Species of Decapod Crustaceans and their distribution in the american marine zoogeographic provinces.

Item Type Journal Contribution

Authors Boschi, E.E.

Citation Revista de Investigación y Desarrollo Pesquero, 13. p. 1-136

Publisher Mar del Plata: Instituto Nacional de Investigacion y Desarrollo Pesquero

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Link to Item http://hdl.handle.net/1834/2606 ISSN N" 0325-6375

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Revista de Investigación y Desarrollo Pesquero N° 13 Mar del Plata, República Argentina

ResumidaJindizada en: Aquatic Sciences & Fisheries Abstracts (ASFA); Agrindex; INFOMARNAP; Marine, Oceanographic & Freshwater Resources; Wildlife Worldwide; Zoological Record Revista de Investigación y Desarrollo Pesquero

N° 13 - Agosto 2000

SPECIES OF DECAPOD CRUSTACEANS AND THEIR DISTRIBUTION IN THE AMERICAN MARINE ZOOGEOGRAPHIC PROVINCES

by

ENRIQUE E. BOSCHI CONTENTS

RESUMEN 7 SUMMARY 7 INTRODUCTION 8 MATERIALANDMETHODS 15 Arctic Province (1) 18 The N ortheastern Pacific 18 Provinces of temperate waters Aleutian Province (2) 18 Oregonian Province (3) 19 Californian Province (4) ,... ,. 19 The Eastern Central Pacific Ocean 19 Provinces and Subprovinces of subtropical and tropical waters Cortés Subprovince (5) 20 Panamic Province (6) 21 Galápagos Subprovince (7) " ,. 22 The Southeastern Pacific Ocean ,. .,... 22 Province of temperate waters Perú-Chilean Province (8) 22 The Southeastern Pacific Ocean and Southwestern Atlantic Ocean 23 Province of cold-temperate waters Magellanic Province (9) 23 The Southwestern Atlantic Ocean 24 Provinces of warm-temperate and subtropical-tropical waters Argentinian Province (10) 24 Brazilian Province (11) 25 The Caribbean and the Gulf ofMexico 25 Provinces of tropical, subtropical and temperate waters Caribbean Province (12) 26 Texan Province (13) 27 The N011hwestern Atlantic Ocean ,. 27 Provinces of temperate and cold-temperate waters Carolinian Province (14) 28 Virginian Province (15) 28 Boreal Province (16) 29 CONCLUSIONS 29 ACKNOWLEDGEMENTS 30 NOTE ADDED IN PROOF ,. 30 REFERENCES 32 APPENDIX ..,. 64 CONTENIDO

RESUMEN 7 SUMMARY 7 INTRODUCCION 8 MATERIALy METODOS 15 Provincia Artica (1) 18 Océano Pacífico Noreste 18 Provincias de aguas templadas

Provincia Aleutiana (2) '''''''''''''''''''''''''''''''''''''''''''''''' 18 Provincia Oregoniana (3) 19 Provincia Califomiana (4) 19 Océano Pacífico Central Este 19 Provincias y Subprovincias de aguas tropicales y subtropicales Subprovincia Cortés (5) 20 Provincia Panameña (6) 21 Subprovincia Galápagos (7) 22 Océano Pacífico Sudeste 22 Provincia de aguas templadas Provincia Peruano-Chilena (8) 22 Océanos Pacífico Sudeste y Atlántico Sudoeste 23 Provincia de aguas templadas frías Provincia Magallánica (9) 23 Océano Atlántico Sudoeste 24 '"''''''''''''''''''''''''''''''''''''''''''''''''''''''''''''''''"""""'''''''''''''''''' Provincias de aguas templadas cálidas y de aguas subtropicales y tropicales Provincia Argentina (10) 24 Provincia Brasileña (11 ) 25

Caribey Golfode México """'''''''''''''''''''''''''''''''''''' 25 Provincias de aguas tropicales, subtropicales y templadas Provincia Caribeña (12) 26 Provincia Texana (13) 27 Océano Atlántico Noroeste 27 Provincias de aguas templadas y templadas frías Provincia Caroliniana (14) 28 Provincia Virginiana (15) 28 Provincia Boreal (16) 29 CONCLUSIONES . 29 AGRADECIMIENTOS 30 ADDENDA 30 BIBILOGRAFIA 32 APENDICE 64 REv. INVEST.DES. PESQ. N° 13: 7-136 (2000) 7

SPECIES OF DECAPOD CRUSTACEANS AND THEIR DISTRIBUTION IN THE AMERICAN MARINE ZOOGEOGRAPHIC PROVINCES*

by

ENRIQUE E. BOSCHI

Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires and Instituto Nacional de Investigación y Desarrollo Pesquero (INIDEP) Paseo Victoria Ocampo N"1, Escollera Norte, (P.O. Box 175), 7600, Mar del Plata, Argentina. E-mail: [email protected]

RESUMEN

Las especies de Crustáceos Decápodos y su distribución en las provincias zoogeográficas marinas ameri- canas. Se han establecido catorce provincias y dos subprovincias zoogeográficas en las Américas sobre la base de la distribución de los crustáceos decápodos marinos de las aguas de las plataformas que incluyen el suprali- toral, mediolitoral, infralitoral y circalitoral. El número total de especi!:s en esta amplia región, que abarca ambas costasoceánicas,desdeel Cabode Hornosen el sur hastael OcéanoArticoen el norte,se estimaen 2.472.Los resultados de este estudio sobre biodiversidad y distribución de las especies de decápodos se ajusta al carac- terístico de los grupos de la epifauna marina con una gran amplitud latitudinal y un pronunciado decrecimiento del número de especies desde las regiones tropicales hacia las polares y con una cierta distribución clinal. En la Provincia Caribeña se halló la más alta densidad de especies que, en un número estimado de 1.058, fue mayor que el de la Provincia Panameña del Pacífico tropical donde se registraron 825 especies. En lo que concierne a la relación con la riqueza en especies según los grandes grupos de Decápodos, el número más alto se halló en los Brachyura con 1.090 especies seguido por los Caridea con 536 especies, los Anomura con 509, los Thalassinidea con 157 y los Penaeoidea con 96. Los restantes grupos están representados por un número inferior.

SUMMARY

Fourteen marine zoogeographic provinces and two subprovinces were established for the Americas. The num- ber of species of marine decapod crustaceans recorded in the supralittoral, intertidal, eulittoral and sublittoral (continental shelf to 200-300 m) including both, continental coasts from the Arctic Polar region in the north to Cabo de Hornos in the south is estimated at 2472 species. The results of this study on the biodiversity and dis- tribution of decapod species call attention to a common characteristic of typical epifaunal groups of organisms with a large range of latitudinal geographic distribution: a high number of species in tropical regions on both coasts and a pronounced decrease of taxa towards the polar regions that generates a clear clinal distribution. The Caribbean Province showed the largest diversity which, with a total of 1058 species was higher than that of the tropical Panamic Province where 825 species were registered. With regard to the species richness in the nine major groups of Decapods, the largest number was found in the Brachyura hovering 1090 estimated species fol- lowed by the Caridea with 536 species, the Anomura with 509 species, the Thalassinidea with 157 and the Penaeoidea with 96. The remaining four decapod taxa are represented only by a small number of species.

Key words: Marine zoogeography, Crustacea Decapoda, North, Central and South America. Palabras clave: Zoogeografía marina, Crustacea Decapoda, América del Norte, Central y del Sur.

Contribución INIDEP 1152 * N° 8 REV. INVEST.DES. PESQ. N° 13: 7-136 (2000)

lability, patterns of tides and sea level, composi- INTRODUCTION tion of the community, prey-predator relation- ships, interactions among species, reproduction strategies, etc. (Abele, 1974, Abele, 1976 a, The decapod crustaceans are invertebrates Brusca and Wallerstein, 1979, Gore et al., 1978, very frequently found in marine littoral and Kohn, 1997). In continental shelf waters, the brackish waters of the Americas. Most species are oceanic fronts originating in areas between cu- found in tropical and subtropical regions with a rrents and tidal fronts may constitute a barrier for marked decrease in numbers towards temperate the distribution of species (Fig 1,2). and cold regions. Distribution may depend on the The catalogue of species of decapod crus- influence of environmental factors. On the other taceans from littoral and coastal seas of both mar- hand, it may also be attributed to the stability of gins of the American continent has allowed to the ecosystems which, in the historical sense of determine the limits of different zoogeographical the evolution of the earth, may account for the provinces and to quantify the number of species existence of a large number of species in the tro- found in each of them. In Table 1 the distribution pical regions (Margalef, 1963). Nevertheless, it of species in the extensive areas under study is does not seem to be true in the presently indicated. Due to the fact that many of them are constrained biotopes (Abele, 1976 b). In more present in more than one province, it can be lirnited areas, with a few latitudinal differences, observed that the total data relative to species per variations and richness of the species may depend province and subprovince reach 5,350(1) entries on the characteristics of the bottom, food avai- (Fig 3,4,5).

(1)The number of species and records per province has changed during the course of this study due to publications of ta- xonomic revisions and changes in the distributions of species. For this reason, the numbers mentioned by the author in pre- vious papers are slightly different from the ones appearing in this study.

Figure 1. Zoogeographical provinces and subprovinces of the Americas and their limits. Arctic (1) between Nunivak Is. to Strait of Belle Isle; Aleutian (2) Nunivak Is. to Puget Sound; Oregonian (3) Puget Sound to Point Conception; Californian (4) Point Conception to Bahía Magdalena; Cortés (5) subprovince, from Tiburón and Angel de la Guarda Islands (Gulf of California) to north end of the Gulf; Panamic (6) Bahía Magdalena to Gulf of GuayaquilJBahía Sechura; Galápagos (7) subprovince, Lat. 0040'N and 1030'S-long. 89°20'W and 91°50'W; Perú-Chilean (8) Bahía Sechura to the north of Chiloé Is.; Magellanic (9) Chiloé Is. to 35° S Southwest Atlantic; Argentinian (10) 43/44° S to Cabo Frío; Brazilian (11) Cabo Frío to Delta Río Orinoco; Caribbean (12) Delta Río Orinoco to Cabo Rojo, Gulf of Mexico, Caribbean Islands and Cape Romano to Cape Canaveral, Florida Peninsula, Bermuda Is.; Texan (13) Cabo Rojo to Cape Romano, G. of Mexico; Carolinian (14) Cape Canaveral to Cape Hatteras; Virginian (15) Cape Hatteras to Cape Cod; Boreal (16) Cape Cod to Strait of Belle Isle. Figura 1. Provincias y subprovincias zoogeográficas de las Américas y sus límites. Artica (1) entre la Isla Nunivak al Estrecho de Belle Isle; Aleutiana (2) Isla Nunivak a Puget Sound; Oregoniana (3) Puget Sound a Punta Concepción; Californiana (4) Punta Concepción a Bahía Magdalena; subprovincia Cortés (5), en el golfo de California, desde las Islas Tiburón y Angel de la Guarda hasta el extremo norte del golfo; Panameña (6) Bahía Magdalena al golfo de Guayaquil/Bahía Sechura; subprovincia Galápagos (7), Lat. 0°40 'N y 1030'S-long. 89°20'W y 9r50'W; Peruano-Chilena (8) Bahía Sechura hasta el norte de la isla Chiloé; Magallánica (9) isla Chiloé hasta 35° S en el Atlántico Sudoeste; Argentina (10) 43/44° S a Cabo Frío; Brasileña (11) Cabo Frío hasta el delta del río Orinoco; Caribeña (12) delta del río Orinoco a Cabo Rojo, golfo de México, islas del Caribe y cabo Romano a cabo Cañaveral, península de Florida, islas Bermudas; Texana (13) cabo Rojo a cabo Romano, golfo de México; Caroliniana (14) cabo Cañaveral a cabo Hatteras; Virginiana (15) cabo Hatteras a cabo Cod; Boreal (16) cabo Cod a estrecho de Belle Isle. BOSCHI: DISTRIBUTION OF DECAPOD CRUSTACEANS 9

20"

O"

20" 10 REV. INVEST.DES. PEsQ. N" 13: 7-136 (2000)

1200

1000

Q)en 800

'üQ) g. 600 Z

400

200

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 (.) e c: c: (1) (J UJ r:::: CJ c: c: c: c: c: c: (ij ~ ~ .~ .~ ~'E&m.~.~~ « j 8~~:EQ):g~ ..c~~:~:~~F &.gQ) COCOUoQ)L... "'5 ~ tD « -te .~ ~ := [L~2~E>aJ () ()ea:> O ()'" Q) « . 11. Zoogeographic provinces/*subprovinces

Figure 2. Total number of decapod species per province and subprovince. Figura 2. Número total de especies de crustáceos decápodos por provincia y subprovincia.

Table 1.Distribution per province/*subprovince and group of Decapod Crustacean species of the Americas. Tabla l. Distribución por provincia/*subprovincia por grupos de crustáceos decápodos en las Américas.

Prov. Penaeoidea Sergestoidea Stenopodidea CarideaAstacideaThalassinidea Palinura Anomura Brachyura Total

2 1 85 5 1 56 32 182

4 4 74 8 2 67 126

7 3 161 34 7 165

12 3 41 3 7 43

19 5 43 2 12 3 52 194 330

50 15 8 224 4 83 15 205 454

4 2 82 1 8 7 60

4 37 1 3 1 11 Total 292 6 21 1167 15 234 73 997 2485 5350 BOSCHI: D¡STRIBUTION OF DECAPOD CRUSTACEANS II

Penaeoidea

Q) "(3Q)'" a. Z'"

2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Sergestoidea

16 14 12

Q) 10 "(3Q)'" a. Z'" 6 4

2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Stenopodidea

9 8 7 6 Q) 5 "(3Q)'" a. 4 Z'" 3 2 1 O 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Figure 3. Distribution of Decapod Crustacean species by group for each province. Figura 3. Distribución de las especies de crustáceos decápodos por grupos y provincia. 12 REV. INVEST. DES. PESQ. N° 13: 7-136 (2000)

Caridea

250

200

(/) ID 150 '13 ID (/)c. z 100

50

2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Astacidea

6

5

4 (/) ID '13 ID 3 (/)C. z 2

...:::::::::::"..:.:::::::V':..:::::~::::".,:.:::::::::"",.:::::::::::.,,...:::::::::...,, ,.:::::::::::., ..,::::~ o 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Thalassinidea

80 70 60 (/) 50 ID '13 ID 40 (/)c.

Z 30 20 10 O 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 provinces

Figure 4. Distribution of Decapod Crustacean species by group for each province. Figura 4. Distribución de las especies de crustáceos decápodos por grupos y provincia. BOSCHI: DISTRIBUTIONOF DECAPODCRUSTACEANS 13

Palinura

16 14 12

Q)V) 10 .13Q) 8 V)o. z 6 4 2 O 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Anomura

200 180 160 140 ~120 .~ 100 V)o. z 80 60 40 20 O 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Brachyura

500 -/í 450-vr 400-r 350 ~300 .~ o. 250 ~200 150 100 50 O 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Provinces

Figure 5. Distribution of Decapod Crustacean species by group for each province. Figura 5. Distribución de las especies de crustáceos decápodos por grupos y provincia. 14 REY. INYEST.DES. PEsQ. N" 13: 7-136 (2000)

The total number of species by group is 2472 classical biogeography so they are not followed in (Table 2). In addition, data on the distribution of the present analysis. decapods are compared to those of other marine The extreme variation of the size of continen- orgamsms. tal shelves in the marine littoral of the Americas is a phenomenon that appears to have an influence Table 2. Total number of marine decapod crustacean on the presence and distribution of different species found on continental shelf of Americas by group. graups of decapod species. In the eastern margin Tabla 2. Número total de crustáceos decápodos marinos of the continent, in the Southwestern Atlantic, hallados en las plataformas continentales de las Américas por grupos. there is a tendency to find extensive continental shelves. In consequence, the adjacent sea is not deep, with a smooth transition to slope depths of Groups Number of species only 150-200 m. For example, in the South Atlantic of the Patagonian region of Argentina the Penaeoidea 96 largest extension of the shelf reaches 850 km. In Sergestoidea 29 the nor-theastern region of Brazil, from Salvador, Stenopodidea 12 Bahia State, to Río Grande do Norte, the conti- Caridea 536 nental shelf is significantly reduced. The same 7 Astacidea happens in the southeastern side of the Florida Thalassinidea 157 Palinura 36 Peninsula. In contrast, the continental shelves off Anomura 509 west Florida and Yucatán Peninsulas are wide Brachyura 1090 (Martinez-López and Parés-Sierra, 1998). In the western coasts of the continent, in the Total 2472 East Pacific Ocean, shelves are markedly reduced; in consequence, the seas adjacent to the continent are very deep near the coast. In the li- The fact that component species vary in their ttoral of Chile, fram Cabo de Hornos to Chiloé physiological tolerances and that, for that reason Island, the shelf is wider in some areas. Along the they are found in more than one pravince, makes rest of the coast the shelf is reduced or nonexis- it difficult to establish boundaries. AIso, opinions tent. This feature extends to the Alaska Peninsula. and criteria of biologists on what constitutes a In northwestern Alaska, Bering Sea, the area of pravince can vary significantly. Eurytopic species continental shelf is extensive. In the Gulf of show an extensive distribution in the latitudinal California, the continental shelves along the sense and are present in vast regions of the littoral coasts of Hermosillo and Nayarit are relatively so they are not very usefui for biogeographical wide with a smooth decline. In contrast, on the studies. Depth represents another confounding west coast, between Angel de la Guardia Island factor; therefore, only littoral and coastal species up to Cape San Lucas, the continental shelf is are considered (see Material and Methods). It can very narrow or absent. In the northern part of the be generally stated that the number of pravinces Gulf, north of Tiburón and Angel de la Guardia and their limits in the American continent coin- Islands, the continental shelf is generally smooth, cide. reaching depths of 150-200 m (Roden and On the other hand, Longhurst (1998), referring Graves, 1959, Parker, 1963). to the pelagic biogeography, established biomes Inaccuracy of geographical data pravided in and provinces based on oceanographic character- many studies makes it extremely difficult to istics and algal ecology that do not agree with establish species distributions with some precision. BOSCHI: DrSTRIBUTION OF DECAPOD CRUSTACEANS 15

In some cases, there is lack of data on latitude, correspond to floristic and faunal units with pro- longitude and depth. This is especially observed nounced endemism of taxa at the generic or hi- in the Gulfs of Mexico and California where, gher levels (Balech and EhrIich, MS). sometimes, it is not indicated if species are found In this paper, a Province is defined as a part of in the north or south, making it difficult to esta- the neritic zone with a relatively narrow range of blish which province the species belong to. temperatures where the fauna shows certain Considering that temperature is an important homogeneity. The term "subregion" is used as the factor for the distribution of many species, data equivalent of a province. In many cases, the li- on the temperature where the species are found mits between provinces are flexible due to fluc- may provide useful information on environmental tuations that occur in the distribution of species in conditions. different developmental stages such as larvae, The main aim of this contribution is to esti- postlarvae, juveniles and adults. mate the number of species of marine decapod They are produced by environmental changes, crustaceans living in continental shelf waters pre- colonization of new areas, geographic expansion sent in the Americas and surnmarize their known of the species, etc. The "subprovince" denomina- distributions. tion was given to areas with high species affinity between adjacent provinces. In species found in various provinces it is MATERIAL AND METHODS observed that the distribution limit includes only a part of the last province occupied or the transi- tion area between them. The data used in this study are based on infor- The stenohaline and stenothermal species are mation from the literature available to the author the most sensitive and useful to determine the on decapod crustaceans and their distribution on limits of each province. The euryhaline and eury- the continental shelves of the eastern and western thermal species are not as useful for these pur- coasts of North, Central and South America, poses. Although the above mentioned difficulties between latitudes 70° N and 56° S, approximate- are encountered, the best method is to measure ly. In addition, studies on the biogeography of the the number of endemic species for each province littoral system of other groups of marine organ- (Table 3). Briggs (1974) considered that an area isms were also considered. defined with 10% or more endemic species may The area included consisted of the continental be considered as a province. Nevertheless, this shelf to a depth of 200-300 m which included the criterion is not acceptable because said value is supra, medium, infralittoral and, in some areas, established in an arbitrary manner. the circalittoral (Péres, 1961, Sverdrup et al., 1955). Although in some cases the names of the For authors of English language, this co- provinces are different, the results of this study on rresponds to the supralittoral, eulittoral and sub- the number and characteristics of the zoogeo- littoral zones. In some cases, species that are graphical provinces coincide to a certain degree found up to 400 m are included. Naturally, the with other studies on this subject (e.g., Ekman, neritic species found at greater depths were also 1953, Briggs, 1974, etc.). Moreover, according to included. Additionally, species which are present the author's criterion, some variations in the geo- in both Americas as well as in other regions of the graphic limits and the groups of organisms under WorId Ocean are considered. From a biogeo- consideration were observed. The areas about graphical point of view, the major divisions of the which most differences of opinions exist are the sea are designated as Regions which, in general, Gulfs of California and Mexico. 16 REV. 1NVEST.DES. PESQ. N" 13: 7-136 (2000)

Table 3. Endemic species by group (* Subprovince). Tabla 3. Especies endémicas por grupo (* Subprovincia).

Prov. Records Endemics % Anomura % Astacidea % Brachyura % Endemic Endemic Endemic

41 22.53 14 7.69 4

48 15.95 12 3.99

62 7.52

24

31.95 79 7.47

5 1.30 O 0.00 1

4 5.19 O 0.00 O

Prov. Records Caridea % Pa1inura % Penaeoidea % Endemic Endemic Endemic

2 22 12.09 O 0.00 O

4 301 11 3.65 O 3

6 7.03 O 8

8 13 6.13 6

10 1.82 1 0.30

12 1058 93 8.79 1 0.09 5

14 2 0.52 O 0.00 O

16 4 5.19 O 0.00 O 0.00 BOSCHI: DISTRIBUTION OF DECAPOD CRUSTACEANS 17

Table 3. Continued. Tabla 3. Continuación.

Prov. Records Sergestoidea % Stenopodidea % Thalassinidea % Endernic Endemic Endernic

2 182 O 0.00 O 0.00 0.55

4 301 1 0.33 O 0.00 4 1.33

6 825 3 0.36 1 0.12 21 2.55

8 212 1 0.47 O 0.00 2 0.94

10 330 1 0.30 O 0.00 2 0.61

12 1058 10 0.95 4 0.38 52 4.91

16 77 O 0.00 O 0.00 O 0.00

The enumeration of most decapods of the Provinces N spp by prov. regions under study and their known and current scientific name required consultation of many 1. Arctic 37 studies published until 1999 which are included 2. Aleutian 182 in the bibliography. It is possible that some new 3. Oregonian 193 species are not included in the checklist because 4. Californian 301 265 they could have be en published in joumals of 5. Cortés* 6. Panamic 825 limited distribution or not accessible to the 7. Galápagos* 253 author. 8. Peru-Chilean 212 Moreover, there could be undescribed species 9. Magellanic 79 which could significantly increase the total num- 10. Argentinian 330 ber. In this study, a total of fourteen provinces 11. Brazilian 572 and two subprovinces were established (Table 4, 12. Caribbean 1058 Fig. 1, 2). 13. Texan 422 14. Carolinian 386 Table 4. Total decapod species per province and sub- 15. Virginian 158 province (* Subprovince). 16. Boreal 77 Tabla 4. Total de especies de decápodos por provincia y Record s 5350 subprovincia (* Subprovincia). 18 REY. 1NYEST.DES. PEsQ. N" 13: 7-136 (2000)

Arctic Province (1) The Northeastern Pacific Ocean Provinces oi temperate waters The Arctic Province of the Americas extends from the center of the Strait of Bering, Nunivak Although different names were used and some Island (600N) to the Labrador Peninsula, north disparity in limits was observed, the temperate Strait of Belle Isle (Lat. 51°37'N). coastal waters of the Northeastern Pacific Ocean The ecological conditions there do not vary as far south as the California Peninsula (Bahía much due to very small temperature oscillations Magdalena) were divided into three zoogeograph- maintaining values always around O°c. ic provinces according to the criteria of several For the subdivisions of the region the position authors. These are: the Aleutian, Oregonian and of the isotherms of O°C and 5°C between the high Californian Provinces. Arctic and low Arctic was taken into account The whole region which extends between 60° (Ekman, 1953, Briggs, 1974). and 25° N is under the intense influence of the In this study, the distribution of decapods in magnitude and direction of the Alaska and the whole province will not be discussed. Only California Currents originating in the Subarctic the presence of relatively few species of decapod Current and its corresponding oceanographic con- crustaceans known in the American area, consi- ditions. The changes in the hydrological regime, dered as a very young region of very recent set- temperature and salinity of the water, isolation, tlement, will be mentioned (Bernard et al., 1991, stability and persistence of these factors during Clarke and Crame, 1997). the year are primary factors that determine the A fact that supports the idea of different zoo- ranges of distribution and life cydes of species. geographical subprovinces is that various species restricted to the eastern and western waters of the Aleutian Province (2) region were observed. The total estimated number of species, none of which endemic, is 37 (Fig. 6, In this province the most northwestern coast of Table 1, 3, 4). North America is included. It begins in Nunivak

25 90 80 20 70 (/) (/) Q) Q) 60 .0 15 .0 Q) Q) 50 Q. Q. (/) 10 (/) 40 z z 30 5 20 10 O O tUQ) tUQ) tUQ) tUQ) tUQ) tUQ) tU tU tU tU~Q) tUQ) tUQ) tUQ) tUQ) tUQ) tU tU :; :; :; :; :; :;J "O "O "O "O "O "O "O "O "O "O "O "O .0 .0 .0 .c .5 .0 .0 .0 .c Q) .;¡¡ E .J::>- Q) .~ .5 E .J::>- t) '5o tU '¡¡; c¡¡ o u t) '5o tU '¡¡; c¡¡ o u tU Q) Q. Ü t) (/) !l.. <: tU tU ~Q) Q. Ü t) (/) !l.. <: <:Q) OJ o tU <1: Q)<: OJ o tU <1: [D Qj Q)<: <1: c¡¡ aJ Qj Q)<: <1: c¡¡ !l.. Cf) .J:: !l.. Cf) .J:: U5 1- U5 1- groups groups

Figure 6. Distribution of Decapod Crustacean species in Figure 7. Distribution of Decapod Crustacean species in Artic province for each group. Aleutian proYince for each group. Figura 6. Distribución de las especies de crustáceos Figura 7. Distribución de las especies de crustáceos decápodos, por grupos, en la provincia Artica. decápodos, por grupos, en la provincia Aleutiana. BOSCHI: DISTRIBUTION OF DECAPOD CRUSTACEANS 19

Island (Lat. 60° N) on the Alaskan Peninsula, Californian Province (4) includes the Aleutian Islands and extends to Puget Sound (Lat. 47°50'N), Washington. It is a This province extends from Point Conception province of cold-temperate waters. The minimum to Bahía Magdalena (Lat. 24°40'N) in south Baja surface temperature reaches 5°C, sometimes O°C, California, Mexico. Surface temperature varia- and the maximum is around 14°C. tion is between 13° and 25° C and corresponds to The variations of the province limits agree warm-temperate waters. with changes in current systems that dominate the The Califomian Current transports cold waters area (Briggs, 1974, Brusca & Wallerstein, 1979, along the coasts of California to Point Conception Foster el al., 1991, Hall 1964, Valentine, 1966, where the temperature decreases markedly and Sverdrup el al., 1955). the current deftects offshore. The total number of species is estimated at 182, This determines a sharp temperature gradient 41 of which are endemic (Fig. 7, Table 1,3,4). with a rapid increase towards the south and changes in faunal composition that form a transi- Oregonian Province (3) tional zone. The total number of estimated species is 301, This province extends from Puget Sound, 48 of which are endemic (Fig. 9, Table 1, 3,4). Washington, to Point Conception, California (Lat. 34°35'N) and corresponds to cool-temperate waters with surface temperatures between 9° and The Eastern Central Pacific Ocean 16°C. The southern portion of the Oregonian Provinces and Subprovinces of Province is considered a transitional zone subtropical and tropical waters between this province and the north of the Californian Province (Foster el al., 1991). In the first part of this zone there is a region of The total number of species is estimated at subtropical and tropical waters that cover the 193,4 ofwhich are endemic (Fig. 8, Table 1,3,4). coasts of Baja California south of Bahía

70 60

en 50 .~ 40 ID 5} 30 z 20 10 ::::::::,:3,:, O ea ea ea ea ea ea ea ea ea ea ciI ciI ea ea ID ~ID ~~ea ID ID ID ID ::; ::; :J ID ID ID ID ID ID :J :J ::; '"O '"O '"O '"O '"O '"O '"O '"O '"O '"O '"O '"O '0 '0 .~ '0 .5 E .r:>- .~ .5 E >- ID '6 ea .¡¡;'c o '0 '0 '6 '0 'c o .r:() tí o e « ro ID (¡¡C> e « (ij ro o... ID

Figure 8. Distribution of Decapod Crustacean species in Figure 9. Distribution of Decapod Crustacean species in Oregonian province for each group. Californian province for each group. Figura 8. Distribución de las especies de crustáceos Figura 9. Distribución de las especies de crustáceos decápodos, por grupos, en la provincia Oregoniana. decápodos, por grupos, en la provincia Californiana.

~ .r: (¡j U .- t:: Q .¡¡; § o .g '"O m .~ -;;:¡ n ~' ~ en ,~ 1ii - " 20 REY. INYEST.DES. PESQ. W 13: 7-136 (2000)

Magdalena to Gulf of Guayaquil, Ecuador. and tides, with very pronounced seasonal The provinces and subprovinces included are: changes. Naturally, in deep areas, changes in tem- Panamic, Cortés and Galápagos. perature are not so pronounced. Together they determine the warm region of In the north of the Gulf, at depths of 30 - 40 m, the Eastern Pacific with a high degree of temperatures are around 15.0°C in January and endemism in various groups of marine organisms 27.0°C in August. Between 60 and 75 m, temper- (Brusca and Wallerstein, 1979). atures are 14.SOCin January and 20. 8°C in August (Hendrickx, 1992). Cortés Subprovince (5) With regard to diversity, distribution and abundance of species of demersal and benthic The Gulf of California is situated east of the decapods, temperatures at the bottom produce a Central Eastern Pacific Ocean, between latitudes significant effect on these communities. However, 31°40' and 20°4 7'N and between the California the total life cycle has to be considered and the Peninsula to the west and the American continent important larval phase included. to the east. It is a semienclosed, narrow body of Generally, larvae reach the epipelagic zone in water with a north-south extension. their daily vertical movements and, for this rea- In the northern part the waters are as deep as son, the surface and subsurface temperatures may in the central and southern parts where the system be limiting distribution factors. is open and in direct contact with the Pacific As previously mentioned, the oceanographic Ocean. The limit between both may be consi- conditions in the northern part of the Gulf and the dered as an imaginary line between Cape San origin of its biota is very distinct from those in the Lucas and Cape Corrientes. south of the Gulf and adjacent Pacific Ocean. The deepest depths oscillate from about 200 m These characteristics determine the composi- in the north to some 1800 m in the center, fre- tion of the communities of marine organisms that quently reaching more than 3000 m in the south establish in these zones. Hendrickx (1992) (Roden and Groves, 1959, Parker, 1963, observed an important decrease in species abun- Hendrickx, 1992, 1993, 1995, 1996, Villalobos el dance in the north of the Gulf, especially in the al., 1992). number of tropical species of decapods. The northern part of the Gulf is the most iso- However, Hendrickx (1992) did not consider it lated one in the East Pacific and under the great- sufficient enough to justify an independent zoo- est influence of the continental climate. geographic unity. Following Briggs (1974), he The warm and dry winds of the north generate defined the area as the Mexican Province, from an intense surface evaporation and high tempera- Bahía Magdalena to Cape San Lucas, in the west- tures that create strong stratification in surnmer ern coast of Baja California, that includes the and produce a considerable alteration in the water en tire coast of Mexico to the Gulf of that, in Puerto Peñasco (Lat. 31°21 'N), reaches a Tehuantepec. temperature of more than 31°C and a salinity of There is a large area from 28° 30'N stemming around 36.0 ppt. Winter temperatures in this part south of Tiburón Island and going to the northern of the Gulf may decrease to 12-13°C (Brusca, limit of the Gulf. 1980). As proposed by Briggs (1974), this area has a The wind regime and topographic characteris- transitional fauna with subtropical predominance tics of the Gulf of California have a significant and some degree of endemism that may justify influence on the circulation of water masses, for- the category of a subprovince denominated mation of upwellings, temperature distribution "Cortés" . BOSCHl: DISTRIBUTION OF DECAPOD CRUSTACEANS 21

Garth (1960), when working on the distribu- coral s of the Mexican Pacific, determined the tion of Brachyura in the Gulf of California, point- existence of two independent groups. ed out that north of Cedros Island the environ- In one of them, in the central and northern part ment is temperate with high temperatures in sum- of the Gulf, there are only two species of Porites mero In contrast, although it is not possible to adapted to the frequent seasonal environmental establish the northern limit of the Panamic fauna, changes in temperature, salinity and light which the southern part contiguous to the Pacific Ocean coincide with the formerly mentioned is tropical. Soule (1960), when studying affinities Subprovince of Cortés. The remaining parts of of the littoral marine Bryozoa, established zones the region include other zoogeographical for species in the Gulf of California which includ- provmces. ed subtropical species in the north followed by a Bernard et al. (1991), who studied the zoo- transition zone from around 28° N. geography of bivalve mollusks of the East Pacific Moreover, a tropical zone in the south extend- Ocean, established at least nine species belonging ed to the west coast of the Californian Peninsula to the north of the Californian Gulf. These authors up to approximately Bahía Magdalena. considered their possible connection with the Correa Sandoval and Carvacho Bravo (1992), ocean during the Pleistocene, a fact that could based on studies on the brachyuran crabs, divided explain isolation in the n0l1hern part of the Gulf the Gulf of California into two parts: high and of California. low Gulf. This is due to the biogeographical bar- Accordingly, the northern part of the Gulf rier of Angel de la Guarda and Tiburón Islands from the Tiburón and Angel de La Guarda Islands that represent a 69% filter for species from the to the north makes up the Cortés Subprovince. south to the north with seven endemic pin- The number of decapod species is 265, 25 of notherids in the high Gulf. which are endemic (Fig. 10, Table 1, 3,4). On the other hand, it is interesting to point out that Reyes-Bonilla and López-Pérez (1998), Panamic Province (6) based on studies on the distribution of calcareous The Panamic Province extends from Bahía Magdalena, Baja California, Mexico, to the Gulf 160 of Guayaquil-Bahía Sechura (Lat. 4°38'S). 140 It includes the entire Mexican littoral, the west 120 ~ 100 coast of Central America, Colombia and Ecuador 'g 80 in South America and the oceanic islands of the 5J- 60 Z region (Revillagigedo, Clipperton, Coco, 40 Malpelo, Gorgona, etc). ;;,;,;,;,;;;;;; 2~ These islands are connected to the continent Q)C\J C\JQ) Q)C\J Q)C\J Q)C\J Q)C\J C\J C\J C\J ::; ::; ::; by a permanent system of surface currents 1:J 1:J :g 1:J 1:J 1:J ,s 'oQ) 'o 1:J .~ 'o 'e: E ..c>- en e C\J '¡¡; (ij O ü (Lemaitre and Alvarez León, 1992). C\J Q) Q. Ü en en el. e C\J Q)e OJ o C\J « Q)e « (ij m In the Panamic Province the largest diversity el. ID ..c Cf' U5 1- of species in the entire East Pacific Ocean occurs. groups Temperatures oscillate between 19° and 28°C although, in winter, they may decrease to around Figure 10. Distributian af Decapad Crustacean species in Cortés subprovince far each group. 14°C in the northern part and in summer reach Figura 10. Distribución de las especies de crustáceos 30°- 32°C in some extreme areas. decápodos, por grupos, en la subprovincia Cortés. The estimated species are 825, 315 of which 22 REV. INVEST.DES. PESQ. N" 13: 7-136 (2000) are endemic (Fig. 11, Table 1, 3, 4). The Southeastern Pacific Ocean Province 01 temperate waters Galápagos Subprovince (7) Perú-Chilean Province (8) The Galápagos Subprovince inc1udes the islands situated at about 1,000 km from the coasts The Perú-Chilean Province extends from the of Ecuador. They are also known as Archipiélago Gulf of Guayaquil-Bahía Sechura (Lat. 4°38'S) to de Colón. This subprovince is situated between the north of Chiloé Island (Lat. 41°48'S). the following coordinates: Lat. 0040'N and 1°30'S- The coastal topography of this extended Long. 89°20' W and 91°50' W. region is regular. It is totally exposed to winds Due to the large number of decapod species and waves with narrow and open beaches and a and the endernism of some of them it is consid- few protected bays. At the Valparaiso latitude the ered as a province (Garth, 1946). Briggs (1974), continental shelf is very narrow, generally with- taking into consideration its high level of out sediments and with a substrate of hard rocks endemism, considered it as a different province. derived from the ridges of coastal mountains Kim and Abele (1988) studied the snapping (Mordojovich, 1983). Normally, this continental shrimp of the genus Alpheus of the Eastem shelf has no more than 5-10 km. in width and only Pacifico They considered these islands as a subre- in some parts reaches 30 km (Brattstróm and gion but, on the other hand, suggested that the Johanssen, 1983). area under study, between 34°N and 2°S, which The Pacific coasts are under the influence of includes three subregions, could be considered as the Peruvian Current (also called Humboldt) that a provmce. transports cold water rich in nutrients from the The total number of estimated species of Chiloé Island norther1y to Perú with temperatures decapods is 253, 41 of which are endernic (Fig. ranging between 11°C and 19°C. These oceano- 12, Table 1,3,4). graphic conditions of the Chilean and Peruvian

400 140 350 120 300 100 ~ 250 .~ 200 .~(} 80 5} 150 ~ 60 z 100 ~ 40 50 ,.,. 20 ., o """""", O Q)ti! Q)ti! Q)ti! Q)ti! Q)ti! Q)ti! ti! Q)ti! ti! Q)ti! Q)ti! Q)ti! ti! ti! ti! ti! ~ ~ Q) Q) :J :5 :J :5 :5 :5 ""O ""O ""O ""O ""O :Q ""O ""O ""O ""O ""O ""O .~ .S >- .S >- '0Q) '0 'ü e E .r:(} '0Q) '0 .~ 'ü 'c E .r:(} (jj '5 ti! .¡¡; '¡¡:j o (jj '5 ti! .¡¡; '¡¡:j o (jj o o ti! Q) Q. O (f) e ti! ti! Q) Q. O (jj (f) e ti! c.. c.. Q)e OJ o ti! « Q)e OJ o ti! « e « '¡¡:j iD Q)e « '¡¡:j iD Q) c.. (j)ID c.. (j)ID .r: U5 .r: iJ5 1- 1- groups groups

Figure 11. Distribution of Decapod Crustacean species in Figure 12. Distribution of Decapod Crustacean species in Panamic province for each group. Galápagos subprovince for each group. Figura 11. Distribución de las especies de crustáceos Figura 12. Distribución de las especies de crustáceos decápodos, por grupos, en la provincia Panameña. decápodos, por grupos, en la subprovincia Galápagos. BOSCHI: DISTRIBUTION OF DECAPOD CRUSTACEANS 23 coasts are disturbed at irregular intervals by a The Southeastern Pacific Ocean and strong penetration of subsurface warm waters of Southwestern Atlantic Ocean the "El Niño" Current which produce important Province of cold-temperate waters changes in the coastal fauna. These alterations in the temperature have been recorded up to latitude Magellanic Province (9) 35°S (Santelices, 1991). This province also includes warm-temperate The Magellanic Province extends from the water s with an area of transition in the south, from north of Chiloé Island, on the Eastem Pacific Valparaiso (33°S) to the north of the Chiloé Island Ocean, follows the Magellanic and Tierra del (42°S) Brattstrom and Johanssen (1983). Fuego region through the Cabo de Hornos, reach- In this province 212 species of decapod crus- es the coastal area of the Southwestem Atlantic taceans are found, 77 of which are endemic Ocean off the Patagonian region, Argentina, and (Fig.13, Table 1,3,4). includes the Malvinas Islands to the Valdés It is evident that the largest number of species Peninsula. The Province deflects from the conti- in the southeastem Pacific is observed northerly nent at Lat. 43°- 44°S (Rawson, Chubut) going of Chiloé Island. In the Juan Femández Islands, north, reaching latitude 35°S (see Fig.1) at a dis- situated about 600 km from the coast, the decapod tance of 100-150 km from the coast, with a depth fauna is mainly related to species of the South of 60-200 m and limits difficult to establish. The Pacifico temperatures in winter range from 4°C in the The average surface temperature in the region south to 14-16°C in the north. At that latitude, is 18°C (Retamal, pers. comm.). Briggs (1974) between the Patagonian region and Buenos Aires, considered that this could be an independent zoo- it reaches the Argentinian Province (see Fig.1). geographic area. Many authors have treated the topic relative to the biogeographical region of the south of South America. There is agreement on the faunal and floristic relations between the Southeastem Pacific Ocean and the Southwestem Atlantic Ocean. Ekman (1953), who studied the 120 Antiboreal region, considers that the fauna of the 100 south of Chile goes from the Chiloé Island C/) 80 Q) through Tierra del Fuego in the Patagonian and .~ 60 Malvinas waters (Argentina). He argued that, due C/)c. z 40 to the scant information available on the South 20 ,m;¡¡. Atlantic region, it was not known how far north O this fauna extends. Q)tU Q)tU Q)tU Q)tU Q)tU Q)tU tU tU tU :; :; :; :Q :Q Carcelles and Williamson (1951) and Balech .5 E >- "'0Q) "'0 ".~ "'0 .¡¡;e (ij .r: en o o u tU Q) "c. ü .sC/) C/) a.. e (1954) established the limits of the Magellanic Q)e CJ o tU - '5 (¡j .s:::() ID tí o

Figure 14. Distribution of Decapod Crustacean species in (') In this regard, the existence of mangroves, which re{J- Magellanic proYince for each group. resent a very significative community in the area could Figura 14. Distribución de las especies de crustáceos be taken as a limit since they do not reach latitude of decápodos, por grupos, en la provincia Magallánica. 28°30'S (Cintrón and Schaeffer-Novelli, 1983). BOSCHI: DISTRIBUTION OF DECAPOD CRUSTACEANS 25

It is important to point out that this province of transition reaching, inclusively, the littoral of includes geographical features like the Rio de la Buenos Aires. Plata, the Patos Lagoon in Brazil and other water This province can be characterized as a warm- bodies. The estuaries influence the primary and temperate province with 330 species, 42 of which secondary production in the area and, con se- are endemic (Fig. 15, Table 1, 3,4). quently, the distribution and abundance of d~ca- pod species. Brazilian Province (11) Moreover, in the whole region there is a per- manent interaction off the coasts between the The BraziHan Province occupies a large exten- Malvinas Current flowing on the slope from the sion of the tropical and subtropical Httoral of the south with cold subantarctic waters rich in nutri- north and northwestem of Brazil, French Guiana, ents and the Brazilian Current, out o-!"shelf, from Suriname, Guyana and a small part of Venezuela, the north, with temperatures higher than 20°C and from Cabo Frío to the mouth of the Orinoco Ri ver salinity over 36.0 ppt (Costa and Costa (Lat. 8°56'N). Femández, 1993). In this extended area temperatures vary from Disregarding the delimitation of the sub- 22°C in the south to 30°C in the north. There are provinces, there is affinity among various deca- different opinions as regards the area of this pod crustacean species of the coastal littoral of extensive province. Balech (1954) named it the Buenos Aires and the south of Brazil at latitude Antillian Province with the limits previously indi- 23°S. cated and divided into three districts. Coelho Of the total of 93 decapod species indicated (1967/69), Coelho and Ramos (1972), and Coelho for the littoral of Buenos Aires (Boschi et al., and dos Santos (1980), who studied the distribu- 1992), 36 reach latitude 23°S. tion of decapod crustaceans of the Brazilian shelf As occurs with some caridean shrimp and proposed a subdivision of the province into sub- brachyuran crabs (Christoffersen, 1982, Melo, provinces. However, they accepted the southem 1996), some tropical species enter this large zone limit of 23°S. Forest and Saint-Laurent (1967), referring to the pagurids of the South Atlantic, pointed out some zoogeographic regions. 200 Briggs (1974) estabHshed the limits of the Brazilian Province from the mouth of the Orinoco Q)en 150 .~ River to Cabo Frío, accepted in this study. 100 enel. The catalogue of all species of marine Z 50 decapods of Brazil appeared in Young (1998). The estimated number of decapod crustacean species is 572, 64 of which are endemic (Fig. 16, Q)- '0Q) '0 '5 '0 .r:: ID o

300 500 450 250 400 350 en Q) 200 ~ 300 .~ 150 '0 250 el. en~ 200 ~ 100 z 150 50 100 50 @)':¡, O r:~:~:~:~:~. O '.'.'.'"'' (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j (¡j Q) Q) Q) Q) Q) Q) Q) Q) Q) Q) Q) Q) -o -O -O -O -O ~ ~ ~ -o -O -o -o ~~-O -O ~ :2 .S >- '0Q) .~ '0(¡j e .S E .c>- '0Q) '0 .~ '0(¡j E .c() '0 '6 .¡¡; -¡¡¡ O () '6O .¡¡;'c -¡¡¡ O (¡j U;Q) O e (¡j U;Q) Ü e (¡j el. Ü U; (¡jen a.. ~ el. U; (¡jen a.. Q)e O> O « Q)e O> O « « Q; Q)e « -¡¡¡ te Q; Q)e -¡¡¡ c!5 a.. a.. C/) .c C/) .c Cñ 1- Cñ 1- groups groups

Figure 16. Distribution of Decapod Crustacean species in Figure 17. Distribution ofDecapod Crustacean species in Brazilian province for each group. Caribbean province for each group. Figura 16. Distribución de las especies de crustáceos Figura 17. Distribución de las especies de crustáceos decápodos, por grupos, en la provincia Brasileña. decápodos, por grupos, en la provincia Caribeña. BOSCHI: DISTRIBUTION OF DECAPOD CRUSTACEANS 27

Texan Province (13) The 20°C difference in the winter temperature between the northern and southern areas is a fac- The Texan Province corresponds to a large tor that really limits the distribution of stenotype area of the Gulf of Mexico which includes part of speCles. the northwestern, northern and southeastern Gulf The variety of hard and soft substrates in the of Mexico from Cabo Rojo, south of Tampico, Gulf of Mexico, the outer continental shelf in Mexico, to Cabo Romano, south of the Florida Mississippi, Alabama and eastern Louisiana, Peninsula, (Lat. 25°54'N), USA. It also includes exerts a high influence on the diversity of inverte- part of the littoral of the States of Veracruz, brate assemblages (Gittings et al., 1992). Tamaulipas (Mexico) and those of Texas, Due to the number of endemic species, this Louisiana, Mississippi, Alabama and part of area of the Gulf of Mexico may be considered as northern Florida, USA (Briggs, 1974). an independent province with warm-temperate The climate and oceanographic conditions of characteristics; nevertheless, during part of the this are a of the Gulf of Mexico are quite different year (summer) subtropical conditions are from the tropical part. Winds from the north are observed. As proposed by Briggs (1974), this area generated by polar masses of air which, in autumn can be called the Texan Province. and winter, (November through March) produce The number of species is about 422, 20 of low surface and shallow water temperatures. which are endemic (Fig. 18, Table 1, 3, 4). It is The lowest water surface temperatures that interesting to mention the study by Williams and drop as low as 11°C are found in the north of the Felder (1986) on species of Mennippe in the Gulf Gulf in winter. To the south, in the same season, of Mexico. These authors considered a new they are around 21°C. species, Mennippe adina, distributed in the Texan In summer the highest surface temperature Province, different from Mennippe mercenaria of that reaches 30°C is very homogeneous in the Cape Lookout, North Carolina, the Florida whole Gulf (Leipper, 1954). Peninsula and the Caribbean.

250 The Northwestern Atlantic Ocean Provinces of temperate and 200 cold-temperate waters

.~() 150 Q)

fE- 100 This large region, which includes the zoogeo- z 50 graphic provinces of the coastal waters of the northwestern Atlantic Ocean, extends from south

(tj (tj (tj (tj (tj (tj (tj (tj Q) Q) Q) Q) Q) of the Labrador Peninsula (Island of ~Q) :; :; ~'0""O ""O ""O ""O ""O ""O .S: Newfoundland) in the Strait of Belle Isle Q) '0 .~ '0(tj 'c E .I::()>- 'Oo .¡¡; '¡¡j o (tj 1ñQ) a. Ü c: 1ñ (tjen a. (51°37'N) to the Florida Peninsula (25°1O'N). Q)~c: O> o « [D Q)c: « '¡¡j a. (j')ID .I:: The region includes a considerable variation U5 1- groups of climates, from polar to tropical, with water temperatures ranging between O°C and 30°C. Figure 18. Distribution of Decapod Crustacean species in The region is influenced by two important cur- Texan province for each group. rent systems: The Gulf Stream, with warm waters, Figura 18. Distribución de las especies de crustáceos flowing northerly and the Labrador Current, with decápodos, por grupos, en la provincia Texana. cold waters, flowing southerly.

u. en a. ~.Jl'.J 11~) (O me 1'lOnaa renmSUla ~.LY lUI~). c:'" ~ o ~ m ~ -~ 28 REY. INYEST. DES. PEsQ. N° 13: 7-136 (2000)

The provinces included from south to north a zone of zoogeographic changes greater than in are the following: the Carolinian, the Virginian Cape Hatteras. and the Boreal (Hedgpeth, 1957, Briggs, 1974, The number of species is estimated at 386, 5 of Williams, 1984). which are endemic (Fig. 19, Table 1, 3,4).

Carolinian Provine e (14) Virginian Provinee (15)

The Carolinian Province begins at Cape The Virginian Province extends from Cape Canaveral, Florida and extends up to Cape Hatteras, North Carolina, to Cape Cod (Lat. Hatteras, in North Carolina (Lat. 35°17'N), on the 41°48'N), Massachusetts. These capes are the eastern coast of USA. It is an area that can be con- most significant points of deflection of the Gulf sidered as warm-temperate (Briggs, 1974). Stream and the Labrador Current. The temperature ranges between 20° and 25°C In this area, changes in the characteristics of in winter and between 28° and 30°C in summer water temperature and in the patterns of circula- (Dawes et al., 1991). Ray et al. (1997) drew tion, very important for the ranges of distribu- attention to the importance of the estuaries in the tion of the organisms (Ray et al., 1997) take biodiversity of species, especially in the area of place. Chesapeake Bay. They also mentioned the break The range of surface water temperatures is in faunal associations of invertebrates and fishes high in the region - between 20° and 25°C. at the latitudes of Cape Cod and Cape Hatteras Temperatures vary between -1° and 23° C in which confirms the value of these areas when Massachusetts and between 3° and 30° C in North establishing limits of the Northwestern American Carolina. Provinces. Salinities in open areas vary between 33.0 and On the other hand, Herbst et al. (1979) point- 37.0 ppt. (Orth et al., 1991). Ray et al. (1997) ed out that Cape Lookout, North Carolina, marks agreed with the nomenclature of traditional

200 180 160 Cf) 140 .~ 120 á5 100 ~ 80 z 60 40 2~ tW{ .

Q)tU Q)tU Q)tU Q)tU Q)tU Q)tU tU ~ ~ ~Q)tU Q)tU Q)tU Q)tU Q)tU Q)tU tU tU :; ::> ::> :; :; ::> '"O '"O '"O '"O '"O '"O .!;; '"O '"O '"O '"O '"O '"O .!;; >- .~ '(3 E .r:::.>- Q) .~ '(3 'c (¡j E .r:::. '0Q) '0 '5 .¡¡;'c (¡j O 'o '0 '5 tU .¡¡; o 1i) O tU o 1i)Q) o o tU Q) el. Ü 1i) Cf) a.. e tU tU el. Ü 1i) Cf) a.. e tU Q)e O> O tU « Q)e O> O « tU « (¡¡ Q)e « (¡j as (¡¡ Q)e (¡j as a.. a.. C/) .r:::. C/) .r:::. Cñ 1- Cñ 1- groups groups

Figure 19. Distribution of Decapod Crustacean species in Figure 20. Distribution of Decapod Crustacean species in Carolinian province for each group. Virginian proYince for each group. Figura 19. Distribución de las especies de crustáceos Figura 20. Distribución de las especies de crustáceos decápodos, por grupos, en la provincia Caroliniana. decápodos, por grupos, en la provincia Virginiana. BOSCHI: DISTRlBUTION OF DECAPOD CRUSTACEANS 29 provinces on the eastern coast of USA. They also attached importance to the estuaries CONCLUSIONS (as in the Carolinian Province) in the composition of associations of organisms and in the species biodiversity of the area and emphasized the For the first time, an inventory of the known importance of the oyster reefs in the formation of species of Decapod Crustaceans living in conti- biotopes in the coastal fauna. nental shelf waters of the Americas (North, The number of decapod species in the Central and South) was completed. province is estimated at 158, 1 of which is ende- The distribution of species in this extensive mic (Fig. 20, Table 1, 3, 4). region, from the Arctic in the north to Cabo de Hornos in the south is analyzed in relation to Boreal Province (16) environmental conditions. In this region, the structure and extension of The Boreal Province extends from Cape Cod the continental shelves, from the coast to the in the eastern coast of U.S.A. to Newfoundland slope, is very variable. The extension exceeds 850 Island and the Strait of Belle Isle, in Canada (Lat. km in the Argentinian Patagonia while in the cen- 51°37'N). The coasts are under the influence of tral and northern coast of Chile, n011h of Brazil, the cold Labrador Current which keeps tempera- east coast of Florida, etc., is very reduced or tures quite low. nonexistent. The variable dimensions of the The limits of surface temperatures are esta- shelves covered by epicontinental seas determine blished by the isotherms of 10°C in winter and 15°C different environmental conditions. in summer in the south and of O°C in the north. The characteristics of the bottom, marine cur- The number of species of decapod crustaceans rents with variable temperatures and salinities, is relatively low; 77 recorded to date, 4 of which transparency of waters, primary and secondary are endemic (Fig. 21, Table 1,3,4). production, nutrients, etc. are factor s that deter- mine the characteristics of the flora and fauna pre- sent in each region. In the case of decapod crustaceans, different characteristics were observed between species living in the sea with a wide continental shelf and a smooth depth gradient and those living at greater depths, close to the coast and without shel f. The limits of the isotherms were considered as

(\1 (\1 ~Q) Q)(\1 Q)(\1 Q) Q)(\1 Q)(\1 (\1 (\1 a very important factor used in the definition of :; :; :J 'C 'C 'C 'C 'C 'C .~ E >- margins between the zoogeographic provinces. "oQ) "o '5 "~ "ü(\1 'c -¡¡¡ .r::() tí o "¡¡; o (\1 Q) Q. Ü tí UJ c.. e (\1 The number of described species in the entire eQ) OJ O (\1

degree with those established for other groups of B. Holthuis, A. Kameya Kameya, B. Kensley, R. . . manne orgamsms. Lalana, R. Lemaitre, R. B. Manning, 1. W. Maliin, The Iargest discrepancy is observed in the G. A. S. Melo, J. L. Orensanz, M. Ortiz, 1. Pérez Subprovince of Cortés in the Gulf of Califomia Farfante, M. Retamal, G. Rodríguez, S. de A. and in the Texan Province in the Gulf of Mexico. Rodrigues, L. Soto, M. Tavares, B. Werding, M. The provinces with the largest number of K. Wicksten and A. B. Williams. I also thank J. L. species are the Caribbean with 1058 species and Swidzinsky for computer work. In particular, I the Panamic with 825 species. wish to acknowledge the help of my wife Elba These provinces show a high level of and I thank an anonymous reviewer for comments endemism, the former with 338 species or 32% and useful suggestions. and the Iatter with 315 species or 39%.

NOTE ADDED IN PROOF ACKNOWLEDGEMENTS

Frorn the time this paper was subrnitted changes The author is very grateful to many colleagues were introduced in the narne of sorne species and fa- who provided advice and literature: L. G. Abele, rnilies, new ones were described and sorne others, not R. Bauer, E. Campos, N. Campos, P. A. Coelho, registered in the originallist, incorporated. Table 5, 6. S. Gomez-Aguirre, D. Guinot, M. Hendrickx, L.

Table 5. Species not included in appendix. Tabla 5. Especies /70 incluidas en el apéndice.

Species Family Proy. References

Plesionika macropoda Chace, 1939 Pandalidae 12-13 Memor. SocoCubana Hist. Nat 13(1), 1939 Alpheus polysticlllS Knowlton and Keller, 1985 Alpheidae 12 Rey. Ayicennia O, 1993 Cuba Paguristes depressus Stimpson, 1858 Diogenidae 12 Rey. Inyest. Marina 5(1), Venezuela Paguristes maclauglinae M.Iglesias and Gómez, 1989 Diogenidae 12 Poeyana, Cuba N°379 Porcellana lillyae Lemaitre and Campos, 2000 Porcellanidae 12 JCB 20(2), 2000 Petrolisthes cessacii (A. Milne Edwards, 1878) Porcellanidae 11-12 Anal. Inyest. Marinas 9, 1977 Venezuela PontoniamanningiFransen,2000 Palaemonidae12-13-14JCB SpeciaI20(2),2000 Lepidophthalmus manningi Felder and Staton, 2000 Callianassidae 12 JCB Special 20(2), 2000 Naushonia manningi Alyarez, Villalobos and Lliffe, 2000 Laomediidae 12 JCB Special 20(2), 2000 Calliasmata nohochi Escobar, Camacho and Alcocer, 1997 Hippolytidae 12 JCB 17(4), 1997 *Notiax santarita Thatje, Romera and Tapella Callianassidae 9 1Jornadas Arg.Carcinología, 1999 BA, Arg. *Upogebia australis Thatje, Romero and Tapella Upogebiidae 9 1Jornadas Arg.Carcinología, 1999 BA, Arg. Cyclodorippe longifrons Campos Jr. and Melo, 1999 Cyclodorippidae 10 Atlántida 21(1),1999 Río Grande Clythrocerus bidentatus Campos Jr. and Melo, 1999 Cyclodorippidae 10 Atlántida 21(1), 1999 Río Grande **Bermudacaris harti Anker and IIiffe, 2000 Alpheidae 12 Prac. B. SocoWashington 113(3), 2000 Fabia insularis Melo, 1971 Pinnotheidae 10 Rey. Brasilera Zool. 13(1), 1996 Planes marinus Rathbun, 1914 Grapsidae 10 JCB 19(1), 1999 Calliasmata rimolii Chace, 1975 Hippolytidae 12 Prac. B. SocoWashington 88,1975

* Beagle channel (Southern South America). ** Bermuda Islands. BOSCHI: DrSTRIBUTION OF DECAPOD CRUSTACEANS 31

Table 6. Last changes in the families and species names. Tabla 6. Ultimos cambios en la denominación de especies y familias.

Previous classification

Species Family

1 Benthochascon schmitii (Rathbun, 1931) Portunidae 2 Iliacantha intermedia Miers, 1886 Leucosiidae 3 Anacalliax agassizi (Biffar, 1971) Callianassidae 4 Anacalliax argentinensis (Biffar, 1971) Callianassidae 5 Dawsonius latispina (Dawson, 1967) Callianassidae 6 Gourretia biffari Blanco and Liñero, 1994 Callianassidae 7 Gourretia laresi Blanco and Liñero, 1994 Callianassidae 8 Corallichirus longiventris (A. M. Edwards, 1870) Callianassidae 9 Porcellana stimpsoni A. Milne Edwards, 1880 Porcellanidae

Updated classification

Species Family Reference

1 Raymanninus schmitt (Rathbun, 1931) Portunidae JCB Special 20/2/2000 2 Acanthilia intermedia (Miers, 1886) Leucosiidae PBSW 113(2), 2000 3 Anacalliax agassizi (Biffar, 1971) Ctenochelidae JCB Special 20/2/2000 4 Anacalliax argentinensis (Biffar, 1971) Ctenochelidae JCB Special 20/2/2000 5 Dawsonius latispina (Dawson, 1967) Ctenochelidae JCB Special 20/2/2000 6 Gourretia biffari Blanco and Liñero, 1994 Ctenochelidae JCB Special 20/2/2000 7 Gourretia laresi Blanco and Liñero, 1994 Ctenochelidae JCB Special 20/2/2000 8 Corallianassa longiventris (A. M. Edwards, 1870) Callianassidae JCB Special 20/2/2000 9 Porcellana sayana (Leach, 1820) Porcellanidae JCB 20/2/2000 32 REV. INVEST.DES. PESQ. W 13: 7-136 (2000)

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'-'AlYIYU;:',D. lYYV. ~atypTraeOl1leres, a new genus nus tor the Central Amencan crab Pinnixa cos- 36 REY. INYEST.DES. PESQ. N° 13: 7-136 (2000)

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séum National D'Histoire Naturelle 4° Série 11 In: Guía FAO para la identificación de especies (1): 193-202. para los fines de la pesca. Pacífico centro-Orien- HENDRICKX,M. E. 1990. New geographic distribu- tal. 1. Plantas e invertebrados 646pp.W. Fischer, tions and bathymetric records of Processidae F. Krupp, W. Schneider, C. Sommer, K.E. Car- (Caridea) and Penaeidae (Penaeoidea) in the penter y V. H. Niem (Eds.) FAO Roma, Italia. Gulf of California, Mexico. Investigaciones HENDRICKX,M. E. 1995. Anomuros. 539-564 pp. Marinas CICIMAR 5 (1): 93-95. In: Guía FAO para la identificación de especies HENDRICKX,M. E. 1992. Distribution and zoogeo- para los fines de la pesca. Pacífico centro-Orien- graphic affinities of decapod crustaceans of tal. 1.Plantas e invertebrados 646 pp. W. Fischer, the Gulf of California, México. Proceedings of F. Krupp, W. Schneider, C. Sornmer, K.E. Car- the San Diego Society Natural History 20: 1-12. penter y V. H. Niem (Eds.) FAO Roma, Italia. HENDRICKX,M. E. 1993. Distribution of Petrolis- HENDRICKX,M. E. 1995. Cangrejos. 565-636 pp. thes lewisi (Crustacea: Porcellanidae) in the In: Guía FAO para la identificación de especies Eastem Tropical Pacifico Revista de Biología para los fines de la pesca. Pacífico centro-Orien- Tropical 41 (2): 287-290. tal. 1.Plantas e invertebrados 646 pp. W. Fischer, HENDRICKX,M. E. 1993. Crustáceos decápodos F. Krupp, W. Schneider, C. Sommer, K.E. Car- del Pacífico Mexicano. 271-318 pp. In: Biodi- penter y V. H. Niem (Eds.) FAO Roma, Italia. versidad marina y costera de México.S. 1. Sa- HENDRICKX,M. E. 1995. Checklist of brachyuran lazar- Vallejo y N. E. González (Eds.). Comi- crabs (Crustacea: Decapoda) from the eastern sión Nacional de Biodiversidad y CIQRO, Mé- tropical Pacifico Bulletin de L'lnstitut Royal XICO. des Sciences Naturelles de Belgique, Biologie HENDRICKX,M. E. 1995. Checklist of lobster-like 65: 125-150. decapod crustaceans (Crustacea: Decapoda: HENDRICKX,M. E. 1995. Restitution de Pseudo- Thalassinidea, Astacidea and Palinuridea) from rhombila xanthiformis Garth, 1940, pour Na- the eastern tropical Pacifico Anales del Institu- noplax garthi Guinot, 1969 (Decapoda, Gone- to de Biología. Universidad Nacional Autóno- placidae). Crustaceana 68 (1): 12-20. ma de México, Series Zoológicas 66 (2): 151- HENDRICKX,M. E. 1996. Habitats and biodiversi- 163. ty of decapod crustaceans in the SE Gulf of HENDRICKX,M. E. 1995. Introducción. 1-7 pp. In: California, México. Revista de Biología Tropi- Guía FAO para la identificación de especies cal 44 (2): 603-617. para los fines de la pesca. Pacífico Centro O- HENDRICKX,M. E. 1996. Los camarones Penaeoi- riental. I . Plantas e invertebrados.646 pp. W. dea bentónicos (Crustacea: Decapoda: Dendro- Fisher, F. Krupp, W. Schneider, C. Sommer. branchiata) del Pacífico Mexicano. CONABIO/ K.E. Carpenter y V. H. Niem (Eds.) FAO Ro- Instituto de Ciencias del Mar y Limnología, ma, Italia. Universidad Nacional Autónoma de México HENDRICKX,M. E. 1995. Langostas (Langostas 1-147,21am. espinosas bogavantes y cigarras y zapateras, HENDRICKX,M. E. 1997. Los cangrejos braquiuros langostas de lodo, etc.). 383-415 pp. In: Guía (Crustacea: Brachyura: Dromiidae hasta Leu- FAO para la identificación de especies para los cosiidae). CONABIO/ Instituto de Ciencias fines de la pesca. Pacífico centro-Oriental. 1. del Mar y Limnología, Universidad Nacional Plantas e invertebrados 646 pp. W. Fischer, F. Autónoma de México 1-178. Krupp, W. Schneider, C. Sommer, K.E. Carpen- HENDRICKX,M. E. 1997. Los cangrejos braquiuros ter y V. H. Niem (Eds.) FAO Roma, Italia. (Crustacea:Majoidea y Parthenopoidea). CONABIO HENDRICKX,M. E. 1995. Camarones. 417-537 pp. /Instituto de Ciencias del Mar yLimnología,

y C1>plllU1><:11> UUb<:1 V<111le; 1> ~lb<111 <11>Y L<1p<1l~"''', .1..1.Ll'1jl.Il'\ l'---'.1~, .1,..L..L.J. .L/./ l. .L...I'--'~ """,..U.J.6.1."""'J'-'U """"."-"1"-..I.",,",o'-JLJ 46 REY. INYEST.DES. PESQ. N° 13: 7-136 (2000)

Universidad Nacional Autónoma de México 1- HENDRICKX,M. E. & A. W. HARVEY.1999. Check- 274, 13 lám. list of anomuran crabs (Crustacea: Decapoda) HENDRICKX,M. E. 1998. A new genus and species from the eastern tropical Pacifico Belg. J. Zool. of "goneplacid-like" brachyuran crab (Crusta- 129 (2): 363-389. cea: Decapoda) from the Gulf of California, HEREU,C. M. 1999. Aspectos biológicos e ecoló- Mexico, and a proposal for the use of the fami- gicos durante o período pelagial de Libinia spi- ly Pseudorhombilidae Alcock, 1900. Procee- nasa (Milne Edwards 1834) (Brachyura: Ma- dings of the Biological Society of Washington jidae) no litoral do Rio Grande do Sul, Brasil. 111 (3): 634-644. Disertac;ao de Mestrado. Fundac;ao Universi- HENDRICKX,M. E. & F. D. ESTRADANAVARRETE. dade do Rio Grande, BraziIl35 pp. 1996. Los camarones pelágicos del Pacífico HERBST,G. N., A. B. WILLIAMS& B. B. BOOTHE, Mexicano (Dendrobranchiata y Caridea). JR. 1979. Reassessment of northern geographic CONABIO/ Instituto de Ciencias del Mar y limits for decapod crustacean species in the Limnología, Universidad Nacional Autónoma Carolinian Province, USA; some major range de México 1-157. extensions itemized. Proceedings of the Biolo- HENDRICKX,M. E. & J. A. ESPARZA-HARO.1997. gical Society of Washington 91 (4): 989-998. A new species of Clibanarius (Crustacea, Ano- HERNÁNDEZAGUILERA,J. L. 1998. Sobre una co- mura, Diogenidae) from the eastern tropical lección de talasínidos (Crustacea: Decapoda) Pacifico Zoosystema, Paris 19 (1): 111-119. de la costa del Pacífico de México, con la des- HENDRICKX,M. E. & V. LANDA- JAIME. 1997. A cripción de una especie nueva del género Biffarius. new species of Parthenape Weber (Crustacea: Ciencias Marinas 24 (3):303-312. Brachyura: Parthenopidae) from the Pacific HERNÁNDEZAGUILERA,J. L. & J. L. VILLALOBOS- coast of Mexico. Bulletin de L'Institut des Ro- HIRIART.1980. Contribución al conocimiento yal des Sciences NatureIles de Belgique, Biolo- de los crustáceos decápodos y estomatópodos gie 67: 95-100. de la Sonda de Campeche. Investigaciones 0- HENDRICKX,M. E., M. DEMESTRE,A. ESPARZA- ceanográficas lB 80-07: 1-47. HARO& J. SALGADOBARRAGÁN.1997. Stoma- HERNÁNDEZAGUILERA,J. L & L. A. MARTÍNEZ topod and decapod crustaceans collected du- GUZMÁN.1992. Notas acerca de la distribución ring the Ceemex P7 cruises to the Gulf of Te- de los estomatópodos y décapodos de la isla huantepec, Mexico. Oceánides 11 (2): 1-28. Clarión, Archipiélago Revillagigedo, Colima, HENDRICKX,M. E., R. PÉREZ- GONZÁLEZ,L. M. México. Proceedings of the San Diego Society FLORES-CAMPAÑA& M. AYÓN-PARENTE.1997. ofNaturalHistory 19: 1-6. Nuevas capturas de dos especies raras de can- HERNÁNDEZAGUILERA,J. L., R. E. TORALALMA- grejos braquiuros (Crustacea: Decapoda: Bra- ZÁN& J. A. RUIZNuÑo. 1996. Especies catalo- chyura) para la costa del Pacífico Este tropical. gadas de crustáceos estomatópodos y decapo- Ciencias del Mar, Universidad Autónoma de dos para el Golfo de México, Rio Bravo, Tamps. Sinaloa 15: 51-53. a Progreso, Yuc. Dirección General de Ocea- HENDRICKX,M. E. & L. A. V ÁZQUEZ-CUREÑO. nografía Naval y CONABIO 98 pp., 17 lam. 1998. Composition and zoogeographical affi- HERNÁNDEZ,G. & J. BOLAÑOS.1995. Additions to nities of stomatopod and decapod crustacean the decapod crustacean fauna of northeastern fauna coIlected during the Ceemex P4 Cruise venezuelan Islands, with the description of the in the Gulf of Tehuantepec, Mexico. Bulletin male of Pinnatheres maseri Rathbun, 1918 (De- de L'Institut Royal des Sciences Naturelles de capoda: Brachyura: Pinnotheridae). Nauplius, Belgique, Biologie 68: 135-144. Rio Grande 3: 75-81. BOSCHI: DISTRIBUTION OF DECAPOD CRUSTACEANS 47

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