New records of anchialine fauna from the Yucatan Peninsula, Mexico

Abstract

New records for 17 species of crustaceans from anchialine systems in the Yucatan Peninsula, Mexico, are presented. The records come from explorations in Dzilam de Bravo, Yucatan, and from Puerto Aventuras and the Nohoch Nah Chich and Ox Bel Ha cave systems near Tulum in Quintana Roo, Mexico. For five of the 17 species dealt with here, the records presented constitute the second time those species are reported after their original descriptions. For the alpheid shrimp Yagerocaris cozumel, we present the first record of the species for continental Yucatan and for the atyid shrimp Jonga serrei, the second record from Mexico. Depth data are provided for all species.

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Check List the journal of biodiversity data
10TH ANNIVERSARY ISSUE
Check List | www.biotaxa.org/cl Volume 11 | Number 1 | Article 1505 1
New records of anchialine fauna from the Yucatan Peninsula,
Mexico
Fernando Alvarez1*
, Thomas M. Iliffe2, Sergio Benitez1, David Brankovits2 and José Luis Villalobos1
1 Colección Nacional de Crustáceos, Instituto de Biología, Universidad Nacional Autónoma de México, Apartado Postal 70-153, México 04510 D.F., México
2 Texas A&M University at Galveston, Department of Marine Biology, 200 Seawolf Parkway, OCSB #251, Galveston, Texas 77553, USA
* Corresponding author. E-mail: falvarez@unam.mx
Check List 11(1): 1505, January 2015 doi: http://dx.doi.org/10.15560/11.1.1505
ISSN 1809-127X © 2015 Check List and Authors
Abstract: New records for  species of crustaceans from
anchialine systems in the Yucatan Peninsula, Mexico, are
presented. e records come from explorations in Dzilam de
Bravo, Yucatan, and from Puerto Aventuras and the Nohoch
Nah Chich and Ox Bel Ha cave systems near Tulum in Quin-
tana Roo, Mexico. For ve of the  species dealt with here, the
records presented constitute the second time those species
are reported after their original descriptions. For the alpheid
shrimp Yagerocaris cozumel, we present the rst record of
the species for continental Yucatan and for the atyid shrimp
Jonga serrei, the second record from Mexico. Depth data are
provided for all species.
Key words: Crustacea, anchialine, cenote, sinkhole, Yucatan,
Quintana Roo
INTRODUCTION
e anchialine fauna of the Yucatan Peninsula, composed
mainly of crustacean species, has been intensely studied for 
years since the diving explorations of the Nohoch Nah Chich
and Sac Actun systems in Quintana Roo began in  (Ger-
rard ). However, a few widely distributed species in the
peninsula were described years before (e.g., Creaseria morleyi
and Typhlatya pearsei in ) because they could be captured
from siphons inside caves with a dry initial section like Balan
Canche Cave in Yucatan (Creaser ). Several publications
have summarized what is known about the anchialine fauna
of the Yucatan in dierent moments (Reddell ; Ilie ,
; Alvarez and Ilie ); in most cases a large proportion
of the information has come from previous studies with very
few new records appearing in each new occasion. For ve species
dealt with here the only previous records were those contained
in the original description (i.e., Tuluweckelia cernua, Holsinger
; Metacirolana mayana, Bowman ; Typhlatya dzilamensis,
Alvarez et al. ; Yagerocaris cozumel, Kensley ; Calliasmata
nohochi, Escobar-Briones et al. ); and for one species, the
atyid Jonga serrei (Bouvier ), the record presented in this
contribution is the second one for Mexico.
We present herein new records for  species from  cenotes
(Figure ). Although the distribution data are important to bet-
ter dene particular distribution ranges, these records, mostly
obtained during , are evidence of the current presence of the
species in a region that is being rapidly developed. e northeast-
ern portion of the Yucatan Peninsula, where the Cancún-Tulum
tourist corridor known as the “Riviera Maya” is located, is also
the area where the largest anchialine systems are found. e
rapid growth of the tourist infrastructure may compromise in
the near future the preservation of these underground reser-
voirs. e northern coast of the peninsula is also being rapidly
developed from the Port of Progreso to the east, where a number
of cenotes are located. e periodic monitoring of anchialine
species will become increasingly important in the future as the
present period of intense urban development continues and
compromises the availability of freshwater that is drawn from
the same aquifer that constitutes the anchialine systems.
MATERIALS AND METHODS
Collection of organisms was conducted by cave divers
using plankton nets or individually with glass vials. A Hydro-
lab Multiparameter Sonde or a YSI Multiparameter Water
Quality Analyzer were used to obtain water column proles
of temperature, salinity, pH and dissolved oxygen concentra-
tion. e depth at which every specimen was collected was
recorded; all vials were numbered and after an organism was
caught the depth was recorded for that vial, plankton tows
were also conducted at depths that divers recorded above an
below the halocline.
e localities where organisms were collected are referred
to as “cenotes” or sinkholes; however, all organisms were col-
lected from the aphotic (dark) zone of the submerged caves
associated to each one of the cenotes mentioned. e records
are arranged chronologically. Species identication was done
with the aid of stereomicroscopes and in some cases using a
compound microscope too. All specimens were collected under
the scientic collector’s license issued to FA (FAUT ) by
the Mexican environmental authority (SEMARNAT). All
organisms are deposited in the National Crustacean Collec-
tion (CNCR) of the Institute of Biology, UNAM, Mexico City.
RESULTS
A summary of the species recorded, the number of speci-
mens per species, the number of cenotes they were found in
and if they were collected in freshwater or saltwater appears
in Table . Following is an individual account for each species
recorded.
LISTS OF SPECIES

Alvarez et al. | Anchialine fauna from Yucatan
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Figure 1. Map of the Yucatan Peninsula showing sites of collection: 1, Cenote Bang; 2, Cenote Muknal; 3, Cenote Odyssey; 4, Cenote Na’ach Wennen Ha;
5, Cenote Nohoch Nah Chich; 6, Cenote Eden; 7, Cenote Cervera.
Table 1. List of crustacean species and number of specimens collected from cenotes (caves) in the Yucatan Peninsula of Mexico. CB, Cenote Bang; CM,
Cenote Muknal; CO, Cenote Odyssey; CW, Cenote Na’ach Wennen Ha; CN, Cenote Nohoch Nah Chich; CE, Cenote Eden; CC, Cenote Cervera.
Taxa Salinity CB CM CO CW CN CE CC Total Specimens
Remipedia         
Xibalbanus tulumensis (Yager, 1987) SW 1 2 2 5
Ostracoda         
Humphreysella mexicana(Kornicker and Ilie, 1989) SW 3 1 4
Thermosbaenacea         
Tulumella unidens Bowman and Ilie, 1988 FW 6 9 6 22 43
Mysida         
Antromysis cenotensis Creaser, 1936 FW 4 10 14
Stygiomysis cokei Kallmeyer and Carpenter, 1996 FW 1 7 8
Stygiomysis cf. holthuisi (Gordon, 1958) FW 2 1 1 4 8
Amphipoda         
Mayaweckelia cenoticolaHolsinger, 1977 FW 1 1
Tuluweckelia cernuaHolsinger, 1990 FW 4 1 3 10 18
Isopoda         
Metacirolana mayana (Bowman, 1987) SW 3 6 9
Creaseriella anops (Creaser, 1936) FW 6 8 3 1 18
Decapoda         
Jonga serrei (Bouvier, 1909) FW 7 7
Typhlatya dzilamensis Alvarez, Ilie and Villalobos, 2005 SW 2 2
Typhlatya mitchelli Hobbs and Hobbs, 1976 FW 5 10 9 11 35
Typhlatya pearseiCreaser, 1936 FW/SW 1 6 5 12
Yagerocaris cozumel Kensley, 1988 — 1 1
Creaseria morleyi (Creaser, 1936) FW 1 2 1 4
Calliasmata nohochi Escobar-Briones, Camacho and Alcocer, 1997 SW 1 1
Subphylum Crustacea
Class Remipedia
Order Nectiopoda
Family Speleonectidae
Xibalbanus tulumensis (Yager, )
(Figure )
M :  organisms; Cenote Muknal,  m
depth, Tulum, Quintana Roo, Mexico;  July , colls. T.M.
Ilie, D. Brankovits; CNCR , .  organism, Cenote
Bang,  m depth, Tulum, Quintana Roo, Mexico;  August
; colls. T.M. Ilie, D. Brankovits; CNCR .  organ-
isms; Cenote Odyssey, .– m depth, Tulum, Quintana
Roo, Mexico;  August ; colls. T.M. Ilie, D. Brankovits;
CNCR , .
D: e known range of the species is from Cenote
Crustacea, south of Puerto Morelos, to cenotes around Tulum,
Quintana Roo, Mexico (Yager ; Alvarez and Ilie ).

Alvarez et al. | Anchialine fauna from Yucatan
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R: Previously known as Speleonectes tulumensis
Yager, , this species has now been placed in the new genus
Xibalbanus based on a molecular phylogenetic analysis of the
entire class Remipedia (Hoenemann et al. ). It occurs at
depths below  m, beneath the halocline, at salinities above
 ppt. e records presented herein correspond to cenotes 
to  km inland from the coastline.
e species is listed in the Mexican Red List of reatened
Species (Semarnat ) as in danger of extinction; although
it can be relatively common in a few cenotes, its distribution
range is restricted to some cenotes in the Puerto Morelos–
Tulum area.
Class Ostracoda
Family aumatocyprididae
Humphreysella mexicana(Kornicker & Ilie, )
(Figure )
M :  organism; Cenote Muknal,  m
depth, Tulum, Quintana Roo, Mexico;  February ;
colls. T.M. Ilie, D. Brankovits; CNCR .  organisms;
Cenote Bang, – m depth, Tulum, Quintana Roo, Mexico;
 February , coll. D. Brankovits; CNCR , . 
organism; Cenote Bang,  m depth, Tulum, Quintana Roo,
Mexico;  August , coll. T.M. Ilie; CNCR .
D: e type locality is Cenote Cristal, Tulum,
Quintana Roo, Mexico, but also known from Cenote Pon-
derosa in Puerto Aventuras, and Cenote  Steps in Akumal,
Quintana Roo, Mexico (Kornicker and Ilie ).
R: Previously known as Danielopolina mexicana,
this species was reassigned to Humphreysella together with
other species from around the Caribbean (Iglikowska and
Boxshall ). e present records expand the presence of
this species into the Ox Bel Ha System, showing that it prefers
moderate salinities near the halocline and cenotes that are 
to  km inland from the coastline.
Class Malacostraca
Superorder Peracarida
Order ermosbaenacea
Family Tulumellidae
Tulumella unidens Bowman & Ilie, 
(Figure )
M :  organisms; Cenote Muknal, depth
 m, Tulum, Quintana Roo, Mexico;  February ; colls.
T.M. Ilie, D. Brankovits, B. Phillips; CNCR , . 
organisms; Cenote Na’ach Wennen Ha, – m depth, Tulum,
Quintana Roo, Mexico;  February ; colls. T.M. Ilie, D.
Brankovits, B. Phillips; CNCR -, , . 
organisms; Cenote Bang, depth – m, Tulum, Quintana
Roo, Mexico;  February ; colls. T.M. Ilie, D. Brankov-
its, B. Phillips; CNCR , , , .  organisms;
Cenote Muknal, depth  m; Tulum, Quintana Roo, Mexico;
 July ; colls. T.M. Ilie, D. Brankovits, T. Winkler;
CNCR , .  organism, Cenote Odyssey, depth  m,
Tulum, Quintana Roo, Mexico;  August ; colls. T.M. Ilie,
D. Brankovits, T. Winkler; CNCR .  organisms; Cenote
Na’ach Wennen Ha, .–. m depth, Tulum, Quintana Roo,
Mexico;  August ; colls. T.M. Ilie, D. Brankovits, T. Win-
kler; CNCR , , , , .  organisms;
Cenote Muknal, depth .– m; Tulum, Quintana Roo,
Mexico;  December ; colls. T.M. Ilie, D. Brankovits, T.
Winkler; CNCR , , , , .  organ-
isms; Cenote Bang, depth – m, Tulum, Quintana Roo,
Mexico;  December ; colls. T.M. Ilie, D. Brankovits, T.
Winkler; CNCR , .  organisms; Cenote Odyssey,
depth .–. m, Tulum, Quintana Roo, Mexico;  December
; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR ,
, .  organisms; Cenote Na’ach Wennen Ha, depth
–. m, Tulum, Quintana Roo, Mexico;  December ;
colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR ,
.
D: Cenotes Cristal, Calavera and Actun Ha,
Tulum, Quintana Roo, Mexico (Bowman and Ilie ; Alva-
rez and Ilie ).
R: is species is locally abundant in cenotes
around the town of Tulum, and is present throughout the Ox
Bel Ha System in cenotes that range from  to  km from
the coastline. Inside the caves, T. unidens is found at depths
ranging from . to  m, always above the halocline.
Order Mysida
Family Mysidae
Antromysis cenotensis Creaser, 
(Figure )
M :  organisms; Cenote Muknal, depth
– m, Tulum, Quintana Roo, Mexico;  February ;
coll. T.M. Ilie; CNCR , , , .  organisms;
Cenote Odyssey, depth  m, Tulum, Quintana Roo, Mexico;
 February , coll. T.M. Ilie, D. Brankovits, B. Phillips;
CNCR , .  organisms; Cenote Odyssey, depth .
m, Tulum, Quintana Roo, Mexico;  August ; colls. T.M.
Ilie, D. Brankovits, T. Winkler; CNCR -.
D: Widely distributed in the central and
northern portions of the Yucatan Peninsula (Reddell ;
Alvarez and Ilie ). Present in Quintana Roo from a
number of cenotes in Sian Ka’an and the Tulum-Cobá region.
In Yucatan it has been recorded from Grutas de Balancanché,
Tinúm; and a number of cenotes that form the ring of cenotes
around the city of Merida (Reddell ; Pérez-Aranda c).
R: Although this species is listed in the Mexican
Red List of reatened Species (Semarnat ) as threatened,
it is widely distributed and very abundant, often forming
swarms of hundreds of organisms. A. cenotensis occurs prefer-
entially above the halocline, from the surface to  m depths;
however, it can occasionally be found below the halocline in
moderate salinities.
Family Stygiomysidae
Stygiomysis cokei Kallmeyer & Carpenter, 
(Figure )
M :  organisms; Cenote Na’ach Wennen
Ha, depth – m, Tulum, Quintana Roo, Mexico;  February
; colls. T.M. Ilie, D. Brankovits, B. Phillips; CNCR ,
, .  organism; Cenote Muknal, depth . m,
Tulum, Quintana Roo, Mexico;  July ; coll. T.M. Ilie;
CNCR .  organisms; Cenote Na’ach Wennen Ha, depth
–. m, Tulum, Quintana Roo, Mexico;  August ; colls.

Alvarez et al. | Anchialine fauna from Yucatan
Check List | www.biotaxa.org/cl Volume 11 | Number 1 | Article 1505 4
Figure 2. A, Xibalbanus tulumensis (Remipedia); B, Humphreysella mexicana (Ostracoda); C, Tulumella unidens (Thermosbaenacea); D, Antromyisis cenoten-
sis (Mysida); E, Stygiomysis cokei (Mysida); F, Stygiomysis cf. holthuisi (Mysida); G, Mayaweckelia cenoticola (Amphipoda); Tuluweckelia cernua (Amphipoda).

Alvarez et al. | Anchialine fauna from Yucatan
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T.M. Ilie, D. Brankovits, T. Winkler; CNCR , ,
.
D: In Quintana Roo: Cenotes Calavera, Escon-
dido, Actun Ha, Cristal and Actun Ko; in Yucatan: Cenotes
Pabakal, San Eduardo, Kankirixché, Mucuyché and Dzonotila
(Pesce and Ilie ).
R: e present records show that S. cokei occurs in
low salinities (– ppt) above the halocline in cenotes  to 
km inland from the coastline.
Stygiomysis cf. holthuisi (Gordon, )
(Figure )
M :  organisms; Cenote Na’ach Wennen
Ha, depth – m, Tulum, Quintana Roo, Mexico;  February
; colls. T.M. Ilie, D. Brankovits, B. Phillips; CNCR ,
, .  organisms; Cenote Bang, depth  m, Tulum,
Quintana Roo, Mexico;  February ; colls. T.M. Ilie, D.
Brankovits, B. Phillips; CNCR , .  organism; Cenote
Muknal, depth . m, Tulum, Quintana Roo, Mexico;  July
; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR .
 organism; Cenote Odyssey, depth . m, Tulum, Quintana
Roo, Mexico;  August ; colls. T.M. Ilie, D. Brankovits,
T. Winkler; CNCR .  organism; Cenote Na’ach Wennen
Ha, depth . m, Tulum, Quintana Roo, Mexico;  August
; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR .
D: Stygiomysis holthuisi was described from
material collected in St. Martin, Lesser Antilles (Gordon
), and was subsequently reported from Puerto Rico,
Anguilla and the Bahamas (Bowman et al. ). Pesce and
Ilie () reported the species from cenotes in Quintana
Roo (Escondido and Casa Cenote) and Yucatan (Mucuyché),
Mexico.
R: e organisms identied as Stygiomysis holthuisi
from the Yucatan Peninsula agree well, but not completely,
with the description of the species. Pesce and Ilie () had
already pointed out several dierences seen in the Yucatan
organisms, to which we could add a larger uropod length
in relation to the telson length. It seems very possible that
a new species will be erected to accommodate the Yucatan
specimens.
Order Amphipoda
Family Hadziidae
Mayaweckelia cenoticolaHolsinger, 
(Figure )
M :  organism; Cenote Odyssey, depth
. m, Tulum, Quintana Roo, Mexico;  December ; coll.
T.M. Ilie; CNCR .
D: Along the northern portion of the Yucatan
Peninsula (Holsinger ; Reddell ).
R: Although this species has been regarded as com-
mon in other parts of the Yucatan Peninsula, it is rare in the
Ox Bel Ha System, appearing only once at moderate depth in
freshwater.
Tuluweckelia cernuaHolsinger, 
(Figure )
M :  organisms; Cenote Na›ach Wen-
nen Ha, depth – m, Tulum, Quintana Roo, Mexico; 
February ; colls. T.M. Ilie, D. Brankovits; CNCR ,
, , , , .  organisms; Cenote Bang,
depth – m, Tulum, Quintana Roo, Mexico;  February
; coll. T.M. Ilie; CNCR , .  organism; Cenote
Muknal, depth . m, Tulum, Quintana Roo, Mexico; 
July ; colls. T.M. Ilie; CNCR .  organisms; Cenote
Bang, depth – m, Tulum, Quintana Roo, Mexico;  August
; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR ,
.  organism; Cenote Odyssey, depth . m, Tulum,
Quintana Roo, Mexico;  August ; coll. D. Brankovits;
CNCR .  organisms; Cenote Na›ach Wennen Ha, depth
-. m, Tulum, Quintana Roo, Mexico;  August ; colls.
T.M. Ilie, D. Brankovits, T. Winkler; CNCR , ,
.  organisms; Cenote Odyssey, depth .–. m, Tulum,
Quintana Roo, Mexico;  December ; colls. T.M. Ilie, D.
Brankovits; CNCR , .
D: In Quintana Roo in the following cenotes
around Tulum: Calavera, Aktun Ha, Escondido, Mojarra and
Cristal (Holsinger ).
R: e records presented herein are the rst to be
published about T. cernua after Holsinger (). e species is
common in the Ox Bel Ha System; it is clearly a freshwater spe-
cies occurring always above the halocline to depths of  m.
Order Isopoda
Family Cirolanidae
Metacirolana mayana (Bowman, )
(Figure )
M :  organism; Cenote Bang, depth 
m, Tulum, Quintana Roo, Mexico;  February , coll. T.M.
Ilie; CNCR .  organisms; Cenote Bang, depth – m,
Tulum, Quintana Roo, Mexico;  August ; colls. T.M. Ilie,
D. Brankovits; CNCR , .  organisms; Cenote Odys-
sey, depth – m, Tulum, Quintana Roo, Mexico;  August
; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR ,
, .  organisms; Cenote Odyssey, depth  m,
Tulum, Quintana Roo, Mexico;  December ; colls. T.M.
Ilie, D. Brankovits, T. Winkler; CNCR , , .
D: Known from Quebrada Cave in Chank-
anaab Park, Cozumel and Cenote Calavera, Tulum, Quintana
Roo, Mexico (Bowman ).
R: e records presented herein are the rst to be
published after Bowman () original description of the
species. Metacirolana mayana was always collected at or below
the halocline in salinities of  ppt or higher.
Creaseriella anops (Creaser, )
(Figure )
M :  organisms; Cenote Muknal, - m
depths, Tulum, Quintana Roo, Mexico;  February ; colls.
T.M. Ilie, D. Brankovits, B. Phillips; CNCR , , ,
, , , .  organisms; Cenote Odyssey, depth
– m, Tulum, Quintana Roo, Mexico;  February ; colls.
T.M. Ilie, D. Brankovits; CNCR , .  organim; Cenote
Na’ach Wennen Ha, depth  m, Tulum, Quintana Roo, Mexico;
 February , coll. D. Brankovits; CNCR .  organisms;
Cenote Bang, depth – m, Tulum, Quintana Roo, Mexico; 
February ; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR
, , , , .  organism; Cenote Bang,

Alvarez et al. | Anchialine fauna from Yucatan
Check List | www.biotaxa.org/cl Volume 11 | Number 1 | Article 1505 6
Figure 3. A, Metacirolana mayana (Isopoda); B, Creaseriella anops (Isopoda); C, Jonga serrei (Atyidae); D, Typhlatya dzilamensis (Atyidae); E, Typhlatya
mitchelli (Atyidae); F, Typhlatya pearsei (Atyidae); G, Yagerocaris cozumel (Alpheidae); H, Creaseria morleyi (Palaemonidae); I, Calliasmata nohochi (Hip-
polytidae).

Alvarez et al. | Anchialine fauna from Yucatan
Check List | www.biotaxa.org/cl Volume 11 | Number 1 | Article 1505 7
depth  m, Tulum, Quintana Roo, Mexico;  August ; coll. D.
Brankovits; CNCR .  organism; Cenote Odyssey, depth 
m, Tulum, Quintana Roo, Mexico;  August ; coll. T.M. Ilie;
CNCR .  organism; Cenote Muknal, depth  m, Tulum,
Quintana Roo, Mexico;  December ; coll. D. Brankovits;
CNCR .
D: roughout the central and northern por-
tions of the Yucatan Peninsula, Mexico (Pérez-Aranda b;
Alvarez and Ilie ).
R: Creaseriella anops is listed in Mexican Red List of
reatened Species (Semarnat ) as threatened; however,
the species is usually common within its range and depending
on the sampling method, it can be abundant. It was present in
all cenotes studied in the Ox Bel Ha System.
Order Decapoda
Family Atyidae
Jonga serrei (Bouvier, )
(Figure )
M :  organisms; Cenote Nohoch Nah
Chich, depth - m, Quintana Roo, Mexico;  June ;
colls. T.M. Ilie, J.L. Villalobos, S. Benitez; CNCR .
D: Cuba, Jamaica, Puerto Rico, Dominica,
Barbados, Costa Rica and Mexico (Chace and Hobbs ;
Alvarez and Ilie ).
R: Alvarez and Ilie () rst reported this spe-
cies from Cenote del Mar, south of Tulum, Quintana Roo. e
present record is the second for the species in Mexico. Although
considered to be a coastal species, the two records from Mexico
are from caves. e organisms collected are pigmented with nor-
mal eyes, not showing any obvious adaptation to cave life, and, as
pointed out by Alvarez and Ilie (), the Mexican organisms
are larger ( mm of total length (TL)) than those previously
reported from around the Caribbean ( mm TL). In Cenote
Nohoch Nah Chich, this species occurs in the cave entrance in
semi-darkness and inside the cave.
Typhlatya dzilamensis Alvarez, Ilie & Villalobos, 
(Figure )
M :  organisms; Cenote Cervera, depth
– m, Dzilam de Bravo, Yucatan, Mexico;  December ;
colls. T.M. Ilie, D. Brankovits; CNCR .
D: Only known from two cenotes in the town
of Dzilam de Bravo, Yucatan and one from Buya Uno, a marine
cave  m oshore from the town in the Gulf of Mexico
(Alvarez et al. ).
R: e record presented herein represents the sec-
ond time the species is collected. e rst collections of the
species used to describe it were made in  (Alvarez et al.
). Typhlatya dzilamensis was collected again in the type
locality at depths ranging between - m in fully marine
water, while T. mitchelli was also found at depths shallower
than  m in freshwater.
Typhlatya mitchelli Hobbs & Hobbs, 
(Figure )
M :  organisms; Cenote Muknal, depth
– m, Tulum, Quintana Roo, Mexico;  February ;
colls. T.M. Ilie, D. Brankovits, B. Phillips; CNCR , ,
, , .  organisms; Cenote Odyssey, depth 
m, Tulum, Quintana Roo, Mexico;  February ; colls.
T.M. Ilie, D. Brankovits, B. Phillips; CNCR , . 
organisms; Cenote Na’ach Wennen Ha, depth – m, Tulum,
Quintana Roo, Mexico;  February , colls. T.M. Ilie, D.
Brankovits; CNCR , , , .  organisms,
Cenote Na’ach Wennen Ha, depth – m, Tulum, Quintana
Roo, Mexico;  February ; colls. T.M. Ilie, D. Branko-
vits, B. Phillips; CNCR , .  organisms; Cenote
Muknal, depth – m, Tulum, Quintana Roo, Mexico; 
July ; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR
, , .  organisms; Cenote Bang, depth –
m, Tulum, Quintana Roo, Mexico;  August ; colls. T.M.
Ilie, D. Brankovits; , .  organisms; Cenote
Odyssey, depth .–. m, Tulum, Quintana Roo, Mexico;
 August ; colls. T.M. Ilie, D. Brankovits, T. Winkler;
CNCR , , , , .  organisms; Cenote
Na’ach Wennen Ha, depth .–. m, Tulum, Quintana
Roo, Mexico;  August ; colls. T.M. Ilie, D. Brankovits,
T. Winkler; CNCR , , , .  organisms;
Cenote Muknal, depth .–. m, Tulum, Quintana Roo,
Mexico;  December ; colls. T.M. Ilie, D. Brankovits;
CNCR , .  organisms; Cenote Bang, depth – m,
Tulum, Quintana Roo, Mexico;  December ; colls. T.M.
Ilie, D. Brankovits, T. Winkler; CNCR , , .
 organisms; Cenote Odyssey, depth – m, Tulum, Quintana
Roo, Mexico;  December ; colls. T.M. Ilie, D. Brankovits;
CNCR , .  organism; Cenote Na’ach Wennen Ha,
depth . m, Tulum, Quintana Roo, Mexico;  December ;
coll. T Winkler; CNCR .
D: roughout the Yucatan Peninsula, from
the western portion at Maxacanú, south of Merida, Yucatan,
to the cenotes between Akumal and Tulum, Quintana Roo;
to the north it occurs near Tizimin, Yucatan, Mexico (Pérez-
Aranda a; Alvarez and Ilie ).
R: is species is listed in the Mexican Red List of
reatened Species (Semarnat ) as threatened; however,
similar to other anchialine species, it is very common within
its range. It occurs from the surface in cave pools to depths
of  m, always in freshwater. T. mitchelli is considered as a
common species in the Ox Bel Ha System.
Typhlatya pearseiCreaser, 
(Figure )
M :  organisms; Cenote Odyssey, depth
– m, Tulum, Quintana Roo, Mexico;  February ;
colls. T.M. Ilie, D. Brankovits, B. Phillips; CNCR , .
 organism; Cenote Na›ach Wennen Ha, depth  m, Tulum,
Quintana Roo, Mexico;  February ; coll. D. Brankovits;
CNCR .  organism; Cenote Bang, depth  m, Tulum,
Quintana Roo, Mexico;  February ; coll. D. Brankovits;
CNCR .  organisms; Cenote Na›ach Wennen Ha, depth
.-. m, Tulum, Quintana Roo, Mexico;  August ; colls.
T.M. Ilie, D. Brankovits; CNCR , .  organisms;
Cenote Odyssey, depth – m, Tulum, Quintana Roo, Mex-
ico; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR ,
, , .  organisms; Cenote Na›ach Wennen Ha,
depth .–. m, Tulum, Quintana Roo, Mexico;  December
; colls. T.M. Ilie, D. Brankovits, T. Winkler; CNCR ,
.

Alvarez et al. | Anchialine fauna from Yucatan
Check List | www.biotaxa.org/cl Volume 11 | Number 1 | Article 1505 8
D: Widely distributed in the northern por-
tion of the Yucatan Peninsula. It occurs from Champoton,
Campeche, to Telchac Puerto in the northern coast of Yucat-
an, to the cenotes in the area of Tulum and Akumal, Quintana
Roo (Perez-Aranda b).
R: is species is listed in Mexican Red List of
reatened Species (Semarnat ) as threatened; however,
this is the species of Typhlatya in the Yucatan Peninsula with
the largest distribution range. It occurs in freshwater above
the halocline to depths of  m.
Family Alpheidae
Yagerocaris cozumel Kensley, 
(Figure )
M :  organism; Cenote Eden, Soli-
daridad, Quintana Roo, Mexico;  July ; coll. T.M. Ilie;
CNCR .
D: Previously known from Cenote Aerolito
which is the type locality of the species, and Cueva la Que-
brada, Chankanaab Park; both sites in the island of Cozumel,
Quintana Roo, Mexico (Kensley ).
R: e present record extends the distribution of the
species to continental Yucatan, since it was previously consid-
ered an insular species from the island of Cozumel (Alvarez and
Ilie ). e present record is also the rst one to appear
since the description of the species by Kensley ().
Family Palaemonidae
Creaseria morleyi (Creaser, )
(Figure )
M :  organism; Cenote Bang, depth 
m, Tulum, Quintana Roo, Mexico;  February ; coll. D.
Brankovits; CNCR .  organisms; Cenote Muknal, depth
– m, Tulum, Quintana Roo, Mexico;  July ; colls.
T.M. Ilie, D. Brankovits; CNCR , .  organism;
Cenote Na’ach Wennen Ha, depth  m, Tulum, Quintana
Roo, Mexico;  August ; coll. T.M. Ilie; CNCR .
D: Widely distributed in the northern section
of the Yucatan Peninsula in Yucatan and Quintana Roo. e
species has been recorded from many cenotes composing the
“Ring of Cenotes” around the City of Merida, but also from
the cenotes along the coast of Quintana Roo (Perez-Aranda
a; Botello and Alvarez , ).
R: is species is listed in Mexican Red List of
reatened Species (Semarnat ) as threatened. C. mor-
leyi is one of the species with the widest distribution in the
Yucatan Peninsula, it occurs mostly in freshwater at shallow
depths to  m.
Family Hippolytidae
Calliasmata nohochi Escobar-Briones, Camacho & Alcocer, 
(Figure )
M :  organism; Cenote Muknal, depth
 m, Tulum, Quintana Roo, Mexico;  July ; coll. T.M.
Ilie; CNCR .
D: Previously known only from two caves:
Cenote Crack House (type locality), in the Nohoch Nah Chich
System; and Cenote Escondido, in the Ox Bel Ha System.
R: is is a very rare species that has been collected
only three times, the material from the rst two collections is
mentioned in the description of the species (Escobar-Briones
et al. ). e new record comes from Cenote Muknal,
which is also part of the Ox Bel Ha System, to the southeast
of Cenote Escondido.
DISCUSSION
Until now,  anchialine species of crustaceans from the
Yucatan Peninsula have been recognized (Alvarez and Ilie
; Alvarez et al. ; Neiber et al. ; Boxshall et al.
); other anchialine fauna include an undescribed species
of gastropod, one or possibly two echinoderms (Solís-Marín
and Laguarda-Figueras ) and two species of sh (Alvarez
and Ilie ). We estimate that , out of the known 
crustacean species, could have been collected during the pres-
ent survey as they are widely distributed in Quintana Roo or
have more ample distribution ranges. e other  species are
known only from their type locality in the states of Yucatan
and Campeche or are restricted to the island of Cozumel. We
report on  species,  of the possible  species, however
no copepods are included in this paper as they will be treated
in an upcoming contribution on anchialine plankton.
Regarding the identity of the organisms collected, all could
be readily identied to species, except the mysid shrimp
referred to here as Stygiomysis cf. holthuisi. In this case,
several characters do not agree completely with the original
description; a comparative study with samples from other
locations in the Yucatan Peninsula and from around the
Caribbean region would be necessary to determine if this is a
wide ranging species with a signicant degree of morphologi-
cal variation or a species complex. In the case of Jonga serrei,
even when the closest records are those from Cuba and Costa
Rica, no signicant variation from the description was noted.
Our results conrm a wide distribution for several species,
a pattern that is consistent with the degree of connectivity
that has become apparent in the anchialine systems of the
eastern portion of Quintana Roo, where cenotes, kilometers
of passageways, and new connections between systems, are
being discovered and explored each year (Bauer-Gottwein et
al. ). e records presented herein expand the distribution
ranges of  of the  reported species and represent for ve
species the second time they are collected after their original
description. Six of the  species reported herein are included
in the Mexican Red List of reatened Species (Semarnat
). Considering the new records, the hadziid Tuluweckelia
cernua, the cirolanid Metacirolana mayana, the atyid Typhlatya
dzilamensis, the alpheid Yagerocaris cozumel, and the hippoly-
tid Calliasmata nohochi, should be included in the Red List due
to their conrmed reduced distribution range and/or to the
low numbers that have been collected that suggest very small
natural populations.
ACKNOWLEDGMENTS
e rst author gratefully acknowledges the funding
awarded through CONACYT grant () “Processes
that create and maintain the biodiversity in an extreme
environment: the anchialine systems of Yucatan”. Additional
funding was provided to the rst two authors through a Texas

Alvarez et al. | Anchialine fauna from Yucatan
Check List | www.biotaxa.org/cl Volume 11 | Number 1 | Article 1505 9
A&M University — CONACYT collaborative science grant
 “What promotes species diversication in anchialine
habitats?”. S. Benitez acknowledges the support received
through a CONACYT scholarship. D. Brankovits sincerely
thanks for the support of the Ruord Foundation and the
National Speleological Society’s Ralph W. Stone Scholarship.
We sincerely appreciate the assistance provided by Texas
A&M University students and sta Tyler Winkler and Brett
Gonzalez, UNAM student Olinka Cortés and local cave divers
Bil Phillips and Robbie Schmittner.
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starsh (Echinodermata: Asteroidea) from an anchialine cave
in the Mexican Caribbean. Revista Mexicana de Biodiversidad :
– (http://www.scielo.org.mx/scielo.php?script=sci_artte
xt&pid=S-).
Yager, J. . Speleonectes tulumensis, n. sp. (Crustacea, Remipedia)
from two anchialine cenotes of the Yucatan Peninsula, Mexico.
Stygologia : –.
Authors’ contribution statement: All authors made the species
identications; FA, TMI and JLV prepared the text; TMI, DB and SB
conducted the eld sampling; FA and SB prepared the gures.
Received: August 
Accepted: November 
Editorial responsibility: Luis E. ArrudaBezerra