, -
sites nearby the type locality.
Known from the type locality and a few E, -
sites nearby the type locality.
Known from the type locality and a few DD
sites nearby the type locality.
Sacttered localities throughout Cuba. DD
Known from the type locality and a few -
sites nearby the type locality.
Known from the type locality and a few DD
sites nearby the type locality.
The Crayfish of Middle America 453
Northern Guatemala, Belize and southern LC
Mexico.
Known from the type locality, a few sites DD
nearby the type locality and from Lake
Yojoa, Honduras.
-
Lake Yojoa. -
Southern Tamaulipas, Media Luna
Lagoon in San Luis Potosi, southern
Morelos, Mexico.
 454 Freshwater Crayfish

Crayfish are a conspicuous component of freshwater ecosystems co-occurring

in many areas of the Neotropics with freshwater shrimps of the families Atyidae
and Palaemonidae and with freshwater crabs of the families Pseudothelphusidae and
Trichodactylidae (Villalobos et al. 2010, Alvarez et al. 2012). Due to the dependence
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of brackish water to complete their development, freshwater shrimp are restricted to

the continental slopes where many crayfish species occur, especially along the Gulf of
Mexico coastal plain. Crayfish are also common in the central portion of the continent,
where they have colonized lakes and streams, habitats that are also inhabited by
pseudothelphusid crabs (Alvarez et al. 2012).
In general, most of the species of crayfish dealt with here occur in low densities
and are not commercially exploited. However, a few species can reach very high
densities and are consumed seasonally and locally; examples of this are: Cambarellus
montezumae, in the Valley of Mexico and adjacent areas, where it has been consumed
by native Mexicans from well before the Spanish conquest (Arredondo et al. 2011);
Procambarus llamasi, consumed in small towns of southern Quintana Roo, where
it becomes very abundant at the beginning of the rainy season; and Procambarus
acanthophorus, consumed in towns along the Coatzacoalcos River, Veracruz, where it
is sold live in local markets.

Systematics
The taxonomy of the Cambaridae is largely based on the morphology of the modified
first pair of abdominal appendages of males, which are used as copulatory organs and
referred to as gonopods. Males in the family Cambaridae can have two morphological
forms according to their reproductive phase: form I males, which are in a reproductive
stage, are those that present a first gonopod with well defined apical processes and
sharp and well developed hooks on the ischia of the second, third or fourth pairs of
pereiopods; form II males, or non-reproductive males, exhibit a gonopod in which the
apical processes are blunt as well as the hooks on the ischia of the pereiopods. The
morphology of the form I male gonopod is the most important structure taxonomically,
whereas the genital plate of the female between the coxae of the fifth pair of pereiopods,
the annulus ventralis, can sometimes be used too for species identification (Villalobos
1983).
The three genera of crayfish present in Middle America can be distinguished
by the structure of the form I male gonopod, and to some extent by their size. The
species of Cambarellus are typically small-sized, reaching up to 4 cm in total length.
The gonopod of Cambarellus has three processes that can be slender and elongated or
somewhat truncated, grooved or spatulated (Fig. 18.1). The hooks are present on the
ischia of the second and third pairs of pereiopods. The annulus ventralis of females,
composed of two rami, has an inverted U shape oriented caudally and is articulated.
The 10 species distributed in Middle America (Table 18.1) belong to the subgenus
Cambarellus characterized by a male gonopod with curved apical processes and a
mesial process with a longitudinal groove, and females with a postannular sternite as
wide as long and a symmetrical annulus ventralis (Fitzpatrick 1983).
In Orconectes the ischium of the third maxilliped bears teeth on the mesial margin,
the palm of the chela has a row of usually less than 12 tubercles, the ischium of the

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 The Crayfish of Middle America 455
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Fig. 18.1 Crayfish species from Middle America: (A) Procambarus clarkii, Durango, Mexico; (B)
Cambarellus montezumae, Mexico City, Mexico; (C) Procambarus llamasi, Quintana Roo, Mexico; (D)
Procambarus sp., Chiapas, Mexico; (E) Procambarus xochitlanae, Puebla, Mexico; (F) Procambarus
toltecae, Veracruz, Mexico; (G) Orconectes virilis, Chihuahua, Mexico; (H) Procambarus mirandai,
Chiapas, Mexico.

third, rarely third and fourth, pereiopod of male with hook. The gonopod of the form
I male has usually two apical processes, rarely three, divergent, straight or curved
caudodistally or caudally (Hobbs 1974). Orconectes virilis is the only species of the

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 456 Freshwater Crayfish

genus present in Middle America. It was introduced in northern Mexico in the state of
Chihuahua, where a population has subsisted for several decades.
Procambarus is the most species rich genus in the Cambaridae, and is represented
by 50 species in Middle America (Table 18.1). Procambarus has been organized
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in 17 subgenera, of which nine are distributed in Middle America. In alphabetical

order, the main morphological characteristics of the subgenera, presence of hooks on
pereiopods and morphology of the form I male gonopod, taken from Hobbs (1972) are:
Austrocambarus, hooks on ischia of third pair of pereiopods and absent from the fourth
pair, gonopod with lanceolate mesial process directed distally or laterodistally, central
projection acute or blade-like, and caudal process absent, vestigial or adopting different
shapes; Girardiella, hooks on ischia of third pair of pereiopods and absent from the
fourth pair, gonopod without subapical setae, with prominent cephalic shoulder, mesial
process with spicule-like shape, cephalic process acute or subacute on cephalomesial
angle, compressed caudal process and blade-like central projection; Mexicambarus,
hooks on ischia of third pair of pereiopods and absent form the fourth pair, gonopod
with reduced and acute mesial process, prominent, curved, lamelliform cephalic process
and small central projection; Ortmannicus, hooks on ischia of third and fourth pairs of
pereiopods, gonopod usually without cephalic shoulder, subapical setae almost always
present, mesial process spiculate to lanceolate, cephalic process from blade-like to
setiform, caudal process and central projection variable in size, shape and position;
Paracambarus, hooks on ischia of fourth pair of pereiopods and absent form the third
pair, first gonopods asymmetrical, gonopod with reduced cephalic shoulder, without
subapical setae, large caudodistal or small caudolateral mesial process, cephalic process
reduced or absent, central projection beak-like, caudal process vestigial; Pennides, hooks
on ischia of third and fourth pairs of pereiopods, first gonopods asymmetrical, gonopod
usually without cephalic shoulder, subapical setae present, slender and acute mesial
process, cephalic process in mesial to cephalolateral position, caudal process variable
in shape, reduced or absent, and central projection dentiform; Procambarus, hooks on
ischia of third pair of pereiopods and absent form the fourth pair, gonopod with cephalic
shoulder, without subapical setae, mesial process setiform, cephalic process lamelliform,
caudal process reduced, cental projection blade-like; Scapulicambarus, hooks on
ischia of third and fourth pairs of pereiopods, gonopod with well developed cephalic
shoulder, subapical setae present, acute mesial process, cephalic process lamelliform,
caudal process subspatulate and central projection small; Villalobosus, hooks on ischia
of fourth pair of pereiopods, gonopods asymmetrical, gonopod usually without setae,
mesial process variable in size and shape, cephalic process vestigial, flattened or
spiniform, caudal process tooth-like to prominent and wide, central projection varying
from acute to tuberculiform.

Phylogenetic Relationships
Cambarid crayfish have been the subject of a number of phylogenetic studies using
different types of data, morphological and molecular, and including also different
numbers of taxa. The general consensus is that the taxonomy at higher levels is
adequate, reflecting well phylogentic relationships (Crandall et al. 2000, Sinclair et al.
2004, Johnson et al. 2011); however, starting at the subgenus level, a category that has

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 The Crayfish of Middle America 457

been widely used in cambarid taxonomy, relationships are more complex than what is
reflected in current taxonomic arrangements.
The genus Orconectes was studied by Crandall and Fitzpatrick (1996) who
found a certain degree of incongruence between current taxonomy, especially of the
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subgenera, and phylogenetic relationships, suggesting that a new revision to reconcile

both approaches would be necessary. For Cambarellus, Pedraza-Lara et al. (2012) found
two main groups within the genus and several cryptic lineages that likely represent
new species. The first group recovered was the ‘Gulf group’ that includes species from
Texas, Louisiana and Florida, while the second group, the ‘Mexican group’ included
all the species in Mexico, from Chihuahua, Coahuila and Nuevo Leon, to those found
along the Trans-Mexican Volcanic Belt.
We are lacking a detailed phylogenetic study for Procambarus, and for the
whole family Cambaridae, that will consider morphological and molecular characters.
Current taxonomy has to be thoroughly revised and the usefulness of the subgenera in
Procambarus has to be reevaluated. Breinholt et al. (2012) found for Cambarus mixed
results regarding the monophyly of its subgenera and identified several convergent
characters that confound direct relationships.

Distribution
The crayfish of Middle America occur from sea level wetlands and even salt marshes,
to altitudes above 3,000 m in central Mexico. Among the species that can be found in
coastal plains very close to the coast line are: Procambarus caballeroi, P. hoffmanni and
P. gonopodocristatus in Veracruz; and P. llamasi and P. maya in the Yucatan Peninsula.
Although the natural habitat of crayfish is generally limited to freshwater, some species
like P. clarkii have been recorded in brackish water (Huner and Barr 1991); so is the
case for P. maya, collected in a salt marsh 1 km from the coast at a salinity of 5.5 ppt
in the Sian Ka’an Nature Reserve in Quintana Roo, Mexico (Alvarez et al. 2007).
Other species occur at high altitudes, in particular three of the genus Cambarellus.
Cambarellus montezumae is very common in the Valley of Mexico at 2,300 m asl, but it
could have been very common at higher altitudes in the states of Puebla and Tlaxcala.
Cambarellus zempoalensis in the state of Morelos, is endemic of the Zempoala Lagoons
at 2,800 m asl; while C. lermensis, from Lerma, state of Mexico, occurs at 3,000 m
asl (Alvarez et al. 2012).
Crayfish are not evenly distributed throughout Middle America. In Mexico more
species are found along the Gulf of Mexico slope, then a less diverse group occurs along
the Trans-Mexican Volcanic Belt, and a third disjunct group with only a few species is
distributed on the Pacific versant. A diversity hotspot for crayfish has been identified
along the Gulf of Mexico slope in a region where the states of Veracruz, Hidalgo and
Puebla, come together (Armendáriz 2011). Around 25 species occur in this area, with
representatives of several subgenera of Procambarus: Ortmannicus, Villalobosus and
Austrocambarus. A second area of high species diversity is located in southern Veracruz
and northern Oaxaca, where mainly species of Procambarus (Austrocambarus) occur
(Armendáriz 2011). A third area with high diversity of crayfish is located in Michoacan,
where several species of Cambarellus (C. montezumae, C. patzcuarensis, C. chapalanus)
are distributed together with Procambarus digueti and P. bouvieri.

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 458 Freshwater Crayfish

Ecology
Habitat
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Crayfish occur preferentially in lentic habitats, in places with little or no current, with
muddy bottoms or with a certain amount of organic matter (Rojas et al. 1997). They can
be found in floodplains and seasonal ponds where they thrive during the rainy season,
remaining in burrows during the dry period (Villalobos 1983). The burrowing behavior
of crayfish has been classified into three types according to the time the organisms
remain in the burrows and to the length and complexity of the burrows (Hobbs 2001).
Cambarellus montezumae, the most common species along the Trans-Mexican
Volcanic Belt is generally associated to the roots and branches of the riparian vegetation
in lakes, canals, ditches, dams and ponds. The range of water quality conditions where
the species is found are: pH 7.0–10.0, dissolved oxygen concentration 0.25–14.8 mg/l
and temperature 13.0–25.0ºC (Arana-Magallón et al. 1998, Alvarez and Rangel 2007).
Sixteen species of crayfish have been recorded from caves and hypogean waters;
however only six (Procambarus cavernicola, P. niveus, P. oaxacae oaxacae, P. oaxacae
reddelli, P. rodriguezi and P. xilitlae) have developed clear adaptations to cave life,
such as: reduced eyes, elongation of appendages and loss of pigmentation (Hobbs and
Villalobos 1964, Hobbs et al. 1977, Mejía-Ortiz et al. 2003). In the other 10 species
either their presence in caves is accidental, since they occur also in epigean bodies of
water, or they have invaded the cave environment recently as they don’t yet show any
obvious adaptations to cave life.

Reproduction
The reproductive cycle of cambarid crayfish has been extensively studied focusing
mainly on species with commercial value or species that have been introduced
in different parts of the world. Regarding species distributed in Middle America,
Cambarellus montezumae has been the subject of most of the studies examining
reproductive aspects. In central Mexico C. montezumae is relatively abundant all year
long, form II males are smaller (15–30 mm TL) than form I males (21–36 mm TL) and
both forms are always present, ovigerous females range from 21 to 24 mm TL with a
mean fecundity of 50 eggs, and the estimated lifespan is 15 months, but they could live
up to two years in the laboratory (Alvarez and Rangel 2007). Arredondo et al. (2011)
estimated a mean fecundity of 45 eggs per spawn with an 84% eclosion success.

Aquaculture
Few species of crayfish in Mexico and in Middle America have been used for
aquacultural purposes. The red swamp crayfish Procambarus clarkii has been introduced
into a number of lakes and ponds outside its native range in northern Mexico, under an
extensive aquacultural scheme. As a result P. clarkii has become an introduced species
in a number of sites through the states of Tamaulipas, Nuevo Leon, Coahuila, Durango,
Chihuahua, Sonora, Baja California and Chiapas (Campos and Rodríguez-Almaraz
1992, Hernández et al. 2008, Torres and Alvarez 2012).

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 The Crayfish of Middle America 459

For Procambarus acanthophorus, which is abundant along the coastal plain of the
Gulf of Mexico coast in southern Veracruz and Tabasco, important contributions have
been published on its suitability for culture. Aspects of its reproductive performance,
growth rate, mortality, and responses to selection under controlled conditions in
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experimental ponds and in the laboratory have been studied (Cervantes et al. 2010a,b,
Díaz et al. 2012).
Procambarus llamasi is another species that has shown attractive characteristics
for culturing. It is abundant in the Yucatan peninsula, attains large sizes and can be
found in high densities in the wild. The fecundity, growth, gonadal development and
spawning characteristics of this species have been studied (Rodríguez-Serna et al. 2000,
Carmona-Osalde et al. 2002, 2004a,b).

Conservation
Two initiatives have revised the conservation status of crayfish in Mexico. First, the
IUCN published in 2010 (IUCN 2013) the assessments for all cambarid species. The
Mexican environmental authority (SEMARNAT) also published the Mexican Red List of
threatened species in 2010 which is known as Nom-059-Semarnat-2010 (SEMARNAT
2010). All the assessments are included in Table 18.1; it has to be noted that only
one species of crayfish, Procambarus regiomontanus, is considered in the Nom-059-
Semarnat-2010 (SEMARNAT 2010). A first salient fact is that two species, Cambarellus
alvarezi and C. chihuahuae, are considered extinct. The former was endemic to a small
spring in El Potosí, a small village in southern Nuevo León. The species went extinct
when the spring was depleted for agricultural use (Contreras-Balderas and Lozano-
Vilano 1996, Alvarez et al. 2010a). Cambarellus chihuahuae experienced a similar
situation in an agricultural district near Villa Ahumada, Chihuahua, where groundwater
was overexploited dessicating most or all of the springs where this species was found
(Alvarez et al. 2010b). Another case where the same process might be operating is that
of C. areolatus in southern Coahuila, near the town of Parras, where several recent
surveys have failed to produce new samples of the species (Rodríguez-Almaraz and
Muñiz-Martínez 2008).
Six species are listed as critically endangered: Cambarellus areolatus, C. prolixus,
Procambarus catemacoensis, P. regiomontanus, P. ortmannii and P. paradoxus.
Cambarellus prolixus, Procambarus catemacoensis, and P. paradoxus are known from
a single site or a highly reduced area in places where human activities rapidly destroy
their environment. Procambarus ortamnni is known also from only the type locality
and from a reduced number of organisms, and has not been collected in recent years.
Procambarus regiomontanus populations have been declining due to environmental
degradation but also due to the introduction of P. clarkii in much of its range (Rodríguez-
Almaraz and Muñiz-Martínez 2008). The latter species is becoming more abundant
and expanding its range in Nuevo Leon (Hernández et al. 2008). It is noteworthy that
P. regiomontanus is listed in the Nom-059-Semarnat-2010 (SEMARNAT 2010) as
critically endangered.
Seven species are considered as endangered: Cambarellus patzcuarensis,
Procambarus bouvieri, P. digueti, P. roberti, P. contrerasi, P. hortonhobbsi and
P. zihuateutlensis. In all these cases the species are distributed in bodies of water that are
becoming heavily polluted and where canalization and water extraction is increasing.

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 460 Freshwater Crayfish

In Michoacan, Lake Patzcuaro (C. patzcuarensis), the springs and dams around the city
of Uruapan (P. digueti) and the Duero River in its last section before reaching Lake
Chapala (P. digueti) are heavily impacted. Media Luna Lagoon (P. roberti) in San Luis
Potosi is also threatened as it is a unique body of water in a desert area. For P. contrerasi,
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P. hortonhobbsi and P. zihuateutlensis, the main threat is habitat degradation as they

are distributed in areas of Puebla with intense agricultural activity.
Four species are classified as near threatened: Cambarellus chapalanus,
Procambarus vazquezae, P. zapoapensis and P. cuetzalanae. Cambarellus chapalanus
is distributed in a large area of Lake Chapala, Jalisco, which is the largest lake in
Mexico with a maximum length of 90 km; however, in spite of the large extension
of the lake, pollution and constant water level changes are becoming critical issues.
Procambarus vazquezae, distributed in Lake Catemaco has similar threats regarding
pollution from agrochemicals but is also facing an overexploitation of several key
species in the lake. Procambarus zapoapensis and P. cuetzalanae, occur in rural areas
in Los Tuxltas, Veracruz and in Cuetzalan, Puebla, where agriculture is expected to
intensify in future years.
Three species are placed as vulnerable: Procambarus cavernicola, P. citlaltepetl
and P. ruthveni. Procambarus cavernicola is a cave adapted species, known only from
one cave (Mejía-Ortiz et al. 2003), making it vulnerable in case the land around the
cave is impacted or modified. Procambarus citlaltepetl is known only from La Doncella
Park, an aquatic park in Ciudad Mendoza, Veracruz; thus its reduced distribution and
the potential for disturbances to occur in this location make it vulnerable (Rojas et al.
1999). Procambarus ruthveni also presents a highly reduced distribution range in Los
Tuxtlas region, Veracruz, where intense agricultural activity can become a threat to this
species (Alvarez et al. 2010c).
The largest group of species is that for which there is ‘least concern’. Nineteen
species appear in this category, of which 15 have relatively large distribution areas
suggesting that even if they are impacted at one locality there will be other populations
of the same species that will remain under favorable conditions. The other four species
of this group (C. occidentalis, C. zempoalensis, P. oaxacae oaxacae, P. hidalgoensis)
occur in one or a few nearby sites, but their populations are stable as no threats have
been noted.
The second largest group, with 16 species, corresponds to those classified as ‘data
deficient’. For most of the species in this group, all that is known comes from the original
paper where they were described and no later collections or published information are
available. Twelve of these species are distributed in Mexico and the other four occur
in Cuba, Guatemala and Honduras (Table 18.1).

Introduced Species
Three species have been recognized as introduced into Middle America: the northern
crayfish Orconectes virilis, the red swamp crayfish Procambarus clarkii and the
parastacid Australian redclaw crayfish Cherax quadricarinatus. The presence of
O. virilis, native to the north-central portion of the United States, was first reported in
Mexico by Campos and Contreras (1985) based on samples collected near the town
of Meoqui, Chihuahua in 1982. Samples of O. virilis from a stream near the town of
Cuauhtemoc, Chihuahua, were collected in 2009 and are deposited in the National

© 2016 by Taylor & Francis Group, LLC

 The Crayfish of Middle America 461

Crustacean Collection (CNCR) of the Institute of Biology at UNAM. This latter

record, 27 years after the first one, suggest that the species has become established in
the northern portion of Chihuahua.
The red swamp crayfish is probably the most widely introduced species of
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freshwater invertebrate in the world. It is native to the southern United States from
Texas, Louisiana, from Florida to Ohio, and from southern Illinois, and to northern
Mexico in northern Tamaulipas and Nuevo Leon (Hobbs 1972, Rodríguez-Almaraz and
Muñiz 2008). Although native to northern Mexico, it has been introduced in Durango,
Sonora, Baja California and Chiapas (Hernandez et al. 2008, Alvarez et al. 2011).
Torres and Alvarez (2012) found that populations within the native range were more
similar genetically to each other than to introduced populations in Mexico and Costa
Rica, although overall genetic variation was low.
The Australian redclaw crayfish was brought into Mexico in 1995 to start
experimental cultures and a few years later commercial farms were established to culture
them in Morelos and Tamaulipas. By 2007 the farms had been closed at least for two
years; however, wild populations of C. quadricarinatus were found in the proximity
to them, suggesting that these were the sources for the invasive populations (Bortolini
et al. 2007). In Morelos the C. quadricarinatus population is slowly spreading out
from Las Estacas, an aquatic park near the town of Tlaltizapan, to the south along the
Yautepec River, which is a tributary of the Balsas River. In Tamaulipas, the redclaws
have spread about 185 km from the town of Llera de Canales to the coastal lagoon
system in the Port of Tampico, mainly along the Tamesi River basin (CNCR records).
An additional occurrence of redclaws in the Media Luna Lagoon, San Luis Potosi, was
recorded in 2013 (CNCR records). Mendoza-Alfaro et al. (2011) reviewed the status of
the introduction of C. quadricarinatus in Mexico; however, no studies have focused on
the impact that this exotic species is having on the native fauna, especially in Tamaulipas
and San Luis Potosi where an important diversity of native crayfish occurs.

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