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Revista de Biología Tropical, ISSN: 2215-2075, Vol. 73(S1): e63625, enero-diciembre 2025 (Publicado Mar. 03, 2025)
Diversity of freshwater crab: new distributional data for four species
(Decapoda: Pseudothelphusidae) from Meso- and South American countries
Célio Magalhães 1,2; https://orcid.org/0000-0003-4858-2575
Ingo S. Wehrtmann 3,4; https://orcid.org/0000-0002-6826-7938
1. Laboratory of Bioecology and Crustacean Systematics (LBSC), Department of Biology, Faculty of Philosophy,
Sciences and Letters at Ribeirão Preto (FFCLRP), University of São Paulo (USP), São Paulo, Brazil; celiomag@usp.br
(*Correspondence)
2. Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Brazil.
3. Museo de Zoología of the Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa
Rica, San José, Costa Rica; ingo.wehrtmann@ucr.ac.cr
4. Centro de Investigación en Ciencias del Mar y Limnología (CIMAR), Universidad de Costa Rica, San José, Costa Rica.
Received 07-III-2024. Corrected 30-VIII-2024. Accepted 24-I-2025.
ABSTRACT
Introduction: The Pseudothelphusidae is a very diverse family of primary freshwater crabs widely distributed
throughout the Neotropical region, but the true extent of the geographic distribution of several species is poorly
known because many of them occur in very remote areas with only a single or very few records available.
Objective: Here we present new country records for four species of pseudothelphusid crabs from Meso- and
South American countries.
Methods: The data about the geographic distribution are based on specimens deposited in crustacean collections
of institutions from Brazil, Germany, and USA; illustrations of the male first gonopod were prepared in stereo-
microscope equipped with a camera lucida.
Results: Occurrence records of Phygiopilus acanthophallus from Mexico, Fredius stenolobus from Guyana, and
Kunziana irengis from Brazil are published for the first time, and the occurrence of Raddaus mertensi from
Honduras is confirmed. Comments on the morphology of the male first gonopod of each species are given,
their range extensions are discussed, and the importance of these new records for conservation assessments is
highlighted.
Conclusions: The new data expand the area of occurrence of these four species of Pseudothelphusidae freshwater
crabs, providing new data that are relevant for taxonomic and biogeographical studies as well as for the develop-
ment of better regional and national assessments of the conservation status of the aquatic fauna.
Key words: Amazonia; biogeography; conservation; distribution records; geographical range; Guayana Shield;
Neotropical region; Kingsleyinae; Raddausinae.
RESUMEN
Diversidad de cangrejos de agua dulce: nuevos datos de distribución para cuatro especies
(Decapoda: Pseudothelphusidae) de países de Meso y Sudamérica
Introducción: Pseudothelphusidae es una familia muy diversa de cangrejos primarios de agua dulce ampliamen-
te distribuida por toda la región Neotropical, pero la verdadera extensión de la distribución geográfica de varias
especies es poco conocida porque muchas de ellas viven en zonas muy remotas de las que sólo se dispone de un
único registro o de muy pocos.
https://doi.org/10.15517/rev.biol.trop..v73iS1.63625
SUPPLEMENT
2Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63625, enero-diciembre 2025 (Publicado Mar. 03, 2025)
INTRODUCTION
Freshwater covers less than 1 % of the
worlds surface area but harbors almost 10 %
of all described species (Darwall et al., 2018;
Dijkstra et al., 2014; Dudgeon et al., 2006;
Mittermeier et al., 2010). Despite the indisput-
able importance of freshwater ecosystems in
economic, cultural, aesthetic, scientific, and
educational terms (see Dudgeon et al., 2006),
we are facing a freshwater biodiversity crisis
due to catastrophic declines in freshwater spe-
cies and the degradation of these hydrological
ecosystems (Darwall et al., 2018; Harrison et
al., 2018; World Wildlife Fund [WWF], 2020).
The Living Planet Report 2020 revealed a 94
% decline of the Living Planet Index for Latin
America and the Caribbean, the most striking
result observed in any region (WWF, 2020).
True or primary freshwater crabs (see
Cumberlidge & Ng, 2009; Yeo et al., 2008) form
with approximately 1 400 species an important
part of the freshwater biodiversity in tropi-
cal and subtropical regions around the world
(Yeo et al., 2008). The freshwater crab species
diversity in the Neotropics — 311 species in
three families (Trichodactylidae, Epiloboceri-
dae and Pseudothelphusidae) — is only sur-
passed by that of east and southeastern Asia
(Cumberlidge et al., 2014). According to the
IUCN Red List conservation assessment, 34 %
of the pseudothelphusids have an elevated risk
of extinction, and 56 % of these species are too
poorly known to assess (see Cumberlidge et
al., 2014). Moreover, a recent assessment of the
freshwater crabs in Colombia, which harbors
the highest number of species in the Neo-
tropics, revealed that pseudothelphusids are
the most threatened freshwater crab species in
this country (Acevedo-Alonso & Cumberlidge,
2021, Acevedo-Alonso & Cumberlidge, 2022).
These data highlight the urgent need to obtain
additional information about freshwater crab
species inhabiting the Neotropics, especially
about Pseudothelphusidae.
The Pseudothelphusidae is a very diverse
family of primary freshwater crabs widely dis-
tributed throughout the Neotropical region,
but the true extent of the geographic distribu-
tion of several species is poorly known because
many of them occur in very remote areas with
only a single or very few records available. For
such species, therefore, all data relating to new
occurrences are relevant as they are important
for both taxonomic and biogeographic studies
as well as for the development of regional and
national assessments of the conservation status
of the fauna. Here, we present new country
records for three species of pseudothelphusid
crabs: Phygiopilus acanthophallus Smalley, 1970
Objective: Aquí presentamos nuevos registros nacionales para cuatro especies de cangrejos pseudotelfusidos de
países de Meso- y Sudamérica.
Métodos: Los datos sobre la distribución geográfica se basan en especímenes depositados en colecciones de crus-
táceos de instituciones de Brasil, Alemania y Estados Unidos; se prepararon ilustraciones del primer gonópodo
masculino en un microscopio estereoscópico equipado con una cámara lúcida.
Resultados: Se publican por primera vez los registros de ocurrencia de Phygiopilus acanthophallus en México,
Fredius stenolobus en Guyana y Kunziana irengis en Brasil, y se confirma la presencia de Raddaus mertensi en
Honduras. Se dan comentarios sobre la morfología del primer gonópodo masculino de cada especie, se discuten
sus extensiones de distribución y se destaca la importancia de estos nuevos registros para las evaluaciones de
conservación.
Conclusiones: Los nuevos datos de la distribución geográfica amplían las áreas de ocurrencia de estas cuatro
especies de cangrejos dulceacuícolas de la familia Pseudothelphusidae, proporcionando nuevos datos que son
relevantes para estudios taxonómicos y biogeográficos, así como para el desarrollo de mejores evaluaciones regio-
nales y nacionales del estado de conservación de la fauna acuática.
Palabras clave: Amazonia; biogeografía; conservación; registros de distribución; rango geográfico; escudo de
Guayana; región Neotropical; Kingsleyinae; Raddausinae.
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from Mexico and two from South Ameri-
can countries: Fredius stenolobus Rodríguez
& Suárez, 1994 from Guyana and Kunziana
irengis (Pretzmann, 1971) from Brazil, as well as
the confirmation of the occurrence of Raddaus
mertensi (Bott, 1956) in Honduras. Consider-
ing that accurate information on the geographic
occurrence and distribution of these species is
scarce, here we discuss these new records and
range extensions and highlight the importance
of such data for the conservation assessment of
freshwater crabs in the Neotropics.
MATERIALS AND METHODS
The new records are based on specimens
that are preserved in ethanol 70 % and deposited
at the following institutions: Forschungsinstitut
und Naturmuseum Senckenberg (SMF), Frank-
furt am Main, Germany; Instituto Nacional
de Pesquisas da Amazônia (INPA), Manaus;
Museu de Zoologia, Universidade de São Paulo
(MZUSP), São Paulo, Brazil; and Nation-
al Museum of Natural History, Smithsonian
Institution (USNM), Washington D.C., U.S.A.
Measurements of carapace are presented in
millimeters; carapace width (cw) was mea-
sured across the carapace at its widest point,
and carapace length (cl) was measured along
the midline, from the frontal to the posterior
carapace margin. The abbreviation “G1” refers
to the male first gonopod and “coll.”/“colls.” to
collector(s). The Portuguese word “igarapé
means creek or a small forest stream. The only
locality without geographic coordinates origi-
nally informed in the label was tentatively geo-
referenced using Google Earth® software and
these coordinates are given in brackets. Line
drawings were made by CM using a Wild M8
(Wild, Heerbrugg, Switzerland) stereomicro-
scope equipped with a drawing tube attached,
then scanned and edited in Adobe Photoshop®
CS 2 software. The plates were mounted in
CorelDraw® X3 software.
RESULTS
Distributional data of four species of pseu-
dothelphusid crabs, two of Raddausinae and
two of Kingsleyinae, are presented for four
Meso- and South American countries: P. ac an -
thophallus in southeastern Mexico, R. mertensi
in western Honduras, F. stenolobus in west-cen-
tral Guyana, and K. irengis in northern Brazil.
Raddausinae Álvarez, Ojeda, Souza-Carvalho,
Villalobos, Magalhães, Wehrtmann
& Mantelatto, 2020
Phrygiopilus acanthophallus Smalley, 1970
(Fig. 1A)
Material examined: 2 males (cw 11.8,
cl 8.0; cw 24.3, cl 15.2), 2 females (cw 19.5,
cl 11.3; damaged), USNM 1180974, Mexico,
Chiapas, near San Jose [San Jose de las Pal-
mas, ~16°08’15”N 91°38’27”W], 28 miles ESE
of Comitán [Comitán de Domínguez], 7–20.
IV.1950, F.A. Pitelka coll.
Diagnosis: See Smalley (1970) and Rodri-
guez (1982).
Distribution: Guatemala (Alta Verapaz,
Baja Verapaz, Zacapa) (Rodriguez, 1982; Smal-
ley, 1970; Wehrtmann et al., 2016) and Mexico
(present study).
Remarks: The identity of present speci-
mens was determined by Martha R. Cam-
pos in June 1997, but this information has
never been formally published (http://n2t.
net/ark:/65665/3f6df1a83-408a-4140-8897-
1ae4806b9974).
The G1 of the largest male from Chiapas,
Mexico (USNM 1180974) resembles that of the
holotype regarding its general morphology and,
particularly, the proportions of the supra-apical
process (Fig. 1A).
Raddaus mertensi (Bott, 1956)
(Fig. 1B, Fig. 1C)
Material examined: 1 male (cw 35.1, cl
22.5), 1 female (cw 24.5, cl 16.6), SMF 31690,
Honduras, Ocotepeque, Guarin, cerro El Pital,
14°23.92’N 89°09.19’W, 14.V.1997, G. Köhler
coll.
Diagnosis: See Rodriguez (1982).
Distribution: Guatemala (Chiquimula)
(Wehrtmann et al., 2016), El Salvador (Santa
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Ana) (Bott, 1956), and Honduras (Ocotepeque)
(Acevedo-Alonso & Cumberlidge, 2022; pres-
ent study).
Remarks: The occurrence of R. mertensi
in Honduras has already been reported by
Acevedo-Alonso & Cumberlidge (2022) in a
study on the conservation status of mountain
species. However, these authors neither speci-
fied the distributional data of the specimen(s)
nor reported the voucher material on which the
record was based. The record presented herein
confirms the occurrence of the species in the
Ocotepeque department of Honduras.
The G1 of the specimen from Honduras
is similar to that of the holotype except for the
crenulation in the distal margin of the mesial
process, which is slightly more pronounced
in the Honduras specimen; furthermore, the
small, rounded protuberance on the cephalic
surface of the mesial process, which is present
Fig. 1. Phrygiopilus acanthophallus, USNM 1180974, left male first gonopod, entire stem in mesial view (A); Raddaus
mertensi, SMF 31690, right male first gonopod, distal portion in mesocaudal view (B) and laterocephalic view (C); Fredius
stenolobus, MZUSP 24496, left male first gonopod, distal portion in mesocaudal view (D); Kunziana irengis, INPA 2565, left
male first gonopod, distal portion in mesial view (E) and caudolateral view (F).
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in the G1 of the holotype, is barely noticeable in
the Honduras specimen (Fig. 1C).
Kingsleyinae Bott, 1970
Fredius stenolobus Rodríguez & Suárez, 1994
(Fig. 1D)
Material examined: 1 male (cw 42.2, cl
28.3), MZUSP 24492, Guyana, Potaro-Siparuni,
creek tributary of Kuribrong river, 05º27’N
59º31’W, 28.III.2011, F.C.T. Lima coll.
Diagnosis: See Rodríguez & Suárez (1994).
Distribution: Venezuela (Bolívar)
(Magalhães & Pereira, 2003; Mora-Day & Blan-
co-Belmonte, 2008; Rodríguez & Suárez, 1994),
Brazil (Roraima) (Zanetti et al., 2018), and
Guyana (present study).
Remarks: The G1 of the present specimen
is partially damaged, with a fissure at the base
of the apical complex, just between the mesial
process and the cephalic spine (Fig. 1D). Even
so, it can be seen that the G1 morphology of
the Guyana specimen is very similar to that of
specimens from the Orinoco (see Rodríguez
& Campos, 1998: 766, fig. 2A, B) and Amazon
basins (see Zanetti et al., 2018: 4, fig. 1G). The
only noticeable difference is in the G1 mar-
ginal process: in the present specimen, there is
a small subdistal concavity whereas this con-
cavity is very shallow in the specimen from the
Amazon basin and indistinct in the specimen
from the Orinoco basin.
Kunziana irengis (Pretzmann, 1971)
(Fig. 1E, Fig. 1F)
Material examined. 1 male (cw 23.0, cl
15.3), 1 female (soft carapace), MZUSP 45521,
Brazil, Roraima, Uiramutã, Serra do Sol, Parque
Nacional do Monte Roraima (= Mount Ror-
aima National Park), igarapé Caramambai, near
the bridge to the community Caramambatai,
05º07’58.8”N 60º35’08.8”W, 1005 m elev., col-
lected with dip net, captured while copulating,
10.XII.2019, J. Zuanon, G.T. Vilara and R. Bold-
rini colls.; 1 male (cw 13.8, cl 8.4) 1 female (cw
19.5, cl 12.5), MZUSP 45522, Brazil, Roraima,
Uiramutã, Serra do Sol, Parque Nacional do
Monte Roraima (= Mount Roraima Nation-
al Park), unnamed 2nd/3rd order stream, on
the trail to the Ingarikó farm, 05º06’52.1”N
60º36’34.0”W, 1003 m elev., 05.XII.2019, J.
Zuanon and D.A. Bastos colls.; 2 males (cw
20.0, cl 14.0; cw 21.3, cl 14.3) 1 female (cw 15.0,
cl 10.5), CCDB 7811, Brazil, Roraima, Uira-
mutã, Serra do Sol, Parque Nacional do Monte
Roraima (= Mount Roraima National Park),
unnamed 3rd order stream, 05º07’07.5”N
60º36’21.0”W, 1043 m elev., 30.XI.2019, J.
Zuanon and D.A. Bastos colls.; 1 male (cw
30.5, cl 19.7) 2 females (cw 23.3, cl 15.4; cw
30.3, cl 19.2), INPA 2565, Brazil, Roraima,
Uiramutã, Serra do Sol, Parque Nacional do
Monte Roraima (= Mount Roraima Nation-
al Park), 05º07’36.2”N 60º35’36.7”W, 1008 m
elev., 29.XI.2019, J. Zuanon and D.A. Bastos
colls.; 1 female (cw 22.8, cl 16.2), INPA 2566,
Brazil, Roraima, Uiramutã, Serra do Sol, Parque
Nacional do Monte Roraima (= Mount Rorai-
ma National Park), unnamed 1st order stream,
at the beginning of the trail to the Ingarikó
farm, 05º07’37.7”N 60º35’58.6”W, ~1000 m
elev., 30.XI.2019, J. Zuanon and D.A. Bastos
colls.; 2 females (cw 22.6, cl 15.1; cw 22.7, cl
15.7), INPA 2567, Brazil, Roraima, Uiramutã,
Serra do Sol, Parque Nacional do Monte Rorai-
ma (= Mount Roraima National Park), igarapé
Sokopi (3nd/4th order stream), 05º07’27.7”N
60º35’49.3”W, 1005 m elev., 01.XII.2019, J.
Zuanon and D.A. Bastos colls.
Diagnosis: See Magalhães et al. (2009).
Distribution. Guyana (Potaro-Siparuni)
(Magalhães et al., 2009; Pretzmann, 1971) and
Brazil (Roraima) (present study).
Remarks. The monospecific genus Kun-
ziana is characterized by a very unusual G1
morphology in which the apical processes
exhibit a combined strong twist in cephalic
direction with a longitudinal bending towards
the basal portion of the stem, causing the apical
field of spines to face downwards (Magalhães et
al., 2009: 42, fig. 1E–I). Such peculiar morphol-
ogy can also be seen in the G1 of the Brazilian
specimen of K. irengis examined herein, which,
however, exhibits a slightly more developed
patch of apical spinules on the distal portion
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of the caudal surface, and the apical plate is
not so distinctly bilobed (Fig. 1F). These small
differences could be attributed to the differ-
ent sizes of the specimens examined or intra-
specific variability.
DISCUSSION
Species distribution: This is the first pub-
lished record of P. acanthophallus from Mexico.
The species was previously known only from
Guatemala, in the departments of Alta Verapaz
(Rodriguez, 1982; Smalley, 1970) and Zacapa
(Wehrtmann et al., 2016), from the Caribbean
Sea drainage area. The location of the present
record in the department of Chiapas is located
in the upper-median portion of the Usumacinta
River basin, which empties into the Gulf of
Mexico. The species, therefore, has a distribu-
tion that encompasses river systems draining
into two versants. This new occurrence in
Chiapas, near the border with Guatemala, also
suggests that the species must have a wider
distribution in northwestern Guatemala, since
it would also be expected to occur in the upper
portion of the Usumacinta River basin.
Documented records of R. mertensi were
so far available only from its type locality, the
Department of Santa Ana, in northwestern El
Salvador (Bott, 1956) and from the Department
of Chiquimula, eastern Guatemala, in the Pacif-
ic versant (Wehrtmann et al., 2016). Although
the occurrence of the species has already been
mentioned from Honduras (Acevedo-Alonso &
Cumberlidge, 2022), this is the first document-
ed record from the country, and such occur-
rence is not unexpected since the three known
occurrences of the species are situated relatively
close to each other in the region where the
borders of the three countries converge, all in
aquatic systems of the Pacific drainage.
Fredius stenolobus had already been record-
ed from the Caroni and Caura River basins in
the state of Bolívar, Venezuela (Magalhães &
Pereira, 2003; Mora-Day & Blanco-Belmonte,
2008; Rodríguez & Suárez, 1994), which are
part of the Orinoco River basin, and from the
Branco River basin in the state of Roraima,
Brazil (Zanetti et al., 2018), an affluent of the
Negro River, one of the main northern tributar-
ies of the Amazon basin. Its occurrence in the
Kuribrong River, an affluent of the Potaro River,
which is part of the Essequibo River basin in
west-central Guyana, is the first record of the
species from this country. The present new
record, therefore, reveals that F. stenolobus has
a distribution that covers three different river
basins draining the Guayana Shield region in
northern South America.
Kunziana irengis has been originally found
in the Guyanase side of the Maú/Ireng River
basin (Magalhães et al., 2009; Pretzmann, 1971),
the river that delineates the border between
western Guyana and Brazil. The present record
in creeks that are tributaries of the Contigo
River (tributary of the Surumu → Tacutu →
Branco → Negro → Amazon Rivers) confirms
that it also occurs in Brazilian territory, as
suggested by Magalhães et al. (2009). Both the
Contigo River and the Maú/Ireng River, which
also flows into the Tacutu River, are some of
the northernmost tributaries of the Amazon
basin. Although the known occurrences of the
species in Guyanese territory are located on
the southern flank of the Wokomung Massif,
whose streams drain into the Maú/Ireng River
basin, Magalhães et al. (2009) did not rule out
the possibility that the species is also present
in the Essequibo River basin, in streams that
form the headwaters of the Potaro River in the
northern portion of the aforementioned massif.
The presence of common elements of the
aquatic fauna between the Essequibo and Bran-
co River basins, as it is the case of F. stenolobus
and K. irengis reported herein, as well as for
other species of pseudothelphusid crabs (Man-
telatto et al., 2022), can be understood in the
context of the hydrogeomorphologic complex
history of the region. This region was drained
by the Proto-Berbice River basin that flew
during Late Cretaceous to Early Quaternary
from the southwest of the state of Roraima to
the northeast of present-day Guyana. Tectonic
and climatic events during the Plio-Pleistocene,
however, promoted a rearrangement of the
basin, especially the southward reversion of the
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Branco River and its entire northern catchment
area, including the Maú/Ireng river drainage,
into the Amazon basin, with a still existing con-
nection area, the Rupununi Portal, a wetland
region between the Tacutu and upper Esse-
quibo rivers (Cremon et al., 2016; Ferreira et
al., 2007; Lujan & Armbruster, 2011; Schaefer
& Vale, 1999).
Conservation assessment: The IUCN
global conservation assessment carried out in
2008 contains information only for three of
the four species treated in this contribution:
P. acanthophallus and R. mertensi were listed
as “Vulnerable, while F. stenolobus was clas-
sified as “Least Concern. The fourth species,
K. irengis, was not included in the 2008 IUCN
global assessment, since it was considered as
incertae sedis and as a non-valid species (C.
Magalhães, personal observation, Sept. 2007).
Only after this assessment, Magalhães et al.
(2009) confirmed Kunziana as a valid genus
and redescribed K. irengis. In the case of the
three freshwater crab species included in the
IUCN conservation assessment, all of them
are lacking information about population size,
population trends, and abundance. Moreover,
their assessment was based on only three (R.
mertensi and F. stenolobus) or four (P. a c a n -
thophallus) locations. In all three cases, the
results of our study not only add new geo-
graphical reports/locations, but also include a
new country previously not recorded for the
species. Such information is important since
it is the basis for the calculation of two evalu-
ation criteria (Extent of occurrence and Area
of occupancy) used for the assessment of the
IUCN categories (International Union for Con-
servation of Nature [IUCN], 2012). Moreover,
regular updating of the data is essential, as
shown by the example of Colombia, the country
with the highest number of freshwater crab in
the Neotropics: between the first (2008) and
the latest (2018) assessment, the percentage
of threatened species in Colombia showed a
dramatic increase from 26 % in 2008, to 34
% in 2015, to 62 % in 2018 (Acevedo-Alonso
& Cumberlidge, 2021). Therefore, additional
studies are constantly necessary not only to
update and expand our knowledge about the
different freshwater crab species but also to
broaden and improve the information about
threats affecting these macroinvertebrates.
A relatively high number of data-defi-
cient species could lead to underestimations of
the numbers of threatened freshwater species
(Cumberlidge et al., 2014). In the case of K.
irengis, this species has not even been assessed
yet and only few specimens have been col-
lected so far. The limited information available
about the habitat of this species indicates that
it occurs in clean rocky mountain streams
with elevations ranging from 1 077 to 1 485 m
(Magalhães et al., 2009). Just as in Colombia
(see Acevedo-Alonso & Cumberlidge, 2021;
Acevedo-Alonso & Cumberlidge, 2022), these
montane environments might be threatened
by increasing rates of deforestation, pollution,
and human population growth. Serra do Sol, in
the setentrional region of the state of Roraima,
Brazil, is a mountainous region covered by
savannah and forest situated within the Rapo-
sa-Serra do Sol Indigenous Land inhabited by
indigenous people from several ethnic groups,
an area that has experienced intense political
conflicts between developers and the native
population due to the regions mineral, agricul-
tural, and tourist potential (Lauriola, 2003; Sar-
tori & Bethonico, 2018). Therefore, there is an
urgent need to obtain additional information
about K. irengis and to identify current threats
to its habitat to develop adequate conservation
measures for this species.
Although these new records only slightly
expand the geographic distribution areas of
these four Pseudothelphusidae freshwater crab
species, they provide relevant new data for
taxonomic and biogeographic studies, as well
as for the development of better regional and
national assessments of the conservation status
of the aquatic fauna.
Ethical statement: the authors declare
that they all agree with this publication and
made significant contributions; that there is
no conflict of interest of any kind; and that we
8Revista de Biología Tropical, ISSN: 2215-2075 Vol. 73(S1): e63625, enero-diciembre 2025 (Publicado Mar. 03, 2025)
followed all pertinent ethical and legal proce-
dures and requirements. All financial sources
are fully and clearly stated in the acknowledg-
ments section. A signed document has been
filed in the journal archives.
ACKNOWLEDGEMENTS
We thank Jansen Zuanon and Douglas
A. Bastos for making available the crustacean
material collected during their expedition to
Serra do Sol for our study. CM also acknowl-
edges the assistance of Marcos Tavares
(MZUSP), the late Michael Türkay (SMF), and
Rafael Lemaitre and Karen Reed (USNM) dur-
ing study visits to their respective institutions.
ISW would like to acknowledge the support of
the Universidad de Costa Rica through project
No. C2721. We also acknowledge the valuable
comments of two anonymous reviewers, which
contributed to improving the manuscript.
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