This Article Was Originally Published in a Journal Published by Elsevier, and the Attached Copy Is Provided by Elsevier For

This article was originally published in a journal published by Elsevier, and the attached copy is provided by Elsevier for the author’s benefit and for the benefit of the author’s institution, for non-commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues that you know, and providing a copy to your institution’s administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution’s website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier’s permissions site at:

http://www.elsevier.com/locate/permissionusematerial Fisheries Research 85 (2007) 68–73

Demersal fish density in the upwelling ecosystem off Colombia, Caribbean Sea: Historic outlook Camilo B. Garc´ıa a,∗, Luis O. Duarte b,c, Jairo Altamar b, Luis M. Manjarres´ b a Departamento de Biolog´ıa, CECIMAR/INVEMAR, Universidad Nacional de Colombia, A.A. 1016, Santa Marta, Colombia b Laboratorio de Investigaciones Pesqueras Tropicales, Universidad del Magdalena, Cra. 32 #22-08, Santa Marta, Colombia c Programa de Postgrado en Oceanograf´ıa, Departamento de Oceanograf´ıa, Universidad de Concepción, Casilla 160-C, Concepción, Chile Received 14 July 2006; received in revised form 11 December 2006; accepted 21 December 2006

Abstract We conducted a review of historic demersal fish biomass estimates (1970–2001) in the upwelling area off northern Colombia. A clear pattern of decline in demersal fish biomass has been identified. Total demersals and most selected species groups showed a negative trend in mean biomass. Partial recovery in biomass by year 2001 as compared to 1970 was detected, concomitantly with reductions in fishing effort by the shrimp trawling fleet. Triggerfishes and puffers showed an increasing trend in biomass probably as a result of good survivorship as discards from the shrimp fishery. These findings are discussed in the context of current general concerns on the health of world marine ecosystems. © 2007 Elsevier B.V. All rights reserved.

Keywords: Biomass trends; Tropical ﬁsh; Upwelling; Colombia; Caribbean Sea

1. Introduction 1980–1984 period, declining afterwards to the end of their study period (1990–1994). In Colombian Caribbean waters the only Deterioration of ecosystems should ultimately be reflected published register of a decline in historic landings is that of in steady reductions of biomass (Pitcher, 1998). Recent global Garc´ıa and Solano (1995) with regard to Tarpon atlanticus. assessments of marine fish biomass conclude exactly that. For The present work compiles and presents historic changes instance, Myers and Worm (2003) estimate that large preda- (1970–2001) in demersal fish biomass in the upwelling area off tory fish biomass today is only about 10% of pre-industrial Colombia, Caribbean Sea (Fig. 1). Information from scientific levels. Watson and Pauly (2001) showed that global landings cruises which had the general purpose of estimating biomass lev- tended to decline by 0.36 million tonnes per year since 1988 els of demersal fishes, was compiled and analysed for biomass after correcting for misreporting to FAO. trends. Although there is a general concern about the conservation status of Western Atlantic marine ecosystems (Jackson, 2. Methods 2001), regional or local assessments of fish biomass trends in the Caribbean Sea have been scarce. The only regional study A total of 13 cruises have been conducted in the last 30 years known to us that quantitatively explores catch trends is that of in the study area, several of them in the same year. Temporal Baum et al. (2003) that reviewed the status of sharks in the coverage of the cruises is uneven. Cruises were conducted in north Atlantic including the Caribbean Sea. They show that 1970 (3 cruises), 1988 (4 cruises), 1992 (2 cruises), 1995 (2 coastal and oceanic shark populations have rapidly declined. cruises), 1996 (1 cruise) and 2001 (1 cruise) yielding a total Caddy et al. (1998) found for FAO statistical area 31 including of 237 tows. Tow speed and duration, and depth, that may the Caribbean that landings of shelf dependent species (dem- influence composition of capture (Garc´ıa et al., 1998; Winger ersal fish plus commercialAuthor's benthic invertebrates) peaked in thepersonalet al., 1999; Manjarres´ et al.,copy 2001), varied short as well as mesh size (Table 1). In order to assure comparability among cruises, data are expressed as biomass density, i.e., kg per 2 ∗ Corresponding author. Tel.: +57 13165237; fax: +57 13165310. km , estimated by the swept area method. Because all research E-mail address: [email protected] (C.B. Garc´ıa). vessels were otter trawlers using similar scientific fishing gears

Fig. 1. The upwelling ecosystem off Colombia showing tows carried out during the sampling dates. Data recorded in the 1990s (1992–1996) were grouped for the analysis. and simple random sampling was applied in all cruises with gies (e.g., Calamus penna), spadefishes (only Chaetodipeturs the only constraint posed by bottoms not amenable to trawling, faber), boxfishes (e.g., Acanthostracion quadricornis), grunts we assume a catchability of 1 in this calculation as we do not (e.g., Haemulon aurolineatum) and other demersals (remain- expect it to change over time in the context of this study. ing species, e.g., Polydactilus virginicus, Priacanthus arenatus, In the analysis species groups were preferred over individ- etc.). Species that were caught in the demersal nets but are ual species in order to reduce stochastic fluctuations. These considered of pelagic habit were excluded from analysis. groups constitute the bulk of demersal fish biomass, are of par- In order to picture time trends arithmetic mean biomass per ticular interest due to their alleged sensitivity to overfishing, year for each group were plotted against year. Data recorded like rays and sharks (e.g., Casey and Myers, 1998; Stobutzki et in the 1990s were grouped for this plot. Nonparametric 95% al., 2002; Baum et al., 2003), or support a long standing fish- confidence intervals of the biomass were derived by the bias- ing pressure, like snappers (Manjarres´ et al., 2004a,b). Species corrected and accelerated bootstrap method (Efron, 1987). groups included are: total demersals, snooks (e.g., Centropomus Lineal trends with time were sought by correlating (Pear- undecimalis), mojarras (e.g., Eucinostomus spp.), croakers (e.g., son’s coefficient) year and yearly mean biomass (no grouping Micropogonias furnieri), goatfishes (e.g., Upeneus parvus), of data recorded in the 1990s). Correlation between historic sharks (e.g., Rhizoprionodon porosus), triggerfishes (e.g., Bal- effort (days at sea) exerted by the shrimp fleet and yearly mean istes capriscus), puffers (e.g., Diodon holacanthus), snappers biomass was also investigated. Significance of correlation was (e.g., Lutjanus analis), rays (e.g., Dasyatis americana), por- tested via a bootstrap procedure that tests the null hypothesis

Table 1 Historical scientiﬁc cruises carried out in the upwelling ecosystem off Colombia, Caribbean Sea Year Vessel Headrope length (m) Codend mesh (mm) Towing velocity (knots) Towing duration (h) Depth (m) Number of tows Source

1970 Choco´ 43.0Author's and 54.0 36 personal 2.5 0.1–2.0 copy 13–134 32 1 1988 Fridtjof Nansen 31.0 20 1.0–4.5 0.2–0.6 11–151 96 2 1992 Malpelo/Ancon´ 33.3/26.6 50/54 3.4–3.9 0.5–3.0 25–89 21 3 and 4 1995 Ancon´ 26.6 45 1.7–4.4 0.5–0.6 13–90 33 5 1996 Malpelo 33.3 50 2.9–4.5 0.5–0.6 14–146 16 5 2001 Ancon´ 26.6 45 2.8–3.8 0.5 10–88 39 6

Source:1,Testaverde and R´ıos (1972);2,Strømme and Saetersdal (1989);3,Quintero (1992);4,Zuniga˜ and Escobar (1993);5,Manjarres´ et al. (2005a,b,c);6, Manjarres´ (2002). 70 C.B. Garc´ıa et al. / Fisheries Research 85 (2007) 68–73 that the observed correlation value is to be expected by chance. Snooks were scarce, probably due to their rather near coast- Mean biomass of year 1970, the 1990s (data grouped for this estuarine habit (Orrell, 2002), and hence were not amenable to analysis) and 2001 was contrasted to each other via a bootstrap statistical analysis. Nevertheless, it is noticeable that in practical procedure that tests the null hypothesis that the observed differ- terms they disappear in the 1990s (Fig. 2). ence between means is to be expected by chance. Routines for Triggerfishes and puffers share the feature that year 2001 these tests were written in Resampling Stats®. biomass is significantly higher than 1970 biomass level (p < 0.05 in both cases; for puffers the contrast is against 1988 as in 3. Results 1970 puffers were registered in only one tow). For triggerfishes biomass level in 2001 is also significantly higher than in the Temporal change in mean annual biomass and their boot- 1990s (p < 0.05), whereas for puffers it is not (p > 0.05). This strap confidence intervals is shown in Fig. 2. The biomass of suggests that the increase in biomass for triggerfishes found in total demersals is high in 1970, gets somewhat reduced in 1988, 2001 has been abrupt, whereas in the case of puffers the increase declines severely in the 1990s and shows some recovery in 2001. has been more uniform. This pattern is followed by a number of the groups but not all For other demersals mean biomass in 1970 is significantly of them, suggesting that not all demersal fish respond the same higher than in the 1990s and 2001 (p < 0.05), whereas mean way to the vagaries of time. biomass of 2001 is not significantly higher (p > 0.05) than that Negative lineal trends of mean biomass with time were found in the 1990s. This suggests that for this group biomass has not for total demersals, goatfishes, croakers, goatfishes, rays, sharks, recovered from the low level in the 1990s. mojarras, snappers, porgies, boxfishes, spadefishes, and other demersals. Of these, the negative lineal trend was significant 4. Discussion (p < 0.05) for total demersals, croakers, goatfishes, sharks and other demersals. Positive lineal trends were found for grunts, A clear pattern of decline in demersal fish biomass has been triggerfishes, and puffers, of which only that of puffers was identified. So strong that the recovery found in 2001 was not significant (p < 0.05). enough to disperse statistical significance of the negative linear Correlation between historic effort exerted by the shrimp fleet trend with time in total demersals, several fish groups and other (Fig. 3) and mean yearly biomass (ρ = −0.78) turned out to be demersals. With few exceptions, the fish group biomass was significant (p < 0.05) signalling a probable cause of the observed much lower in the 1990s than in 1970 and showed different demersal fishes time trajectories. levels of recovery (including no recovery at all) in 2001, thus When years 1970, the 1990s and 2001 were contrasted the suggesting that the different fish groups responded differently following results were obtained: for total demersals, croakers, and at different speeds to factors affecting biomass. goatfishes and, marginally for rays (p = 0.09), mean biomass Additional evidence of deterioration of fish biomass in the in year 1970 is significantly higher than both in the 1990s Colombian Caribbean comes from National fishery statistics. and in 2001 (p < 0.05). At the same time for all four groups Landings by the artisanal fleets in the last decade have decreased mean biomass in 2001 is significantly higher than in the 1990s from an average of 9546 tonnes per year between 1990 and 1992 (p < 0.05, p = 0.08 for rays). This suggests that for the demer- to 3937 tonnes per year between 1998 and 2000 (INPA, 1999, sal fish assemblage, in general, and for croakers, goatfishes and 2001). It is interesting to note that landings by bottom trawls rays, in particular, although a significant recovery of biomass in in the Caribbean in general have followed a similar pattern with 2001 has taken place as compared to the 1990s, its level is still peaks in the period of 1983–1997 and a steady decline thereafter far from the biomass level in 1970. (http://www.seaaroundus.org/). For sharks no significant difference in biomass was found. A number of factors may well be responsible for the patterns This finding stands in contrast to the significant negative lin- above. Ranging high is fishing pressure. The significant negative eal trend found (see above). This suggest that although mean correlation between effort by the shrimp fleet and yearly mean biomass did decrease with time, individual tows were highly biomass reinforces the argument. Thus, the recovery of biomass variable. detected in year 2001 is concomitant with a reduction of effort For mojarras, snappers, porgies and grunts biomass in 1970 by the industrial shrimp fleet in terms of number of operational is not significantly higher than in 2001 (p > 0.05), but biomass trawlers due to low yields and profitability. Historically effort in both those years is significantly higher than in the 1990s in the period 1975–1980 amounted to 4206 (±215 S.E.) days at (p < 0.05). This suggests for these groups that biomass in year sea per year, peaked to 11215 (±730 S.E.) days at sea per year 2001 has recovered to the level found in 1970. between 1985 and 1993 and declined to 3260 (±352 S.E.) days Boxfishes biomass in the 1990s was marginally higher at sea per year between 1995 and 2000 (Mora, 1988; INDER- (p = 0.07) than in 2001 butAuthor's similar to that in 1970 (p > 0.05). personalENA/INPA unpublished data, copy see Fig. 3). It is well known that This suggest that after a moderate burst in the 1990s biomass shrimp trawling has profound effects in demersal communities has returned to 1970 levels by 2001. Spadefishes biomass could due to direct mortality in the form of bycatch (Alverson et al., only be contrasted for years 1988 and 2001 because no spade- 1994) and destruction of habitat (Jennings and Kaiser, 1998). In fish was registered in 1970 and it was so in only one tow in a recent simulation study that covers part of our geographical 1990. Biomass in 1988 was marginally higher (p = 0.08) than in area, Criales-Hernandez et al. (2006) showed that the reduction 2001. of bycatch from shrimp trawling was conductive to rapid rebuild- C.B. Garc´ıa et al. / Fisheries Research 85 (2007) 68–73 71

Fig. 2. Historic biomass densities of demersal fishes in the upwelling ecosystem off Colombia. Confidence intervals were derived by the bias-corrected and accelerated bootstrap method. ing of biomass in sensitive fish groups like croakers, snappers, reduction in fishing is necessary but is not sufficient for recov- rays and sharks, among others. ery and discusses a number of variables that might correlate Hence, and assuming a causal relationship between reduc- with recovery including several aspects of life history like age at tion of fishing pressure and recovery of biomass levels in the maturity, fecundity, size at maturity and individual growth rate. upwelling area of the Colombian Caribbean, the finding that Tropical fishes are expected to have shorter generation times in about 5 years (1996–2001)Author's biomass for several fish groupspersonal(Longhurst and Pauly, 1987 copy), i.e., they will mature, grow and may recover to levels before massive shrimp trawling, stands die at a faster rate than their temperate counterparts and these in contrast to findings in higher latitudes that biomass recov- features correlate positively with potential maximum popula- ery may take decades or may just not happened after release tion growth (Hutchings, 2004). Thus, if populations have not from fishing. Hutchings (2000) found that the time required for been reduced to the level of seriously affecting interspecific recovery after long-term declines appears to be considerable, competition, mating success, genetic variation, etc., on the one particularly in demersal fishes. Hutchings (2004) concludes that hand, and essential habitats maintain a minimum integrity, on 72 C.B. Garc´ıa et al. / Fisheries Research 85 (2007) 68–73

Our ﬁndings are very much in line with current diagnostics of the state of world living marine resources and add to the need for regulatory actions (Jackson and Sala, 2001; Pauly et al., 2003) if we are to maintain the biological basis that sustains protein production in the sea for humankind.

Acknowledgements

This work was supported by COLCIENCIAS grant 3135- 09-11245 for the project “Dinamica´ espacio-temporal del ecosistema de afloramiento del area´ Bocas de Ceniza- Fig. 3. Time series of shrimp trawling effort (days at sea) reported in the Punta Espada (Caribe Colombiano) y sus implicaciones upwelling ecosystem off Colombia, Caribbean Sea. para un regimen´ de pesca responsable” and by Universi- dad del Magdalena, Universidad Nacional de Colombia and the other, tropical life history traits might explain the findings INPA/INCODER. L.O. Duarte was also supported by Deutscher here. Akademisher Austauschdienst DAAD under a PhD scholarship. In line with findings that elasmobranches have low sustain- C.B. Garc´ıa and L.O. Duarte acknowledge additional financial ability faced with fish trawling (e.g., Stobutzki et al., 2002; Baum support by European Community grant 003739 to the project et al., 2003) we found that rays and sharks are among the affected “Integrating multiple demands on coastal zones with emphasis groups. So, for instance, the biomass of rays by year 2001 had on aquatic ecosystems and fisheries”, INCOFISH. We are grate- not yet returned to 1970 levels and mean biomasses of sharks ful to Paul Gomez,´ Fabian´ Escobar and Jorge Viana˜ for help in steadily decreased in the period. gathering and processing of historic data. Suggestions by two Triggerfishes and puffers showed their highest biomass level anonymous referees helped to improve the paper. by year 2001, although for triggerfishes another peak was found in 1988. These fishes have no economic value in the region, thus References are systematically discarded or not targeted by fishermen. This, combined with resistance to periods out of the water, may rep- Alverson, D.L., Freeberg, M.H., Murawski, S.A., Pope, J.G., 1994. A global resent a plausible mechanism explaining the observed trajectory assessment of fisheries bycatch and discards. FAO Fish. Tech. Pap. 339, in time of their biomass. 1–233. Baum, J.K., Myers, R.A., Kehler, D.G., Worm, B., Harley, S.J., Doherty, P.A., B. capriscus constitutes the bulk of biomass among the 2003. Collapse and conservation of shark populations in the northwest triggerfishes. This fish is amphiatlantic. Interestingly, it was Atlantic. Science 299, 389–392. reported to proliferate in West African waters in the 1970s Caddy, J.F., Carocci, F., Coppola, S., 1998. Have peak fishery production levels as a consequence of heavy exploitation by trawlers in the been passed in continental shelf areas? Some perspectives arising from his- 1960s (Cavariviere,` 1982; Koranteng, 2001). Longhurst and torical trends in production per shelf area. J. Northwest Atl. Fish. Sci. 23, 191–219. Pauly (1987) list some features that may render B. capriscus Casey, J.M., Myers, R.A., 1998. Near extinction of a large widely distributed (synonym Balistes carolinensis) a winner under disruption of fish. Science 281, 690–692. demersal communities due to trawling: feeding of plankton in Cavariviere,` A., 1982. Les balistes des cotesˆ africaines (Balistes carolinen- midwater at night, construction of nests and parental care, resis- sis). Biologie, proliferation et posibilites´ dˇıexplotation. Oceanol. Acta 5, tance to desiccation due to its leathery skin and a tendency 453–460. Criales-Hernandez, M., Duarte, L.O., Garc´ıa, C.B., Manjarres,´ L., 2006. Ecosys- of being rapidly discarded (before other fish) due to its dan- tem impacts of the introduction of bycatch reduction devices in a tropical gerous dorsal spine. Weather a combination of above features shrimp trawl fishery: insights through simulation. Fish. Res. 77, 333–342. explains the biomass trajectory of triggerfishes in our waters Efron, B., 1987. Better bootstrap confidence intervals. J. Am. Stat. Assoc. 82, is open to question. Nevertheless, the coincidence is remark- 79–89. able. Garc´ıa, C.B., Solano, O., 1995. Tarpon atlanticus in Colombia: a big fish in trouble. Naga ICLARM Q. 18 (3), 47–49. Parrish (1995) postulated that small, “ugly”, fishes would sur- Garc´ıa, C.B., Duarte, L.O., Schiller, D., 1998. Demersal fish assemblages of the vive better in present times of industrial fishing. We see some Gulf of Salamanca Colombia (southern Caribbean Sea). Mar. Ecol. Prog. of his predictions becoming reality. The future of the bottom Ser. 174, 13–25. fishery both industrial and artisanal in the upwelling ecosys- Hutchings, J.A., 2000. Collapse and recovery of marine fishes. Nature 406, tem off Colombia, Caribbean Sea, looks compromised. Clearly 882–885. Hutchings, J.A., 2004. Marine fish population collapses: consequences for recov- radical management decisions must be taken including tem- Author's personalery and extinction risk. BioScience copy 54 (4), 297–309. poral and spatial closures (National Research Council, 2001), INPA, 1999. Instituto Nacional de Pesca y Acuicultura, Bolet´ın estad´ıstico substantial reduction of fishing effort, substantial reduction of pesquero 1997–1998, INPA, Bogota.´ bycatch due to shrimp trawling (Kennelly and Broadhurst, 2002; INPA, 2001. Instituto Nacional de Pesca y Acuicultura, Bolet´ın estad´ıstico Criales-Hernandez et al., 2006) and other regulations, in order pesquero colombiano 1999–2000, INPA, Bogota.´ Jackson, J.B.C., 2001. What was natural in the coastal oceans? PNAS 98 (10), to reverse trends of destruction of demersal fish biomass. Once 5411–5418. this is done and in the light of the findings here, there is evidence Jackson, J.B.C., Sala, E., 2001. Unnatural oceans. Sci. Mar. 65 (Suppl. 2), that recovery may proceed at good speed. 273–281. C.B. Garc´ıa et al. / Fisheries Research 85 (2007) 68–73 73

Jennings, S., Kaiser, M.J., 1998. The effects of fishing on marine ecosystems. yendo condiciones oceanograficas.´ Parte III: Crucero INPA-VECEP/ Adv. Mar. Biol. 34, 201–352. UE/DEMER/9604 (abril de 1996). Rev. Intropica. 2 (1), 117–149. Kennelly, S.J., Broadhurst, M.K., 2002. By-catch begone: changes in the phi- Mora, J.H., 1988. Analisis´ de la Pesca de Camaron´ de Aguas Someras (Penaeus losophy of fishing technology. Fish Fish 3, 340–355. f. notialis y Penaeus brasiliensis) Efectuada por la Flota Camaronera del Koranteng, K., 2001. Structure and dynamics of demersal assemblages on the Caribe Colombiano. Technical Report, INDERENA, Cartagena. continental shelf and upper slope off Ghana, West Africa. Mar. Ecol. Prog. Myers, R.A., Worm, B., 2003. Rapid worldwide depletion of predatory fish Ser. 220, 1–12. communities. Nature 423, 280–283. Longhurst, A.R., Pauly, D., 1987. Ecology of Tropical Oceans. Academic Press, National Research Council, 2001. Marine Protected Areas: Tools for Sustaining London. Ocean Ecosystems. National Academy Press, Washington, DC. Manjarres,´ L., Garc´ıa, C.B., Acero, A., 2001. Caracterizacion´ ecologica´ de las Orrell, T.M., 2002. Centropomidae. In: Carpenter, K.E. (ed.), The Living Marine asociaciones de peces demersales del Caribe colombiano norte con enfasis´ Resources of the Western Central Atlantic, vol. 2. Bony Fishes Part 1. FAO en pargos. Bol. Inst. Invest. Mar. 30, 77–107. Species Identification Guide for Fisheries Purposes and Americas Society Manjarres,´ L., 2002. Evaluacion´ de las Pesquer´ıas Demersales del Area´ Norte of Ichthyologist and Herpetologist Special Publication No. 5. FAO, Rome, del Caribe Colombiano y Parametros´ Ecologicos,´ Biologico-Pesqueros´ y pp. 1286–1293. Poblacionales del Recurso Pargo. Technical Report Code 3135-09-550-98, Parrish, R.H., 1995. Laternfish heaven: the future of world fisheries? Naga COLCIENCIAS, Santa Marta. ICLARM Q. 18 (3), 7–9. Manjarres,´ L., Arevalo,´ J.C., Rodr´ıguez, D.J., Gomez-Canchong,´ P., 2004a. Pauly, D., Alder, J., Bennett, E., Christensen, V., Tyedmers, P., Watson, R., 2003. Dinamica´ poblacional y manejo de stock de pargo rayado (Lutjanus syna- The future for fisheries. Science 302, 1359–1361. gris Linneaus, 1758) de La Guajira (Caribe colombiano). In: Manjarres,´ Pitcher, T., 1998. A cover story: fisheries may drive stocks to extinction. Rev. L. (Ed.), Pesquer´ıas demersales de area´ norte del Mar Caribe de Colom- Fish Biol. Fish. 8, 367–370. bia y parametros´ biologico-pesqueros´ y poblacionales del recurso pargos. Quintero, R., 1992. Proyecto Reconocimiento Pesquero Mediante Prospeccion´ Universidad del Magdalena, Santa Marta, pp. 267–295. Acustica´ en Areas´ Mar´ıtimas Colombianas. Fase I. Ministerio de Defensa Manjarres,´ L., Arevalo, J.C., Rodr´ıguez, D.J., Gomez-Canchong,´ P., 2004b. Nacional, Armada Nacional. Technical Report, CIOH, Cartagena. Mortalidad y estrategias de manejo del stock de pargo palmero (Lutjanus Stobutzki, I.C., Miller, M.J., Heales, D.S., Brewer, D.T., 2002. Sustainability of analis Cuvier, 1828) del area´ norte del Caribe Colombiano. In: Manjarres,´ elasmobranches caught as bycatch in a tropical prawn (shrimp) trawl fishery. L. (Ed.), Pesquer´ıas demersales de area´ norte del Mar Caribe de Colom- Fish. Bull. 100, 800–821. bia y parametros´ biologico-pesqueros´ y poblacionales del recurso pargos. Strømme, T., Saetersdal, G., 1989. Prospecciones de los Recursos Pesqueros de Universidad del Magdalena, Santa Marta, pp. 297–317. las Areas´ de la Plataforma Entre Surinam y Colombia 1988. IMR, Bergen. Manjarres,´ L., Vergara, A., Torres, J., Rodr´ıguez, G., Arteaga, E., Viana,˜ J., Testaverde, S.A., R´ıos, C.E., 1972. Collection of fish from fish cruises of the R/V Arevalo,´ J., Galvis, R., 2005a. Evaluacion´ de peces demersales e ictioplanc- Choco´ along the Caribbean Coast of Colombia during 1970. Proyecto para ton en el Mar Caribe de Colombia, incluyendo condiciones oceanograficas.´ el desarrollo de la pesca mar´ıtima en Colombia. PNUD, FAO, INDERENA. Parte I: Crucero INPA-VECEP/UE/DEMER/9507 (julio de 1995). Rev. Bol. Inf. 2 (4), 63–110. Intropica. 2 (1), 51–86. Watson, R., Pauly, D., 2001. Systematic distortions in world fisheries catch Manjarres,´ L., Rodr´ıguez, G., Torres, J., Vergara, A., Arteaga, E., Arevalo,´ J., trends. Nature 414, 534–536. Galvis, R., Viana,˜ J., 2005b. Evaluacion´ de peces demersales e ictioplancton Winger, P.D.,He, P., Walsh, S.J., 1999. Swimming endurance of American plaice en el Mar Caribe de Colombia, incluyendo condiciones oceanograficas.´ Parte (Hippoglossoides paltessoides) and its role in fish capture. ICES J. Mar. Sci. II: Crucero INPA-VECEP/UE/DEMER/9510 (octubre–noviembre de 1995). 56 (3), 253–265. Rev. Intropica. 2 (1), 87–115. Zuniga,˜ H., Escobar, M., 1993. Analisis´ de la Operatividad del Equipo de Pesca Manjarres,´ L., Rodr´ıguez, G., Vergara, A., Torres, J., Arteaga, E., Arevalo,´ y Estimacion´ de Abundancia Relativa de Demersales. Zona norte de Caribe J., Viana,˜ J., Galvis, R., Rodr´ıguez, D.Y., Barros, M., 2005c. Evaluacion´ colombiano. Informe complementario del crucero INPA/DEMER/9210. de peces demersales e ictioplancton en el Mar Caribe de Colombia, inclu- Technical Report, INPA/Universidad del Magdalena, Santa Marta.

Author's personal copy