All Superfamilies, Families, and Species of Brachyurans Found in the Mangroves Are Listed

Appendix 1

All superfamilies, families, and species of brachyurans found in the mangroves are listed below, together with their geographical distribution, habitat, behavior, and life cycle. The information about geographical distribution and certain ecological aspects was taken from Melo (1996).

Superfamily: Xanthoidea MacLeay 1838

Family: Panopeidae Ortmann 1893

Eurytium limosum (Say 1818)

Geographical distribution: Western Atlantic – Bermuda, Florida, Gulf of

Mexico, Central America, Antilles, northern South America, and Brazil

(from Pará to Santa Catarina).

Habitat and behavior: Mud bottoms of mangroves. These crabs live in

burrows constructed by themselves or by other species. The burrows are

partly filled with water and are located beneath mangrove roots or

stones. They are scavengers and can also consume invertebrates,

mainly mollusks.

Life cycle: Four zoea stages and one megalopa (Kurata et al. 1981;

Messerknecht et al. 1991). At the fourth juvenile stage it is possible to

determine the sex of the crab by the external morphology of the

abdominal appendages (Guimarães and Negreiros-Fransozo 2005).

Panopeus austrobesus (Williams 1983)

Geographical distribution: Western Atlantic – Brazil (from Rio de Janeiro to Rio Grande do Sul).

Habitat and behavior: Builds burrows in mud sediments of mangroves,

beneath stones or mangrove roots. The chelipeds are adapted to

manipulate and open mollusk shells. Mollusks may be the main food

Life cycle: Four zoea stages and one megalopa (Montú et al. 1988). The

juvenile development was described by Hebling et al. (1982; as P.

herbstii). It is possible to recognize the sex beginning with the 4th juvenile

stage.

Panopeus occidentalis (Saussure, 1857)

Geographical distribution: Western Atlantic – from North Carolina to

Florida, Gulf of Mexico, Central America, Antilles, northern South

America, Guyana, and Brazil (from Ceará to Santa Catarina).

Habitat and behavior: This species lives in sand bottoms, under rocks or

mangrove roots. They are scavengers, and can be predators of

invertebrates.

Life cycle: Four zoea stages and one megalopa (Ingle, 1985).

Superfamily: Pinnotheroidea De Haan 1833

Family: Pinnotheridae De Haan 1833

Austinixa patagoniensis (Rathbun 1918)

Geographical distribution: Western Atlantic – Brazil (from Rio de Janeiro

to Rio Grande do Sul), Uruguay, and Argentina.

Habitat and behavior: Lives on sandy bottoms, and can live in tubes of Callichirus (Thalassinidea, Crustacea) and Arenicola (Polychaeta).

Life cycle: Five zoea stages and one megalopa (Boschi 1981).

Zaops ostreuus (Say 1817)

Geographical distribution: Western Atlantic – from Massachusetts to

southern Florida, Gulf of Mexico, Antilles, and Brazil (from Pernambuco

to Santa Catarina).

Habitat and behavior: The adult lives commensally in bivalves, mainly

oysters of the genus Crassostrea. Probably it feeds on food particles

from inhalant currents.

Life cycle: Four zoea stages and one megalopa (Sandifer 1972).

Superfamily: Grapsoidea MacLeay 1839

Family: Gecarcinidae MacLeay 1839

Cardisoma guanbumi Latreille 1825

Mexico, Antilles, Colombia, Venezuela, and Brazil (from Ceará to São

Paulo).

Habitat and behavior: Builds deep burrows in the upper (inland) zone of

the mangroves, but in areas where seawater may reach. Mainly

herbivorous, but may also act as a scavenger.

Life cycle: Five zoea stages and one megalopa (Costlow and Bookhout

1968).

Family: Grapsidae MacLeay 1838 Goniopsis cruentata (Latreille 1803)

Mexico, Antilles, Guyana, and Brazil (Fernando de Noronha and from

Pará to Santa Catarina). Eastern Atlantic – from Senegal to Angola.

Habitat and behavior: In mangroves, on or under the roots and trunks. It does not build burrows. It has predates upon invertebrates, including other crabs.

Life cycle: Only the first zoea stage is known (Fransozo et al. 1998).

Pachygrapsus gracilis (Saussure 1858)

Geographical distribution: Western Atlantic – North Carolina, Florida,

Gulf of Mexico, Antilles, northern South America, Guyana, and Brazil

(from Ceará to Rio Grande do Sul). Eastern Atlantic – Cabo Verde Island to Angola; Mediterranean Sea; Eastern Pacific (California to Peru).

Habitat and behavior: Lives on rocky bottoms, and occasionally among mangrove roots and on sandy beaches. Herbivorous, feeds on algae.

Life cycle: Only 13 zoea stages are known (Brossi-Garcia and Rodrigues

1993).

Pachygrapsus transversus (Gibbes 1850)

Geographical distribution: Western Atlantic – Bermuda, from North

Carolina to Florida, Gulf of Mexico, Antilles, northern South America,

Brazil (from Ceará to Rio Grande do Sul).

Habitat and behavior: In the intertidal zone, under stones, on sandy beaches, and among mangrove roots. Life cycle: Eleven zoea stages (Cuesta and Rodriguez 1994) and one

megalopa. It is possible to recognize the sex beginning with the second

juvenile stage (Flores et al. 1998).

Family: Sesarmidae Dana 1851

Aratus pisonii (H. Milne Edwards 1837)

Geographical distribution: Western Atlantic – Florida, Gulf of Mexico,

Antilles, northern South America, Guyana, and Brazil (from Piauí to São

Paulo). Eastern Pacific – from Nicaragua to Peru.

Habitat and behavior: Lives on roots, branches, and trunks of

mangroves. It is mainly herbivorous.

Life cycle: Four zoea stages and one megalopa (Díaz and Bevilacqua

1987).

Armases angustipes (Dana 1852)

Geographical distribution: Western Atlantic – Mexico, Antilles, and Brazil

(from Ceará to Santa Catarina).

Habitat and behavior: Lives in the supratidal zone of estuaries, near a

freshwater source. Also lives on rocky bottoms and in bromeliads.

Omnivorous, feeds on animal remains and plants.

Life cycle: Four zoea stages and one megalopa (Cuesta & Anger 2001).

Armases rubripes (Rathbun 1897)

Geographical distribution: Western Atlantic – Central America, northern

South America, Guyana, Brazil (from Ceará to Rio Grande do Sul), Uruguay, and Argentina.

Habitat and behavior: Lives on the sediment or among the roots,

branches, and trunks of mangroves. It feeds on algae that live on

mangrove roots.

Life cycle: Five zoea stages and one megalopa (Díaz and Ewald 1968;

Montú et al. 1990).

Sesarma rectum Randall 1840

Geographical distribution: Western Atlantic – Venezuela, Guyana, and

Brazil (from Amapá to Santa Catarina).

Habitat and behavior: Lives in the upper zone of the mangroves. It

constructs a shallow burrow under trees (Hibiscus tiliaceus L.), in grassy

areas or in drier zones. It feeds on animal remains and plants.

Life cycle: Three zoea stages and one megalopa (Fransozo and Hebling

1986). From the 12th juvenile stage it is possible to distinguish the sex

(Fransozo 1987) by the secondary sexual characters (abdomen shape,

and number of abdominal appendages).

Family: Varunidae H. Milne Edwards 1837

Neohelice granulata (Dana 1851)

Geographical distribution: Western Atlantic – Brazil (from Rio de Janeiro

to Rio Grande do Sul), Uruguay, and Argentina.

Habitat and behavior: Lives on muddy sediments, where it constructs a

shallow burrow. Semi-terrestrial and can remain for several hours under

stones or tree trunks. Life cycle: Four or five zoea stages and one megalopa (Boschi et al.

1967).

Cyclograpsus integer (H. Milne Edwards 1853)

Central America, northern South America, and Brazil (from Ceará to

Santa Catarina); Eastern Atlantic (from Cabo Verde Islands to Senegal);

Indo-Pacific.

Habitat and behavior: Lives in burrows in the marine supratidal zone, and

in the intertidal zone on stony beaches. Also lives in estuaries.

Life cycle: Five or six zoea stages and one megalopa (Gore and Scotto

1982).

Superfamily: Ocypodoidea Rafinesque 1815

Family: Ocypodidae Rafinesque 1815

Uca (Leptuca) cumulanta Crane 1943

Geographical distribution: Western Atlantic – Central America, northern

South America, Guyana, and Brazil (from Pará to Rio de Janeiro). This is

the first record of this species for the state of São Paulo.

Habitat and behavior: Lives on mud or sand-mud beaches, near

mangrove trees, but not beneath the canopy. Populations are generally

concentrated below mid-tide level.

Life cycle: Beginning with the 3rd juvenile stage it is possible to

distinguish the sex (Hirose et al. 2010) by the secondary sex characters

(abdomen and cheliped shape). Uca (Leptuca) leptodactylus Rathbun 1898

Antilles, Venezuela, and Brazil (from Maranhão to Santa Catarina).

Habitat and behavior: Lives on sandy bottoms, near the sea. Deposit feeder.

Life cycle: Unknown.

Uca (Leptuca) uruguayensis Nobili 1901

Geographical distribution: Western Atlantic – Brazil (from Rio de Janeiro to Rio Grande do Sul), Uruguay, and Argentina.

Habitat and behavior: Lives on sandy bottoms with small amounts of mud, near the sea and river mouths. It lives in burrows at the edges of mangroves. Deposit feeder.

Life cycle: Five to six zoea stages and one megalopa (Rieger 1992).

Uca (Minuca) burgersi Holthuis 1967

Antilles, Venezuela, and Brazil (from Maranhão to São Paulo).

Habitat and behavior: Burrows in low vegetation (mainly Poaceae) of mangrove edges. Deposit feeder.

Life cycle: Four zoea stages and one megalopa (Rieger 1992). Its juvenile development was recently described by Vieira et al. (2010). It is possible to recognize the sex from the 4th juvenile stage on. Uca (Minuca) mordax (Smith 1870)

Geographical distribution: Western Atlantic – Gulf of Mexico, Central

America, northern South America, Guyana, and Brazil (from Pará to São

Paulo).

Habitat and behavior: Burrows in low vegetation (mainly Poaceae) of the middle zone in mangroves. Deposit feeder.

Life cycle: Five to six zoea stages and one megalopa (Rieger 1992).

Uca (Minuca) rapax (Smith 1870)

Antilles, Venezuela, and Brazil (from Pará to Santa Catarina).

Habitat and behavior: Burrows in sediments of different grain sizes. It lives among vegetation in mangrove edges. Deposit feeder.

Uca (Minuca) thayeri Rathbun 1900

Antilles, Guatemala, Panama, Venezuela, and Brazil (from Maranhão to

Santa Catarina).

Habitat and behavior: Lives mainly in mud or sand-mud bottoms, near rivers, sometimes in shaded areas. The burrow is generally elevated above the sediment like a chimney. Deposit feeder.

Life cycle: Four zoea stages and one megalopa (Anger et al. 1990).

Uca (Minuca) vocator (Herbst 1804) Geographical distribution: Western Atlantic – Gulf of Mexico, Central

America, Antilles, northern South America, Guyana, and Brazil (from

Pernambuco to Santa Catarina).

Habitat and behavior: Lives mainly in mud sediments, near rivers,

generally in shaded zones. The burrow has two openings, 10 cm apart.

Deposit feeder.

Life cycle: Four, five, or six zoea stages and one megalopa (Rieger

1992).

Family: Ucididae Števčić 2005

Ucides cordatus (Linnaeus 1763)

Central America, Antilles, northern South America, Guyana, and Brazil

(from Pará to Santa Catarina).

Habitat and behavior: Lives on mud sediments among mangrove roots.

The burrows are generally deep and straight. It feeds on remains of

animals and plants.

Life cycle: Five or six zoea stages and one megalopa (Rodrigues and

Hebling 1989).

Superfamily: Portunoidea Rafinesque 1815

Family: Portunidae Rafinesque 1815

Callinectes boucorti A. Milne Edwards 1879

Antilles, Colombia, Venezuela, Guyana, and Brazil (from Amapá to Santa Catarina).

Habitat and behavior: Lives on subtidal unconsolidated sediments of estuaries, and in low-salinity water. Lives on mud, sand, and rocky bottoms, and can live on bottoms with mollusk shells, at depths to 20 m.

An opportunist predator, feeds on invertebrates and small vertebrates.

Callinectes danae Smith 1869

Mexico, Antilles, Colombia, Venezuela, and Brazil (from Paraíba to Rio

Grande do Sul).

Habitat and behavior: Lives on unconsolidated subtidal sediments and tolerates changes in salinity. Present in estuaries. An opportunist predator, feeds on invertebrates and small vertebrates.

Callinectes exasperatus (Gerstaecker 1856)

Mexico, Antilles, Colombia, Venezuela, and Brazil (from Maranhão to

Santa Catarina).

Habitat and behavior: Lives on unconsolidated subtidal sediments and tolerates changes in salinity. Present in estuaries and creeks to depths of

8 m. An opportunist predator, feeds on invertebrates and small vertebrates.

Life cycle: Unknown. Callinectes sapidus Rathbun 1896

Geographical distribution: Western Atlantic – east coast of USA to

Florida, Gulf of Mexico, Antilles, Central America, Venezuela, Brazil (from

Bahia to Rio Grande do Sul), and Argentina. Eastern Atlantic, North Sea,

Mediterranean Sea, Adriatic Sea, Black Sea, and Indo-Pacific (Japan).

Habitat and behavior: Lives on unconsolidated subtidal sediments. An

opportunist predator, feeds on invertebrates and small vertebrates.

Life cycle: Seven or eight zoea stages and one megalopa (Costlow and

Bookhout 1959).

Literature cited

Anger K, Montú M, Bakker C, et al. (1990) Larval development of Uca thayeri

Rathbun, 1900 (Decapoda: Ocypodidae) reared in the laboratory.

Meeresforschung 32:276-294.

Boschi EE (1981) Morphology of the first zoeal stage of the commensal southwestern Atlantic crab Austinixa aidae (Righi 1967) (Brachyura:

Pinnotheridae), hatched in the laboratory. In: Boltovskoy D (ed) Atlas del

Zooplancton del Atlántico Sudoccidental, vol 38. Contribuciones del Instituto

Nacional de Investigaciones y Desarrollo Pesquero, Mar del Plata, pp 699–757.

Boschi EE, Scelzo MA, Goldstein B (1967) Desarrollo larval de dos especies de

Crustáceos Decápodos en el laboratorio, Pachycheles haigae Rodrigues da

Costa (Porcellanidae) y Chasmagnathus granulata Dana (Grapsidae). Bol Inst

Biol Mar (Mar del Plata) 12:4-46.

Brossi-Garcia AL, Rodrigues MD (1993) Zoeal morphology of Pachygrapsus gracilis (Saussure, 1858) (Decapoda, Grapsidae) reared in the laboratory. Invert

Reprod Dev 24:197-204.

Costlow JD Jr, Bookhout CG (1959) The larval development of Callinectes sapidus Rathbun reared in the laboratory. Biol Bull 116:373-396.

Costlow JD Jr, Bookhout CG (1968) The complete larval development of the land-crab, Cardisoma guanhumi Latreille in the laboratory (Brachyura,

Gecarcinidae). Crustaceana (Suppl. 2):259-270.

Cuesta JA, Rodríguez A (1994) Early zoeal stages of Pachygrapsus marmoratus (Fabricius), P. transversus (Gibbes) and P. maurus (Lucas)

(Decapoda, Brachyura, Grapsidae) reared in the laboratory. Sci Mar 58:323-

Cuesta JA, Anger K. (2001) Larval morphology of the Sesarmid crab Armases angustipes Dana, 1852 (Decapoda, Brachyura, Grapsoidea). J.Crust Biol

21(3):821-838

Díaz H, Bevilacqua M (1987) Early developmental sequences of Aratus pisonii

(H. Milne Edwards) (Brachyura: Grapsidae) under laboratory conditions. J

Coastal Res 3:63-70

Díaz H, Ewald JJ (1968) A comparison of the larval development of

Metasesarma rubripes (Rathbun) and Sesarma ricordi H. Milne Edwards

(Brachyura, Grapsidae) reared under similar laboratory conditions. Crustaceana

2:225-248.

Flores AAV, Negreiros- Fransozo ML, Fransozo A (1998) The megalopa and juvenile development of Pachygrapsus transversus (Gibbes, 1850) (Decapoda,

Brachyura) compared with other grapsid crabs. Crustaceana 71(2):197-222.

Fransozo A (1987) Desenvolvimento larval de Eriphia gonagra (Fabricius, 1781) (Decapoda, Xanthidae), em laboratório. Rev Bras Zool 4:165-179.

Fransozo A, Cuesta JA, Negreiros-Fransozo ML (1998) First zoeal stage of two species of Grapsidae (Decapoda, Brachyura) and a key to such larvae from the

Brazilian coast. Crustaceana 71(3): 331-343.

Fransozo A, Hebling NJ (1986) Desenvolvimento larval de Sesarma

(Holometopus) rectum Randall, 1840 (Decapoda, Grapsidae), em laboratório.

Rev Bras Biol 46:353-364.

Gore RH, Scotto LE (1982) Cyclograpsus integer H. Milne Edwards, 1837

(Brachyura, Grapsidae): the complete larval development in the laboratory, with notes on larvae of the genus Cyclograpsus. Fish Bull 80:501-521.

Guimarães FJ, Negreiros-Fransozo ML (2005) Juvenile development and growth patterns in the mud crab Eurytium limosum (Say, 1818) (Decapoda,

Brachyura, Xanthidae) under laboratory conditions. J Nat Hist 39:2145-2161.

Hebling NJ, Fransozo A, Negreiros-Fransozo ML (1982) Desenvolvimento dos primeiros estágios juvenis de Panopeus herbstii H. Milne-Edwards, 1834

(Crustacea, Decapoda, Xanthidae) criados em laboratório. Naturalia 7:177-188.

Hirose GL, Bolla Jr EA, Negreiros-Fransozo ML (2010) Post-larval morphology, growth, and development of Uca cumulanta Crane, 1943 (Crustacea,

Decapoda, Ocypodidae) under laboratory conditions. Invertebr Reprod Dev

54(2):95-109.

Ingle RW (1985) Larval development of the mud crab Panopeus occidentalis de

Saussure, from Bermuda (Crustacea: Xanthoidea: Panopeidae). Bulletin of the

British Museum of Natural History (Zoology) 48:233–248.

Kurata H, Heard RW, Martin JW (1981) Larval development under laboratory conditions of the xanthid mud crab Eurytium limosum (Say, 1818) (Brachyura: Xanthidae) from Georgia. Gulf Res Rep 7:19-25.

Melo GAS (1996) Manual de Identificação dos Brachyura (Caranguejos e Siris) do Litoral Brasileiro. Plêiade, São Paulo.

Messerknecht I, Montú M, Anger K (1991) Larval development and salinity tolerance of Brazilian Eurytium limosum (Say, 1818) (Brachyura: Xanthidae) reared in the laboratory. Meeresforschung 33:275-296.

Montú M, Anger K, Bakker C (1990) Variability in the larval development of

Metasesarma rubripes (Decapoda, Grapsidae) reared in the laboratory. Nerítica

5:113-128.

Montú M, Anger K, Bakker C et al (1988) Larval development of the Brazilian mud crab Panopeus austrobesus Williams, 1983 (Decapoda: Xanthidae) reared in the laboratory. J Crust Biol 8:594-613.

Rieger PJ (1992) Desenvolvimento larval de Uca (Celuca) uruguayensis Nobili,

1901, Uca (Minuca) burgersi Holthuis, 1967, Uca (Minuca) mordax (Smith,

1870) e Uca (Minuca) vocator (Herbst, 1804) (Crustacea, Decapoda,

Ocypodidae), em laboratório. Doctoral thesis, Biosciences Institute, Rio Claro,

São Paulo, Brazil, Universidade Estadual Paulista.

Rodrigues MD, Hebling NJ (1989) Ucides cordatus (Linnaeus, 1763)

(Crustacea, Decapoda). Complete larval development under laboratory conditions and its systematic position. Rev Bras Zool 6:147-166.

Sandifer PA (1972) Morphology and ecology of Chesapeake Bay decapod crustacean larvae. Dissertation, University of Virginia.

Vieira, RRR, Pinho GLL, Rieger PJ (2010). Juvenile development of Uca

(Minuca) burgersi Holthuis, 1967 (Crustacea, Brachyura Ocypodidae) in the laboratory. Atlântica 32(1):59-70.