Ferm et al.: Foraging habits and dietary overlap of Limanda aspera and Lepidopsetta polyxystra in the Bering Sea 11 
and 3 anonymous reviewers provided valuable comments 
on this manuscript. 
Literature cited 
Abookire, A. A., and C. S. Rose. 
2005. Modifications to a plumb staff beam trawl for sampling 
uneven, complex habitats. Fish. Res. 71:247—254. 
Barnes, C. L., A. H. Beaudreau, and R. N. Yamada. 
2021. The role of size in trophic niche separation between two 
groundfish predators in Alaskan waters. Mar. Coast. Fish. 
13:69-84. 
Beverton, R. J. H. 
1995. Spatial limitation of population size; the concentration 
hypothesis. Neth. J. Sea Res. 34:1-6. 
Brown, S. C., J. J. Bizzarro, G. M. Cailliet, and D. A. Ebert. 
2012. Breaking with tradition: redefining measures for 
diet description with a case study of the Aleutian skate 
Bathyraja aleutica (Gilbert 1896). Environ. Biol. Fishes 
95:3-20. 
Carter, C. G., D. J. Grove, and D. M. Carter. 
1991. Trophic resource partitioning between two coexisting 
flatfish species off the north coast of Anglesey, North Wales. 
Neth. J. Sea Res. 27:325-335. 
Chao, A., N. J. Gotelli, T. C. Hsieh, E. L. Sande, K. H. Ma, 
R. K. Colwell, and A. M. Ellison. 
2014. Rarefaction and extrapolation with Hill numbers: 
a framework for sampling and estimation in species diver- 
sity studies. Ecol. Monogr. 84:45-67. 
Corcobado Onate, F. 
1991. Food and daily ration of the rock sole Lepidopsetta 
bilineata (Pleuronectidae) in the eastern Bering Sea. Mar. 
Biol. 108:185-191. 
Fissel, B., M. Dalton, B. Garber-Yonts, A. Hayne, S. Kasperski, 
J. Lee, D. Lew, C. Seung, K. Sparks, M. Szymkowiak et al. 
2021. Stock assessment and fishery evaluation report for the 
groundfish fisheries of the Gulf of Alaska and Bering Sea/ 
Aleutian Islands area: economic status of the groundfish 
fisheries off Alaska, 2019, 271 p. Alaska Fish. Sci. Cent., 
Natl. Mar. Fish. Serv., Seattle, WA, [Available from website.] 
Gibson, R. N. 
2005. The behavior of flatfishes. In Flatfishes: biology and 
exploitation (R. N. Gibson, ed.), p. 218-239. Blackwell 
Science, Oxford, UK. 
Holladay, B. A., and B. L. Norcross. 
1995. Diet diversity as a mechanism for partitioning nurs- 
ery grounds of pleuronectids. In Proceedings of the inter- 
national symposium on North Pacific flatfish. Alaska Sea 
Grant College Program Rep. AK-SG-95-04, p. 177-203. 
University of Alaska, Fairbanks, AK. 
Hsieh, T. C., K. H. Ma, and A. Chao. 
2016. iNEXT: an R package for rarefaction and extrapolation 
of species diversity (Hill numbers). Methods Ecol. Evol. 
7:1451-1456. 
2020. iNEXT: interpolation and extrapolation for species 
diversity. R package, vers. 2.0.20. [Available from website, 
accessed June 2021.] 
Hurst, T. P. 
2016. Shallow-water habitat use by Bering Sea flatfishes along 
the central Alaska Peninsula. J. Sea Res. 111:37—45. 
Hurst, T. P., and T. A. Duffy. 
2005. Activity patterns in northern rock sole are mediated 
by temperature and feeding history. J. Exp. Mar. Biol. Ecol. 
325:201-213. 
Hurst, T. P., C. H. Ryer, J. M. Ramsey, and S. A. Haines. 
2007. Divergent foraging strategies of three co-occurring 
north Pacific flatfishes. Mar. Biol. 151:1087—1098. 
Lang, G. M., P. A. Livingston, and B.S. Miller. 
1995. Food habits of three congeneric flatfishes: yellowfin sole 
(Pleuronectes asper), rock sole (P. bilineatus), and Alaska 
plaice (P. quadrituberculatus) in the eastern Bering Sea. 
In Proceedings of the international symposium on North 
Pacific flatfish. Alaska Sea Grant College Program Rep. AK- 
SG-95-04, p. 225-245. University of Alaska, Fairbanks, AK. 
Lemke, J. L., and C. H. Ryer. 
2006. Risk sensitivity in three juvenile (age-0) flatfish spe- 
cies: does estuarine dependence promote risk-prone behav- 
ior? J. Exp. Mar. Biol. Ecol. 333:172—180. 
Linehan, J. E., R.S Gregory, and D. C. Schneider. 
2001. Predation risk of age-0 cod (Gadus) relative to depth 
and substrate in coastal waters. J. Exp. Mar. Biol. Ecol. 
263:25-44. 
Livingston, M. E. 
1987. Morphological and sensory specializations of five New 
Zealand flatfish species, in relation to feeding behavior. 
J. Fish Biol. 31:775—-795. 
Mariani, S., C. Boggan, and D. Balata. 
2011. Food resource use in sympatric juvenile plaice and floun- 
der in estuarine habitats. Mar. Ecol. 32:96—-101. 
Matta, M. E., B. A. Black, and T. K. Wilderbuer. 
2010. Climate-driven synchrony in otolith growth-increment 
chronologies for three Bering Sea flatfish species. Mar. 
Ecol. Prog. Ser. 413:137-145. 
Mecklenburg, C. W., T. A. Mecklenburg, and L. K. Thorsteinson. 
2002. Fishes of Alaska, 1116 p. Am. Fish. Soc., Bethesda, MD. 
Mees, J., and M. B. Jones. 
1997. The hyperbenthos. Oceangr. Mar. Biol. 35:221—255. 
Nash, R. D. M., and A. J. Geffen. 
2012. Mortality through the early life-history of fish: what 
can we learn from European plaice (Pleuronectes platessa 
L.)? J. Mar. Syst. 93:58-68. 
Neutel, A. M., J. A. P. Heesterbeek, and P. C. de Ruiter. 
2002. Stability in real food webs: weak links in long loops. 
Science 296:1120-1123. 
Paterson, A. W., and A. K. Whitfield. 
2000. Do shallow-water habitats function as a refugia for juve- 
nile fishes? Estuar. Coast. Shelf Sci. 51:359-364. 
Poiesz, S. S. H., A. van Leeuwen, K. Soetaert, J. I. J. Witte., 
D.S. C., Zaat, and H. W. van der Veer. 
2020. Is summer growth reduction related to feeding guild? 
A test for a benthic juvenile flatfish sole (Solea solea) in a 
temperate coastal area, the western Wadden Sea. Estuar. 
Coast. Shelf Sci. 235:106570. 
R Core Team. 
2021. R: a language and environment for statistical com- 
puting. R Foundation for Statistical Computing, Vienna, 
Austria. [Available from website, accessed March 2021.] 
Ross, S. T. 
1986. Resource partitioning in fish assemblages: a review of 
field studies. Copeia 1986:352-388. 
Russo, T,, D. Pulcini, A. O’Leary, S. Cataudella, and S. Mariani. 
2008. Relationship between body shape and trophic niche 
segregation in two closely related sympatric fishes. J. Fish 
Biol. 73:809-828. 
Schabetsberger, R., R. Brodeur, L. Ciannelli, J. M. Napp, and 
G. L. Swartzman. 
2000. Diel vertical migration and interaction of zooplankton 
and juvenile walleye pollock (Theragra chalcogramma) at 
a frontal region near the Pribilof Islands, Bering Sea. ICES 
J. Mar. Sci. 57:1283-1295. 
