Temporal and Spatial Patterns in the Abundance of Jellyfish in the Northern Benguela Upwelling Ecosystem and their Link to Thwarted Pelagic Fishery Recovery

African Journal of Marine Science

Volumn 34, Issue 1, 2012, Pages 131-146

  Flynn, B A ^a   Richardson, A J ^b,c   Brierley, A S ^d   Boyer, D C ^e,f   Axelsen, B E ^g   Scott, L ^h   Moroff, N E ^e   Kainge, P I ^e   Tjizoo, B M ^e   Gibbons, M J ^a  

^a UNIVERSITY OF THE WESTERN CAPE   (South Africa)

^b CSIRO   (Australia)

^c UNIVERSITY OF QUEENSLAND   (Australia)

^d UNIVERSITY OF ST ANDREWS   (United Kingdom)

^e MINISTRY OF FISHERIES AND MARINE RESOURCES   (Namibia)

^f Orchard Farm   (United Kingdom)

^g INSTITUTE OF MARINE RESEARCH   (Norway)

^h ASCLME House   (South Africa)

Author keywords

Aequorea; Chrysaora; fish recruitment; jellyfish joyride; overfishing

Indexed keywords

ABUNDANCE; DEMERSAL FISH; JELLYFISH; LITERATURE REVIEW; OCEANOGRAPHY; OVERFISHING; PELAGIC FISHERY; PREDATION; RECRUITMENT (POPULATION DYNAMICS); SPATIAL VARIATION; SPAWNING; TEMPORAL ANALYSIS; TRAWLING; UPWELLING; ZOOPLANKTON;

ATLANTIC OCEAN; BENGUELA CURRENT; NAMIBIA;

AEQUOREA; CHRYSAORA; SCYPHOZOA;

EID: 84862193406     PISSN: 1814232X     EISSN: 18142338     Source Type: Journal    
DOI: 10.2989/1814232X.2012.675122     Document Type: Review

References (124)

1. Albert, DJ. 2005. Reproduction and longevity of Aurelia aurita in Roscoe Bay, a small bay on the Pacific coast of Canada. Journal of the Marine Biological Association, 85: 575-581.
2. Arai, MN. 1997. A functional biology of Scyphozoa, London: Chapman and Hall.
3. Bailey, KM and Batty, RS. 1983. Laboratory study of predation by Aurelia aurita on larval herring (Clupea harengus): experimental observations compared with model predictions. Marine Biology, 72: 295-301.
4. Bakun, A and Weeks, SJ. 2006. Adverse feedback sequences in exploited marine systems: are deliberate interruptive actions warranted?. Fish and Fisheries, 7: 316-333.
5. Bakun, A and Weeks, SJ. 2008. The marine ecosystem off Peru: what are the secrets of its fishery productivity and what might its future hold?. Progress in Oceanography, 79: 290-299.
6. Barange M, Boyd AJ. 1992. Life history, circulation and maintenance of Nyctiphanes capensis (Euphausiacea) in the northern Benguela upwelling system. In: Payne AIL, Brink KH, Mann KH, Hilborn R (eds), Benguela trophic functioning. South African Journal of Marine Science 12: 95-106.
7. Bartholomae, CH and van der Plas, AK. 2007. Towards the development of environmental indices of the Namibian shelf, with particular reference to fisheries management. African Journal of Marine Science, 29: 25-35.
8. Boyd AJ, Salat J, Masó M. 1987. The seasonal intrusion of relatively saline water on the shelf off northern and central Namibia. In: Payne AIL, Gulland JA, Brink KH (eds), The Benguela and comparable ecosystems. South African Journal of Marine Science 5: 107-120.
9. Boyer DC, Boyer HJ, Fossen I, Kreiner A. 2001. Changes in abundance of the northern Benguela sardine stock during the decade 1990-2000, with comments on the relative importance of fish and the environment. In: Payne AIL, Pillar SC, Crawford RJM (eds), A decade of Namibian fisheries science. South African Journal of Marine Science 23: 67-84.
10. Boyer DC, Hampton I. 2001. An overview of the living marine resources of Namibia. In: Payne AIL, Pillar SC, Crawford RJM (eds), A decade of Namibian fisheries science. South African Journal of Marine Science 23: 5-35.
11. Boyer, DC, Cole, J and Bartholomae, C. 2000. Southwestern Africa: northern Benguela Current region. Marine Pollution Bulletin, 41: 123-140.
12. Brewer, RH and Feingold, JS. 1991. The effect of temperature on the benthic stages of Cyanea (Cnidaria: Scyphozoa), and their seasonal distribution in the Niantic River estuary, Connecticut. Journal of Experimental Marine Biology and Ecology, 152: 49-60.
13. Brierley, AS, Axelsen, BE, Boyer, DC, Lynam, CP, Didcock, CA, Boyer, HJ, Sparks, CAJ, Purcell, JE and Gibbons, MJ. 2004. Single-target echo detections of jellyfish. ICES Journal of Marine Science, 61: 383-393.
14. Brierley, AS, Axelsen, BE, Buecher, E, Sparks, C, Boyer, H and Gibbons, MJ. 2001. Acoustic observations of jellyfish in the Namibian Benguela. Marine Ecology Progress Series, 210: 55-66.
15. Brierley, AS, Boyer, DC, Axelsen, BE, Lynam, CP, Sparks, CAJ, Boyer, H and Gibbons, MJ. 2005. Towards the acoustic estimation of jellyfish abundance. Marine Ecology Progress Series, 295: 105-111.
16. Brodeur, RD. 1998. In situ observations of the association between juvenile fishes and scyphomedusae in the Bering Sea. Marine Ecology Progress Series, 163: 11-20.
17. Brodeur, RD, Suchman, CL, Reese, DC, Miller, TW and Daly, EA. 2008. Spatial overlap and trophic interactions between pelagic fish and large jellyfish in the northern California Current. Marine Biology, 154: 649-659.
18. Brodeur, RD, Sugisaki, H and Hunt, GL. 2002. Increases in jellyfish biomass in the Bering Sea: implications for the ecosystem. Marine Ecology Progress Series, 233: 89-103.
19. Brotz L, Cheung WWL, Kleisner K, Pakhomov E, Pauly D. In press. Increasing jellyfish populations: trends in large marine ecosystems. Hydrobiologia.
20. Brown, PC, Painting, SJ and Cochrane, KL. 1991. Estimates of phytoplankton and bacterial biomass and production in the northern and southern Benguela ecosystems. South African Journal of Marine Science, 11: 537-564.
21. Buecher, E, Sparks, C, Brierley, A, Boyer, H and Gibbons, MJ. 2001. Biometry and size distribution of Chrysaora hysoscella (Cnidaria, Scyphozoa) and Aequorea aequorea (Cnidaria, Hydrozoa) off Namibia, with some notes on their parasite Hyperia medusarum. Journal of Plankton Research, 23: 1073-1080.
22. Butterworth, DS. 1983. "Assessment and management of pelagic stocks in the southern Benguela region". In Proceedings of the expert consultation to examine changes in abundance and species composition of neritic fish resources, San José, Costa Rica, April 1983, Edited by: Sharp, GD and Csirke, J. 329-405. San José, Costa Rica: Food and Agriculture Organization.
23. Chavez, FP, Ryan, JP, Lluch-Cota, SE and Niquen, M. 2003. From anchovies to sardines and back: multidecadal change in the Pacific Ocean. Science, 299: 217-221.
24. Colombo, GA, Mianzan, H and Madirolas, A. 2003. Acoustic characterization of gelatinous-plankton aggregations: four case studies from the Argentine continental shelf. ICES Journal of Marine Science, 60: 650-657.
25. Cram, DL and Visser, GA. 1973. SWA pilchard stock shows first signs of recovery (summary of results of Phase III of the Cape Cross programme). South African Shipping News and Fishing Industry Review, 28: 56-63.
26. Crawford, RJM, Shannon, LV and Pollock, DE. 1987. The Benguela ecosystem. Part IV. The major fish and invertebrate resources. Oceanography and Marine Biology: an Annual Review, 25: 353-505.
27. Cury, P, Bakun, A, Crawford, RJM, Jarre, A, Quiñones, RA, Shannon, LJ and Verheye, HM. 2000. Small pelagics in upwelling systems: patterns of interaction and structural changes in ''wasp-waist'' ecosystems. ICES Journal of Marine Science, 57: 603-618.
28. Cury, P and Shannon, L. 2004. Regime shifts in upwelling ecosystems: observed changes and possible mechanisms in the northern and southern Benguela. Progress in Oceanography, 60: 223-243.
29. di Camillo, CG, Betti, F, Bo, M, Martinelli, M, Puce, S and Bavestrello, G. 2010. Constribution to the understanding of the seasonal cycle of Aurelia aurita (Cnidaria: Scyphozoa) scyphopolyps in the northern Adriatic Sea. Journal of the Marine Biological Association of the United Kingdom, 90: 1105-1110.
30. Doyle, TK, de Haas, H, Cotton, D, Dorschel, B, Cummins, V, Houghton, JDR and Hays, GC. 2008. Widespread occurrence of the jellyfish Pelagia noctiluca in Irish coastal and shelf waters. Journal of Plankton Research, 30: 963-968.
31. Elmhirts, R. 1925. Lunar periodicity in Obelia. Nature, 116: 358-359.
32. Engås, A, Skeide, R and West, CW. 1997. The 'Multisampler': a system for remotely opening and closing multiple codends on a sampling trawl. Fisheries Research, 29: 295-298.
33. Estrada M, Marrasé C. 1987. Phytoplankton biomass and productivity off the Namibian coast. In: Payne AIL, Gulland JA, Brink KH (eds), The Benguela and comparable ecosystems. South African Journal of Marine Science 5: 347-356.
34. Fearon, JJ, Boyd, AJ and Schülein, FH. 1992. Views on the biomass and distribution of Chrysaora hysoscella (Linné, 1766) and Aequorea aequorea (Forskål, 1775) off Namibia, 1982-1989. Scientia Marina, 56: 75-85.
35. Feigenbaum, DL and Kelly, M. 1984. Changes in the lower Chesapeake Bay food chain in the presence of the sea nettle Chrysaora quinquecirrha (Scyphomedusae). Marine Ecology Progress Series, 19: 39-47.
36. Flynn, BA and Gibbons, MJ. 2007. A note on the diet and feeding of Chrysaora hysoscella in Walvis Bay Lagoon, Namibia, during September 2003. African Journal of Marine Science, 29: 303-307.
37. Gammelsrød T, Bartholomae CH, Boyer DC, Fillipe VLL, O'Toole MJ. 1998. Intrusion of warm surface water along the Angolan-Namibian coast in February-March 1995: the 1995 Benguela Niño. In: Pillar SC, Moloney CL, Payne AIL, Shillington FA (eds), Benguela dynamics: impacts of variability on shelf-sea environments and their living resources. South African Journal of Marine Science 19: 41-56.
38. Gibbons, MJ. 1993. Vertical migration and feeding of Euphausia lucens at two 72 h stations in the southern Benguela upwelling region. Marine Biology, 116: 257-268.
39. Gliwicz, MZ. 1986. Predation and the evolution of vertical migration in zooplankton. Nature, 320: 746-748.
40. Goy, J. 1973. Gonionemus suavensis: structural characters, developmental stages and ecology. Publications of the Seto Marine Biological Laboratory, 20: 525-536.
41. Graham, WM and Bayha, KM. 2007. "Biological invasions by marine jellyfish". In Ecological studies, Edited by: Nentwig, W. 239-256. Berlin: Springer-Verlag.
42. Graham, WM, Pagès, F and Hamner, WM. 2001. A physical context for gelatinous zooplankton aggregations: a review. Hydrobiologia, 451: 199-212.
43. Gröndahl, F. 1988. Interactions between polyps of Aurelia aurita and planktonic larvae of scyphozoans: an experimental study. Marine Ecology Progress Series, 45: 87-93.
44. Hart, TJ and Currie, RI. 1960. The Benguela Current. Discovery Reports, 31: 123-298.
45. Hastie, T and Tibshirani, R. 1990. Generalized additive models, London: Chapman and Hall.
46. Hays, GC, Bastian, T, Doyle, TK, Fossette, S, Gleiss, AC, Gravenor, MB, Hobson, VJ, Humphries, NE, Lilley, MKS, Pade, NG and Sims, DW. 2011. High activity and Lévy searches: jellyfish can search the water column like fish. Proceedings of the Royal Society B [London], 279: 465-473.
47. Hays, GC, Doyle, TK, Houghton, JDR, Lilley, MKS, Metcalf, JD and Righton, D. 2008. Diving behaviour of jellyfish equipped with electronic tags. Journal of Plankton Research, 30: 325-331.
48. Heymans, JJ, Shannon, LJ and Jarre, A. 2004. Changes in the northern Benguela ecosystem over three decades: 1970's, 1980's and 1990's. Ecological Modeling, 172: 175-195.
49. Hutchings, L, Pillar, SC and Verheye, HM. 1991. Estimates of standing stock, production and consumption of meso-and macrozooplankton in the Benguela ecosystem. South African Journal of Marine Science, 11: 499-512.
50. Il'inskii EN, Zavolokin AV. 2007. The qualitative composition and autumn-winter distribution of scyphomedusae (Scyphozoa) in the Sea of Okhotsk epipelagial. Zoologicheskii Zhurnal 86: 1168-1176 (translated from Russian).
51. Kaartvedt, S, Klevjer, TA, Torgersen, T, Sørnes, TA and Røstad, A. 2007. Diel vertical migration of individual jellyfish (Periphylla periphylla). Limnology and Oceanography, 52: 975-983.
52. Kainge, P, Kjesbu, OS, Thorsen, A and Salvanes, AG. 2007. Merluccius capensis spawn in Namibian waters, but do M. paradoxus?. African Journal of Marine Science, 29: 379-392.
53. King DPF, O'Toole MJ. 1973. A preliminary report on the findings of the South West African pelagic egg and larval surveys. SFRI Internal Report, Cape Cross Progress Phase III. Sea Fisheries Research Institute, Cape Town.
54. Kramp, PL. 1957. Hydromedusae of the Discovery Collections. Discovery Report, 29: 1-128.
55. Kramp, PL. 1959. The Hydromedusae of the Atlantic Ocean and adjacent waters. Dana Report, 46: 1-283.
56. Kreiner, A, Yemane, D, Stenevik, EK and Moroff, NE. 2011. The selection of spawning location of sardine (Sardinops sagax) in the northern Benguela after changes in stock structure and environmental conditions. Fisheries Oceanography, 20: 560-569.
57. Lampert, W. 1989. The adaptive significance of diel vertical migration of zooplankton. Functional Ecology, 3: 21-27.
58. Le Clus, F, Melo, YC, Cooper, RM and Malan, PE. 1987. The multispecies pelagic fishery off Namibia 1982-1986 and stock assessments for pilchard and anchovy. Collection of Scientific Papers of the International Commission for Southeast Atlantic Fisheries, 14: 7-25.
59. Leloup, E. 1934. Siphonophores Calycophorides de l'Océan Atlantique tropical et austral, Bulletin du Musée Royal d. Histoire Naturelle de Belgique, 10: 1-87.
60. Liu, WC, Lo, WT, Purcell, JE and Chang, HH. 2009. Effects of temperature and light intensity on asexual reproduction of the scyphozoan, Aurelia aurita (L.) in Taiwan. Hydrobiologia, 616: 247-258.
61. Lucas, CH. 2001. Reproduction and life history strategies of the common jellyfish, Aurelia aurita, in relation to its ambient environment. Hydrobiologia, 451: 229-246.
62. Lynam, CP, Attrill, MJ and Skogen, MD. 2010. Climatic and oceanic influences on the abundance of gelatinous zooplankton in the North Sea. Journal of the Marine Biological Association of the United Kingdom, 90: 1153-1159.
63. Lynam, CP, Gibbons, MJ, Axelsen, BA, Sparks, CAJ, Coetzee, J, Heywood, BG and Brierley, AS. 2006. Jellyfish overtake fish in a heavily fished ecosystem. Current Biology, 16: 492-493.
64. Lynam, CP, Heath, MR, Hay, SJ and Brierley, AS. 2005. Evidence for impacts by jellyfish on North Sea herring recruitment. Marine Ecology Progress Series, 298: 157-167.
65. Mills, CE. 1993. Natural mortality in NE Pacific coastal Hydromedusae: grazing predation, wound healing and senescence. Bulletin of Marine Science, 53: 194-203.
66. Mills, CE. 2001. Jellyfish blooms: are populations increasing globally in response to changing ocean conditions?. Hydrobiologia, 451: 55-68.
67. Mohrholz, V, Schmidt, M, Lutjeharms, JRE and John, HC. 2004. Space-time behavior of Angola-Benguela Frontal Zone during the Benguela Niño of April 1999. International Journal of Remote Sensing, 25: 1337-1340.
68. Möller, H. 1984. Reduction of a larval herring population by jellyfish predator. Sciences, 224: 621-622.
69. Moser, F. 1925. Die Siphonophoren der Deutschen Südpolar Expedition 1901-1903. Deutschen Südpolar Expedition, 17: 1-541.
70. Neethling S. 2010. Re-descriptions of some southern African Scyphozoa: out with the old and in with the new. MSc thesis, University of the Western Cape, South Africa.
71. Nogueira, M Jnr, Nagata, RM and Haddad, MA. 2010. Seasonal variation of macromedusae (Cnidaria) at North Bay, Florianópolis, southern Brazil. Zoologia, 27: 377-386.
72. Oguz, T, Fach, B and Salihoglu, B. 2008. Invasion dynamics of the alien ctenophore Mnemiopsis leidyi and its impact on anchovy collapse in the Black Sea. Journal of Plankton Research, 30: 1385-1397.
73. Olivar, MP and Barange, M. 1990. Zooplankton of the northern Benguela region in a quiescent upwelling period. Journal of Plankton Research, 12: 1023-1044.
74. Olivar, MP, Rubiés, P and Salat, J. 1988. Early life history and spawning of Merluccius capensis Castelnau in the northern Benguela Current. South African Journal of Marine Science, 6: 245-254.
75. Olivar, MP and Shelton, PA. 1993. Larval fish assemblages of the Benguela Current. Bulletin of Marine Science, 53: 450-474.
76. O'Toole MJ. 1977. Investigations into some important fish larvae in the South East Atlantic in relation to the hydrographical environment. PhD thesis, University of Cape Town, South Africa.
77. O'Toole, MJ. 1978. Aspects of the early life history of the hake Merluccius capensis Castelnau off South West Africa. Fisheries Bulletin South Africa, 10: 20-36.
78. Pagès, F and Gili, JM. 1992. Influence of the thermocline on the vertical migration of medusa during a 48 h sampling period. South African Journal of Zoology, 27: 50-59.
79. Pagès, F, Gili, JM and Bouillon, J. 1992. Planktonic cnidarians of the Benguela Current. Scientia Marina, 56: 1-144.
80. Pearre, S. 1973. Vertical migration and feeding in Sagitta elegans Verrill. Ecology, 54: 300-314.
81. Pearre, S. 1979. Problems of detection and interpretation of vertical migration. Journal of Plankton Research, 1: 29-44.
82. Pillar, SC, Armstrong, DA and Hutchings, L. 1989. Vertical migration, dispersal and transport of Euphausia lucens in the southern Benguela Current. Marine Ecology Progress Series, 53: 179-190.
83. Purcell, JE and Arai, MN. 2001. Interactions of pelagic cnidarians and ctenophores with fish: a review. Hydrobiologia, 451: 27-44.
84. Purcell, JE and Sturdevant, MV. 2001. Prey selection and dietary overlap among zooplanktivorous jellyfish and juvenile fishes in Prince William Sound, Alaska. Marine Ecology Progress Series, 210: 67-83.
85. Purcell, JE, Uye, SI and Lo, WT. 2007. Anthropogenic causes of jellyfish blooms and direct consequences for humans: a review. Marine Ecology Progress Series, 350: 153-174.
86. Richardson, AJ, Bakun, A, Hays, GC and Gibbons, MJ. 2009. The jellyfish joyride: causes, consequences and management responses to a more gelatinous future. Trends in Ecology & Evolution, 24: 312-322.
87. Roosen-Runge, EC. 1970. Life cycle of the hydromedusae Phialidium gregarium (A. Agassiz, 1862) in the laboratory. The Biological Bulletin, 139: 203-221.
88. Rouault, M, Illig, S, Bartholomae, C, Reason, CJC and Bentamy, A. 2007. Propagation and origin of warm anomalies in the Angola Benguela upwelling system in 2001. Journal of Marine Systems, 68: 473-488.
89. Sætersdal, G, Bianchi, G, Strømme, T and Venema, SC. 1999. The Dr Fridtjof Nansen Programme 1975-1993. Investigations of fishery resources in developing countries. History of the programme and review of results. FAO Fisheries Technical Paper, 391: 1-434.
90. Schülein, FH, Butterworth, DS and Cram, DL. 1978. An assessment of the southeast Atlantic pilchard population in ICSEAF divisions 1.4 and 1.5, 1953-1977. Collection of Scientific Papers of the International Commission for Southeast Atlantic Fisheries, 5: 35-44.
91. Schuyler, Q and Sullivan, BK. 1997. Light responses and diel migration of the scyphomedusa Chrysaora quinquecirrha in mesocosms. Journal of Plankton Research, 19: 1417-1428.
92. Sexton, MA, Hood, RR, Sarkodee-adoo, J and Liss, AM. 2010. Response of Chrysaora quinquecirrha medusa to low temperature. Hydrobiologia, 645: 125-133.
93. Shackleton LY. 1987. A comparative study of fossil fish scales from three upwelling regions. In: Payne AIL, Gulland JA, Brink KH (eds), The Benguela and comparable ecosystems. South African Journal of Marine Science 5: 79-84.
94. Shannon, LV. 1985. The Benguela ecosystem. I. Evolution of the Benguela, physical features and processes. Oceanography and Marine Biology: an Annual Review, 23: 105-182.
95. Shannon, LV, Boyd, AJ, Brundrit, GB and Taunton-Clarke, J. 1986. On the existence of an El Niño-type phenomena in the Benguela system. Journal of Marine Research, 44: 495-520.
96. Shannon, LV and Pillar, SC. 1986. The Benguela Ecosystem. Part III. Plankton. Oceanography and Marine Biology: an Annual Review, 24: 65-170.
97. Shoji, J, Mizuno, K-I, Yamamoto, M, Miller, TW, Hamaoka, H and Omori, K. 2009. Spatial distribution and dietary overlap between Japanese anchovy Engraulis japonicus and moon jellyfish Aurelia aurita in the Seto Inland Sea, Japan. Scientia Marina, 73S1: 191-198.
98. Sommer, U, Stibor, H, Katechakis, A, Sommer, F and Hansen, T. 2002. Pelagic food web configurations at different levels of nutrient richness and their implications for the ratio fish production: primary production. Hydrobiologia, 484: 11-20.
99. Sparks, C, Brierley, AS, Buecher, E, Boyer, D, Axelsen, BE and Gibbons, MJ. 2005. Submersible observations on the daytime vertical distribution of Aequorea forskalea off the west coast of South Africa. Journal of the Marine Biological Association of the United Kingdom, 85: 519-522.
100. Sparks, C, Buecher, E, Brierley, AS, Boyer, H, Axelsen, BE and Gibbons, MJ. 2001. Observations on the distribution and relative abundance of the scyphomedusan Chrysaora hysoscella (Linné, 1766) and the hydrozoan Aequorea aequorea (Forskål, 1775) in the northern Benguela ecosystem. Hydrobiologia, 451: 275-286.
101. Stander, GH and De Decker, AHB. 1969. Some physical and biological aspects of an oceanographic anomaly off South West Africa in 1963, Investigational Report, Division of Sea Fisheries. South Africa, 81: 1-46.
102. Stenevik EK, Sundby S, Cloete R. 2001. Influence of buoyancy and vertical distribution of sardine Sardinops sagax eggs and larvae on their transport in the northern Benguela ecosystem. In: Payne AIL, Pillar SC, Crawford RJM (eds), A decade of Namibian fisheries science. South African Journal of Marine Science 23: 85-97.
103. Stiasny, G. 1934. Scyphomedusae. Discovery Report, 8: 329-396.
104. Stiasny, G. 1939. ber Dactylometra fulgida (Reynaud) von der Walfischbai. Zoologischer Anzeiger, 126: 172-185.
105. Strømme T, Iilende T. 2001. Precision in systematic trawl surveys as assessed from replicate sampling by parallel trawling off Namibia. In: Payne AIL, Pillar SC, Crawford RJM (eds), A decade of Namibian fisheries science. South African Journal of Marine Science 23: 385-396.
106. Suchman, CL and Brodeur, RD. 2005. Abundance and distribution of large medusa in surface waters of the northern California Current. Deep-Sea Research II, 52: 51-72.
107. Suchman CL, Brodeur RD, Emmert RL, Daly EA In press. Large medusa in surface waters of the northern California Current: variability in relation to environmental conditions. Hydrobiologia.
108. Sundby S, Boyd AJ, Hutchings L, O'Toole MJ, Thorisson K, Thorsen A. 2001. Interaction between Cape hake spawning and the circulation in the northern Benguela upwelling ecosystem. In: Payne AIL, Pillar SC, Crawford RJM (eds), A decade of Namibian fisheries science. South African Journal of Marine Science 23: 317-336.
109. Thomas, RM. 1986. The Namibian pilchard: the 1985 season, assessment for 1952-1985 and recommendations for 1986. Collection of Scientific Papers of the International Commission for Southeast Atlantic Fisheries, 13: 243-269.
110. Unterüberbacher, HK. 1964. Zooplankton studies in the waters off Walvis Bay with special reference to the Copepoda, Investigational Reports of the Marine Research laboratory. South West Africa, 11: 1-42.
111. Utne-Palm, AC, Salvanes, AGV, Currie, B, Kaartvedt, S, Nilsson, GE, Baithwaite, V, Stecyk, JAW, Hundt, M, Flynn, B, van der Bank, M, Peard, KR, Lunde, IG, Sandvik, GK, Klevjer, TA, Pittman, K, Sweetman, AK, Strandabø, RAU and Gibbons, MJ. 2010. Trophic structure and community stability in an overfished ecosystem. Science, 329: 333-336.
112. Valdemarsen, JW and Misund, OA. 1994. "Trawl designs and techniques used by Norwegian research vessels to sample fish in the pelagic zone". In Proceedings of the sixth PRlW-PEW0 symposium, Edited by: Hylen, A. 135-144. Bergen: Institute of Marine Research.
113. van der Bank, MG, Utne-Palm, AC, Pittman, K, Sweetman, AK, Richoux, NB, Brüchert, V and Gibbons, MJ. 2011. Dietary success of a 'new' key fish in an overfished ecosystem: evidence from fatty acid and stable isotope signatures. Marine Ecology Progress Series, 428: 219-233.
114. van der Lingen, CD, Shannon, LJ, Cury, P, Kreiner, A, Moloney, CL, Roux, J-P and Vaz-Velho, F. 2006. "Resource and ecosystem variability, including regime shifts, in the Benguela Current system". In Benguela: predicting a large marine ecosystem, Edited by: Shannon, V, Hempel, G, Malanotte-Rizzoli, P, Moloney, C and Woods, J. 147-184. Amsterdam: Elsevier.
115. van der Westhuizen A. 2001. A decade of exploitation and management of the Namibian hake stocks. In: Payne AIL, Pillar SC, Crawford RJM (eds), A decade of Namibian fisheries science. South African Journal of Marine Science 23: 307-315.
116. Vanhöffen, E. 1902a. Die acraspeden medusen der Deutschen Tiefsee-Expedition 1898-1899. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer "Valdivia", 3: 1-52.
117. Vanhöffen, E. 1902b. Die craspeden medusen der Deutschen Tiefsee-Expedition 1898-1899. 1. Trachymedusen. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer "Valdivia", 3: 55-86.
118. Vanhöffen, E. 1908. Die Lucernariden und Skyphomedusen der Deutschen Südpolar Expedition 1901-1903. Deutschen Südpolar Expedition, 10: 25-59.
119. Vanhöffen, E. 1911. Die Anthoathecata und Leptothecata der Deutschen Tiefsee Expedition 1898-1899. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer "Valdivia", 19: 193-233.
120. Vanhöffen, E. 1912. Die craspedoten Medusen der Deutschen Südpolar Expedition 1901-1903. Deutschen Südpolar Expedition, 13: 351-396.
121. Venter, GE. 1988. Occurrence of jellyfish on the west coast off South West Africa/Namibia. Report of the South African National Science Programmes, 157: 56-61.
122. Verheye, HM, Hutchings, L and Peterson, WT. 1991. Life history and population maintenance strategies of Calanoides carinatus (Copepoda: Calanoida) in the southern Benguela ecosystem. South African Journal of Marine Science, 11: 179-191.
123. Widmer, CL. 2004. The hydroid and early medusa stages of Mitrocomella cellularia (Hydrozoa, Mitrocomidae). Marine Biology, 145: 315-321.
124. Wood, SN. 2006. Generalized additive models: an introduction, London: Chapman & Hall/CRC Press.