Lindholm et al . : Ecological effects of bottom trawling on fish habitat along the central California outer continental shelf 
95 
assemblages on a cobble bottom ecosystem on West- 
ern Bank (northwest Atlantic). Mar. Ecol. Prog. Ser. 
306:63-78. doi: 10.3354/meps306063. 
Hiddink, J. G., S. Jennings, M. J. Kaiser, A. M. Queiros, D. E. 
Duplisea, and G. J. Piet. 
2006. Cumulative impacts of seabed trawl disturbance 
on benthic biomass, production, 
and species richness in different habitats. Can. J. Fish. 
Aquat. Sci. 63:721-736. doi: 10.1139/f05-266 
Hiddink, J. G., S. Jennings, and M. J. Kaiser. 
2007. Assessing and predicting the relative ecologi- 
cal impacts of disturbance on habitats with differ- 
ent sensitivities. J. Appl. Ecol. 44:405-413. doi: 
0.1111/j. 1365-2664. 2007. 01274.x. 
Hilborn, R., I. J. Stewart, T. A. Branch, and O. P. Jensen. 
2012. Defining trade-offs among conservation, profit- 
ability, and food security in the California Current 
bottom-trawl fishery. Conserv. Biol. 26:257-266. doi: 
10.1111/j. 1523- 1739. 20 11. 01800.x. 
Hixon, M. A., and B. N. Tissot. 
2007. Comparison of trawled vs. untrawled mud seafloor 
assemblages of fishes and macroinvertebrates at Co- 
quille Bank, Oregon. J. Exp. Mar. Biol. Ecol. 344:23- 
34. doi: 10.1016/j.jembe.2006.12.026. 
Hourigan, T. F. 
2009. Managing fishery impacts on deep-water coral 
ecosystems of the USA: emerging best practices. Mar. 
Ecol. Prog. Ser. 397:333-340. doi: 10.3354/meps08278. 
Hurlbert, S. H. 
1984. Pseudoreplication and the design of ecological 
field experiments. Ecol. Monogr. 54:187-211. doi: 
10.2307/1942661. 
Jackson, J. B. C., M. X. Kirby, W. H. Berger, K. A. Bjorndal, 
L. W. Botsford, B. J. Bourque, R. H. Bradbury, R. Cooke, J. 
Erlandson, J. A. Estes, T. P. Hughes, S. Kidwell, C. B. Lange, 
H. S. Lenihan, J. M. Pandolfi, C. H. Peterson, R. S. Steneck, 
M. J. Tegner, and R. R. Warner. 
2001. Historical overfishing and the recent collapse 
of coastal ecosystems. Science 293:629-637. doi: 
10. 1126/science. 1059199. 
Kaiser, M. J., and B. E. Spencer. 
1994. Fish scavenging behavior in recently trawled ar- 
eas. Mar. Ecol. Prog. Ser. 112:41-49. 
Kaiser, M. J., J. S. Collie, S. J. Hall, S. Jennings, and I. R. 
Poiner. 
2002. Modification of marine habitats by trawling activi- 
ties: prognosis and solutions. Fish Fish. 3:114-136. doi: 
10. 1046/j. 1467-2979.2002. 00079.x. 
Kaiser, M. J., D. B. Edwards, P. J. Armstrong, K. Radford, N. 
E. L. Lough, R. P. Flatt, and H. D. Jones. 
1998. Changes in megafaunal benthic communities in 
different habitats after trawling disturbance. ICES J. 
Mar. Sci. 55:353-361. doi: 10.1006/jmsc.l997.0322. 
Kaiser, M. J., K. Ramsay, C. A. Richardson, F. E. Spence, and 
A. R. Brand. 
2000. Chronic fishing disturbance has changed shelf sea 
benthic community structure. J. Anim. Ecol. 69:494- 
503. doi: 10. 1046/j. 1365-2656. 2000. 00412.x. 
Kitaguchi, B. D. 
2011. Benthic community structure in trawled and un- 
trawled areas of the outer continental shelf. M.S. the- 
sis, 114 p. San Jose State Univ., San Jose, CA. 
Koslow, J. A., K. Gowlett-Holmes, J. K. Lowry, T. O’Hara, G. C. 
B. Poore, and A. Williams. 
2001. Seamount benthic macrofauna off southern Tas- 
mania: community structure and impacts of trawl- 
ing. Mar. Ecol. Prog. Ser. 213:111-125. doi: 10.3354/ 
meps2131 11. 
Lindholm, J. B., P. J. Auster, and L. S. Kaufman. 
1999. Habitat-mediated survivorship of juvenile (0- 
year) Atlantic cod Gadus morhua. Mar. Ecol. Prog. Ser. 
180:247-255. doi: 10.3354/mepsl80247. 
Lindholm, J. B., P. J. Auster, M. Ruth, and L. Kaufman. 
2001. Modeling the effects of fishing and impli- 
cations for the design of marine protected ar- 
eas: juvenile fish responses to variations in sea- 
floor habitat. Conserv. Biol. 15:424-437. doi: 
10. 1046/j. 1523-1739. 2001. 015002424.x. 
Lindholm, J., P. Auster, and P. Valentine. 
2004. Role of a large marine protected area for conserv- 
ing landscape attributes of sand habitats on Georges 
Bank (NW Atlantic). Mar. Ecol. Prog. Ser. 269:61-68. 
doi: 10.3354/meps269061. 
Lindholm, J., M. Kelly, D. Kline, and J. de Marignac. 
2009. Patterns in the local distribution of the sea whip 
(Halipteris willemoesi), in an area impacted by mo- 
bile fishing gear. Mar. Technol. Soc. J. 42:64-68. doi: 
10.4031/002533208787157651. 
Lipsey, M. 
1990. Design sensitivity: statistical power for experi- 
mental research, 207 p. Sage Publications, Newbury 
Park, CA. 
Lohrer, A. M., S. F. Thrush, and M. M. Gibbs. 
2004. Bioturbators enhance ecosystem function 
through complex biogeochemical interactions. Nature 
431:1092-1095. doi: 10.1038/nature03042. 
Machias, A., V. Vassilopoulou, D. Vatsos, P. Bekas, A. Kalliani- 
otis, C. Papaconstantinou, and N. Tsimenides. 
2001. Bottom trawl discards in the northeastern Medi- 
terranean Sea. Fish. Res. 53:181-195. doi: 10.1016/ 
SO 165-7836(00)00298-8. 
Malatesta, R. J., and P. J. Auster. 
1999. The importance of habitat features in low-relief 
continental shelf environments. Oceanol. Acta 22:623- 
626. doi: 10. 10 16/S0399- 1784(00)88953-7. 
Malecha, P. W., and R. P. Stone. 
2009. Response of the sea whip Halipterus willomoesi 
to simulated trawl disturbance and its vulnerability to 
subsequent predation. Mar. Ecol. Prog. Ser. 388:197- 
206. doi: 10.3354/meps08145. 
Mason, J., R. Kosaka, A. Mamula, and C. Speir. 
2012. Effort changes around a marine reserve: the case 
of the California Rockfish Conservation Area. Mar. Pol- 
icy 36:1054-1063. doi: 10. 1016/j.marpol. 2012. 03.002. 
McConnaughey, R. A., K. L. Mier, and C. B. Drew. 
2000. An examination of chronic trawling effects on 
soft-bottom benthos of the eastern Bering Sea. ICES 
J. Mar. Sci. 57:1377-1388. doi: 10.1006/jmsc.2000.0906. 
Meysman, F. J. R., J. J. Middelburg, and C. H. R. Heip. 
2006. Bioturbation: a fresh look at Darwin’s last 
idea. Trends Ecol. Evol. 21:688-695. doi: 10.1016/j. 
tree. 2006. 08. 002. 
Morrisey, D. J., L. Howitt, A. J. Underwood, and J. S. Stark. 
1992. Spatial variation in soft-sediment benthos. Mar. 
Ecol. Prog. Ser. 81:197-204. 
NRC (National Research Council). 
1999. Sustaining marine fisheries, 164 p. National 
Academies Press, Washington, D.C. 
2002. Effects of trawling and dredging on seafloor habi- 
tat, 126 p. National Academy Press, Washington, D.C. 
