Volume 9, Manuscript ID
es20260001, p. 01-12, 2026
Doi: https://doi.org/10.32435/envsmoke-2026-0001
Environmental
Smoke, e-ISSN: 2595-5527
“A leading multidisciplinary
peer-reviewed journal”
Full
Article:
RANGE EXTENSION OF TWO
BRACHYURAN CRABS COLLECTED FROM THE AMAZON CONTINENTAL SHELF
Déborah Elena Galvão Martins1,2* (https://orcid.org/0000-0002-3829-4388); Flavio de Almeida Alves-Júnior1,2
(https://orcid.org/0000-0003-3002-6845); Israel Hidenburgo Aniceto Cintra1 (https://orcid.org/0000-0001-5822-454X); Bianca Bentes2 (https://orcid.org/0000-0002-4089-7970)
1Laboratório de Crustáceos (LabCrus),
Socio-Environmental and Water
Resources Institute
(ISARH), Federal Rural University of Amazônia (UFRA),
Avenida Presidente Tancredo Neves, nº 2501, Terra Firme, CEP: 66077-830, Belém,
Pará, Brasil
2Núcleo de Ecologia
Aquática e Pesca da Amazônia (NEAP), Post-Graduate
Programme in Aquatic Ecology and Fisheries
(PPGEAP), Federal University of Pará (UFPA), Avenida
Perimetral, 2651, Terra Firme, 66077-530 Belém, Pará, Brasil
*Corresponding author: deborah.martins@ufra.edu.br
Submitted on: 17 Nov. 2025
Accepted on: 11 Dec. 2025
Published on: 20 Jan. 2026
License:
https://creativecommons.org/licenses/by/4.0/
Abstract
The Amazon Continental Shelf (ACS) is composed of numerous benthic
habitats, which shelter a high diversity of invertebrates, especially
crustaceans. However, due to the complexity of ecosystems, knowledge of
crustacean fauna is still incomplete. We herein report the northernmost
Brazilian record of two brachyuran crab species, Teleophrys pococki
Rathbun, 1924 (Mithracidae) and Euchirograpsus
americanus A. Milne-Edwards, 1880 (Plagusiidae),
associated with the Great Amazon Reef System (GARS) (State of Amapá),
additionally, providing a description of their first gonopods (G1). Both male
specimens E. americanus and T. pococki, were accidentally
collected during commercial fishing operation of the red snapper Lutjanus
purpureus (Poey, 1866) in the ACS (northern Brazil), at depths up to 70 m,
in August 2023 and September 2025. The presence of both species in the GARS
increases crustacean biodiversity in this ecosystem and highlights the need for
further studies in the region to assess its true diversity and to develop
management plans focused on species conservation.
Keywords: Amazon
Reefs. Bycatch. State of Amapá. Geographic distribution. New record.
1
Introduction
The Amazon continental shelf (ACS) is classified as a hotspot of diversity,
due to the elevated complexity of habitats, the wide range of ecological
interactions and adequate environmental conditions for the establishment of
species (KLAUTAU et al., 2025); much of this biodiversity is due to the
presence of the Amazon River plume, which transports organic matter,
establishing a high energy flow in marine food chains (WEBER et al., 2017;
DRAKE et al., 2021).
In this area, there is also an ecosystem composed of
rhodoliths, and coral beds designated as the Great Amazon Reef System (GARS),
which is distributed along mesophotic areas of ACS, ranging from 20 to ~100 m
(VALE et al., 2022).
The GARS is the target of several studies, especially
on geological mapping, ecological functionality and the diversity associated
with this environment (MOURA et al., 2016; VALE et al., 2022).
However, this region has already been affected by
human activities, such as fishing, marine litter and the effects of climate
change (ALVES-JÚNIOR et al., 2024a), even before the knowledge about the
species diversity (including crustaceans, which are usually recorded as part of
fishing bycatch) was completely available (MARTINS; ALVES-JÚNIOR, CINTRA,
2025).
The family Mithracidae
MacLeay, 1838 corresponds to 15 genera and 68 species, occurring from the
coastal zones to the continental shelf break, associated with sand, gravel or
coral bottoms (MELO, 1996; WINDSOR; FELDER, 2014; DECANET, 2025a).
Into this family, the genus Teleophrys
Stimpson, 1860 comprises five recent species: T. cristulipes Stimpson,
1860, T. ornatus Rathbun, 1901, T. pococki Rathbun, 1924, T. ruber (Stimpson, 1871), T. tumidus
(Cano, 1889), besides T. acornis Portell & Collins, 2004
as a fossil record (DECANET, 2025a).
From the Brazilian waters, only the species T.
ornatus and T. pococki have been recorded in the northeastern and
southeastern regions (MELO, 1996). Despite the occurrence of T. pococki
in coastal zones of Brazil, this species was reported in a few localities,
represented only by occasional records.
Additionally, crabs of the family Plagusiidae
Dana, 1851 are found along
the coastal zones and continental slope areas (including Oceanic
Islands), associated with bottoms of sand, gravel and coral (MELO, 1996; 1998).
These organisms are widely reported along the Atlantic
and Indo-Pacific Oceans, which are classified in 21 species distributed into
seven genera: Caligoplagusia Fujita & Narusa, 2014, Davusia
Guinot, 2007, Euchirograpsus H. Milne Edwards,
1853, Guinusia Schubart & Cuesta, 2010, Miersiograpsus Türkay, 1978 and Plagusia
Latreille, 1804, as well as the extinct species Petrusia
Beschin, Busulini, Tessier & Zorzin,
2016 (GUINOT, 2007; NG; GUINOT; DAVIE, 2008; DECANET, 2025b).
Currently, the genus Euchirograpsus is
composed of nine valid species, occurring from 10 to 510 m of depth in the
bottoms of sand, mud, rock and corals, associated with gorgonians and sponges
(TÜRKAY, 1975; MANNING; HOLTHUIS, 1981, ALVES-JÚNIOR et al., 2016; WILSON;
POHLE, 2016). In Brazilian waters, only two species have been recorded: Euchirograpsus antillensis
Türkay, 1975 and Euchirograpsus
americanus A. Milne-Edwards, 1880, both occurring along the continental
shelf and shelf break of the northeastern and southeastern regions (MELO, 1996;
1998; ALVES-JÚNIOR et al., 2016). Thus, herein we report the Brazilian
northernmost records of two brachyuran species: Teleophrys pococki and Euchirograpsus
americanus from the state of Amapá, associated with the GARS.
2 Material and Methods
The crab species were obtained incidentally attached
to an iron cage trap called "manzuá" (mesh
size of 5 cm) during commercial fishing operations of the red snapper Lutjanus
purpureus (Poey, 1866). These fishing activities were performed between the
depths of 70 and 100 m, along the continental shelf of the Brazilian state of
Amapá (Figure 1) from August
2023 to September 2025, covering areas above the Great Amazon Reef System
(GARS).
Figure 1. Map of the study area, covering the Amazon continental
shelf (ACS), indicating the occurrence sites of Teleophrys pococki
Rathbun, 1924 and Euchirograpsus americanus
A. Milne-Edwards, 1880. Shapefile of the GARS from Moura et al. (2016).
In the field, the rhodoliths adhered to the external
surface of the trap were manually removed, related invertebrate species were
fixed in 70% ethanol, and then transported to the Laboratório
de Crustáceos (LabCrus). All
sampling procedures were supervised by the
Centro Nacional de Pesquisa e Conservação da Biodiversidade Marinha do Norte
(CEPNOR) under the SISBIO Number 44915–3.
In the laboratory, both species were identified following Türkay (1975)
and Melo (1996), photographed, measured in carapace width (cw.), carapace
length (cl.) and wet weight (ww.). After the procedures, the specimens were
deposited under voucher number at the Coleção
Carcinológica do Laboratório
de Crustáceos (LabCrus) / Universidade
Federal Rural da Amazônia (UFRA).
3 Results
and Discussion
One adult male of T. pococki (Figure 2A-C) (cw.
11.6 mm, cl. 10.3 mm and ww. 0.57 g) (Voucher Number: 2.16.1A) was collected at
a depth of 70 m in the GARS area (August 2023), associated with rhodoliths in
the gravel bottom, state of Amapá (Brazil) (04°43’59” N; 050°39’18” W).
According to Tavares and Mendonça Jr. (2022), the bathymetric distribution of T.
pococki covers continental shelf areas, from shallow waters to 100 m;
however, it is more commonly found up to 25 m of depth (MELO, 1996; CASTAÑO;
CAMPOS, 2003; CARMONA-SUÁREZ; POUPIN, 2016).
Figure 2. Teleophrys pococki Rathbun, 1924. (A) Male in dorsal
view; (B) ventral view; (C) right first gonopod in pleonal
view, in stereomicroscopy (Scale bars = 5mm (A,B), 1mm
(C).
Our specimen fits well with the original description provided by Rathbun
(1924) and additional information provided by Rathbun (1925) and Melo (1996),
which include the main characteristics: carapace larger than long, smooth,
containing few granules or small tubercles which are more visible in the
branchial and cardiac regions; rostrum bifid, also larger than long, with one
antero-lateral teeth; carpus of chelipeds with two lobes only in the inner
face, chelipeds smooth forming an open angle in the centre, with curved edges.
Ambulatory legs with several tubercles forming crests on the upper surface (Figure 2A). Free
abdominal somites with a median elevation in somites 3 to 6 (Figure 2B).
The first gonopod (G1) is long and robust, base with plumose setae,
lateral margin with simple setae sparsely distributed, a suture present from
the base to the distal margin of the apical plate, rounded protuberance on the
lateral margin; apical plate with two lobes covered with several acute spines (Figure 2C).
Originally, the species T. pococki was designated as a new
species in a revision of the type material of T. cristulipes provided by
Rathbun (1924), being these both closely related species, which can be easily
separated by main characters: rostrum short and large, slightly bifid and not
reaching the first antennal peduncle in T. pococki vs. rostrum
with frontal margin large, bifid and overcoming the first antennal peduncle in T.
cristulipes; carapace with only one antero-lateral tooth in T. pococki
vs. carapace with 3-4 antero-lateral teeth in T. cristulipes;
carapace smooth, with small granules and tubercles only in the branchial and
cardiac regions in T. pococki vs. all carapace regions covered
with small granules and tubercles in T. cristulipes. Ambulatory legs are
slender, with tubercles forming a crest in the superior margin in T. pococki
vs. ambulatory legs robust with a visible lobe at propodus in T.
cristulipes (see more details in RATHBUN, 1925). Rathbun (1900), analyzing samples from Brazil, indicated T. cristulipes
as occurring in the region; however, the same author posteriorly described the
Brazilian species as T. pococki, indicating T. cristulipes as
native from the Pacific Ocean. In addition, the distribution of T.
cristulipes covers the Pacific Ocean, Mexico (Baja California), Costa Rica
(Golfo Dulce), Panama, Colombia, Ecuador (Galapagos) (HOLTHUIS, 1979;
HENDRICKX, 2010).
The species T. pococki is restricted to Western Atlantic,
occurring in the United States (Louisiana), Bonaire, Curaçao, Honduras, Bahamas, Lesser Antilles, Colombia
(Santa Marta), Venezuela and Brazil: (Amapá [this study], Rocas Atoll, Fernando
de Noronha Archipelago, Pernambuco, Alagoas, Bahia and São Paulo, Trindade and Martin Vaz
Archipelago) (GARTH, 1978; GOUVÊA, 1986; COELHO; ALMEIDA; BEZERRA, 2008;
MELO, 1996; 1998; ALVES et al., 2012; FLEEGER; COWAN-JR.; PASCAL, 2013;
CARMONA-SUÁREZ; POUPIN, 2016). In Brazilian waters, only two Teleophrys
species (T. pococki and T. ornatus) have been reported; however, T.
ornatus is recorded only in Fernando de Noronha Archipelago, Bahia and São
Paulo (MELO, 1996; ALVES et al., 2012). The main differences between these two species
are: T. pococki has the carapace unarmed, covered with granules and
wider than long, and the ambulatory legs with a crest only on the upper
surface; while T. ornatus has the carapace armed with tubercles and
small spines and longer than wide, and the ambulatory legs showing a crest on
both sides (superior and inferior margins) (see MELO, 1996, pags.
248, 249).
The mithracid species T. pococki is considered an
ecosystem engineer, especially living in association with coral reefs (e.g. Mussismilia
hispida (Verrill, 1901) in shallow habitats), or living in burrows in dead
coral or artificial substrates such as artificial reefs, oil platforms and ship
hulls (NOGUEIRA; NEVES; JOHNSON, 2015).
Furthermore, this species may be found in calcareous algae beds;
nonetheless, it is commonly found using seaweed of the genera Centroceras Kützing,
1841, Ceramium Roth, 1797 and Chaetomorpha Kützing,
1845 as ornamentation on areas of the carapace and walking legs (GOUVÊA, 1986;
LÓPEZ; SOLANO, 2005; TAVARES; MENDONÇA JR., 2022). Studies performed by
Vera-Caripe et al. (2017) indicated the commensal association of T. pococki
with the sea urchin Tripneustes ventricosus (Lamarck, 1816), being the echinoderm a protection
against predators.
Into our grapsid specimen, we collected one
male of E. americanus (Figure 3 A-D) (cw.
7.10 mm, cl. 7.50 mm and ww. 0.24 g) (Voucher Number: 22.2.1 A), at 85 m of
depth, in areas of the GARS (September 2025), state of Amapá (Brazil)
(03°00’27” N; 049°06’20” W), associated with rhodoliths in the gravel bottom.
Our specimen agrees with the original description provided by A. Milne-Edwards
(1880) and additional posterior descriptions of A. Milne-Edwards and Bouvier
(1894), Rathbun (1918), Türkay (1975) and Melo (1996).
Figure 3. Euchirograpsus americanus A. Milne-Edwards, 1880. (A) Male in
dorsal view; (B) inferior margin of the merus
of the second pair of ambulatory legs (P2); (C) Carapace; (D) right
first gonopod in pleonal view, in stereomicroscopy
(Scale bars = 5mm (A), 1mm (D).
The main characteristics for the identification of this species are: carapace subquadrate, flattened posteriorly, with the
surface covered with small granules and clumps of stout setae spread
non-uniformly, including numerous short setae covering the carapace. The
bilobed front reaches the middle of the first antennular article (Figure 3A). Three
antero-lateral acute spines (excluding the orbital), being the second more
pronounced/acute than the others (Figure 3A-C).
Chelipeds thin, covered by stout setae and small granules, slightly
curved downward. Merus of the chelipeds contain five acute spines on the
ventral-distal angle. Ambulatory legs covered with stout setae and small
granules; merus of pereopods 2 to 5 (P2-P5) ending in
a sharp distal tooth, merus of the second ambulatory
leg (P2) containing three to
four subdistal teeth (a higher and sharper) (Figure 3B). Dactyl of P2-P5 with five to
six robust spines, being the tip trifurcated (similar to
a hook). Abdominal somites one and two are reduced,
and somites 4-6 are slightly fused. First gonopod
(G1) robust, slightly curved towards the mesial line, with sparse setae in its
mid-distal portion and dense pubescence at the tip (see Figure
3D) (modified from TÜRKAY, 1975; MELO, 1996).
In the Western
Atlantic, the species most closely related to E. americanus is E. antillensis.
Both species occur in the same habitats (sympatric species),
however they are morphologically distinguished according to Türkay (1975, pag. 113, Fig. 16) and Alves-Júnior et al. (2016, pag. 3, Fig. 1) following the characters: the individuals
of E. americanus have three to four subdistal teeth on the inferior
margin of the merus of the second pair of ambulatory
legs (P2), being the first or second (sometimes the last) larger and higher
pitched than the others; while in E. antillensis
all subdistal teeth on the inferior margin of the merus
of P2 are of the same size. Additionally, Türkay (1975) indicated that the
anterolateral teeth of E. americanus are less developed than in E. antillensis. Studies developed by Poupin (1994; 2018)
indicated this character as valid for male and female specimens
identification.
Until now, several
specimens were erroneously assigned as E. americanus, especially in
Indo-Pacific regions; however, after the revision provided by Türkay (1975) and
additional observations of Manning and Holthuis
(1981); these authors indicated that only two Euchirograpsus
species (E. americanus and E. antillensis)
are restricted to the Western Atlantic. The distribution of E. americanus covers: Canada (Bay of
Fundy), USA (towards the Carolina’s in Toms/ Meys
Canyon and Baltimore Canyon, Massachusetts in Canyon at the edge of Georges
Bank, Delaware, New Jersey, Louisiana, Florida), Gulf of Mexico, Dry Tortugas,
Cuba, Lesser Antilles, West Indies, Guadalupe, Barbados, Colombia, Venezuela,
French Guiana and Brazil (Amapá [this study], Maranhão, Rio Grande do Norte and
Rio Grande do Sul states) (ZARIQUIEY ALVAREZ, 1968; TÜRKAY, 1975; MELO, 1996; 1998; CORBARI et al. 2015;
ALVES-JÚNIOR et al., 2016; WILSON; POHLE, 2016; POUPIN, 2024). According to
Wilson and Pohle (2016), this species is normally found in tropical areas;
however, possibly due to climate change events, E. americanus has
expanded into subtropical areas, as observed in cold waters (~8.3 °C) of Canada
(Bay of Fundy).
The presence of these
species in the State of Amapá may be associated with the GARS, once this region
encompasses a wide range of micro- and macro habitats beneath the Amazon River
plume, characterized by high energy flow and acting as connection between South
America and the Caribbean Sea (MOURA et al., 2016; VALE et al., 2022;
ALVES-JÚNIOR; MARTINS; CINTRA, 2024b). Along the GARS, the
migration/distribution of invertebrate and vertebrate species may occur in a
stepping-stones process, especially in areas below 70 m (ALVES-JÚNIOR; MARTINS;
CINTRA, 2024b; MARTINS; ALVES-JÚNIOR; CINTRA, 2025). This explains the
widespread occurrence of T. pococki and E. americanus throughout
the Western Atlantic.
Two other additional hypotheses
that may explain the presence of both species in northern Brazil are: 1) their
transport (larvae and adults) is associated with ships on route between Brazil
and the Caribbean Sea; 2) these species were probably already distributed
throughout the Amazon continental shelf region, but the lack of researchers and
specialized sampling efforts along the GARS have hindered knowledge of the
distribution of T. pococki and E. americanus.
4 Conclusions
Despite the high biological
diversity found in the ACS, the region is suffering severe anthropogenic
impacts, especially associated with fishing activities, climate change, marine
debris and ghost fishing. As a result, several species of marine invertebrates
and vertebrates are included in threatened species lists, even before they have
been catalogued or their biological characteristics fully understood. The
presence of only a single specimen of T. pococki and E. americanus in
the northern region limits our understanding of their biology and conservation
status. In conclusion, we herein update the geographic distribution of two
brachyuran crab species from the Amazon continental shelf; however, further
studies in the Amazon reef region are needed to improve the knowledge of
crustacean diversity in northern Brazil.
CREDIT AUTHORSHIP CONTRIBUTION STATEMENT
Conceptualization:
D.E.G.M., I.H.A.C., F.A.A.J. and B.B. Funding: I.H.A.C., F.A.A.J. and B.B. Data
curation, Methodology: D.E.G.M., I.H.A.C. and F.A.A.J. Investigation: D.E.G.M.
and F.A.A.J. Visualization: D.E.G.M. Writing – original draft: D.E.G.M., I.H.A.C.
and F.A.A.J. Writing – Review & Editing: D.E.G.M. and B.B.
DECLARATION
OF INTEREST
The authors disclose that they
have no known competing financial interests or personal relationships that
could have appeared to influence the study reported in this manuscript.
FUNDING
SOURCE
This research was financed by the
Conselho Nacional de Desenvolvimento
Científico e Tecnológico – CNPq
(Brazil), through the productivity grant award, reference numbers: 304401/2025-0
(author Israel Cintra) and 302175/2025-3 (Bianca Bentes).
ETHICAL PROCEDURES
No live animal treatments were
carried out in this study. All specimens were retrieved from regulated
commercial fishing operations and were dead upon examination.
ACKNOWLEDGEMENTS
The authors would like to thank the Centro Nacional de Pesquisa e Conservação da Biodiversidade
Marinha do Norte (CEPNOR, Brazil) for the sampling of the examined material
and the support in the laboratory. Additionally, we would like to thank the anonymous
reviewers for their valuable comments throughout the manuscript.
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