Research Article |
Corresponding author: Setuko Masunari ( set_mas@yahoo.com.br ) Academic editor: Ingo S. Wehrtmann
© 2020 Setuko Masunari, Salise Brandt Martins, André Fernando Miyadi Anacleto.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Masunari S, Martins SB, Anacleto AFM (2020) An illustrated key to the fiddler crabs (Crustacea, Decapoda, Ocypodidae) from the Atlantic coast of Brazil. ZooKeys 943: 1-20. https://doi.org/10.3897/zookeys.943.52773
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Fiddler crabs are one of the most notable animal groups in Brazilian estuarine environments, due to their high density and characteristic waving of males. An illustrated key to the ten species recorded as far in the country is provided using only clearly visible characters of males. Furthermore, additional recognition characters, information about geographic distribution and biology of each species are presented. Most examined crabs were collected in Guaratuba Bay, southern Brazil.
Biological notes, distribution, mangrove, recognition characters, tidal flats
Studies on fiddler crabs began more than 300 years ago, certainly because the observers were attracted to the immense claw of males that were tirelessly waving in a typical movement and rhythm. Fiddler crabs are semi-terrestrial decapod crustaceans and inhabit shaded substrates of mangrove forest or sunny tidal flats adjacent to it. During high tides they hide in individual burrows that are the center of a fiddler crab life (
Ten species of fiddler crabs are known from the Brazilian Atlantic coast (
Despite the valuable identification keys for Brazilian fiddler crabs elaborated by
Ten species of fiddler crabs recorded along Brazilian coast were analyzed, each one represented by numerous individuals. Most of them (seven species) were collected in various habitats of Guaratuba Bay, municipality of Guaratuba, southern Brazil: Minuca burgersi (Holthuis, 1967), M. mordax (Smith, 1870), M. rapax (Smith, 1870), Leptuca leptodactyla (Rathbun, 1898), L. thayeri (Rathbun, 1900), L. uruguayensis (Nobili, 1901), and Uca maracoani (Latreille, 1802–1803). These crabs were deposited in the Natural Museum of Natural History of Capão da Imbuia located in Curitiba, Paraná State, southern Brazil. The remaining three species were obtained from other locations in Brazil: L. cumulanta (Crane, 1943) from Natal (Rio Grande do Norte state, northeastern Brazil), M. vocator (Herbst, 1804) from Cananeia (São Paulo state, southeastern Brazil) and M. victoriana (von Hagen, 1987) from Guarapari (Espírito Santo state, southeastern Brazil); the specimens of the latter species were deposited at the Museum of Zoology of University of São Paulo.
The illustrated key was elaborated as simple as possible, and only clearly visible characters were selected. The key is exclusively based on adult male individuals, as they are provided with the diagnostic characters of the species. As fiddler crabs have gregarious habits, male individuals are hardly absent in the populations.
Line drawings were prepared using a drawing tube attached to a stereoscopic microscope. The systematic nomenclature was based on
The pile, an important morphological feature, is a wooly pubescence on the surface of carapace and ambulatory legs of some species. It is a somewhat difficult to be recognized by beginners, especially in crabs that were preserved in liquids. Drying the specimens in the open air is a practical clue to facilitate the visualization: the piles appear as clear and rough patches on the darkened surface of the carapace or ambulatory legs. It is highly recommended that beginners learn to distinguish these piles, since some morphologically similar species can be easily identified by observing the distribution of these patches.
Geographic distribution of the fiddler crab species along the Atlantic coast of Brazil. The states were organized by increasing southern latitudes (from left to right), except Amapá that is located northern to Equator. Abbreviations: AL = Alagoas, AP = Amapá, BA = Bahia, CE = Ceará, ES = Espírito Santo, MA = Maranhão, PA = Pará, PB = Paraíba, PE = Pernambuco, PI = Piauí, PR = Paraná, RJ = Rio de Janeiro, RN = Rio Grande do Norte, RS = Rio Grande do Sul, SC = Santa Catarina, SE = Sergipe, SP = . Black circle = occurrence recorded; white circle = probable occurrence but not officially recorded; black square = presence of mangrove.
States of Brazil | AP | Equator | PA | MA | PI | CE | RN | PB | PE | AL | SE | BA | ES | RJ | Tropic of Capricorn | SP | PR | SC | RS |
Mangrove | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | ■ | |||
Minuca mordax | • | • | • | ○ | • | • | ○ | • | • | • | • | • | • | • | • | • | • | ||
Minuca rapax | • | • | • | • | • | • | • | • | • | • | • | • | • | • | • | • | |||
Uca maracoani | • | • | • | ○ | • | • | • | • | • | • | • | • | • | • | • | • | |||
Leptuca leptodactyla | • | • | • | • | • | • | • | • | • | • | • | • | • | • | • | ||||
Leptuca thayeri | • | • | ○ | • | • | • | • | • | • | • | • | • | • | • | • | ||||
Minuca burgersi | • | • | ○ | • | • | • | • | • | • | • | • | • | • | • | • | ||||
Minuca vocator | • | • | • | ○ | • | ○ | • | • | • | • | • | ○ | • | • | |||||
Leptuca cumulanta | • | • | • | ○ | • | • | • | • | • | • | • | ○ | • | ||||||
Minuca victoriana | • | • | • | • | • | • | |||||||||||||
Leptuca uruguayensis | • | • | • | • | • |
1 | Narrow front, width less than or equal to 15% of front–orbital breadth (Fig. |
2 |
– | Wide and triangular front, width more than 15% of front–orbital breadth (Fig. |
3 |
2 | Spatulate front (Fig. |
Uca maracoani |
– | Triangular front, base ca. 15% of front–orbital breadth; patches of pile (= woolly pubescence, easily detached) on dorsal surface of carapace (Fig. |
Leptuca thayeri |
3 | Carapace provided with major and minor pairs of postero–lateral striae (Figs |
4 |
– | Carapace provided with a single pair of postero–lateral striae (Figs |
8 |
4 | Exuberant pile on the dorsal surface of the carapace forming a typical pattern (Fig. |
Minuca vocator |
– | Carapace with discrete pile or without any, but segments of ambulatory legs (at least from 1st to 3rd pairs) with pile (Figs |
5 |
5 | Male major chela provided with a short and straight depression filled with pile at the base of pollex (Fig. |
Minuca victoriana |
– | Male major chela without any depression at the base of pollex (Fig. |
6 |
6 | Pile on dorsal surface of carpus and merus and around the entire surface of manus of ambulatory legs (1st to 3rd pairs) (Fig. |
Minuca mordax |
– | Pile limited to dorsal surface of carpus and manus of ambulatory legs (1st to 3rd pairs); merus without pile (Fig. |
7 |
7 | Ambulatory legs with wide merus and dorsal margin convex; merus of last pair of ambulatory legs more than two times wider than respective carpus in its maximum breadth (Fig. |
Minuca rapax |
– | Ambulatory legs with narrow merus and dorsal margin almost straight; merus of last pair of ambulatory legs less than two times wider than respective carpus in its maximum breadth (Fig. |
Minuca burgersi |
8 | Abdomen with middle somites fused (Fig. |
9 |
– | Abdomen with all somites distinct (Fig. |
Leptuca cumulanta
(Fig. |
9 | Manus of major claw provided with a long groove on dorsal surface following its margin, mostly filled with dirt (Fig. |
Leptuca uruguayensis |
– | Manus of major claw without any groove on dorsal surface (Fig. |
Leptuca leptodactyla |
Leptuca uruguayensis. A carapace with a pair of postero–lateral striae, dorsal view B male major claw with dorsal groove (seta), dorso-frontal view. Leptuca leptodactyla C carapace with a pair of postero-lateral striae, dorsal view D male major claw without dorsal groove, dorso-frontal view.
Carapace pentagonal moderately arched in the anteroposterior direction and dorsal surface without pile (Fig.
The species reproduces year-round in the population of Ubatuba, southeastern Brazil (
The species is morphologically close to its congeners M. rapax and M. mordax; the distinguishing characters among these species are treated in the subsequent items.
Carapace pentagonal moderately arched and with naked dorsal surface, without pile (Fig.
Ovigerous females were collected inside burrows that were ornamented with poorly structured chimney at Guaratuba Bay, southern Brazil, during a warm month (November) (
The most conspicuous morphological character of M. mordax is the presence of piles covering the entire surface (dorsal, lateral and ventral) of the manus of 1st to 3rd ambulatory legs. This feature allows to easily distinguish M. mordax from two other closely related species, M. burgersi and M. rapax. As no piles are present on carapace of these three species, they hardly will be confused with Leptuca thayeri or Minuca vocator. In the field, these species can also be distinguished by its respective habitat: while M. mordax is mostly found in freshwater or oligohaline areas such as river banks and tidal flats near river mouth, M. rapax and M. burgersi are mainly found in mesohaline tidal flats, often in co-occurrence.
Carapace pentagonal moderately arched and provided with small and scarce tubercles in the antero-lateral corner (Fig.
The species reproduces year-round in the populations from northern and southeastern Brazil (
Morphologically very similar to M. burgersi and M. mordax. Minuca rapax can be distinguished from M. mordax in not having a pile around the entire surface of manus of 1st to 3rd ambulatory legs. The distinction between M. rapax and M. burgersi, however, requires an extra attention: both species have piles limited to the dorsal surface of carpus and manus of 1st to 3rd ambulatory legs. The easiest way to distinguish these two species is to compare the last ambulatory leg: while M. rapax has a wide merus with convex dorsal margin (Fig.
Carapace pentagonal moderately arched and provided with few tubercles on the surface of antero-lateral corners and on mesogastric area (Fig.
Although with a wide geographical distribution, the species forms sparse populations constituted by small individuals in impacted mangroves of southeastern Brazil (
The easiest way to recognize this species is to examine the presence of a short and straight depression filled with a pile at the pollex base in the male major claw (Fig.
Carapace pentagonal moderately arched; profuse pile on dorsal surface forming a characteristic pattern mostly on hepatic and branchial regions (Fig.
The species forms one of the densest populations composed by large crabs, and its reproductive period coincides with the rainy period in northern Brazil (
Characteristic pubescence pattern on the carapace and dense piles on dorsal surface of ambulatory legs are the best diagnostic characters for distinguishing it from other Minuca species recorded in Brazil. Another Brazilian fiddler crab that has an exuberant pile on the carapace surface is Leptuca thayeri, easily distinguishable from M. vocator by a very narrow front of the former species (compare Figs
Carapace semi-cylindrical, width ca. 1.6 times the length; strongly arched and dorsal surface without pile; lateral margins almost parallel (Fig.
The species reproduces year-round in populations of northern Brazil and the crabs reach the largest CW among all other populations (
In sympatric area of Brazilian coast L. cumulanta can be confused with L. leptodactyla (from Pará to Rio de Janeiro) and L. uruguayensis (Rio de Janeiro state): they are similar in size and the major male claw is provided with a very curved dactyl forming a wide gap with the pollex. The best ways to distinguish these three species is described when referring to L. leptodactyla and L. uruguayensis (see below).
Carapace semi-cylindrical, width ca. 1.7 times the length; strongly arched and dorsal surface without any pile or other ornaments; lateral margins almost parallel (Fig.
One of the most common fiddler crabs in sandy substrate of estuarine intertidal zone, L. leptodactyla reproduces year-round in the population of Ceará state (Bezerra and Matthews-Cascon, 2007), but only in warmer months at the southern coast (
Leptuca leptodactyla may be confused with L. cumulanta at the coast from Pará to Rio de Janeiro states. These two species, however, can be distinguished by features of the male abdomen: the middle somites are fused in the former species (Fig.
Carapace trapezoidal weakly arched and covered with exuberant pile (pubescence easily detached) (Fig.
Populations living in the northeastern Brazilian coast reproduce only in the rainy season (Ceará state) (
This species is hardly confused with other species of Brazilian fiddler crabs, as it has a very narrow triangular front (see Fig.
Carapace semi-pentagonal strongly arched and dorsal surface without pile or other ornaments (Fig.
The species reproduces year-round in southeastern (
Leptuca uruguayensis can be confused with L. cumulanta and L. leptodactyla in Rio de Janeiro state coast (these species are sympatric) due to the small size attained by these three species. The easiest way to separate them is by observing the male abdominal segments: among these three species only L. cumulanta has all somites distinct (see Fig.
Carapace trapezoidal moderately arched and naked dorsal surface, without any ornaments. Dorso-lateral margins well marked, long and weakly converging posteriorly; postero-lateral striae absent (Fig.
The species reproduces year-round in northern (
In the field, these crabs are unmistakable recognizable by the flattened fingers of the male major claw. Furthermore, they are visibly larger than any other Brazilian fiddler crab species.
Most Brazilian fiddler crabs occur along the coastal estuaries in environments closely related to mangroves that are distributed in the country from Amapá state to Laguna do Imaruí in the municipality of Laguna (20°30’S), Santa Catarina state (
We hypothesized that in Piauí State and others, where records of some common fiddler crab species are missing (Table
We are grateful to Prof. Marcelo Antonio Amaro Pinheiro from Biosciences Institute of São Paulo State University-UNESP for providing us specimens of the fiddler crabs Minuca victoriana and M. vocator, and for Prof. Luís Ernesto Arruda Bezerra from Marine Science Institute of the Federal University of Ceará for critical review. Thanks are also due to Prof. Luís Amilton Foerster from Zoology Department of Federal University of Paraná for the English revision. Some undergraduate students were invited to test the present key, to whom we are so grateful. All biological sampling of the present study complies with the current laws of Paraná State and Brazilian Federal Government, which was conducted with the permission of SISBIO (Authorization system and information on biodiversity license N° 16140-1).