Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica

Wehrtmann, Ingo S.; Echeverría-Sáenz, Silvia fauna (Stomatopoda: ) associated with the deepwater fishery of Heterocarpus vicarius (Decapoda: Pandalidae) along the Pacific coast of Costa Rica Revista de Biología Tropical, vol. 55, núm. 1, 2007, pp. 121-130 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica

Available in: http://www.redalyc.org/articulo.oa?id=44909916

How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Crustacean fauna (Stomatopoda: Decapoda) associated with the deepwater fishery of Heterocarpus vicarius (Decapoda: Pandalidae) along the Pacific coast of Costa Rica

Ingo S. Wehrtmann 1,2,3 & Silvia Echeverría-Sáenz 1 1 Escuela de Biología, 2 Museo de Zoología y 3 Centro de Investigaciones en Ciencias del Mar y Limnología (CIMAR), Universidad de Costa Rica, 2060 San José, Costa Rica; [email protected]

Received 11-X-2005. Corrected 08-VIII-2006. Accepted 16-III-2007.

Abstract: Commercial bottom trawling is a successful and commonly used method to catch marine . However, the fishing gears are poorly selective, and in addition to the target species they catch and retain large quantities of non-target species (bycatch). This study presents data concerning species composition and depth distribution of the crustacean fauna (stomatopods and decapods) associated with Heterocarpus vicarius catches from Pacific Costa Rica. A total of 74 samples (three to five 20 min-tows each month) were taken between January 2004 and December 2005 with commercial shrimp trawlers at depths varying between 192 and 350 m. In all depth ranges analyzed, total catch of was significantly higher than that of fishes. A total of 28 decapods and two stomatopod species were identified. In comparison to other bycatch composi- tion of comparable fisheries in Latin America, the crustacean fauna of the H. vicarius fishery in Costa Rica is highly diverse. Most common species were Solenocera agassizii (Solenoceridae), Squilla biformis (Squillidae), Plesionika trispinus (Pandalidae), and Pleuroncodes sp. (Galatheidae), reaching total catch percentages of 57.2 %, 81.5 %, 91.8 %, and 99.6 % of individual catches, respectively. The results presented herein may contribute to the development of responsible management strategies for the deepwater fisheries in Costa Rica and Central America. Rev. Biol. Trop. 55 (Suppl. 1): 121-130. Epub 2007 June, 29.

Key words: Heterocarpus vicarius , deepwater fishery, bycatch, Central America, eastern tropical Pacific.

Commercial bottom trawling is a suc- and 10:1 (FAO 1982, Harris and Poiner 1990). cessful and commonly used method to catch Bycatch may be used as food in some coun- marine shrimps. These fishing activities are tries, however, in many others it is returned to considered as one of the primary causes of the sea, either because of its size or because physical perturbation to the seabed on the of poor commercial value (FAO 1982, Hill continental shelf and upper slope (Watling and Wassenberg 2000, Machias et al. 2001). and Norse 1998, Bozzano and Sardà 2002). Information concerning bycatch composition Typically, the shrimp fishing gears are poorly and discard rates are necessary to develop selective, and in addition to the target spe- more intelligent ways to harvest our marine cies they catch and retain large quantities resources (Hall et al. 2000). of non-target species, so-called “bycatch” The ecology of deepwater shrimp spe- (Andrew and Pepperell 1992, Hall et al. cies, their fishery and impact on the environ- 2000). Tropical shrimp trawl fisheries are ment have been especially well studied in the considered to have the highest discard rate in Mediterranean (Marsan et al. 2000, Belcari et the world’s marine fisheries (Kelleher 2005). al. 2003, Sardà et al. 2003a, b, 2004, Castro The ratio of bycatch to shrimps in tropical et al. 2005, Chilari et al. 2005). In contrast, and subtropical zones is usually between 5:1 our knowledge regarding these topics in Latin

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 121 America is far from complete. Concerning MATERIALS AND M ETHODS the Pacific, several studies have been pub- lished from Chilean waters, focusing on the Study site and sampling: The study area pandalid shrimp Heterocarpus reedi (e.g., extended approximately from 08°38’02’’ to Bahamonde and Henríquez 1965, Andrade 10°32’10’’ N and 84°05’54’’ to 86°10’42’’ W, and Báez 1980, Roa and Ernst 1996, Arana et covering the main fishing grounds for the target al. 2003b), the red royal shrimp Haliporoides species, Heterocarpus vicarius ; most samples diomedeae (Arana et al. 2003a and references were collected in the central Pacific region of therein), and the razor shrimp Campylonotus Costa Rica (09°26’46’’ to 09°20’54’’ N and semistriatus (Arana and Ahumada 2006). 85°05’50’’ to 84°36’16’’ W). Bottom water Hendrickx (1996, 2001, 2003) provided temperatures and salinities were around 11.3- insights concerning the presence of deep- 13.5°C and 34.9-35.1 psu, respectively (I.S. water shrimps of the Gulf of California, and Wehrtmann, unpubl. data). Hendrickx et al. (1998) reported data con- A total of 22 monthly samples (Fig. cerning growth, abundance, distribution and 1) were taken between January 2004 and biochemical composition of H. vicarius from December 2005 with commercial shrimp the southeastern Gulf of California. To our trawlers equipped with two standard epiben- knowledge, there is no published informa- thic nets; each net was 20.5 m long and had tion available regarding the fishery-biology a mouth opening of 5.35 x 0.85 m. The mesh of shrimps from deepwater zones of Pacific size was 4.45 cm, that of the codend 3.81 cm. Central America. During each month three to five 20 minute- Three deepwater decapod species are tows were carried out during daylight (74 sam- known to be commercially exploited along ples in total) at a speed of 0.8-1.2 nm. All catch the Pacific coast of Central America (Holthuis values refer to kg per 20-minute haul. The 1980): Solenocera agassizii , Heterocarpus initial and final positions were recorded by vicarius , and H. affinis . While the former a portable global satellite positioning system two species may co-occur in depths rang- (GARMIN GPS 12 XL). Fishing depths were ing between 200 and 400 m, H. affinis is obtained from an installed echo-sounder, and fished in considerably deeper waters (700–1 varied between 192 and 350 m. The location 400 m; Holthuis 1980, Hendrickx 1995b). of the first haul was decision of the captain Heterocarpus vicarius is known to occur along and was a place where he expected to find H. the Pacific coast from the Gulf of California to vicarius , generally between 250-300 m depth. Peru, and is commercially exploited in Costa The other hauls were carried out in the same Rica and Panama (Holthuis 1980, Hendrickx fishing ground, but at least one slightly deeper 1995b). Commercial landings of H. vicarius (ca. 300-350 m) and another one slightly in Costa Rica peaked in 1993 with roughly shallower (ca. 200-250 m) than the first tow. 400 tons, then decreased substantially well Depending on the amount of the catch, we below 100 tons per year, and increased again sorted the entire material of both nets or only in 2004, reaching almost 300 tons (H. Araya, representative portions of the catch. The mate- Instituto Costarricense de Pesca y Acuicultura, rial was separated on deck into the following pers. comm.). groups: H. vicarius (Pandalidae), Plesionika In order to facilitate the development of trispinus (Pandalidae), Solenocera agassizii responsible management strategies for deep- (Solenoceridae), Stomatopoda, Galatheidae, water resources in the area, here we present fish and others. These groups were weighted data concerning species composition and depth on board, and weight values were expressed as distribution of the crustacean fauna (stomato- percentage of the total catch. From each haul, pods and decapods) associated with H. vicarius several collected individuals of Stomatopoda, catches from Pacific Costa Rica. Galatheidae and the other uncommon groups

122 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 Fig. 1. Typical catch of Heterocarpus vicarius and associated fauna on board the shrimp trawler “Onuva”, December 2004, Pacific coast of Costa Rica. were separated and frozen for further identifica- were representatives of two different families tion in the Museo de Zoología of the Universidad (Hemisquillidae and Squillidae). Associated de Costa Rica in San José. For the identifica- decapod species belonged to 22 families; tion we used the keys provided by Hendrickx Penaeoidea were represented by two species (1995a, b, c, d) as well as other specific publica- from two families, Caridea by four species tions (e.g., Garth 1958, Haig 1960, Hendrickx from four families, Anomura by eight species and Salgado-Barragán 1991, Hendrickx and from four families, and Brachyura by 14 species Estrada Navarette 1996, Hendrickx 1997, 2000, from ten families (Table 1). The family with the Martin and Davis 2001). highest number of species encountered in the material was Galatheidae (4 spp.), followed by Calappidae (3 spp.). Most common species RESULTS (occurring in more than 60 % of all hauls; see Table 1) were Squilla biformis (Squillidae), Mean total length of the target species, H. Solenocera agassizii (Solenoceridae), Plesionika vicarius , and the other commercially exploited trispinus (Pandalidae), and Pleuroncodes sp. species, S. agassizii , was 89.5 mm (n= 14785) (Galatheidae). and 94.1 mm TL (n= 9898), respectively. Table 3 provides an overview of the total A total of 28 decapods and two stomatopod catch per group expressed as mean percentage species were identified (Table 1); the latter from all hauls obtained between January 2004

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 123 TABLE 1 Composition, depth range and occurrence of the associated crustacean fauna with the Heterocarpus vicarius -fishery along the Pacific coast of Costa Rica (January 2004 – December 2005).

Group Family Species Depth range (m) Occurrence

Stomatopoda Hemisquillidae Hemisquilla californiensis 273 rare (Stephenson, 1967) Squillidae Squilla biformis Bigelow, 1891 201-331 very common

Penaeoidea Sicyoniidae Sicyonia picta Faxon, 1893 201-274 common

Solenoceridae Solenocera agassizi Faxon, 1893 229-309 very common

Caridea Glyphocrangonidae Glyphocrangon alata Faxon, 1893 209 rare

Hippolytidae Lysmata sp. 232-275 rare

Pandalidae Plesionika trispinus Squires 229-326 very common &Barragán, 1976 Pasiphaeidae Pasiphaea americana Faxon, 1893 216-339 common

Anomura Diogenidae Paguristes bakeri Holmes, 1900 273 rare

Diogenidae Petrochirus californiensis Bouvier, 293 occasional 1895 Dorippidae Ethusa ciliatifrons Faxon, 1893 220-311 common

Galatheidae Munida gracilipes Faxon, 1893 220-300 common

Galatheidae Munida obesa Faxon, 1893 229-313 common

Galatheidae Munida refulgens Faxon, 1893 229-274 occasional

Galatheidae Pleuroncodes sp . 229-339 very common

Porcellanidae Porcellana hancocki Glassel, 1937 256 rare

Brachyura Calappidae Acanthocarpus alexandri Stimpson, 256-284 common 1871 Calappidae Hepatus kossmanni Newmann, 339 rare 1878 Calappidae Platymera gaudichaudii H. Milne 256-293 common Edwards, 1837 johngarthi Carvacho, 1989 201-339 common

Goneplacidae Oediplax granulata Rathbun, 1893 339 occasional

Inachidae Paradasygyius depressus (Bell, 311-339 occasional 1835) Inachidae Stenorynchus debilis (Smith, 1871) 293 rare

Leucosiidae Iliacantha hancocki Rathbun, 1935 298-339 occasional

Leucosiidae Persephona orbicularis Bell, 1855 220-339 common

Majidae Maiopsis panamensis Faxon, 1893 201-313 occasional

124 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 TABLE 1 (Continued) Composition, depth range and occurrence of the associated crustacean fauna with the Heterocarpus vicarius -fishery along the Pacific coast of Costa Rica (January 2004 – December 2005).

Mithracidae Stenocionops ovata (Bell, 1835) 216-256 occasional

Palicidae Palicus fragilis (Rathbun, 1893) 273-275 occasional

Parthenopidae Parthenope (Platylambrus ) exilipes 311 rare (Rathbun, 1893) Portunidae Portunus iridescens (Rathbun, 216-276 occasional 1893) very common = in >60 % of hauls; common = in >30 % of hauls; occasional = in >15 % of hauls; rare = in <15 % of hauls

TABLE 2 Comparison of crustacean species and family numbers recorded as bycatch from the Heterocarpus reedi – fishery in Chile (J.C. Orellana, unpubl. data) and H. vicarius – fishery in Costa Rica.

Group Species Families Chile Costa Rica Chile Costa Rica Stomatopoda 1 2 1 2 Penaeoidea 1 2 1 2 Caridea 2 4 2 4 Palinura 2 0 2 0 Anomura 4 8 2 4 Brachyura 5 14 5 10 Total 15 30 13 22

TABLE 3 Representation of taxonomic groups (mean weight percentage of total catch during January 2004- December 2005) associated with the Heterocarpus vicarius - fishery along the Pacific coast of Costa Rica.

Fish 25 % Stomotopoda 19 % Heterocarpus vicarius 18 % Galatheidae 17 % Solenocera agassizi 5 % Others 2.5 % Plesionika trispinus 14 % and December 2005. Fishes, stomatopods and Crustaceans were predominant (in terms the target species H. vicarius , were the most of weight of the total catch) in all three depth important taxa regarding total catch, followed ranges (Fig. 2; 200-250 m: ANOVA, F 5.19; by the squat (Galatheidae), and the p<0.05; 250-300 m: ANOVA, F 5.64; p<0.05; pandalid species, P. trispinus . Solenocera agas- 300-350 m: ANOVA, F 9.33; p<0.01). Highest sizii comprised 5 % of the total catch, and all total catch percentages from deeper waters the remaining species represented 2.5 %. (>300 m) were obtained for stomatopods,

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 125 2004: 91.8 %; 1 585 kg; 242 m; August 2005: 30 Crustaceans Fish 87.6 %, 495 kg, 201 m). 25 The target species, H. vicarius , predomi- 20 nated in terms of weight in catches carried out 15 between 250 and 300 m depth (Fig. 4). The 10 highest concentration of the target species was 5 in an individual haul at 267 m (October 2004; 0

Weight of Weight total catch (mean %) 200-249 250-300 301-350 106 kg; 70.5 % of total catch) closely followed Depth range (m) by a haul carried out in December 2004 at 280 Fig. 2. Mean total catch percent per depth range of crusta- m with 70.2 % of the total catch (135 kg). In ceans and fish associated with the Heterocarpus vicarius - six additional occasions H. vicarius constituted fishery along the Pacific coast of Costa Rica (January 2004 – December 2005). more than 60 % (range: 61.1-65.8 %) of the total catch per haul obtained from depths vary- 30 ing between 284 and 311 m. 25 30 20 Galatheidae 25 15 Stomatopoda P. trispinus 20 10 H. vicarius 15

Biomass (mean %) 5 S. agassizi 10 0 200-249 250-300 301-350 5 Biomass (mean %) Depth range (m) 0 200-249 250-300 301-350 Fig. 3. Mean total catch percent per depth range of Depth range (m) Galatheidae, Stomatopoda, and Plesionika trispinus asso- ciated with the Heterocarpus vicarius -fishery along the Fig. 4. Mean total catch percent per depth range of Pacific coast of Costa Rica (January 2004 – December Heterocarpus vicarius and Solenocera agassizi in trawls 2005). carried out along the Pacific coast of Costa Rica (January 2004 – December 2005). principally represented by Squilla biformis The second commercially important spe- (Squillidae). The mean weight percentages for cies, S. agassizii , showed a gradual decrease stomatopods increased gradually with depth of mean weight with increasing depth (Fig. 4), and reached a maximum in hauls carried out at varying between 8.7 % and 5.3 % of the total depth >300 m (24.5 % of the total catch caught catch per depth range. The highest concentra- in this depth range; Fig. 3). Maximum total tion refers to a haul carried out in August 2005 catch percentage for stomatopods was 81.5 % at 238 m depth, where S. agassizii comprised in a haul carried out in May 2005 at 331 m. 57.2 % of the total catch; in additional five Although the mean weight percentage of squat occasions, the species made up roughly a third lobsters (Galatheidae) was almost identical in of the total catch (range: 26.4 - 31.6 %) from all three depth ranges (varying between 12.3 depths ranging between 229 and 293 m. and 13.5 %; Fig. 3), in three occasions they made up practically the entire catch of the haul: February 2004 (92.8 %; 318 kg; 339 DISCUSSION m), November 2004 (99.6 %; 2 288 kg; 276 m), and April 2005 (92.3 %; 479 kg; 220 m). Crustaceans clearly dominated the catches For P. trispinus , highest (16.2 %) and lowest of Heterocarpus vicarius : 75 % of the total mean percentage values (9.1 %) were obtained catch were decapods and stomatopods, and from hauls between 301-350 m and 250-300 m, only 25 % was comprised by fishes (Table 3). respectively. However, almost entire individual Almost identical percentages have been report- hauls could consist of P. trispinus (September ed from the fishery of Heterocarpus reedi in

126 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 Chile (Arana et al. 2003b), a species which is shallower waters, Branco and Fracasso (2004) commercially exploited in similar depth ranges reported 28 decapod species associated with as H. vicarius (Bahamonde and Henríquez the fishery in southern 1965, Arana et al. 2003b). Although fish and Brazil, and Pérez-Barros et al. (2004) found 16 crustaceans were the dominant megafaunal decapod species in catches of Munida spp. in groups in deep sea areas of the Mediterranean, the Beagle Channel, Argentina. decapod biomass was higher only at depths The crustacean fauna of the H. vicarius >1 500 m (Company et al. 2004). These fishery in Costa Rica is highly diverse when discrepancies with the above-mentioned data compared to other bycatch compositions of regarding Heterocarpus spp. fisheries may be comparable fisheries in Latin America. Species related to the different environmental condi- numbers of decapods are almost three times tions, especially to the oligotrophic nature of higher than in the H. reedi fishery off Chile the Mediterranean. where the associated decapod fauna is com- Wicksten (1989) listed a total of 183 prised by a total of 11 species (Arana et al. deepwater (>200 m) crustacean species for 2003b). More recently, however, J.C. Orellana the eastern Pacific Ocean, of which 100 occur (pers. comm.) listed a total of 14 decapod spe- within the limits of the eastern tropical Pacific cies as bycatch from the Chilean H. reedi fish- (Hendrickx 1996). Thus, species encountered ery (Table 2). Interestingly, Andrade and Báez as bycatch in H. vicarius hauls in Costa Rica (1980) studied the decapod crustaceans caught represent 30 % of the crustacean deepwater with the H. reedi fishery during 1976 and 1979 fauna of the entire eastern tropical Pacific. in Chile and reported a total of 23 decapods, Hendrickx (1996) analyzed the biodiversity considerably more species than encountered of decapod crustaceans in the SE Gulf of during the more recent surveys (Arana et al. California, , and reported 19 decapod 2003b, J.C. Orellana, pers. comm.). species from the upper slope (200- 1 200 The crustacean fauna associated with the m). The major difference in species numbers two Heterocarpus fisheries in Latin America between both studies refers to Brachyura: while (H. reedi in Chile, H. vicarius in Costa Rica) only three species of Brachyura were found in has only one species in common: the caridean deepwater areas of the SE Gulf of California, shrimp Glyphocrangon alata , known to occur Mexico (Hendrickx 1996), our study revealed in Pacific deepwater zones from Mexico to the presence of 14 species (Table 1). More Chile. According to Hendrickx (1995b), this recently, Hendrickx (2001) reported a total of species can be found between 600 and 1 325 31 species collected between 550 and 2 250 m m, while our single individual was caught at in the SE Gulf of California, Mexico, including a depth of 209 m, substantially shallower than just one brachyuran species and 13 species the previously indicated depth range. Apart of Caridea, considerably more than in our study from G. alata , representatives of three genera (4 spp.). However, both studies conducted in (Pleuroncodes , Munida , and Cancer ) were the SE Gulf of California (Hendrickx 1996, encountered in both fisheries. In general, the 2001) included the fauna of considerably deep- associated fauna in Costa Rica contained twice er depth intervals than our study, thus making as many species per taxonomic group as the more detailed comparisons difficult. Species Heterocarpus -fishery in Chile (Table 2). There number can also be compared to the crustacean are, however, two exceptions: (1) two species fauna associated to Pandalus platyceros catches of Palinura were reported from Chile, while in Baja California, Mexico, which were carried none representative of this infraorder was col- out in depths between 90 and 290 m. According lected in our study; (2) the species number of to Flores et al. (2004), the associated fauna was brachyuran was three times higher than comprised by a total of nine crustacean spe- in Chile (Table 2). The absence of representa- cies: two stomatopods and seven decapods. In tives of Palinura in our study is not surprising,

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 127 since these species have not been reported Costa Rica. Entre enero del 2004 y diciembre de 2005, se from deepwater zones along the Pacific coast tomaron 74 muestras en profundidades de 192-350 m (tres of Costa Rica. The fact that considerably more a cinco arrastres de 20 min cada mes), utilizando redes de arrastre comercial dirigidas a la pesca de camarones. En brachyuran species were encountered in our todos los intervalos de profundidad analizados, las capturas samples compared to those in Chile may reflect (kg) de crustáceos fueron significativamente mayores que higher species diversity in deepwater zones of las de los peces. Se identificaron 28 especies de decápodos Central America. y dos de estomatópodos. Al comparar la composición de The information presented herein may especies de fauna acompañante de H. vicarius en Costa Rica con otras pesquerías de camarones en América Latina, contribute to a better understanding of the se encuentra que la fauna de crustáceos es muy diversa. deepwater fishery in Central America and its Las especies más comunes fueron Solenocera agassizii impacts on benthic communities, thus favoring (Solenoceridae), Squilla biformis (Squillidae), Plesionika the development of responsible management trispinus (Pandalidae) y Pleuroncodes sp. (Galatheidae), strategies for deepwater resources in the area. alcanzando porcentajes máximos de concentración - en un sólo arrastre - de hasta 57.2 %, 81.5 %, 91.8 % y 99.6 % de la captura, respectivamente. Los resultados presentados aquí pueden contribuir al desarrollo de estrategias de mane- ACKNOWLEDGMENTS jo responsable para las pesquerías de aguas profundas en Costa Rica y en Centroamérica. The study was part of a project about stan- Palabras clave: Heterocarpus vicarius , pesquería dards for a sustainable deepwater shrimp fishery de aguas profundas, fauna acompañante, América Central, in Pacific Costa Rica, which received funding Pacífico este tropical. from the German Government (Ministerium für wirtschaftliche Zusammenarbeit und Entwicklung, BMZ) and the Ristic AG REFERENCES (Oberferrieden, Germany). Additional support came from the Universidad de Costa Rica (proj- Andrade, H. & P. Báez. 1980. Crustáceos decápodos ect # V.I. 111-A4-508). Special thanks to the cap- asociados a la pesquería de Heterocarpus reedi Bahamonde, 1955 en la zona central de Chile. Bol. tain (Rigoberto Villalobos Cruz) and the crew of Mus. Nac. Hist. Nat. Chile 37: 261-267. the shrimp trawler “Onuva” as well as to Ronny Gründler, The Rainbow Jewels, (Puntarenas, Andrew, N.L. & J.G. Pepperell. 1992. The by-catch of Costa Rica). We are more than grateful to all the shrimp trawl fisheries, p. 527-565. In M. Barnes, students who helped us during the fieldwork. We A.D. Ansell & R.N. Gibson (eds.). Oceanogr. Mar. Biol. Ann. Rev. 30. University of California Press, appreciate the help provided by Juan Bautista Berkeley, California, USA. Chavarría (Universidad de Costa Rica, Escuela de Estadística) and Rita Vargas (Universidad Arana, P.M. & M. Ahumada. 2006. Camarón navaja de Costa Rica, Museo de Zoología) concerning (Campylonotus semistriatus ), crustáceo de aguas pro- statistical analyses and species identification, fundas frente a la costa central de Chile (Crustacea, Decapoda, Campylonotidae). Invest. Mar., Valparaíso respectively. Patricio Arana sent us valuable 34: 3-14. literature for the discussion. Arana, P.M., M. Ahumada & A. Guerrero. 2003a. Distribución y abundancia de la gamba Haliporoides diomedeae RESUMEN (Crustacea: Decapoda: Penaeidae) frente a la costa central de Chile. Invest. Mar., Valparaíso 31: 57-71. Las redes de arrastre son un método exitoso y común- mente utilizado para la pesca de camarones marinos. Sin Arana, P., M. Ahumada, A. Guerrero, S. Palma, T. embargo, son poco selectivas y, junto con las especies Melo, D. Queirolo, R. Bahamonde, M. A. Barbieri comerciales, se pesca también grandes cantidades de otras & C. Canales. 2003b. Evaluación directa de especies (fauna acompañante). Este estudio presenta infor- camarón nailon entre la II y VII Regiones, año mación acerca de la composición y distribución batimétrica 2003 (Proyecto FIP Nº 2003-05). Informe Pre- de los crustáceos (estomatópodos y decápodos) asociados Final. Estud. Doc, Pont. Univ. Católica Valparaíso con las capturas de Heterocarpus vicarius en el Pacífico de 25/2003. 297 p.

128 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 Bahamonde, N. & G. Henríquez. 1965. Sinopsis de datos Haig, J. 1960. The Porcellanidae (Crustacea Anomura) of biológicos sobre el camarón nailon Heterocarpus the eastern Pacific. Allan Hancock Pac. Exp. Univ. reedi Bahamonde, 1955. Santiago, Chile, FAO Fish. South. Cal. Press. Vol 24. 440 p. Synop. 107: 1607-1627. Hall, M., D.L. Alverson & K.I. Metuzals. 2000. By-catch: Belcari, B., C. Viva, M. Mori & S. De Ranieri. 2003. problems and solutions. Mar. Pollut. Bull. 41: 204-219. Fishery and biology of Aristaeomorpho foliacea (Risso, 1827) (Crustacea: Decapoda) in the northern Harris, A.N. & I.R. Poiner. 1990. By-catch of the Tyrrhenian Sea (western Mediterranean). J. Northw. fishery of Torres Strait; composition and partitioning Atl. Fish. Sci. 31: 195-204. of the discards into components that float or sink. Aust. J. Mar. Freshwater Res. 41: 37-52. Bozzano, A. & F. Sardà. 2002. Fishery discard consump- tion rate and scavenging activity in the northwestern Hendrickx, M.E. 1995a. Estomatópodos, p. 355-382. In W. Mediterranean Sea. ICES J. Mar. Sci. 59: 15-28. Fischer, F. Krupp, W. Schneider, C. Sommer, K.E. Carpenter & V.H. Niem (eds.). Guía FAO para la Branco, J.O. & H.A.A. Fracasso. 2004. Ocorrencia e abun- identificación de especies para los fines de la pesca. dancia da carcinofauna acompanhante na pesca do Pacífico centro-oriental. Volumen I. Plantas e inver- camarao sete-barbas, Xiphopenaeus kroyeri Heller tebrados. FAO, Rome, Italy. (Crustacea, Decapoda), na Armacao do Itapocoroy, Penha, Santa Catarina, Brasil. Rev. Brasil. Zool. 21: Hendrickx, M.E. 1995b. Camarones, p. 417-538. In W. 295-301. Fischer, F. Krupp, W. Schneider, C. Sommer, K.E. Carpenter & V.H. Niem (eds.). Guía FAO para la Castro, M., A. Araújo & P. Monteiro. 2005. Fate of discards identificación de especies para los fines de la pesca. from deep water crustacean trawl fishery off the south Pacífico centro-oriental. Volumen I. Plantas e inver- coast of Portugal. New Zealand J. Mar. Freshwat. tebrados. FAO, Rome, Italy. Res. 39: 437-446. Hendrickx, M.E. 1995c. Anomuros, p. 539-564. In W. Chilari, A., M. Thessalou-Legaki & G. Petrakis. 2005. Fischer, F. Krupp, W. Schneider, C. Sommer, K.E. Population structure and reproduction of the Carpenter & V.H. Niem (eds.). Guía FAO para la deep-water shrimp Plesionika martia (Decapoda: identificación de especies para los fines de la pesca. Pandalidae) from the eastern Ionian Sea. J. Crust. Pacífico centro-oriental. Volumen I. Plantas e inver- Biol. 25: 233-241. tebrados. FAO, Rome, Italy.

Company, J.B., P. Maiorano, A. Tselepidies, C.-Y. Politou, Hendrickx, M.E. 1995d. Cangrejos, p. 565-636. In W. W. Plaity, G. Rotllant & F. Sardà. 2004. Deep-sea Fischer, F. Krupp, W. Schneider, C. Sommer, K.E. decapod crustaceans in the western and central Carpenter & V.H. Niem (eds.). Guía FAO para la Mediterranean Sea: preliminary aspects of species identificación de especies para los fines de la pesca. distribution, biomass, and population structure. Sci. Pacífico centro-oriental. Volumen I. Plantas e inver- Mar. 68 (Suppl. 3): 73-86. tebrados. FAO, Rome, Italy.

FAO, 1982. Fish By-Catch – Bonus from the Sea. Report Hendrickx, M.E. 1996. Habitats and biodiversity of deca- of a Technical Consultation on Shrimp By-Catch pod crustaceans in the SE Gulf of California, Mexico. Utilization held in Georgetown, Guayana, 27/30 Rev. Biol. Trop. 44: 603-617. October 1981. IDRC, Ottawa, Ontario, Canada. 163 p. Hendrickx, M.E. 1997. Los cangrejos braquiuros (Crustacea: Decapoda: Dromiidae, hasta Leucosiidae). Inst. Flores, J., M.E. Hendrickx & H. Chapa-Saldaña. 2004. Cienc. Mar Limnol., Univ. Nal. Autón. México y Capturas del camarón mancha blanca, Pandalus CONABIO. México. 178 p. platyceros Brandt, 1851 (Caridea: Pandalidae), en la costa oeste de Baja California, México, con una breve Hendrickx, M.E. 2000. The genus Munida Leach revisión de su biología y su pesca, p. 199-211. In (Crustacea, Decapoda, Galatheidae) in the eastern M.E. Hendrickx (ed.). Contributions to the Study of tropical Pacific, with description of two new species. East Pacific Crustaceans 3, Inst. Cienc. Mar. Limnol, Bull. Inst. Roy. Sci. Nat. Belgique 70: 163-192. UNAM, México. 245 p. Hendrickx, M.E. 2001. Occurence of a continental slope Garth, J.S. 1958. Brachyura of the Pacific coast of America, decapod crustacean community along the edge of Oxyrhyncha. Allan Hancock Pac. Exp. Univ. South. the minimum oxygen zone in south eastern Gulf of Cal. Press. Vol 21, Parts I & II. 854 p. California, Mexico. Belg. J. Zool. 131: 95-109.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007 129 Hendrickx, M.E. 2003. Size and abundance of deep water in the southern Adriatic basin (Mediterranean shrimps on the continental slope of the SE Gulf of Sea), p. 763–769. In J.C. von Vaupel Klein & California, Mexico, p. 227-234. In M.E. Hendrickx F.R. Schram (eds.). The Biodiversity Crisis and (ed.). Contributions to the Study of East Pacific Crustacea. Proceedings of the Fourth International Crustaceans 3, Inst. Cienc. Mar. Limnol, UNAM, Crustacean Congress. A.A. Balkema, Rotterdam, The México. 303 p. Netherlands.

Hendrickx, M.E. & J. Salgado-Barragán. 1991. Los estom- Martin, J. & G. Davis. 2001. An updated classification of atópodos (Crustacea: Hoplocarida) del Pacífico the recent Crustacea. Natural History Museum of Los Mexicano. Inst. Cienc. Mar Limnol., Univ. Nal. Angeles County, Science series no. 39. 132 p. Autón. México, Publ. Esp. 10: 1-200. Pérez-Barros, P., F. Tapella, M.C. Romero, J.A. Calcagno Hendrickx, M.E. & F.D. Estrada Navarrete. 1996. & G.A. Lovrich. 2004. Benthic decapod crustaceans Los camarones pelágicos del Pacífico mexicano associated with captures of Munida spp. (Decapoda: (Dendrobrachiata y Caridea). Inst. Cienc. Mar Anomura) in the Beagle Channel, Argentina. Sci. Limnol., Univ. Nal. Autón. México y CONABIO. Mar. 68: 237-246. México. 157 p. Roa, R. & B. Ernst. 1996. Age structure, annual growth, and Hendrickx, M.E., F. Páez-Osuna & H.M. Zazueta-Padilla. variance of size-at-age of the shrimp Heterocarpus 1998. Biology and biochemical composition of the reedi . Mar. Ecol. Progr. Ser. 137: 59-70. deep-water shrimp Heterocarpus vicarius Faxon (Crustacea: Decapoda: Caridea: Pandalidae) from the Sardà, F., J.B. Company & A. Castellón. 2003a. southeastern Gulf of California, Mexico. Bull. Mar. Intraspecific aggregation structure of a shoal of Sci. 63 : 265-275. a western Mediterranean (Catalan coast) deep-sea shrimp, Aristeus antennatus (Risso, 1816) during the Hill, B.J. & T.J. Wassenberg. 2000. The probable fate of reproductive period. J. Shellfish Res. 22: 569-579. discards from prawn trawlers fishing near coral reefs. A study in the northern Great Barrier Reef, Australia. Sardà, F., J.B. Company & F. Maynou. 2003b. Deep- Fish. Res. 48: 277-286. sea shrimp Aristeus antennatus Risso, 1816 in the Catalan Sea, a review and perspectives. J. Northw. Holthuis, L.B. 1980. Shrimps and of the world. An Atl. Fish Sci. 31: 127-136. annotated catalogue of species of interest to fisheries. FAO Fish. Synop.125: 1-271. Sardà, F., G. D’Onghia, C.Y. Politou, J.B, Company, P. Maiorano & K. Kapiris. 2004. Deep-sea distribu- Kelleher, K. 2005. Discards in the world’s marine fisheries. tion, biological and ecological aspects of Aristeus An update. FAO Fish. Techn. Pap. 470: 1-131. antennatus (Risso, 1816) in the western and central Mediterranean Sea. Sci. Mar. 68 (Suppl. 3): 117- Machias, A., V. Vassilopoulou, D. Vatsos, P. Bekas, A. 127. Kallianiotis, C. Papaconstantinou & N. Tsimenides. Watling, L. & E.A. Norse. 1998. Disturbance of the sea- 2001. Bottom trawl discards in the northeastern bed by mobile fishing gear: a comparison to forest Mediterranean Sea. Fish. Res. 53: 181-195. clearcutting. Conserv. Biol. 12: 1180-1197.

Marsan, R., N. Ungaro, C.A. Marano & M.C. Marzano. Wicksten, M.K. 1989. Ranges of offshores decapod crus- 2000. Remarks on distribution and fishery biology taceans in the eastern Pacific Ocean. Transact. San of some Plesionika species (Decapoda, Pandalidae) Diego Soc. Nat. Hist. 21: 291-316.

130 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 55 (Suppl. 1): 121-130, June 2007