NOAA Technical Memorandum NMFS-AFSC-159


Comparison of Size Selectivity 

Between Marine Mammals and 

Commercial Fisheries with 

Recommendations for Restructuring 

Management Policies 

by 

M. A. Etnier and C. W. Fowler

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration 

National Marine Fisheries Service 

Alaska Fisheries Science Center 

October 2005 
NOAA Technical Memorandum NMFS 

The National Marine Fisheries Service's Alaska Fisheries Science Center 

uses the NOAA Technical Memorandum series to issue informal scientific and 

technical publications when complete formal review and editorial processing 

are not appropriate or feasible. Documents within this series reflect sound 

professional work and may be referenced in the formal scientific and technical 

literature. 

The NMFS-AFSC Technical Memorandum series of the Alaska Fisheries 

Science Center continues the NMFS-F/NWC series established in 1970 by the 

Northwest Fisheries Center. The NMFS-NWFSC series is currently used by 

the Northwest Fisheries Science Center. 

This document should be cited as follows: 

Etnier, M. A., and C. W. Fowler. 2005. Comparison of size selectivity 

between marine mammals and commercial fisheries with 

recommendations for restructuring management policies. U.S. Dep. 

Commer., NOAA Tech. Memo. NMFS-AFSC-159, 274 p. 

Reference in this document to trade names does not imply endorsement by 

the National Marine Fisheries Service, NOAA. 
NOAA Technical Memorandum NMFS-AFSC-159


Comparison of Size Selectivity 

Between Marine Mammals and 

Commercial Fisheries with 

Recommendations for Restructuring 

Management Policies 

by 

M. A. Etnier and C. W. Fowler

Alaska Fisheries Science Center 

7600 Sand Point Way N.E. 

Seattle, WA 98115 

www.afsc.noaa.gov 

U.S. DEPARTMENT OF COMMERCE

Carlos M. Gutierrez, Secretary 

National Oceanic and Atmospheric Administration 

Vice Admiral Conrad C. Lautenbacher, Jr., U.S. Navy (ret.), Under Secretary and Administrator 

National Marine Fisheries Service 

William T. Hogarth, Assistant Administrator for Fisheries 

October 2005 
This document is available to the public through: 

National Technical Information Service 

U.S. Department of Commerce 

5285 Port Royal Road 

Springfield, VA 22161 

www.ntis.gov 
ABSTRACT 

Conventional fisheries management schemes often maintain harvest practices quite different 

from those of natural predators, explicitly or implicitly stressing the importance of avoiding strategies 

similar to those of other species. While this may minimize some of the perceived direct interactions 

between fisheries and marine mammals, such management often exerts strong abnormal influences, 

both on the target species and the ecosystems of which they are a part. It is impossible to exhaustively 

account for the indirect effects of such practices. In contrast, in Systemic Management, it is argued 

that the patterns of predation exhibited by marine mammals, structured by natural selection over 

thousands of generations, are evidence of sustainable resource use in the very long term, thus 

accounting for such complexity. As such, these patterns can be used to guide the ways in which 

humans extract resources from the environment. 

In this study, we compiled prey size data from the food habits literature for 63 marine mammal 

species as a guide for sustainable harvest practices. Our results show that the overwhelming tendency 

of marine mammals is to target prey smaller than 30 cm in length. This pattern of selectivity applies 

regardless of the maximum attainable size of either the predator or the prey species. In contrast, 

commercial fisheries tend to select individuals larger than 30 cm when possible. Thus, the size 

selectivity of the commercial catch of small-bodied prey species fits within the norm of the patterns 

exhibited by marine mammals, but the commercial catch of large-bodied prey species does not. We 

conclude that, in order to minimize the negative effects of abnormal selective pressures that 

commercial fisheries exert directly on prey stocks and indirectly on the ecosystems of which they are a 

part, the targeted size composition of larger prey species should be reduced to more closely mimic the 

patterns exhibited by marine mammals. 

iii 

CONTENTS

Introduction......................................................................................................................................1 

Materials and Methods.....................................................................................................................4 

Caveats....................................................................................................................................5 

Mechanics of obtaining data ...................................................................................................6

Summary statistics for food habits data ..................................................................................7 

Minimum sample size requirements.......................................................................................8 

Comparison with commercial data .........................................................................................9 

Comparison with survey data................................................................................................10 

Maximum reported size ........................................................................................................11

Predator size..........................................................................................................................12 

Number of non-overlapping taxa ..........................................................................................12 

Results, patterns of marine mammal food habits...........................................................................13 

Accuracy of estimating values from figures.........................................................................14 

Size composition of prey consumed, sorted by predator species..........................................20 

Otariid seals .................................................................................................................22 

Phocid seals..................................................................................................................35 

Odontocete whales.......................................................................................................43 

Size composition of prey consumed, sorted by prey species................................................60 

Crustacean prey species...............................................................................................63 

Cephalopod prey species..............................................................................................65 

Fish prey species..........................................................................................................69 

Overall patterns in the size of prey targeted by marine mammals........................................90 

 v 
Evaluation of biases..............................................................................................................90 

Relationship between predator size and prey size ................................................................97 

Relationship between mean size of prey consumed by marine mammals and maximum 

reported prey size..................................................................................................................99 

Results, comparisons with commercial catch data ......................................................................113 

Species-specific patterns of size composition.....................................................................119 

Relationship between mean size of commercial catch and maximum reported size..........129 

Overlap in size distributions ...............................................................................................137

Results, comparisons with survey data and evaluations of selectivity ........................................146 

Evaluation of appropriateness of survey data .....................................................................153 

Potential Applications..................................................................................................................162 

Discussion and Conclusions ........................................................................................................166 

Recommendations for restructuring management policies.................................................167 

Acknowledgements......................................................................................................................174 

Citations.......................................................................................................................................175 

Appendix 1: Summary food habits data, sorted by predator species..........................................213 

Appendix 2: Raw food habits data, sorted by predator species..................................................215 

Appendix 3: Summary food habits data, sorted by prey species................................................236 

Appendix 4: Raw food habits data, sorted by prey species ........................................................247 

Appendix 5: Pairings of food habits and commercial data .........................................................268 

Appendix 6: Pairings of food habits and survey data .................................................................270 

Appendix 7: Key to citations referenced in Appendices 5 and 6................................................273

 vi 
INTRODUCTION 

If we look at managers as those responsible for regulating human impact on other 

systems (e.g., other species, communities, or ecosystems), one of their challenges is that of 

finding a way to incorporate all the impacts humans have into the decision-making process. 

Commercial fisheries have a variety of ecological effects. Commercial fisheries have been 

known to reduce stock biomasses to fractions of their virgin levels (Myers and Worm 2003), and 

cause reductions in mean age, age-at-maturity (Jørgensen 1990), and mean and maximum size 

(Wysokiński 1984). Coincident with these changes, commercial catches shift to lower trophic 

levels (Pauly et al. 1998a). The cumulative effect is that many of the world’s fish stocks are in 

serious jeopardy (Garcia and Newton 1997, Kock 1985, Pauly et al. 2002, Rosenberg 2003). 

Perhaps one of the more serious kinds of ecological effects of commercial fishing is the 

directional selection and/or genetic change caused by selective harvests—harvest practices which 

preferentially target a particular demographic of a resource population while preferentially 

excluding another (Conover 2000, Conover and Munch 2002, Law 2000, Olsen et al. 2004). 

This is particularly problematic for resource species whose various populations are used as 

sources of food and fiber for human consumption. Every species (including humans, Orians 

1990, NRC 1996) has genetic impacts on the species with which it interacts. These impacts 

occur through processes lumped under the term "coevolution" (Thompson 1982, Futuyma and 

Slatkin 1983, Stenseth and Maynard Smith 1984). What is an appropriate or acceptable 

evolutionary influence on another species and what is excessive? How can managers incorporate 

knowledge of the genetic effects of our actions in establishing the size of harvests, establish 

guidelines for selectivity, and account for things like age, size, and sex in the process? How can 

 
this be done so that not only the selectivity is accounted for, but so that the indirect effects on 

other species and ecosystems are also taken into account?

We propose a form of management based on directly relevant, completely consonant 

guiding information that produces recommendations for changing commercial harvest practices 

in such a way that long-term sustainability is an option. Because genetic effects are clearly 

related to selective mortality, it follows that one part of the challenge is that of addressing the 

issue of selectivity. Selectivity, as such, is also too vague and general to be easily tackled in 

specific management action, owing to the fact that selectivity can relate to a wide variety of 

attributes of a species (e.g., sex, age, size, location, growth patterns, and maturation rates). 

Whatever we do in research to supply management advice, it must involve a measurable aspect 

of selectivity and it must be a component of fishing practices that can be managed. 

In this paper, we address the specific issue of size selectivity as one of many facets of 

fisheries management that must be dealt with so as to provide quantitative guidance to managers. 

Commercial fishing, as currently practiced in many of the world’s oceans, is not sustainable in 

the long-term (Fowler 2003). In contrast, predator-prey relationships in marine ecosystems have 

evolved over thousands of generations and persist over evolutionary time scales. These systems 

exhibit emergent properties that reflect the effects of a wide range of variables at various spatial 

and temporal scales. One of the emergent properties is the appropriate, sustainable, or workable 

size composition for catches of any particular prey species. This is one of the many aspects of 

harvest practices that must be addressed as part of any form of management intended to apply to 

the management of our impacts on either individual resource species or ecosystems. 

To explore this issue, we examine the size composition, measured as the length of 

individual prey items (rather than weight or volume, which would also provide valuable 

 2 
information), in the diets of predators that feed on various species of fish, squid, and to a lesser 

extent, crustaceans. We draw upon information presented in the literature from a variety of 

studies of food habits for marine mammals. This is done in light of the potential that the patterns 

of what nonhuman species do, when considered collectively, are evidence of sustainable 

behaviors following the tenets of management reviewed by Fowler (2003). When viewed in this 

way, anything that deviates from these normative patterns exerts abnormal effects on the system 

(as discussed in Fowler 2003, Fowler and Hobbs 2002). Depending on the degree to which the 

abnormal pattern deviates from the norm, the effects will be more or less problematic. 

Within this framework, our objective is to characterize the size composition of prey items 

within the diets of nonhuman mammalian predators as guidance for fisheries managers. 

Regarding the science of this paper, our working null hypothesis is that humans are not 

statistically significantly different from other predatory species in regard to the size composition 

of their takes (i.e., commercial fishing) in comparison to the size composition of the takes of 

other predatory species (in parallel with other non-human/human comparisons: Fowler and 

Hobbs 2003). Thus, we intend to use existing data on the size composition of marine mammal 

dietary preferences to draw conclusions on what is sustainable to then provide quantitative 

guidance to fisheries managers. This approach will help to avoid any abnormal effects on any 

particular resource species or the ecosystems of which both humans as well as the other species 

are parts. 

 3 
MATERIALS AND METHODS 

The literature regarding the food habits of marine mammals is substantial (see Pauly et al. 

1998b), and has increased exponentially over the past two decades (Fig. 1). For the purposes of 

this study, we restricted our analysis to studies that reported size composition of marine mammal 

prey based on hard or soft tissues recovered from scats, stomachs, or regurgitations. We have 

focused on those studies that report prey lengths, rather than weights or volumes, and we have 

excluded from analysis those food habits studies that quantified only prey species composition 

(i.e., that lack information on size composition for specific prey species). The information in this 

report was obtained through examination of peer-reviewed articles, “gray” literature 

(unpublished progress reports, Ph.D. dissertations, conference proceedings, etc.), and personal 

communications when data representations necessitated access to the raw data (see below). 

0

5

10

15

20

25

30

1950 1960 1970 1980 1990 2000

Year

Frequency of publications

Figure 1.--Number of marine mammal food habits studies published up to 2002, by year (based 

on citations analyzed in Pauly et al. 1998b). 

 4 
Caveats 

The data analyzed here consist of size estimates of prey items identified primarily to the 

species level, with a few cases identified to the genus- or family-level. In almost all the cases 

presented here, prey size (specifically, prey length) has been estimated based on correlations 

relating body size to the size of those tissue types that preserve well in the digestive systems of

marine mammals (e.g., cephalopod beaks, fish otoliths, and bones; Fitch and Brownell 1968). 

This approach to estimating the size composition of marine mammal diets is known to have 

several biases (Bigg and Faucett 1985, Dellinger and Trillmich 1988, Pierce and Boyle 1991). 

Specifically, a host of measures can be under-represented: seasonal variability in the diet, 

relative importance of small-bodied taxa and small-bodied individuals (Reid and Arnould 1996), 

relative importance of large-bodied individuals (Kiyota et al. 1999, Gudmundson et al. in press). 

In addition, body size can be underestimated due to partial or complete digestion of characteristic 

skeletal elements (Bowen 2000, Tollit et al. 2004a, 2004b; Zeppelin et al. 2004). 

Finally, there is some suggestion that estimating body size based on calibration with growth 

curves (e.g., inverse regression) will systematically bias against values in the tails of the 

distribution (the so-called “regression towards the mean”; see Konigsberg et al. 1997). Despite 

the biases inherent in food habits studies, the use of regression to calibrate body size against 

element size is the standard approach in the literature and provides the basis for the 

overwhelming majority of the available data. 

One of the idealized goals of this research is to characterize marine mammal food habits 

in the absence of bias due to anthropogenic perturbations to the ecosystem. However, it is 

almost always the case that food habits studies post-date the development of commercial 

harvests of the marine mammal species, their prey, or both in any particular ecosystem. Thus, 

 5 
food habits studies based on what marine mammals are eating now may not provide an unbiased 

view of “normal” behavior of the predators (what they would eat in the absence of abnormal 

human impacts - a point we will revisit our Discussion section). 

Finally, a third bias relates to our goal of providing spatially and temporally relevant

comparisons among food habits studies (size composition of prey in the diets of non-human 

predators), commercial catch data (size composition of harvests), and survey data (size 

composition of available prey resources). The commercial catch data are typically limited to 

target species, with size composition of bycatch only rarely reported. An unfortunate parallel is 

that trawl survey data are also heavily biased towards commercially valuable taxa. Thus, 

examples that enable comparisons among food habits studies, trawl survey data, and commercial 

catch data in the same season and area are relatively rare. 

Mechanics of Obtaining Data 

Prey size data are presented in a variety of formats in the literature we reviewed. Prey

size data were occasionally listed in the text or in tabular form. These were transcribed directly. 

Information on otolith or beak size was used only if the appropriate regression equations were 

available to allow conversion to prey body size. 

Other data were presented in graphic form (e.g., length frequency [LF] distributions, box 

plots), and presented more of a challenge. In these instances, figures were digitally scanned at a 

resolution of 300 dpi or higher, then digitized using Didger 3 software. This software allowed us 

to estimate the plotted values by calibrating the axes directly from the figure. We found that 

calibration of the vertical axis (e.g., frequency value) is typically more important than calibration 

 6 
of the horizontal axis, since the horizontal axis (e.g., prey size, in size bins ranging from 1 to 10 

cm) tends to be spaced at uniform intervals (though this was not always the case). 

When using graphic information, the format of the original figure exerts a strong 

influence on the accuracy with which data can be estimated. For our purposes, the format that 

resulted in the best estimates was two-dimensional histograms representing individual series of 

data. Histograms representing multiple series of data in two dimensions were somewhat more 

difficult, while any representation in three dimensions was extremely difficult or impossible to 

use. LF distributions represented as curves were arbitrarily estimated at discrete intervals along 

the curve, with the total values re-scaled such that they summed to 100%. 

Body size for crustaceans is listed as total length for shrimp and krill, carapace width for 

crabs, and carapace length for lobsters. Body size for cephalopods is listed as dorsal mantle 

length (DML). Conventions on the measure of body size used for fish (e.g., total length, fork 

length, standard length) tend to be species-specific. As a consequence, we have not consistently 

used one over the other, but have been consistent within species. Within the text of this 

document, we use the terms “length” and “prey size” interchangeably, regardless of the 

taxonomic group. 

Summary Statistics for Food Habits Data 

If the mean size and size range of prey species were not listed in the text of the food 

habits study, they were estimated from the LF distributions when available. Means were 

calculated by weighting each size interval by the appropriate scaling factor (either frequencies or

percents). It was not always possible to determine whether midpoints or end-points were used 

for the size intervals (i.e., the bin size used for bars plotted in LF distributions). Thus, estimates 

 7 
of means calculated this way can be in error by as much as ± 0.5 of the interval of each size class 

(e.g., Augustyn 1991, Torok 1994). In cases where prey size was presented in both tabular and 

graphical format, the error rate in estimating the mean from the LF distributions was calculated. 

In cases where the range of sizes preyed upon by marine mammals was estimated from 

the LF distributions, the possibility exists that low values in the tails of the distributions do not 

appear in graphic representations. Thus, these data represent minimum estimates of the range of 

prey sizes consumed. 

Digitized data were also used to estimate the shape parameters (standard deviation, skew, 

kurtosis) of the LF distribution. In cases where these statistics were presented in both tabular 

and graphic form, the accuracy of our estimation from the graphic data was evaluated. 

Minimum Sample Size Requirements

Although we chose to extract data in all cases where they represented prey size 

information, regardless of sample size (see Appendices 1-4), the overall analysis has been 

arbitrarily limited to those cases where mean prey size was based on samples of 10 or more 

individuals of any given prey taxon. In cases where we evaluate the central tendency of the 

distribution of mean prey sizes, all cases have been weighted equally. That is, we calculate the 

unweighted mean of means. 

The exclusion of cases with sample sizes less than 10 from the overall analysis may 

inadvertently bias against large-bodied prey items simply because fewer large-bodied prey items 

may be consumed in a foraging trip. To test the hypothesis that our minimum sample size 

requirement biases against large-bodied prey items, the LF distribution of single prey items was 

 8 
generated by combining data for sample sizes of one, two (if the size range was reported), or 

three (if the mean and range was reported). 

Comparison with Commercial Data 

As stated in the Introduction, the objective of the study is to characterize the patterns of 

prey size selectivity among marine mammals as a guide to what is sustainable over the long 

term. The claim that commercial fishing practices need to change to match the behavioral 

patterns of marine mammals first requires a demonstration that there are substantive differences 

between the two (see Fowler et al. 1999, Livingston 1993 regarding comparisons of total 

removals). 

Toward this end, data describing the size composition of commercial catches were 

compiled for comparison with the marine mammal food habits data to test the null hypothesis of 

no difference. This is not as straightforward as it might at first seem. An obvious first step is to 

eliminate from consideration all those taxa that are not commercially important

1

. Once this is 

done, however, careful attention must still be paid to obtaining commercial data that are 

comparable both spatially and temporally. Because many of the world’s fisheries actively 

attempt to minimize the degree of spatial overlap with marine mammals (Kaschner and Pauly 

2004), this is typically only feasible at a regional level. 

It is particularly difficult to obtain comparable data for prey taxa with highly variable 

year-class strength because a strong year class will “pulse” through the system for several 

subsequent years (cf. Castonguay and Mercille 1988, Livingston 1993, Overholtz and Waring 

1991, Reid et al. 1996). This will also vary in importance as a function of the periodicity of 

reproductive cycles. Annual breeders with a very highly constrained breeding period will have 

 

1

 This excludes, almost entirely, bycatch of non-target species. 

 9 
much more distinctive peaks pulsing through the LF distribution than taxa that have either a 

much shorter periodicity or a highly protracted breeding period. Prey taxa with short life cycles, 

like shrimp, squid, and some fish, provide serious complications to the standard approach of 

using age frequency or LF data to describe/monitor the population being exploited (Pauly 1985). 

This is because the annual (or seasonal) contribution of recruitment to the overall catch can be 

substantial. For seasonally/spatially/annually variable LF data, frequencies for each sampling 

period were first estimated (if possible), and then an overall average frequency calculated for 

each size interval. 

Comparison with Survey Data 

In addition to an interest in comparing the size composition of marine mammal prey with 

that of the commercial fishing industry, it is also important to compare both of these with what is 

available. This requires having survey data estimating age/size structure of stocks being 

exploited. Such data provide a means to evaluate the degree to which both marine mammals and 

commercial fisheries are selective in the size of prey they target, and then to compare this 

selectivity. Marine mammals have been characterized as “opportunistic” foragers based on 

comparisons of the prey species in their diets with those found where they forage (Kajimura 

1985), suggesting that they might simply consume any prey item that they encounter. However, 

several studies suggest that the decision-making process is much more elaborate (Sinclair et al. 

1994, Croxall and Pilcher 1984, Reid et al. 1996). Thus, survey data provide an index of what 

marine mammals (as well as the commercial fisheries) are taking relative to what is available and 

whether it relates to the choice of species consumed or to the size of prey items chosen. 

 10 
As with the commercial catch data, survey data tend to be fairly limited in scope (either 

temporally or spatially) relative to the foraging behavior of marine mammals. In an attempt to 

maximize the comparability of the data, survey data were used in this study only if they

corresponded to prey species from the same region and time span as the corresponding food 

habits study. Ideally, stock structure data derive from fisheries-independent surveys. However, 

stock structure data are occasionally inferred from analysis of commercial catch data. In no 

cases were acoustic survey data used for comparison with either the commercial catch data or the

food habits data. 

Maximum Reported Prey Size 

Another metric of interest to this study is the relationship between the targeted size and 

the maximum reported size of each prey species. For smaller-bodied prey species such as krill 

(Euphausia spp.), we might expect targeting of similar size classes by marine mammals and 

commercial fisheries alike. In contrast, specific size classes of prey species with larger overall 

body size such as Atlantic cod (Gadus morhua) may be differentially selected by either marine 

mammals or commercial fisheries. In most cases, maximum reported size was obtained from online searchable databases hosted by FishBase (Froese and Pauly 2003) and CephBase (Wood and 

Day 1998). In cases where the maximum reported size was either not listed in the databases, or 

was smaller than the maximum reported size from the food habits study, the upper end of the size 

range listed in the food habits studies was used2

. 

 

2

 Because the size estimates from the food habits literature are typically based on regressions, it may be the case that 

estimated maximum sizes fail to account for lack-of-fit for extreme values in the growth curves. 

 11 
Predator Size 

Finally, we are also interested in testing the hypothesis that there is a relationship 

between prey size targeted by marine mammals and the body size of the predator. Information 

relating gape size of the predators to body size of the prey may provide a more appropriate 

comparison (Nilsson and Brönmark 2000, Scharf et al. 2000), but those data are not readily 

available for all marine mammal taxa. Unless otherwise provided in the text of the relevant food 

habits study, predator size was obtained through internet sources such as the University of 

Michigan’s “Animal Diversity Web” (University of Michigan 1995) for the age and/or size 

classes of marine mammals included in the study (if known). 

Number of Non-overlapping Taxa 

In this report, we frequently use the terms “species” and “taxon” in ways that may seem 

synonymous. The two are not, however, strictly interchangeable. Whereas “species” refers to 

only one level of taxonomic specificity, “taxon” (or its plural, taxa) can include multiple 

taxonomic levels simultaneously. With particular reference to this report, identifications of 

predators and prey vary in taxonomic specificity from sub-species to family. 

Counts of how many predators and prey are represented in this study are based on 

numbers of non-overlapping taxa. To illustrate, a listing of Enoploteuthis anaspis and 

Enoploteuthis sp. is tallied as one taxon because of the possibility that the two are not mutually 

exclusive. In contrast, Gobiidae is listed as a prey item for Phocoena phocoena (Appendix 2), 

possibly representing multiple species. But because it is the only prey entry from that family, it 

was counted as a single taxon. In speciose genera like Sebastes spp., genus-level identifications 

 12 
`were counted multiple times only if they derived from different ocean basins (e.g., Northwest 

Atlantic versus Northeast Pacific). 

Using this protocol, we report the number of non-overlapping taxa of predators for which 

we have prey size data (regardless of sample size) and the distribution of prey size means (for 

sample sizes ≥ 10). We also report the total number of prey taxa for which we have size 

information deriving from marine mammal food habits studies. Taxonomy for marine mammals 

follows Rice (1998); taxonomy for crustaceans follows the European Register of Marine Species 

(Costello and Emblow 2004); taxonomy for cephalopods follows Wood and Day (1998); 

taxonomy for fish follows Froese and Pauly (2003). 

RESULTS: PATTERNS OF MARINE MAMMAL FOOD HABITS 

 

Our search of peer-reviewed articles and “gray” literature yielded 135 citations that 

specified prey size for 63 marine mammal taxa, resulting in 1,166 prey entries (Appendices 1-4). 

The breakdown, by predator type, is: 13 species of otariid seals, 15 species of phocid seals, 4 

taxa of mysticete whales, and 31 species of odontocete whales. From these citations, estimated 

prey size was obtained for: 17 species of crustaceans, (representing 12 families), 140 taxa of 

cephalopods (representing 28 families), and 223 taxa of fish (representing 71 families). 

This is decidedly not an exhaustive survey of the literature. Nevertheless, we feel 

confident that the large number of cases for which we have data ensure that the overall patterns 

are well documented in this report (with the exception, perhaps, of the part represented by 

mysticete whales). 

 13 
Accuracy of Estimating Values from Figures 

As discussed above, prey size data for marine mammals are often presented graphically, 

in a variety of different formats. The accuracy with which figures can be used to provide the 

data needed for our study is a function of sample size, resolution, and format of the original 

figure. Sample size dictates the minimum detectable difference between integer values on the 

histograms. For small (n ≤ 200) samples, the difference between one and two individuals (or 33 

versus 34) is easily detectable, with summed totals usually accurate to within ± 1%. Larger 

samples (n > 200) yield variable results, depending on the quality of the original image and the 

accuracy of the data/figure. 

Other problems encountered include unlisted sample sizes or unlabeled vertical axes. In 

these cases, it was still possible to estimate the shape of the distribution accurately (e.g., the 

relative contribution of each size bin), rescaling the values such that the total summed to 100%. 

In cases where the estimated values were in error by ≥ 1%, the whole data series was rescaled to 

100%. For instance, if the initial estimate of the summed total of the digitized LF distribution 

was 93%, each histogram bin was rescaled by dividing by 0.93. This is a systematic rescaling 

across all size bins; it cannot correct for random errors within a given size interval. Rather, 

random errors were detectable only through visual comparison of the estimated distribution and 

the original published distribution. 

Unless otherwise listed in the text of the original article, summary statistics (mean prey 

size, variance, skew, and kurtosis) were estimated from digitized LF distributions. The accuracy 

with which we could estimate mean prey size was quite high. Of the 121 cases where mean prey 

size was reported along with the LF distribution, our estimate of the mean was within ± 2 % of 

 14 
the reported mean 85% of the time (Fig. 2). In practical terms, this translates to 83% of the cases 

being estimated to within ± 0.50 cm (Fig. 3). 

0

5

10

15

20

25

30

35

40

45

50

-10 -8 -6 -4 -2 0 2 4 6 8 10

Percent difference between reported and estimated means

Frequenc

y

n = 121

Figure 2.--Distribution of errors in estimated mean prey size from LF distributions, as the percent 

difference between the reported and the estimated values. 

0

10

20

30

40

50

60

70

80

90

0.00 0.50 1.00 1.50 2.00 2.50 3.00 3.50 4.00

Absolute difference between reported and estimated means (cm)

Frequenc

y

n = 121

Figure 3.--Distribution of errors in estimated mean prey size determined from graphic LF 

distributions, as the absolute difference between the reported and estimated values. 

 15 
Our estimates of the variance of the LF distributions (typically reported as standard 

deviation, but occasionally reported as standard error) suffered a reduction in accuracy relative to 

the estimates of the mean. The bulk (81%) of estimated variances fell within ± 10% of reported 

values (Fig. 4). It is likely that this loss of accuracy is an artifact of the calculation for variance, 

often referred to as the second moment of the mean, because it involves summing the squared 

differences between observations and the (estimate of the) mean. Thus, any inaccuracies in 

estimating the mean will be amplified in calculations of the variance. 

0

5

10

15

20

25

30

-50 -40 -30 -20 -10 0 10 20 30 40 50

Percent difference between reported and estimated variance

Frequenc

y

n = 77

Figure 4.--Distribution of errors in estimated variance from LF distributions, as the percent 

difference between the reported and estimated values. 

 16 
With specific reference to the analysis presented here, two biases in reporting must be 

considered. The first is that prey size information is typically presented in terms of mean prey 

size. Less commonly, the range of size classes is presented or, less common still, the estimated 

size frequency distribution of the prey is presented in tabular or graphical form. In most cases 

where the size frequency distribution of the prey is presented, the distributions are decidedly 

asymmetric. Specifically, they are typically positively skewed, in which case mean prey size is 

higher than the modal, or most commonly encountered, prey size. 

In a series of 242 LF distributions, relatively few (22%) were significantly different from 

a normal distribution (Kolmogorov-Smirnov [K-S] test, P-values ≥ 0.050; Fig. 5). However, 

33% of the 242 distributions had skew values of 1.0 or higher, indicating that the mean is higher 

than the mode (Fig. 6). 

The tendency toward asymmetry in the LF distributions from food habits studies is 

perhaps more intuitively shown in the relationship between the mean and the midpoint of the size 

range for 504 sets of prey data. When the mean is subtracted from the midpoint of the range, the 

mean is within ± 2 cm of the midpoint of the range in 70% of the cases (Fig. 7). The mean is 2 

cm or more lower than the midpoint (resulting in a positive difference, indicative of positive 

skew) in 23% of the cases. It is 2 cm or greater than the midpoint (indicative of negative skew) 

in only 7% of the cases. There is an overall mean difference between the midpoint of the size 

ranges and their means of 1.21 cm. There is a clear but non-uniform tendency toward a positive 

skew. 

 

 17 
0

10

20

30

40

50

60

70

80

90

100

0.000 0.010 0.020 0.030 0.040 ≥ 0.050

P-value of K-S statistic

Frequency

n = 242

Figure 5.--Distribution of P-values for K-S statistic in 242 LF distributions for prey taxa in 

marine mammal food habits data, tested against a normal distribution. 

0

10

20

30

40

50

60

70

80

90

100

-3.0 -2.0 -1.0 0.0 1.0 2.0 3.0 4.0 5.0 6.0 7.0

Skew values

Frequenc

y

n = 242

Figure 6.--Distribution of skew values for 242 LF distributions (same sample as represented in 

Fig. 5). 

 18 
0

5

10

15

20

25

30

-10 -8 -6 -4 -2 0 2 4 6 8 ≥10

Midpoint of range minus mean

Percent of total

n = 504

mean difference = 1.21 cm

Figure 7.--Distribution of absolute differences between midpoint of range of values and the mean 

for 504 prey size distributions. 

In a few cases in the food habits literature, the authors document the shape of the 

distribution; in even fewer cases, median or modal sizes are used (Fea et al. 1999); and in even 

fewer cases still, the distributions are tested for normality (e.g., using a K-S statistic; e.g., 

Börjesson et al. 2003, McGarvey and Fowler 2002, Poulson et al. 2000). It is important to note 

that some LF distributions are distinctly bimodal, in which case the mean value may not be 

represented by a single individual.

 19 
Size Composition of Prey Consumed, Sorted by Predator Species 

As stated above, we have compiled prey size data for 63 marine mammal taxa (Appendix 

1, 2). However, many of the prey size data were based on sample sizes of less than 10 

individuals of a particular prey taxon and will not be included in the main analysis. Likewise, 

even with sample sizes of ≥ 10, many of the entries do not have mean prey size listed. For 

instance, of the 63 marine mammal taxa for which we have prey size data, seven of them lack 

data on mean prey size and/or the mean was based on a sample size of less than 10 prey items

(Fig. 8). At the other end of the spectrum, 20 marine mammal taxa have 10 or more entries of 

mean prey size, each of which was based on a sample size of ≥ 10 (Fig. 8; Appendix 1, 2). It is 

to the specifics of those 20 marine mammal taxa that we will now turn. 

8

5

7

3

0

5

2

4 3

20

7

0

5

10

15

20

25

0 1 2345 6789 ≥10

Number of entries per predator taxon with n ≥ 10

Frequenc

y

Figure 8.--Frequency distribution of the number of sets of data for prey from studies of marine 

mammal predator taxa where mean prey size data are available with sample sizes of 

10 or more prey items. Note that seven predator taxa had prey size data available, but 

did not list mean prey size and/or the mean was based on a sample size of less than 

10. 

 20 
For each of those 20 marine mammal taxa, we provide information on the number of taxa 

of different prey types (i.e., crustaceans, cephalopods, fish) identified in the literature, and the 

nature of the samples from which the food habits data derive. We also report the number of prey 

entries (n) that meet our sample size requirements and for which mean prey size is reported or 

could be estimated graphically. Finally, we also present the LF distribution of prey means and 

report the overall mean of means ( ̄x ) for those prey entries. All of these data except the type of 

samples utilized (scats versus stomach samples, etc.) are also provided in raw form in Appendix 

1. 

 21 
Otariid Seals 

Otariidae: Arctocephalus gazella, Antarctic fur seal; n = 99,  ̄x= 12.33 cm

Prey size information for the Antarctic fur seal was recorded for one crustacean species, 

11 cephalopod species, and 35 fish species, based on the citations listed in Table 1. There are 99

entries of mean prey size for A. gazella that meet our minimum sample size requirement, with an 

overall mean of means of 12.33 cm. The modal mean prey size for A. gazella is 5-10 cm, with a 

positively skewed distribution (Fig. 9). 

Table 1.--Sources of prey size data for Arctocephalus gazella. Cells with “n.a.” indicate cases 

 where the sample type is known but the sample size was not reported. 

Citation Scats Stomachs Regurge Lavage Age/sex

Casaux et al. 1998 51 non-breeding 

males

Cherel et al. 1997 22 

Croxall and Pilcher 1984 24 

Croxall et al. 1999 n.a. n.a. 

Daneri 1996 34 

Daneri and Carlini 1999 70 

Daneri and Coria 1993 105 

Daneri et al. 1999 412 

Doidge and Croxall 1985 238 adult females 

 22 
dos Santos and Haimovici 

2001 

1

Goldsworthy et al. 1997 138* 

Green et al. 1989 563 

Green et al. 1997 560 mostly males 

Kirkman et al. 2000 123 90 

Klages and Bester 1998 224 

Klages et al. 1999 93 breeding females, 

some juvenile males 

Lea et al. 2002 131 

McCafferty et al. 1998 436 

North 1996 55 

Reid 1995 ≥ 376 adult, sub-adult 

males

Reid and Arnould 1996 497 

Reid and Brierly 2001 n.a. 

Reid et al. 1996 16 adult females 

(lactating) 

*138 scat from an unspecified mix of A. tropicalis and A. gazella 

 23 
0

5

10

15

20

25

30

35

40

45

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequency

n = 99; mean of means = 12.33

Figure 9.--Frequency distribution of mean prey size for taxa eaten by Arctocephalus gazella. 

 24 
Otariidae: Arctocephalus pusillus doriferus, Australian fur seal; n = 11,  ̄x= 19.97 cm

Prey size information for the Australian fur seal was recorded for nine cephalopod 

species and five fish species, based on the two citations listed in Table 2. Note that these two 

publications are based on the same samples, but represent different analyses (cephalopods in one, 

fish in the other). Eleven entries of mean prey size for A. p. doriferus met our minimum sample

size requirement, with an overall mean of means of 19.97 cm. The modal mean prey size for A. 

p. doriferus is 15-20 cm, and the distribution is approximately normal (Fig. 10). 

Table 2.--Sources of prey size data for Arctocephalus pusillus doriferus

Citation Scats Stomachs Regurge Lavage Age/sex

Gales et al. 1993 317 15 40 

Gales and Pemberton 1994 317 40 

 25 
0

1

2

3

4

5

6

7

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 11; mean of means = 19.97

Figure 10.--Frequency distribution of mean prey size for taxa eaten by Arctocephalus pusillus 

 doriferus. 

 26 
Otariidae: Arctocephalus pusillus pusillus, South African fur seal; n = 49,  ̄x= 17.98 cm

Prey size information for the South African fur seal was recorded for one species of 

crustacean, 5 cephalopod species, and 23 taxa of fish, based on the three citations listed in Table 

3. There are 49 entries of mean prey size for A. p. pusillus that meet our minimum sample size 

requirement, with an overall mean of means of 17.98 cm. The modal mean prey size for A. p. 

pusillus is 10-20 cm, with a positively skewed distribution (Fig. 11).

Table 3.--Sources of prey size data for Arctocephalus pusillus pusillus. 

Citation Scats Stomachs Regurge Lavage Age/sex

Castley et al. 1991 49 

David 1987 997 

Lipinski and David 1990 2,195 1,522 adults; 

673 pups 

 27 
0

2

4

6

8

10

12

14

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 49; mean of means = 17.98

Figure 11.--Frequency distribution of mean prey size for taxa eaten by Arctocephalus pusillus 

 pusillus. 

 28 
Otariidae: Arctocephalus tropicalis sub-Antarctic fur seal; n = 12,  ̄x= 9.47 cm

Prey size information for the sub-Antarctic fur seal was recorded for two cephalopod 

species and nine species of fish based on the four citations listed in Table 4. Twelve entries of 

mean prey size for A. tropicalis met our minimum sample size requirement, with an overall 

mean of means of 9.47 cm. The modal mean prey size for A. tropicalis is 5-10 cm, and 

approximates a normal distribution (Fig. 12).

Table 4.--Sources of prey size data for Arctocephalus tropicalis. 

Citation Scats Stomachs Regurge Lavage Age/sex

Bester and Laycock 1985 220 

dos Santos and Haimovici 2001 8 

Goldsworthy et al. 1997 <138*

Klages and Bester 1998 245 

*138 scat from an unspecified mix of A. tropicalis and A. gazella

 29 
0

1

2

3

4

5

6

7

8

9

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 12; mean of means = 9.47

Figure 12.--Frequency distribution of mean prey size for taxa eaten by Arctocephalus tropicalis. 

 30 
Otariidae: Callorhinus ursinus, northern fur seal; n = 18,  ̄x= 12.90 cm

Prey size information for the northern fur seal was recorded for seven taxa of 

cephalopods and one species of fish based on the citations listed in Table 5. Eighteen entries of 

mean prey size for C. ursinus met our minimum sample size requirement, with an overall mean 

of means of 12.90 cm. The modal mean prey size for C. ursinus is 5-10 cm, with a positively 

skewed distribution (Fig. 13). 

Table 5.--Sources of prey size data for Callorhinus ursinus. Cells with “n.a.” indicate cases 

 where the sample type is known but the sample size was not reported. 

Citation Scats Stomachs Regurge Lavage Age/sex

Kiyota et al. 1999 107 6 

Lowry et al. 1989 n.a.a 

McAlister et al. 1976 n.a.b

Mori et al. 2001 89 

Sinclair et al. 1994 73 61 adult F; 12 juvenile M 

a Lowry et al. 1989 cite T. R. Loughlin, pers. com., as the source of data. 

b

 McAlister et al. 1976 obtained their sample during one year of the pelagic sampling program of 

the National Marine Fisheries Service, which took over 16,000 females between 1956 and 

1974. 

 31 
0

1

2

3

4

5

6

7

8

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 18; mean of means = 12.90

Figure 13.--Frequency distribution of mean prey size for taxa eaten by Callorhinus ursinus. 

 32 
Otariidae: Zalophus californianus, California sea lion; n = 12,  ̄x= 14.27 cm

Prey size information for California sea lions was recorded for one cephalopod species 

and four species of fish based on the citations listed in Table 6. Twelve entries of mean prey size 

for Z. californianus met our minimum sample size requirement, with an overall mean of means 

of 14.27 cm. The modal mean prey size for Z. californianus is 10-15 cm, with a slightly

positively skewed distribution (Fig. 14).

Table 6.--Sources of prey size data for Zalophus californianus. Cells with “n.a.” indicate cases 

 where the sample type is known but the sample size was not reported. 

Citation Scats Stomachs Regurge Lavage Age/sex

Antonelis et al. 1984 224 

Bailey and Ainley 1982 n.a. n.a. 

Lowry and Caretta 1999 6930 187 

Melin 2002 120 

Morejohn et al. 1978 34 n.a. 

 33 
0

1

2

3

4

5

6

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 12; mean of means = 14.27

Figure 14.--Frequency distribution of mean prey size for taxa eaten by Zalophus californianus. 

 34 
Phocid Seals 

Phocidae: Halichoerus grypus, gray seal; n = 52,  ̄x= 24.64 cm

Prey size information for gray seals was recorded for one cephalopod species and 18 

species of fish based on the citations listed in Table 7. There are 52 entries of mean prey size for

H. grypus that meet our minimum sample size requirement, with an overall mean of means of 

24.64 cm. The modal mean prey size for H. grypus is 15-30 cm, with a slightly positively 

skewed distribution (Fig. 15).

Table 7.--Sources of prey size data for Halichoerus grypus. 

Citation Scats Stomachs Regurge Lavage Age/sex

Benoit and Bowen 1990 295* 

Bowen and Harrison 1994 393 

Bowen et al. 1993 528 

Hammond et al. 1994a 993 

Hammond et al. 1994b 749 

Hauksson and Bogason 1997 1,059 

Murie and Lavigne 1992 82 

Prime and Hammond 1990 481 

*count includes only food-containing stomachs 

 35 
0

2

4

6

8

10

12

14

16

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 52; mean of means = 24.64

Figure 15.--Frequency distribution of mean prey size for taxa eaten by Halichoerus grypus. 

 36 
Phocidae: Mirounga leonina, southern elephant seal; n = 12,  ̄x= 14.72 cm

Prey size information for southern elephant seals was recorded for nine cephalopod 

species and one species of fish based on the three citations listed in Table 8. Twelve entries of 

mean prey size for M. leonina met our minimum sample size requirement, with an overall mean 

of means of 14.72 cm. The modal mean prey size for M. leonina is 15-20 cm, with a distribution 

that is approximately normal (Fig. 16).

Table 8.--Sources of prey size data for Mirounga leonina. 

Citation Scats Stomachs Regurge Lavage Age/sex

Daneri and Carlini 2002 153 

Daneri et al. 2000 25 

Rodhouse et al. 1992 51 

 37 
0

1

2

3

4

5

6

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 12; mean of means = 14.72

Figure 16.--Frequency distribution of mean prey size for taxa eaten by Mirounga leonina. 

 38 
Phocidae: Phoca groenlandica, harp seal; n = 11,  ̄x= 20.31 cm

Prey size information for harp seals was recorded for seven species of fish based on the 

citations listed in Table 9. Eleven entries of mean prey size for P. groenlandica met our 

minimum sample size requirement, with an overall mean of means of 20.31 cm. The modal 

mean prey size for P. groenlandica is 15-20 cm, with a slightly positively skewed distribution 

(Fig. 17). 

Table 9.--Sources of prey size data for Phoca groenlandica. 

Citation Scats Stomachs Regurge Lavage Age/sex

Beck et al. 1993 247 

Finley et al. 1990 157 

Hauksson and Bogason 1997 1,059 pups and 1-year olds

Murie and Lavigne 1991 25 

Nilssen et al. 1990 59 

 39 
0

1

2

3

4

5

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 11; mean of means = 20.31

Figure 17.--Frequency distribution of mean prey size for taxa eaten by Phoca groenlandica. 

 40 
Phocidae: Phoca vitulina, harbor seal; n = 33,  ̄x= 15.45 cm

Prey size information for harbor seals was recorded for one cephalopod species and 23 

species of fish based on the seven citations listed in Table 10. There are 33 entries of mean prey 

size for P. vitulina that meet our minimum sample size requirement, with an overall mean of 

means of 15.45 cm. The modal mean prey size for P. vitulina is 5-20 cm, with a slightly 

positively skewed distribution (Fig. 18).

Table 10.--Sources of prey size data for Phoca vitulina. 

Citation Scats Stomachs Regurge Lavage Age/sex

Behrends 1982 185 

Bowen and Harrison 1996 338 pups, yearlings, and 

>1 year old 

Brown and Mate 1983 150 

Frost and Lowry 1986 5 

Harvey et al. 1995 30 

Pitcher 1981 548 

Torok 1994 215 

 41 
0

1

2

3

4

5

6

7

8

9

10

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 33; mean of means = 15.45

Figure 18.--Frequency distribution of mean prey size for taxa eaten by Phoca vitulina. 

 42 
Odontocete Whales 

Delphinidae: Globicephala melas = G. melaena, long-finned pilot whale; n = 10,  ̄x= 16.43 cm

Prey size information for long-finned pilot whales was recorded for seven cephalopod 

species and one species of fish based on the citations listed in Table 11. Ten entries of mean 

prey size for G. melas met our minimum sample size requirement, with an overall mean of 

means of 16.43 cm. The modal mean prey size for G. melas is 15-20 cm, with a slightly 

positively skewed distribution (Fig. 19).

Table 11.--Sources of prey size data for Globicephala melas. 

Citation Scats Stomachs Regurge Lavage Age/sex

Clarke and Goodall 1994 4 1 M; 3 F 

Desportes and Mouritsen 1988 720 

dos Santos and Haimovici 2001 5 

Gannon et al. 1997a 30 

Gannon et al. 1997b 8 4 M; 4 F 

calf to mature

 43 
0

1

2

3

4

5

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 10; mean of means = 16.43

Figure 19.--Frequency distribution of mean prey size for taxa eaten by Globicephala melas. 

 44 
Delphinidae: Grampus griseus, Risso's dolphin; n = 15,  ̄x= 12.08 cm

Prey size information for Risso’s dolphins was recorded for 14 cephalopod species based 

on the two citations listed in Table 12. Fifteen entries of mean prey size for G. griseus met our 

minimum sample size requirement, with an overall mean of means of 12.08 cm. The modal 

mean prey size for G. griseus is 5-10 cm, with a slightly positively skewed distribution (Fig. 20).

Table 12.--Sources of prey size data for Grampus griseus. 

Citation Scats Stomachs Regurge Lavage Age/sex

Clarke and Young 1998 1 

Sekiguchi et al. 1992 18 

0

1

2

3

4

5

6

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 15; mean of means = 12.08

Figure 20.--Frequency distribution of mean prey size for taxa eaten by Grampus griseus. 

 45 
Delphinidae: Sotalia fluviatilis, tucuxi; n = 10,  ̄x= 9.71 cm

Prey size information for the tucuxi was recorded for three cephalopod species and six 

species of fish based on the two citations listed in Table 13. Ten entries of mean prey size for S. 

fluviatilis met our minimum sample size requirement, with an overall mean of means of 9.71 cm. 

The mean prey size for S. fluviatilis has weak modes at 0-5 cm and 10-15 cm (Fig. 21).

Table 13.--Sources of prey size data for Sotalia fluviatilis. 

Citation Scats Stomachs Regurge Lavage Age/sex

de Oliveira Santos et al. 2002 9 

dos Santos and Haimovici 2001 56 

0

1

2

3

4

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 10; mean of means = 9.71

Figure 21.--Frequency distribution of mean prey size for taxa eaten by Sotalia fluviatilis. 

 46 
Delphinidae: Stenella attenuata, pan-tropical spotted dolphin; n = 37,  ̄x= 12.74 cm

Prey size information for pan-tropical spotted dolphins was recorded for five cephalopod 

species and nine species of fish based on the three citations listed in Table 14. There are 37 

entries of mean prey size for S. attenuata that meet our minimum sample size requirement, with 

an overall mean of means of 12.74 cm. The modal mean prey size for S. attenuata is 15-20 cm, 

with a slightly positively skewed distribution (Fig. 22).

 

Table 14.--Sources of prey size data for Stenella attenuata. 

Citation Scats Stomachs Regurge Lavage Age/sex

Perrin et al. 1973 22 

Robertson and Chivers 1997 428 

Sekiguchi et al. 1992 3 

Wang et al. 2003 45 

 47 
0

1

2

3

4

5

6

7

8

9

10

11

12

13

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequency

n = 37; mean of means = 12.74

Figure 22.--Frequency distribution of mean prey size for taxa eaten by Stenella attenuata. 

 48 
Phocoenidae: Phocoena phocoena, harbor porpoise; n = 31,  ̄x= 16.99 cm

Prey size information for harbor porpoises was recorded for two cephalopod species and 

22 species of fish based on the citations listed in Table 15. There are 31 entries of mean prey 

size for P. phocoena that meet our minimum sample size requirement, with an overall mean of 

means of 16.99 cm. The modal mean prey size for P. phocoena is 15-20 cm, with an 

approximately normal distribution (Fig. 23).

Table 15.--Sources of prey size data for Phocoena phocoena. 

Citation Scats Stomachs Regurge Lavage Age/sex

Börjesson et al. 2003 112 60 M; 52 F 

juvenile to adult 

Fontaine et al. 1994 111* 

Gannon et al. 1998 95 

Morejohn et al. 1978 15 

Recchia and Read 1989 127 

Walker et al. 1998 25 14 M; 11 F 

*count of food-containing stomachs only 

 49 
0

1

2

3

4

5

6

7

8

9

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 31; mean of means = 16.99

Figure 23.--Frequency distribution of mean prey size for taxa eaten by Phocoena phocoena. 

 50 
Phocoenidae: Phocoenoides dalli, Dall's porpoise; n = 37,  ̄x= 12.88 cm

Prey size information for Dall’s porpoises was recorded for 11 cephalopod species and 12 

taxa of fish, based on the five citations listed in Table 16. There are 37 entries of mean prey size 

for P. dalli that meet our minimum sample size requirement, with an overall mean of means of 

12.88 cm. The modal mean prey size for P. dalli is 5-15 cm, with a slightly positively skewed 

distribution (Fig. 24).

Table 16.--Sources of prey size data for Phocoenoides dalli. 

Citation Scats Stomachs Regurge Lavage Age/sex

Crawford 1981 457 

Morejohn et al. 1978 27 

Ohizumi et al. 2000 150 

Walker 1996 85 

Walker et al. 1998 22 11 M; 11 F 

 51 
0

2

4

6

8

10

12

14

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 37; mean of means = 12.88

Figure 24.--Frequency distribution of mean prey size for taxa eaten by Phocoenoides dalli. 

 52 
Physeteridae: Kogia breviceps, pygmy sperm whale; n = 12,  ̄x= 15.85 cm

Prey size information for pygmy sperm whales was recorded for 10 cephalopod species 

and one species of fish based on the three citations listed in Table 17. Twelve entries of mean 

prey size for K. breviceps met our minimum sample size requirement, with an overall mean of 

means of 15.85 cm. The modal mean prey size for K. breviceps is 10-25 cm, with an 

approximately normal distribution (Fig. 25).

Table 17.--Sources of prey size data for Kogia breviceps. 

Citation Scats Stomachs Regurge Lavage Age/sex

dos Santos and Haimovici 2001 3 

Sekiguchi et al. 1992 24 

Wang et al. 2002 6 

 53 
0

1

2

3

4

5

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 12; mean of means = 15.85

Figure 25.--Frequency distribution of mean prey size for taxa eaten by Kogia breviceps. 

 54 
Physeteridae: Physeter macrocephalus = P. catadon, sperm whale; n = 47,  ̄x= 31.35 cm

Prey size information for sperm whales was recorded for 31 cephalopod species based on 

the citations listed in Table 18. There are 47 entries of mean prey size for P. macrocephalus that

meet our minimum sample size requirement, with an overall mean of means of 31.35 cm. The 

modal mean prey size for P. macrocephalus is 20-25 cm, with a positively skewed distribution 

(Fig. 26).

Table 18.--Sources of prey size data for Physeter macrocephalus. Cells with “n.a.” indicate 

cases where the sample type is known but the sample size was not reported. 

Citation Scats Stomachs Regurge Lavage Age/sex

Best 1999 1,268 

Clarke 1997 1 

Clarke and Young 1998 2 

Clarke et al. 1993 17 15 M; 2 F 

Nemoto et al. 1985 n.a. 

Nemoto et al. 1987 n.a. 

Santos et al. 1999 17 

 55 
0

1

2

3

4

5

6

7

8

9

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 47; mean of means = 31.35

Figure 26.--Frequency distribution of mean prey size for taxa eaten by Physeter macrocephalus. 

 56 
Ziphiidae: Berardius bairdii, Baird's beaked whale; n = 15,  ̄x= 29.35 cm

Prey size information for Baird's beaked whales was recorded for three cephalopod 

species and six species of fish based on the two citations listed in Table 19. There are 15 entries 

of mean prey size for B. bairdii that meet our minimum sample size requirement, with an overall 

mean of means of 29.35 cm. The modal mean prey size for B. bairdii is 35-40 cm, with an 

approximately normal distribution (Fig. 27).

Table 19.--Sources of prey size data for Berardius bairdii. 

Citation Scats Stomachs Regurge Lavage Age/sex

Ohizumi et al. 2003 26 11 M; 15 F 

Walker et al. 2002 127 

0

1

2

3

4

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 15; mean of means = 36.64

Figure 27.--Frequency distribution of mean prey size for taxa eaten by Berardius bairdii. 

 57 
Ziphiidae: Hyperoodon planifrons, southern bottlenosed whale; n = 44,  ̄x= 25.04 cm

Prey size information for southern bottlenosed whales was recorded for 25 cephalopod 

species based on the three citations listed in Table 20. There are 44 entries of mean prey size for 

H. planifrons that meet our minimum sample size requirement, with an overall mean of means of 

25.04 cm. The modal mean prey size for H. planifrons is 5-25 cm, with a positively skewed 

distribution (Fig. 28).

Table 20.--Sources of prey size data for Hyperoodon planifrons. 

Citation Scats Stomachs Regurge Lavage Age/sex

Clarke and Goodall 1994 2 

Sekiguchi et al. 1993 2 1 M; 1 F 

Slip et al. 1995 1 F 

 58 
0

1

2

3

4

5

6

7

8

9

 0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequenc

y

n = 44; mean of means = 25.04

Figure 28.--Frequency distribution of mean prey size for taxa eaten by Hyperoodon planifrons. 

 59 
Size Composition of Prey Consumed, Sorted by Prey Species 

In addition to knowing predator-specific patterns in the size of prey targeted by predators, 

it is also useful to determine the size composition of a single prey species targeted by multiple 

predators. Most of the prey species for which we have data are represented by fewer than five 

data entries (Fig. 29; Appendix 3, 4). We use the same minimum sample size requirement for

determining which cases were acceptable for examination here as we did to characterize the diets 

of predators. That is, prey species covered here have ≥ 10 entries for which mean targeted size is 

known, with each entry based on sample sizes of ≥ 10 individuals of that prey species. Note that 

individual entries (information from the literature that serves as our data points) may represent 

multiple studies of the same predator species or independent studies of different predator species.

46

27 21

5 4 4 4 1

13

157

0

20

40

60

80

100

120

140

160

180

1 2 3 4 5 6 7 8 9 ≥10

Number of entries per prey taxon with n ≥ 10

Frequency

Figure 29.--Frequency distribution of the number of cases per prey taxon for which 

 mean prey size was reported based on a sample size of 10 or larger. 

For each of the following prey species, we also present data on what is known about the 

maximum size individuals are likely to attain. Although most of our data on maximum prey size 

 60 
come from studies where examination of whole animals was possible (Froese and Pauly 2003, 

Wood and Day 1998), a large proportion of data are based on reconstructed sizes estimated 

through regressions of body size against the size of hard parts such as squid beaks or fish otoliths 

likely to survive the digestion process. Specifically, of the 379 non-overlapping taxa for which 

data on maximum reported size were available, 100 (26%) derive from food habits studies. As 

discussed in our Methods section, food habits data were used when no other source of data was 

available, or when the food habits length data exceeded other available data. The effect this is 

likely to have on subsequent analyses is highly variable. For instance, krill (Euphausia superba) 

are generally regarded as reaching a maximum size of 6.2 cm (FAO 2000). The reconstructed 

maximum size based on regressions of carapace length to total length has been reported by Reid 

et al. (1996) to be 6.6 cm—an increase of 6%. In contrast, the maximum size of market squid 

(Loligo opalescens) reported by Lowry and Caretta (1999) was estimated by regressions of beak 

length to dorsal mantle length to be 23.5 cm, exceeding the largest whole squid measured by 

Kashiwada et al. (1979) by 4.0 cm—an increase of roughly 21%. 

There are several potential explanations for these discrepancies. First, there may be 

problems with the regressions used to estimate body size. Regressions used for estimating body 

size are rarely presented with plots of the original data, making it impossible to evaluate if 

appropriate transformations have been used and whether or not the error structure is uniform

throughout the distribution. As a consequence, regressions may be providing biased estimates in 

the tails of the distribution (Konigsberg et al. 1997). 

This assumes, of course, that the prey remains have been identified correctly. If the 

wrong regressions are used then the estimated size will be unreliable. This is particularly 

problematic for cephalopod species, since the taxonomy of the beaks, let alone the whole 

 61 
animals, can be problematic even for specialists in the field. By way of example, 53 of the 100 

cases for which we used size information from the food habits literature are for cephalopod 

species. In contrast, fish and crustaceans represent 36/100 and 11/100 cases, respectively. 

Finally, we need to consider the theoretical nature of maximum reported size. Most, if 

not all, of the species considered here are indeterminate in their growth. That is, they grow 

throughout their lifetime, albeit at a steadily decreasing rate. Therefore, if any of these species 

were allowed to grow indefinitely, the maximum reported size may well be exceeded. 

Nevertheless, the available data on maximum reported size provide a valuable reference point for 

evaluating the size composition of targeted prey species. 

 62 
Crustacean Prey Species 

Euphausiidae: Euphausia superba, Antarctic krill; n = 15,  ̄x = 4.53 cm

Krill reach a maximum size of about 6.6 cm total length (maximum reported size from

Reid et al. 1996). The modal mean size of marine mammal predation is 5 cm, with a mean of 

means of 4.53 cm (Fig. 30). 

Table 21.--Predators for which mean size of Euphausia superba is known from sample sizes of 

10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus gazella South Georgia Island, S Atlantic 1972-1977 Croxall and Pilcher 1984

A. gazella South Georgia Island, S Atlantic 1986 Croxall et al. 1999 

A. gazella South Georgia Island, S Atlantic 1994 Croxall et al. 1999 

A. gazella South Georgia Island, S Atlantic 1982-1983 Doidge and Croxall 

1985 

A. gazella Bouvetøya Island, Southern Ocean 1998-1999 Kirkman et al. 2000 

A. gazella South Georgia Island, S Atlantic 1994-1996 McCafferty et al. 1998 

A. gazella South Georgia Island, S Atlantic 1992 Reid 1995 

A. gazella South Georgia Island, S Atlantic 1993 Reid 1995 

A. gazella South Georgia Island, S Atlantic 1991-1994 Reid and Arnould 1996 

A. gazella South Georgia Island, S Atlantic 1994-1999 Reid and Brierly 2001 

A. gazella South Georgia Island, S Atlantic 1986 Reid et al. 1996 

Balaenoptera spp.* Southern Ocean 1950s?? Mackintosh 1974 

Balaenoptera spp.* South Georgia Island, S Atlantic 1929-1930 Marr 1962 

 63 
Balaenoptera spp.* South Georgia Island, S Atlantic 1932-1933 Marr 1962 

Balaenoptera spp.* South Georgia Island, S Atlantic 1939-1940 Marr 1962 

* samples consist of an unspecified mix of blue (B. musculus) and fin (B. physalus) whales 

0

1

2

3

4

5

6

7

8

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

Total length (cm)

Frequency

n = 15, mean of means= 4.53

Figure 30.--Frequency distribution of mean size of Euphausia superba consumed by 

 marine mammals. 

 64 
Cephalopod Prey Species 

Loliginidae: Loligo opalescens, market squid; n = 10,  ̄x = 11.58 cm

Market squid reach a maximum size of 23.50 cm (maximum reported size from Lowry 

and Caretta 1999). The size of market squid consumed by marine mammals was determined 

from food habits studies of three species of pinnipeds and three species of odontocetes (Table 

22). The modal mean size of market squid eaten by marine mammals is 10-15 cm, with an 

overall mean of means of 11.58 cm (Fig. 31). 

Table 22.--Predators for which mean size of Loligo opalescens is known from sample sizes of 10 

or more. 

Predator Sampling Region Dates Citation 

Callorhinus ursinus N Pacific 1958-1974 Perez and Bigg 1986 

Globicephala macrorhynchus Channel Islands, NE Pacific 1980 Seagars and Henderson 1985

G. macrorhynchus S California Bight, NE Pacific 1977 Sinclair 1992

G. macrorhynchus S California Bight, NE Pacific 1969-1977 Sinclair 1992

Mirounga angustirostris S California Bight, NE Pacific 1969-1983 Sinclair 1994

Phocoena phocoena Washington and British Columbia 1990-1997 Walker et al. 1998 

Phocoenoides dalli Washington and British Columbia 1990-1997 Walker et al. 1998 

Zalophus californianus San Miguel Island, NE Pacific 1978-1979 Antonelis et al. 1984 

Z. californianus Channel Islands, NE Pacific 1981-1985 Lowry and Caretta 1999 

Z. californianus San Miguel Island, NE Pacific 1993, 1996 Melin 2002 

Z. californianus Monterey Bay, NE Pacific n.a. Morejohn et al. 1978 

 65 
0

1

2

3

4

5

6

7

5 10 15 20 25 30 35 40 45 50 55 60

Dorsal mantle length (cm)

Frequency

n = 10, mean of means= 11.58

Figure 31.--Frequency distribution of mean size of Loligo opalescens consumed by marine 

 mammals.

 66 
Loliginidae: Loligo vulgaris reynaudii, Cape Horn squid; n = 11,  ̄x = 14.04 cm

Cape Horn squid reach a maximum size of 40 cm (Wood and Day 1998). The size of 

Cape Horn squid consumed by marine mammals is based on food habits studies of one species of 

pinniped and eight species of odontocetes (Table 23). The modal mean size of Cape Horn squid 

eaten by marine mammals is 10-20 cm, with an overall mean of means of 14.04 cm (Fig. 32). 

Table 23.--Predators for which mean size of Loligo vulgaris reynaudii is known from sample 

sizes of 10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus pusillus pusillus Benguela, South Africa 1976-1990 Castley et al. 1991 

A. p. pusillus Benguela, South Africa 

(West Coast) 

1974-1985 Lipinski and David 1990

A. p. pusillus Benguela, South Africa 

(South Coast) 

1974-1985 Lipinski and David 1990

Cephalorhynchus heavisidii Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

Delphinus delphis Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

Grampus griseus Benguela, South Africa 1975-1990 Sekiguchi et al. 1992 

Kogia breviceps Benguela, South Africa 1975-1990 Sekiguchi et al. 1992 

K. sima Benguela, South Africa 1975-1988 Sekiguchi et al. 1992 

Lagenorhynchus obscurus Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

Stenella coeruleoalba Benguela, South Africa 1975-1989 Sekiguchi et al. 1992 

Tursiops truncatus Benguela, South Africa 1975-1989 Sekiguchi et al. 1992 

 67 
0

1

2

3

4

5

 0-5 10-15 20-25 30-35 40-45 50-55

Dorsal mantle length (cm)

Frequency

n = 11, mean of means = 14.04

Figure 32.--Frequency distribution of mean size of Loligo vulgaris reynaudii consumed by 

 marine mammals. 

 68 
Fish Prey Species 

Clupeidae: Clupea harengus, Atlantic herring; n = 10,  ̄x = 27.49 cm

Atlantic herring reach a maximum size of 45 cm (Froese and Pauly 2003). The size of

Atlantic herring consumed by marine mammals was determined from food habits studies of three 

species of pinniped and one odontocete (Table 24). The modal mean size of Atlantic herring 

eaten by marine mammals is 25-30 cm, with an overall mean of means of 27.49 cm (Fig. 33). 

Table 24.--Predators for which mean size of Clupea harengus is known from sample sizes of 10 

or more. 

Predator Sampling Region Dates Citation 

Halichoerus grypus Gulf of St. Lawrence, NW Atlantic 1982-1987 Benoit and Bowen 1990 

H. grypus Scotian Shelf, NW Atlantic 1988-1990 Bowen et al. 1993 

H. grypus Gulf of St. Lawrence, NW Atlantic 1983 Murie and Lavigne 1992 

Phoca groenlandica Iceland 1992-1993 Hauksson and Bogason 1997 

P. vitulina Nova Scotia, Canada 1988-1992 Bowen and Harrison 1996 

P. vitulina Bay of Fundy, NW Atlantic 1988-1992 Bowen and Harrison 1996 

Phocoena phocoena Kattegat Sea and Skagerrak Sea 1989-1996 Börjesson et al. 2003 

P. phocoena Gulf of St. Lawrence, NW Atlantic 1989 Fontaine et al. 1994 

P. phocoena Gulf of Maine, NW Atlantic 1989-1994 Gannon et al. 1998 

P. phocoena Bay of Fundy, NW Atlantic 1985-1987 Recchia and Read 1989 

 69 
0

1

2

3

4

5

6

7

 0-5 10-15 20-25 30-35 40-45

Length (cm)

Frequency

n = 10, mean of means = 27.49

Figure 33.--Frequency distribution of mean size of Clupea harengus consumed by marine 

 mammals.

 70 
Clupeidae: Sardinops sagax = S. ocellatus, pilchard/sardine; n = 11,  ̄x = 16.72 cm

Pilchards reach a maximum size of 39.50 cm (Froese and Pauly 2003). The size of 

pilchards consumed by marine mammals was determined from food habits studies of four taxa of 

pinnipeds and three odontocetes (Table 25). The modal mean size of pilchards eaten by marine 

mammals is 15-20 cm, with an overall mean of means of 16.72 cm (Fig. 34). 

Table 25.--Predators for which mean size of Sardinops sagax is known from sample sizes of 10 

or more. 

Predator Sampling Region Dates Citation 

Arctocephalus galapagoensis Galapagos Islands, E Tropical Pacific 1983 Dellinger and Trillmich 1999

A. pusilus doriferus Tasmania 1984-1986 Gales and Pemberton 1994

A. pusillus pusillus Benguela, South Africa 1974-1985 David 1987 

A. p. pusillus Benguela, South Africa 1974-1985 David 1987 

A. p. pusillus Benguela, South Africa 1974-1985 David 1987 

Delphinus delphis Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

Lagenorhynchus obscurus Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

Phocoenoides dalli Sea of Okhotsk and Sea of Japan 1988-1996 Ohizumi et al. 2000 

P. dalli Sea of Okhotsk 1988-1989 Walker 1996

Zalophus californianus woelbecki Galapagos Islands, E Tropical Pacific 1983 Dellinger and Trillmich 1999

Z. c. woelbecki Galapagos Islands, E Tropical Pacific 1983-1986 Dellinger and Trillmich 1999

 71 
0

1

2

3

4

5

6

7

8

 0-5 10-15 20-25 30-35 40-45

Length (cm)

Frequency

n = 11, mean of means = 16.72

Figure 34.--Frequency distribution of mean size of Sardinops sagax consumed by marine 

 mammals.

 72 
Myctophidae: Electrona antarctica, unnamed lanternfish; n = 12,  ̄x = 7.83 cm

Electrona antarctica reaches a maximum size of 13.26 cm (maximum reported size from

Kirkman et al. 2000). The size of E. antarctica consumed by marine mammals was determined 

from the results of 11 food habits studies of Arctocephalus gazella (Table 26). The modal mean 

size of E. antarctica eaten by A. gazella is 8 cm, with an overall mean of means of 7.83 cm (Fig. 

35). 

Table 26.--Predators for which mean size of Electrona antarctica is known from sample sizes of 

10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus gazella South Shetland Islands, S Atlantic 1996-1997 Casaux et al. 1998 

A. gazella Iles Kerguelen, S Atlantic 1994 Cherel et al. 1997 

A. gazella South Shetland Islands, S Atlantic 1992 Daneri 1996 

A. gazella South Shetland Islands, S Atlantic 1993-1994 Daneri and Carlini 1999 

A. gazella South Orkney Islands, S Atlantic 1988 Daneri and Coria 1993 

A. gazella Heard Island, Indian Ocean 1987-1988 Green et al. 1989 

A. gazella Heard Island, Indian Ocean 1987-1988 Green et al. 1989 

A. gazella Heard Island, Indian Ocean 1993-1993 Green et al. 1997 

A. gazella Bouvetøya Island, Southern Ocean 1998-1999 Kirkman et al. 2000 

A. gazella Bouvetøya Island, Southern Ocean 1996-1997 Klages et al. 1999 

A. gazella Iles Kerguelen, S Atlantic 1998-2000 Lea et al. 2002 

A. gazella South Georgia Island, S Atlantic 1983 North 1996 

 73 
0

1

2

3

4

5

6

12345678 9 10 11 12 13 14 15

Length (cm)

Frequency

n = 12, mean of means = 7.83

Figure 35.--Frequency distribution of mean size of Electrona antarctica consumed by marine 

 mammals.

 74 
Mycophidae: Electrona subaspera, unnamed lanternfish; n = 10,  ̄x = 8.04 cm

Electrona subaspera reach a maximum size of 12.75 cm (maximum reported size from 

Green et al. 1997). The size of E. subaspera consumed by marine mammals was determined 

from the results of food habits studies of three species of fur seal (Table 27). The modal mean 

size of E. subaspera eaten by Arctocephalus spp. is 8 cm, with an overall mean of means of 

8.04 cm (Fig. 36). 

Table 27.--Predators for which mean size of Electrona subaspera is known from sample sizes of 

10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus forsteri Macquarie Island, S Atlantic 1988-1989 Green et al. 1990 

A. forsteri Macquarie Island, S Atlantic 1988-1989 Green et al. 1990 

A. gazella Macquarie Island, S Atlantic 1990-1991 Goldsworthy et al. 1997 

A. gazella Heard Island, Indian Ocean 1993-1993 Green et al. 1997 

A. gazella Marion Island, S Atlantic 1989-1995 Klages and Bester 1998

A. gazella Iles Kerguelen, S Atlantic 1998-2000 Lea et al. 2002 

A. tropicalis Macquarie Island, S Atlantic 1990-1991 Goldsworthy et al. 1997 

A. tropicalis Marion Island, S Atlantic 1989-1995 Klages and Bester 1998

Arctocephalus spp.* Macquarie Island, S Atlantic 1988-1989 Green et al. 1990 

Arctocephalus spp.* Macquarie Island, S Atlantic 1988-1989 Green et al. 1990 

*includes an unspecified mix of A. forsteri, A. gazella, and A. tropicalis

 75 
0

1

2

3

4

5

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

Length (cm)

Frequency

n = 10, mean of means = 8.04

Figure 36.--Frequency distribution of mean size of Electrona subaspera consumed by marine 

 mammals.

 76 
Myctophidae: Gymnoscopelus nicholsi, unnamed lanternfish; n = 15,  ̄x = 12.06 cm

Gymnoscopelus nicholsi reach a maximum size of 18.90 cm (maximum reported size 

from Casaux et al. 1998). The size of G. nicholsi consumed by marine mammals was 

determined from food habits studies of three species of fur seal and one phocid (Table 28). The 

modal mean size of G. nicholsi eaten by marine mammals is 13 cm, with an overall mean of 

means of 12.06 cm (Fig. 37). 

Table 28.--Predators for which mean size of Gymnoscopelus nicholsi is known from sample sizes 

of 10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus gazella Bouvetøya Island, Southern Ocean 1998-1999 Kirkman et al. 2000 

A. gazella South Shetland Islands, S Atlantic 1996-1997 Casaux et al. 1998 

A. gazella Iles Kerguelen, S Atlantic 1994 Cherel et al. 1997 

A. gazella South Shetland Islands, S Atlantic 1992 Daneri 1996 

A. gazella South Shetland Islands, S Atlantic 1993-1994 Daneri and Carlini 1999

A. gazella South Orkney Islands, S Atlantic 1988 Daneri and Coria 1993 

A. gazella Heard Island, Indian Ocean 1987-1988 Green et al. 1989 

A. gazella Heard Island, Indian Ocean 1987-1988 Green et al. 1989 

A. gazella Marion Island, S Atlantic 1989-1995 Klages and Bester 1998

A. gazella Bouvetøya Island, Southern Ocean 1996-1997 Klages et al. 1999 

A. gazella Iles Kerguelen, S Atlantic 1998-2000 Lea et al. 2002 

A. tropicalis Marion Island, S Atlantic 1989-1995 Klages and Bester 1998

 77 
Arctocephalus spp.* Macquarie Island, S Atlantic 1988-1989 Green et al. 1990 

Arctocephalus spp.* Macquarie Island, S Atlantic 1988-1989 Green et al. 1990 

Mirounga leonina South Shetland Islands, S Atlantic 1993-2000 Daneri and Carlini 2002

*includes an unspecified mix of A. forsteri, A. gazella, and A. tropicalis 

0

1

2

3

4

5

6

7

8

123 45678 9 10 11 12 13 14 15

Length (cm)

Frequency

n = 15, mean of means = 12.06

Figure 37.--Frequency distribution of mean size of Gymnoscopelus nicholsi consumed by marine 

 mammals.

 78 
Gadidae: Gadus morhua, Atlantic cod; n = 19,  ̄x = 26.15 cm

Atlantic cod reach a maximum size of 200 cm (Froese and Pauly 2003). The size of 

Atlantic cod consumed by marine mammals was determined from food habits studies of five 

species of pinnipeds and one odontocete (Table 29). The modal mean size of Atlantic cod eaten 

by marine mammals is 25-30 cm, with an overall mean of means of 26.15 cm (Fig. 38). 

Table 29.--Predators for which mean size of Gadus morhua is known from sample sizes of 10 or 

more. 

Predator Sampling Region Dates Citation 

Cystophora cristata Iceland 1992-1993 Hauksson and Bogason 1997 

Halichoerus grypus Gulf of St. Lawrence, NW Atlantic 1982-1987 Benoit and Bowen 1990 

H. grypus Scotian Shelf, NW Atlantic 1991-1993 Bowen and Harrison 1994 

H. grypus Scotian Shelf, NW Atlantic 1988-1990 Bowen et al. 1993 

H. grypus Orkney Island, NE Atlantic 1985 Hammond et al. 1994a 

H. grypus Orkney Island, NE Atlantic 1985 Hammond et al. 1994a 

H. grypus Hebrides Islands, NE Atlantic 1985 Hammond et al. 1994b 

H. grypus Hebrides Islands, NE Atlantic 1985 Hammond et al. 1994b 

H. grypus Hebrides Islands, NE Atlantic 1985 Hammond et al. 1994b 

H. grypus Gulf of St. Lawrence, NW Atlantic 1983 Murie and Lavigne 1992 

H. grypus Donna Nook, SW North Sea 1985 Prime and Hammond 1990

Phoca groenlandica Gulf of St. Lawrence, NW Atlantic 1988-1990 Beck et al. 1993 

P. groenlandica Iceland 1992-1993 Hauksson and Bogason 1997 

P. groenlandica Ullsford, Norway 1986 Nilssen et al. 1990 

P. groenlandica Ullsford, Norway 1988 Nilssen et al. 1990 

 79 
P. vitulina Waddensea, North Sea 1975-1981 Behrends 1982 

P. vitulina Bay of Fundy, NW Atlantic 1988-1992 Bowen and Harrison 1996 

P. vitulina Nova Scotia, Canada 1988-1992 Bowen and Harrison 1996 

Phocoena phocoena Kattegat Sea and Skagerrak Sea 1989-1996 Börjesson et al. 2003 

0

1

2

3

4

5

6

7

 0-5 10-15 20-25 30-35 40-45

Length (cm)

Frequency

n = 19, mean of means = 26.15

Figure 38.--Frequency distribution of mean size of Gadus morhua consumed by marine 

 mammals. 

 80 
Gadidae: Theragra chalcogramma, walleye pollock; 91.00 n = 26,  ̄x = 24.10 cm

Walleye pollock reach a maximum size of 91 cm (Froese and Pauly 2003). The size of

pollock consumed by marine mammals was determined from food habits studies of six species of 

pinnipeds, two odontocetes, and one mysticete (Table 30). The frequency distribution of mean 

size of pollock eaten by marine mammals is tri-modal, with strong modes at 10-15 cm and 25-30 

cm, and a weak mode at 45-50 cm. The overall mean of means is 24.10 cm (Fig. 39). 

Table 30.--Predators for which mean size of Theragra chalcogramma is known from sample 

sizes of 10 or more. 

Predator Sampling Region Dates Citation 

Balaenoptera acutorostrata Bering Sea 1975 Frost and Lowry 1986 

Berardius bairdii Abahiri, Japan, Sea of Okhotsk Jul - Aug 1999 Ohizumi et al. 2003 

Callorhinus ursinus St. Paul Island, Bering Sea 1996 Kiyota et al. 1999 

C. ursinus Bering Sea 1981-1982 Lowry et al. 1989 

C. ursinus Bering Sea 1974 McAlister et al. 1976 

C. ursinus Bering Sea 1981 Sinclair et al. 1994 

C. ursinus Bering Sea 1982 Sinclair et al. 1994 

C. ursinus Bering Sea 1985 Sinclair et al. 1994 

Erignathus barbatus Bering Sea 1981 Antonelis et al. 1994b 

Eumetopias jubatus Kodiak Island, Gulf of Alaska 1985-1986 Calkins and Goodwin 1988

E. jubatus SE Alaska, Gulf of Alaska 1985-1986 Calkins and Goodwin 1988

E. jubatus Pribilof Islands, Bering Sea 1976, 1979 Frost and Lowry 1986 

 81 
E. jubatus Bering Sea 1981 Frost and Lowry 1986 

E. jubatus Gulf of Alaska 1975-1978 Pitcher 1981

E. jubatus SE Alaska 1994-1999 Tollit et al. 2004a 

E. jubatus Aleutian Is. (western stock) 1998-2000 Zeppelin et al. 2004 

Phoca fasciata Bering Sea 1976-1979 Frost and Lowry 1980 

P. fasciata Bering Sea 1976-1978 Frost and Lowry 1986 

P. largha Bering Sea 1976-1978 Bukhtiyarov et al. 1984 

P. largha Bering Sea 1978 Frost and Lowry 1986 

P. vitulina Bering Sea 1981 Frost and Lowry 1986 

P. vitulina Pribilof Islands, Bering Sea 1979 Frost and Lowry 1986 

P. vitulina Gulf of Alaska 1975-1978 Pitcher 1981

Phocoenoides dalli Sea of Okhotsk and Sea of Japan 1988-1996 Ohizumi et al. 2000 

P. dalli Sea of Okhotsk 1988-1989 Walker 1996

P. dalli Washington and British Columbia 1990-1997 Walker et al. 1998 

 82 
0

1

2

3

4

5

6

7

8

9

 0-5 10-15 20-25 30-35 40-45

Length (cm)

Frequency

n = 26, mean of means = 24.10

Figure 39.--Frequency distribution of mean size of Theragra chalcogramma consumed by 

 marine mammals.

 83 
Merlucciidae: Merluccius spp. (M. capensis and M. paradoxus), shallow-water and deep-water

Cape hakes, respectively; n = 10,  ̄x = 20.84 cm

Food habits studies rarely distinguish between the two species of hake common in the 

Benguela Current of South Africa. However, the shallow-water hake (maximum size = 140 cm; 

Froese and Pauly 2003) is most abundant in waters shallower than 200 m, whereas the deepwater hake (maximum size = 115 cm; Froese and Pauly 2003) is most abundant in waters 300 m 

and deeper (Botha 1985). The size of Cape hakes consumed by marine mammals is based on 

food habits studies of one species of pinniped and seven odontocetes (Table 31). The modal 

mean size of Cape hakes eaten by marine mammals is 15-25 cm, with an overall mean of means 

of 20.84 cm (Fig. 40). 

Table 31.--Predators for which mean size of Merluccius capensis and M. paradoxus is known 

from sample sizes of 10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus pusillus pusillus Benguela, South Africa 1974-1985 David 1987 

A. p. pusillus Benguela, South Africa 1974-1985 David 1987 

A. p. pusillus Benguela, South Africa 1974-1985 David 1987 

Cephalorhynchus heavisidii Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

Delphinus delphis Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

Kogia breviceps Benguela, South Africa 1975-1990 Sekiguchi et al. 1992 

K. sima Benguela, South Africa 1975-1988 Sekiguchi et al. 1992 

Lagenorhynchus obscurus Benguela, South Africa 1969-1990 Sekiguchi et al. 1992 

 84 
Stenella coeruleoalba Benguela, South Africa 1975-1989 Sekiguchi et al. 1992 

Tursiops truncatus Benguela, South Africa 1975-1989 Sekiguchi et al. 1992 

0

1

2

3

4

5

 0-5 10-15 20-25 30-35 40-45

Length (cm)

Frequency

n = 10, mean of means = 20.84

Figure 40.--Frequency distribution of mean size of Merluccius capensis and M. paradoxus 

 consumed by marine mammals. 

 85 
Nototheniidae: Pleuragramma antarcticum, Antarctic silverfish; 25.91 n = 11,  ̄x = 14.40 cm

Antarctic silverfish reach a maximum size of 25.91 cm (maximum reported size from

Burns et al. 1998). The size of Antarctic silverfish consumed by marine mammals is based on 

food habits studies of four species of pinnipeds (Table 32). The modal mean size of Antarctic 

silverfish eaten by marine mammals is 10-15 cm, with an overall mean of means of 14.40 cm

(Fig. 41). 

Table 32.--Predators for which mean size of Pleuragramma antarcticum is known from sample 

sizes of 10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus gazella South Shetland Islands, S Atlantic 1992 Daneri 1996 

A. gazella South Shetland Islands, S Atlantic 1993-1994 Daneri and Carlini 1999 

Hydrurga leptonyx Davis Station, Antarctica 1984 Green and Williams 1986

Leptonychotes weddelii McMurdo Sound, Antarctica 1989-1993 Burns et al. 1998 

L. weddelii Mawson, Antarctica 1983-1985 Green and Burton 1987 

L. weddelii McMurdo Sound, Antarctica 1983-1985 Green and Burton 1987 

L. weddelii Davis Station, Antarctica 1983-1985 Green and Burton 1987 

L. weddelii Weddell Sea (eastern shore), Antarctica 1983, 1985 Plotz 1986 

L. weddelii Weddell Sea (southern shore), Antarctica 1983, 1985 Plotz 1986 

L. weddelii Weddell Sea, Antarctica 1998 Plotz et al. 2001 

Mirounga leonina South Shetland Islands, S Atlantic 1993-2000 Daneri and Carlini 2002 

 86 
0

1

2

3

4

5

6

7

8

 0-5 10-15 20-25 30-35 40-45 50-55

Length (cm)

Frequency

n = 10, mean of means = 14.40

Figure 41.--Frequency distribution of mean size of Pleuragramma antarcticum consumed by 

 marine mammals. 

 87 
Channichthyidae: Champsocephalus gunnari, mackerel icefish; n = 10,  ̄x = 24.85 cm

Mackerel icefish reach a maximum size of 66 cm (Froese and Pauly 2003), however Gon 

and Heemstra (1990) indicate that fish from Heard and Kerguelen Islands reach a maximum size 

of 45 cm. The size of mackerel icefish consumed by marine mammals is based on reports of 

several food habits studies of Arctocephalus gazella (Table 33). The frequency distribution of 

mean size of mackerel icefish eaten by A. gazella is strongly modal at 20-25 cm, with an overall 

mean of means of 24.85 cm (Fig. 42). 

Table 33.--Predators for which mean size of Champsocephalus gunnari is known from sample 

sizes of 10 or more. 

Predator Sampling Region Dates Citation 

Arctocephalus gazella Heard Island, Indian Ocean 1987-1988 Green et al. 1989 

A. gazella Heard Island, Indian Ocean 1993-1993 Green et al. 1997 

A. gazella Heard Island, Indian Ocean 1993-1993 Green et al. 1997 

A. gazella Iles Kerguelen, Indian Ocean 1998-2000 Lea et al. 2002 

A. gazella South Georgia Island, S Atlantic 1983 North 1996 

A. gazella South Georgia Island, S Atlantic 1993 Reid 1995 

A. gazella South Georgia Island, S Atlantic 1992 Reid 1995 

A. gazella South Georgia Island, S Atlantic 1991 Reid and Arnould 1996 

A. gazella South Georgia Island, S Atlantic 1993 Reid and Arnould 1996 

A. gazella South Georgia Island, S Atlantic 1994 Reid and Arnould 1996 

 88 
0

1

2

3

4

5

6

7

8

 0-5 10-15 20-25 30-35 40-45

Length (cm)

Frequency

n = 10, mean of means = 24.85

Figure 42.--Frequency distribution of mean size of Champsocephalus gunnari consumed by 

 marine mammals. 

 89 
Overall Patterns in the Size of Prey Consumed By Marine Mammals 

The species-specific examinations of prey size consumed by marine mammals suggest 

that the majority of prey consumed is smaller than 30 cm, as exemplified in the studies

summarized above (Figs. 9-28). This conclusion is further supported when the entire set of data 

for mean prey size is expanded to include the collective set of data that meet our sample size 

criteria (Fig. 43). Indeed, of the 726 cases plotted in Figure 43, 493 (68%) are ≤ 20 cm, while 

625 (86%) are ≤ 30 cm. 

0

10

20

30

40

50

60

70

80

90

100

0-5 10-15 20-25 30-35 40-45 50-55 60-65 70-75 80-85 90-95 ≥100

Mean prey length (cm)

Frequency

cetaceans: n = 343; mean of means = 18.45

pinnipeds: n = 383; mean of means = 16.89

Figure 43.--Frequency distribution of prey size means for cases based on all samples from

 marine mammal food habits studies with sample sizes ≥ 10. 

Evaluation of Biases 

Several potential biases were listed previously. One of these is that our minimum sample 

size requirement might inadvertently bias against large-bodied prey items. Intuitively, marine 

mammals can consume fewer large prey items than small prey items in any given foraging trip. 

To evaluate this potential bias, the LF of prey size was generated for rare taxa (defined here as 

 90 
having samples sizes from 1to 3). The data were generated as follows: for taxa where prey size 

was based on a single specimen, the data were transcribed directly. For taxa where prey size was 

based on two specimens, the sizes of the individual specimens contributing to that sample could 

only be calculated if the lower and upper ends of the size range were reported. Finally, 

individual prey size was calculated for cases where n = 3, but only if mean size was reported 

along with the lower and upper ends of the size range. 

There is no clear pattern to the differences in means (Table 34). Note, however, that the 

LF distributions are broadly similar to the distributions of means based on large sample sizes in 

all cases (Figs. 44 – 46 in comparison with Fig. 43). 

Table 34.--Mean prey size based on large (n ≥ 10) versus small (n = 1-3) sample sizes. 

cephalopods fish

 large small large small ceph diffs fish diffs 

odontocetes 19.90 14.61 18.55 15.88 5.29 2.67

otariids 12.86 11.80 17.35 17.50 1.06 -0.15

phocids 15.77 20.07 20.11 23.44 -4.30 -3.33

 91 
0

2

4

6

8

10

12

14

16

18

20

5 15 25 35 45 55 65 75 85 ≥100

Length (cm)

Frequency

cephalopods: n = 82; mean = 14.61

fish: n = 33; mean = 15.88

Figure 44.--Length frequency distribution of individual prey items consumed by odontocete 

 whales. 

0

2

4

6

8

10

12

14

16

5 15 25 35 45 55 65 75 85 ≥100

Length (cm)

Frequency

cephalopods: n = 34; mean = 11.80

fish: n = 46; mean = 17.50

Figure 45.--Length frequency distribution of individual prey items consumed by otariid seals. 

 92 
0

1

2

3

4

5

6

7

8

5 15 25 35 45 55 65 75 85 ≥100

Length (cm)

Frequenc

y

cephalopods: n = 16; mean = 20.07

fish: n = 31; mean = 23.44

Figure 46.--Length frequency distribution of individual prey items consumed by phocid seals. 

A second potential bias relates to the methodology of the majority of food habits studies, 

wherein prey size is estimated based on regressions relating body size to the size of hard parts 

(fish bones and otoliths, and cephalopod beaks) that 1) survive the digestive systems of marine 

mammals and 2) are recovered in the laboratory. 

The issue of differential recovery as a source of bias is important, as illustrated by Reid 

and Arnould’s (1996) study of the food habits of lactating female Antarctic fur seals at South 

Georgia. The samples from 1991 were sorted visually without the aid of magnification, and they 

found no mackerel icefish (Champsocephalus gunnari) otoliths from fish smaller than 26 cm --

fish that are approximately 1 year old. In subsequent years, they sorted samples with the aid of a 

stereomicroscope and found that otoliths from fish smaller than 26 cm comprised 77% of the 

sample in 1993 and 7% in 1994. Clearly, year-class strength could be playing a role in this 

 93 
example. However, the point remains that even if small otoliths survive the digestive process, 

they are more likely to be overlooked than are the larger specimens. 

Thus, prey size estimates can be too large if small elements are not recovered or are too 

badly eroded for identification and/or measurement (Reid and Arnould 1996). However, 

estimates of prey size can also be too small if prey are incompletely consumed (e.g., if heads are 

discarded in favor of body parts with more protein and fat) and/or the specific elements used for 

the regressions are reduced in size through partial or complete digestion (Bowen 2000, Tollit et 

al. 2004a, 2004b; Zeppelin et al. 2004). Finally, prey size estimates can also be too small if 

large skeletal elements are differentially regurgitated rather than passed through the digestive 

system, thus failing to appear in samples restricted to fecal material (Kiyota et al. 1999, 

Gudmundson et al. in press). 

If we assume that all the prey taxa that are part of the diet are recovered and correctly 

identified, then the distribution of the maximum reported sizes of those prey taxa observed in the 

diet can provide an indication of the patterns of size selectivity at a very general level. Keeping 

in mind that our database is limited to those prey taxa for which targeted size was reported, the 

overall distribution of maximum potential prey size is remarkably similar to the distribution of

targeted prey means. This is true regardless of whether all prey entries (482 cephalopods; 637 

fish) are plotted (Figs. 47 and 48), or only the list of non-overlapping prey taxa (140 cephalopod 

taxa; 222 fish taxa) is plotted (Figs. 49 and 50). Although the modal value of maximum

potential size is larger than the modal size of prey means (35-40 cm versus 5-10 cm), the overall 

pattern suggests that, in general, marine mammals target a fairly consistent size category of prey 

the majority of the time. 

 94 
0

10

20

30

40

50

60

70

80

5 25 45 65 85 105 125 145 165 185 ≥200

Dorsal mantle length (cm)

Frequenc

y

n = 482, mean = 46.08

Figure 47.--Frequency distribution of maximum potential cephalopod size for all taxonomic 

 entries consumed by marine mammals. 

0

10

20

30

40

50

60

70

80

5 25 45 65 85 105 125 145 165 185 ≥200

Length (cm)

Frequenc

y

n = 637, mean = 60.43

Figure 48.--Frequency distribution of maximum potential fish size for all taxonomic entries 

 consumed by marine mammals. 

 95 
0

5

10

15

20

25

5 25 45 65 85 105 125 145 165 185 ≥200

Dorsal mantle length (cm)

Frequenc

y

n = 140, mean = 40.53

Figure 49.--Frequency distribution of maximum potential size for non-overlapping cephalopod 

 prey taxa consumed by marine mammals. 

0

5

10

15

20

25

5 25 45 65 85 105 125 145 165 185 ≥200

Length (cm)

Frequenc

y

n = 222, mean = 55.25

Figure 50.--Frequency distribution of maximum potential size for non-overlapping fish prey taxa 

 consumed by marine mammals. 

 96 
Relationship Between Predator Size and Prey Size 

Despite the fact that the majority of prey targeted by marine mammals is smaller than 30 

cm (Fig. 43), examination of the relationship between the size of the predator and mean size of 

their prey shows a significant positive correlation (P < 0.001). This is true for all predator 

species considered together (Fig. 51), as well as for odontocete whales (Fig. 52, 

P < 0.001) and otariid seals (Fig. 53, P < 0.001) considered separately. Surprisingly, the 

relationship is not significant for phocid seals (Fig. 54; P = 0.87). 

y = 1.537x + 13.145

R2

 = 0.1398

0

20

40

60

80

100

120

140

0 5 10 15 20

Predator length (m)

Mean prey length (cm)

Figure 51.--Relationship between predator length and mean prey length for 699 cases, all marine 

 mammal species combined (P < 0.001). 

 97 
y = 1.7757x + 10.341

R2

 = 0.2104

0

20

40

60

80

100

120

140

0 5 10 15 20

Predator length (m)

Mean prey length (cm)

Figure 52.--Relationship between predator length and mean prey length for 146 odontocete 

 entries (P < 0.001). 

. 

y = 8.8329x + 4.4547

R2

 = 0.1286

0

10

20

30

40

50

60

70

80

90

0.0 0.5 1.0 1.5 2.0 2.5 3.0

Predator length (m)

Mean prey length (cm)

Figure 53.--Relationship between predator length and mean prey length for 237 otariid entries 

 (P < 0.001). 

 

 98 
y = 0.2115x + 18.207

R2

 = 0.0002

0

10

20

30

40

50

60

0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0

Predator length (m)

M

ean prey length (c

m)

Figure 54.--Relationship between predator length and mean prey length for 146 phocid entries 

 (P = 0.87). 

Relationship Between Mean Size of Prey Consumed by Marine Mammals and Maximum

Reported Prey Size 

Although the relationship between predator size and prey size is significant in most cases, 

recall from the information presented above that the overall mean prey size is relatively small for 

each of these groups of predators when compared to the maximum reported size of each prey 

species. Examination of the relationship between mean prey size targeted by marine mammals 

and the maximum reported size of the prey species shows that the two are only weakly 

correlated. Consider, for example, the case of gray seals (Halichoerus grypus) foraging around 

Anticosti Island (Benoit and Bowen 1990). The largest taxon they prey upon is the Atlantic cod 

(Gadus morhua), with a maximum reported length of 200 cm (Froese and Pauly 2003). 

 99 
However, gray seals restrict their consumption to cod ranging from 6.7 to 79.7 cm, with a mean 

size of 28.2 cm (Fig. 55). We will evaluate the degree to which that reflects size selectivity 

versus the relative availability of different size classes of cod when we consider the available 

survey data below. 

T.a. C.h. M.b.

Sa.sa.

Ma.am.

R.h.

P.v.

M.v.

C.l.

H.p.

G.m.

Me.ae. Sc.sc.

0

50

100

150

200

Prey size (cm)

Increasing maximum reported size

Figure 55.--Relationship between targeted size (shown as mean and range) and increasing 

maximum reported size (shown as dash above each prey entry) for various prey taxa 

consumed by gray seal (Halichoerus grypus, from Benoit and Bowen 1990). Key to 

prey species abbreviations: M.v. = Mallotus villosus, T.a. = Tautogolabrus 

adspersus, C.h. = Clupea harengus, C.l. = Cyclopterus lumpus, Sc.sc. = Scomber 

scombrus, M.b. = Merluccius bilinearis, R.h. = Reinhardtius hippoglossoides, H.p. = 

Hippoglossoides platessoides, Me.ae. = Melangrammus aeglefinus, Ma.am. = 

Macrozoarces americanus, P.v. = Pollachius virens, Sa.sa. = Salmo salar, G.m. = 

Gadus morhua. 

 100 
There is a clear positive relationship between mean size and maximum reported size for 

prey consumed by gray seals (Fig. 56), but it appears to be a curvilinear relationship with mean 

prey size reaching an asymptote well below the maximum reported size for the larger prey 

species. We have characterized this asymptotic relationship using the Holling disk equation 

(Holling 1959), which has the form: 

bx

ax

y

+ = 1 , 

where y = mean prey size and x = maximum reported prey size. The shape parameters a and b 

were estimated in SPSS using non-linear regression (Norusis 1998). In addition to the parameter 

estimates, SPSS provides the asymptotic standard errors of the parameter estimates (Table 35). 

These will be examined in more detail below in comparison with commercial fisheries.

 101 
Table 35.--Parameter estimates for non-linear estimation of the Holling disk equation (Holling 1959) describing 

between mean prey size and the maximum reported size of each prey taxon, for pairings of different pprey types. Lower and upper CI = 95% confidence limits. 

Pairing n Parameter

Parameter 

estimate 

Asymptotic 

Standard error Lower C

gray seals X fish 13 A 1.7489 1.5502 -1.6631 gray seals X fish 13 B 0.0438 0.0492 -0.0646 Cephalopods, all prey entries 275 A 0.4915 0.0264 0.4395 Cephalopods, all prey entries 275 B 0.0033 0.0006 0.0022 Odontocetes X Cephalopods 219 A 0.5037 0.0319 0.4408 Odontocetes X Cephalopods 219 B 0.0034 0.0007 0.0021 Odontocetes X Cephalopods, 600 cm outlier removed 218 A 0.5575 0.0439 0.4711 Odontocetes X Cephalopods, 600 cm outlier removed 218 B 0.0052 0.0012 0.0028 Otariids X Cephalopods 38 A 0.7197 0.2466 0.2196 Otariids X Cephalopods 38 B 0.0225 0.0160 -0.0099 Phocids X Cephalopods 18 A 1.5031 1.1567 -0.9489 Phocids X Cephalopods 18 B 0.0612 0.0697 -0.0865 Fish, all prey entries 417 A 0.8678 0.0791 0.7123 

102
Fish, all prey entries 417 B 0.0227 0.0032 0.0163 

Odontocetes X Fish 114 A 0.8230 0.1676 0.4910 Odontocetes X Fish 114 B 0.0199 0.0069 0.0063 Otariids X Fish 181 A 0.7283 0.0766 0.5772 Otariids X Fish 181 B 0.0132 0.0027 0.0078 Phocids X Fish 120 A 0.8828 0.1520 0.5817 Phocids X Fish 120 B 0.0275 0.0067 0.0143 

103
0

10

20

30

40

50

60

70

0 50 100 150 200 250

Maximum reported prey size (cm)

Mean prey size (cm)

Figure 56.--Relationship between mean size of consumed prey and maximum reported size of 

 individuals in the populations of 13 prey species eaten by gray seals. The curve 

 describing the relationship is based on the Holling disk equation (Holling 1959), 

 estimated using non-linear regression. 

104
0

20

40

60

80

100

120

140

160

180

200

0 100 200 300 400 500 600 700

Maximum reported prey size (cm)

Mean prey size (cm)

Figure 57.--Relationship between mean size of consumed prey and maximum reported size of 

 individuals in the populations of 219 taxonomic entries of cephalopods eaten by 

 odontocete whales. The curve describing the relationship is based on the Holling disk 

 equation (Holling 1959), estimated using non-linear regression.

105
0

20

40

60

80

100

120

0 50 100 150 200 250 300

Maximum reported prey size (cm)

Mean prey size (cm)

Figure 58.--Relationship between mean size of consumed prey and maximum reported size of 

individuals in the populations of 218 taxonomic entries of cephalopods eaten by 

odontocete whales, minus one 600 cm outlier (Architeuthis dux). The curve 

describing the relationship is based on the Holling disk equation (Holling 1959), 

estimated using non-linear regression.

106
0

5

10

15

20

25

30

35

0 20 40 60 80

Maximum reported prey size (cm)

Mean prey size (cm)

100

Figure 59.--Relationship between mean size of consumed prey and maximum reported size of 

 individuals in the populations of 38 taxonomic entries of cephalopods eaten by 

 otariid seals. The curve describing the relationship is based on the Holling disk 

 equation (Holling 1959), estimated using non-linear regression.

107
0

5

10

15

20

25

30

35

40

45

50

0 20 40 60 80

Maximum reported prey size (cm)

Mean prey size (cm)

100

Figure 60.--Relationship between mean size of consumed prey and maximum reported size of 

 individuals in the populations of 18 taxonomic entries of cephalopods eaten by 

 phocid seals. The curve describing the relationship is based on the Holling disk 

 equation (Holling 1959), estimated using non-linear regression.

108
0

10

20

30

40

50

60

70

80

90

100

0 50 100 150 200 250

Maximum reported prey size (cm)

Mean prey size (cm)

Figure 61.--Relationship between mean size of consumed prey and maximum reported size of 

 individuals in the populations of 114 taxonomic entries of fish eaten by odontocete 

 whales. The curve describing the relationship is based on the Holling disk equation 

 (Holling 1959), estimated using non-linear regression.

109
0

10

20

30

40

50

60

70

80

90

100

0 50 100 150 200 250

Maximum reported prey size (cm)

Mean prey size (cm)

walleye pollock

Figure 62.--Relationship between mean size of consumed prey and maximum reported size of 

 individuals in the populations of 181 taxonomic entries of fish eaten by otariid seals. 

 The curve describing the relationship is based on the Holling disk equation (Holling 

 1959), estimated using non-linear regression. The series of points at 91 cm 

 maximum reported size represents walleye pollock (Theragra chalcogramma). 

110
0

10

20

30

40

50

60

70

80

90

100

0 50 100 150 200 250

Maximum reported prey size (cm)

Mean prey size (cm)

Atlantic cod and

Atlantic ling

Figure 63.--Relationship between mean size of consumed prey and maximum reported size of 

 individuals in the populations of 120 taxonomic entries of fish eaten by phocid seals. 

 The curve describing the relationship is based on the Holling disk equation (Holling 

 1959), estimated using non-linear regression. The series of points at 200 cm 

 maximum reported size represents Atlantic cod (Gadus morhua) and Atlantic ling 

 (Molva molva). 

As each of the preceding figures (Figs. 57 – 63) shows, the size of prey targeted by 

marine mammals is only weakly correlated with the maximum reported size of each prey taxon. 

This is particularly evident as prey size increases above 100 cm. That is, the majority of targeted 

prey with maximum size of 100 cm or larger tends to average under 50 cm. This relationship 

(which we will call a functional response, different from the functional response involving prey 

density/availability typical of other literature) is likely due to a complex interaction of factors, 

including swimming speed and gape size of the predator (Nilsson and Brönmark 2000, Scharf et 

al. 2000), swimming speed and body depth of the prey (Nilsson and Brönmark 2000), and size 

111
composition of the available prey. However, we cannot evaluate the extent to which these 

patterns represent size selectivity on the part of the predators until we have examined the 

available commercial and survey data in the following sections, which will provide an indication 

of the size composition of the prey available. 

Another feature evident from the preceding figures, and figures to follow, is the distinct 

clustering of points at certain sizes along the x-axis. Specifically, there is a cluster at 91 cm

maximum reported size corresponding to walleye pollock (Theragra chalcogramma; Fig. 62) 

and 200 cm maximum reported size corresponding to Atlantic cod (Gadus morhua) and Atlantic 

ling (Molva molva; Fig. 63). These clusters re-emphasize that when enough data are available 

for a particular prey species or species group, distinct patterns emerge (also illustrated in Figs. 

30-42) showing the central tendency, as well as the variability around that central tendency, in 

the size selectivity of marine mammals. These clusters of points can be used to inform on singlespecies applications of the management approach advocated here.

112
RESULTS: COMPARISONS WITH COMMERCIAL CATCH DATA 

The ease with which we were able to locate commercial data that corresponded with a 

particular food habits study relates to the sampling of the food habits study, the prey species 

involved, and the various ways in which commercial data are presented. Food habits studies 

conducted over a single season provide easy comparisons, as long as the commercial and survey 

data were also generated that same season in the same region. In contrast, some food habits 

studies span multiple sampling seasons, resulting in time-averaged values for the size

composition of targeted prey. Consequently, any commercial data used for comparison should, 

to the extent possible, span the same temporal period as the food habits studies. In many cases 

this is clearly impossible to achieve. As a result, we have been able to draw direct comparisons 

between the size selectivity of marine mammals and commercial fisheries in relatively few cases 

(118 of 1,138 entries: see Appendix 5). 

There are multiple ways of comparing food habits data with commercial data. In the 

context of interactions between commercial fishing and the food habits of birds, Duffy and 

Schneider (1994) have outlined five specific methods that have been used in the literature to 

quantify levels of competition. Of these, we will focus on two: the “Horn Ratio,” which 

calculates the overlap in species composition, and a modification of the “Schaefer Ratio,” which 

calculates the total removals of a prey species by a predator relative to the total catch of the 

commercial fishery. 

Species composition is one measure of overlap between commercial and predator

interests, and has arguably been either the explicit or implicit focus of the majority of food habits 

studies to date (Nilssen et al. 1990, Kaschner and Pauly 2004, Baraff and Loughlin 2000). With 

regard to the list of prey taxa compiled here, it may be the case that because we have limited our 

113
compilation to only those taxa for which prey size is available, we may be providing a biased 

species list. For instance, commercially unimportant taxa may not have been studied sufficiently 

to have regressions for calibrating body size estimates from measurements of preserved prey 

remains. 

Examination of what is known about the food habits of northern fur seals (Callorhinus 

ursinus) suggests that this is not a significant source of bias. We have compiled prey size data 

for 28 non-overlapping prey species consumed by northern fur seals. Of these, 19, or 68%, are 

commercially important. For comparison, Kajimura’s (1984) summary of food habits studies 

based on analysis of over 16,000 stomach samples listed 73 non-overlapping prey species for 

northern fur seals but did not present data on prey size. Of these, 48, or 66%, are commercially 

important. Thus, limiting our analysis to prey taxa for which size selectivity data are available 

does not appear to be biased towards commercially important taxa. 

Our database contains data from a large number of independent studies, many of which 

cover the same predator and/or prey species. Consequently, we have a measure of how often 

various prey species are listed in food habits studies (a modified form of frequency of 

occurrence). We can also distill this data to include only the non-overlapping taxa. Either way, 

the results are broadly similar (Table 36). When all taxonomic entries are considered, 756/1,166, 

or 65% are, or potentially are, commercially harvested (Appendix 3). When only the nonoverlapping taxa are considered, that number drops to 57% (218/380). 

114
Table 36.--Levels of commercial importance for prey species in this study. Crustacean data from

original food habits studies. Cephalopod data from Roper et al. 1984 and Nesis 1987. 

Fish data from FishBase (Froese and Pauly 2003). 

Level of All taxonomic entries Non-overlapping taxonomic entries

commercial 

importance crustaceans cephalopods fish crustaceans cephalopods fish

potential -- 98 10 -- 24 4 

minor -- 15 120 -- 5 43 

yes 22 115 360 6 25 110

no 12 221 140 11 81 61

indet. -- 20 5 -- 3 -- 

sub-total 34 469 635 17 138 218

TOTAL 1138 373

We also examine the overlap between food habits and commercial fisheries by examining 

modifications of the Schaefer Ratio (Duffy and Schneider 1994). Although we will generally be 

comparing total removals, we will be using size-structured models to do so. Specifically, we will 

focus on a comparison between the targeted prey size of marine mammal food habits and the size 

composition of the commercial take of the same prey species. We will do this in two ways. 

First, we will draw a comparison between the mean prey size taken by marine mammals and the 

mean size of the commercial catch. Second, we will evaluate the degree of overlap in the LF 

distributions for marine mammals compared to commercial fisheries. 

115
Keeping in mind the caveats discussed previously concerning the appropriateness of 

using mean prey size as the measure of central tendency of size in distributions that are often 

asymmetrical, the tendency for the size distributions of commercial catch to be asymmetric were 

evaluated. We have not calculated any of the shape statistics for the commercial data. However, 

we calculated the same index of symmetry as for the food habits data: the mean is subtracted 

from the midpoint of the range. When this is done for 118 entries for commercial catch data 

(each of which corresponds to a food habits entry), the results are broadly similar to what was 

seen for the food habits data (see Fig. 7), but with a higher tendency for the mean to be less than 

the midpoint of the size range (i.e., positively skewed). Specifically, the overall mean difference 

between the midpoint of the size range of and the mean of commercial data is 2.69 cm 

(compared to 1.21 in the food habits size distributions). The mean is within ± 2 cm of the 

midpoint of the range in 51% of the cases (Fig. 64). In contrast, the mean is 2 cm or more below 

the midpoint (resulting in a positive difference, indicative of positive skew) in 41% of the cases 

while it is 2 cm or more above the midpoint (indicative of negative skew) in 8% of the cases. 

116
0

5

10

15

20

25

-10 -8 -6 -4 -2 0 2 4 6 8 ≥10

Midpoint of range minus mean

Percent of total

mean difference = 2.69 cm

Figure 64.--Distribution of absolute differences between midpoint of range of values and the 

 mean for 120 commercial catch size distributions. Compare this figure with 

 Figure 7. 

Regardless of whether or not the mean size of prey species is the most appropriate 

summary statistic to use, that is the form of the data that are most readily available from the food 

habits literature for comparison with commercial fisheries data. For our next analysis, we 

calculated the difference in mean prey size by subtracting the food habits mean from the 

commercial mean for pairings of data that are from the same region and the same time period. 

The mean difference in means is 7.37 cm, with the overwhelming majority (71%) being positive 

values. That is, the mean size of commercial catches was greater than mean size of prey found in 

the food habits of marine mammals (Fig. 65). This shows a clear tendency for commercial 

fisheries to target larger individuals than are being targeted by marine mammal predators. 

Furthermore, the magnitude of the difference increases significantly (P < 0.001) with increasing 

overall body size of the target species (Fig. 66). 

117
118

0

5

10

15

20

25

30

35

-50 -40 -30 -20 -10 0 10 20 30 40 50

Absolute difference, commercial mean minus food habits mean (cm)

Frequency

mean difference of means = 7.37 cm

Figure 65.--Distribution of differences in means (commercial mean minus food habits mean) for 

 120 prey entries. 

y = 0.1046x

R2

 = 0.2108

-60

-50

-40

-30

-20

-10

0

10

20

30

40

50

60

0 50 100 150 200 250

Maximum reported prey size (cm)

Difference in means (cm)

crustaceans: n = 5

cephalopods: n = 17

fish: n = 98

mean difference in means = 7.37 cm

T.c.

M.c.

G.m.

Figure 66.--Distribution of differences in means (commercial mean minus food habits mean) for 

 120 prey entries, as a function of the maximum reported size for each prey species. 

 Dashed horizontal line indicates the mean difference of means (7.37 cm). 

 Abbreviations in plot refer to prey taxa with multiple entries (T.c. = Theragra 

 chalcogramma, M.c. = Merluccius capensis, G.m. = Gadus morhua). 
Species-specific Patterns of Size Composition 

Examination of many of the previous figures shows several clusters of points with the 

same maximum reported prey size (e.g., Fig. 66). In some cases these clusters represent multiple 

prey taxa that coincidentally have the same maximum reported size. For example, the cluster at 

60 cm maximum reported prey size in Figure 66 is composed of data from three prey taxa: 

Agulhas sole (Austroglossus pectoralis), Atlantic mackerel (Scomber scombrus), and Cape horse 

mackerel (Trachurus trachurus capensis). However, some of the clusters are comprised solely, 

or predominantly, of one prey taxon (e.g., the cluster at 91 cm represents only walleye pollock, 

Theragra chalcogramma). The prey taxa represented by 10 or more cases are the same taxa that 

appeared in our discussion of patterns of predation by marine mammals on selected prey taxa 

(Figs. 30-42). 

Here we draw a direct comparison between the mean size of prey consumed by marine 

mammals (Fig. 30-42) and the corresponding commercial data. For many of the cases we have 

commercial data that match reasonably well in terms of spatial and temporal coverage. 

However, it is often the case that a single record of commercial catch must be used for 

comparison with multiple food habits entries. For example, our database (Appendix 2) contains

24 entries of mean prey size for odontocete whales stranded on the South Africa coast between 

1969 and 1990 (Sekiguchi et al. 1992). These data cover predation by eight species of 

odontocetes on seven prey taxa. However, because the food habits data were all collected from

the same general area and span approximately the same time period, a single source of data was 

used for comparison with the commercial catch (Sekiguchi et al. 1992). Consequently, the 

frequency distribution of means that we use here will include only independent commercial 

means. That is, rather than repeating the identical commercial catch data for each of the 

119
corresponding food habits studies, each commercial data point will only appear once in the 

following LF distributions. 

Of the 15 entries of mean prey size we have for krill (Euphausia superba), we were able 

to locate only two cases of commercial catch data that correspond temporally and spatially with 

the food habits studies (Fig. 67). These two cases fall within the distribution of mean krill size 

targeted by marine mammals. 

0

1

2

3

4

5

6

7

8

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

Total length (cm)

Frequency

food habits n = 15

commercial n = 2

Figure 67.--Frequency distribution of mean size of krill (Euphausia superba) targeted by marine 

 mammals (solid bars) versus commercial fisheries (hatched bars).

120
0

1

2

3

4

5

6

7

 0-5 10-15 20-25 30-35 40-45 50-55

Dorsal mantle length (cm)

Frequency

food habits n = 10

commerical n = 3

Figure 68.--Frequency distribution of mean size of market squid (Loligo opalescens) targeted by 

 marine mammals (solid bars) versus commercial fisheries (hatched bars). 

The situation is similar for market squid (Loligo opalescens), with only 3 of the 10 food 

habits entries with comparable commercial data available for comparison (Fig. 68). As with 

krill, the mean size of the commercial catch of market squid is broadly comparable with that of 

marine mammals. However, it is important to note that the mean size of the commercial catch 

declined steadily over the latter half of the 20th century (Leos 1998). 

The situation for Cape Horn squid (L. vulgaris reynaudii) is also similar to the first two 

examples. The mean size of the commercial catch is similar to the mean size targeted by marine 

mammals (Fig. 69). One of the two commercial catch records (10.52 cm) corresponds well with 

eleven separate food habits prey entries (Appendix 5). 

121
0

1

2

3

4

5

 0-5 10-15 20-25 30-35 40-45 50-55

Dorsal mantle length (cm)

Frequency

food habits n = 11

commercial n = 2

Figure 69.--Frequency distribution of mean size Cape Horn squid (Loligo vulgaris reynaudii) 

 targeted by marine mammals (solid bars) versus commercial fisheries (hatched bars). 

The case of Atlantic herring (Clupea harengus), in contrast to the examples above, shows 

hints of a difference between the commercial catch data and the mean size targeted by marine 

mammals (Fig. 70). Although we only have two independent cases of commercial data, they 

correspond to 8 of the 10 food habits entries (Appendix 5), showing a slight tendency for 

commercial fisheries to target larger herring than do marine mammals. 

122
0

1

2

3

4

5

6

 0-5 10-15 20-25 30-35 40-45 50-55

Length (cm)

Frequency

food habits n = 10

commercial n = 2

Figure 70.--Frequency distribution of mean size of Atlantic herring (Clupea harengus) targeted 

 by marine mammals (solid bars) versus commercial fisheries (hatched bars). 

We located 3 commercial catch records of pilchard (Sardinops sagax) for comparison 

with 7 of the 11 food habits entries (Fig. 71; Appendix 5). As with the commercial targeting of 

herring, there is a slight tendency for commercial fisheries to target larger pilchard than do 

marine mammals.

123
0

1

2

3

4

5

6

7

8

 0-5 10-15 20-25 30-35 40-45 50-55

Length (cm)

Frequenc

y

food habits n = 11

commercial n = 3

Figure 71.--Frequency distribution of mean size of pilchard (Sardinops sagax) targeted by 

 marine mammals (solid bars) versus commercial fisheries (hatched bars). 

A clear indication of a difference between the mean size of prey targeted by commercial 

fisheries and marine mammals is seen in the case of Atlantic cod (Gadus morhua; Fig. 72). The 

2 commercial data points correspond to 5 of the 19 food habits cases, with the mean commercial 

size larger than the food habits mean in every case. 

124
0

1

2

3

4

5

6

7

 0-5 10-15 20-25 30-35 40-45 50-55

Length (cm)

Frequency

food habits n = 19

commercial n = 2

Figure 72.--Frequency distribution of mean size of Atlantic cod (Gadus morhua) targeted by 

 marine mammals (solid bars) versus commercial fisheries (hatched bars). 

Walleye pollock (Theragra chalcogramma) is one of the few prey species for which we 

have a substantial number (14) of independent commercial cases for comparison with the food 

habits data. Indeed, the temporal and spatial coverage of the commercial data corresponds with 

19 of the 25 food habits entries. Recall from the discussion above that the distribution of means 

targeted by marine mammals has strong modes at 10-15 cm and 25-30 cm, with a weaker mode 

at 45-50 cm. The distribution of commercial means has a strong mode at 30-35 cm, with a 

weaker mode at 40-45 cm (Fig. 73). In the 19 pair-wise comparisons between the commercial 

mean and the food habits mean, the commercial mean is larger than the food habits mean in 18 

cases, with an overall mean difference in means of 17.36 cm (Fig. 74; Appendix 5). The single 

case where the food habits mean was larger than the commercial mean is for Steller sea lions 

(Eumetopias jubatus) sampled in the Pribilof Islands in 1976 and 1979 (Frost and Lowry 1986). 

In those 2 years, the mean size of pollock targeted by Steller sea lions was 46.9 cm. In contrast, 

125
the mean size of the commercial catch in the eastern Bering Sea in 1976 was 33.1 cm (Bakkala 

1989). 

0

1

2

3

4

5

6

7

8

9

10

 0-5 10-15 20-25 30-35 40-45 50-55

Length (cm)

Frequency

food habits n = 25

commercial n = 14

Figure 73.--Frequency distribution of mean size of walleye pollock (Theragra chalcogramma) 

 targeted by marine mammals (solid bars) versus commercial fisheries (hatched bars). 

126
0

1

2

3

4

5

6

 -15 to -10 -5 to 0 5 to 10 15 to 20 25 to 30 35 to 40

Commercial mean minus food habits mean (cm)

Frequency

mean difference = 17.36 cm

Figure 74.--Distribution of differences in means (commercial mean minus food habits mean) for 

 19 prey entries for walleye pollock (Theragra chalcogramma). 

All of the data for marine mammal predation on Cape hakes (Merluccius spp.) derive 

from the Benguela Current of South Africa (n = 10), spanning the years 1969 to 1990 (David 

1987, Sekiguchi et al. 1992). Thus, as with the case of Cape Horn squid, a single, time-averaged 

value for the period 1984 to 1986 (Fig. 75) has been used for the commercial catch (Payne et al. 

1985, 1986; Sekiguchi et al. 1992). This value (27.19 cm) is situated in the upper end of the 

distribution of means from food habits studies. However, the difference of means in the pairwise comparisons is positive in 8 of the 10 cases, with an overall mean difference of 6.35 cm

(Fig. 76). 

127
0

1

2

3

4

5

6

 0-5 10-15 20-25 30-35 40-45 50-55

Length (cm)

Frequenc

y

food habits n = 10

commercial n = 1

Figure 75.--Frequency distribution of mean size of Cape hakes (Merluccius spp.) targeted by 

 marine mammals (solid bars) versus commercial fisheries (hatched bars). 

0

1

2

3

4

 -4 - -2 0-2 4-6 8-10 12-14

Commercial mean minus food habits mean (cm)

Frequency

mean difference = 6.35 cm

Figure 76.--Distribution of differences in means (commercial mean minus food habits mean) for 

 10 prey entries for Cape hakes (Merluccius spp.). 

128
The final example we use to show the relationship between the mean size of prey targeted 

by marine mammals and the mean size taken by commercial fisheries is that of the mackerel 

icefish (Champsocephalus gunnari). For this species, we have 3 independent cases of 

commercial data (Fig. 77) corresponding to 4 of the 10 food habits cases. Generally speaking, 

the commercial means fall within the distribution of food habits means. However, in each of the 

four pair-wise comparisons, the commercial mean is larger than the food habits mean (mean 

difference = 4.9 cm). 

0

1

2

3

4

5

6

7

8

 0-5 10-15 20-25 30-35 40-45 50-55

Length (cm)

Frequenc

y

food habits n = 10

commercial n = 3

Figure 77.--Frequency distribution of mean size of mackerel icefish (Champsocephalus gunnari) 

 targeted by marine mammals (solid bars) versus commercial fisheries (hatched bars). 

Relationship Between Mean Size of Commercial Catch and Maximum Reported Size 

Recall from the previous discussion that the size of prey targeted by marine mammals is 

positively correlated in a non-linear fashion with the maximum reported size that a particular 

prey species attains. Specifically, as maximum prey size increases, the mean size targeted by 

marine mammals increases asymptotically, but to a point substantially smaller than the 

129
maximum size of the prey (Figs. 57-63). This functional response is likely to be constrained by a 

combination of factors, many of which probably do not apply to the capture techniques used in 

commercial fisheries. Thus, it is instructive to examine the same relationship for taxa for which 

there is a commercial fishery to determine the extent to which human removals are, or are not, 

similar to the removals of other mammalian predators. 

We use the same approach here as we did with the food habits data, plotting mean 

targeted size against maximum reported size for each target species. As before, this relationship 

is characterized with the Holling disk equation (Holling 1959) using non-linear regression in 

SPSS (Norusis 1998). Given the relatively small number of commercial fisheries cases we have 

found that correspond to specific food habits studies, we only have results for targeted species of 

fish3

. 

We have already shown a tendency for commercial fisheries to target larger individuals 

in any pair-wise comparison with food habits data. Thus, we would expect that the functional 

response curve for commercial fisheries would be higher than that of marine mammal predation. 

Indeed, plotting all available data (i.e., all pair-wise comparisons, plus food habits data for which 

we do not have comparable fisheries data) show that this is the case, with the commercial curve 

significantly higher than the food habits curves (Fig. 78; Table 37). The same pattern emerges 

when food habits data are considered separately for Odontocetes, Otariids, and Phocids (Figs. 

79-81; Table 38). 

 

3

 Although there are 19 entries for commercial catches of cephalopods, 10 of these are the same commercial catch 

record for Loligo vulgaris reynaudii repeated for comparison with multiple marine mammal predation reported in

Castley et al. 1991, Lipinski and David 1990, and Sekiguchi et al. 1992, all of which span the period of

approximately 1969-1990. 

130
0

10

20

30

40

50

60

70

80

90

0 50 100 150 200 250

Maximum reported length (cm)

Mean prey size (cm)

food habits

commercial

Figure 78.--Relationship between mean prey size and maximum reported size of 417 taxonomic 

 entries of fish eaten by marine mammals (diamonds, solid line) compared with 94 

 commercial fisheries cases (triangles, dashed line). The curves describing the 

 relationships are based on the Holling disk equation (Holling 1959), estimated using 

 non-linear regression. 

131
0

10

20

30

40

50

60

70

80

90

100

0 50 100 150 200 250

Maximum reported prey size (cm)

Mean prey size (cm)

food habits

commercial

Figure 79.--Relationship between mean prey size and maximum reported size of 114 taxonomic 

 entries of fish eaten by odontocetes (diamonds, solid line) compared with 27 

 commercial fisheries cases (triangles, dashed line). The curves describing the 

 relationships are based on the Holling disk equation (Holling 1959), estimated using 

 non-linear regression. 

132
0

10

20

30

40

50

60

70

80

90

100

0 50 100 150 200 250

Maximum reported prey size (cm)

Mean prey size (cm)

food habits

commercial

walleye pollock

Figure 80.--Relationship between mean prey size and maximum reported size of 181 taxonomic 

 entries of fish eaten by otariids (diamonds, solid line) compared with 44 commercial 

 fisheries cases (triangles, dashed line). The curves describing the relationships are 

 based on the Holling disk equation (Holling 1959), estimated using non-linear 

 regression. The series of points at 91 cm maximum reported size represents walleye 

 pollock (Theragra chalcogramma). 

133
0

10

20

30

40

50

60

70

80

90

100

0 50 100 150 200 250

Maximum reported prey size (cm)

Mean prey size (cm)

food habits

commercial

Atlantic cod and

Atlantic ling

Figure 81.--Relationship between mean prey size and maximum reported size of 120 taxonomic 

 entries of fish eaten by phocids (diamonds, solid line) compared with 23 commercial 

 fisheries cases (triangles, dashed line). The curves describing the relationships are 

 based on the Holling disk equation (Holling 1959), estimated using non-linear 

 regression. The series of points at 200 cm maximum reported size represents Atlantic 

 cod (Gadus morhua) and Atlantic ling (Molva molva). 

134
Table 37.--Parameter estimates for non-linear estimation of the Holling disk equation (Holling 1959) describing 

between mean fish size targeted by commercial fisheries and the maximum reported size of each prey commercial data paired with specific food habits studies. Lower and upper CI = 95% confidence lim

Pairing n Parameter

Parameter

estimate 

Asymptotic 

Standard error Lower CI Uppe

Fish, all commercial entries 94 a 0.917399 0.155654 0.608256 1.22Fish, all commercial entries 94 b 0.014262 0.004157 0.006006 0.02

Commercial X Fish X Odontocetes 27 a 1.285662 0.698803 -0.153550 2.724

Commercial X Fish X Odontocetes 27 b 0.028906 0.022554 -0.017544 0.07

Commercial X Fish X Otariids 44 a 0.598275 0.113277 0.369672 0.82

Commercial X Fish X Otariids 44 b 0.005124 0.002762 -0.000450 0.01

Commercial X Fish X Phocids 23 a 1.418855 0.291042 0.813599 2.024

Commercial X Fish X Phocids 23 b 0.025172 0.007162 0.010279 0.04

135 
Table 38.--Significance of statistical comparison of parameter estimates using two-sample t-test 

for each pair of parameters. Note that while the estimates of b in the pairing between 

Phocids and Commercial targeting of fish is not significant, the curves as a whole are. 

Pairing n Parameter

Parameter

estimate 

Asymptotic 

Standard error P-value

Fish, all prey entries 417 a 0.8678 0.0791 P < 0.01

Fish, all commercial entries 94 a 0.9174 0.1557

Fish, all prey entries 417 b 0.0227 0.0032 P < 0.01

Fish, all commercial entries 94 b 0.0143 0.0042

Odontocetes X Fish 114 a 0.8230 0.1676 P < 0.01

Commercial X Fish X Odontocetes 27 a 1.2857 0.6988

Odontocetes X Fish 114 b 0.0199 0.0069 P < 0.01

Commercial X Fish X Odontocetes 27 b 0.0289 0.0226

Otariids X Fish 181 a 0.7283 0.0766 P < 0.01

Commercial X Fish X Otariids 44 a 0.5983 0.1133

Otariids X Fish 181 b 0.0132 0.0027 P < 0.01

Commercial X Fish X Otariids 44 b 0.0051 0.0028

Phocids X Fish 120 a 0.8828 0.1520 P < 0.01

Commercial X Fish X Phocids 23 a 1.4189 0.2910

Phocids X Fish 120 b 0.0275 0.0067 P = 0.07

Commercial X Fish X Phocids 23 b 0.0252 0.0072

136
Overlap in Size Distributions 

If the commercial fishery tends to target larger individuals than predators do, then it 

stands to reason that the LF distributions will also be different. However, calculating the amount 

of overlap, or the degree to which they are different (or similar) is surprisingly complicated. 

Calculating the degree of overlap when the distributions do not intersect is straightforward 

enough because there is zero overlap (Fig. 82). This is the ideal situation sought under 

conventional management schemes, because under such a harvest regime there should be no 

(direct) competition between different stake holders (adding the ever-important caveat “as long 

as recruitment is not affected”; Frost and Lowry 1986). 

Figure 82.--Hypothetical situation where the LF distribution of the prey size targeted by marine 

 mammals does not overlap with the LF distribution of the commercial fishery. 

137
Although calculating the degree of overlap in this first example is deceptively simple, the 

situation changes quickly once the LF distributions intersect. Indeed, as we show below, there 

are at least six different ways to calculate overlap. Each of these will be discussed briefly. 

Of the six approaches to calculating overlap, there are two main categories: those that 

calculate the overlap two dimensionally by summing the area of overlap for the two curves; and 

those that calculate the overlap in only one dimension as the range of overlap along the axis of 

prey size. Both of these approaches can also be sub-divided further depending on whether they 

calculate the overlap with respect to the total size range of the prey species, the size range 

targeted by the fishery, or the size range targeted by the predator. 

To illustrate further, we demonstrate three cases where the upper end of the LF 

distribution of the predator overlaps with the lower end of the LF distribution of the fishery 

between 20 and 90 cm, with the maximum length of 160 cm for the hypothetical prey species. In 

the first example (Fig. 83), the overlap is calculated with respect to the total size range of the 

prey species. Thus, the area of overlap calculated in this manner is 24%, while the range of 

overlap is 44%. If the two curves overlaid one another perfectly, the area and the range of 

overlap would be 100%. 

136

 
Figure 83.--Overlap in size composition of a hypothetical prey species taken by both marine 

 mammals and commercial fisheries, considered with respect to the total size range of 

 the prey species. The area of the intersection of the two LF distribution curves 

 (shaded area) is 24%, while the range of overlap (a/b = 70/160 X 100) is 44%. 

If we examine the range of overlap from the standpoint of the fishery, the picture is much 

different. Using the same curves as in Figure 83, the two distributions still overlap between 20 

and 90 cm. However, if the area of overlap is considered with respect to the fishery (Fig. 84), 

32% of the size range targeted by the fishery is also targeted by the predator. Similarly, the two 

distributions overlap across 70 cm of a total size range of 140 cm targeted by the fishery, for a 

range of overlap of 50%. 

Finally, our hypothetical example changes yet again if we examine the overlap with 

respect to the size range targeted by the predator (Fig. 85). We still use the same curves, but 

from the standpoint of the predator, 85% of the area of the LF distribution of the predator’s size 

137

 
selectivity is also targeted by the commercial fishery. When examined at the one-dimensional 

level of the range of overlap, 78% of the size range targeted by the predator is also targeted by 

the fishery. 

Figure 84.--Overlap in size composition of a hypothetical prey species taken by both marine 

 mammals and commercial fisheries, considered with respect to the size range 

 targeted by the fishery. The area of the intersection of the two LF distribution curves 

 (shaded area) is 32%, while the range of overlap (a/b X 100 = 70/140 X 100) is 

 50%. 

 

138

 
Figure 85.--Overlap in size composition of a hypothetical prey species taken by both marine 

 mammals and commercial fisheries, considered with respect to the size range 

 targeted by the predator. The area of the intersection of the two LF distribution 

 curves (shaded area) is 85%, while the range of overlap (a/b X 100 = 70/90 X 100) is 

 78%. 

As our series of hypothetical examples illustrates, there are numerous approaches to 

quantifying overlap between the size selectivity of predators and commercial fisheries, all of 

which yield widely different values. With specific reference to the calculations of the area of 

overlap, it is important to note that interpretation of the data requires detailed information on the 

magnitude of each of the sources of removals. For example, if the fishery and predators are 

removing comparable amounts of biomass, interpretation of the area of overlap is 

straightforward. However, if information on total removals is lacking, clear interpretation of the 

area of overlap is difficult or impossible. In contrast, the calculations of the range of overlap are 

139

 
independent of the magnitude of the total removals. Although this approach suffers the loss of a 

significant source of information, the relationship between a and b in the examples above does 

not change with different vertical scaling. 

Because detailed information on total removals is not available for most food habits 

studies, we have chosen to focus our analyses on the range of overlap with respect to the total 

size of the prey, the size range taken by the fishery, and the size range taken by the predators. 

We will briefly examine overlap in terms of the area of intersection between LF distributions, but 

that will be discussed below in relationship to size structure of the available prey as indicated by 

fisheries survey data. 

Following the sequence used above, the first relationship we examine is between the 

range of overlap of the commercial and the food habits distributions with respect to the 

maximum recorded length of the prey species (as in Fig. 84). There is a significant negative 

correlation (P = 0.004) between the range of overlap and the size of the prey species (Fig. 86). 

That is, as the size of prey species increases, the range of overlap between commercial fisheries 

and marine mammal food habits decreases. This makes sense intuitively, considering the fact 

that marine mammals tend to focus on the lower end (< 30 cm, as shown earlier) of the size 

distribution of prey, regardless of the maximum length of the prey species targeted. The 

relationship presented here simply documents that commercial fisheries tend to target larger 

individuals of a given prey species, and the discrepancy between marine mammals and 

commercial fisheries increases with body size of the target species. 

140

 
y = -0.1072x + 35.027

R2

 = 0.0908

0

20

40

60

80

100

0 50 100 150 200 250

Maximum reported length of prey species (cm)

Overlap, as % of maximum length

crustaceans: n = 5

cephalopods: n = 13

fish: n = 61

T.c.

M.c. G.m.

L.v.r.

Figure 86.--Overlap between the size range of prey targeted by marine mammals and the size 

 range targeted by commercial fisheries, represented with respect to the maximum 

 reported length of the prey species. Abbreviations in plot refer to prey taxa with 

 multiple entries (L.v.r. = Loligo vulgaris renaudii, T.c. = Theragra chalcogramma, 

 M.c. = Merluccius capensis, G.m. = Gadus morhua). 

When viewed with respect to the commercial fishery (Fig. 87), there is also a significant 

negative correlation (P = 0.002) between the range of overlap and the size of the prey species. 

This follows directly from the information in Figure 86 because it would be impossible for a 

fishery to target a wider size range of the target species than was available. Indeed, it simply 

reflects the fact that commercial fisheries selectively target large fish when it is an option. Thus 

there is less overlap with marine mammal size selectivity for larger prey species and more 

overlap for small species where commercial selectivity is less of an option. 

141

 
The third way in which we examine overlap in the size range of prey consumption is 

conducted with respect to consumption by marine mammals (Fig. 88). Here, we see that there is 

no relationship between the degree of overlap and the maximum prey size (P = 0.90). This is 

perhaps due to the fact that, in general, marine mammals tend to target a much narrower range of 

prey sizes than do comparable fisheries focused on the same prey species (mean range of food 

habits = 23.82 cm; mean range of commercial fisheries = 35.96), even though the range of sizes 

targeted by marine mammals is significantly correlated (P = 0.05) with the maximum size of the 

prey species (Fig. 89). Furthermore, the correlation between targeted size range and maximum

prey size has a significantly higher slope (P < 0.01) for the commercial fisheries data (Fig. 90) 

than for the marine mammals food habits data (Fig. 89). 

142

 
y = -0.2137x + 73.929

R2

 = 0.1034

0

20

40

60

80

100

0 50 100 150 200 250

Maximum reported length of prey species (cm)

Overlap, as % of commercial range

crustaceans: n = 5

cephalopods: n = 13

fish: n = 61

T.c.

G.m.

M.c.

L.v.r.

Figure 87.--Overlap between the size range of prey targeted by marine mammals and the size 

 range targeted by commercial fisheries, represented with respect to the commercial 

 fishery. Abbreviations in plot refer to prey taxa with multiple entries (L.v.r. = Loligo 

 vulgaris renaudi, T.c. = Theragra chalcogramma, M.c. = Merluccius capensis, G.m. 

 = Gadus morhua). 

143

 
y = 0.0026x + 74.018

R2

 = 2E-05

0

20

40

60

80

100

0 50 100 150 200 250

Maximum reported length of prey species (cm)

Overlap, as % of food habits range

crustaceans: n = 5

cephalopods: n = 13

fish: n = 61

T.c.

M.c. G.m.

L.v.r.

Figure 88.--Overlap between the size range of prey targeted by marine mammals and the size 

 range targeted by commercial fisheries, represented with respect to the predators. 

 Abbreviations in plot refer to prey taxa with multiple entries (L.v.r. = Loligo vulgaris 

 renaudi, T.c. = Theragra chalcogramma, M.c. = Merluccius capensis, G.m. = Gadus 

 morhua). 

144

 
y = 0.2708x

R2

 = 0.0831

0

20

40

60

80

100

0 50 100 150 200 250

Maximum reported length of prey species (cm)

Size range targeted (cm)

crustaceans: n = 5

cephalopods: n = 13

fish: n = 61 T.c.

G.m.

M.c.

L.v.r.

Figure 89.--Relationship between size range of prey (in cm) targeted by marine mammals as a 

 function of the maximum reported length of the prey species. Mean range = 23.82 

 cm. Regression line is forced through the origin. Abbreviations in plot refer to prey 

 taxa with multiple entries (L.v.r. = Loligo vulgaris renaudi, T.c. = Theragra 

 chalcogramma, M.c. = Merluccius capensis, G.m. = Gadus morhua). 

145

 
y = 0.4632x

R2

 = 0.8314

0

20

40

60

80

100

0 50 100 150 200 250

Maximum reported length of prey species (cm)

Size range targeted (cm)

crustaceans: n = 5

cephalopods: n = 13

fish: n = 61

T.c.

Figure 90.--Relationship between size range of prey (in cm) targeted by commercial fisheries as 

 a function of the maximum reported length of the prey species. Mean range = 35.96 

 cm. Regression line is forced through the origin. Abbreviation in plot refers to prey 

 taxon with multiple entries (T.c. = Theragra chalcogramma). 

RESULTS: COMPARISONS WITH SURVEY DATA AND EVALUATIONS OF 

SELECTIVITY 

The comparisons above clearly indicate that commercial fisheries have a tendency to 

target larger individual fish and squid than do marine mammals foraging in the same areas at the 

same time. This suggests that size selectivity is occurring. There are several ways this can 

occur: 

1. commercial fisheries may be systematically selecting for the larger size classes available 

2. marine mammals may be systematically selecting for the smaller size classes available 

146

 
3. marine mammals may be opportunistically eating prey in proportion to their natural 

abundance, which would tend to favor small prey; or 

4. combinations of 1 and either 2 or 3 above may be occurring. 

We believe that a combination of 1 and 2 above is being observed, but the actual selectivity in 

each case is not yet clear. It is possible that marine mammals are more selective than 

commercial fisheries. That is, marine mammals may be systematically targeting a much 

narrower range of available size classes than do commercial fisheries. 

As with the comparisons between mean prey size from food habits studies and the mean 

targeted catch of commercial fisheries, we can examine the relationship between mean prey size 

and the mean size of the available prey population (as indicated by survey data). It is only 

through examination of the survey data that we can be more conclusive about the size selectivity 

of marine mammals. We proceed with the caveat that surveys are often conducted with gear and 

procedures so similar to those of commercial fishing operations as to obscure any selectivity on 

the part of commercial fisheries. 

In common with the commercial data needed for comparison with specific food habits 

studies, the availability of survey data is limited (Appendix 6). Survey data are typically 

generated in support of stock assessments for commercial fisheries. Thus, there is a heavy bias 

towards commercially valuable taxa. Furthermore, many research programs have predetermined sampling regions which are surveyed on a regular basis. It is typically not known (or 

knowable) the degree to which these sampling regions overlap with the foraging habitat of 

marine mammal taxa for any given food habits study (but see Kaschner and Pauly 2004). 

147

 
To evaluate the degree to which commercial fisheries and marine mammals exert size

selectivity on their target species, we use the same type of pairings as above to ensure that we 

used survey data with comparable temporal and spatial coverage to that of the food habits data 

and/or the commercial data. The mean size of prey from food habits studies and those from

commercial fisheries were subtracted from the means from surveys (Appendix 5 and 6). Thus, 

positive differences reflect situations where mean size of targeted prey was systematically 

smaller than the mean size of what was available, while negative differences reflect situations 

where mean size targeted was systematically larger than the mean size of the prey available. As 

we show below, however, negative values may reflect directional size selectivity, or they may 

reflect the lack of appropriate survey data for comparison (see Figs. 98 and 99 below). 

For the pairings between food habits means and survey means, the difference is 

consistently positive, indicating that in general, marine mammals systematically target smaller 

individuals than are present in the prey population (Fig. 91). The magnitude of the difference 

between food habits means and survey means is significantly correlated with maximum reported 

body size of the prey species (Fig. 92; P < 0.001). That is, the level of selectivity increases as 

maximum reported size increases for prey species.

148

 
0

5

10

15

20

25

30

35

40

45

50

-50 -40 -30 -20 -10 0 10 20 30 40 50

Absolute difference, survey mean minus food habits mean (cm)

Frequency

mean difference of means = 4.67 cm

Figure 91.--Distribution of differences in means (survey mean minus food habits mean) for 152 

 prey entries. 

149

 
y = 0.0969x

R2

 = 0.3413

-40

-30

-20

-10

0

10

20

30

40

50

60

0 50 100 150 200 250

Maximum reported prey size (cm)

Difference in means (cm)

crustaceans: n = 17

cephalopods: n = 18

fish: n = 117

mean difference in means = 4.67 cm

T.c.

M.c.

G.m.

Figure 92.--Distribution of differences in means (survey mean minus food habits mean) for 152 

 prey entries, as a function of the maximum reported size for each prey species. Solid 

 horizontal line is y = 0. Dashed horizontal line indicates the mean difference of 

 means (4.67 cm). Abbreviations in plot refer to prey taxa with multiple entries (T.c. 

 = Theragra chalcogramma, M.c. = Merluccius capensis, G.m. = Gadus morhua). 

In contrast to the case of food habits studies, the difference between mean size of prey as 

indicated by surveys and the mean size targeted by commercial fisheries is dominated by 

negative values and values close to zero (Fig. 93), with an overall mean difference of -0.4 cm. 

150

 
0

5

10

15

20

25

-50 -40 -30 -20 -10 0 10 20 30 40 50

Absolute difference, survey mean minus commercial mean (cm)

Frequenc

y

mean difference of means = - 0.43 cm

Figure 93.--Distribution of differences in means (survey mean minus commercial mean) for 85 

 taxonomic entries. 

151

 
y = 0.0112x

R2

 = 0.008

-40

-30

-20

-10

0

10

20

30

40

0 50 100 150 200 250

Maximum reported prey size (cm)

Difference in means (cm)

crustaceans: n = 5

cephalopods: n = 11

fish: n = 69

T.c.

Figure 94.--Distribution of differences in means (survey mean minus commercial mean) for 85 

 prey entries, as a function of the maximum reported size for each prey species. The 

 slope of the regression line is non-significant (P = 0.37). Solid horizontal line is 

 y = 0. The mean difference of means (-0.43 cm) has not been plotted. Abbreviation 

 in plot refers to prey taxon with multiple entries (T.c. = Theragra chalcogramma). 

This suggests that commercial fisheries tend to take prey size classes in close proportion to their 

natural abundance as indicated by survey data. Indeed, although there is variability in the 

magnitude of the differences, there is no relationship between the difference in means between 

survey and commercial data and the maximum reported size of the target species (Fig. 94). 

Another way to examine the relationships described in Figures 91-94 is to examine 

scatter plots of mean targeted size (either by marine mammals or by commercial fisheries) as a 

function of the mean size indicated by survey data. The mean size targeted by commercial 

152

 
fisheries shows a 1:1 relationship with the mean size available according to survey data, whereas 

marine mammals consistently target smaller individuals than are available (Fig. 95). 

0

10

20

30

40

50

60

70

80

90

100

0 20 40 60 80

Survey mean (cm)

Mean targeted size (cm)

100

food habits

commercial

y = x

Figure 95.--Relationship between survey means and mean targeted size for commercial and food 

 habits entries considered separately. The slope of the commercial regression (heavy 

 dashed line) is not significantly different from 1.0 (fine dashed line; P < 0.001); the 

 slope of the food habits regression (solid line) is significantly different from 1.0 

 (P < 0.001) and is also significantly different from the slope of the commercial 

 regression (P < 0.001). 

Evaluation of Appropriateness of Survey Data 

The preceding analyses show that there are consistent patterns to the size of prey targeted 

by both commercial fisheries and marine mammals. However, there are some notable outliers 

that require detailed examination. Next, we will examine a series of LF distributions to illustrate 

the general patterns of selectivity that we have documented thus far, and to highlight cases where 

the match between food habits data, commercial data, and survey data needs improvement. 

153

 
The first set of LF distributions we will examine is for round herring (Etrumeus 

whiteheadi) from the Benguela Current off of South Africa, where this species reaches a 

maximum size of 23.6 cm (Sekiguchi et al. 1992). Our previous analyses have shown that 

commercial fisheries and marine mammals target small-bodied prey taxa similarly. Examination 

of the LF distributions of the round herring targeted as compared to what are available, as 

indicated by survey data, shows that, in addition to a high degree of overlap between commercial 

fisheries and marine mammals, both are taking round herring roughly in proportion to their 

availability (Fig. 96). 

0

5

10

15

20

25

30

35

0 5 10 15 20 25 30

Length (cm)

Percent of total

food habits

survey

commercial

Figure 96.--Comparison of LF distributions of round herring (Etrumeus whiteheadi) from food 

 habits data (solid line) from 1969 to 1990 for common dolphin (Delphinus delphis), 

 survey data (fine dashed line) from 1984 to 1989, and commercial catch (heavy 

 dashed line) from 1986. Food habits and commercial data are from Sekiguchi et al. 

 (1992) and survey data are from Waldron et al. 1991. 

154

 
In contrast to the case of round herring, a mid-sized flatfish shows a slightly different 

pattern. Specifically, American plaice (Hippoglossoides platessoides), which reach a maximum 

reported length of 82 cm (Froese and Pauly 2003), are targeted slightly differently by marine 

mammals versus commercial fisheries (Fig. 97). Food habits data for gray seal (Halichoerus 

grypus) show a bimodal distribution of size selectivity, with an overall mean size of 24.3 cm

(Benoit and Bowen 1990). The mean size targeted by the commercial fishery is substantially 

larger (33.2 cm), but there is a high degree of overlap in the two LF distributions. Note, 

however, that the survey data clearly show that marine mammals and commercial fisheries are 

each targeting a slightly different component of the plaice population: marine mammals tend to 

target fish 35 cm and smaller, while commercial fisheries tend to target fish 25 cm and larger. 

155

 
0

5

10

15

20

25

30

35

40

0 10 20 30 40 50 60 70 8

Length (cm)

Percent of total

0

food habits

survey

commercial

Figure 97.--Comparison of LF distributions of American plaice (Hippoglossoides platessoides) 

 from food habits data (solid line) from 1982, 1983, 1986 and 1987 for gray seal 

 (Halichoerus grypus), survey data (fine dashed line) from 1975 to 1979, and 

 commercial catch (heavy dashed line) from 1984. Food habits data are from Benoit 

 and Bowen (1990), survey data are from Beacham (1982), and commercial data are 

 from Chouinard and Metuzals (1985). 

156

 
Moving to a much larger prey species, the LF distributions of Cape hake (Merluccius 

paradoxus and M. capensis), with maximum reported sizes of 115 cm and 140 cm, respectively 

(Froese and Pauly 2003), show a more striking pattern (Fig. 98). In particular, the LF 

distribution of hake targeted by Heaviside’s dolphin (Cephalorhynchus heavisidii) overlaps only 

slightly with the LF distribution of the same two species targeted by the commercial fishery 

(Sekiguchi et al. 1992). The survey data, however, seem to indicate that the commercial fishery 

is missing a substantial portion of the larger-bodied fish. In this particular case, the only 

available survey data are for M. capensis, the larger of the two species. Thus, although the 

commercial and food habits data are comparable, the only available survey data probably are not 

appropriate for comparison in this case. 

157

 
0

5

10

15

20

25

30

35

40

0 20 40 60 80

Length (cm)

Percent of total

100

food habits

survey

commercial

Figure 98.--Comparison of LF distributions of Cape hake (Merluccius capensis and M. 

 paradoxus) from food habits data (solid line) from 1969 to 1990 for Heaviside’s 

 dolphin (Cephalorhynchus heavisidii), survey data (fine dashed line) from 1987, and 

 commercial catch (heavy dashed line) from 1984 to 1986. Food habits and 

 commercial data are from Sekiguchi et al. (1992) and survey data are from Morales- 

 Nin (1991). 

158

 
Another situation in which the available data provide a misleading comparison comes 

again from the Benguela Current in South Africa. European anchovy (Engraulis encrasicolus) 

reach a maximum size of 20 cm (Froese and Pauly 2003). As a small-bodied prey species of 

high commercial importance, we would expect to see almost perfect overlap in the sizes targeted. 

In contrast, the available data seem to indicate that marine mammals systematically target larger 

individuals than the commercial fishery (11.2 cm versus 8.27 cm; Fig. 99). Note also, however, 

that the survey data indicate that the available anchovies do not exceed 10 cm in length. There is 

a clear inconsistency. Indeed, the available survey and commercial data are from along the 

Namibian coast (Cruikshank et al. 1990, Sekiguchi et al. 1992), which is dominated, for 

oceanographic reasons, by juvenile anchovies. The marine mammal food habits data, on the 

other hand, are primarily from marine mammals stranded along the southern coast of West 

Africa (Sekiguchi et al. 1992), which tends to have higher concentrations of adult anchovies. 

159

 
0

5

10

15

20

25

30

35

0 5 10 15 20

Length (cm)

Percent of total

food habits

survey

commercial

Figure 99.--Comparison of LF distributions of European anchovy (Engraulis encrasicolus) from 

 food habits data (solid line) from 1969-1990 for common dolphin (Delphinus 

 delphis), survey data (fine dashed line) from 1983, and commercial catch (heavy 

 dashed line) from 1986. Food habits and commercial data are from Sekiguchi et al. 

 (1992) and survey data are from Cruikshank et al. (1990). 

160

 
Finally, we consider the case of walleye pollock (Theragra chalcogramma) in the eastern 

Bering Sea. Pollock reach a maximum reported size of 91 cm (Froese and Pauly 2003). As with 

most other prey species, the size range targeted by marine mammals (here, northern fur seals 

0

5

10

15

20

25

0 10 20 30 40 50 60

Length (cm)

Percent of total

70

food habits

survey

commercial

Figure 100.--Comparison of LF distributions of walleye pollock (Theragra chalcogramma) from 

 food habits data (solid line) from 1974 for northern fur seals (Callorhinus ursinus), 

 survey data (fine dashed line) from 1975, and commercial catch (heavy dashed 

 line) from 1974. Food habits data are from McAlister and Perez (1977); survey 

 and commercial data are from Bakkala (1988). 

(Callorhinus ursinus)) is primarily smaller than 30 cm (Fig. 100). The commercial and survey 

data show a high degree of concordance, and both clearly indicate that northern fur seals are 

selectively targeting small individuals. 

On the basis of the overall patterns shown above, we conclude that marine mammals are 

more selective in the size composition of fish and cephalopods they consume than are 

commercial fisheries when food habits and fishery catches are compared to what is available, as 

161

 
indicated in surveys. We have not accounted for the selectivity of surveys (or commercial 

fisheries) relative to the size distribution across the full spectrum of sizes with the populations of 

resource species. This is because few stock assessment surveys and no commercial fisheries use 

gear that samples individuals that are only hours (even days or weeks) old – individuals that 

often fall in the size ranges of species ordinarily considered part of the plankton community. 

POTENTIAL APPLICATIONS 

Although we have documented consistent patterns in the targeted size composition both 

across and among different marine mammal taxa, most of these examples cover vast expanses of 

geographical range. Taken together, the consistency of the patterns suggests that the foraging 

behavior of marine mammals worldwide is under broadly similar selective pressures. From that, 

we argue that management should strive to generate the same patterns. We caution, however, 

that applying a pattern that appears to be global to a resource managed at a local level, under 

perhaps unique conditions, is unwise without reference to more detailed data. We provide two 

more examples of the kinds of data that are required under the systemic management approach to 

inform on local management decisions. 

The first example is ecosystem-specific, and examines the size selectivity of marine 

mammals in the Benguela Current. The Benguela Current flows along the west coast of South 

Africa and Namibia, and features a strong, nutrient-rich upwelling system that supports one of 

the most productive ecosystems in the world (O’Toole et al. 2001). In addition to supporting a 

large commercial fishery, the Benguela Current is also home to hosts of marine mammals. The 

large set of food habits data from the Benguela Current (David 1987, Sekiguchi et al. 1992) 

162

 
allows examination of the pattern of selectivity exerted by marine mammals on prey stocks in 

that system. 

Using the same criteria as before, 99 food habits cases meet our minimum sample size 

requirements, including data from one otariid seal and 10 odontocete whales, and their 

consumption of one crustacean taxon, 25 cephalopods taxa, and 22 fish taxa (Appendix 1, 3). 

The distribution of mean prey sizes for marine mammals foraging in the Benguela current 

strongly resembles most of the patterns we have documented above. Specifically, the 

distribution is distinctly positively skewed (Fig. 101), with a modal mean size of 15-20 cm, and 

an overall mean of means of 18.26 cm. Because many of the prey species represented in this 

example are harvested commercially (1/1 crustacean species; 4/25 cephalopod taxa; 21/22 fish 

taxa), the application to fisheries management is direct: in order to fit within the norm of

predation pressure in the Benguela Current, commercial fisheries would need to target the same

size distribution as presented here. Within this overarching pattern, patterns like those presented

in Figures 30-42 would help guide the species-specific applications. 

163

 
0

5

10

15

20

25

5 15 25 35 45 55 65 75 85 ≥100

Length (cm)

Frequenc

y

n = 99; mean of means = 18.26 cm

Figure 101.--Frequency distribution of prey size means for cases based on samples the Benguela 

Current with sample sizes ≥ 10. 

The second example spans adjoining oceanic domains highlighting a distinction between 

near-shore, or coastal, and off-shore, or pelagic systems. Several odontocete whale species have 

multiple ecotypes, often with distinctive coloration, dorsal fin morphology, and population 

genetics (Perrin et al. 2002). Not surprisingly, these ecotypes also tend to have distinctly 

different diets. Specifically, off-shore ecotypes have a tendency to target small-bodied prey taxa, 

whereas the near-shore ecotypes tend to target taxa with a larger range of sizes. For instance, 

Crawford (1981) found that 78% of all fish eaten by Dall’s porpoise (Phocoenoides dalli) in 

pelagic waters of the North Pacific were Protomyctophum thompsoni, which reach a maximum

total length of 5.4 cm (Froese and Pauly 2003). The fish taxa measured by Crawford (1981), 

which did not include P. thompsoni (Appendix 1, 3), averaged 10.02 cm in length (5 cases with 

n ≥ 10). If P. thompsoni had been included in Crawford’s measurements, the mean size would 

have been even smaller. In contrast, fish taxa consumed by coastal populations of Dall’s 

164

 
porpoise averaged 16.94 cm in length (14 cases with n ≥ 10; Ohizumi et al. 2000, Walker 1996, 

Walker et al. 1998). 

Examining the distribution of mean prey size for pan-tropical spotted dolphin (Stenella 

attenuata, Fig. 102), we begin to see how these differences are manifested. Food habits data 

from pelagic waters (Robertson and Chivers 1998) show a pattern similar to that of Dall’s 

porpoise, with a mean prey length of 9.45 cm (23 cases with n ≥ 10; Appendix 1, 3). Samples 

from coastal waters (Wang et al. 2003), which average 15.53 cm (13 cases with n ≥ 10),

nevertheless show a strong mode at 10-15 cm (Fig. 102). While it is clear that mean prey size 

targeted in coastal systems is larger than that targeted in pelagic systems, the difference is 

accounted for by a positively skewed distribution of means in the coastal system. Thus, while 

the bulk of the prey are less than 20 cm, a few prey taxa up to 60 cm mean length are also 

targeted. 

165

 
0

1

2

3

4

5

6

7

8

9

0-5 20-25 40-45 60-65 80-85 ≥100

Length (cm)

Frequency

near-shore: n = 13; mean of means = 15.53

off-shore: n = 23; mean of means = 9.45

Figure 102.--Frequency distribution of mean prey size for taxa eaten by Stenella attenuata. 

Near-shore data are from Sekiguchi et al. (1992) and Wang et al. (2003). Off-shore 

data are from Robertson and Chivers (1998). 

We have shown several different examples of how the specificity of the existing data can 

be sub-divided to highlight particular patterns, and emphasize that all of these patterns should be 

accounted for simultaneously in fisheries management. It is equally important to consider the 

single-species patterns (e.g., Figs. 30-42) as it is the ecosystem-level patterns (Fig. 101). 

Because the emergent properties of the components contribute to the emergent properties of the 

whole, management decisions based on this information can be consonant across all levels of 

organization. 

DISCUSSION AND CONCLUSIONS 

Our study crosses a wide diversity and size range of both marine mammal and prey taxa. 

The emergent pattern is that marine mammals selectively target small-bodied prey. Because the 

166

 
pattern is so pervasive, it seems safe to argue that it is stable much as other macroecological 

patterns, and is a reflection of the contributing factors that make it a characteristic of marine 

ecosystems (e.g., it includes the contribution of evolutionary and coevolutionary factors, as well 

as the selectivity of extinction processes to include macroevolutionary factors). As such, it 

follows that it is indicative of what is sustainable over very long periods, automatically 

incorporating and integrating the effects of a wide range of variables at various spatial and 

temporal scales (including regional and global climate change, changes in ocean circulation or

upwelling patterns, etc.). The main caveat to be recognized in this regard is that patterns 

observed in biotic systems today also reflect human influence. In other words, among the 

multitude of factors integrated in the patterns shown above are the numerous effects that we 

humans have, and have had, on our environment (e.g., climate change, pesticides, and especially 

in many of the cases we present, the effects of intensive fishing pressure).

Recommendations for Restructuring Management Policies 

Our results show that, as body size for prey species increases, the size selectivity of 

commercial harvests of fish exhibit an increasing departure from the size selectivity observed 

among other mammalian species (Fig. 66). This pattern demonstrates several things 

simultaneously. First, it provides a measure of the problem of abnormal selectivity exhibited by 

commercial fishing measured as the difference from what is normally experienced by prey 

species in these marine systems. Second it shows that the problem is more extreme with the 

larger-bodied species of prey. Third, it allows for taking the mean or maximum size of prey into 

account in the advice offered to fisheries managers. 

167

 
In order to avoid abnormal effects, managers are advised to preferentially target fish and 

squid such that the average size of the catch is ≤ 30 cm. For small-bodied taxa (e.g., market 

squid (Loligo opalescens), herring (Clupea harengus), etc.), the size composition achieved 

through current management practices match the patterns of marine mammal predation (Figs. 66-

71) and, as such, need not be changed. However, in order for management to achieve 

consistency with marine mammal predation on large-bodied taxa (e.g., Atlantic cod (Gadus 

morhua), walleye pollock (Theragra chalcogramma), etc.), substantial changes would need to be 

made to shift the emphasis towards targeting the small size classes of those taxa (i.e., mimic the 

selectivity of marine mammals). 

Special harvesting strategies (gear type, season, location) and regulations would be 

necessary to conduct fishing consistent with the selectivity normally experienced by each of the 

prey species included in this synthesis and collectively by the ecosystems of which they are a 

part. In the absence of information for individual species, guidance can be tentatively applied in 

the harvest of other species based on knowledge of their maximum body size. 

This advice is in direct contrast to current fishing practices (Figs. 66, 72-77) wherein 

larger fish are targeted in preference to smaller fish. Indeed, development of fisheries directed at 

forage (i.e., small, schooling) fish have been banned in some regions (NMFS 1998). In 

conventional approaches, catching larger fish can be justified on the grounds that it helps avoid 

direct competition with other consumers such as marine mammals—competition that would 

occur if small fish were caught. Or it might be argued that harvests with size composition 

similar to that of wild resource populations is non-selective and minimizes genetic effects. 

While there is undoubtedly something to such arguments, they fail to account for the complexity 

of the situation and introduce other problems, such as abnormal selectivity. Competition is a 

168

 
factor to be taken into account and must be dealt with on its own merits. Here we are dealing 

directly with the issue of selectivity and what might appear to be non-selective from one 

perspective is actually abnormally selective when evaluated on the basis of natural patterns. 

Bias caused by human influence cannot be ignored in the differences we have measured 

between the selectivity of marine mammals and commercial fisheries. Given that most or all of 

our data were derived from marine systems that have experienced varying degrees of 

anthropogenic perturbation, the possibility exists that the patterns of marine mammal size 

selectivity we observe now do not represent normal or natural behavior. This hypothesis can 

only be tested by the removal of abnormal anthropogenic influences and recovery of the affected 

systems. There are also the biases inherent to different approaches to documenting the food 

habits of marine mammals. These may eventually be remedied (Gudmundson et al. in press, 

Tollit et al. 2004b, Zeppelin et al. 2004). Removing the bias against large specimens in food 

habits studies may lead to the conclusion that the pattern we have documented is less clear than 

the published data show. Therefore, it is incumbent upon us as scientists to continue to 

document, to the best of our ability, the patterns of marine mammal foraging behavior. This will 

be particularly important if/when the management recommendations that emerge from studies 

such as this are implemented. 

The matter of taking into account the body size of prey (Fig. 65) is an example of using 

correlative information to help account for complexity (Fowler 2003). Accounting for the body 

size of a resource species (prey) is only one aspect of using correlative information and 

accounting for the body size of the predator is another. Accounting for the body size of humans 

(as the predator being managed) would be accomplished by using data such as that shown in 

Figures 51-54 if part of the variance is explained by a relationship between body size of the 

169

 
predator and selectivity in observed consumption patterns. If further research reveals such a 

correlation, the species chosen to provide the best examples of sustainable selectivity would be 

of a body size similar to that of humans (or, alternatively, the correlation would be used to 

estimate the size distribution of prey in a sustainable harvest by species of human body size). 

Many modern commercial fishing practices result in systemic effects that are 

different from those exerted by marine mammals. In applying the principles of management 

(and systemic management in particular; Fowler 2003), fishing effort would mimic the patterns 

seen in the feeding habits of marine mammals. This would reduce the mortality on the larger 

size classes, but not necessarily to zero. It would also avoid the abnormal effects (especially the 

risks) of either preferentially harvesting larger fish or trying to harvest non-selectively. 

Of particular concern are the abnormal genetic effects human harvest practices are likely 

to be exerting on commercial fish stocks. One might think that accounting for such effects 

would be of little concern owing to the fact that evolution occurs over relatively long time scales 

compared to behavioral effects or density dependent responses by a prey species. There are 

several reasons why it is important to take seriously the need to manage for selectivity typically 

experienced within the natural system. First, selective effects, and especially abnormal selective 

effects, can take their toll on time scales measured in just a few generations of the prey species 

(often less than a decade in lapsed time: Slatkin 1983, Allen 1987, Bradshaw and Hardwick 

1989, Conover 2000, Conover and Munch 2002, Cury 1993, Kirkpatrick 1993, Levinton 1992, 

Mitchell and Arthur 1991, Orians 1975, Reznick et al. 1997, Sutherland 1990). Second, the 

intensity of fishing rates makes the selective pressure particularly strong (Law 2000). This fact 

also emphasizes the importance of management to reduce harvests (which themselves can be 

brought to within the normal range of natural variation through systemic management: Fowler et 

170

 
al. 1999, Fowler 1999, Fowler and Hobbs 2002, 2003; Fowler 2003). Third, one of the goals of 

management is that of accounting for longer time scales whether or not they seem important to 

us in daily, weekly, or monthly time scales (Fowler 2003). As mentioned in the literature cited 

in our introduction, the need is clear and not to be avoided. Genetic effects count as one of the 

many factors to be addressed in accounting for complexity. This includes not only direct effects 

but also indirect effects (Conover 2000). 

On the surface, it might seem that a benign approach to managing the size composition of 

harvests would be to harvest such that each age class is represented in the harvest in proportion 

to its abundance in the resource population. In effect, this would introduce a form of selection 

different from that experienced in the natural system – selection that is among the forces that 

cause a system to have its observed form and function (e.g., the patterns in normal variation for 

the more inclusive systems, such as ecosystems). If we insist on taking larger individuals from

the populations of the large-bodied resource species (e.g., adult salmon, Oncorhynchus spp., for 

aesthetic or economic reasons), the appropriate information for managing the harvest (e.g., and 

especially, the consumption/harvest rate) would come from non-human consumer species that 

restrict their consumption to adult fish (e.g., bears, Ursus spp., in the case of salmon). 

It is important to keep in mind that we are not offering the results of our work as a 

means simply to allow for a Darwinian approach to management. The "evolutionary enlightened 

manager" acting independent of (and especially instead of) the "ecologically enlightened 

manager" (Brown and Parman 1993) does not achieve the holism needed in accounting for 

complexity. Both have to be involved, not so much as separate disciplines, but as a matter of 

taking into account the elements of natural systems involved in the respective studies. Both the 

ecological and the evolutionary are taken into account when management is based on mimicking 

171

 
natural patterns (along with all other elements of the complexity taken into account (Fowler and 

Crawford 2004)). 

There is a distinction to be made in single-species applications of the advice from studies 

such as this and ecosystem applications of the resulting advice. Single species applications 

would be exemplified by management ensuring that the harvests of walleye pollock (Theragra

chalcogramma) have a size composition similar to that of the food habits of marine mammals. 

This would be based on information such as shown in Figure 73. Such management would entail 

reducing the size of pollock taken by the commercial fishery so that the LF distribution would 

correspond to that of the food habits of marine mammals (e.g., similar mean and variance). An 

ecosystem application, however, would entail similar approaches for all harvested species in any 

particular ecosystem, as shown in Figure 101. Likewise, the slope of relationships such as that 

shown in Figure 66 would be reduced to zero and the slope of the line for commercial fisheries in 

Figure 95 would be reduced to correspond to that for marine mammals. The pattern for 

commercial fisheries at the ecosystem level would correspond to the pattern for marine mammals 

-- a pattern that demonstrates what works within marine ecosystems and a pattern important to 

the health of these ecosystems. 

Changing commercial fishing practices to fit more closely with the patterns documented 

for marine mammals would, especially for larger bodied prey, require dramatic shifts in the 

targeted size composition. However, such changes are among those needed to account for the 

complexity of large systems and must be implemented simultaneously with other changes such 

as large reductions required in the total catch. Reduction in the size composition of commercial 

catches would, at a minimum:

172

 
1. increase direct competition between commercial fisheries and some marine mammal 

species -- a step toward consistency in the competition observed in natural systems, 

2. raise the question of sustainable harvest rates for large bodied prey if we persist with 

current selectivity, 

3. raise the issue of marketability of small-bodied fish, and 

4. force reconsideration of the driving economic factors associated with large-scale 

commercial fishing practices. 

Many of the world’s commercial fish stocks are clearly in serious trouble. Although 

sustainable harvests are typically the stated goal in fisheries management, previous practices 

have, in large part, failed to meet that goal. Systemic management is based on directly relevant 

(completely consonant) guiding information that produces recommendations for changing 

commercial harvest practices in such a way that long-term sustainability is an option. 

As has been argued before (Fowler and Hobbs 2002), systemic management

automatically incorporates all uncertainties. In contrast, conventional management attempts the 

difficult, or perhaps impossible task of parameterizing all the variables that are both a) thought to 

be important and b) measurable. Inasmuch as we agree that it is important to identify the 

appropriate variables (Fowler and Crawford 2004), we believe that our documentation of the size 

selectivity of marine mammals provides an elegant solution to the challenge of sufficient 

parameterization faced by managers today. 

173

 
ACKNOWLEDGEMENTS 

We would like to thank Sandy McFarlane, Loretta O’Brien, Kristin Laidre and Beth 

Sinclair for access to unpublished data. The manuscript has benefited substantially from

thoughtful reviews provided by Gary Duker, Jenny Etnier, Liz Etnier, Anne Hollowed, Jim Lee, 

Pat Livingston, Beth Sinclair, and Bill Walker. Finally, MAE would like to thank CWF for his 

constant enthusiasm and support during the process of assembling this report. Is has been an 

unforgettable experience collaborating on such a fascinating topic. 

174

 
CITATIONS
 

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212 
 
Appendix 1: Summary food habits data, sorted phylogenetically by predator. 

Total N of cases Mean of prey 

Common name N of cases in analysis* means (cm) 

Otariid seals 361 237 15.82 

Predator species 

Arctocephalus australis Arctocephalus forsteri Arctocephalus galapagoensis Arctocephalus gazella Arctocephalus pusillus doriferus Arctocephalus pusillus pusillus Arctocephalus tropicalis Arctocephalus spp. 

Callorhinus ursinus Eumetopias jubatus Otaria flavescens Phocarctos hookeri Zalophus californianus Zalophus californianus woelbecki 

Phocid seals 

South American fur seal 

New Zealand fur seal Galapagos fur seal 

Antarctic fur seal Australian fur seal South African fur seal sub-Antarctic fur seal 

sub-Antarctic and Antarctic fur seals, mixed northern fur seal Steller sea lion South American sea lion Hooker's sea lion California sea lion Galapagos sea lion 

2 1 9.80 

14 7 13.72 

1 1 14.10 

142 99 12.33 

14 11 19.97 72 49 17.98 22 12 9.47 

4 2 6.20 

46 18 12.90 11 8 33.35 

6 6 26.48 

11 9 36.11 

14 12 14.27 

2 2 13.80 

234 146 18.64 

213

Cystophora cristata Erignathus barbatus Halichoerus grypus Hydrurga leptonyx Leptonychotes weddelii Lobodon carcinophagus Mirounga angustirostris Mirounga leonina Monachus monachus Ommatophoca rossii Phoca fasciata Phoca groenlandica Phoca hispida Phoca largha Phoca vitulina 

hooded seal bearded seal grey seal leopard seal Weddell seal crabeater seal northern elephant seal southern elephant seal Mediterranean monk seal Ross seal ribbon seal harp seal ringed seal spotted seal harbor seal 

2 2 34.40 4 3 10.30 

78 52 24.64 

1 1 13.76 

23 9 11.79 

2 0 n.a. 

3 2 16.92 

33 12 14.72 

2 1 30.00 

6 3 14.70 3 3 12.44 

26 11 20.31 

9 9 10.55 6 5 13.13 

36 33 15.45 

Mysticete whales 7 5 5.90 

Balaenoptera acutorostrata minke whale 1 1 14.50 

Balaenoptera pysalus fin whale 1 0 n.a. 

Balaenoptera spp. rorquals, mix of fin and blue whales 4 4 4.45 

Megaptera novaeangliae humpback whale 1 0 n.a. 
Appendix 1: Summary food habits data, sorted phylogenetically by predator. 

Total N of cases Mean of prey 

Common name N of cases in analysis* means (cm) Predator species 

Odontocete whales 556 338 18.64 

214

Berardius bairdii Baird's beaked whale 19 15 36.64 

Cephalorhynchus commersonii Commerson's dolphin 3 3 9.50 

Cephalorhynchus heavisidii Heaviside's dolphin 6 4 14.83 Delphinapterus leucas beluga whale 3 2 13.51 Delphinus delphis common dolphin 6 6 16.33 Delphinus sp. common dolphin 4 3 4.20 

Feresa attenuata pygmy killer whale 4 0 n.a. 

Globicephala macrorhynchus short-finned pilot whale 3 3 14.27 Globicephala melas = G. melaena long-finned pilot whale 21 10 16.43 

Globicephala sp. pilot whale, species not specified 2 2 24.86 

Grampus griseus Risso's dolphin 24 15 12.08 

Hyperoodon planifrons southern bottlenosed whale 71 44 25.04 Kogia breviceps pygmy sperm whale 27 12 15.85 

Kogia sima dwarf sperm whale 24 8 9.89 

Lagenodelphis hosei Fraser's dolphin 27 8 8.92 

Lagenorhynchus obliquidens Pacific white-sided dolphin 2 0 n.a. 

Lagenorhynchus obscurus dusky dolphin 9 8 12.86 

Mesoplodon densirostris Blainville's beaked whale 1 0 n.a. 

Mesoplodon layardii strap-toothed whale 28 6 16.59 

Monodon monoceros narwhal 8 6 20.55 

Orcinus orca killer whale 9 1 12.60 

Phocoena phocoena harbor porpoise 50 31 16.99 

Phocoenoides dalli Dall's porpoise 43 37 12.88 Physeter macrocephalus = P. catadon sperm whale 69 47 31.35 

Pontoporia blainvillei Franciscana 9 6 9.12 

Pseudorca crassidens false killer whale 2 0 n.a. 

Sotalia fluviatilis tucuxi 20 10 9.71 Stenella attenuata pan-tropical spotted dolphin 42 37 12.74 

Stenella coeruleoalba striped dolphin 2 2 12.10 

Stenella frontalis Atlantic spotted dolphin 1 1 8.90 Stenella longirostris spinner dolphin 9 7 11.44 Steno bredanensis rough-toothed dolphin 1 1 21.50 Tursiops truncatus bottlenosed dolphin 7 3 23.77 

mixed taxa** 8 0 n.a. 

OVERALL 1166 726 17.9 

*includes only those cases where mean prey size was based on a sample size of >=10 **an unspecified mix of northern fur seal, Dall's porpoise, and Pacific white-sided dolphin 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

215

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

Arctocephalus spp. Electrona carlsbergi 1.38 5.87 0.35 30 4.85 6.79 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 

Arctocephalus spp. Electrona subaspera 1.38 6.53 0.64 22 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Arctocephalus spp. Electrona subaspera 1.38 7.94 0.57 5 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Arctocephalus spp. Gymnoscopelus piabilis 1.38 8.94 3.04 2 6.79 11.09 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 

A. australis Loligo sanpaulensis 1.65 9.80 37 4.50 18.50 dos Santos and Haimovici 2001 Brazil 1985-1998 A. australis Argonauta nodosa 1.65 1.50 3 1.00 1.80 dos Santos and Haimovici 2001 Brazil 1985-1998 

A. forsteri Nototodarus sloanii 1.40 § 19.00 386 5.00 43.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

A. forsteri Electrona subaspera 1.20 6.94 1.01 42 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 A. forsteri Electrona subaspera 1.20 8.55 0.70 57 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 A. forsteri Electrona subaspera 1.20 7.65 1.40 15 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 A. forsteri Electrona sp. 1.40 § 5.00 280 2.00 7.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

A. forsteri Gymnoscopelus nicholsi 1.20 13.35 1.51 52 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

A. forsteri Gymnoscopelus nicholsi 1.20 12.53 2.33 99 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

A. forsteri Lampanyctodes hectoris 1.40 § 4.60 1801 2.00 6.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

A. forsteri Symbolophorus sp. 1.40 § 8.50 421 2.00 14.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 A. forsteri Auchenoceros punctatus 1.40 § 11.50 871 5.00 15.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

A. forsteri Trachurus declivis 1.40 31.00 5.00 145 Lake 1997 Tasmania 1992-1993 

A. forsteri Trachurus sp. 1.40 3 80.00 90.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

A. forsteri Trachurus sp. 1.40 10 50.00 60.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 A. forsteri Emmelichthys nitidus 1.40 16.00 4.00 315 Lake 1997 Tasmania 1992-1993 

A. galapagoensis Sardinops sagax 1.37 14.10 33 Dellinger and Trillmich 1999 Galapagos Islands, E Tropical Pacific 1983 

A. gazella Loligo sanpaulensis 1.25 11.10 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

A. gazella Brachioteuthis cf. picta 1.00 7.08 26 5.69 8.65 Daneri et al. 1999 South Orkney Islands, S Atlantic 1988 

A. gazella Brachioteuthis cf. picta 1.00 8.09 9 7.69 8.55 Daneri et al. 1999 South Shetland Islands, S Atlantic 1992-1994 

A. gazella Brachioteuthis cf. riisei 0.75 5.84 31 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Brachioteuthis spp. 0.80 5.60 3 4.66 6.27 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Gonatus antarcticus 0.75 8.55 4 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Histioteuthis machrohista 0.80 8.41 1 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Mastigoteuthis sp. A 0.75 7.65 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Martialia hyadesi 1.25 17.46 1.13 4 15.80 18.30 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Martialia hyadesi 0.75 22.07 39 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Kondakovia longimana 0.80 20.20 1 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Moroteuthis ingens 0.75 16.45 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Moroteuthis knipovitchi 0.80 20.44 8 13.40 27.10 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Moroteuthis knipovitchi 0.75 9.18 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 A. gazella Psychroteuthis glacialis 1.00 9.07 7 7.83 10.76 Daneri et al. 1999 South Orkney Islands, S Atlantic 1988 

A. gazella Psychroteuthis glacialis 1.00 10.12 13 7.83 12.83 Daneri et al. 1999 Scotia Arc, Antarctica 1988-1994 A. gazella Psychroteuthis glacialis 1.00 11.26 6 9.98 12.83 Daneri et al. 1999 South Shetland Islands, S Atlantic 1992-1994 

A. gazella Euphausia superba 0.75 5.54 324 3.80 6.60 Reid et al. 1996 South Georgia Island, S Atlantic 1986 A. gazella Euphausia superba 0.75 4.45 308 3.60 6.20 Croxall et al. 1999 South Georgia Island, S Atlantic 1994 A. gazella Euphausia superba 0.75 5.54 324 3.80 6.60 Croxall et al. 1999 South Georgia Island, S Atlantic 1986 A. gazella Euphausia superba 0.55 5.13 0.31 301 3.90 6.10 Croxall and Pilcher 1984 South Georgia Island, S Atlantic 1972-1977 

A. gazella Euphausia superba 0.55 5.14 388 3.50 6.50 Doidge and Croxall 1985 South Georgia Island, S Atlantic 1982-1983 

A. gazella Euphausia superba 0.75 4.67 5587 3.20 6.40 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

A. gazella Euphausia superba 1.50 4.03 781 2.40 5.60 Reid 1995 South Georgia Island, S Atlantic 1992 A. gazella Euphausia superba 1.50 4.47 1387 3.60 5.80 Reid 1995 South Georgia Island, S Atlantic 1993 A. gazella Euphausia superba 0.70 4.26 n.a.** McCafferty et al. 1998 South Georgia Island, S Atlantic 1994-1996 

A. gazella Euphausia superba 0.75 4.63 n.a.** 3.20 6.10 Reid and Brierly 2001 South Georgia Island, S Atlantic 1994-1999 

A. gazella Euphausia superba 0.80 5.10 241 4.20 5.90 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Notolepis coatsi 1.00 27.18 12 20.72 30.89 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

A. gazella Electrona antarctica 1.25 9.80 1.00 22 7.70 11.00 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Electrona antarctica 1.00 7.24 114 5.50 9.00 Daneri and Coria 1993 South Orkney Islands, S Atlantic 1988 

A. gazella Electrona antarctica 1.00 8.65 0.92 38 7.15 10.36 Daneri 1996 South Shetland Islands, S Atlantic 1992 

A. gazella Electrona antarctica 0.75 5.61 16 4.13 8.10 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Electrona antarctica 1.25 5.99 112 4.80 8.80 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

216

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

A. gazella Electrona antarctica 1.25 7.32 92 5.00 17.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

A. gazella Electrona antarctica 1.50 7.68 2202 3.75 9.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 A. gazella Electrona antarctica 1.00 7.81 75 5.75 9.50 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

A. gazella Electrona antarctica 1.25 10.40 78 8.60 11.60 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

A. gazella Electrona antarctica 1.00 6.20 0.70 340 4.00 8.00 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

A. gazella Electrona antarctica 0.80 10.88 51 6.40 13.26 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Electrona antarctica 0.75 6.35 92 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Electrona carlsbergi 1.25 7.10 1.33 56 3.40 9.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Electrona carlsbergi 1.25 8.10 18 7.30 8.40 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

A. gazella Electrona carlsbergi 1.00 8.50 1.00 80 6.00 10.10 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 A. gazella Electrona carlsbergi 0.80 8.01 23 6.30 8.98 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Electrona carlsbergi 0.75 8.02 26 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Electrona subaspera 0.75 7.14 8 4.84 8.41 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Electrona subaspera 1.25 8.00 1.34 98 3.90 10.60 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. gazella Electrona subaspera 1.25 8.52 1.59 103 1.69 12.59 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 A. gazella Electrona subaspera 1.50 10.82 2030 6.25 12.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 A. gazella Electrona subaspera 0.75 7.40 120 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Gymnoscopelus bolini 0.75 6.50 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Gymnoscopelus bolini 1.25 11.70 1.41 12 9.70 14.20 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Gymnoscopelus bolini 0.75 18.50 17 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Gymnoscopelus braueri 0.75 6.59 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Gymnoscopelus braueri 1.00 8.20 1.10 5 6.90 9.60 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

A. gazella Gymnoscopelus braueri 1.25 10.30 6 8.60 11.60 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

A. gazella Gymnoscopelus braueri 0.80 10.63 19 8.16 12.49 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Gymnoscopelus braueri 0.75 9.59 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Gymnoscopelus fraseri 0.75 7.90 15 6.77 8.56 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Gymnoscopelus fraseri 1.25 8.80 2.08 189 5.80 13.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. gazella Gymnoscopelus fraseri 0.75 7.62 156 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Gymnoscopelus microlampas 1.25 9.00 2 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Gymnoscopelus nicholsi 1.00 12.28 109 10.00 14.00 Daneri and Coria 1993 South Orkney Islands, S Atlantic 1988 

A. gazella Gymnoscopelus nicholsi 1.00 14.42 0.77 134 12.60 16.55 Daneri 1996 South Shetland Islands, S Atlantic 1992 A. gazella Gymnoscopelus nicholsi 0.75 8.90 295 5.50 12.89 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Gymnoscopelus nicholsi 1.25 12.92 114 5.20 8.80 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

A. gazella Gymnoscopelus nicholsi 1.25 12.78 474 6.00 17.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

A. gazella Gymnoscopelus nicholsi 1.25 8.40 2.22 46 5.70 14.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Gymnoscopelus nicholsi 1.00 13.36 105 11.51 15.29 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

A. gazella Gymnoscopelus nicholsi 1.25 12.90 38 10.80 15.10 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

A. gazella Gymnoscopelus nicholsi 1.00 14.70 2.30 369 6.20 18.90 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

A. gazella Gymnoscopelus nicholsi 0.80 11.34 14 8.79 12.80 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

A. gazella Gymnoscopelus nicholsi 0.75 11.74 144 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Gymnoscopelus piabilis 0.75 1.30 32 8.22 14.29 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Gymnoscopelus piabilis 1.25 12.30 1.80 447 4.70 15.50 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Gymnoscopelus piabilis 0.75 12.84 139 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Krefftichthys anderssoni 1.25 3.30 2 3.30 3.30 North 1996 South Georgia Island, S Atlantic 1983 A. gazella Krefftichthys anderssoni 0.75 3.90 7 2.90 5.40 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Krefftichthys anderssoni 1.25 5.60 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. gazella Krefftichthys anderssoni 1.50 4.55 2790 2.75 7.25 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 A. gazella Krefftichthys anderssoni 1.50 3.38 350 2.75 8.25 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 A. gazella Krefftichthys anderssoni 1.25 5.60 1 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

A. gazella Krefftichthys anderssoni 1.00 8.50 0.30 3 8.20 8.70 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

A. gazella Krefftichthys anderssoni 0.75 3.52 47 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Lampichthys procerus 1.25 6.90 0.64 4 6.00 7.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Metelectrona ventralis 0.75 6.18 16 4.90 7.16 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Metelectrona ventralis 1.25 7.10 1.26 78 3.80 10.20 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. gazella Metelectrona ventralis 0.75 6.90 11 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

217

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

A. gazella Protomyctophum andriashevi 0.75 5.12 2 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Protomyctophum bolini 1.25 4.80 1.60 3 3.70 6.70 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Protomyctophum bolini 0.75 3.70 17 3.25 4.14 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Protomyctophum bolini 1.25 5.70 1.94 11 2.80 9.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Protomyctophum bolini 0.75 3.99 81 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Protomyctophum choriodon 1.25 7.10 0.50 4 6.50 7.50 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Protomyctophum choriodon 0.75 7.00 11 5.78 7.85 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Protomyctophum choriodon 1.25 8.60 2.07 82 4.00 13.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. gazella Protomyctophum choriodon 0.75 8.52 261 7.25 9.25 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

A. gazella Protomyctophum choriodon 0.75 8.73 241 7.25 9.75 Reid and Arnould 1996 South Georgia Island, S Atlantic 1994 

A. gazella Protomyctophum choriodon 0.75 9.25 215 7.25 11.25 Reid and Arnould 1996 South Georgia Island, S Atlantic 1993 

A. gazella Protomyctophum choriodon 0.75 6.68 114 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Protomyctophum tenisoni 0.75 4.02 2 3.52 4.52 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Protomyctophum tenisoni 1.25 4.60 0.51 322 1.90 6.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Protomyctophum tenisoni 0.75 4.16 32 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Muraenolepis microps 1.25 38.30 0.70 4 37.50 39.10 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Dissostichus eleginoides 1.25 45.40 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Gobionotothen acuta 0.75 7.32 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Gobionotothen gibberifrons 1.25 31.60 11.70 55 5.90 54.00 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Gobionotothen gibberifrons 1.50 42.10 49 23.00 59.00 Reid 1995 South Georgia Island, S Atlantic 1992 A. gazella Gobionotothen gibberifrons 1.50 43.22 46 21.00 59.00 Reid 1995 South Georgia Island, S Atlantic 1993 A. gazella Gobionotothen gibberifrons 1.00 37.00 1 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

A. gazella Gobionotothen marionensis 1.25 11.60 2.59 4 9.70 15.30 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Lepidonotothen larseni 1.25 14.70 3.30 91 7.80 22.00 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Lepidonotothen larseni 0.75 16.20 142 10.50 23.50 Reid and Arnould 1996 South Georgia Island, S Atlantic 1994 

A. gazella Lepidonotothen larseni 0.75 14.04 110 10.50 23.50 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

A. gazella Lepidonotothen larseni 0.75 18.55 65 13.50 25.50 Reid and Arnould 1996 South Georgia Island, S Atlantic 1993 

A. gazella Lepidonotothen larseni 1.50 16.79 193 4.00 26.00 Reid 1995 South Georgia Island, S Atlantic 1992 

A. gazella Lepidonotothen larseni 1.50 20.30 187 11.00 28.00 Reid 1995 South Georgia Island, S Atlantic 1993 

A. gazella Lepidonotothen squamifrons 0.75 3.53 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Notothenia squamifrons 1.25 11.40 4.96 12 3.60 22.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. gazella Paranotothenia magellanica 1.25 14.00 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. gazella Pleuragramma antarcticum 1.00 14.26 1.24 136 11.59 17.80 Daneri 1996 South Shetland Islands, S Atlantic 1992 A. gazella Pleuragramma antarcticum 1.00 13.87 33 11.90 16.88 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

A. gazella Pleuragramma antarcticum 1.00 15.00 1.60 6 12.90 17.00 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 A. gazella Chaenocephalus aceratus 1.25 49.30 4.50 7 41.40 53.40 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Champsocephalus gunnari 1.25 23.80 8.60 201 10.00 46.00 North 1996 South Georgia Island, S Atlantic 1983 

A. gazella Champsocephalus gunnari 0.75 21.50 7 18.49 22.94 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Champsocephalus gunnari 1.25 21.09 530 12.00 34.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

A. gazella Champsocephalus gunnari 0.75 31.88 328 15.00 45.00 Reid and Arnould 1996 South Georgia Island, S Atlantic 1994 

A. gazella Champsocephalus gunnari 0.75 24.27 158 15.00 47.00 Reid and Arnould 1996 South Georgia Island, S Atlantic 1993 

A. gazella Champsocephalus gunnari 0.75 38.11 18 27.00 49.00 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

A. gazella Champsocephalus gunnari 1.50 21.72 1522 12.00 44.00 Reid 1995 South Georgia Island, S Atlantic 1992 

A. gazella Champsocephalus gunnari 1.50 25.09 660 16.00 46.00 Reid 1995 South Georgia Island, S Atlantic 1993 

A. gazella Champsocephalus gunnari 1.50 20.80 1263 18.75 23.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 

A. gazella Champsocephalus gunnari 1.50 22.42 877 18.25 27.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 

A. gazella Champsocephalus gunnari 0.75 19.29 9.80 161 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. gazella Channichthys rhinoceratus 1.25 32.52 101 20.00 44.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

A. gazella Chionodraco rastrospinosus 1.00 15.10 2.20 2 13.60 16.70 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

A. gazella Cryodraco antarcticus 1.00 36.60 2.00 3 35.30 38.90 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 A. gazella Pseudochaenichthys georgianus 1.25 28.80 2.40 26 24.10 34.80 North 1996 South Georgia Island, S Atlantic 1983 A. gazella Paradiplospinus gracilis 0.75 3.97 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

A. gazella Icichthys australis 0.75 31.81 17 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

A. p. doriferus Sepia apama 1.70 30.10 5.67 43 21.00 41.30 Gales et al. 1993 Tasmania 1989-1990 

A. p. doriferus Sepia novaehollandiae 1.70 15.50 4.51 23 3.70 20.10 Gales et al. 1993 Tasmania 1989-1990 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

218

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

A. p. doriferus Sepioteuthis australis 1.70 32.80 11.33 29 13.50 53.10 Gales et al. 1993 Tasmania 1989-1990 

A. p. doriferus Nototodarus gouldi 1.70 23.90 6.99 213 8.80 41.50 Gales et al. 1993 Tasmania 1989-1990 

A. p. doriferus Argonauta nodosa 1.70 5.70 3.58 3 1.60 8.20 Gales et al. 1993 Tasmania 1989-1990 A. p. doriferus Octopus australis 1.70 10.10 2.63 13 4.50 15.50 Gales et al. 1993 Tasmania 1989-1990 

A. p. doriferus Octopus maorum 1.70 15.30 3.03 13 11.70 22.70 Gales et al. 1993 Tasmania 1989-1990 

A. p. doriferus Octopus pallidus 1.70 11.70 1.40 2 10.70 12.70 Gales et al. 1993 Tasmania 1989-1990 

A. p. doriferus Octopus superciliosus 1.70 5.02 1 Gales et al. 1993 Tasmania 1989-1990 

A. p. doriferus Sardinops sagax cf. neopilchardus 1.70 17.30 1.00 15 15.00 19.00 Gales and Pemberton 1994 Tasmania 1984-1986 A. p. doriferus Pseudophycis bachus 1.70 19.80 5.70 13 15.00 35.00 Gales and Pemberton 1994 Tasmania 1984-1986 

A. p. doriferus Trachurus declivis 1.70 21.20 2.80 48 16.00 31.00 Gales and Pemberton 1994 Tasmania 1984-1986 

A. p. doriferus Emmelichthys nitidus 1.70 17.00 1.20 194 14.00 21.00 Gales and Pemberton 1994 Tasmania 1984-1986 

A. p. doriferus Seriolella brama 1.70 16.70 1.50 15 14.00 19.00 Gales and Pemberton 1994 Tasmania 1984-1986 

A. p. pusillus Sepia officinalis 1.70 16.31 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Sepia sp. 2.05 14.29 76 4.80 21.70 Lipinski and David 1990 Benguela, South Africa 1974-1985 

A. p. pusillus Loligo vulgaris reynaudii 2.05 11.50 51 7.10 18.90 Lipinski and David 1990 Benguela, South Africa 1974-1985 

A. p. pusillus Loligo vulgaris reynaudii 2.05 15.60 596 4.10 32.40 Lipinski and David 1990 Benguela, South Africa 1974-1985 

A. p. pusillus Loligo vulgaris reynaudii 2.05 16.06 183 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Lycoteuthis lorigera 2.05 8.71 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Ommastrephes bartramii 2.05 20.82 9 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Todarodes angolensis 2.05 14.60 22 3.40 29.10 Lipinski and David 1990 Benguela, South Africa 1974-1985 

A. p. pusillus Todaropsis eblanae 2.05 8.30 94 3.50 16.10 Lipinski and David 1990 Benguela, South Africa 1974-1985 A. p. pusillus Onychoteuthis banksii 2.05 14.95 2 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Argonauta nodosa 2.05 7.21 3 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Octopus sp. 2.05 7.42 15 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Jasus lalandii 0.85 4.18 89 1.00 7.00 David 1987 Benguela, South Africa 1981 

A. p. pusillus Jasus lalandii 0.85 5.20 38 1.00 7.00 David 1987 Benguela, South Africa 1984 

A. p. pusillus Jasus lalandii 0.85 2.50 253 1.00 6.00 David 1987 Benguela, South Africa 1985 

A. p. pusillus Etrumeus whiteheadi 0.85 14.70 17 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Etrumeus whiteheadi 0.85 13.20 32 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Sardinops sagax = S. ocellatus 2.05 21.66 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Sardinops sagax cf. ocellatus 0.85 14.90 24 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Sardinops sagax cf. ocellatus 0.85 19.80 134 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Sardinops sagax cf. ocellatus 0.85 18.70 214 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Engraulis encrasicolus 0.85 12.70 1799 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Engraulis encrasicolus 0.85 9.90 2911 David 1987 Benguela, South Africa 1974-1985 A. p. pusillus Engraulis encrasicolus 0.85 9.40 15871 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Engraulis encrasicolus 2.05 13.47 1 Castley et al. 1991 Benguela, South Africa 1976-1990 A. p. pusillus Maurolicus muelleri 0.85 3.40 1 David 1987 Benguela, South Africa 1974-1985 A. p. pusillus Maurolicus muelleri 0.85 3.40 1257 David 1987 Benguela, South Africa 1974-1985 A. p. pusillus Lampanyctodes hectoris 0.85 4.80 595 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Lampanyctodes hectoris 0.85 5.00 1849 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Macrouridae 0.85 26.10 1 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Macrouridae 0.85 26.10 144 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Merluccius capensis 2.05 28.19 64 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Merluccius spp. 0.85 29.50 116 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Merluccius spp. 0.85 17.40 350 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Merluccius spp. 0.85 21.30 1177 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Genypterus capensis 0.85 45.90 25 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Genypterus capensis 2.05 52.73 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Chorisochismus dentex 0.85 9.50 1 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Scomberesox saurus 0.85 24.20 29 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Helicolenus dactylopterus 0.85 20.00 1 David 1987 Benguela, South Africa 1974-1985 A. p. pusillus Helicolenus dactylopterus 0.85 23.20 13 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Helicolenus dactylopterus 0.85 20.00 145 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Chelidonichthys capensis 0.85 15.00 10 David 1987 Benguela, South Africa 1974-1985 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

219

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

A. p. pusillus Trachurus capensis 0.85 10.80 66 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Trachurus capensis 0.85 16.30 449 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Trachurus capensis 0.85 14.80 3542 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Trachurus capensis 2.05 27.15 56 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Argyrozona argyrozona 0.85 16.10 2 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Argyrozona argyrozona 2.05 37.03 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Cheimerius nufar 2.05 11.75 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Pagellus natalensis 2.05 22.87 7 Castley et al. 1991 Benguela, South Africa 1976-1990 A. p. pusillus Pterogymnus laniarius 0.85 20.00 15 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Pterogymnus laniarius 2.05 28.36 15 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Argyrosomus hololepidotus 2.05 34.68 2 Castley et al. 1991 Benguela, South Africa 1976-1990 A. p. pusillus Mugil cephalus 0.85 13.80 38 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Gymnammodytes capensis 0.85 25.00 12 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Gymnammodytes capensis 0.85 25.00 25 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Clinus sp. 0.85 11.30 7 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Callionymidae 0.85 21.50 86 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Sufflogobius bibarbatus 0.85 6.10 64 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Sufflogobius bibarbatus 0.85 6.90 48078 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Thyrsites atun 0.85 80.00 1 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Thyrsites atun 0.85 80.00 2 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Thyrsites atun 0.85 80.00 10 David 1987 Benguela, South Africa 1974-1985 A. p. pusillus Lepidopus caudatus 0.85 75.00 1 David 1987 Benguela, South Africa 1974-1985 A. p. pusillus Lepidopus caudatus 0.85 75.00 5 David 1987 Benguela, South Africa 1974-1985 A. p. pusillus Scomber japonicus 0.85 32.50 6 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Scomber japonicus 0.85 24.70 15 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Pleuronectidae 0.85 21.10 32 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Pleuronectidae 0.85 14.60 144 David 1987 Benguela, South Africa 1974-1985 

A. p. pusillus Austroglossus pectoralis 2.05 27.90 10 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. p. pusillus Cynoglossus zanzibarensis 2.05 20.20 117 Castley et al. 1991 Benguela, South Africa 1976-1990 

A. tropicalis Loligo sanpaulensis 1.50 9.30 14 5.80 13.50 dos Santos and Haimovici 2001 Brazil 1985-1998 A. tropicalis Illex argentinus 1.50 33.20 6 29.70 35.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

A. tropicalis Ommastrephes bartramii 1.50 22.10 23 9.30 34.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

A. tropicalis Moroteuthis knipovitchi 1.50 10.54 6 Bester and Laycock 1985 South Georgia Island, S Atlantic 1977-1978 

A. tropicalis Argonauta nodosa 1.50 10.30 2 7.30 13.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

A. tropicalis Electrona antarctica 1.50 7.20 0.30 7 6.90 7.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Electrona carlsbergi 1.50 6.50 0.87 114 3.80 8.80 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Electrona subaspera 1.50 7.80 1.34 70 5.00 12.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Electrona subaspera 1.50 8.21 1.48 58 1.69 12.59 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 

A. tropicalis Gymnoscopelus bolini 1.50 10.60 2.71 40 6.50 14.50 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. tropicalis Gymnoscopelus fraseri 1.50 7.80 1.46 275 3.50 13.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Gymnoscopelus microlampas 1.50 8.60 0.34 2 8.40 8.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Gymnoscopelus nicholsi 1.50 8.50 2.83 62 4.80 15.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. tropicalis Gymnoscopelus piabilis 1.50 11.60 2.17 283 5.80 16.00 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. tropicalis Krefftichthys anderssoni 1.50 4.40 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Metelectrona ventralis 1.50 7.10 1.44 188 3.20 12.20 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Protomyctophum bolini 1.50 4.00 0.54 9 2.90 5.00 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Protomyctophum choriodon 1.50 9.50 1.37 73 5.40 13.10 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Protomyctophum tenisoni 1.50 4.60 0.60 225 2.60 7.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

A. tropicalis Paranotothenia magellanica 1.50 17.90 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. tropicalis Champsocephalus gunnari 1.50 14.10 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 A. tropicalis Paradiplospinus gracilis 1.50 21.50 2.19 2 19.90 23.00 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 C. ursinus Loligo bleekeri 1.12 12.45 8 6.93 17.97 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Loligo opalescens 1.40 43 7.00 15.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Loliolus japonica 1.12 9.61 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Chiroteuthis imperator 1.12 6.14 28 4.48 7.18 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

220

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

C. ursinus Enoploteuthis chunii 1.12 10.75 3 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Watasenia scintillans 1.12 4.40 99 3.00 5.60 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Watasenia scintillans 1.12 4.20 978 3.00 5.80 Mori et al. 2001 Honshu, NW Pacific 1987-1994 C. ursinus Watasenia scintillans 1.12 4.30 1248 2.52 6.80 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Berryteuthis anonychus 0.93 29.68 3 27.21 31.99 Mori et al. 2001 N Pacific 1987-1991 C. ursinus Gonatopsis borealis 0.93 8.22 8 5.96 10.36 Mori et al. 2001 N Pacific 1987-1991 C. ursinus Gonatopsis borealis 1.12 10.40 37 5.56 16.68 Mori et al. 2001 Honshu, NW Pacific 1987-1994 C. ursinus Gonatus berryi 1.12 9.21 5 6.45 12.80 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Gonatus onyx 1.12 11.40 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Gonatidae 1.40 >59 5.00 24.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Mastigoteuthis sp. 1.12 13.73 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Octopoteuthis sp. 1.12 6.68 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Eucleoteuthis luminosa 1.12 5.07 10 Mori et al. 2001 Honshu, NW Pacific 1987-1994 C. ursinus Ommastrephes bartramii 0.93 8.61 10 7.09 9.96 Mori et al. 2001 N Pacific 1987-1991 

C. ursinus Ommastrephes bartramii 1.12 16.24 24 12.65 21.76 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Todarodes pacificus 1.12 14.15 3 5.04 33.55 Mori et al. 2001 Honshu, NW Pacific 1987-1994 C. ursinus Moroteuthis loennbergi 1.12 21.19 2 13.88 29.20 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Onychoteuthis borealijaponica 0.93 15.64 42 6.86 27.58 Mori et al. 2001 N Pacific 1987-1991 

C. ursinus Onychoteuthis borealijaponica 1.12 11.06 9 1.54 35.29 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

C. ursinus Onychoteuthidae 1.40 >3 14.00 22.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Clupea harengus pallasi 1.40 >27 10.00 25.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Engraulis mordax 1.40 27 9.00 18.00 Perez and Bigg 1986 N Pacific 1958-1974 C. ursinus Bathylagidae 1.40 986 8.00 12.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Mallotus villosus 1.40 64 7.00 14.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Thaleichthys pacificus 1.40 11 12.00 21.00 Perez and Bigg 1986 N Pacific 1958-1974 C. ursinus Salmonidae 1.40 >26 15.00 41.00 Perez and Bigg 1986 N Pacific 1958-1974 

Fiscus et al. 1964, cited in Frost and Lowry 1986 Bering Sea 1962 C. ursinus Theragra chalcogramma 1.40 tly <30-35 cm (p. 36)" 

C. ursinus Theragra chalcogramma 1.40 19.30 n.a.** 10.00 35.00 McAlister et al. 1976 Bering Sea 1974 

C. ursinus Theragra chalcogramma 1.21 32.97 39 6.00 41.00 Sinclair et al. 1994 Bering Sea 1981 

C. ursinus Theragra chalcogramma 1.16 6.70 1191 3.00 42.00 Sinclair et al. 1994 Bering Sea 1982 C. ursinus Gonatopsis/Berryteuthis 1.16 9.71 166 7.00 17.00 Sinclair et al. 1994 Bering Sea 1982 

C. ursinus Gonatopsis/Berryteuthis 1.24 9.22 136 5.00 17.00 Sinclair et al. 1994 Bering Sea 1985 

C. ursinus Gonatus madokai/G. middendorffi 1.24 6.67 86 5.00 9.00 Sinclair et al. 1994 Bering Sea 1985 

C. ursinus Theragra chalcogramma 1.24 13.03 1428 4.00 47.00 Sinclair et al. 1994 Bering Sea 1985 C. ursinus Theragra chalcogramma 1.40 29.19 59 6.80 48.00 Kiyota et al. 1999 St. Paul Island, Bering Sea 1996 

C. ursinus Theragra chalcogramma 1.40 1721 4.00 40.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Theragra chalcogramma 1.40 n.a.** 6.00 20.00 Swartzman and Haar 1983 NE Pacific & Bering Sea 1960-1974 

C. ursinus Theragra chalcogramma 1.40 30.40 n.a.** Lowry et al. 1989 Bering Sea 1981-1982 

C. ursinus Merluccius productus 1.40 15.00 2 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Cololabis saira 1.40 25.00 4 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Sebastes spp. 1.40 >19 11.00 31.00 Perez and Bigg 1986 N Pacific 1958-1974 

C. ursinus Pleurogrammus monopterygius 1.40 >5 15.00 23.00 Perez and Bigg 1986 N Pacific 1958-1974 

E. jubatus Theragra chalcogramma 2.55 25.20 1135 8.30 64.20 Frost and Lowry 1986 Bering Sea 1981 E. jubatus Theragra chalcogramma 2.55 21.80 10.30 51.60 Lowry et al. 1989 St. Matthew Island, Bering Sea 1985 

E. jubatus Theragra chalcogramma 2.55 33.50 20.80 44.50 Lowry et al. 1989 St. Paul Island, Bering Sea 1985 

E. jubatus Theragra chalcogramma 2.55 29.80 11.60 2030 5.60 62.90 Pitcher 1981 Gulf of Alaska 1975-1978 

E. jubatus Theragra chalcogramma 2.55 46.90 280 18.40 61.40 Frost and Lowry 1986 Pribilof Islands, Bering Sea 1976, 1979 

E. jubatus Theragra chalcogramma 2.55 29.90 1.70 42.70 Lowry et al. 1989 E Aleutian Islands, Gulf of Alaska 1981-1982 

E. jubatus Theragra chalcogramma 2.20 25.50 80 4.80 55.70 Calkins and Goodwin 1988 Kodiak Island, Gulf of Alaska 1985-1986 

E. jubatus Theragra chalcogramma 2.25 25.40 1064 7.90 54.20 Calkins and Goodwin 1988 SE Alaska, Gulf of Alaska 1985-1986 

E. jubatus Theragra chalcogramma 2.55 42.40 11.6 909 10.00 78.10 Tollit et al. 2004a SE Alaska 1994-1999 

E. jubatus Theragra chalcogramma 2.28 39.30 14.3 666 3.70 70.80 Zeppelin et al. 2004 Aleutian Is. (western stock) 1998-2000 

E. jubatus Pleurogrammus monopterygius 2.28 32.30 5.9 1685 15.30 49.60 Zeppelin et al. 2004 Aleutian Is. (western stock) 1998-2000 

O. flavescens Loligo gahi 1.85 9.70 n.a.** 4.00 23.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

221

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

O. flavescens Illex argentinus 1.85 20.90 n.a.** 14.00 30.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 O. flavescens Illex argentinus 1.85 25.00 n.a.** 18.00 30.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 O. flavescens Macruronus magellanicus 1.89 47.50 8.90 206 20.00 70.00 George-Nascimento et al. 1985 Chile 1979-1981 

O. flavescens Merluccius hubbsi 1.85 26.70 n.a.** 10.00 55.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 O. flavescens Merluccius hubbsi 1.85 29.10 n.a.** 15.00 60.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 P. hookeri Octopus maorum 2.00 120.00 33 30.00 150.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Ovalipes catharus 2.00 7.00 118 4.00 11.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Raja spp. 2.00 68.00 13 53.00 93.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Auchenoceros punctatus 2.00 10.00 166 7.00 13.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 P. hookeri Pseudophycis sp. 2.00 36.00 20 20.00 50.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Genypterus blacodes 2.00 4 68.00 124.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Trachurus sp. 2.00 38.00 47 27.00 45.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Notolabrus sp. 2.00 23.00 21 9.00 40.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Odax pullus 2.00 31.00 36 22.00 49.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Thyrsites atun 2.00 77.00 14 49.00 86.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

P. hookeri Rhombosolea sp. 2.00 35.00 54 16.00 50.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Z. californianus Loligo opalescens 2.00 12.89 300 7.50 22.50 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

Z. californianus Loligo opalescens 2.00 9.50 17.50 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

Z. californianus Loligo opalescens 2.00 12.70 1.70 76 6.20 18.50 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 

Z. californianus Loligo opalescens 1.80 12.70 25386 1.00 23.50 Lowry and Caretta 1999 Channel Islands, NE Pacific 1981-1985 Z. californianus Loligo opalescens 1.80 5.20 164 4.20 6.70 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Z. californianus Sardinops sagax 1.80 9.20 5 8.20 11.50 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Z. californianus Engraulis mordax 2.00 9.50 0.80 94 5.50 14.10 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 Z. californianus Engraulis mordax 1.80 10.00 12 9.00 11.10 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Z. californianus Merluccius productus 2.20 30.70 1856 13.00 58.00 Bailey and Ainley 1982 Farallon Islands, NE Pacific 1974-1978 

Z. californianus Merluccius productus 2.00 16.60 6.00 222 8.90 26.10 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 Z. californianus Merluccius productus 1.80 18.60 35 4.90 26.80 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Z. californianus Sebastes sp. 2.00 17.10 2.20 155 12.90 22.70 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 

Z. californianus Sebastes spp. 1.80 12.10 31 9.20 14.60 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Z. californianus Trachurus symmetricus 1.80 13.10 12 10.90 16.20 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Z. c. woelbecki Sardinops sagax 2.00 14.20 66 Dellinger and Trillmich 1999 Galapagos Islands, E Tropical Pacific 1983 

Z. c. woelbecki Sardinops sagax 2.00 13.40 334 Dellinger and Trillmich 1999 Galapagos Islands, E Tropical Pacific 1983-1986 

C. cristata Gadus morhua 2.25 36.40 68 14.00 73.00 Hauksson and Bogason 1997 Iceland 1992-1993 

C. cristata Sebastes sp. 2.25 32.40 n.a.** 23.00 43.00 Hauksson and Bogason 1997 Iceland 1992-1993 

E. barbatus Chionoectes opilio 2.25 5.70 1.20 336 3.20 12.30 Antonelis et al. 1994b Bering Sea 1981 E. barbatus Boreogadus saida 2.25 13.40 272 6.90 23.60 Finley and Evans 1983 Baffin Bay, NW Atlantic 1978-1979 

E. barbatus Theragra chalcogramma 2.25 11.80 0.15 56 6.90 14.30 Antonelis et al. 1994b Bering Sea 1981 

Antonelis pers. com., 

E. barbatus Theragra chalcogramma 2.25 8.20 cited in Frost and Lowry 1986 St. Matthew Island, Bering Sea <1983 

H. grypus Illex illecebrosus 2.08 18.50 78 10.00 25.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Raja sp. 2.08 70.00 2 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Alosa pseudoharengus 2.08 22.60 10 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Clupea harengus 2.08 25.40 5.60 48 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Clupea harengus 2.08 24.90 111 20.00 37.30 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Clupea harengus 2.08 34.50 160 5.00 40.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Clupea harengus 2.08 n.a.** 25.00 35.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Mallotus villosus 2.08 16.30 0.40 219 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Mallotus villosus 2.08 13.90 891 8.30 17.40 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Mallotus villosus 2.08 n.a.** 10.00 15.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Osmerus mordax 2.08 12.00 1 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Salmo salar 2.08 43.00 5 35.70 52.10 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Gadus morhua 2.08 17.30 11.40 101 10.00 40.00 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 H. grypus Gadus morhua 2.08 32.14 63 5.00 65.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Gadus morhua 2.08 32.21 61 5.00 75.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Gadus morhua 2.08 28.27 49 5.00 75.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

SE 0.32 

SE 0.47 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

222

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

H. grypus Gadus morhua 2.08 29.75 180 2.50 72.50 Prime and Hammond 1990 Donna Nook, SW North Sea 1985 

H. grypus Gadus morhua 2.08 31.01 172 2.50 67.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Gadus morhua 2.08 32.29 145 2.50 77.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Gadus morhua 2.08 28.20 269 6.70 79.70 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Gadus morhua 2.08 25.80 SE 0.92 115 10.00 55.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Gadus morhua 2.08 18.80 81 2.50 37.50 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

H. grypus Gadus morhua 2.08 468 30.00 50.00 Hauksson and Bogason 1997 Iceland 1992-1993 H. grypus Melanagrammus aeglefinus 2.08 23.50 2 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Melanagrammus aeglefinus 2.08 32.40 2 14.80 57.20 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Melanagrammus aeglefinus 2.08 20 20.00 40.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Merlangius merlangus 2.08 21.25 1318 9.00 37.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Merlangius merlangus 2.08 26.50 203 9.00 43.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Pollachius virens 2.08 37.94 79 7.50 72.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Pollachius virens 2.08 45.71 63 22.50 92.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Pollachius virens 2.08 10.90 2 8.10 13.80 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Pollachius virens 2.08 20.00 55 5.00 35.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Pollachius virens 2.08 48 10.00 60.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Molva molva 2.08 28.06 81 5.00 85.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Molva molva 2.08 32.83 60 5.00 85.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Molva molva 2.08 32.53 77 5.00 105.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Molva molva 2.08 33.30 256 2.50 77.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Urophycis tenuis 2.08 22.20 12 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Merluccius bilinearis 2.08 30.30 2 21.10 42.30 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Merluccius bilinearis 2.08 25.40 43 10.00 45.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Merluccius bilinearis 2.08 25.67 30 15.00 35.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

H. grypus Sebastes sp. 2.08 n.a.** 20.00 30.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Sebastes spp. 2.08 14.15 71 10.00 25.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

H. grypus Myoxocephalus scorpius 2.08 24.60 1 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Myoxocephalus scorpius 2.08 n.a.** 15.00 25.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Cyclopterus lumpus 2.08 15.50 0.50 15 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Cyclopterus lumpus 2.08 24.90 26 15.00 29.20 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Tautogolabrus adspersus 2.08 25.40 1 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Zoarces americanus 2.08 53.30 12 45.70 61.00 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Anarhichas lupus 2.08 29.60 3 22.90 35.60 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Anarhichas lupus 2.08 n.a.** 20.00 60.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Ammodytes dubius 2.08 19.10 SE 0.47 139 5.00 25.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Ammodytes sp. 2.08 18.10 62 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Ammodytes sp. 2.08 n.a.** 15.00 35.00 Hauksson and Bogason 1997 Iceland 1992-1993 

H. grypus Ammodytes spp. 2.08 16.72 324 5.00 31.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Ammodytes spp. 2.08 17.50 1878 5.00 43.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Ammodytes spp. 2.08 16.29 418 3.00 43.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Ammodytes spp. 2.08 16.82 1220 3.00 45.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

H. grypus Ammodytes spp. 2.08 13.93 6752 3.00 25.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Ammodytes spp. 2.08 13.96 5943 1.00 25.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Ammodytes spp. 2.08 16.67 1536 3.00 31.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Ammodytes spp. 2.08 15.00 876 5.00 30.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

H. grypus Ammodytidae 2.08 16.37 2341 1.50 34.50 Prime and Hammond 1990 Donna Nook, SW North Sea 1985 

H. grypus Scomber scombrus 2.08 32.60 24 25.00 47.90 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Scomber scombrus 2.08 20.90 8 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 H. grypus Glyptocephalus cynoglossus 2.08 31.30 1 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Hippoglossoides platessoides 2.08 34.40 5.20 6 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

H. grypus Hippoglossoides platessoides 2.08 24.30 269 11.10 46.80 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Hippoglossoides platessoides 2.08 14.70 9 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Hippoglossoides platessoides 2.08 25.91 23 15.00 40.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

H. grypus Limanda ferruginea 2.08 37.50 1 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

SE 1.04 SE 1.48 

SE 1.03 

SE 5.05 

SE 1.56 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

223

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

H. grypus Limanda ferruginea 2.08 20.00 14 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Limanda ferruginea 2.08 26.76 27 10.00 40.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

H. grypus Pleuronectes americanus 2.08 24.30 2 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

H. grypus Pleuronectes platessa 2.08 31.99 247 2.50 77.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Pleuronectes platessa 2.08 24.07 51 7.50 77.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

H. grypus Reinhardtius hippoglossoides 2.08 46.10 2 43.00 49.10 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

H. grypus Solea solea 2.08 24.04 550 2.50 57.50 Prime and Hammond 1990 Donna Nook, SW North Sea 1985 

H. leptonyx Pleuragramma antarcticum 2.80 13.76 2.43 226 8.80 17.50 Green and Williams 1986 Davis Station, Antarctica 1984 L. weddelii Psychroteuthis glacialis 2.00 § 35.10 5 12.30 38.30 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Chorismus antarcticus 3.00 1.33 0.14 1496 0.71 2.00 Green and Burton 1987 McMurdo Sound, Antarctica 1983-1985 

L. weddelii Notocrangon antarcticus 3.00 1.41 0.14 836 Green and Burton 1987 McMurdo Sound, Antarctica 1983-1985 

L. weddelii Pleuragramma antarcticum 3.00 16.24 80 12.00 21.00 Plötz et al. 1991 Weddell Sea, Antarctica 1998 

L. weddelii Pleuragramma antarcticum 3.00 15.41 1296 5.00 21.50 Plötz 1986 Weddell Sea, Antarctica 1983, 1985 

L. weddelii Pleuragramma antarcticum 3.00 16.05 1034 6.00 22.00 Plötz 1986 Weddell Sea, Antarctica 1983, 1985 

L. weddelii Pleuragramma antarcticum 3.00 11.64 2.33 26 5.20 19.10 Green and Burton 1987 Mawson, Antarctica 1983-1985 

L. weddelii Pleuragramma antarcticum 3.00 15.03 1.69 169 7.20 21.00 Green and Burton 1987 Davis Station, Antarctica 1983-1985 

L. weddelii Pleuragramma antarcticum 3.00 14.97 2.33 330 5.20 25.00 Green and Burton 1987 McMurdo Sound, Antarctica 1983-1985 L. weddelii Pleuragramma antarcticum 3.00 14.07 4.27 168 5.07 25.91 Burns et al. 1998 McMurdo Sound, Antarctica 1989-1993 L. weddelii Trematomus eulepidotus 2.00 § 18.70 68 14.40 23.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Trematomus lepidorhinus 2.00 § 14.90 12 12.30 19.30 Plötz et al. 1991 Weddell Sea, Antarctica 1986 L. weddelii Trematomus nicolai 2.00 § 18.90 3 16.70 20.00 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Trematomus spp. 3.00 23.34 7.89 6 12.73 34.10 Burns et al. 1998 McMurdo Sound, Antarctica 1989-1993 

L. weddelii Artedidraco loennbergi 2.00 8.70 2 8.50 8.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Dolloidraco longedorsalis 2.00 § 9.40 5 7.70 9.60 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Histiodraco velifer 2.00 9.85 2 8.90 10.80 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Gerlachea australis 2.00 § 20.60 7 19.00 22.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Gymnodraco acuticeps 2.00 § 24.60 7 18.00 29.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Racovitzia glacialis 2.00 § 19.10 65 12.20 22.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 L. weddelii Chionodraco myersi 2.00 § 27.30 107 20.10 34.40 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

L. weddelii Cryodraco antarcticus 2.00 § 27.00 28 20.80 32.40 Plötz et al. 1991 Weddell Sea, Antarctica 1986 L. weddelii Pagetopsis maculatus 2.00 § 16.90 66 13.30 20.50 Plötz et al. 1991 Weddell Sea, Antarctica 1986 L. carcinophagus Antarctomysis maxima 2.40 75 2.00 2.50 Green and Williams 1986 Davis Station, Antarctica 1984 L. carcinophagus Pleuragramma antarcticum 2.40 15.50 4 12.20 18.40 Green and Williams 1986 Davis Station, Antarctica 1984 

M. angustirostris Loligo opalescens 1.69 11.84 90 6.17 20.51 Sinclair 1994 S California Bight, NE Pacific 1969-1983 

M. angustirostris Gonatopsis borealis 3.60 22.00 5.40 22 13.60 28.00 Antonelis et al. 1994a San Miguel Island, NE Pacific 1978-1979 

M. angustirostris Merluccius productus 3.60 28.74 252 15.40 51.80 Antonelis et al. 1994a San Miguel Island, NE Pacific 1978-1979 

M. leonina Batoteuthis skolops 3.45 10.30 3 8.90 10.90 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Brachioteuthis cf. picta 3.45 8.10 192 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Brachioteuthis cf. picta 3.45 7.28 26 Daneri et al. 2000 South Shetland Islands, S Atlantic 1995-1996 

M. leonina Chiroteuthis veranyi 3.45 14.10 1 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1994 

M. leonina Chiroteuthis sp. 3.45 13.50 6 10.20 15.60 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Galiteuthis glacialis 3.45 18.70 7 13.00 23.60 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Mesonychoteuthis hamiltoni 3.45 36.20 2 31.90 40.50 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Gonatus antarcticus 3.45 13.90 67 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Gonatus antarcticus 3.45 18.53 12 Daneri et al. 2000 South Shetland Islands, S Atlantic 1995-1996 

M. leonina Histioteuthis sp. B 3.45 4.70 58 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Histioteuthis sp. 3.45 5.30 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

M. leonina Lycoteuthis lorigera 3.45 7.80 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

M. leonina Mastigoteuthis psychrophila 3.45 11.00 5 10.10 13.00 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Alluroteuthis antarctica 3.45 14.50 51 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Alluroteuthis antarctica 3.45 13.13 13 Daneri et al. 2000 South Shetland Islands, S Atlantic 1992-1994 

M. leonina Illex argentinus 3.45 26.30 2 26.00 26.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

M. leonina Martialia hyadesi 3.45 27.10 43 19.3 42.6 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Kondakovia longimana 3.45 26.19 7 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1995 

M. leonina Moroteuthis knipovitchi 3.45 23.70 121 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

SE 1.95 SE 2.01 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

224

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

M. leonina Moroteuthis knipovitchi 3.45 22.13 3 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1996 

M. leonina Psychroteuthis glacialis 3.45 19.78 231 Daneri et al. 2000 South Shetland Islands, S Atlantic 1995-1996 

M. leonina Pareledone charcoti 3.45 4.80 5 4.20 4.80 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

M. leonina Pareledone cf. charcoti 3.45 5.71 5 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1997 

M. leonina Pareledone cf. turqueti 3.45 7.25 1 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1999 

M. leonina Pareledone polymorpha 3.45 3.60 6 3.00 4.50 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 M. leonina Pareledone cf. polymorpha 3.45 4.51 1 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1998 

M. leonina Electrona subaspera 3.45 10.70 1 Green and Burton 1993 Macquarie Island, Southern Ocean 1988 

M. leonina Gymnoscopelus braueri 3.45 11.20 1 Green and Burton 1993 Heard Island, Indian Ocean 1988 

M. leonina Gymnoscopelus nicholsi 3.45 12.71 1.85 72 7.95 16.71 Daneri and Carlini 2002 South Shetland Islands, S Atlantic 1993-2000 

M. leonina Pleuragramma antarcticum 3.45 13.15 4.14 16 9.06 22.62 Daneri and Carlini 2002 South Shetland Islands, S Atlantic 1993-2000 

M. leonina Champsocephalus gunnari 3.45 30.11 6 Green and Burton 1993 Heard Island, Indian Ocean 1988 

M. leonina Channichthys rhinoceratus 3.45 37.83 4 Green and Burton 1993 Heard Island, Indian Ocean 1988 

M. leonina mixed fish taxa 3.45 6.23 27.82 Daneri and Carlini 2002 South Shetland Islands, S Atlantic 1993-2000 

M. monachus Trachurus sp. 2.40 30.00 50 Sergeant et al. 1979 Madeira, North Atlantic 1957 

M. monachus Pagrus africanus 2.40 35.00 1 Sergeant et al. 1979 Madeira, North Atlantic 1957 

O. rossii Galiteuthis armata 1.85 19.60 14 13.50 23.70 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

O. rossii Alluroteuthis antarctica 1.85 12.20 57 6.10 18.40 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

O. rossii Kondakovia longimana 1.85 41.20 1 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

O. rossii Moroteuthis knipovitchi 1.85 28.40 3 23.20 33.70 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

O. rossii Psychroteuthis glacialis 1.85 12.30 97 5.80 34.40 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

O. rossii Pleuragramma antarcticum 1.85 15.10 9 11.20 18.50 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

P. fasciata Boreogadus saida 1.60 15.87 92 12.00 21.00 Frost and Lowry 1980 Bering Sea 1976-1979 P. fasciata Theragra chalcogramma 1.60 10.26 447 6.00 20.00 Frost and Lowry 1980 Bering Sea 1976-1979 

P. fasciata Theragra chalcogramma 1.60 11.20 468 6.50 34.40 Frost and Lowry 1986 Bering Sea 1976-1978 

P. groenlandica Alosa pseudoharengus 1.70 10.10 3 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Clupea harengus 1.70 30.70 n.a.** 23.00 36.00 Hauksson and Bogason 1997 Iceland 1992-1993 

P. groenlandica Mallotus villosus 1.70 16.00 2244 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Mallotus villosus 1.70 13.20 30 8.80 15.70 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Osmerus mordax 1.70 12.00 4 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Salmo salar 1.70 29.10 1 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Arctozenus risso 1.70 13.00 1 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Nezumia bairdi 1.70 30.50 9 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Arctogadus glacialis 1.70 21.10 338 Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

P. groenlandica Boreogadus saida 1.70 13.40 648 4.80 23.60 Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

P. groenlandica Boreogadus saida 1.70 13.30 7 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Gadus morhua 1.70 23.80 2 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Gadus morhua 1.70 24.70 25 15.00 40.00 Nilssen et al. 1990 Ullsford, Norway 1986 

P. groenlandica Gadus morhua 1.70 33.40 99 5.00 65.00 Nilssen et al. 1990 Ullsford, Norway 1988 

P. groenlandica Gadus morhua 1.70 14.00 43 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Gadus morhua 1.70 15.90 38 3.00 48.00 Hauksson and Bogason 1997 Iceland 1992-1993 

P. groenlandica Gadus ogac 1.70 14.00 3 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Melanagrammus aeglefinus 1.70 16.80 1 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Enchelyopus cimbrius 1.70 25.90 5 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Urophycis tenuis 1.70 27.30 1 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Merluccius albidus 1.70 29.60 24 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Merluccius bilinearis 1.70 26.00 1 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Sebastes sp. 1.70 11.40 257 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

P. groenlandica Cottidae 1.70 12.00 7 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Scophthalmus aquosus 1.70 23.50 1 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. groenlandica Pleuronectidae 1.70 25.40 5 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

P. hispida Gammarus setosus 1.30 1.53 35 1.20 2.00 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

P. hispida Themisto libellula 1.30 1.66 279 1.40 3.40 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 P. hispida Thysanoessa inermis 1.30 1.75 189 1.20 2.60 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 P. hispida Sabinea septemcarinata 1.30 5.02 19 3.60 5.60 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

225

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

P. hispida Pandalus borealis 1.30 7.47 51 4.80 9.20 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

P. hispida Boreogadus saida 1.30 10.50 4047 4.80 23.60 Finley et al. 1983, Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

P. hispida Boreogadus saida 1.30 10.90 2.00 4847 4.90 16.10 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 P. hispida Sebastes marinus 1.30 45.00 3286 10.00 120.00 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

P. hispida Myoxocephalus scorpius 1.30 11.14 35 8.00 18.00 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 P. largha Boreogadus saida 1.75 14.90 326 7.60 20.50 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

P. largha Eleginus gracilis 1.75 16.70 131 6.20 25.10 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

P. largha Theragra chalcogramma 2.25 10.90 21 8.00 15.00 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

P. largha Theragra chalcogramma 1.75 10.90 21 8.00 15.00 Frost and Lowry 1986 Bering Sea 1978 

P. largha Cottidae 1.75 10.00 19 3.30 16.50 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 P. largha Lycodes sp. 1.75 27.30 6 22.30 30.90 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

P. vitulina Illex illecebrosus 1.55 17.60 139 5.00 20.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

P. vitulina Illex illecebrosus 1.55 19.10 51 15.00 25.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 

P. vitulina Clupea harengus 1.55 22.00 197 10.00 35.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

P. vitulina Clupea harengus 1.55 25.70 84 15.00 35.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 P. vitulina Engraulis mordax 1.55 12.90 81 4.00 15.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 

P. vitulina Osmerus eperlanus 1.55 10.20 41 Behrends 1982 Waddensea, North Sea 1975-1981 

P. vitulina Gadus morhua 1.55 14.40 49 Behrends 1982 Waddensea, North Sea 1975-1981 

P. vitulina Gadus morhua 1.55 35.20 33 5.00 50.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

P. vitulina Gadus morhua 1.55 19.00 46 10.00 35.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 

P. vitulina Microgadus proximus 1.55 14.00 8 4.00 23.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 P. vitulina Pollachius virens 1.55 20.50 19 5.00 30.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

P. vitulina Pollachius virens 1.55 17.70 122 10.00 35.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 

P. vitulina Theragra chalcogramma 1.55 31.80 23 10.30 56.30 Frost and Lowry 1986 Pribilof Islands, Bering Sea 1979 

P. vitulina Theragra chalcogramma 1.55 10.60 12 8.20 12.60 Frost and Lowry 1986 Bering Sea 1981 

P. vitulina Theragra chalcogramma 1.55 19.20 9.60 2180 4.20 53.20 Pitcher 1981 Gulf of Alaska 1975-1978 

P. vitulina Porichthys notatus 1.55 21.80 73 7.00 39.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 P. vitulina Leptocottus armatus 1.55 11.00 85 4.00 21.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 P. vitulina Leptocottus armatus 1.55 14.03 72 5.00 25.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 P. vitulina Genyonemus lineatus 1.55 22.20 40 13.00 29.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 

P. vitulina Cymatogaster aggregata 1.55 8.50 31 6.50 11.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Anarrhichthys ocellatus 200.00 1 Baldridge and Rogers 1991 Point Lobos State Reserve, CA 1986 

P. vitulina Ammodytes hexapterus 1.55 9.50 621 8.00 13.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 P. vitulina Acanthogobius flavimanus 1.55 9.20 2.30 1021 3.50 20.20 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 

P. vitulina Pomatoschistus spp. 1.55 3.84 641 Behrends 1982 Waddensea, North Sea 1975-1981 

P. vitulina Citharichthys sordidus 1.55 6.00 74 4.00 21.50 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Citharichthys stigmaeus 1.55 6.50 29 5.00 10.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Glyptocephalus zachirus 1.55 16.50 113 5.00 28.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Isopsetta isolepis 1.55 18.00 10 7.00 26.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Limanda limanda 1.55 8.13 219 < 7.0 20.00 Behrends 1982 Waddensea, North Sea 1975-1981 

P. vitulina Lyopsetta exilis 1.55 13.50 21 8.00 20.50 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Microstomus pacificus 1.55 15.00 62 7.00 21.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Parophrys vetulus 1.55 17.20 26 12.00 32.00 Harvey et al. 1995 Elkhorn Slough, NE Pacific 1975-1977 

P. vitulina Parophrys vetulus 1.55 7.00 140 4.00 24.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

P. vitulina Platichthys flesus 1.55 16.00 139 Behrends 1982 Waddensea, North Sea 1975-1981 

P. vitulina Pleuronectes platessa 1.55 10.00 199 Behrends 1982 Waddensea, North Sea 1975-1981 P. vitulina Psettichthys melanostictus 1.55 14.00 2 10.00 18.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 Balaenoptera spp. Euphausia superba 23.50 2.66 2745 1.10 6.00 Marr 1962 South Georgia Island, S Atlantic 1932-1933 

Balaenoptera spp. Euphausia superba 23.50 3.38 3562 1.10 6.00 Marr 1962 South Georgia Island, S Atlantic 1939-1940 

Balaenoptera spp. Euphausia superba 23.50 3.94 39081 1.10 6.00 Marr 1962 South Georgia Island, S Atlantic 1929-1930 

Balaenoptera spp. Euphausia superba 23.50 5.00 >2000 1.10 6.50 Mackintosh 1974 Southern Ocean 1950s?? 

B. acutorostrata Theragra chalcogramma 7.30 14.50 121 11.80 17.50 Frost and Lowry 1986 Bering Sea 1975 

B. pysalus Theragra chalcogramma 20.70 <=30.00 n.a.** Nemoto 1959 N Pacific 1952-1958 

M. novaeangliae Theragra chalcogramma 13.00 >=30.00 n.a.** Nemoto 1959 N Pacific 1952-1958 

B. bairdii Taonius borealis 11.10 25.20 240 14.26 49.11 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

SE 0.17 SE 0.36 

SE 0.36 SE 0.51 

SE 2.08 SE 0.75 SE 1.56 SE 0.38 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

226

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

B. bairdii Taonius borealis 11.10 32.09 85 13.85 45.06 Walker et al. 2002 Sea of Okhotsk 1988-1989 

B. bairdii Berryteuthis magister 11.10 21.37 697 11.12 33.43 Walker et al. 2002 Sea of Okhotsk 1988-1989 

B. bairdii Gonatopsis borealis 11.10 17.70 56 9.06 31.02 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

B. bairdii Albatrossia pectoralis 11.10 21.28 5 10.18 34.58 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

B. bairdii Albatrossia pectoralis 11.10 21.91 7 16.90 28.16 Walker et al. 2002 Sea of Okhotsk 1988-1989 

B. bairdii Coryphaenoides acrolepis 9.95 73.30 11.08 72 45.00 95.00 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

B. bairdii Coryphaenoides acrolepis 11.10 22.07 55 4.06 28.74 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

B. bairdii Coryphaenoides cinereus 11.10 11.92 330 6.99 17.92 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

B. bairdii Coryphaenoides cinereus 10.32 35.84 3.1 3 30.00 35.00 Ohizumi et al. 2003 Abahiri, Japan, Sea of Okhotsk Jul - Aug 1999 

B. bairdii Coryphaenoides cinereus 9.95 35.33 4.25 40 25.00 45.00 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

B. bairdii Coryphaenoides cinereus 11.10 10.85 5 9.13 12.13 Walker et al. 2002 Sea of Okhotsk 1988-1989 

B. bairdii Coryphaenoides longifilis 9.95 75.70 6.54 158 55.00 100.00 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

B. bairdii Antimora microlepis 11.10 46.94 13 28.74 83.80 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

B. bairdii Laemonema longipes 11.10 51.01 766 18.73 71.26 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

B. bairdii Laemonema longipes 10.32 17.79 6.3 27 5.00 30 Ohizumi et al. 2003 Abahiri, Japan, Sea of Okhotsk Jul - Aug 1999 

B. bairdii Laemonema longipes 9.95 36.04 9.09 55 15.00 55 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

B. bairdii Laemonema longipes 11.10 35.82 41 30.76 42.78 Walker et al. 2002 Sea of Okhotsk 1988-1989 

B. bairdii Theragra chalcogramma 9.95 47.32 8.44 13 35.00 60.00 Ohizumi et al. 2003 Abahiri, Japan, Sea of Okhotsk Jul - Aug 1999 C. commersonii Loligo gahi 1.48 7.60 n.a.** 2.00 18.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

C. commersonii Illex argentinus 1.48 11.00 n.a.** 6.00 28.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

C. commersonii Merluccius hubbsi 1.48 9.90 n.a.** 0.10 20.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

C. heavisidii Loligo vulgaris reynaudii 1.57 17.40 4.70 11 9.30 24.40 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

C. heavisidii Sardinops sagax = S. ocellatus 1.57 20.60 2 20.60 20.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

C. heavisidii Engraulis encrasicolus 1.57 10.90 1 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 C. heavisidii Merluccius spp. 1.57 19.50 4.20 160 4.90 28.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 C. heavisidii Trachurus capensis 1.57 17.10 2.40 23 10.70 22.20 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 C. heavisidii Sufflogobius bibarbatus 1.57 5.30 2.00 n.a.** Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

D. leucas Boreogadus saida 4.25 14.54 125 9.00 21.50 Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

D. leucas Eleginus gracilis 4.25 12.48 530 5.50 30.50 Seaman et al. 1982 Bering Sea 1978 

D. leucas Pungitius pungitius 4.25 3.20 1 Seaman et al. 1982 Bering Sea and Chukchi Sea 1978 

D. delphis Loligo vulgaris reynaudii 2.08 17.30 6.80 322 2.50 34.50 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

D. delphis Etrumeus whiteheadi 2.08 16.20 2.70 171 8.50 23.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 D. delphis Sardinops sagax = S. ocellatus 2.08 19.00 1.80 163 12.60 24.30 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

D. delphis Engraulis encrasicolus 2.08 11.20 1.80 413 5.30 18.10 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

D. delphis Merluccius spp. 2.08 18.10 8.00 203 5.10 39.30 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 D. delphis Trachurus capensis 2.08 16.20 6.20 134 4.60 34.30 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 Delphinus sp. Semirossia tenera 2.15 1.70 69 1.00 2.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

Delphinus sp. Loligo plei 2.15 18.00 2 17.70 18.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Delphinus sp. Loligo sanpaulensis 2.15 5.00 58 2.80 8.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Delphinus sp. Illex argentinus 2.15 5.90 20 2.10 27.60 dos Santos and Haimovici 2001 Brazil 1985-1998 F. attenuata Loligo plei 2.35 14.90 2 13.60 16.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

F. attenuata Illex argentinus 2.35 22.70 2 18.70 26.70 dos Santos and Haimovici 2001 Brazil 1985-1998 

F. attenuata Ornithoteuthis antillarum 2.35 13.10 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

F. attenuata Merluccius spp. 2.33 6.30 0.50 6 5.70 7.20 Sekiguchi et al. 1992 Benguela, South Africa 1990 

Globicephala sp. Loligo pealeii 6.00 13.40 68 Overholtz and Waring 1991 Mid-Atlantic Bight, USA 1989 

Globicephala sp. Scomber scombrus 6.00 36.32 22 Overholtz and Waring 1991 Mid-Atlantic Bight, USA 1989 

G. macrorhynchus Loligo opalescens 4.40 13.80 11 11.00 17.09 Sinclair 1992 S California Bight, NE Pacific 1977 

G. macrorhynchus Loligo opalescens 4.63 12.80 11 11.20 14.20 Seagars and Henderson 1985 Channel Islands, NE Pacific 1980 G. macrorhynchus Loligo opalescens 5.52 16.20 609 8.70 23.20 Sinclair 1992 S California Bight, NE Pacific 1969-1977 

G. melas Loligo gahi 4.06 12.80 34 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

G. melas Loligo pealeii 3.40 16.07 145 5.00 42.00 Gannon et al. 1997b Mid-Atlantic Bight, USA 1973-1993 

G. melas Loligo pealeii 5.90 19.60 578 3.70 43.60 Gannon et al. 1997a NW Atlantic 1989-1991 

G. melas Ancistrocheirus lesueurii 5.90 17.20 3 6.10 24.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

G. melas Brachioteuthis riisei 4.06 13.60 9 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 G. melas Chiroteuthis veranyi 4.06 16.10 5 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

227

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

G. melas Galiteuthis armata 4.06 26.80 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 G. melas Mesonychoteuthis hamiltoni 4.06 22.00 7 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

G. melas Gonatus antarcticus 4.06 15.10 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

G. melas Histioteuthis eltaninae 4.06 5.20 32 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

G. melas Histioteuthis sp. 5.90 5.60 68 4.80 11.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

G. melas Lycoteuthis lorigera 5.90 11.00 45 9.30 14.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

G. melas Alluroteuthis antarctica 4.06 29.30 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 G. melas Octopoteuthis sp. 5.90 14.40 6 13.00 17.30 dos Santos and Haimovici 2001 Brazil 1985-1998 G. melas Illex argentinus 5.90 22.10 6 15.00 33.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

G. melas Todarodes sagittatus 1.50 17.00 n.a.** 25.00 Desportes and Mouritsen 1988 Faroe Islands, NE Atlantic 1986-1987 G. melas Todarodes sagittatus 3.50 20.00 n.a.** 50.00 Desportes and Mouritsen 1988 Faroe Islands, NE Atlantic 1986-1987 G. melas Moroteuthis ingens 4.06 21.20 19 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

G. melas Octopodidae 4.06 16.00 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 G. melas Squalus acanthias 3.40 75.00 6 Gannon et al. 1997b Mid-Atlantic Bight, USA 1973-1993 

G. melas Scomber scombrus 5.90 35.80 45 26.50 42.10 Gannon et al. 1997a NW Atlantic 1989-1991 

G. griseus Loligo vulgaris reynaudii 2.79 13.00 7.30 67 2.10 32.90 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 G. griseus Ancistrocheirus lesueurii 3.33 15.80 55 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Bathyteuthis abyssicola 3.33 4.00 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Brachioteuthis sp. 3.33 7.30 7 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Chiroteuthis sp. 3.33 8.00 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Planctoteuthis sp. 3.33 5.80 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Megalocranchia fisheri A 3.33 20.80 240 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Megalocranchia sp. B 3.33 23.20 45 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Abraliopsis sp. A 3.33 4.40 241 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Abraliopsis sp. B 3.33 3.20 46 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Abraliopsis sp. C 3.33 3.40 19 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Abraliopsis sp. D 3.33 5.30 12 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Enoploteuthis cf. anaspis 3.33 7.70 208 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Enoploteuthis sp. A 3.33 9.50 279 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Histioteuthis sp. A 3.33 3.10 10 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Pholidoteuthis boschmai 3.33 19.60 2 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Lycoteuthis lorigera 3.33 6.80 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Octopoteuthis spp. 3.33 5.70 3 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Octopoteuthis spp. 3.33 12.70 3 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Ommastrephes bartramii 3.33 35.40 17 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Onychoteuthis sp. 1 3.33 8.10 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Onychoteuthis sp. 2 3.33 20.30 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Pyroteuthis addolux 3.33 4.50 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

G. griseus Trachurus capensis 2.79 32.60 1 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

H. planifrons Bathyteuthis abyssicola 6.55 7.33 4.61 4 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Bathyteuthis abyssicola 7.20 3.60 6 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Batoteuthis skolops 7.20 9.60 6 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Brachioteuthis riisei 7.20 43.40 32 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Brachioteuthis sp. 6.30 8.57 0.86 5 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Chiroteuthis veranyi 7.20 12.40 5 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Chiroteuthis sp. 6.30 10.38 1.05 74 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Chiroteuthis sp. 6.49 10.61 1.34 388 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Galiteuthis armata 7.20 27.30 154 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Galiteuthis glacialis 6.30 19.52 1.54 375 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Galiteuthis glacialis 6.49 19.25 1.75 394 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Liocranchia reinhardti 6.55 24.90 3.33 9 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Liocranchia sp. 6.43 36.01 21.15 2 Sekiguchi et al. 1993 South Africa, east coast 1975 

H. planifrons Liocranchia sp. 6.30 25.40 4.46 334 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Megalocranchia sp. 6.49 19.65 5.98 77 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Mesonychoteuthis hamiltoni 6.30 92.51 16.12 5 Slip et al. 1995 Heard Island, Indian Ocean 1992 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

228

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

H. planifrons Mesonychoteuthis hamiltoni 6.49 69.58 25.78 65 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Mesonychoteuthis hamiltoni 7.20 96.40 22 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Taonius megalops 6.43 40.55 4.92 2 Sekiguchi et al. 1993 South Africa, east coast 1975 H. planifrons Taonius pavo 6.30 41.55 7.18 178 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Taonius pavo 6.49 34.91 8.27 614 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Taonius pavo 7.20 37.90 1663 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Teuthowenia megalops 6.55 22.17 4.66 160 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Teuthowenia pellucida 6.43 18.55 1.25 8 Sekiguchi et al. 1993 South Africa, east coast 1975 H. planifrons Teuthowenia pellucida 7.20 18.40 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Cycloteuthis sp. 6.55 46.19 1.32 2 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Discoteuthis discus 6.55 10.33 0.85 60 Sekiguchi et al. 1993 South Africa, west coast 1990 H. planifrons Discoteuthis cf. laciniosa 7.20 7.50 13 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Enoploteuthidae 6.55 5.01 1 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Enoploteuthis sp. 7.20 18.00 2 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Gonatus antarcticus 6.30 20.54 2.07 191 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Gonatus antarcticus 6.49 25.32 5.93 304 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Gonatus antarcticus 7.20 21.50 274 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Gonatidae 6.43 23.63 3.86 3 Sekiguchi et al. 1993 South Africa, east coast 1975 H. planifrons Histioteuthis bonnellii corpuscula 6.43 4.90 0.73 6 Sekiguchi et al. 1993 South Africa, east coast 1975 

H. planifrons Histioteuthis eltaninae 6.30 4.85 0.46 77 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Histioteuthis eltaninae 6.49 5.90 0.90 132 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Histioteuthis eltaninae 7.20 5.50 752 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Histioteuthis machrohista 6.43 4.42 0.77 8 Sekiguchi et al. 1993 South Africa, east coast 1975 H. planifrons Histioteuthis meleagroteuthis 6.43 5.49 0.73 397 Sekiguchi et al. 1993 South Africa, east coast 1975 

H. planifrons Histioteuthis sp. 6.49 5.40 1.71 506 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Pholidoteuthis boschmai 6.49 29.60 2.51 7 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Lycoteuthis lorigera 6.55 8.53 1 Sekiguchi et al. 1993 South Africa, west coast 1990 H. planifrons Mastigoteuthis sp. A 7.20 18.80 4 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Mastigoteuthis sp. 6.55 10.79 1.48 20 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Mastigoteuthis sp. 6.30 8.57 0.57 12 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Alluroteuthis antarctica 6.30 14.41 1.51 49 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Alluroteuthis antarctica 6.49 40.13 14.06 39 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Alluroteuthis antarctica 7.20 30.80 18 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Octopoteuthis rugosa 6.55 15.02 2.20 17 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Octopoteuthis sp. 6.43 16.64 2.34 8 Sekiguchi et al. 1993 South Africa, east coast 1975 

H. planifrons Taningia danae 6.49 18.92 13.04 36 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Taningia danae 7.20 44.40 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Martialia hyadesi 6.30 22.60 1.52 65 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Martialia hyadesi 7.20 30.10 25 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? H. planifrons Todarodes sagittatus 6.55 39.82 1 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Todaropsis eblanae 6.43 7.14 1 Sekiguchi et al. 1993 South Africa, east coast 1975 

H. planifrons Kondakovia longimana 6.55 46.94 6.95 59 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Kondakovia longimana 6.30 49.36 9.67 99 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Kondakovia longimana 7.20 10.40 77 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Moroteuthis ingens 6.55 70.36 5.02 5 Sekiguchi et al. 1993 South Africa, west coast 1990 H. planifrons Moroteuthis ingens 7.20 67.20 13 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Moroteuthis knipovitchi 6.30 24.70 8.85 37 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Moroteuthis knipovitchi 6.49 22.10 3.49 34 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Moroteuthis knipovitchi 7.20 17.00 16 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

H. planifrons Moroteuthis robsoni 6.49 20.71 8.36 60 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Moroteuthis sp. 6.49 22.42 4.42 74 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

H. planifrons Onychoteuthis banksii 6.55 21.36 9.73 8 Sekiguchi et al. 1993 South Africa, west coast 1990 H. planifrons Onychoteuthis sp. 6.55 29.26 5.24 10 Sekiguchi et al. 1993 South Africa, west coast 1990 

H. planifrons Psychroteuthis glacialis 6.30 9.87 1.96 1513 Slip et al. 1995 Heard Island, Indian Ocean 1992 

H. planifrons Dissostichus eleginoides 6.30 35.23 16.34 3 Slip et al. 1995 Heard Island, Indian Ocean 1992 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

229

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

K. breviceps Loligo vulgaris reynaudii 2.52 11.60 5.20 21 5.30 20.90 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 K. breviceps Chiroteuthis veranyi 3.05 10.40 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. breviceps Chiroteuthis cf. veranyi 3.05 15.82 1 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Taonius pavo 3.05 30.94 70 17.63 49.02 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Teuthowenia sp. 3.05 18.03 5 15.04 20.55 Wang et al. 2002 Taiwan 1998-2000 K. breviceps Discoteuthis sp. 3.05 14.01 2 8.71 19.31 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Abralia sp. 3.05 3.60 5 3.00 4.30 dos Santos and Haimovici 2001 Brazil 1985-1998 K. breviceps Enoploteuthis chunii 3.05 7.17 180 5.59 8.78 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Histioteuthis hoylei 3.05 10.47 7 7.81 15.52 Wang et al. 2002 Taiwan 1998-2000 K. breviceps Histioteuthis miranda 3.05 9.44 17 6.39 10.83 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Histioteuthis reversa 3.05 5.31 6 4.86 5.79 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Histioteuthis spp. 3.05 7.10 16 5.70 9.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. breviceps Pholidoteuthis boschmai 3.05 29.24 5 25.70 36.63 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Lycoteuthis lorigera 3.05 8.90 17 7.50 12.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. breviceps Idioteuthis hjorti 3.05 11.00 17 7.58 14.62 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Octopoteuthis cf. deletron 3.05 16.65 6 14.26 18.57 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Octopoteuthis sp. 3.05 16.20 4 14.50 19.70 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. breviceps Octopoteuthis sp. 3.05 16.89 2 16.01 17.78 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Eucleoteuthis luminosa 3.05 13.35 4 12.02 14.65 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Illex argentinus 3.05 21.70 25 14.60 28.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. breviceps Ornithoteuthis antillarum 3.05 8.10 2 6.90 9.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. breviceps Ornithoteuthis volatilis 3.05 20.12 13 13.28 25.24 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Sthenoteuthis oualaniensis 3.05 23.35 20 16.48 30.53 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Moroteuthis loennbergi 3.05 17.58 11 11.15 21.21 Wang et al. 2002 Taiwan 1998-2000 

K. breviceps Onychoteuthis banksii 3.05 16.63 2 10.77 22.49 Wang et al. 2002 Taiwan 1998-2000 K. breviceps Engraulis encrasicolus 2.52 10.00 3 10.00 10.00 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 

K. breviceps Merluccius spp. 2.52 21.30 2.60 11 16.20 25.50 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 

K. sima Semirossia tenera 2.40 2.00 2 1.80 2.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. sima Loligo vulgaris reynaudii 2.23 7.10 1.40 11 5.30 8.90 Sekiguchi et al. 1992 Benguela, South Africa 1975-1988 

K. sima Chiroteuthis veranyi 2.40 11.90 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. sima Chtenopteryx sicula 2.40 7.07 1 Wang et al. 2002 Taiwan 1998-2000 

K. sima Chtenopteryx sp. 2.40 7.04 6 6.36 7.91 Wang et al. 2002 Taiwan 1998-2000 K. sima Galiteuthis armata 2.40 7.93 2 7.34 8.52 Wang et al. 2002 Taiwan 1998-2000 

K. sima Leachia dislocata 2.40 2.27 2 2.25 2.29 Wang et al. 2002 Taiwan 1998-2000 

K. sima Taonius pavo 2.40 20.59 14 10.01 34.83 Wang et al. 2002 Taiwan 1998-2000 

K. sima Discoteuthis sp. 2.40 13.33 1 Wang et al. 2002 Taiwan 1998-2000 

K. sima Abralia redfieldi 2.40 2.90 23 2.20 3.60 dos Santos and Haimovici 2001 Brazil 1994, 1998 

K. sima Enoploteuthis chunii 2.40 7.13 72 5.63 8.84 Wang et al. 2002 Taiwan 1998-2000 

K. sima Histioteuthis miranda 2.40 8.13 13 3.04 10.63 Wang et al. 2002 Taiwan 1998-2000 

K. sima Histioteuthis spp. 2.40 6.00 94 2.60 13.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. sima Lycoteuthis lorigera 2.40 9.40 7 8.10 10.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. sima Octopoteuthis sp. 2.40 14.70 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. sima Eucleoteuthis luminosa 2.40 13.06 4 6.98 18.22 Wang et al. 2002 Taiwan 1998-2000 

K. sima Illex argentinus 2.40 22.40 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. sima Ornithoteuthis antillarum 2.40 5.70 24 3.00 9.60 dos Santos and Haimovici 2001 Brazil 1985-1998 

K. sima Ornithoteuthis volatilis 2.40 17.80 3 9.64 23.10 Wang et al. 2002 Taiwan 1998-2000 

K. sima Sthenoteuthis oualaniensis 2.40 15.18 1 Wang et al. 2002 Taiwan 1998-2000 

K. sima Moroteuthis loennbergi 2.40 10.77 1 Wang et al. 2002 Taiwan 1998-2000 

K. sima Onychoteuthis banksii 2.40 13.76 3 9.25 16.20 Wang et al. 2002 Taiwan 1998-2000 K. sima Engraulis encrasicolus 2.23 11.60 2.80 7 9.30 17.60 Sekiguchi et al. 1992 Benguela, South Africa 1975-1988 

K. sima Merluccius spp. 2.23 21.60 3.20 20 16.60 26.80 Sekiguchi et al. 1992 Benguela, South Africa 1975-1988 

L. hosei Loligo sanpaulensis 2.33 12.90 19 5.10 21.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

L. hosei Abraliopsis cf. lineata 2.33 5.09 30 3.70 6.70 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

L. hosei Argonauta nodosa 2.33 0.60 1 dos Santos and Haimovici 2001 Brazil 1985-1998 L. hosei Acanthephyra carinata 2.33 3.76 5 3.00 5.00 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

SE 0.14 SE 0.35 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

230

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

L. hosei Acanthephyra quadrispinsoa 2.33 1.47 19 1.00 1.90 Dolar et al. 2003 Sulu Sea, Philippines n.a. L. hosei Notostomus elegans 2.33 3.74 5 2.30 4.50 Dolar et al. 2003 Sulu Sea, Philippines n.a. L. hosei Gonostoma cf. elongatum 2.33 184 8.30 22.50 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Argyropelecus affinis 2.33 177 4.00 7.00 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Maurolicus cf. muelleri 2.33 112 2.50 3.50 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Ichthyococcus cf. irregularis 2.33 120 4.00 6.90 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Chauliodus cf. barbatus 2.33 18.00 238 8.50 20.00 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Rosenblattichthys cf. volucris 2.33 67 6.50 14.00 Robison and Craddock 1983 E Tropical Pacific 1972 

L. hosei Scopelarchus guentheri 2.33 127 8.00 16.00 Robison and Craddock 1983 E Tropical Pacific 1972 

L. hosei Paralepididae 2.33 87 20.00 30.00 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Evermanella cf. ahlstromi 2.33 119 3.50 9.00 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Ceratoscopelus warmingii 2.33 5.93 171 4.20 6.90 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

L. hosei Lampadena luminosa 2.33 255 7.50 10.50 Robison and Craddock 1983 E Tropical Pacific 1972 

L. hosei Lampadena luminosa 2.33 10.20 245 8.10 14.60 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

L. hosei Lampadena sp. 2.33 9 6.00 7.00 Robison and Craddock 1983 E Tropical Pacific 1972 

L. hosei Lampanyctus cf. idostigma 2.33 208 6.00 9.50 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Lampanyctus cf. nobilis 2.33 103 7.00 12.00 Robison and Craddock 1983 E Tropical Pacific 1972 

L. hosei Myctophum nitidulum 2.33 4.50 3 3.70 5.10 Dolar et al. 2003 Sulu Sea, Philippines n.a. L. hosei Symbolophorus evermanni 2.33 6.48 43 5.80 7.60 Dolar et al. 2003 Sulu Sea, Philippines n.a. L. hosei Melanonus sp. 2.33 71 12.00 22.00 Robison and Craddock 1983 E Tropical Pacific 1972 

L. hosei Scopelogadus mizolepis bispinosus 2.33 143 3.00 6.50 Robison and Craddock 1983 E Tropical Pacific 1972 

L. hosei Diretmus argenteus 2.33 363 6.00 28.50 Robison and Craddock 1983 E Tropical Pacific 1972 L. hosei Cubiceps pauciradiatus 2.33 11.30 55 8.50 14.20 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

L. obliquidens Loligo opalescens 2.05 4.70 17.60 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

L. obliquidens Ommastrephes bartramii 2.05 32.10 8 27.00 39.50 Walker and Jones 1993 N Pacific 1990 

L. obscurus Loligo gahi 2.05 7.50 n.a.** 2.00 23.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

L. obscurus Loligo vulgaris reynaudii 1.71 12.90 6.60 23 1.70 27.00 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 L. obscurus Illex argentinus 2.05 10.40 n.a.** 4.00 28.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 L. obscurus Etrumeus whiteheadi 1.71 11.20 3.50 5 9.20 17.40 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

L. obscurus Sardinops sagax = S. ocellatus 1.71 16.60 3.60 21 13.80 25.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

L. obscurus Engraulis encrasicolus 1.71 9.10 1.80 188 4.40 15.80 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 L. obscurus Merluccius hubbsi 2.05 20.00 n.a.** 0.10 45.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 L. obscurus Merluccius spp. 1.71 16.50 7.20 216 4.80 42.10 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

L. obscurus Trachurus capensis 1.71 9.90 3.00 723 4.20 24.90 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

M. densirostris Merluccius spp. 3.52 6.90 1.20 6 5.90 8.50 Sekiguchi et al. 1992 Benguela, South Africa 1981 

M. layardii Chiroteuthis sp. 6.15 11.17 0.42 3 Sekiguchi et al. 1996 New Zealand 1966 

M. layardii Chiroteuthis sp. 5.41 10.64 1.12 9 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Galiteuthis armata 5.41 19.60 2 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Megalocranchia sp. 5.41 25.40 6.44 3 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Taonius pavo 6.15 25.8 1 Sekiguchi et al. 1996 New Zealand 1966 

M. layardii Taonius pavo 5.41 30.56 10.84 39 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Taonius sp. 5.41 27.34 1.38 10 Sekiguchi et al. 1996 South Africa 1965-1995 M. layardii Teuthowenia megalops 5.41 25.27 2 Sekiguchi et al. 1996 South Africa 1965-1995 M. layardii Teuthowenia pellucida 6.15 22.22 0.86 2 Sekiguchi et al. 1996 New Zealand 1966 

M. layardii Teuthowenia pellucida 5.41 18.12 2.39 12 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Teuthowenia sp. 5.41 34.25 1 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Cycloteuthis sp. 5.41 24.80 1 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Discoteuthis discus 5.41 9.19 1 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Discoteuthis laciniosa 5.41 12.78 1.97 5 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Gonatus antarcticus 5.41 34.49 0.66 5 Sekiguchi et al. 1996 South Africa 1965-1995 M. layardii Gonatidae 5.41 12.17 4.55 2 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Histioteuthis machrohista 5.41 6.30 1.12 21 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Histioteuthis meleagroteuthis 6.15 7.75 1 Sekiguchi et al. 1996 New Zealand 1966 

M. layardii Histioteuthis meleagroteuthis 5.41 5.22 0.30 5 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Histioteuthis miranda 6.15 15.96 1 Sekiguchi et al. 1996 New Zealand 1966 

SE 0.06 

SE 0.43 

SE 0.04 SE 0.09 

SE 0.44 

SE 0.05 SE 0.17 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

231

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

M. layardii Histioteuthis sp. 5.41 5.83 1.25 58 Sekiguchi et al. 1996 South Africa 1965-1995 M. layardii Mastigoteuthis sp. 5.41 11.37 1.18 22 Sekiguchi et al. 1996 South Africa 1965-1995 M. layardii Octopoteuthis rugosa 5.41 7.93 2 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Octopoteuthis sp. 5.41 13.48 2 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Ommastrephidae 5.41 6.00 1 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Moroteuthis knipovitchi 5.41 5.80 4.30 3 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Moroteuthis robsoni 5.41 28.40 1 Sekiguchi et al. 1996 South Africa 1965-1995 

M. layardii Vampyroteuthis infernalis 5.41 5.00 1.26 3 Sekiguchi et al. 1996 South Africa 1965-1995 

M. monoceros Gonatus fabricii 4.45 22.00 5473 6.80 40.25 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 

M. monoceros Boreomysis sp. 4.45 n.a.** 3.00 5.00 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 M. monoceros Arctogadus glacialis 4.45 18.60 376 9.30 26.60 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 

M. monoceros Arctogadus glacialis 4.45 20.00 616 10.00 40.00 Heide-Jorgensen et al. 1994 Inglefield Bredning, Greenland 1984, 1985 

M. monoceros Boreogadus saida 4.45 11.30 607 4.80 21.50 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 

M. monoceros Boreogadus saida 4.45 15.70 557 12.00 24.00 Heide-Jorgensen et al. 1994 Inglefield Bredning, Greenland 1984, 1985 

M. monoceros Reinhardtius hippoglossoides 4.45 n.a.** 45.00 61.00 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 

M. monoceros Reinhardtius hippoglossoides 4.45 35.70 9.00 140 13.80 54.70 Laidre and Heide-Jorgensen 2005 Disko Bay, Greenland 2002-2003 

O. orca Loligo plei 7.65 14.30 2 12.40 16.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

O. orca Loligo sanpaulensis 7.65 12.60 10 5.80 18.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

O. orca Ancistrocheirus lesueurii 7.65 29.30 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

O. orca Gonatus antarcticus 7.65 20.10 6 18.80 21.40 dos Santos and Haimovici 2001 Brazil 1985-1998 O. orca Histioteuthis spp. 7.65 9.60 7 6.90 13.70 dos Santos and Haimovici 2001 Brazil 1985-1998 

O. orca Lycoteuthis lorigera 7.65 11.00 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

O. orca Octopoteuthis sp. 7.65 16.50 4 15.60 19.00 dos Santos and Haimovici 2001 Brazil 1985-1998 O. orca Ommastrephes bartramii 7.65 28.70 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

O. orca Ornithoteuthis antillarum 7.65 4.40 1 dos Santos and Haimovici 2001 Brazil 1985-1998 P. phocoena Rossia macrosoma 1.65 4.40 0.20 2 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Sepietta oweniana 1.65 2.60 9 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Loligo opalescens 1.65 5.60 13.50 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

P. phocoena Loligo opalescens 1.65 8.14 1673 5.80 12.50 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. phocoena Loligo pealeii 1.65 12.90 3.00 8 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Loligo subulata 1.65 7.10 1.10 71 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Loligo vulgaris vulgaris 1.65 2.90 2 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Bathypolypus arcticus 1.65 5.20 1 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Myxine glutinosa 1.65 31.60 1 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Anguilla anguilla 1.65 26.40 10.00 9 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Clupea harengus 1.65 33.70 47 29.00 38.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 

P. phocoena Clupea harengus 1.65 26.50 136 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 

P. phocoena Clupea harengus 1.65 25.40 3.60 507 18.00 33.00 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Clupea harengus 1.65 26.10 4.90 103 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Clupea harengus pallasi 1.65 17.86 53 12.30 22.80 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. phocoena Sprattus sprattus 1.65 14.00 0.90 166 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Mallotus villosus 1.65 13.80 0.80 23 13.00 15.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 

P. phocoena Thaleichthys pacificus 1.65 15.30 3 12.40 18.10 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. phocoena Maurolicus muelleri 1.65 5.00 0.50 146 3.60 6.80 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Maurolicus weitzmani 1.65 5.00 0.40 5898 3.00 5.50 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Gadus morhua 1.65 29.00 5 26.00 32.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 

P. phocoena Gadus morhua 1.65 15.90 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 P. phocoena Gadus morhua 1.65 24.10 13.30 5 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Gadus morhua 1.65 28.10 5.80 30 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Melanagrammus aeglefinus 1.65 18.20 3.00 4 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Merlangius merlangus 1.65 18.70 4.00 65 14.00 29.00 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Merlangius merlangus 1.65 5.77 2.60 449 2.00 12.00 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Micromesistius poutassou 1.65 27.80 13.70 5 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Pollachius pollachius 1.65 5.60 0.10 2 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Pollachius virens 1.65 19.50 10.10 76 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

SE 0.32 

SE 1.72 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

232

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

P. phocoena Pollachius virens 1.65 17.40 6.40 91 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Theragra chalcogramma 1.65 18.64 3 18.50 18.80 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. phocoena Trisopterus esmarkii 1.65 12.40 2.20 381 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Trisopterus minutus 1.65 14.70 1.30 14 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Enchelyopus cimbrius 1.65 23.60 4.70 18 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Urophycis spp. 1.65 15.90 14.60 474 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Merluccius bilinearis 1.65 19.60 173 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 

P. phocoena Merluccius bilinearis 1.65 16.40 9.60 1605 3.00 38.00 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Merluccius merluccius 1.65 14.50 1.50 4 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Merluccius productus 1.65 35.30 15 33.80 37.10 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. phocoena Sebastes spp. 1.65 3.70 0.30 47 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Lycodopsis pacifica 1.65 10.43 1786 8.00 11.00 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. phocoena Ammodytes hexapterus 1.65 15.70 2 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. phocoena Ammodytidae 1.65 13.10 6.20 13 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Gobiidae 1.65 4.50 2.00 79 1.50 9.00 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

P. phocoena Scomber scombrus 1.65 37.70 1.60 10 35.00 40.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 

P. phocoena Scomber scombrus 1.65 29.60 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 P. phocoena Scomber scombrus 1.65 22.40 5.30 15 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Peprilus triacanthus 1.65 9.70 1.20 38 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

P. phocoena Citharichthys sordidus 1.65 15.20 10 12.50 17.60 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. dalli Loligo opalescens 1.95 4.70 17.40 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? P. dalli Loligo opalescens 1.95 9.52 44 6.90 12.70 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. dalli Taonius pavo 1.89 21.10 12 13.98 30.61 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Watasenia scintillans 1.89 3.39 462 2.44 4.51 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Berryteuthis magister 1.95 13.31 538 5.50 32.50 Walker 1996 Sea of Okhotsk 1988-1989 

P. dalli Berryteuthis magister 1.89 8.30 1.20 22 5.50 10.50 Ohizumi et al. 2000 Sea of Japan May-89 

P. dalli Berryteuthis magister 1.89 7.10 4.20 90 0.50 18.50 Ohizumi et al. 2000 Sea of Japan May-95 P. dalli Berryteuthis magister 1.89 11.50 3.60 69 2.50 18.50 Ohizumi et al. 2000 Sea of Japan May-96 P. dalli Berryteuthis magister 1.89 9.30 3.30 62 4.50 17.50 Ohizumi et al. 2000 Sea of Okhotsk Oct-94 

P. dalli Berryteuthis magister 1.89 10.60 4.20 120 4.50 19.50 Ohizumi et al. 2000 Sea of Okhotsk Aug-95 

P. dalli Berryteuthis magister 1.89 12.80 3.60 70 3.50 25.50 Ohizumi et al. 2000 Sea of Okhotsk Jun-88 

P. dalli Gonatopsis borealis 1.89 14.97 28 10.86 21.21 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Gonatopsis makko 1.89 9.96 647 2.10 28.91 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Gonatopsis octopedatus 1.89 9.11 560 1.00 15.14 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 P. dalli Gonatus berryi 1.89 13.99 10 4.87 22.48 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Gonatus madokai 1.89 10.12 459 2.36 17.10 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Gonatus onyx 1.89 8.59 191 5.22 10.76 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Gonatus sp. 1.89 8.24 872 4.00 14.77 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 P. dalli Todarodes pacificus 1.89 7.37 2125 3.09 22.21 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 P. dalli Clupea harengus pallasi 1.95 15.10 66 10.20 21.70 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. dalli Sardinops sagax melanostictus 1.95 18.01 1419 13.25 23.25 Walker 1996 Sea of Okhotsk 1988-1989 

P. dalli Sardinops sagax cf. melanosticus 1.89 17.89 372 12.53 22.46 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 P. dalli Engraulis japonicus 1.89 12.73 978 9.77 17.16 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Nansenia candida 1.95 7.07 6 2.70 14.80 Crawford 1981 N Pacific 1978-1979 

P. dalli Bathylagidae 1.95 9.43 60 5.00 17.90 Crawford 1981 N Pacific 1978-1979 

P. dalli Bathylagidae 1.89 11.77 315 11.12 12.46 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Thaleichthys pacificus 1.95 15.80 21 12.80 17.60 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. dalli Maurolicus muelleri 1.89 2.27 661 1.48 3.01 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Scopelosaurus harryi 1.95 15.34 37 7.50 21.50 Crawford 1981 N Pacific 1978-1979 

P. dalli Paralepididae 1.95 43.00 2 38.00 48.00 Crawford 1981 N Pacific 1978-1979 

P. dalli Myctophidae 1.95 6.54 166 3.50 10.50 Crawford 1981 N Pacific 1978-1979 

P. dalli Theragra chalcogramma 1.95 22.61 135 6.50 60.50 Walker 1996 Sea of Okhotsk 1988-1989 

P. dalli Theragra chalcogramma 1.89 33.64 639 15.08 52.48 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Theragra chalcogramma 1.95 25.90 113 12.70 36.30 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. dalli Merluccius productus 1.95 41.75 7 39.70 43.80 Walker et al. 1998 Washington and British Columbia 1990-1997 

SE 0.18 

SE 1.44 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

233

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

P. dalli Melamphaes sp. 1.95 8.38 8 7.80 9.00 Crawford 1981 N Pacific 1978-1979 

P. dalli Pleurogrammus azonus 1.89 21.60 154 13.70 28.94 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli Pleurogrammus monopterygius 1.95 13.10 1 Crawford 1981 N Pacific 1978-1979 P. dalli Lycodopsis pacifica 1.95 9.50 10175 8.00 10.50 Walker et al. 1998 Washington and British Columbia 1990-1997 P. dalli Ammodytes hexapterus 1.95 10.90 61 9.30 13.30 Walker et al. 1998 Washington and British Columbia 1990-1997 

P. dalli Ammodytes personatus 1.89 19.44 277 12.75 22.21 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

P. dalli mixed fish taxa 1.95 10.22 98 3.00 33.00 Crawford 1981 N Pacific 1978-1979 P. dalli mixed fish taxa 1.95 8.56 378 2.70 48.00 Crawford 1981 N Pacific 1978-1979 P. macrocephalus Ancistrocheirus lesueurii 15.75 20.45 60 14.00 30.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Ancistrocheirus lesueurii 10.00 21.66 73 10.00 40.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Ancistrocheirus lesueurii 15.75 16.40 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Ancistrocheirus lesueurii 12.40 23.00 204 16.70 28.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Architeuthis cf. dux 12.40 115.90 135 34.60 186.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Architeuthis sp. 15.75 107.75 4 82.00 151.00 Best 1999 Donkergat, South Africa 1962-1963 P. macrocephalus Architeuthis sp. 10.00 82.56 9 42.00 116.00 Best 1999 Donkergat, South Africa 1962-1963 P. macrocephalus Architeuthis sp. 15.75 166.30 2 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Chiroteuthis picteti 15.75 13.90 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Chiroteuthis sp. 12.40 12.60 168 10.90 20.70 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Helicocranchia sp. ? 12.40 25.80 4 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Liguriella sp. 12.40 12.90 1 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 P. macrocephalus Liocranchia reinhardti 12.40 25.80 4 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 P. macrocephalus Megalocranchia fisheri 15.75 96.00 36 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Megalocranchia sp. A 12.40 50.50 200 28.30 69.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Megalocranchia sp. G 12.40 81.70 30 49.50 140.80 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Taonius pavo 15.75 41.10 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Taonius pavo 12.40 41.20 12 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Teuthowenia maculata 12.40 29.90 200 19.20 37.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Teuthowenia megalops 12.40 30.10 200 20.40 35.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Teuthowenia megalops 12.35 23.71 110 14.50 31.50 Santos et al. 1999 North Sea 1990-1996 

P. macrocephalus Teuthowenia sp. 12.40 19.90 8 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Cycloteuthis akimushkini 15.75 38.90 18 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Cycloteuthis akimushkini 12.40 37.00 152 18.90 50.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Discoteuthis laciniosa 15.75 16.10 . 2 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Discoteuthis laciniosa 12.40 13.20 200 9.80 15.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Gonatus berryi 15.75 28.50 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Gonatus fabricii 14.80 21.80 2000 Clarke 1997 North Sea 1994 

P. macrocephalus Gonatus cf. fabricii 13.12 22.30 2116 15.50 28.50 Santos et al. 1999 North Sea 1990-1996 

P. macrocephalus Gonatus cf. fabricii 12.35 22.02 14468 14.50 29.50 Santos et al. 1999 North Sea 1990-1996 

P. macrocephalus Gonatus steenstrupi 12.40 14.90 200 11.50 18.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Histioteuthis arcturi 12.40 11.10 200 8.30 13.00 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Histioteuthis bonnellii bonnellii 12.40 17.50 200 7.80 29.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Histioteuthis bonnellii corpuscula 12.40 5.90 20 4.10 6.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Histioteuthis cf. celetaria 12.40 9.10 200 7.50 13.30 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Histioteuthis hoylei 15.75 11.60 129 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Histioteuthis meleagroteuthis 12.40 8.10 59 5.20 9.60 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Histioteuthis miranda 15.75 16.67 6 13.00 22.00 Best 1999 Donkergat, South Africa 1962-1963 P. macrocephalus Histioteuthis miranda 10.00 17.16 19 14.00 21.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Histioteuthis cf. miranda 12.40 15.00 200 11.20 18.90 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Histioteuthis reversa 12.40 5.20 19 4.40 6.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 P. macrocephalus Histioteuthis sp. 10.00 4.67 6 3.00 8.00 Best 1999 Donkergat, South Africa 1962-1963 P. macrocephalus Histioteuthis sp. 15.75 5.50 6 5.00 6.00 Best 1999 Donkergat, South Africa 1962-1963 P. macrocephalus Lepidoteuthis grimaldii 12.40 57.90 200 30.60 102.40 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Pholidoteuthis boschmai 15.75 26.91 11 16.00 39.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Pholidoteuthis boschmai 15.75 29.85 47 14.00 63.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Pholidoteuthis boschmai 10.00 28.84 57 20.00 36.00 Best 1999 Donkergat, South Africa 1962-1963 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

234

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

P. macrocephalus Pholidoteuthis boschmai 12.40 41.10 200 19.60 38.40 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Pholidoteuthis cf. boschmai 15.75 28.00 6 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Idioteuthis famelica 15.75 14.20 21 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Mastigoteuthis sp. A 12.40 15.50 83 12.60 20.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Mastigoteuthis sp. B 12.40 18.60 2 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Octopoteuthis cf. neilseni 15.75 2.50 3 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Octopoteuthis rugosa 15.75 15.75 4 15.00 17.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Octopoteuthis rugosa 10.00 21.36 45 9.00 54.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Octopoteuthis rugosa 15.75 2.70 13 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Octopoteuthis rugosa 12.40 20.00 200 13.30 24.70 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Octopoteuthis sp. G 12.40 32.50 115 23.00 41.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Taningia danae 15.75 42.30 13 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Taningia danae 12.40 75.80 200 18.00 158.70 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Ommastrephes bartramii 15.75 46.60 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Ommastrephes bartramii 12.40 48.60 6 15.50 75.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Todarodes sagittatus 12.40 48.40 200 25.80 69.60 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Kondakovia longimana 16.50 53.50 81 26.00 80.00 Nemoto et al. 1985, 1987 Southern Ocean 1972-1973 

P. macrocephalus Moroteuthis robsoni 15.75 30.90 10 19.00 42.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Moroteuthis robsoni 10.00 35.87 23 17.00 56.00 Best 1999 Donkergat, South Africa 1962-1963 

P. macrocephalus Onychoteuthis borealijaponica 12.40 31.70 110 16.30 44.00 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. macrocephalus Onychoteuthis sp. 15.75 28.00 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

P. macrocephalus Vampyroteuthis infernalis 12.40 10.40 5 9.60 12.90 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

P. blainvillei Semirossia tenera 1.50 3.80 2 3.20 4.50 dos Santos and Haimovici 2001 Brazil 1985-1998 P. blainvillei Loligo plei 1.50 15.40 27 6.80 21.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

P. blainvillei Loligo plei 1.50 16.60 155 6.60 26.60 dos Santos and Haimovici 2001 Brazil 1985-1998 

P. blainvillei Loligo sanpaulensis 1.50 5.10 593 2.00 21.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

P. blainvillei Loligo sanpaulensis 1.50 10.30 2686 2.20 22.00 dos Santos and Haimovici 2001 Brazil 1985-1998 

P. blainvillei Lolliguncula brevis 1.50 4.90 134 2.40 8.40 dos Santos and Haimovici 2001 Brazil 1985-1998 P. blainvillei Argonauta nodosa 1.50 2.40 55 0.50 4.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

P. blainvillei Eledone gaucha 1.50 2.10 1 dos Santos and Haimovici 2001 Brazil 1985-1998 P. blainvillei Octopus tehuelchus 1.50 2.70 3 2.40 2.90 dos Santos and Haimovici 2001 Brazil 1985-1998 P. crassidens Loligo vulgaris reynaudii 3.73 21.10 4.70 3 16.00 25.30 Sekiguchi et al. 1992 Benguela, South Africa 1971-1982 P. crassidens Ommastrephes bartramii 5.50 28.20 5 19.10 32.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

S. fluviatilis Loligo plei 1.75 15.20 137 4.10 26.60 dos Santos and Haimovici 2001 Brazil 1985-1998 

S. fluviatilis Loligo sanpaulensis 1.75 4.50 260 1.40 19.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

S. fluviatilis Lolliguncula brevis 1.75 4.10 199 2.50 6.00 dos Santos and Haimovici 2001 Brazil 1985-1998 

S. fluviatilis Lolliguncula brevis 1.78 4.01 0.79 172 2.84 5.84 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Anchoa sp. 1.78 7.34 1.31 17 5.52 10.60 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Eucinostomus argenteus 1.78 11.91 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Orthopristis ruber 1.78 3.73 0.73 2 3.21 4.24 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Pomadasys corvinaeformis 1.78 9.06 1.98 7 4.92 10.80 de Oliveira Santos et al. 2002 Brazil 1995-1998 S. fluviatilis Cynoscion jamaicensis 1.78 9.35 5.16 2 5.70 13.00 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Cynoscion leiarchus 1.78 24.80 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 S. fluviatilis Isopisthus parvipinnis 1.78 13.98 2.73 56 10.24 19.81 de Oliveira Santos et al. 2002 Brazil 1995-1998 S. fluviatilis Larimus breviceps 1.78 10.51 1.60 6 8.71 12.98 de Oliveira Santos et al. 2002 Brazil 1995-1998 S. fluviatilis Macrodon ancylodon 1.78 28.10 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 S. fluviatilis Micropogonias furnieri 1.78 16.23 7.16 11 5.68 33.08 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Paralonchurus brasiliensis 1.78 14.93 3.96 57 3.93 20.81 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Stellifer brasiliensis 1.78 10.61 1.82 52 6.09 15.54 de Oliveira Santos et al. 2002 Brazil 1995-1998 S. fluviatilis Stellifer rastrifer 1.78 6.19 0.99 657 3.48 10.84 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Mugil spp. 1.78 14.76 0.35 2 14.51 15.00 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Trichiurus lepturus 1.78 18.29 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 

S. fluviatilis Paralichthys orbignyanus 1.78 9.31 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 S. attenuata Loligo vulgaris reynaudii 2.03 7.10 2.90 7 4.20 11.50 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 S. attenuata Ancistrocheirus lesueurii 2.05 4.69 2.96 139 0.88 20.31 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 2. Raw food habits data, sorted phylogenetically by predator species. See Appendix 1 for common and full scientific names of predator species. 

235

Predator Mean prey range range 

Predator species Prey species length (m) length (cm) SD N low high Reference Locality Dates 

S. attenuata Architeuthis sp. 2.05 6.29 2.58 2 4.47 8.12 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Chtenopteryx sicula 2.05 3.47 0.63 39 2.17 5.16 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Leachia dislocata 2.05 12.79 3.34 628 4.96 24.81 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Liocranchia reinhardti 2.05 12.54 3.44 238 4.22 19.96 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Megalocranchia sp. 2.05 10.91 5.42 479 0.24 25.39 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Abraliopsis affinis 2.05 3.00 0.49 3671 1.87 4.54 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Enoploteuthis chunii 2.05 7.12 877 6.10 8.20 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Pholidoteuthis boschmai 2.05 9.73 2.55 150 4.29 18.19 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Mastigoteuthis dentata 2.05 6.91 1.42 870 2.06 13.24 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Octopoteuthis deletron 2.05 4.56 2.35 63 1.03 13.02 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Dosidicus gigas 2.05 52.27 596 46.71 59.95 Perrin et al. 1973 E Tropical Pacific 1968 

S. attenuata Dosidicus gigas 2.05 18.08 4.31 207 9.92 31.94 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Eucleoteuthis luminosa 2.05 11.27 3.39 475 5.16 21.94 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Eucleoteuthis luminosa 2.05 21.03 23 19.14 24.55 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Hyaloteuthis pelagica 2.05 7.53 1.27 114 5.06 11.79 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Nototodarus hawaiiensis 2.05 7.08 0.99 5 6.5 8.79 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Ommastrephes bartramii 2.05 12.92 3.53 1158 6.97 27.35 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Sthenoteuthis oualaniensis 2.05 14.42 4.21 287 7.34 32.09 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Sthenoteuthis oualaniensis 2.05 12.36 23 9.31 14.31 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Onychoteuthis banksii 2.05 7.43 2 646 0.86 13.99 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Onychoteuthis banksii 2.05 11.37 47 6.01 19.17 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Pterygioteuthis giardi 2.05 2.18 0.34 65 1.64 3.17 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Engraulis japonicus 2.05 10.77 115 9.30 12.84 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Ceratoscopelus warmingii 2.05 4.93 0.48 114 3.38 6.15 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Diaphus schmidti 2.05 4.20 396 2.65 5.30 Wang et al. 2003 Taiwan 1994-1995 S. attenuata Diaphus watasei 2.05 12.64 167 8.62 16.69 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Lampanyctodes hectoris 2.05 5.66 88 4.43 6.71 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Myctophum asperum 2.05 7.60 675 6.63 8.42 Wang et al. 2003 Taiwan 1994-1995 S. attenuata Myctophum nitidulum 2.05 5.64 0.52 14 4.67 6.31 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Symbolophorus spp. 2.05 6.66 0.7 273 4.09 8.24 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 S. attenuata Oxyporhamphus micropterus 2.05 14.34 1.51 40 12.22 18.01 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Exocoetus moncirrhus 2.05 16.47 1.68 19 13.69 19.37 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Exocoetus volitans 2.05 15.87 2.22 68 9.09 18.74 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Decapterus macrosoma 2.05 20.06 55 16.11 22.20 Wang et al. 2003 Taiwan 1994-1995 S. attenuata Trichiurus lepturus 2.05 55.50 74 49.62 65.65 Wang et al. 2003 Taiwan 1994-1995 S. attenuata Scomber australasicus 2.05 2.76 29 16.39 33.27 Wang et al. 2003 Taiwan 1994-1995 

S. attenuata Scomber japonicus 2.05 30.77 15 28.00 33.50 Wang et al. 2003 Taiwan 1994-1995 S. attenuata Auxis thazard 2.05 21.15 2.29 8 19 26 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Cubiceps baxteri 2.05 14.09 0.49 3 13.61 14.58 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. attenuata Cubiceps pauciradiatus 2.05 10.91 0.98 233 7.46 12.98 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

S. coeruleoalba Loligo vulgaris reynaudii 2.03 9.30 4.00 183 2.40 22.30 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 

§ = median prey size; n.a.** = sample size not available, but probably large. 
236

Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum

Prey Species size

Crustaceans

Mysidae

Antarctomysis maxima 4.30 

Boreomysis sp. 5.00

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

34 29 3.99 

Brierly et al. 1998 1 0 n.a. no

Finley and Gibb 1982 1 0 n.a. yes

Gammaridae 

Gammarus setosus 2.00 Weslawski et al. 1994 1 1 1.53 76.50 no 

Hyperiidae 

Themisto libellula 3.40 Weslawski et al. 1994 1 1 1.66 48.82 no 

Euphausiidae 

Euphausia superba 6.60 Reid et al. 1996 15 15 4.53 68.64 yes

Thysanoessa inermis 3.20 Nicol and Endo 1997 1 1 1.75 54.69 yes

Oplophoridae 

Acanthephyra carinata 5.00Acanthephyra quadrispinsoa 1.90Notostomus elegans 4.50

Dolar et al. 2003 1 0 n.a. no 

Dolar et al. 2003 2 2 1.45 76.32 no Dolar et al. 2003 1 0 n.a. no 

Majidae 

Chionoectes opilio 15.00 Kaimmer et al. 1976 1 1 5.70 38.00 yes 

Hippolytidae 

Chorismus antarcticus 2.00 Gorny et al. 1993 1 1 1.33 66.50 no 

Aristeidae 

Aristeus sp. = Hemipenaeus sp. 1.60 Dolar et al. 2003 1 0 n.a. no 

Penaeidae 

Hemipenaeus sp. (see Aristeus sp.) no 

Palinuridae 

Jasus lalandii 11.50 Schoeman et al. 2002 3 3 3.96 34.43 yes 

Crangonidae 

Notocrangon antarcticus 2.50 Bluhm and Brey 2001 1 1 1.41 56.40 no

Sabinea septemcarinata 5.60 Weslawski et al. 1994 1 1 5.02 89.64 no

Portunidae 

Ovalipes catharus 11.00 Lalas 1997 1 1 7.00 63.64 no 

Pandalidae 

Pandalus borealis 9.20 Weslawski et al. 1994 1 1 7.47 81.20 yes 

Cephalopods 

Sepiidae

Sepia apama 50.00

Sepia novaehollandiae 20.10Sepia officinalis 45.00Sepia sp. 50.00

487 275 18.94

Wood and Day 1998 1 1 30.10 60.20 minor 

Gales et al. 1993 1 1 15.50 77.11 no 

Wood and Day 1998 1 0 n.a. minor 

Wood and Day 1998 1 1 14.29 28.58 minor 

Sepiolidae 

Rossia macrosoma 8.00Semirossia tenera 5.00Sepietta oweniana 7.00

Wood and Day 1998 1 0 n.a. minor 

Wood and Day 1998 3 1 1.70 34.00 no 

Wood and Day 1998 1 0 n.a. no 

Loliginidae 

Alloteuthis subulata (see Loligo subulata)

Loligo bleekeri 40.00Loligo gahi 28.00Loligo japonica (see Loliolus japonica)Loligo opalescens 23.50Loligo pealeii 50.00Loligo plei 35.00

Loligo sanpaulensis 22.00Loligo subulata = Alloteuthis subulata 20.00

Loligo vulgaris reynaudii 40.00Loligo vulgaris vulgaris 64.00

n.a. 

Wood and Day 1998 1 0 n.a. yes 

Wood and Day 1998 4 4 9.40 33.57 yes 

yes 

Lowry and Caretta 1999 15 10 11.58 49.27 yes 

Wood and Day 1998 4 3 16.36 32.71 yes 

Wood and Day 1998 10 6 16.32 46.62 yes 

dos Santos and Haimovici 2001 9 8 8.69 39.49 yes 

Wood and Day 1998 1 1 7.10 35.50 yes 

Wood and Day 1998 13 11 14.04 35.10 yes Wood and Day 1998 1 0 n.a. yes 
Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum 

Prey Species size 

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

237

Loliolus japonica = Loligo japonica

Lolliguncula brevisSepioteuthis australis

Ancistrocheiridae 

Ancistrocheirus alessandrini (see A. lesueurii) 

Ancistrocheirus lesueurii = A. alessandrini 

Architeuthidae 

Architeuthis cf. dux

Architeuthis sp.

Bathyteuthidae 

Bathyteuthis abyssicola 

Batoteuthidae 

Batoteuthis skolops 

Brachioteuthidae 

Brachioteuthis cf. pictaBrachioteuthis riiseiBrachioteuthis cf. riisei

Brachioteuthis sp.Brachioteuthis spp.

Chiroteuthidae 

Chiroteuthis imperator Chiroteuthis picteti Chiroteuthis veranyi Chiroteuthis cf. veranyi 

Chiroteuthis sp. Planctoteuthis sp. = Valbyteuthis sp. 

Valbyteuthis sp. (see Planctoteuthis sp.) 

Chtenopterygidae 

Chtenopteryx siculaChtenopteryx sp.

Cranchiidae 

Galiteuthis armataGaliteuthis glacialisHelicocranchia sp. ?Leachia dislocataLiguriella sp.

Liocranchia reinhardtiLiocranchia sp.Megalocranchia fisheriMegalocranchia fisheri A

Megalocranchia sp. A

Megalocranchia sp. BMegalocranchia sp. GMegalocranchia sp.Mesonychoteuthis hamiltoniTaonius borealisTaonius megalopsTaonius pavoTaonius sp.Teuthowenia maculataTeuthowenia megalopsTeuthowenia pellucidaTeuthowenia sp.

Cycloteuthidae 

Cycloteuthis akimushkiniCycloteuthis sp.

12.00 Wood and Day 1998 1 0 n.a. no 12.00 Wood and Day 1998 3 3 4.34 36.14 no 53.10 Gales et al. 1993 1 1 32.80 61.77 yes 

n.a. 

40.00 Best 1999 8 5 17.12 42.80 potential 

600.00 Wood and Day 1998 1 1 115.90 no

600.00 Wood and Day 1998 4 0 n.a. 19.32 no

7.33 Sekiguchi et al. 1993 3 0 n.a. no based on a report of mean size 

27.00 Wood and Day 1998 2 0 n.a. no 

9.00 Wood and Day 1998 4 3 7.49 83.20 indet. 

43.40 Clarke and Goodall 1994 2 1 43.40 100.00 indet. based on a report of mean size 

43.40 Clarke and Goodall 1994 1 1 5.84 13.46 indet. based on a report of mean size 43.40 Clarke and Goodall 1994 1 0 n.a. indet. based on a report of mean size 43.40 Clarke and Goodall 1994 1 0 n.a. indet. based on a report of mean size 

25.00 Wood and Day 1998 1 1 6.14 24.56 no 25.00 Wood and Day 1998 1 0 n.a. no 16.10 Clarke and Goodall 1994 5 0 n.a. no based on a report of mean size 

16.10 Clarke and Goodall 1994 1 0 n.a. no based on a report of mean size 25.00 Wood and Day 1998 7 4 10.40 41.59 no 

8.00 Wood and Day 1998 1 0 n.a. no

no

9.00 Wood and Day 1998 2 1 3.47 38.56 no 

9.00 Wood and Day 1998 1 0 n.a. no 

27.30 Clarke and Goodall 1994 5 2 23.45 85.90 no 50.00 Wood and Day 1998 3 2 19.39 38.77 no 25.80 Clarke et al. 1993 1 0 n.a. no based on a report of mean size 24.81 Robertson and Chivers 1997 2 1 12.79 51.55 no 

24.00 Wood and Day 1998 1 0 n.a. no 25.80 Clarke et al. 1993 3 1 12.54 48.60 no based on a report of mean size 36.01 Sekiguchi et al. 1993 2

180.00 Young and Mangold 1996a 1

180.00 Young and Mangold 1996a 1

180.00 Young and Mangold 1996a 1180.00 Young and Mangold 1996a 1180.00 Young and Mangold 1996a 1180.00 Young and Mangold 1996a 3250.00 Wood and Day 1998 5

1 25.40 70.54 no based on a report of mean size 

1 96.00 53.33 no 

1 20.80 11.56 no 

1 50.50 28.06 no 1 23.20 12.89 no 1 81.70 45.39 no 2 15.28 8.49 no 

2 82.99 33.20 potential 

66.00 Young and Mangold 1996b 2 2 28.65 43.40 no 66.00 Young and Mangold 1996b 1 0 n.a. no maximum for genus 49.02 Wang et al. 2002 10 8 32.34 65.98 no 66.00 Young and Mangold 1996b 1 1 27.34 41.42 no 37.10 Clarke et al. 1993 1 1 29.90 80.59 no 40.00 Wood and Day 1998 4 3 25.33 63.32 no 22.22 Sekiguchi et al. 1996 4 1 18.12 81.55 no based on a report of mean size 

40.00 Wood and Day 1998 3 0 n.a. no 

50.50 Clarke and Young 1998 2 2 37.95 75.15 no 

50.50 Clarke and Young 1998 2 0 n.a. no 
Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum 

Prey Species size 

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

238

Discoteuthis discusDiscoteuthis laciniosaDiscoteuthis cf. laciniosa

Discoteuthis sp.

Enoploteuthidae 

Abralia redfieldiAbralia sp.

Abraliopsis affinisAbraliopsis cf. lineata

Abraliopsis sp. AAbraliopsis sp. BAbraliopsis sp. CAbraliopsis sp. DEnoploteuthis cf. anaspis

Enoploteuthis chuniiEnoploteuthis sp. AEnoploteuthis sp.Watasenia scintillans

Enoploteuthidae 

Gonatidae 

Berryteuthis anonychus Berryteuthis magister Gonatopsis borealis/Berryteuthis magister Gonatopsis borealis Gonatopsis makko Gonatopsis octopedatus Gonatus antarcticus Gonatus berryi Gonatus fabricii Gonatus cf. fabricii Gonatus madokai Gonatus madokai/G. middendorffi Gonatus onyx Gonatus steenstrupi Gonatus sp. Gonatidae 

Histioteuthidae 

Histioteuthis arcturiHistioteuthis bonnellii bonnelliiHistioteuthis bonnellii corpusculaHistioteuthis cf. celetaria

Histioteuthis eltaninaeHistioteuthis hoyleiHistioteuthis macrohista

Histioteuthis meleagroteuthisHistioteuthis mirandaHistioteuthis cf. miranda

Histioteuthis reversaHistioteuthis sp. AHistioteuthis sp. B

Histioteuthis sp.

Histioteuthis spp.

Lepidoteuthidae 

Lepidoteuthis grimaldii 

10.33 Sekiguchi et al. 1993 2 1 10.33 100.00 no based on a report of mean size 16.10 Clarke and Young 1998 3 1 13.20 81.99 no based on a report of mean size 

16.10 Clarke and Young 1998 1 1 7.50 46.58 no based on a report of mean size 

19.31 Wang et al. 2002 2 0 n.a. no 

7.00 Wood and Day 1998 1 1 2.90 41.43 no 

7.00 Wood and Day 1998 1 0 n.a. no 4.54 Robertson and Chivers 1997 1 1 3.00 66.08 no 

6.70 Dolar et al. 2003 2 2 4.46 66.49 no 7.00 Wood and Day 1998 1 1 4.40 62.86 no 7.00 Wood and Day 1998 1 1 3.20 45.71 no 7.00 Wood and Day 1998 1 1 3.40 48.57 no 7.00 Wood and Day 1998 1 1 5.30 75.71 no 7.00 Tsuchiya 2000 1 1 7.70 110.00 no 

10.75 Mori et al. 2001 4 3 7.14 66.42 no 18.00 Clarke and Goodall 1994 1 1 9.50 52.78 no 18.00 Clarke and Goodall 1994 1 0 n.a. no 

7.00 Wood and Day 1998 4 4 4.07 58.14 yes 

n.a. 1 0 n.a. no 

31.99 Mori et al. 2001 1 0 n.a. indet. 33.43 Walker et al. 2002 8 8 11.79 35.27 yes 33.43 Walker et al. 2002 2 2 9.47 28.33 yes 31.02 Walker et al. 2002 5 4 16.27 52.44 potential 28.91 Ohizumi et al. 2000 1 1 9.96 34.46 no 24.00 Wood and Day 1998 1 1 9.11 37.94 no 35.00 Wood and Day 1998 9 5 19.96 57.02 no 28.50 Clarke and Young 1998 3 1 13.99 49.09 no 40.25 Finley and Gibb 1982 2 2 21.90 54.41 potential 

40.25 Finley and Gibb 1982 2 2 22.16 55.06 potential 33.00 Wood and Day 1998 1 1 10.12 30.66 yes 33.00 Wood and Day 1998 1 1 6.67 20.21 yes 18.00 Wood and Day 1998 2 1 8.59 47.70 no 18.20 Clarke et al. 1993 1 1 14.90 81.87 potential 35.00 Wood and Day 1998 1 1 8.24 23.55 yes 

n.a. 4 0 n.a. yes 

20.00 Wood and Day 1998 1 1 11.10 55.50 no 33.00 Wood and Day 1998 1 1 17.50 53.03 potential 33.00 Wood and Day 1998 2 1 5.90 17.88 potential 

26.00 Wood and Day 1998 1 1 9.10 35.00 no 10.00 Wood and Day 1998 4 4 5.36 53.63 no 20.00 Wood and Day 1998 2 1 11.60 58.00 no 

8.41 Kirkman et al. 2000 3 1 6.30 74.91 no 

11.40 Young and Veccione 2000 4 2 6.80 59.61 no size at maturity for females 27.00 Wood and Day 1998 5 3 11.58 42.87 potential 

27.00 Wood and Day 1998 1 1 15.00 55.56 no 19.00 Wood and Day 1998 2 1 5.20 27.37 no 33.00 Wood and Day 1998 1 1 3.10 9.39 potential 

33.00 Wood and Day 1998 1 1 4.70 14.24 potential 

33.00 Wood and Day 1998 6 3 5.61 17.00 potential 

33.00 Wood and Day 1998 3 2 6.55 19.85 potential 

102.40 Clarke et al. 1993 1 1 57.90 56.54 no 
239

Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum

Prey Species size

Pholidoteuthidae

Pholidoteuthis boschmai 63.00 

Pholidoteuthis cf. boschmai 63.00 

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

Best 1999 8 5 27.29 43.32 potential

Best 1999 1 0 n.a. potential

Lycoteuthidae 

Lycoteuthis diadema (See Lycoteuthis lorigera) no

Lycoteuthis lorigera 18.00 Wood and Day 1998 8 2 9.95 55.28 no

Mastigoteuthidae 

Idioteuthis famelica = Mastigoteuthis famelica 14.20

Idioteuthis hjorti = Mastigoteuthis hjorti 14.62

Mastigoteuthis dentata 14.00Mastigoteuthis famelica (see Idioteuthis famelica)Mastigoteuthis hjorti (see Idioteuthis hjorti)

Mastigoteuthis psychrophila 14.30Mastigoteuthis sp. A 47.00

Mastigoteuthis sp. B 47.00

Mastigoteuthis sp. 47.00

Clarke and Young 1998 1 1 14.20 100.00 no based on a report of mean size 

Wang et al. 2002 1 1 11.00 75.24 no 

Wood and Day 1998 1 1 6.91 49.36 no 

no no 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 3 1 15.50 32.98 no 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 4 3 10.24 21.79 no 

Neoteuthidae 

Alluroteuthis antarctica 40.13 Sekiguchi et al. 1993 7 6 20.86 51.99 no based on a report of mean size 

Octopoteuthidae 

Octopoteuthis deletron 20.00Octopoteuthis cf. deletron 20.00

Octopoteuthis cf. neilseni 20.00

Octopoteuthis rugosa 54.00Octopoteuthis sp. G 54.00

Octopoteuthis sp. 54.00

Octopoteuthis spp. 54.00

Taningia danae 158.70

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. no 

Best 1999 6 4 14.77 27.35 no 

Best 1999 1 1 32.50 60.19 no 

Best 1999 8 0 n.a. no 

Best 1999 2 0 n.a. no 

Clarke et al. 1993 4 3 45.67 28.78 potential 

Ommastrephidae 

Dosidicus gigas 150.00Eucleoteuthis luminosa 24.55Hyaloteuthis pelagica 11.79Illex argentinus 35.90Illex illecebrosus 31.00Martialia hyadesi 42.60Nototodarus gouldi 41.50Nototodarus hawaiiensis 16.00Nototodarus sloanii 43.00Ommastrephes bartramii 75.10Ornithoteuthis antillarum 20.00Ornithoteuthis volatilis 31.00Sthenoteuthis oualaniensis 35.00

Todarodes angolensis 35.00Todarodes pacificus 50.00Todarodes sagittatus 75.00Todaropsis eblanae 27.00Ommastrephidae n.a.

Wood and Day 1998 3 3 39.61 26.41 yes 

Wang et al. 2002 5 3 12.46 50.75 no 

Robertson and Chivers 1997 1 1 7.53 63.87 no 

dos Santos and Haimovici 2001 11 6 15.82 44.06 yes 

Wood and Day 1998 3 3 18.40 59.35 yes 

Rodhouse et al. 1992 5 4 25.47 59.78 potential 

Gales et al. 1993 1 1 23.90 57.59 yes 

Wood and Day 1998 1 0 n.a. potential 

Fea et al. 1999 1 0 n.a. potential 

Clarke et al. 1993 11 6 23.65 31.49 yes 

Wood and Day 1998 4 1 5.70 28.50 potential 

Wood and Day 1998 2 1 20.12 64.90 potential Wood and Day 1998 4 3 16.71 47.74 no 

Wood and Day 1998 1 1 14.60 41.71 no 

Wood and Day 1998 2 1 7.37 14.74 yes 

Wood and Day 1998 4 3 28.47 37.96 yes 

Wood and Day 1998 2 1 8.30 30.74 yes 

1 0 n.a. n.a. 

Onychoteuthidae 

Kondakovia longimana 80.00Moroteuthis ingens 94.00Moroteuthis knipovitchi 35.00Moroteuthis loennbergi 30.00Moroteuthis robsoni 75.00Moroteuthis sp. 94.00

Onychoteuthis banksii 30.00Onychoteuthis borealijaponica 44.00Onychoteuthis sp. 1 44.00

Nemoto et al. 1985, 1987 7 4 40.05 50.06 no 

Wood and Day 1998 4 2 44.20 47.02 potential Wood and Day 1998 10 4 21.88 62.50 potential 

Wood and Day 1998 3 1 17.58 58.60 potential Wood and Day 1998 4 3 29.16 38.88 potential 

Wood and Day 1998 1 1 22.42 23.85 potential Wood and Day 1998 6 2 9.40 31.33 minor 

Clarke et al. 1993 3 2 23.67 53.80 potential 

Clarke et al. 1993 1 1 8.10 18.41 potential 
240

Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum

Prey Species size

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

Onychoteuthis sp. 2 44.00 

Onychoteuthis sp. 44.00 

Onychoteuthidae n.a. 

Clarke et al. 1993 1 1 20.30 46.14 potential 

Clarke et al. 1993 2 1 29.26 66.50 n.a. 

1 0 n.a. n.a. 

Psychroteuthidae 

Psychroteuthis glacialis 44.00 Wood and Day 1998 7 4 13.02 29.59 potential 

Pyroteuthidae 

Pterygioteuthis giardi 4.00 Wood and Day 1998 1 1 2.18 54.50 noPyroteuthis addolux 4.50 Clarke and Young 1998 1 0 n.a. no

Argonautidae 

Argonauta nodosa 13.10 dos Santos and Haimovici 2001 6 1 2.40 18.32 minor 

Octopodidae 

Bathypolypus arcticus 10.00 Eledone gaucha 6.50 Octopus australis 15.50 Octopus maorum 30.00 Octopus pallidus 15.00 Octopus superciliosus 5.02 Octopus tehuelchus 20.00 Octopus vulgaris 25.00 Octopus sp. n.a. 

Pareledone charcoti 7.00 Pareledone cf. charcoti 7.00 

Pareledone cf. turqueti 15.00 

Pareledone polymorpha 10.00 Pareledone cf. polymorpha 10.00 

Octopodidae n.a. 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. no 

Gales et al. 1993 1 1 10.10 65.16 no 

Wood and Day 1998 2 1 15.30 51.00 no Wood and Day 1998 1 0 n.a. no 

Gales et al. 1993 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. yes 

Wood and Day 1998 1 0 n.a. yes 

1 1 7.42 yes 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. no 

Wood and Day 1998 1 0 n.a. no 

1 0 n.a. yes 

Vampyroteuthidae 

Vampyroteuthis infernalis 13.00 Wood and Day 1998 2 0 n.a. no 

Bony and cartilaginous fishes 645 421 18.46

Myxinidae

Myxine glutinosa 80.00 Froese and Pauly 2003 1 0 n.a. no 

Squalidae 

Squalus acanthias 160.00 Froese and Pauly 2003 1 0 n.a. yes 

Rajidae 

Raja sp. n.a. 1 1 68.00 minor

Raja spp. 93.00 Lalas 1997 1 0 n.a. minor

Anguillidae 

Anguilla anguilla 133.00 Froese and Pauly 2003 1 0 n.a. yes 

Clupeidae 

Alosa pseudoharengus 40.00 Clupea harengus 45.00 Clupea harengus pallasi 46.00 Etrumeus whiteheadi 23.60 Sardinops ocellatus see S. sagax Sardinops sagax = S. ocellatus 39.50 Sprattus sprattus 16.00 

Froese and Pauly 2003 2 1 22.60 56.50 yes Froese and Pauly 2003 11 10 27.49 61.09 yes Froese and Pauly 2003 4 2 16.48 35.83 yes 

Sekiguchi et al. 1992 4 3 14.70 62.29 yes 

yes 

Froese and Pauly 2003 14 11 16.72 42.33 yes Froese and Pauly 2003 1 1 14.00 87.50 yes 

Engraulidae 

Anchoa sp. 50.00

Engraulis encrasicolus = capensis 20.00

Engraulis japonicus 17.16Engraulis mordax 24.80

Froese and Pauly 2003 1 1 7.34 14.68 yes 

Froese and Pauly 2003 10 5 10.46 52.30 yes 

Ohizumi et al. 2000 2 2 11.75 68.46 yes 

Froese and Pauly 2003 5 3 10.80 43.55 yes 

Microstomatidae 

Nansenia candida 22.20 Froese and Pauly 2003 1 0 n.a. no 

Bathylagidae 

Bathylagidae 25.00 Froese and Pauly 2003 3 2 10.60 42.40 no basin-wide maximum for family 

Osmeridae 
241

Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum

Prey Species size

Mallotus villosus 20.00

Osmerus eperlanus 45.00Osmerus mordax 35.60Thaleichthys pacificus 30.00

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

Froese and Pauly 2003 8 5 14.64 73.20 yes Froese and Pauly 2003 1 1 10.20 22.67 yes Froese and Pauly 2003 2 0 n.a. yes 

Froese and Pauly 2003 4 1 15.80 52.67 yes 

Salmonidae 

Oncorhynchus gorbuscha 76.00Oncorhynchus kisutch 108.00Oncorhynchus tshawytscha 150.00Salmo salar 150.00Salmonidae 150.00

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 2 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. yes basin-wide maximum for genus 

Gonostomatidae 

Gonostoma cf. elongatum 27.50 Froese and Pauly 2003 1 0 n.a. no 

Sternoptychidae 

Argyropelecus affinis 8.40Maurolicus muelleri 8.00Maurolicus cf. muelleri 8.00

Maurolicus weitzmani 5.50

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 4 3 3.56 44.45 minor 

Froese and Pauly 2003 1 0 n.a. minor 

Gannon et al. 1998 1 1 5.00 90.91 no 

Phosichthyidae 

Ichthyococcus cf. irregularis 7.60 Froese and Pauly 2003 1 0 n.a. no 

Stomiidae 

Chauliodus cf. barbatus 35.00 Froese and Pauly 2003 1 1 18.00 51.43 no basin-wide maximum for genus 

Scopelarchidae 

Rosenblattichthys cf. volucris 14.00 Robison and Craddock 1983 1 0 n.a. no

Scopelarchus guentheri 16.00 Robison and Craddock 1983 1 0 n.a. no

Notosudidae 

Scopelosaurus harryi 22.00 Froese and Pauly 2003 1 1 15.34 69.74 no 

Paralepididae 

Arctozenus risso = Notolepis risso 30.00

Notolepis coatsi 38.00Notolepis risso (see Arctozenus risso )

Paralepididae n.a.

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 1 27.18 71.51 no 

no 

2 0 n.a. no 

Evermannellidae 

Evermanella cf. ahlstromi 16.90 Froese and Pauly 2003 1 0 n.a. no 

Myctophidae 

Ceratoscopelus warmingii 8.10Diaphus schmidti 5.30Diaphus watasei 17.00Electrona antarctica 13.26Electrona carlsbergi 10.10Electrona subaspera 12.75Electrona sp. 13.26

Gymnoscopelus bolini 28.00Gymnoscopelus braueri 13.20Gymnoscopelus fraseri 13.90Gymnoscopelus microlampas 11.70Gymnoscopelus nicholsi 18.90Gymnoscopelus piabilis 16.00Krefftichthys anderssoni 8.70

Lampadena luminosa 20.00Lampadena sp. 20.00

Lampanyctodes hectoris 7.00Lampanyctus cf. idostigma 9.60

Lampanyctus cf. nobilis 12.40

Lampichthys procerus 9.50Metelectrona ventralis 12.20Myctophum asperum 8.42

Froese and Pauly 2003 3 3 5.59 69.01 no 

Wang et al. 2003 1 1 4.20 79.25 no 

Froese and Pauly 2003 1 1 12.64 74.35 no 

Kirkman et al. 2000 13 12 7.83 59.05 no 

Casaux et al. 1998 7 7 7.44 73.70 minor 

Green et al. 1997 13 10 8.04 63.07 no 

Kirkman et al. 2000 1 0 n.a. no basin-wide maximum for genus 

Froese and Pauly 2003 4 3 13.60 48.57 minor Froese and Pauly 2003 6 1 10.63 80.53 minor 

Klages and Bester 1998 4 4 8.03 57.76 minor 

Froese and Pauly 2003 2 0 n.a. no 

Casaux et al. 1998 15 15 12.06 63.81 minor 

Klages and Bester 1998 5 4 9.51 59.43 no 

Casaux et al. 1998 9 3 3.81 43.79 no 

Froese and Pauly 2003 3 2 9.22 46.10 no Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 4 3 5.15 73.62 yes 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. no 

Cherel et al. 1997 4 4 6.82 55.90 no 

Wang et al. 2003 1 1 7.60 90.26 no 
242

Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum

Prey Species size

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

Myctophum nitidulum 8.30 Protomyctophum andriashevi 6.00 Protomyctophum bolini 9.70 Protomyctophum choriodon 13.70 Protomyctophum tenisoni 7.90 Symbolophorus evermanni 8.00 Symbolophorus sp. 16.00 

Symbolophorus spp. 14.00 

Mycotophidae n.a. 

Froese and Pauly 2003 3 2 5.68 68.43 no 

Froese and Pauly 2003 1 0 n.a. no 

Cherel et al. 1997 5 3 4.46 46.03 no 

Klages and Bester 1998 8 7 8.33 60.80 no 

Klages and Bester 1998 4 3 4.45 56.36 no 

Froese and Pauly 2003 2 2 6.48 81.00 no 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 1 6.66 47.57 no 

1 1 6.54 n.a. 

Muraenolepididae 

Muraenolepis microps 39.10 North 1996 1 0 n.a. no 

Coryphaenoididae 

see Macrouridae n.a. 

Macrouridae 

Albatrossia pectoralis 150.00 Coryphaenoides acrolepis 87.00 Coryphaenoides cinereus 56.00 Coryphaenoides longifilis 100.00 Nezumia bairdi 40.00 Macrouridae = Coryphaenoididae n.a. 

Froese and Pauly 2003 2 0 n.a. yes 

Froese and Pauly 2003 2 2 47.69 54.82 yes Froese and Pauly 2003 4 2 23.63 42.20 minor 

Ohizumi et al. 2003 1 1 75.70 75.70 minor 

Froese and Pauly 2003 1 0 n.a. minor 

2 1 26.10 no 

Moridae 

Antimora microlepis 83.80Auchenoceros punctatus 15.00Laemonema longipes 71.26Pseudophycis bachus 90.00Pseudopycis sp. 90.00

Walker et al. 2002 1 1 46.94 56.01 yes 

Fea et al. 1999 2 1 10.00 66.67 potential 

Walker et al. 2002 4 4 35.17 49.35 yes 

Froese and Pauly 2003 1 1 19.80 22.00 yes 

Froese and Pauly 2003 1 1 36.00 40.00 yes 

Melanonidae 

Melanonus sp. 26.60 Froese and Pauly 2003 1 0 n.a. no 

Gadidae 

Arctogadus glacialis 40.00 Boreogadus saida 40.00 Eleginus gracilis 55.00 Gadus morhua 200.00 Gadus ogac 77.00 Melanagrammus aeglefinus 100.00 Merlangius merlangus 70.00 Microgadus proximus 30.50 Micromesistius poutassou 50.00 Pollachius pollachius 130.00 Pollachius virens 130.00 Theragra chalcogramma 91.00 Trisopterus esmarkii 35.00 Trisopterus minutus 40.00 

Heide-Jorgensen et al. 1994 3 3 19.90 49.75 yes 

Froese and Pauly 2003 10 9 13.39 33.48 yes Froese and Pauly 2003 2 2 14.34 26.07 yes 

Froese and Pauly 2003 24 19 26.15 13.08 yes 

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 5 0 n.a. yes 

Froese and Pauly 2003 4 4 18.05 25.79 yes Froese and Pauly 2003 1 0 n.a. minor Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. yes Froese and Pauly 2003 9 7 25.54 19.65 yes 

Froese and Pauly 2003 36 26 24.10 26.48 yes Froese and Pauly 2003 1 1 12.40 35.43 yes Froese and Pauly 2003 1 1 14.70 36.75 yes 

Lotidae 

Enchelyopus cimbrius 41.00 Froese and Pauly 2003 2 1 23.60 57.56 minorMolva molva 200.00 Froese and Pauly 2003 4 4 31.68 15.84 yes

Phycidae 

Urophycis tenuis 133.00 Froese and Pauly 2003 2 1 22.20 16.69 yesUrophycis spp. 133.00 Froese and Pauly 2003 1 1 15.90 11.95 yes

Merlucciidae 

Macruronus magellanicus 115.00 Merluccius albidus 70.00 Merluccius bilinearis 76.00 Merluccius capensis 140.00 

Merluccius hubbsi 95.00 Merluccius merluccius 140.00 

Froese and Pauly 2003 1 1 47.50 41.30 yes 

Froese and Pauly 2003 1 1 29.60 42.29 yes Froese and Pauly 2003 6 4 21.14 27.82 yes 

Froese and Pauly 2003 1 1 28.19 20.14 yes 

Froese and Pauly 2003 4 4 21.43 22.55 yes 

Froese and Pauly 2003 1 0 n.a. yes 
Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum 

Prey Species size 

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

243

Merluccius productusMerluccius spp.

Ophidiidae 

Genypterus blacodesGenypterus capensis

Batrachoididae 

Porichthys notatus 

Gobiesocidae 

Chorisochismus dentex 

Scomberesocidae 

Cololabis sairaScomberesox saurus

Hemiramphidae 

Oxyporamphus micropterus 

Exocoetidae 

Exocoetus monocirrhusExocoetus volitans

Melamphaidae 

Melamphaes sp.Scopelogadus mizolepis bispinosus

Diretmidae 

Diretmus argenteus 

Gasterosteidae 

Pungitius pungitius 

Sebastidae 

Helicolenus dactylopterusSebastes marinusSebastes sp.

Sebastes spp.

Triglidae 

Chelidonichthys capensis 

Anoplopomatidae 

Anoplopoma fimbria 

Hexagrammidae 

Pleurogrammus azonusPleurogrammus monopterygius

Cottidae 

Leptocottus armatusMyoxocephalus scorpiusCottidae 

Cyclopteridae 

Cyclopterus lumpus 

Carangidae 

Decapterus macrosomaTrachurus capensisTrachurus declivisTrachurus symmetricusTrachurus sp.

Emmelichthyidae 

Emmelichthys nitidus 

Gerreidae 

Eucinostomus argenteus 

Haemulidae 

Orthopristis ruberPomadasys corvinaeformis

91.00 Froese and Pauly 2003

140.00 Froese and Pauly 2003

200.00 Froese and Pauly 2003180.00 Froese and Pauly 2003

7 4 25.30 27.80 yes 

12 10 20.84 14.89 yes 

1 0 n.a. yes 2 1 45.90 25.50 yes 

39.00 Torok 1994 1 1 21.80 55.90 potential 

30.00 Froese and Pauly 2003 1 0 n.a. no 

40.00 Froese and Pauly 2003 1 0 n.a. yes 50.00 Froese and Pauly 2003 1 1 24.20 48.40 minor 

18.50 Froese and Pauly 2003 1 1 14.34 77.51 no 

20.00 Froese and Pauly 2003 1 1 16.47 82.35 minor 30.00 Froese and Pauly 2003 1 1 15.87 52.90 no 

9.00 Crawford 1981 1 0 n.a. no 

11.00 Froese and Pauly 2003 1 0 n.a. no 

28.50 Robison and Craddock 1983 1 0 n.a. potential 

9.00 Froese and Pauly 2003 1 0 n.a. minor 

47.00 Froese and Pauly 2003 3

120.00 Weslawski et al. 1994 1

100.00 Froese and Pauly 2003 4

111.00 Froese and Pauly 2003 4

2 21.60 45.96 yes 

1 45.00 37.50 yes 

3 20.30 20.30 yes maximum for genus in W. Atlantic 

3 9.98 9.00 yes maximum for genus in NE Pacific 

75.00 Froese and Pauly 2003 1 1 15.00 20.00 yes 

120.00 Froese and Pauly 2003 1 0 n.a. yes 

60.00 Froese and Pauly 2003 1 1 21.60 36.00 yes 50.00 Froese and Pauly 2003 3 1 32.30 yes 

46.00 Froese and Pauly 2003 2 2 12.52 27.21 minor 90.00 Froese and Pauly 2003 3 1 11.14 12.38 no 

n.a. 2 1 10.00 no 

60.00 Froese and Pauly 2003 2 2 20.20 33.67 yes 

35.00 Froese and Pauly 2003 1 1 20.06 57.31 yes 60.00 Froese and Pauly 2003 9 7 16.04 26.73 yes 64.00 Froese and Pauly 2003 2 2 26.10 40.78 yes 81.00 Froese and Pauly 2003 1 1 13.10 16.17 yes 90.00 Fea et al. 1999 4 2 34.00 37.78 yes 

50.00 Froese and Pauly 2003 2 2 16.50 33.00 minor 

20.00 Froese and Pauly 2003 1 0 n.a. minor 

40.00 Froese and Pauly 2003 1 0 n.a. yes 25.00 Froese and Pauly 2003 1 0 n.a. minor 
244

Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum

Prey Species size

Sparidae 

Argyrozona argyrozona 90.00

Cheimerius nufar 75.00Pagellus natalensis 30.00Pagrus africanus 75.00Pterogymnus laniarius 45.00

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

Froese and Pauly 2003 2 0 n.a. minor Froese and Pauly 2003 1 0 n.a. minor Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 2 2 24.18 53.73 yes 

Sciaenidae 

Argyrosomus hololepidotus 200.00Cynoscion jamaicensis 50.00Cynoscion leiarchus 90.80Genyonemus lineatus 29.00Isopisthus parvipinnis 25.00Larimus breviceps 31.00Macrodon ancylodon 45.00Micropogonias furnieri 60.00Paralonchurus brasiliensis 30.00Stellifer brasiliensis 15.54Stellifer rastrifer 20.00

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 1 22.20 76.55 yes Froese and Pauly 2003 1 1 13.98 55.92 minor Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. minor Froese and Pauly 2003 1 1 16.23 27.05 yes Froese and Pauly 2003 1 1 14.93 49.77 minor 

de Oliveira Santos et al. 2002 1 1 10.61 68.28 minor 

Froese and Pauly 2003 1 1 6.19 30.95 minor 

Mugilidae 

Mugil cephalus 120.00 Froese and Pauly 2003 1 1 13.80 11.50 yes Mugil spp. 120.00 Froese and Pauly 2003 1 0 n.a. yes basin-wide maximum for genus 

Embiotocidae 

Cymatogaster aggregata 20.30 Froese and Pauly 2003 1 1 8.50 41.87 minor 

Labridae 

Notolabrus sp. 50.00 Froese and Pauly 2003 1 1 23.00 46.00 no basin-wide maximum for genus Tautogolabrus adspersus 38.00 Froese and Pauly 2003 1 0 n.a. yes 

Odacidae 

Odax pullus 49.00 Lalas 1997 1 1 31.00 63.27 minor 

Zoarcidae 

Lycodes sp. 75.00Lycodopsis pacifica 46.00Zoarces americanus 110.00

Froese and Pauly 2003 1 0 n.a. no basin-wide maximum for genus Froese and Pauly 2003 2 2 9.97 21.66 yes 

Froese and Pauly 2003 1 1 53.30 48.45 yes 

Anarhichadidae 

Anarhichas lupus 150.00 Froese and Pauly 2003 2 0 n.a. yes

Anarrhichthys ocellatus 240.00 Froese and Pauly 2003 1 0 n.a. minor

Nototheniidae 

Dissostichus eleginoides 215.00Gobionotothen acuta 35.00Gobionotothen gibberifrons 59.00Gobionotothen marionensis 20.00Lepidonotothen larseni 28.00Lepidonotothen squamifrons = Notothenia squamifrons 55.00Notothenia squamifrons (see Lepidonotothen squamifrons )

Paranotothenia magellanica 38.00Pleuragramma antarcticum 25.91Trematomus eulepidotus 34.00Trematomus lepidorhinus 31.00Trematomus nicolai 36.00Trematomus spp. 41.00

Froese and Pauly 2003 2 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. no 

Reid 1995 4 3 38.97 66.05 minor 

Froese and Pauly 2003 1 0 n.a. no 

Reid 1995 6 6 16.76 59.86 no 

Froese and Pauly 2003 2 1 11.40 20.73 minor 

minor 

Froese and Pauly 2003 2 0 n.a. yes 

Burns et al. 1998 14 11 14.40 55.58 minor 

Froese and Pauly 2003 1 0 n.a. yes 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. yes basin-wide maximum for genus 

Artedidraconidae 

Artedidraco loennbergi 11.00Dolloidraco longedorsalis 13.70Histiodraco velifer 19.20

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. no 
245

Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type

Family Maximum

Prey Species size

Bathydraconidae

Gerlachea australis 24.00

Gymnodraco acuticeps 34.00Racovitzia glacialis 24.00

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. no 

Channichthyidae 

Chaenocephalus aceratus 72.00Champsocephalus gunnari (Heard Island population) 45.00

Champsocephalus gunnari (South Georgia population) 66.00

Channichthys rhinoceratus 60.00Chionodraco myersi 38.00Chionodraco rastrospinosus 52.00Cryodraco antarcticus 57.00Pagetopsis maculatus 25.00Pseudochaenichthys georgianus 60.00

Froese and Pauly 2003 1 0 n.a. minor 

Gon and Heemstra 1990 6 4 20.90 46.44 minor 

Froese and Pauly 2003 7 6 27.48 41.64 minor Froese and Pauly 2003 2 1 32.52 54.20 minor Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 0 n.a. minor 

Froese and Pauly 2003 2 0 n.a. minor 

Froese and Pauly 2003 1 0 n.a. no 

Froese and Pauly 2003 1 1 28.80 48.00 minor 

Ammodytidae 

Ammodytes dubius 25.00Ammodytes hexapterus 27.00Ammodytes personatus 22.21Ammodytes sp. 45.00Ammodytes spp. 45.00Gymnammodytes capensis = Ammodytes capensis 17.00

Ammodytidae n.a.

Froese and Pauly 2003 1 1 19.10 76.40 potential Froese and Pauly 2003 3 2 10.20 37.78 yes Froese and Pauly 2003 1 1 19.44 87.53 yes 

Hammond et al. 1994b 2 1 18.10 40.22 yes Hammond et al. 1994b 8 8 15.86 35.24 yes 

David 1987 2 2 25.00 147.06 yes 

2 2 14.74 yes 

Clinidae 

Clinus sp. 30.00 Froese and Pauly 2003 1 0 n.a. minor 

Callionymidae 

Callionymidae 15.00 Froese and Pauly 2003 1 1 21.50 no 

Gobiidae 

Acanthogobius flavimanus 30.00Pomatoschistus spp. 11.00Sufflogobius bibarbatus 17.00Gobiidae n.a.

Froese and Pauly 2003 1 1 9.20 30.67 no 

Froese and Pauly 2003 1 1 3.84 34.91 no 

Froese and Pauly 2003 3 3 6.10 35.88 minor 

1 1 4.50 no 

Gempylidae 

Paradiplospinus gracilis 52.00 Froese and Pauly 2003 2 0 n.a. no

Thyrsites atun 200.00 Froese and Pauly 2003 4 2 78.50 39.25 yes

Trichiuridae 

Lepidopus caudatus 210.00 Froese and Pauly 2003 2 0 n.a. yes

Trichiurus lepturus 234.00 Froese and Pauly 2003 2 1 55.50 23.72 yes

Scombridae 

Auxis thazard 65Scomber australasicus 33.27Scomber japonicus 64.00Scomber scombrus 60.00

Froese and Pauly 2003 1 0 n.a. yes 

Wang et al. 2003 1 1 2.76 8.29 yes 

Froese and Pauly 2003 3 2 27.74 43.34 yes Froese and Pauly 2003 7 5 32.96 54.94 yes 

Centrolophidae 

Icichthys australis 81.00 Froese and Pauly 2003 1 1 31.81 39.27 noSeriolella brama 76.00 Froese and Pauly 2003 1 1 16.70 21.97 yes

Nomeidae 

Cubiceps baxteri 100.00 Froese and Pauly 2003 1 0 n.a. no

Cubiceps pauciradiatus 20.00 Froese and Pauly 2003 3 2 11.11 55.55 no

Stromateidae 

Peprilus triacanthus 30.00 Froese and Pauly 2003 1 1 9.70 32.33 yes 

Scophthalmidae 

Scophthalmus aquosus 45.00 Froese and Pauly 2003 1 0 n.a. yes 

Paralichthyidae 

Citharichthys sordidus 41.00 Citharichthys stigmaeus 17.00 

Paralichthys orbignyanus 50.00 

Froese and Pauly 2003 2 2 10.60 25.85 yes Froese and Pauly 2003 1 1 6.50 38.24 minor 

Froese and Pauly 2003 1 0 n.a. minor 
Appendix 3: Summary food habits data, sorted phylogenetically by prey species. 

Prey Type 

Family Maximum 

Prey Species size 

As % of 

Source for Total N of cases Mean of maximum 

maximum size N of cases in analysis means (cm) size Commercial? Comments 

Pleuronectidae 

Glyptocephalus cynoglossus Glyptocephalus zachirus Hippoglossoides platessoides Isopsetta isolepis Limanda ferruginea Limanda limanda Lyopsetta exilis Microstomus pacificus Parophrys vetulus Platichthys flesus Pleuronectes americanus Pleuronectes platessa Psettichthys melanostictus Reinhardtius hippoglossoides Rhombosolea sp. 

Pleuronectidae 

Soleidae 

Austroglossus pectoralis Solea solea 

Cynoglossidae 

Cynoglossus zanzibarensis 

Osteichthys 

mixed fish taxa 

60.00 59.00 82.00 55.00 64.00 40.00 35.00 76.00 49.00 60.00 64.00 

100.00 

63.00 80.00 

50.00 

n.a. 

60.00 70.00 

32.00 

n.a. 

Froese and Pauly 2003 

Froese and Pauly 2003 Froese and Pauly 2003 Froese and Pauly 2003 Froese and Pauly 2003 Froese and Pauly 2003 

Froese and Pauly 2003 Froese and Pauly 2003 Froese and Pauly 2003 Froese and Pauly 2003 Froese and Pauly 2003 

Froese and Pauly 2003 

Froese and Pauly 2003 

Froese and Pauly 2003 

Lalas 1997 

Froese and Pauly 2003 Froese and Pauly 2003 

Froese and Pauly 2003 

1 

1 4 1 3 1 

1 1 2 1 1 

3 

1 

3 

1 

3 

1 1 

1 

3 

0 

1 2 1 2 1 

1 1 2 1 0 

3 

0 

1 

1 

2 

1 1 

1 

2 

n.a. 

16.50 25.41 18.00 23.38 

8.13 

13.50 15.00 12.10 16.00 

n.a. 

22.02 

n.a. 

35.70 

35.00 

17.85 

27.90 31.83 

20.20 

9.39 

27.97 30.99 32.73 36.53 

20.33 

38.57 19.74 24.69 26.67 

22.02 

44.63 

70.00 

46.50 45.47 

63.13 

yes 

yes yes 

minor 

yes 

yes 

yes yes yes yes 

yes 

yes 

yes 

yes 

yes 

yes 

minor

yes

yes 

n.a. size data presented only in aggregate 

Overall Total 1166 725 18.04 

246
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

247

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Antarctomysis maxima L. carcinophagus 2.40 75 2.00 2.50 Green and Williams 1986 Davis Station, Antarctica 1984 

Boreomysis sp. M. monoceros 4.45 n.a.** 3.00 5.00 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 Gammarus setosus P. hispida 1.30 1.53 35 1.20 2.00 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

Themisto libellula P. hispida 1.30 1.66 279 1.40 3.40 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 Euphausia superba A. gazella 0.75 5.54 324 3.80 6.60 Reid et al. 1996 South Georgia Island, S Atlantic 1986 Euphausia superba A. gazella 0.75 4.45 308 3.60 6.20 Croxall et al. 1999 South Georgia Island, S Atlantic 1994 Euphausia superba A. gazella 0.75 5.54 324 3.80 6.60 Croxall et al. 1999 South Georgia Island, S Atlantic 1986 Euphausia superba A. gazella 0.55 5.13 0.31 301 3.90 6.10 Croxall and Pilcher 1984 South Georgia Island, S Atlantic 1972-1977 

Euphausia superba A. gazella 0.55 5.14 388 3.50 6.50 Doidge and Croxall 1985 South Georgia Island, S Atlantic 1982-1983 

Euphausia superba A. gazella 0.75 4.67 5587 3.20 6.40 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

Euphausia superba A. gazella 1.50 4.03 781 2.40 5.60 Reid 1995 South Georgia Island, S Atlantic 1992 Euphausia superba A. gazella 1.50 4.47 1387 3.60 5.80 Reid 1995 South Georgia Island, S Atlantic 1993 Euphausia superba A. gazella 0.70 4.26 n.a.** McCafferty et al. 1998 South Georgia Island, S Atlantic 1994-1996 

Euphausia superba A. gazella 0.75 4.63 n.a.** 3.20 6.10 Reid and Brierly 2001 South Georgia Island, S Atlantic 1994-1999 

Euphausia superba A. gazella 0.80 5.10 241 4.20 5.90 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Euphausia superba Balaenoptera spp. 23.50 2.66 2745 1.10 6.00 Marr 1962 South Georgia Island, S Atlantic 1932-1933 

Euphausia superba Balaenoptera spp. 23.50 3.38 3562 1.10 6.00 Marr 1962 South Georgia Island, S Atlantic 1939-1940 

Euphausia superba Balaenoptera spp. 23.50 3.94 39081 1.10 6.00 Marr 1962 South Georgia Island, S Atlantic 1929-1930 

Euphausia superba Balaenoptera spp. 23.50 5.00 >2000 1.10 6.50 Mackintosh 1974 Southern Ocean 1950s?? 

Thysanoessa inermis P. hispida 1.30 1.75 189 1.20 2.60 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 Acanthephyra carinata L. hosei 2.33 3.76 5 3.00 5.00 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Acanthephyra quadrispinsoa L. hosei 2.33 1.47 19 1.00 1.90 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Acanthephyra quadrispinsoa S. longirostris 1.70 1.42 15 1.20 1.80 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Notostomus elegans L. hosei 2.33 3.74 5 2.30 4.50 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Chionoectes opilio E. barbatus 2.25 5.70 1.20 336 3.20 12.30 Antonelis et al. 1994b Bering Sea 1981 

Chorismus antarcticus L. weddelii 3.00 1.33 0.14 1496 0.71 2.00 Green and Burton 1987 McMurdo Sound, Antarctica 1983-1985 

Hemipenaeus sp. S. longirostris 1.70 1.53 7 1.40 1.60 Dolar et al. 2003 Sulu Sea, Philippines n.a. Jasus lalandii A. p. pusillus 0.85 4.18 89 1.00 7.00 David 1987 Benguela, South Africa 1981 

Jasus lalandii A. p. pusillus 0.85 5.20 38 1.00 7.00 David 1987 Benguela, South Africa 1984 

Jasus lalandii A. p. pusillus 0.85 2.50 253 1.00 6.00 David 1987 Benguela, South Africa 1985 Notocrangon antarcticus L. weddelii 3.00 1.41 0.14 836 Green and Burton 1987 McMurdo Sound, Antarctica 1983-1985 Sabinea septemcarinata P. hispida 1.30 5.02 19 3.60 5.60 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

Ovalipes catharus P. hookeri 2.00 7.00 118 4.00 11.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Pandalus borealis P. hispida 1.30 7.47 51 4.80 9.20 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

Sepia apama A. p. doriferus 1.70 30.10 5.67 43 21.00 41.30 Gales et al. 1993 Tasmania 1989-1990 

Sepia novaehollandiae A. p. doriferus 1.70 15.50 4.51 23 3.70 20.10 Gales et al. 1993 Tasmania 1989-1990 

Sepia officinalis A. p. pusillus 1.70 16.31 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

Sepia sp. A. p. pusillus 2.05 14.29 76 4.80 21.70 Lipinski and David 1990 Benguela, South Africa 1974-1985 

Rossia macrosoma P. phocoena 1.65 4.40 0.20 2 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Semirossia tenera Delphinus sp. 2.15 1.70 69 1.00 2.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

Semirossia tenera K. sima 2.40 2.00 2 1.80 2.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Semirossia tenera P. blainvillei 1.50 3.80 2 3.20 4.50 dos Santos and Haimovici 2001 Brazil 1985-1998 Sepietta oweniana P. phocoena 1.65 2.60 9 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Loligo bleekeri C. ursinus 1.12 12.45 8 6.93 17.97 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Loligo gahi C. commersonii 1.48 7.60 n.a.** 2.00 18.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

Loligo gahi G. melas 4.06 12.80 34 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

Loligo gahi L. obscurus 2.05 7.50 n.a.** 2.00 23.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

Loligo gahi O. flavescens 1.85 9.70 n.a.** 4.00 23.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 

Loligo opalescens C. ursinus 1.40 43 7.00 15.00 Perez and Bigg 1986 N Pacific 1958-1974 

Loligo opalescens G. macrorhynchus 4.40 13.80 11 11.00 17.09 Sinclair 1992 S California Bight, NE Pacific 1977 

Loligo opalescens G. macrorhynchus 4.63 12.80 11 11.20 14.20 Seagars and Henderson 1985 Channel Islands, NE Pacific 1980 

Loligo opalescens G. macrorhynchus 5.52 16.20 609 8.70 23.20 Sinclair 1992 S California Bight, NE Pacific 1969-1977 

Loligo opalescens L. obliquidens 2.05 4.70 17.60 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

Loligo opalescens M. angustirostris 1.69 11.84 90 6.17 20.51 Sinclair 1994 S California Bight, NE Pacific 1969-1983 

Loligo opalescens P. phocoena 1.65 5.60 13.50 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

SE 0.35 

SE 0.06 

SE 0.04 

SE 0.43 

SE 0.03 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

248

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Loligo opalescens P. phocoena 1.65 8.14 1673 5.80 12.50 Walker et al. 1998 Washington and British Columbia 1990-1997 

Loligo opalescens P. dalli 1.95 4.70 17.40 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? Loligo opalescens P. dalli 1.95 9.52 44 6.90 12.70 Walker et al. 1998 Washington and British Columbia 1990-1997 

Loligo opalescens Z. californianus 2.00 12.89 300 7.50 22.50 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

Loligo opalescens Z. californianus 2.00 9.50 17.50 Morejohn et al. 1978 Monterey Bay, NE Pacific ?? 

Loligo opalescens Z. californianus 2.00 12.70 1.70 76 6.20 18.50 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 

Loligo opalescens Z. californianus 1.80 12.70 25386 1.00 23.50 Lowry and Caretta 1999 Channel Islands, NE Pacific 1981-1985 

Loligo opalescens Z. californianus 1.80 5.20 164 4.20 6.70 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Loligo pealeii G. melas 3.40 16.07 145 5.00 42.00 Gannon et al. 1997b Mid-Atlantic Bight, USA 1973-1993 

Loligo pealeii G. melas 5.90 19.60 578 3.70 43.60 Gannon et al. 1997a NW Atlantic 1989-1991 

Loligo pealeii Globicephala sp. 6.00 13.40 68 Overholtz and Waring 1991 Mid-Atlantic Bight, USA 1989 

Loligo pealeii P. phocoena 1.65 12.90 3.00 8 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Loligo plei Delphinus sp. 2.15 18.00 2 17.70 18.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo plei F. attenuata 2.35 14.90 2 13.60 16.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo plei O. orca 7.65 14.30 2 12.40 16.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo plei P. blainvillei 1.50 15.40 27 6.80 21.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo plei P. blainvillei 1.50 16.60 155 6.60 26.60 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo plei S. fluviatilis 1.75 15.20 137 4.10 26.60 dos Santos and Haimovici 2001 Brazil 1985-1998 Loligo plei S. frontalis 1.75 8.90 121 2.10 22.00 dos Santos and Haimovici 2001 Brazil 1985-1998 Loligo plei S. bredanensis 2.35 21.50 38 15.30 27.80 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo plei T. truncatus 2.88 17.10 2 16.60 17.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo plei T. truncatus 2.88 20.30 10 13.50 23.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis A. australis 1.65 9.80 37 4.50 18.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis A. gazella 1.25 11.10 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis A. tropicalis 1.50 9.30 14 5.80 13.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis Delphinus sp. 2.15 5.00 58 2.80 8.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis L. hosei 2.33 12.90 19 5.10 21.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis O. orca 7.65 12.60 10 5.80 18.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis P. blainvillei 1.50 5.10 593 2.00 21.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis P. blainvillei 1.50 10.30 2686 2.20 22.00 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo sanpaulensis S. fluviatilis 1.75 4.50 260 1.40 19.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

Loligo subulata P. phocoena 1.65 7.10 1.10 71 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Loligo vulgaris reynaudii A. p. pusillus 2.05 11.50 51 7.10 18.90 Lipinski and David 1990 Benguela, South Africa 1974-1985 

Loligo vulgaris reynaudii A. p. pusillus 2.05 15.60 596 4.10 32.40 Lipinski and David 1990 Benguela, South Africa 1974-1985 

Loligo vulgaris reynaudii A. p. pusillus 2.05 16.06 183 Castley et al. 1991 Benguela, South Africa 1976-1990 

Loligo vulgaris reynaudii C. heavisidii 1.57 17.40 4.70 11 9.30 24.40 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Loligo vulgaris reynaudii D. delphis 2.08 17.30 6.80 322 2.50 34.50 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Loligo vulgaris reynaudii G. griseus 2.79 13.00 7.30 67 2.10 32.90 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 

Loligo vulgaris reynaudii K. breviceps 2.52 11.60 5.20 21 5.30 20.90 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 

Loligo vulgaris reynaudii K. sima 2.23 7.10 1.40 11 5.30 8.90 Sekiguchi et al. 1992 Benguela, South Africa 1975-1988 

Loligo vulgaris reynaudii L. obscurus 1.71 12.90 6.60 23 1.70 27.00 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Loligo vulgaris reynaudii P. crassidens 3.73 21.10 4.70 3 16.00 25.30 Sekiguchi et al. 1992 Benguela, South Africa 1971-1982 

Loligo vulgaris reynaudii S. attenuata 2.03 7.10 2.90 7 4.20 11.50 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 

Loligo vulgaris reynaudii S. coeruleoalba 2.03 9.30 4.00 183 2.40 22.30 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 

Loligo vulgaris reynaudii T. truncatus 2.72 22.70 2.00 12 20.20 26.10 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 

Loligo vulgaris vulgaris P. phocoena 1.65 2.90 2 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Loliolus japonica C. ursinus 1.12 9.61 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Lolliguncula brevis P. blainvillei 1.50 4.90 134 2.40 8.40 dos Santos and Haimovici 2001 Brazil 1985-1998 Lolliguncula brevis S. fluviatilis 1.75 4.10 199 2.50 6.00 dos Santos and Haimovici 2001 Brazil 1985-1998 Lolliguncula brevis S. fluviatilis 1.78 4.01 0.79 172 2.84 5.84 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Sepioteuthis australis A. p. doriferus 1.70 32.80 11.33 29 13.50 53.10 Gales et al. 1993 Tasmania 1989-1990 

Ancistrocheirus lesueurii G. melas 5.90 17.20 3 6.10 24.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ancistrocheirus lesueurii G. griseus 3.33 15.80 55 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Ancistrocheirus lesueurii O. orca 7.65 29.30 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ancistrocheirus lesueurii P. macrocephalus 15.75 20.45 60 14.00 30.00 Best 1999 Donkergat, South Africa 1962-1963 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

249

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Ancistrocheirus lesueurii P. macrocephalus 10.00 21.66 73 10.00 40.00 Best 1999 Donkergat, South Africa 1962-1963 

Ancistrocheirus lesueurii P. macrocephalus 15.75 16.40 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Ancistrocheirus lesueurii P. macrocephalus 12.40 23.00 204 16.70 28.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Ancistrocheirus lesueurii S. attenuata 2.05 4.69 2.96 139 0.88 20.31 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 Architeuthis cf. dux P. macrocephalus 12.40 115.90 135 34.60 186.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Architeuthis sp. P. macrocephalus 15.75 107.75 4 82.00 151.00 Best 1999 Donkergat, South Africa 1962-1963 Architeuthis sp. P. macrocephalus 10.00 82.56 9 42.00 116.00 Best 1999 Donkergat, South Africa 1962-1963 Architeuthis sp. S. attenuata 2.05 6.29 2.58 2 4.47 8.12 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Architeuthis sp. P. macrocephalus 15.75 166.30 2 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Bathyteuthis abyssicola G. griseus 3.33 4.00 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Bathyteuthis abyssicola H. planifrons 6.55 7.33 4.61 4 Sekiguchi et al. 1993 South Africa, west coast 1990 

Bathyteuthis abyssicola H. planifrons 7.20 3.60 6 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Batoteuthis skolops H. planifrons 7.20 9.60 6 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Batoteuthis skolops M. leonina 3.45 10.30 3 8.90 10.90 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Brachioteuthis cf. picta A. gazella 1.00 7.08 26 5.69 8.65 Daneri et al. 1999 South Orkney Islands, S Atlantic 1988 

Brachioteuthis cf. picta A. gazella 1.00 8.09 9 7.69 8.55 Daneri et al. 1999 South Shetland Islands, S Atlantic 1992-1994 

Brachioteuthis cf. picta M. leonina 3.45 8.10 192 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Brachioteuthis cf. picta M. leonina 3.45 7.28 26 Daneri et al. 2000 South Shetland Islands, S Atlantic 1995-1996 

Brachioteuthis riisei G. melas 4.06 13.60 9 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 Brachioteuthis riisei H. planifrons 7.20 43.40 32 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? Brachioteuthis cf. riisei A. gazella 0.75 5.84 31 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Brachioteuthis sp. G. griseus 3.33 7.30 7 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Brachioteuthis sp. H. planifrons 6.30 8.57 0.86 5 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Brachioteuthis spp. A. gazella 0.80 5.60 3 4.66 6.27 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Chiroteuthis imperator C. ursinus 1.12 6.14 28 4.48 7.18 Mori et al. 2001 Honshu, NW Pacific 1987-1994 Chiroteuthis picteti P. macrocephalus 15.75 13.90 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Chiroteuthis veranyi G. melas 4.06 16.10 5 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

Chiroteuthis veranyi H. planifrons 7.20 12.40 5 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Chiroteuthis veranyi K. breviceps 3.05 10.40 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Chiroteuthis veranyi K. sima 2.40 11.90 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Chiroteuthis veranyi M. leonina 3.45 14.10 1 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1994 

Chiroteuthis cf. veranyi K. breviceps 3.05 15.82 1 Wang et al. 2002 Taiwan 1998-2000 

Chiroteuthis sp. G. griseus 3.33 8.00 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Chiroteuthis sp. H. planifrons 6.30 10.38 1.05 74 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Chiroteuthis sp. H. planifrons 6.49 10.61 1.34 388 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Chiroteuthis sp. M. layardii 6.15 11.17 0.42 3 Sekiguchi et al. 1996 New Zealand 1966 

Chiroteuthis sp. M. layardii 5.41 10.64 1.12 9 Sekiguchi et al. 1996 South Africa 1965-1995 Chiroteuthis sp. M. leonina 3.45 13.50 6 10.20 15.60 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Chiroteuthis sp. P. macrocephalus 12.40 12.60 168 10.90 20.70 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Planctoteuthis sp. G. griseus 3.33 5.80 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Chtenopteryx sp. K. sima 2.40 7.04 6 6.36 7.91 Wang et al. 2002 Taiwan 1998-2000 

Chtenopteryx sicula K. sima 2.40 7.07 1 Wang et al. 2002 Taiwan 1998-2000 

Chtenopteryx sicula S. attenuata 2.05 3.47 0.63 39 2.17 5.16 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Galiteuthis armata G. melas 4.06 26.80 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 Galiteuthis armata H. planifrons 7.20 27.30 154 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Galiteuthis armata K. sima 2.40 7.93 2 7.34 8.52 Wang et al. 2002 Taiwan 1998-2000 

Galiteuthis armata M. layardii 5.41 19.60 2 Sekiguchi et al. 1996 South Africa 1965-1995 

Galiteuthis armata O. rossii 1.85 19.60 14 13.50 23.70 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

Galiteuthis glacialis H. planifrons 6.30 19.52 1.54 375 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Galiteuthis glacialis H. planifrons 6.49 19.25 1.75 394 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Galiteuthis glacialis M. leonina 3.45 18.70 7 13.00 23.60 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Helicocranchia sp. ? P. macrocephalus 12.40 25.80 4 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Leachia dislocata K. sima 2.40 2.27 2 2.25 2.29 Wang et al. 2002 Taiwan 1998-2000 

Leachia dislocata S. attenuata 2.05 12.79 3.34 628 4.96 24.81 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Liguriella sp. P. macrocephalus 12.40 12.90 1 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

250

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Liocranchia reinhardti H. planifrons 6.55 24.90 3.33 9 Sekiguchi et al. 1993 South Africa, west coast 1990 Liocranchia reinhardti P. macrocephalus 12.40 25.80 4 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Liocranchia reinhardti S. attenuata 2.05 12.54 3.44 238 4.22 19.96 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 Liocranchia sp. H. planifrons 6.43 36.01 21.15 2 Sekiguchi et al. 1993 South Africa, east coast 1975 Liocranchia sp. H. planifrons 6.30 25.40 4.46 334 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Megalocranchia fisheri P. macrocephalus 15.75 96.00 36 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Megalocranchia fisheri A G. griseus 3.33 20.80 240 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Megalocranchia sp. H. planifrons 6.49 19.65 5.98 77 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Megalocranchia sp. S. attenuata 2.05 10.91 5.42 479 0.24 25.39 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Megalocranchia sp. M. layardii 5.41 25.40 6.44 3 Sekiguchi et al. 1996 South Africa 1965-1995 Megalocranchia sp. A P. macrocephalus 12.40 50.50 200 28.30 69.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Megalocranchia sp. B G. griseus 3.33 23.20 45 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Megalocranchia sp. G P. macrocephalus 12.40 81.70 30 49.50 140.80 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Mesonychoteuthis hamiltoni G. melas 4.06 22.00 7 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

Mesonychoteuthis hamiltoni H. planifrons 6.30 92.51 16.12 5 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Mesonychoteuthis hamiltoni H. planifrons 6.49 69.58 25.78 65 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Mesonychoteuthis hamiltoni H. planifrons 7.20 96.40 22 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Mesonychoteuthis hamiltoni M. leonina 3.45 36.20 2 31.90 40.50 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Taonius borealis B. bairdii 11.10 25.20 240 14.26 49.11 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

Taonius borealis B. bairdii 11.10 32.09 85 13.85 45.06 Walker et al. 2002 Sea of Okhotsk 1988-1989 

Taonius megalops H. planifrons 6.43 40.55 4.92 2 Sekiguchi et al. 1993 South Africa, east coast 1975 Taonius pavo H. planifrons 6.30 41.55 7.18 178 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Taonius pavo H. planifrons 6.49 34.91 8.27 614 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Taonius pavo H. planifrons 7.20 37.90 1663 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Taonius pavo K. breviceps 3.05 30.94 70 17.63 49.02 Wang et al. 2002 Taiwan 1998-2000 Taonius pavo K. sima 2.40 20.59 14 10.01 34.83 Wang et al. 2002 Taiwan 1998-2000 Taonius pavo M. layardii 6.15 25.8 1 Sekiguchi et al. 1996 New Zealand 1966 

Taonius pavo M. layardii 5.41 30.56 10.84 39 Sekiguchi et al. 1996 South Africa 1965-1995 Taonius pavo P. dalli 1.89 21.10 12 13.98 30.61 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 Taonius pavo P. macrocephalus 15.75 41.10 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Taonius pavo P. macrocephalus 12.40 41.20 12 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Taonius sp. M. layardii 5.41 27.34 1.38 10 Sekiguchi et al. 1996 South Africa 1965-1995 Teuthowenia maculata P. macrocephalus 12.40 29.90 200 19.20 37.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Teuthowenia megalops H. planifrons 6.55 22.17 4.66 160 Sekiguchi et al. 1993 South Africa, west coast 1990 

Teuthowenia megalops M. layardii 5.41 25.27 2 Sekiguchi et al. 1996 South Africa 1965-1995 

Teuthowenia megalops P. macrocephalus 12.40 30.10 200 20.40 35.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Teuthowenia megalops P. macrocephalus 12.35 23.71 110 14.50 31.50 Santos et al. 1999 North Sea 1990-1996 

Teuthowenia pellucida H. planifrons 6.43 18.55 1.25 8 Sekiguchi et al. 1993 South Africa, east coast 1975 

Teuthowenia pellucida H. planifrons 7.20 18.40 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Teuthowenia pellucida M. layardii 6.15 22.22 0.86 2 Sekiguchi et al. 1996 New Zealand 1966 

Teuthowenia pellucida M. layardii 5.41 18.12 2.39 12 Sekiguchi et al. 1996 South Africa 1965-1995 

Teuthowenia sp. K. breviceps 3.05 18.03 5 15.04 20.55 Wang et al. 2002 Taiwan 1998-2000 Teuthowenia sp. M. layardii 5.41 34.25 1 Sekiguchi et al. 1996 South Africa 1965-1995 

Teuthowenia sp. P. macrocephalus 12.40 19.90 8 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 Cycloteuthis akimushkini P. macrocephalus 15.75 38.90 18 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Cycloteuthis akimushkini P. macrocephalus 12.40 37.00 152 18.90 50.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Cycloteuthis sp. H. planifrons 6.55 46.19 1.32 2 Sekiguchi et al. 1993 South Africa, west coast 1990 

Cycloteuthis sp. M. layardii 5.41 24.80 1 Sekiguchi et al. 1996 South Africa 1965-1995 

Discoteuthis discus H. planifrons 6.55 10.33 0.85 60 Sekiguchi et al. 1993 South Africa, west coast 1990 Discoteuthis discus M. layardii 5.41 9.19 1 Sekiguchi et al. 1996 South Africa 1965-1995 

Discoteuthis laciniosa M. layardii 5.41 12.78 1.97 5 Sekiguchi et al. 1996 South Africa 1965-1995 

Discoteuthis laciniosa P. macrocephalus 15.75 16.10 . 2 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Discoteuthis laciniosa P. macrocephalus 12.40 13.20 200 9.80 15.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Discoteuthis cf. laciniosa H. planifrons 7.20 7.50 13 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Discoteuthis sp. K. breviceps 3.05 14.01 2 8.71 19.31 Wang et al. 2002 Taiwan 1998-2000 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

251

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Discoteuthis sp. K. sima 2.40 13.33 1 Wang et al. 2002 Taiwan 1998-2000 

Abralia redfieldi K. sima 2.40 2.90 23 2.20 3.60 dos Santos and Haimovici 2001 Brazil 1994, 1998 

Abralia sp. K. breviceps 3.05 3.60 5 3.00 4.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Abraliopsis affinis S. attenuata 2.05 3.00 0.49 3671 1.87 4.54 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Abraliopsis cf. lineata L. hosei 2.33 5.09 30 3.70 6.70 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Abraliopsis cf. lineata S. longirostris 1.70 3.82 112 1.70 6.30 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Abraliopsis sp. A G. griseus 3.33 4.40 241 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Abraliopsis sp. B G. griseus 3.33 3.20 46 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Abraliopsis sp. C G. griseus 3.33 3.40 19 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Abraliopsis sp. D G. griseus 3.33 5.30 12 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Enoploteuthis cf. anaspis G. griseus 3.33 7.70 208 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Enoploteuthis chunii C. ursinus 1.12 10.75 3 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Enoploteuthis chunii K. breviceps 3.05 7.17 180 5.59 8.78 Wang et al. 2002 Taiwan 1998-2000 

Enoploteuthis chunii K. sima 2.40 7.13 72 5.63 8.84 Wang et al. 2002 Taiwan 1998-2000 Enoploteuthis chunii S. attenuata 2.05 7.12 877 6.10 8.20 Wang et al. 2003 Taiwan 1994-1995 

Enoploteuthis sp. H. planifrons 7.20 18.00 2 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Enoploteuthis sp. A G. griseus 3.33 9.50 279 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Watasenia scintillans C. ursinus 1.12 4.40 99 3.00 5.60 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Watasenia scintillans C. ursinus 1.12 4.20 978 3.00 5.80 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Watasenia scintillans C. ursinus 1.12 4.30 1248 2.52 6.80 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Watasenia scintillans P. dalli 1.89 3.39 462 2.44 4.51 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Enoploteuthidae H. planifrons 6.55 5.01 1 Sekiguchi et al. 1993 South Africa, west coast 1990 

Berryteuthis anonychus C. ursinus 0.93 29.68 3 27.21 31.99 Mori et al. 2001 N Pacific 1987-1991 Berryteuthis magister B. bairdii 11.10 21.37 697 11.12 33.43 Walker et al. 2002 Sea of Okhotsk 1988-1989 

Berryteuthis magister P. dalli 1.95 13.31 538 5.50 32.50 Walker 1996 Sea of Okhotsk 1988-1989 

Berryteuthis magister P. dalli 1.89 8.30 1.20 22 5.50 10.50 Ohizumi et al. 2000 Sea of Japan May-89 Berryteuthis magister P. dalli 1.89 7.10 4.20 90 0.50 18.50 Ohizumi et al. 2000 Sea of Japan May-95 Berryteuthis magister P. dalli 1.89 11.50 3.60 69 2.50 18.50 Ohizumi et al. 2000 Sea of Japan May-96 Berryteuthis magister P. dalli 1.89 9.30 3.30 62 4.50 17.50 Ohizumi et al. 2000 Sea of Okhotsk Oct-94 

Berryteuthis magister P. dalli 1.89 10.60 4.20 120 4.50 19.50 Ohizumi et al. 2000 Sea of Okhotsk Aug-95 

Berryteuthis magister P. dalli 1.89 12.80 3.60 70 3.50 25.50 Ohizumi et al. 2000 Sea of Okhotsk Jun-88 

Gonatopsis/Berryteuthis C. ursinus 1.16 9.71 166 7.00 17.00 Sinclair et al. 1994 Bering Sea 1982 

Gonatopsis/Berryteuthis C. ursinus 1.24 9.22 136 5.00 17.00 Sinclair et al. 1994 Bering Sea 1985 

Gonatopsis borealis B. bairdii 11.10 17.70 56 9.06 31.02 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

Gonatopsis borealis C. ursinus 0.93 8.22 8 5.96 10.36 Mori et al. 2001 N Pacific 1987-1991 Gonatopsis borealis C. ursinus 1.12 10.40 37 5.56 16.68 Mori et al. 2001 Honshu, NW Pacific 1987-1994 Gonatopsis borealis M. angustirostris 3.60 22.00 5.40 22 13.60 28.00 Antonelis et al. 1994a San Miguel Island, NE Pacific 1978-1979 

Gonatopsis borealis P. dalli 1.89 14.97 28 10.86 21.21 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Gonatopsis makko P. dalli 1.89 9.96 647 2.10 28.91 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 Gonatopsis octopedatus P. dalli 1.89 9.11 560 1.00 15.14 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 Gonatus antarcticus A. gazella 0.75 8.55 4 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Gonatus antarcticus G. melas 4.06 15.10 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

Gonatus antarcticus H. planifrons 6.30 20.54 2.07 191 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Gonatus antarcticus H. planifrons 6.49 25.32 5.93 304 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Gonatus antarcticus H. planifrons 7.20 21.50 274 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? Gonatus antarcticus M. layardii 5.41 34.49 0.66 5 Sekiguchi et al. 1996 South Africa 1965-1995 Gonatus antarcticus M. leonina 3.45 13.90 67 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Gonatus antarcticus M. leonina 3.45 18.53 12 Daneri et al. 2000 South Shetland Islands, S Atlantic 1995-1996 

Gonatus antarcticus O. orca 7.65 20.10 6 18.80 21.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

Gonatus berryi C. ursinus 1.12 9.21 5 6.45 12.80 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Gonatus berryi P. dalli 1.89 13.99 10 4.87 22.48 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Gonatus berryi P. macrocephalus 15.75 28.50 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Gonatus fabricii M. monoceros 4.45 22.00 5473 6.80 40.25 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 

Gonatus fabricii P. macrocephalus 14.80 21.80 2000 Clarke 1997 North Sea 1994 

Gonatus cf. fabricii P. macrocephalus 13.12 22.30 2116 15.50 28.50 Santos et al. 1999 North Sea 1990-1996 

SE 0.14 

SE 0.10 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

252

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Gonatus cf. fabricii P. macrocephalus 12.35 22.02 14468 14.50 29.50 Santos et al. 1999 North Sea 1990-1996 

Gonatus madokai P. dalli 1.89 10.12 459 2.36 17.10 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Gonatus madokai/G. middendorffi C. ursinus 1.24 6.67 86 5.00 9.00 Sinclair et al. 1994 Bering Sea 1985 

Gonatus onyx C. ursinus 1.12 11.40 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Gonatus onyx P. dalli 1.89 8.59 191 5.22 10.76 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 Gonatus steenstrupi P. macrocephalus 12.40 14.90 200 11.50 18.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Gonatus sp. P. dalli 1.89 8.24 872 4.00 14.77 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 Gonatidae C. ursinus 1.40 >59 5.00 24.00 Perez and Bigg 1986 N Pacific 1958-1974 

Gonatidae H. planifrons 6.43 23.63 3.86 3 Sekiguchi et al. 1993 South Africa, east coast 1975 Gonatidae M. layardii 5.41 12.17 4.55 2 Sekiguchi et al. 1996 South Africa 1965-1995 

Histioteuthis arcturi P. macrocephalus 12.40 11.10 200 8.30 13.00 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Histioteuthis bonnellii bonnellii P. macrocephalus 12.40 17.50 200 7.80 29.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Histioteuthis bonnellii corpuscula H. planifrons 6.43 4.90 0.73 6 Sekiguchi et al. 1993 South Africa, east coast 1975 

Histioteuthis bonnellii corpuscula P. macrocephalus 12.40 5.90 20 4.10 6.50 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Histioteuthis cf. celetaria P. macrocephalus 12.40 9.10 200 7.50 13.30 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Histioteuthis eltaninae G. melas 4.06 5.20 32 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

Histioteuthis eltaninae H. planifrons 6.30 4.85 0.46 77 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Histioteuthis eltaninae H. planifrons 6.49 5.90 0.90 132 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Histioteuthis eltaninae H. planifrons 7.20 5.50 752 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Histioteuthis hoylei K. breviceps 3.05 10.47 7 7.81 15.52 Wang et al. 2002 Taiwan 1998-2000 Histioteuthis hoylei P. macrocephalus 15.75 11.60 129 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Histioteuthis machrohista A. gazella 0.80 8.41 1 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Histioteuthis machrohista H. planifrons 6.43 4.42 0.77 8 Sekiguchi et al. 1993 South Africa, east coast 1975 Histioteuthis machrohista M. layardii 5.41 6.30 1.12 21 Sekiguchi et al. 1996 South Africa 1965-1995 Histioteuthis meleagroteuthis H. planifrons 6.43 5.49 0.73 397 Sekiguchi et al. 1993 South Africa, east coast 1975 

Histioteuthis meleagroteuthis M. layardii 6.15 7.75 1 Sekiguchi et al. 1996 New Zealand 1966 

Histioteuthis meleagroteuthis M. layardii 5.41 5.22 0.30 5 Sekiguchi et al. 1996 South Africa 1965-1995 

Histioteuthis meleagroteuthis P. macrocephalus 12.40 8.10 59 5.20 9.60 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Histioteuthis miranda K. breviceps 3.05 9.44 17 6.39 10.83 Wang et al. 2002 Taiwan 1998-2000 

Histioteuthis miranda K. sima 2.40 8.13 13 3.04 10.63 Wang et al. 2002 Taiwan 1998-2000 

Histioteuthis miranda M. layardii 6.15 15.96 1 Sekiguchi et al. 1996 New Zealand 1966 

Histioteuthis miranda P. macrocephalus 15.75 16.67 6 13.00 22.00 Best 1999 Donkergat, South Africa 1962-1963 

Histioteuthis miranda P. macrocephalus 10.00 17.16 19 14.00 21.00 Best 1999 Donkergat, South Africa 1962-1963 

Histioteuthis cf. miranda P. macrocephalus 12.40 15.00 200 11.20 18.90 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Histioteuthis reversa K. breviceps 3.05 5.31 6 4.86 5.79 Wang et al. 2002 Taiwan 1998-2000 

Histioteuthis reversa P. macrocephalus 12.40 5.20 19 4.40 6.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Histioteuthis sp. A G. griseus 3.33 3.10 10 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Histioteuthis sp. B M. leonina 3.45 4.70 58 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Histioteuthis sp. G. melas 5.90 5.60 68 4.80 11.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

Histioteuthis sp. H. planifrons 6.49 5.40 1.71 506 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Histioteuthis sp. M. layardii 5.41 5.83 1.25 58 Sekiguchi et al. 1996 South Africa 1965-1995 Histioteuthis sp. M. leonina 3.45 5.30 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Histioteuthis sp. P. macrocephalus 10.00 4.67 6 3.00 8.00 Best 1999 Donkergat, South Africa 1962-1963 

Histioteuthis sp. P. macrocephalus 15.75 5.50 6 5.00 6.00 Best 1999 Donkergat, South Africa 1962-1963 

Histioteuthis spp. K. breviceps 3.05 7.10 16 5.70 9.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Histioteuthis spp. K. sima 2.40 6.00 94 2.60 13.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

Histioteuthis spp. O. orca 7.65 9.60 7 6.90 13.70 dos Santos and Haimovici 2001 Brazil 1985-1998 

Lepidoteuthis grimaldii P. macrocephalus 12.40 57.90 200 30.60 102.40 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Pholidoteuthis boschmai G. griseus 3.33 19.60 2 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Pholidoteuthis boschmai H. planifrons 6.49 29.60 2.51 7 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Pholidoteuthis boschmai K. breviceps 3.05 29.24 5 25.70 36.63 Wang et al. 2002 Taiwan 1998-2000 

Pholidoteuthis boschmai P. macrocephalus 15.75 26.91 11 16.00 39.00 Best 1999 Donkergat, South Africa 1962-1963 

Pholidoteuthis boschmai P. macrocephalus 15.75 29.85 47 14.00 63.00 Best 1999 Donkergat, South Africa 1962-1963 

Pholidoteuthis boschmai S. attenuata 2.05 9.73 2.55 150 4.29 18.19 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 Pholidoteuthis boschmai P. macrocephalus 10.00 28.84 57 20.00 36.00 Best 1999 Donkergat, South Africa 1962-1963 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

253

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Pholidoteuthis boschmai P. macrocephalus 12.40 41.10 200 19.60 38.40 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Pholidoteuthis cf. boschmai P. macrocephalus 15.75 28.00 6 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Lycoteuthis lorigera A. p. pusillus 2.05 8.71 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

Lycoteuthis lorigera G. griseus 3.33 6.80 1 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Lycoteuthis lorigera H. planifrons 6.55 8.53 1 Sekiguchi et al. 1993 South Africa, west coast 1990 

Lycoteuthis lorigera G. melas 5.90 11.00 45 9.30 14.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

Lycoteuthis lorigera K. breviceps 3.05 8.90 17 7.50 12.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

Lycoteuthis lorigera K. sima 2.40 9.40 7 8.10 10.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

Lycoteuthis lorigera M. leonina 3.45 7.80 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Lycoteuthis lorigera O. orca 7.65 11.00 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Idioteuthis famelica P. macrocephalus 15.75 14.20 21 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Idioteuthis hjorti K. breviceps 3.05 11.00 17 7.58 14.62 Wang et al. 2002 Taiwan 1998-2000 

Mastigoteuthis dentata S. attenuata 2.05 6.91 1.42 870 2.06 13.24 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Mastigoteuthis psychrophila M. leonina 3.45 11.00 5 10.10 13.00 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Mastigoteuthis sp. A A. gazella 0.75 7.65 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Mastigoteuthis sp. A H. planifrons 7.20 18.80 4 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Mastigoteuthis sp. A P. macrocephalus 12.40 15.50 83 12.60 20.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Mastigoteuthis sp. B P. macrocephalus 12.40 18.60 2 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Mastigoteuthis sp. C. ursinus 1.12 13.73 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Mastigoteuthis sp. H. planifrons 6.55 10.79 1.48 20 Sekiguchi et al. 1993 South Africa, west coast 1990 

Mastigoteuthis sp. H. planifrons 6.30 8.57 0.57 12 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Mastigoteuthis sp. M. layardii 5.41 11.37 1.18 22 Sekiguchi et al. 1996 South Africa 1965-1995 Alluroteuthis antarctica G. melas 4.06 29.30 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 Alluroteuthis antarctica H. planifrons 6.30 14.41 1.51 49 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Alluroteuthis antarctica H. planifrons 6.49 40.13 14.06 39 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Alluroteuthis antarctica H. planifrons 7.20 30.80 18 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Alluroteuthis antarctica M. leonina 3.45 14.50 51 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Alluroteuthis antarctica M. leonina 3.45 13.13 13 Daneri et al. 2000 South Shetland Islands, S Atlantic 1992-1994 

Alluroteuthis antarctica O. rossii 1.85 12.20 57 6.10 18.40 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

Octopoteuthis deletron S. attenuata 2.05 4.56 2.35 63 1.03 13.02 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Octopoteuthis cf. deletron K. breviceps 3.05 16.65 6 14.26 18.57 Wang et al. 2002 Taiwan 1998-2000 

Octopoteuthis cf. neilseni P. macrocephalus 15.75 2.50 3 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Octopoteuthis rugosa H. planifrons 6.55 15.02 2.20 17 Sekiguchi et al. 1993 South Africa, west coast 1990 

Octopoteuthis rugosa M. layardii 5.41 7.93 2 Sekiguchi et al. 1996 South Africa 1965-1995 

Octopoteuthis rugosa P. macrocephalus 15.75 15.75 4 15.00 17.00 Best 1999 Donkergat, South Africa 1962-1963 

Octopoteuthis rugosa P. macrocephalus 10.00 21.36 45 9.00 54.00 Best 1999 Donkergat, South Africa 1962-1963 

Octopoteuthis rugosa P. macrocephalus 15.75 2.70 13 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Octopoteuthis rugosa P. macrocephalus 12.40 20.00 200 13.30 24.70 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Octopoteuthis sp. G P. macrocephalus 12.40 32.50 115 23.00 41.20 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Octopoteuthis sp. C. ursinus 1.12 6.68 1 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Octopoteuthis sp. G. melas 5.90 14.40 6 13.00 17.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Octopoteuthis sp. H. planifrons 6.43 16.64 2.34 8 Sekiguchi et al. 1993 South Africa, east coast 1975 

Octopoteuthis sp. K. breviceps 3.05 16.20 4 14.50 19.70 dos Santos and Haimovici 2001 Brazil 1985-1998 

Octopoteuthis sp. K. breviceps 3.05 16.89 2 16.01 17.78 Wang et al. 2002 Taiwan 1998-2000 

Octopoteuthis sp. K. sima 2.40 14.70 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Octopoteuthis sp. M. layardii 5.41 13.48 2 Sekiguchi et al. 1996 South Africa 1965-1995 

Octopoteuthis sp. O. orca 7.65 16.50 4 15.60 19.00 dos Santos and Haimovici 2001 Brazil 1985-1998 

Octopoteuthis spp. G. griseus 3.33 5.70 3 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Octopoteuthis spp. G. griseus 3.33 12.70 3 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Taningia danae H. planifrons 6.49 18.92 13.04 36 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Taningia danae H. planifrons 7.20 44.40 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Taningia danae P. macrocephalus 15.75 42.30 13 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Taningia danae P. macrocephalus 12.40 75.80 200 18.00 158.70 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Dosidicus gigas S. attenuata 2.05 52.27 596 46.71 59.95 Perrin et al. 1973 E Tropical Pacific 1968 

Dosidicus gigas S. attenuata 2.05 18.08 4.31 207 9.92 31.94 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

254

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Dosidicus gigas S. longirostris 1.70 48.47 15 Perrin et al. 1973 E Tropical Pacific 1968 

Eucleoteuthis luminosa C. ursinus 1.12 5.07 10 Mori et al. 2001 Honshu, NW Pacific 1987-1994 Eucleoteuthis luminosa K. breviceps 3.05 13.35 4 12.02 14.65 Wang et al. 2002 Taiwan 1998-2000 

Eucleoteuthis luminosa K. sima 2.40 13.06 4 6.98 18.22 Wang et al. 2002 Taiwan 1998-2000 

Eucleoteuthis luminosa S. attenuata 2.05 11.27 3.39 475 5.16 21.94 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 Eucleoteuthis luminosa S. attenuata 2.05 21.03 23 19.14 24.55 Wang et al. 2003 Taiwan 1994-1995 

Hyaloteuthis pelagica S. attenuata 2.05 7.53 1.27 114 5.06 11.79 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 Illex argentinus A. tropicalis 1.50 33.20 6 29.70 35.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

Illex argentinus C. commersonii 1.48 11.00 n.a.** 6.00 28.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Illex argentinus Delphinus sp. 2.15 5.90 20 2.10 27.60 dos Santos and Haimovici 2001 Brazil 1985-1998 Illex argentinus F. attenuata 2.35 22.70 2 18.70 26.70 dos Santos and Haimovici 2001 Brazil 1985-1998 

Illex argentinus G. melas 5.90 22.10 6 15.00 33.20 dos Santos and Haimovici 2001 Brazil 1985-1998 

Illex argentinus K. breviceps 3.05 21.70 25 14.60 28.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

Illex argentinus K. sima 2.40 22.40 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Illex argentinus L. obscurus 2.05 10.40 n.a.** 4.00 28.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Illex argentinus M. leonina 3.45 26.30 2 26.00 26.50 dos Santos and Haimovici 2001 Brazil 1985-1998 

Illex argentinus O. flavescens 1.85 20.90 n.a.** 14.00 30.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Illex argentinus O. flavescens 1.85 25.00 n.a.** 18.00 30.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Illex illecebrosus H. grypus 2.08 18.50 78 10.00 25.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 Illex illecebrosus P. vitulina 1.55 17.60 139 5.00 20.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

Illex illecebrosus P. vitulina 1.55 19.10 51 15.00 25.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 

Martialia hyadesi A. gazella 1.25 17.46 1.13 4 15.80 18.30 North 1996 South Georgia Island, S Atlantic 1983 Martialia hyadesi A. gazella 0.75 22.07 39 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Martialia hyadesi H. planifrons 6.30 22.60 1.52 65 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Martialia hyadesi H. planifrons 7.20 30.10 25 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Martialia hyadesi M. leonina 3.45 27.10 43 19.3 42.6 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 Nototodarus gouldi A. p. doriferus 1.70 23.90 6.99 213 8.80 41.50 Gales et al. 1993 Tasmania 1989-1990 

Nototodarus hawaiiensis S. attenuata 2.05 7.08 0.99 5 6.5 8.79 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Nototodarus sloanii A. forsteri 1.40 § 19.00 386 5.00 43.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

Ommastrephes bartramii A. p. pusillus 2.05 20.82 9 Castley et al. 1991 Benguela, South Africa 1976-1990 

Ommastrephes bartramii A. tropicalis 1.50 22.10 23 9.30 34.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ommastrephes bartramii C. ursinus 0.93 8.61 10 7.09 9.96 Mori et al. 2001 N Pacific 1987-1991 Ommastrephes bartramii C. ursinus 1.12 16.24 24 12.65 21.76 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Ommastrephes bartramii G. griseus 3.33 35.40 17 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Ommastrephes bartramii L. obliquidens 2.05 32.10 8 27.00 39.50 Walker and Jones 1993 N Pacific 1990 

Ommastrephes bartramii O. orca 7.65 28.70 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ommastrephes bartramii P. macrocephalus 15.75 46.60 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Ommastrephes bartramii P. macrocephalus 12.40 48.60 6 15.50 75.10 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Ommastrephes bartramii P. crassidens 5.50 28.20 5 19.10 32.90 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ommastrephes bartramii S. attenuata 2.05 12.92 3.53 1158 6.97 27.35 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Ornithoteuthis antillarum F. attenuata 2.35 13.10 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ornithoteuthis antillarum K. breviceps 3.05 8.10 2 6.90 9.30 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ornithoteuthis antillarum K. sima 2.40 5.70 24 3.00 9.60 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ornithoteuthis antillarum O. orca 7.65 4.40 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Ornithoteuthis volatilis K. breviceps 3.05 20.12 13 13.28 25.24 Wang et al. 2002 Taiwan 1998-2000 

Ornithoteuthis volatilis K. sima 2.40 17.80 3 9.64 23.10 Wang et al. 2002 Taiwan 1998-2000 Sthenoteuthis oualaniensis K. breviceps 3.05 23.35 20 16.48 30.53 Wang et al. 2002 Taiwan 1998-2000 

Sthenoteuthis oualaniensis K. sima 2.40 15.18 1 Wang et al. 2002 Taiwan 1998-2000 

Sthenoteuthis oualaniensis S. attenuata 2.05 14.42 4.21 287 7.34 32.09 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Sthenoteuthis oualaniensis S. attenuata 2.05 12.36 23 9.31 14.31 Wang et al. 2003 Taiwan 1994-1995 

Todarodes angolensis A. p. pusillus 2.05 14.60 22 3.40 29.10 Lipinski and David 1990 Benguela, South Africa 1974-1985 

Todarodes pacificus C. ursinus 1.12 14.15 3 5.04 33.55 Mori et al. 2001 Honshu, NW Pacific 1987-1994 Todarodes pacificus P. dalli 1.89 7.37 2125 3.09 22.21 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 Todarodes sagittatus G. melas 1.50 17.00 n.a.** 25.00 Desportes and Mouritsen 1988 Faroe Islands, NE Atlantic 1986-1987 Todarodes sagittatus G. melas 3.50 20.00 n.a.** 50.00 Desportes and Mouritsen 1988 Faroe Islands, NE Atlantic 1986-1987 

SE 0.32 SE 0.17 SE 0.36 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

255

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Todarodes sagittatus H. planifrons 6.55 39.82 1 Sekiguchi et al. 1993 South Africa, west coast 1990 

Todarodes sagittatus P. macrocephalus 12.40 48.40 200 25.80 69.60 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Todaropsis eblanae A. p. pusillus 2.05 8.30 94 3.50 16.10 Lipinski and David 1990 Benguela, South Africa 1974-1985 Todaropsis eblanae H. planifrons 6.43 7.14 1 Sekiguchi et al. 1993 South Africa, east coast 1975 

Ommastrephidae M. layardii 5.41 6.00 1 Sekiguchi et al. 1996 South Africa 1965-1995 

Kondakovia longimana A. gazella 0.80 20.20 1 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Kondakovia longimana H. planifrons 6.55 46.94 6.95 59 Sekiguchi et al. 1993 South Africa, west coast 1990 

Kondakovia longimana H. planifrons 6.30 49.36 9.67 99 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Kondakovia longimana H. planifrons 7.20 10.40 77 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Kondakovia longimana M. leonina 3.45 26.19 7 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1995 

Kondakovia longimana O. rossii 1.85 41.20 1 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

Kondakovia longimana P. macrocephalus 16.50 53.50 81 26.00 80.00 Nemoto et al. 1985, 1987 Southern Ocean 1972-1973 

Moroteuthis ingens A. gazella 0.75 16.45 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 Moroteuthis ingens G. melas 4.06 21.20 19 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 

Moroteuthis ingens H. planifrons 6.55 70.36 5.02 5 Sekiguchi et al. 1993 South Africa, west coast 1990 Moroteuthis ingens H. planifrons 7.20 67.20 13 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Moroteuthis knipovitchi A. gazella 0.80 20.44 8 13.40 27.10 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Moroteuthis knipovitchi A. gazella 0.75 9.18 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 Moroteuthis knipovitchi A. tropicalis 1.50 10.54 6 Bester and Laycock 1985 South Georgia Island, S Atlantic 1977-1978 

Moroteuthis knipovitchi H. planifrons 6.30 24.70 8.85 37 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Moroteuthis knipovitchi H. planifrons 6.49 22.10 3.49 34 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Moroteuthis knipovitchi H. planifrons 7.20 17.00 16 Clarke and Goodall 1994 Tierra del Fuego, Argentina late 1970s? 

Moroteuthis knipovitchi M. layardii 5.41 5.80 4.30 3 Sekiguchi et al. 1996 South Africa 1965-1995 

Moroteuthis knipovitchi M. leonina 3.45 23.70 121 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Moroteuthis knipovitchi M. leonina 3.45 22.13 3 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1996 

Moroteuthis knipovitchi O. rossii 1.85 28.40 3 23.20 33.70 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

Moroteuthis loennbergi C. ursinus 1.12 21.19 2 13.88 29.20 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Moroteuthis loennbergi K. breviceps 3.05 17.58 11 11.15 21.21 Wang et al. 2002 Taiwan 1998-2000 

Moroteuthis loennbergi K. sima 2.40 10.77 1 Wang et al. 2002 Taiwan 1998-2000 

Moroteuthis robsoni H. planifrons 6.49 20.71 8.36 60 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Moroteuthis robsoni M. layardii 5.41 28.40 1 Sekiguchi et al. 1996 South Africa 1965-1995 

Moroteuthis robsoni P. macrocephalus 15.75 30.90 10 19.00 42.00 Best 1999 Donkergat, South Africa 1962-1963 

Moroteuthis robsoni P. macrocephalus 10.00 35.87 23 17.00 56.00 Best 1999 Donkergat, South Africa 1962-1963 

Moroteuthis sp. H. planifrons 6.49 22.42 4.42 74 Sekiguchi et al. 1993 South Africa, east and west coasts 1975, 1990 

Onychoteuthis banksii A. p. pusillus 2.05 14.95 2 Castley et al. 1991 Benguela, South Africa 1976-1990 

Onychoteuthis banksii S. attenuata 2.05 7.43 2 646 0.86 13.99 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Onychoteuthis banksii H. planifrons 6.55 21.36 9.73 8 Sekiguchi et al. 1993 South Africa, west coast 1990 Onychoteuthis banksii K. breviceps 3.05 16.63 2 10.77 22.49 Wang et al. 2002 Taiwan 1998-2000 

Onychoteuthis banksii K. sima 2.40 13.76 3 9.25 16.20 Wang et al. 2002 Taiwan 1998-2000 Onychoteuthis banksii S. attenuata 2.05 11.37 47 6.01 19.17 Wang et al. 2003 Taiwan 1994-1995 Onychoteuthis borealijaponica C. ursinus 0.93 15.64 42 6.86 27.58 Mori et al. 2001 N Pacific 1987-1991 

Onychoteuthis borealijaponica C. ursinus 1.12 11.06 9 1.54 35.29 Mori et al. 2001 Honshu, NW Pacific 1987-1994 

Onychoteuthis borealijaponica P. macrocephalus 12.40 31.70 110 16.30 44.00 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Onychoteuthis sp. 1 G. griseus 3.33 8.10 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Onychoteuthis sp. 2 G. griseus 3.33 20.30 22 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Onychoteuthis sp. H. planifrons 6.55 29.26 5.24 10 Sekiguchi et al. 1993 South Africa, west coast 1990 

Onychoteuthis sp. P. macrocephalus 15.75 28.00 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific 1978, 1990 

Onychoteuthidae C. ursinus 1.40 >3 14.00 22.00 Perez and Bigg 1986 N Pacific 1958-1974 

Psychroteuthis glacialis A. gazella 1.00 9.07 7 7.83 10.76 Daneri et al. 1999 South Orkney Islands, S Atlantic 1988 

Psychroteuthis glacialis A. gazella 1.00 10.12 13 7.83 12.83 Daneri et al. 1999 Scotia Arc, Antarctica 1988-1994 Psychroteuthis glacialis A. gazella 1.00 11.26 6 9.98 12.83 Daneri et al. 1999 South Shetland Islands, S Atlantic 1992-1994 

Psychroteuthis glacialis H. planifrons 6.30 9.87 1.96 1513 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Psychroteuthis glacialis L. weddelii 2.00 § 35.10 5 12.30 38.30 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Psychroteuthis glacialis M. leonina 3.45 19.78 231 Daneri et al. 2000 South Shetland Islands, S Atlantic 1995-1996 

Psychroteuthis glacialis O. rossii 1.85 12.30 97 5.80 34.40 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

256

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Pterygioteuthis giardi S. attenuata 2.05 2.18 0.34 65 1.64 3.17 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Pyroteuthis addolux G. griseus 3.33 4.50 5 Clarke and Young 1998 Hawaiian Islands, E Tropical Pacific n.a. 

Argonauta nodosa A. australis 1.65 1.50 3 1.00 1.80 dos Santos and Haimovici 2001 Brazil 1985-1998 

Argonauta nodosa A. p. doriferus 1.70 5.70 3.58 3 1.60 8.20 Gales et al. 1993 Tasmania 1989-1990 

Argonauta nodosa A. p. pusillus 2.05 7.21 3 Castley et al. 1991 Benguela, South Africa 1976-1990 

Argonauta nodosa A. tropicalis 1.50 10.30 2 7.30 13.10 dos Santos and Haimovici 2001 Brazil 1985-1998 

Argonauta nodosa L. hosei 2.33 0.60 1 dos Santos and Haimovici 2001 Brazil 1985-1998 

Argonauta nodosa P. blainvillei 1.50 2.40 55 0.50 4.40 dos Santos and Haimovici 2001 Brazil 1985-1998 

Bathypolypus arcticus P. phocoena 1.65 5.20 1 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Eledone gaucha P. blainvillei 1.50 2.10 1 dos Santos and Haimovici 2001 Brazil 1985-1998 Octopus australis A. p. doriferus 1.70 10.10 2.63 13 4.50 15.50 Gales et al. 1993 Tasmania 1989-1990 

Octopus maorum A. p. doriferus 1.70 15.30 3.03 13 11.70 22.70 Gales et al. 1993 Tasmania 1989-1990 

Octopus maorum P. hookeri 2.00 120.00 33 30.00 150.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Octopus pallidus A. p. doriferus 1.70 11.70 1.40 2 10.70 12.70 Gales et al. 1993 Tasmania 1989-1990 

Octopus superciliosus A. p. doriferus 1.70 5.02 1 Gales et al. 1993 Tasmania 1989-1990 

Octopus tehuelchus P. blainvillei 1.50 2.70 3 2.40 2.90 dos Santos and Haimovici 2001 Brazil 1985-1998 Octopus vulgaris T. truncatus 2.88 11.00 5 7.70 14.70 dos Santos and Haimovici 2001 Brazil 1985-1998 

Octopus sp. A. p. pusillus 2.05 7.42 15 Castley et al. 1991 Benguela, South Africa 1976-1990 

Pareledone charcoti M. leonina 3.45 4.80 5 4.20 4.80 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Pareledone cf. charcoti M. leonina 3.45 5.71 5 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1997 

Pareledone cf. turqueti M. leonina 3.45 7.25 1 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1999 

Pareledone polymorpha M. leonina 3.45 3.60 6 3.00 4.50 Rodhouse et al. 1992 South Georgia Island, S Atlantic 1986, 1988-1989 

Pareledone cf. polymorpha M. leonina 3.45 4.51 1 Daneri et al. 2000 South Shetland Islands, S Atlantic 1993-1998 

Octopodidae G. melas 4.06 16.00 1 Clarke and Goodall 1994 Tierra del Fuego, Argentina 1977-1982 Vampyroteuthis infernalis M. layardii 5.41 5.00 1.26 3 Sekiguchi et al. 1996 South Africa 1965-1995 Vampyroteuthis infernalis P. macrocephalus 12.40 10.40 5 9.60 12.90 Clarke et al. 1993 Azores Islands, N Atlantic 1981-1984 

Myxine glutinosa P. phocoena 1.65 31.60 1 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Squalus acanthias G. melas 3.40 75.00 6 Gannon et al. 1997b Mid-Atlantic Bight, USA 1973-1993 

Raja sp. H. grypus 2.08 70.00 2 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Raja spp. P. hookeri 2.00 68.00 13 53.00 93.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Anguilla anguilla P. phocoena 1.65 26.40 10.00 9 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Alosa pseudoharengus H. grypus 2.08 22.60 10 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Alosa pseudoharengus P. groenlandica 1.70 10.10 3 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Clupea harengus H. grypus 2.08 25.40 5.60 48 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Clupea harengus H. grypus 2.08 24.90 111 20.00 37.30 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Clupea harengus H. grypus 2.08 34.50 160 5.00 40.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

Clupea harengus H. grypus 2.08 n.a.** 25.00 35.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Clupea harengus P. groenlandica 1.70 30.70 n.a.** 23.00 36.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Clupea harengus P. vitulina 1.55 22.00 197 10.00 35.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

Clupea harengus P. vitulina 1.55 25.70 84 15.00 35.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 Clupea harengus P. phocoena 1.65 33.70 47 29.00 38.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 

Clupea harengus P. phocoena 1.65 26.50 136 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 

Clupea harengus P. phocoena 1.65 25.40 3.60 507 18.00 33.00 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Clupea harengus P. phocoena 1.65 26.10 4.90 103 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Clupea harengus pallasi C. ursinus 1.40 >27 10.00 25.00 Perez and Bigg 1986 N Pacific 1958-1974 

Clupea harengus pallasi mixed taxa n.a. 11 28.50 31.30 Stroud et al. 1980 NE Pacific 1958-1972 

Clupea harengus pallasi P. phocoena 1.65 17.86 53 12.30 22.80 Walker et al. 1998 Washington and British Columbia 1990-1997 

Clupea harengus pallasi P. dalli 1.95 15.10 66 10.20 21.70 Walker et al. 1998 Washington and British Columbia 1990-1997 

Etrumeus whiteheadi A. p. pusillus 0.85 14.70 17 David 1987 Benguela, South Africa 1974-1985 

Etrumeus whiteheadi A. p. pusillus 0.85 13.20 32 David 1987 Benguela, South Africa 1974-1985 

Etrumeus whiteheadi D. delphis 2.08 16.20 2.70 171 8.50 23.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 Etrumeus whiteheadi L. obscurus 1.71 11.20 3.50 5 9.20 17.40 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Sardinops sagax = S. ocellatus A. p. pusillus 2.05 21.66 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

Sardinops sagax = S. ocellatus C. heavisidii 1.57 20.60 2 20.60 20.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Sardinops sagax = S. ocellatus D. delphis 2.08 19.00 1.80 163 12.60 24.30 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

SE 0.47 SE 0.36 SE 0.51 

SE 0.32 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

257

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Sardinops sagax = S. ocellatus L. obscurus 1.71 16.60 3.60 21 13.80 25.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Sardinops sagax A. galapagoensis 1.37 14.10 33 Dellinger and Trillmich 1999 Galapagos Islands, E Tropical Pacific 1983 

Sardinops sagax Z. californianus 1.80 9.20 5 8.20 11.50 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Sardinops sagax Z. c. woelbecki 2.00 14.20 66 Dellinger and Trillmich 1999 Galapagos Islands, E Tropical Pacific 1983 

Sardinops sagax Z. c. woelbecki 2.00 13.40 334 Dellinger and Trillmich 1999 Galapagos Islands, E Tropical Pacific 1983-1986 

Sardinops sagax cf. neopilchardus A. p. doriferus 1.70 17.30 1.00 15 15.00 19.00 Gales and Pemberton 1994 Tasmania 1984-1986 

Sardinops sagax cf. ocellatus A. p. pusillus 0.85 14.90 24 David 1987 Benguela, South Africa 1974-1985 

Sardinops sagax cf. ocellatus A. p. pusillus 0.85 19.80 134 David 1987 Benguela, South Africa 1974-1985 

Sardinops sagax cf. ocellatus A. p. pusillus 0.85 18.70 214 David 1987 Benguela, South Africa 1974-1985 

Sardinops sagax melanostictus P. dalli 1.95 18.01 1419 13.25 23.25 Walker 1996 Sea of Okhotsk 1988-1989 

Sardinops sagax cf. melanosticus P. dalli 1.89 17.89 372 12.53 22.46 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Sprattus sprattus P. phocoena 1.65 14.00 0.90 166 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Anchoa sp. S. fluviatilis 1.78 7.34 1.31 17 5.52 10.60 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Engraulis encrasicolus A. p. pusillus 0.85 12.70 1799 David 1987 Benguela, South Africa 1974-1985 

Engraulis encrasicolus A. p. pusillus 0.85 9.90 2911 David 1987 Benguela, South Africa 1974-1985 

Engraulis encrasicolus A. p. pusillus 0.85 9.40 15871 David 1987 Benguela, South Africa 1974-1985 

Engraulis encrasicolus A. p. pusillus 2.05 13.47 1 Castley et al. 1991 Benguela, South Africa 1976-1990 Engraulis encrasicolus C. heavisidii 1.57 10.90 1 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Engraulis encrasicolus D. delphis 2.08 11.20 1.80 413 5.30 18.10 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Engraulis encrasicolus K. breviceps 2.52 10.00 3 10.00 10.00 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 

Engraulis encrasicolus K. sima 2.23 11.60 2.80 7 9.30 17.60 Sekiguchi et al. 1992 Benguela, South Africa 1975-1988 

Engraulis encrasicolus L. obscurus 1.71 9.10 1.80 188 4.40 15.80 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Engraulis encrasicolus T. truncatus 2.72 12.20 1 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 Engraulis japonicus P. dalli 1.89 12.73 978 9.77 17.16 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Engraulis japonicus S. attenuata 2.05 10.77 115 9.30 12.84 Wang et al. 2003 Taiwan 1994-1995 

Engraulis mordax C. ursinus 1.40 27 9.00 18.00 Perez and Bigg 1986 N Pacific 1958-1974 

Engraulis mordax mixed taxa n.a. 19 14.50 17.80 Stroud et al. 1980 NE Pacific 1958-1972 Engraulis mordax P. vitulina 1.55 12.90 81 4.00 15.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 Engraulis mordax Z. californianus 2.00 9.50 0.80 94 5.50 14.10 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 

Engraulis mordax Z. californianus 1.80 10.00 12 9.00 11.10 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Nansenia candida P. dalli 1.95 7.07 6 2.70 14.80 Crawford 1981 N Pacific 1978-1979 

Bathylagidae C. ursinus 1.40 986 8.00 12.00 Perez and Bigg 1986 N Pacific 1958-1974 

Bathylagidae P. dalli 1.95 9.43 60 5.00 17.90 Crawford 1981 N Pacific 1978-1979 

Bathylagidae P. dalli 1.89 11.77 315 11.12 12.46 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Mallotus villosus C. ursinus 1.40 64 7.00 14.00 Perez and Bigg 1986 N Pacific 1958-1974 Mallotus villosus H. grypus 2.08 16.30 0.40 219 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Mallotus villosus H. grypus 2.08 13.90 891 8.30 17.40 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Mallotus villosus H. grypus 2.08 n.a.** 10.00 15.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Mallotus villosus mixed taxa n.a. 26 10.90 15.50 Stroud et al. 1980 NE Pacific 1958-1972 Mallotus villosus P. groenlandica 1.70 16.00 2244 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Mallotus villosus P. groenlandica 1.70 13.20 30 8.80 15.70 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Mallotus villosus P. phocoena 1.65 13.80 0.80 23 13.00 15.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 

Osmerus eperlanus P. vitulina 1.55 10.20 41 Behrends 1982 Waddensea, North Sea 1975-1981 

Osmerus mordax H. grypus 2.08 12.00 1 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Osmerus mordax P. groenlandica 1.70 12.00 4 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Thaleichthys pacificus C. ursinus 1.40 11 12.00 21.00 Perez and Bigg 1986 N Pacific 1958-1974 Thaleichthys pacificus mixed taxa n.a. 4 15.30 20.50 Stroud et al. 1980 NE Pacific 1958-1972 

Thaleichthys pacificus P. phocoena 1.65 15.30 3 12.40 18.10 Walker et al. 1998 Washington and British Columbia 1990-1997 

Thaleichthys pacificus P. dalli 1.95 15.80 21 12.80 17.60 Walker et al. 1998 Washington and British Columbia 1990-1997 

Oncorhynchus gorbuscha mixed taxa n.a. 3 25.00 40.50 Stroud et al. 1980 NE Pacific 1958-1972 

Oncorhynchus kisutch mixed taxa n.a. 5 21.00 33.00 Stroud et al. 1980 NE Pacific 1958-1972 

Oncorhynchus tshawytscha mixed taxa n.a. 22.75 2 21.00 24.50 Stroud et al. 1980 NE Pacific 1958-1972 

Salmo salar H. grypus 2.08 43.00 5 35.70 52.10 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Salmo salar P. groenlandica 1.70 29.10 1 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Salmonidae C. ursinus 1.40 >26 15.00 41.00 Perez and Bigg 1986 N Pacific 1958-1974 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

258

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Gonostoma cf. elongatum L. hosei 2.33 184 8.30 22.50 Robison and Craddock 1983 E Tropical Pacific 1972 

Argyropelecus affinis L. hosei 2.33 177 4.00 7.00 Robison and Craddock 1983 E Tropical Pacific 1972 Maurolicus muelleri A. p. pusillus 0.85 3.40 1 David 1987 Benguela, South Africa 1974-1985 Maurolicus muelleri A. p. pusillus 0.85 3.40 1257 David 1987 Benguela, South Africa 1974-1985 

Maurolicus muelleri P. phocoena 1.65 5.00 0.50 146 3.60 6.80 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Maurolicus muelleri P. dalli 1.89 2.27 661 1.48 3.01 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Maurolicus cf. muelleri L. hosei 2.33 112 2.50 3.50 Robison and Craddock 1983 E Tropical Pacific 1972 Maurolicus weitzmani P. phocoena 1.65 5.00 0.40 5898 3.00 5.50 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Ichthyococcus cf. irregularis L. hosei 2.33 120 4.00 6.90 Robison and Craddock 1983 E Tropical Pacific 1972 Chauliodus cf. barbatus L. hosei 2.33 18.00 238 8.50 20.00 Robison and Craddock 1983 E Tropical Pacific 1972 Rosenblattichthys cf. volucris L. hosei 2.33 67 6.50 14.00 Robison and Craddock 1983 E Tropical Pacific 1972 

Scopelarchus guentheri L. hosei 2.33 127 8.00 16.00 Robison and Craddock 1983 E Tropical Pacific 1972 

Scopelosaurus harryi P. dalli 1.95 15.34 37 7.50 21.50 Crawford 1981 N Pacific 1978-1979 Arctozenus risso P. groenlandica 1.70 13.00 1 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Notolepis coatsi A. gazella 1.00 27.18 12 20.72 30.89 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

Paralepididae L. hosei 2.33 87 20.00 30.00 Robison and Craddock 1983 E Tropical Pacific 1972 Paralepididae P. dalli 1.95 43.00 2 38.00 48.00 Crawford 1981 N Pacific 1978-1979 

Evermanella cf. ahlstromi L. hosei 2.33 119 3.50 9.00 Robison and Craddock 1983 E Tropical Pacific 1972 Ceratoscopelus warmingii L. hosei 2.33 5.93 171 4.20 6.90 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Ceratoscopelus warmingii S. attenuata 2.05 4.93 0.48 114 3.38 6.15 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Ceratoscopelus warmingii S. longirostris 1.70 5.91 420 4.50 6.90 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Diaphus schmidti S. attenuata 2.05 4.20 396 2.65 5.30 Wang et al. 2003 Taiwan 1994-1995 

Diaphus watasei S. attenuata 2.05 12.64 167 8.62 16.69 Wang et al. 2003 Taiwan 1994-1995 

Electrona antarctica A. gazella 1.25 9.80 1.00 22 7.70 11.00 North 1996 South Georgia Island, S Atlantic 1983 

Electrona antarctica A. gazella 1.00 7.24 114 5.50 9.00 Daneri and Coria 1993 South Orkney Islands, S Atlantic 1988 

Electrona antarctica A. gazella 1.00 8.65 0.92 38 7.15 10.36 Daneri 1996 South Shetland Islands, S Atlantic 1992 Electrona antarctica A. gazella 0.75 5.61 16 4.13 8.10 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Electrona antarctica A. gazella 1.25 5.99 112 4.80 8.80 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 Electrona antarctica A. gazella 1.25 7.32 92 5.00 17.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

Electrona antarctica A. gazella 1.50 7.68 2202 3.75 9.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 Electrona antarctica A. gazella 1.00 7.81 75 5.75 9.50 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

Electrona antarctica A. gazella 1.25 10.40 78 8.60 11.60 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

Electrona antarctica A. gazella 1.00 6.20 0.70 340 4.00 8.00 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

Electrona antarctica A. gazella 0.80 10.88 51 6.40 13.26 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Electrona antarctica A. gazella 0.75 6.35 92 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Electrona antarctica A. tropicalis 1.50 7.20 0.30 7 6.90 7.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Electrona carlsbergi A. gazella 1.25 7.10 1.33 56 3.40 9.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Electrona carlsbergi A. gazella 1.25 8.10 18 7.30 8.40 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

Electrona carlsbergi A. gazella 1.00 8.50 1.00 80 6.00 10.10 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 Electrona carlsbergi A. gazella 0.80 8.01 23 6.30 8.98 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Electrona carlsbergi A. gazella 0.75 8.02 26 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Electrona carlsbergi Arctocephalus spp. 1.38 5.87 0.35 30 4.85 6.79 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 

Electrona carlsbergi A. tropicalis 1.50 6.50 0.87 114 3.80 8.80 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Electrona subaspera A. forsteri 1.20 6.94 1.01 42 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Electrona subaspera A. forsteri 1.20 8.55 0.70 57 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Electrona subaspera A. gazella 0.75 7.14 8 4.84 8.41 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Electrona subaspera A. gazella 1.25 8.00 1.34 98 3.90 10.60 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Electrona subaspera A. gazella 1.25 8.52 1.59 103 1.69 12.59 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 Electrona subaspera A. gazella 1.50 10.82 2030 6.25 12.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 Electrona subaspera A. gazella 0.75 7.40 120 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Electrona subaspera Arctocephalus spp. 1.38 7.94 0.57 5 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Electrona subaspera Arctocephalus spp. 1.38 6.53 0.64 22 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Electrona subaspera A. forsteri 1.20 7.65 1.40 15 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Electrona subaspera A. tropicalis 1.50 7.80 1.34 70 5.00 12.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Electrona subaspera A. tropicalis 1.50 8.21 1.48 58 1.69 12.59 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 

SE 0.04 

SE 0.03 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

259

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Electrona subaspera M. leonina 3.45 10.70 1 Green and Burton 1993 Macquarie Island, Southern Ocean 1988 

Electrona sp. A. forsteri 1.40 § 5.00 280 2.00 7.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

Gymnoscopelus bolini A. gazella 0.75 6.50 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Gymnoscopelus bolini A. gazella 1.25 11.70 1.41 12 9.70 14.20 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus bolini A. gazella 0.75 18.50 17 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Gymnoscopelus bolini A. tropicalis 1.50 10.60 2.71 40 6.50 14.50 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus braueri A. gazella 0.75 6.59 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Gymnoscopelus braueri A. gazella 1.00 8.20 1.10 5 6.90 9.60 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

Gymnoscopelus braueri A. gazella 1.25 10.30 6 8.60 11.60 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

Gymnoscopelus braueri A. gazella 0.80 10.63 19 8.16 12.49 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Gymnoscopelus braueri A. gazella 0.75 9.59 1 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Gymnoscopelus braueri M. leonina 3.45 11.20 1 Green and Burton 1993 Heard Island, Indian Ocean 1988 

Gymnoscopelus fraseri A. gazella 0.75 7.90 15 6.77 8.56 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Gymnoscopelus fraseri A. gazella 1.25 8.80 2.08 189 5.80 13.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Gymnoscopelus fraseri A. gazella 0.75 7.62 156 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Gymnoscopelus fraseri A. tropicalis 1.50 7.80 1.46 275 3.50 13.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus microlampas A. gazella 1.25 9.00 2 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus microlampas A. tropicalis 1.50 8.60 0.34 2 8.40 8.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus nicholsi A. gazella 1.00 12.28 109 10.00 14.00 Daneri and Coria 1993 South Orkney Islands, S Atlantic 1988 

Gymnoscopelus nicholsi A. gazella 1.00 14.42 0.77 134 12.60 16.55 Daneri 1996 South Shetland Islands, S Atlantic 1992 Gymnoscopelus nicholsi A. gazella 0.75 8.90 295 5.50 12.89 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Gymnoscopelus nicholsi A. gazella 1.25 12.92 114 5.20 8.80 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

Gymnoscopelus nicholsi A. gazella 1.25 12.78 474 6.00 17.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

Gymnoscopelus nicholsi A. gazella 1.25 8.40 2.22 46 5.70 14.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus nicholsi A. gazella 1.00 13.36 105 11.51 15.29 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

Gymnoscopelus nicholsi A. gazella 1.25 12.90 38 10.80 15.10 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

Gymnoscopelus nicholsi A. gazella 1.00 14.70 2.30 369 6.20 18.90 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

Gymnoscopelus nicholsi A. gazella 0.80 11.34 14 8.79 12.80 Kirkman et al. 2000 Bouvetøya Island, Southern Ocean 1998-1999 

Gymnoscopelus nicholsi A. gazella 0.75 11.74 144 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Gymnoscopelus nicholsi A. forsteri 1.20 13.35 1.51 52 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Gymnoscopelus nicholsi A. forsteri 1.20 12.53 2.33 99 Green et al. 1990 Macquarie Island, Southern Ocean 1988-1989 

Gymnoscopelus nicholsi A. tropicalis 1.50 8.50 2.83 62 4.80 15.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus nicholsi M. leonina 3.45 12.71 1.85 72 7.95 16.71 Daneri and Carlini 2002 South Shetland Islands, S Atlantic 1993-2000 

Gymnoscopelus piabilis A. gazella 0.75 1.30 32 8.22 14.29 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Gymnoscopelus piabilis A. gazella 1.25 12.30 1.80 447 4.70 15.50 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Gymnoscopelus piabilis A. gazella 0.75 12.84 139 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Gymnoscopelus piabilis Arctocephalus spp. 1.38 8.94 3.04 2 6.79 11.09 Goldsworthy et al. 1997 Macquarie Island, Southern Ocean 1990-1991 

Gymnoscopelus piabilis A. tropicalis 1.50 11.60 2.17 283 5.80 16.00 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Krefftichthys anderssoni A. gazella 1.25 3.30 2 3.30 3.30 North 1996 South Georgia Island, S Atlantic 1983 Krefftichthys anderssoni A. gazella 0.75 3.90 7 2.90 5.40 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Krefftichthys anderssoni A. gazella 1.25 5.60 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Krefftichthys anderssoni A. gazella 1.50 4.55 2790 2.75 7.25 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 Krefftichthys anderssoni A. gazella 1.50 3.38 350 2.75 8.25 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 Krefftichthys anderssoni A. gazella 1.25 5.60 1 Klages et al. 1999 Bouvetøya Island, Southern Ocean 1996-1997 

Krefftichthys anderssoni A. gazella 1.00 8.50 0.30 3 8.20 8.70 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

Krefftichthys anderssoni A. gazella 0.75 3.52 47 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Krefftichthys anderssoni A. tropicalis 1.50 4.40 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Lampadena luminosa L. hosei 2.33 255 7.50 10.50 Robison and Craddock 1983 E Tropical Pacific 1972 

Lampadena luminosa L. hosei 2.33 10.20 245 8.10 14.60 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Lampadena luminosa S. longirostris 1.70 8.24 241 4.90 11.40 Dolar et al. 2003 Sulu Sea, Philippines n.a. Lampadena sp. L. hosei 2.33 9 6.00 7.00 Robison and Craddock 1983 E Tropical Pacific 1972 

Lampanyctodes hectoris A. forsteri 1.40 § 4.60 1801 2.00 6.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

Lampanyctodes hectoris A. p. pusillus 0.85 4.80 595 David 1987 Benguela, South Africa 1974-1985 

Lampanyctodes hectoris A. p. pusillus 0.85 5.00 1849 David 1987 Benguela, South Africa 1974-1985 

Lampanyctodes hectoris S. attenuata 2.05 5.66 88 4.43 6.71 Wang et al. 2003 Taiwan 1994-1995 

SE 0.09 

SE 0.09 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

260

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Lampanyctus cf. idostigma L. hosei 2.33 208 6.00 9.50 Robison and Craddock 1983 E Tropical Pacific 1972 

Lampanyctus cf. nobilis L. hosei 2.33 103 7.00 12.00 Robison and Craddock 1983 E Tropical Pacific 1972 Lampichthys procerus A. gazella 1.25 6.90 0.64 4 6.00 7.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Metelectrona ventralis A. gazella 0.75 6.18 16 4.90 7.16 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Metelectrona ventralis A. gazella 1.25 7.10 1.26 78 3.80 10.20 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Metelectrona ventralis A. gazella 0.75 6.90 11 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Metelectrona ventralis A. tropicalis 1.50 7.10 1.44 188 3.20 12.20 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Myctophum asperum S. attenuata 2.05 7.60 675 6.63 8.42 Wang et al. 2003 Taiwan 1994-1995 

Myctophum nitidulum L. hosei 2.33 4.50 3 3.70 5.10 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Myctophum nitidulum S. attenuata 2.05 5.64 0.52 14 4.67 6.31 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Myctophum nitidulum S. longirostris 1.70 5.72 166 4.10 7.30 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Protomyctophum andriashevi A. gazella 0.75 5.12 2 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Protomyctophum bolini A. gazella 1.25 4.80 1.60 3 3.70 6.70 North 1996 South Georgia Island, S Atlantic 1983 

Protomyctophum bolini A. gazella 0.75 3.70 17 3.25 4.14 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Protomyctophum bolini A. gazella 1.25 5.70 1.94 11 2.80 9.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Protomyctophum bolini A. gazella 0.75 3.99 81 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Protomyctophum bolini A. tropicalis 1.50 4.00 0.54 9 2.90 5.00 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Protomyctophum choriodon A. gazella 1.25 7.10 0.50 4 6.50 7.50 North 1996 South Georgia Island, S Atlantic 1983 

Protomyctophum choriodon A. gazella 0.75 7.00 11 5.78 7.85 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Protomyctophum choriodon A. gazella 1.25 8.60 2.07 82 4.00 13.70 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Protomyctophum choriodon A. gazella 0.75 8.52 261 7.25 9.25 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

Protomyctophum choriodon A. gazella 0.75 8.73 241 7.25 9.75 Reid and Arnould 1996 South Georgia Island, S Atlantic 1994 

Protomyctophum choriodon A. gazella 0.75 9.25 215 7.25 11.25 Reid and Arnould 1996 South Georgia Island, S Atlantic 1993 

Protomyctophum choriodon A. gazella 0.75 6.68 114 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Protomyctophum choriodon A. tropicalis 1.50 9.50 1.37 73 5.40 13.10 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Protomyctophum tenisoni A. gazella 0.75 4.02 2 3.52 4.52 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Protomyctophum tenisoni A. gazella 1.25 4.60 0.51 322 1.90 6.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Protomyctophum tenisoni A. gazella 0.75 4.16 32 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Protomyctophum tenisoni A. tropicalis 1.50 4.60 0.60 225 2.60 7.90 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Symbolophorus evermanni L. hosei 2.33 6.48 43 5.80 7.60 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Symbolophorus evermanni S. longirostris 1.70 6.48 243 5.30 7.90 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Symbolophorus sp. A. forsteri 1.40 § 8.50 421 2.00 14.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

Symbolophorus spp. S. attenuata 2.05 6.66 0.7 273 4.09 8.24 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Myctophidae P. dalli 1.95 6.54 166 3.50 10.50 Crawford 1981 N Pacific 1978-1979 Muraenolepis microps A. gazella 1.25 38.30 0.70 4 37.50 39.10 North 1996 South Georgia Island, S Atlantic 1983 

Albatrossia pectoralis B. bairdii 11.10 21.28 5 10.18 34.58 Walker et al. 2002 Honshu, NW Pacific 1985-1991 Albatrossia pectoralis B. bairdii 11.10 21.91 7 16.90 28.16 Walker et al. 2002 Sea of Okhotsk 1988-1989 Coryphaenoides acrolepis B. bairdii 9.95 73.30 11.08 72 45.00 95.00 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

Coryphaenoides acrolepis B. bairdii 11.10 22.07 55 4.06 28.74 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

Coryphaenoides cinereus B. bairdii 11.10 11.92 330 6.99 17.92 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

Coryphaenoides cinereus B. bairdii 10.32 35.84 3.1 3 30.00 35.00 Ohizumi et al. 2003 Abahiri, Japan, Sea of Okhotsk Jul - Aug 1999 

Coryphaenoides cinereus B. bairdii 9.95 35.33 4.25 40 25.00 45.00 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

Coryphaenoides cinereus B. bairdii 11.10 10.85 5 9.13 12.13 Walker et al. 2002 Sea of Okhotsk 1988-1989 

Coryphaenoides longifilis B. bairdii 9.95 75.70 6.54 158 55.00 100.00 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

Nezumia bairdi P. groenlandica 1.70 30.50 9 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Macrouridae A. p. pusillus 0.85 26.10 1 David 1987 Benguela, South Africa 1974-1985 

Macrouridae A. p. pusillus 0.85 26.10 144 David 1987 Benguela, South Africa 1974-1985 Antimora microlepis B. bairdii 11.10 46.94 13 28.74 83.80 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

Auchenoceros punctatus A. forsteri 1.40 § 11.50 871 5.00 15.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

Auchenoceros punctatus P. hookeri 2.00 10.00 166 7.00 13.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Laemonema longipes B. bairdii 11.10 51.01 766 18.73 71.26 Walker et al. 2002 Honshu, NW Pacific 1985-1991 

Laemonema longipes B. bairdii 10.32 17.79 6.3 27 5.00 30 Ohizumi et al. 2003 Abahiri, Japan, Sea of Okhotsk Jul - Aug 1999 

Laemonema longipes B. bairdii 9.95 36.04 9.09 55 15.00 55 Ohizumi et al. 2003 Wada, Japan, N Pacific Jul - Aug 1999 

Laemonema longipes B. bairdii 11.10 35.82 41 30.76 42.78 Walker et al. 2002 Sea of Okhotsk 1988-1989 

Pseudophycis bachus A. p. doriferus 1.70 19.80 5.70 13 15.00 35.00 Gales and Pemberton 1994 Tasmania 1984-1986 

SE 0.44 

SE 0.05 SE 0.05 

SE 0.03 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

261

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Pseudophycis sp. P. hookeri 2.00 36.00 20 20.00 50.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Melanonus sp. L. hosei 2.33 71 12.00 22.00 Robison and Craddock 1983 E Tropical Pacific 1972 

Arctogadus glacialis M. monoceros 4.45 18.60 376 9.30 26.60 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 

Arctogadus glacialis M. monoceros 4.45 20.00 616 10.00 40.00 Heide-Jorgensen et al. 1994 Inglefield Bredning, Greenland 1984, 1985 

Arctogadus glacialis P. groenlandica 1.70 21.10 338 Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

Boreogadus saida D. leucas 4.25 14.54 125 9.00 21.50 Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

Boreogadus saida E. barbatus 2.25 13.40 272 6.90 23.60 Finley and Evans 1983 Baffin Bay, NW Atlantic 1978-1979 

Boreogadus saida M. monoceros 4.45 11.30 607 4.80 21.50 Finley and Gibb 1982 Baffin Bay, NW Atlantic 1978-1979 

Boreogadus saida M. monoceros 4.45 15.70 557 12.00 24.00 Heide-Jorgensen et al. 1994 Inglefield Bredning, Greenland 1984, 1985 

Boreogadus saida P. fasciata 1.60 15.87 92 12.00 21.00 Frost and Lowry 1980 Bering Sea 1976-1979 Boreogadus saida P. groenlandica 1.70 13.40 648 4.80 23.60 Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

Boreogadus saida P. groenlandica 1.70 13.30 7 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Boreogadus saida P. hispida 1.30 10.50 4047 4.80 23.60 Finley et al. 1983, Finley et al. 1990 Baffin Bay, NW Atlantic 1978-1979 

Boreogadus saida P. hispida 1.30 10.90 2.00 4847 4.90 16.10 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 Boreogadus saida P. largha 1.75 14.90 326 7.60 20.50 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

Eleginus gracilis D. leucas 4.25 12.48 530 5.50 30.50 Seaman et al. 1982 Bering Sea 1978 

Eleginus gracilis P. largha 1.75 16.70 131 6.20 25.10 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

Gadus morhua C. cristata 2.25 36.40 68 14.00 73.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Gadus morhua H. grypus 2.08 17.30 11.40 101 10.00 40.00 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 Gadus morhua H. grypus 2.08 32.14 63 5.00 65.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Gadus morhua H. grypus 2.08 32.21 61 5.00 75.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Gadus morhua H. grypus 2.08 28.27 49 5.00 75.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Gadus morhua H. grypus 2.08 29.75 180 2.50 72.50 Prime and Hammond 1990 Donna Nook, SW North Sea 1985 

Gadus morhua H. grypus 2.08 31.01 172 2.50 67.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Gadus morhua H. grypus 2.08 32.29 145 2.50 77.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Gadus morhua H. grypus 2.08 28.20 269 6.70 79.70 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Gadus morhua H. grypus 2.08 25.80 SE 0.92 115 10.00 55.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

Gadus morhua H. grypus 2.08 18.80 81 2.50 37.50 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

Gadus morhua H. grypus 2.08 468 30.00 50.00 Hauksson and Bogason 1997 Iceland 1992-1993 Gadus morhua P. groenlandica 1.70 23.80 2 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Gadus morhua P. groenlandica 1.70 24.70 25 15.00 40.00 Nilssen et al. 1990 Ullsford, Norway 1986 

Gadus morhua P. groenlandica 1.70 33.40 99 5.00 65.00 Nilssen et al. 1990 Ullsford, Norway 1988 

Gadus morhua P. groenlandica 1.70 14.00 43 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Gadus morhua P. groenlandica 1.70 15.90 38 3.00 48.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Gadus morhua P. vitulina 1.55 14.40 49 Behrends 1982 Waddensea, North Sea 1975-1981 

Gadus morhua P. vitulina 1.55 35.20 33 5.00 50.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

Gadus morhua P. vitulina 1.55 19.00 46 10.00 35.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 

Gadus morhua P. phocoena 1.65 29.00 5 26.00 32.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 

Gadus morhua P. phocoena 1.65 15.90 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 

Gadus morhua P. phocoena 1.65 24.10 13.30 5 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Gadus morhua P. phocoena 1.65 28.10 5.80 30 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Gadus ogac P. groenlandica 1.70 14.00 3 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Melanagrammus aeglefinus H. grypus 2.08 23.50 2 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Melanagrammus aeglefinus H. grypus 2.08 32.40 2 14.80 57.20 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Melanagrammus aeglefinus H. grypus 2.08 20 20.00 40.00 Hauksson and Bogason 1997 Iceland 1992-1993 Melanagrammus aeglefinus P. groenlandica 1.70 16.80 1 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Melanagrammus aeglefinus P. phocoena 1.65 18.20 3.00 4 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Merlangius merlangus H. grypus 2.08 21.25 1318 9.00 37.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Merlangius merlangus H. grypus 2.08 26.50 203 9.00 43.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Merlangius merlangus P. phocoena 1.65 18.70 4.00 65 14.00 29.00 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Merlangius merlangus P. phocoena 1.65 5.77 2.60 449 2.00 12.00 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Microgadus proximus P. vitulina 1.55 14.00 8 4.00 23.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 Micromesistius poutassou P. phocoena 1.65 27.80 13.70 5 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Pollachius pollachius P. phocoena 1.65 5.60 0.10 2 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Pollachius virens H. grypus 2.08 37.94 79 7.50 72.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

SE 2.08 SE 0.75 

SE 1.72 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

262

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Pollachius virens H. grypus 2.08 45.71 63 22.50 92.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Pollachius virens H. grypus 2.08 10.90 2 8.10 13.80 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Pollachius virens H. grypus 2.08 20.00 55 5.00 35.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

Pollachius virens H. grypus 2.08 48 10.00 60.00 Hauksson and Bogason 1997 Iceland 1992-1993 Pollachius virens P. vitulina 1.55 20.50 19 5.00 30.00 Bowen and Harrison 1996 Bay of Fundy, NW Atlantic 1988-1992 

Pollachius virens P. vitulina 1.55 17.70 122 10.00 35.00 Bowen and Harrison 1996 Nova Scotia, Canada 1988-1992 

Pollachius virens P. phocoena 1.65 19.50 10.10 76 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Pollachius virens P. phocoena 1.65 17.40 6.40 91 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Theragra chalcogramma C. ursinus 1.40 19.30 n.a.** 10.00 35.00 McAlister et al. 1976 Bering Sea 1974 

Theragra chalcogramma C. ursinus 1.21 32.97 39 6.00 41.00 Sinclair et al. 1994 Bering Sea 1981 

Theragra chalcogramma C. ursinus 1.16 6.70 1191 3.00 42.00 Sinclair et al. 1994 Bering Sea 1982 

Theragra chalcogramma C. ursinus 1.24 13.03 1428 4.00 47.00 Sinclair et al. 1994 Bering Sea 1985 

Theragra chalcogramma C. ursinus 1.40 29.19 59 6.80 48.00 Kiyota et al. 1999 St. Paul Island, Bering Sea 1996 

Theragra chalcogramma C. ursinus 1.40 n.a.** 6.00 20.00 Swartzman and Haar 1983 NE Pacific & Bering Sea 1960-1974 

Theragra chalcogramma C. ursinus 1.40 30.40 n.a.** Lowry et al. 1989 Bering Sea 1981-1982 

Theragra chalcogramma E. jubatus 2.55 25.20 1135 8.30 64.20 Frost and Lowry 1986 Bering Sea 1981 

Theragra chalcogramma E. jubatus 2.55 29.80 11.60 2030 5.60 62.90 Pitcher 1981 Gulf of Alaska 1975-1978 

Theragra chalcogramma E. jubatus 2.55 46.90 280 18.40 61.40 Frost and Lowry 1986 Pribilof Islands, Bering Sea 1976, 1979 

Theragra chalcogramma E. jubatus 2.20 25.50 80 4.80 55.70 Calkins and Goodwin 1988 Kodiak Island, Gulf of Alaska 1985-1986 

Theragra chalcogramma E. jubatus 2.25 25.40 1064 7.90 54.20 Calkins and Goodwin 1988 SE Alaska, Gulf of Alaska 1985-1986 

Theragra chalcogramma E. jubatus 2.55 42.40 11.6 909 10.00 78.10 Tollit et al. 2004a SE Alaska 1994-1999 

Theragra chalcogramma E. jubatus 2.28 39.30 14.3 666 3.70 70.80 Zeppelin et al. 2004 Aleutian Is. (western stock) 1998-2000 

Theragra chalcogramma E. barbatus 2.25 11.80 0.15 56 6.90 14.30 Antonelis et al. 1994b Bering Sea 1981 

Theragra chalcogramma P. fasciata 1.60 10.26 447 6.00 20.00 Frost and Lowry 1980 Bering Sea 1976-1979 

Theragra chalcogramma P. fasciata 1.60 11.20 468 6.50 34.40 Frost and Lowry 1986 Bering Sea 1976-1978 

Theragra chalcogramma P. largha 2.25 10.90 21 8.00 15.00 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

Theragra chalcogramma P. largha 1.75 10.90 21 8.00 15.00 Frost and Lowry 1986 Bering Sea 1978 

Theragra chalcogramma P. vitulina 1.55 31.80 23 10.30 56.30 Frost and Lowry 1986 Pribilof Islands, Bering Sea 1979 

Theragra chalcogramma P. vitulina 1.55 10.60 12 8.20 12.60 Frost and Lowry 1986 Bering Sea 1981 

Theragra chalcogramma P. vitulina 1.55 19.20 9.60 2180 4.20 53.20 Pitcher 1981 Gulf of Alaska 1975-1978 

Theragra chalcogramma B. acutorostrata 7.30 14.50 121 11.80 17.50 Frost and Lowry 1986 Bering Sea 1975 

Theragra chalcogramma B. bairdii 9.95 47.32 8.44 13 35.00 60.00 Ohizumi et al. 2003 Abahiri, Japan, Sea of Okhotsk Jul - Aug 1999 Theragra chalcogramma P. dalli 1.95 22.61 135 6.50 60.50 Walker 1996 Sea of Okhotsk 1988-1989 Theragra chalcogramma P. dalli 1.89 33.64 639 15.08 52.48 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Theragra chalcogramma P. dalli 1.95 25.90 113 12.70 36.30 Walker et al. 1998 Washington and British Columbia 1990-1997 

Theragra chalcogramma B. pysalus 20.70 <=30.00 n.a.** Nemoto 1959 N Pacific 1952-1958 

Fiscus et al. 1964, 

Theragra chalcogramma C. ursinus 1.40 tly <30-35 cm (p. 36)" cited in Frost and Lowry 1986 Bering Sea 1962 

Theragra chalcogramma C. ursinus 1.40 1721 4.00 40.00 Perez and Bigg 1986 N Pacific 1958-1974 

Antonelis pers. com., 

Theragra chalcogramma E. barbatus 2.25 8.20 cited in Frost and Lowry 1986 St. Matthew Island, Bering Sea <1983 

Theragra chalcogramma E. jubatus 2.55 21.80 10.30 51.60 Lowry et al. 1989 St. Matthew Island, Bering Sea 1985 

Theragra chalcogramma E. jubatus 2.55 33.50 20.80 44.50 Lowry et al. 1989 St. Paul Island, Bering Sea 1985 

Theragra chalcogramma E. jubatus 2.55 29.90 1.70 42.70 Lowry et al. 1989 E Aleutian Islands, Gulf of Alaska 1981-1982 

Theragra chalcogramma M. novaeangliae 13.00 >=30.00 n.a.** Nemoto 1959 N Pacific 1952-1958 

Theragra chalcogramma P. phocoena 1.65 18.64 3 18.50 18.80 Walker et al. 1998 Washington and British Columbia 1990-1997 

Trisopterus esmarkii P. phocoena 1.65 12.40 2.20 381 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Trisopterus minutus P. phocoena 1.65 14.70 1.30 14 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Enchelyopus cimbrius P. groenlandica 1.70 25.90 5 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Enchelyopus cimbrius P. phocoena 1.65 23.60 4.70 18 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Molva molva H. grypus 2.08 28.06 81 5.00 85.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Molva molva H. grypus 2.08 32.83 60 5.00 85.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Molva molva H. grypus 2.08 32.53 77 5.00 105.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Molva molva H. grypus 2.08 33.30 256 2.50 77.50 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Urophycis tenuis H. grypus 2.08 22.20 12 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

SE 1.04 SE 1.56 SE 0.38 

SE 1.48 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

263

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Urophycis tenuis P. groenlandica 1.70 27.30 1 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Urophycis spp. P. phocoena 1.65 15.90 14.60 474 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Macruronus magellanicus O. flavescens 1.89 47.50 8.90 206 20.00 70.00 George-Nascimento et al. 1985 Chile 1979-1981 

Merluccius albidus P. groenlandica 1.70 29.60 24 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Merluccius bilinearis H. grypus 2.08 30.30 2 21.10 42.30 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Merluccius bilinearis H. grypus 2.08 25.40 43 10.00 45.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

Merluccius bilinearis H. grypus 2.08 25.67 30 15.00 35.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

Merluccius bilinearis P. groenlandica 1.70 26.00 1 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Merluccius bilinearis P. phocoena 1.65 19.60 173 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 

Merluccius bilinearis P. phocoena 1.65 16.40 9.60 1605 3.00 38.00 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Merluccius capensis A. p. pusillus 2.05 28.19 64 Castley et al. 1991 Benguela, South Africa 1976-1990 

Merluccius hubbsi C. commersonii 1.48 9.90 n.a.** 0.10 20.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Merluccius hubbsi L. obscurus 2.05 20.00 n.a.** 0.10 45.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Merluccius hubbsi O. flavescens 1.85 26.70 n.a.** 10.00 55.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Merluccius hubbsi O. flavescens 1.85 29.10 n.a.** 15.00 60.00 Crespo et al. 1997 San Jorge Gulf, Patagonia 1989-1994 Merluccius merluccius P. phocoena 1.65 14.50 1.50 4 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Merluccius productus C. ursinus 1.40 15.00 2 Perez and Bigg 1986 N Pacific 1958-1974 

Merluccius productus M. angustirostris 3.60 28.74 252 15.40 51.80 Antonelis et al. 1994a San Miguel Island, NE Pacific 1978-1979 

Merluccius productus P. phocoena 1.65 35.30 15 33.80 37.10 Walker et al. 1998 Washington and British Columbia 1990-1997 

Merluccius productus P. dalli 1.95 41.75 7 39.70 43.80 Walker et al. 1998 Washington and British Columbia 1990-1997 

Merluccius productus Z. californianus 2.20 30.70 1856 13.00 58.00 Bailey and Ainley 1982 Farallon Islands, NE Pacific 1974-1978 

Merluccius productus Z. californianus 2.00 16.60 6.00 222 8.90 26.10 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 

Merluccius productus Z. californianus 1.80 18.60 35 4.90 26.80 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Merluccius spp. A. p. pusillus 0.85 29.50 116 David 1987 Benguela, South Africa 1974-1985 Merluccius spp. A. p. pusillus 0.85 17.40 350 David 1987 Benguela, South Africa 1974-1985 Merluccius spp. A. p. pusillus 0.85 21.30 1177 David 1987 Benguela, South Africa 1974-1985 

Merluccius spp. C. heavisidii 1.57 19.50 4.20 160 4.90 28.60 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 Merluccius spp. D. delphis 2.08 18.10 8.00 203 5.10 39.30 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 Merluccius spp. F. attenuata 2.33 6.30 0.50 6 5.70 7.20 Sekiguchi et al. 1992 Benguela, South Africa 1990 

Merluccius spp. K. breviceps 2.52 21.30 2.60 11 16.20 25.50 Sekiguchi et al. 1992 Benguela, South Africa 1975-1990 

Merluccius spp. K. sima 2.23 21.60 3.20 20 16.60 26.80 Sekiguchi et al. 1992 Benguela, South Africa 1975-1988 

Merluccius spp. L. obscurus 1.71 16.50 7.20 216 4.80 42.10 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Merluccius spp. M. densirostris 3.52 6.90 1.20 6 5.90 8.50 Sekiguchi et al. 1992 Benguela, South Africa 1981 

Merluccius spp. S. coeruleoalba 2.03 14.90 2.60 12 7.80 17.20 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 

Merluccius spp. T. truncatus 2.72 28.30 6.00 15 17.10 42.30 Sekiguchi et al. 1992 Benguela, South Africa 1975-1989 

Genypterus blacodes P. hookeri 2.00 4 68.00 124.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Genypterus capensis A. p. pusillus 0.85 45.90 25 David 1987 Benguela, South Africa 1974-1985 

Genypterus capensis A. p. pusillus 2.05 52.73 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

Porichthys notatus P. vitulina 1.55 21.80 73 7.00 39.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 

Chorisochismus dentex A. p. pusillus 0.85 9.50 1 David 1987 Benguela, South Africa 1974-1985 

Cololabis saira C. ursinus 1.40 25.00 4 Perez and Bigg 1986 N Pacific 1958-1974 

Scomberesox saurus A. p. pusillus 0.85 24.20 29 David 1987 Benguela, South Africa 1974-1985 

Oxyporhamphus micropterus S. attenuata 2.05 14.34 1.51 40 12.22 18.01 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Exocoetus moncirrhus S. attenuata 2.05 16.47 1.68 19 13.69 19.37 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Exocoetus volitans S. attenuata 2.05 15.87 2.22 68 9.09 18.74 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Melamphaes sp. P. dalli 1.95 8.38 8 7.80 9.00 Crawford 1981 N Pacific 1978-1979 

Scopelogadus mizolepis bispinosus L. hosei 2.33 143 3.00 6.50 Robison and Craddock 1983 E Tropical Pacific 1972 

Diretmus argenteus L. hosei 2.33 363 6.00 28.50 Robison and Craddock 1983 E Tropical Pacific 1972 Pungitius pungitius D. leucas 4.25 3.20 1 Seaman et al. 1982 Bering Sea and Chukchi Sea 1978 

Helicolenus dactylopterus A. p. pusillus 0.85 20.00 1 David 1987 Benguela, South Africa 1974-1985 Helicolenus dactylopterus A. p. pusillus 0.85 23.20 13 David 1987 Benguela, South Africa 1974-1985 

Helicolenus dactylopterus A. p. pusillus 0.85 20.00 145 David 1987 Benguela, South Africa 1974-1985 

Sebastes marinus P. hispida 1.30 45.00 3286 10.00 120.00 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 

Sebastes sp. C. cristata 2.25 32.40 n.a.** 23.00 43.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Sebastes sp. H. grypus 2.08 n.a.** 20.00 30.00 Hauksson and Bogason 1997 Iceland 1992-1993 

SE 1.03 

SE 0.18 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

264

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Sebastes sp. P. groenlandica 1.70 11.40 257 Murie and Lavigne 1991 Gulf of St. Lawrence, NW Atlantic 1983 

Sebastes sp. Z. californianus 2.00 17.10 2.20 155 12.90 22.70 Antonelis et al. 1984 San Miguel Island, NE Pacific 1978-1979 Sebastes spp. C. ursinus 1.40 >19 11.00 31.00 Perez and Bigg 1986 N Pacific 1958-1974 

Sebastes spp. H. grypus 2.08 14.15 71 10.00 25.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

Sebastes spp. P. phocoena 1.65 3.70 0.30 47 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Sebastes spp. Z. californianus 1.80 12.10 31 9.20 14.60 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Chelidonichthys capensis A. p. pusillus 0.85 15.00 10 David 1987 Benguela, South Africa 1974-1985 

Anoplopoma fimbria mixed taxa n.a. 25.00 1 Stroud et al. 1980 NE Pacific 1958-1972 Pleurogrammus azonus P. dalli 1.89 21.60 154 13.70 28.94 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Pleurogrammus monopterygius C. ursinus 1.40 >5 15.00 23.00 Perez and Bigg 1986 N Pacific 1958-1974 

Pleurogrammus monopterygius E. jubatus 2.28 32.30 5.9 1685 15.30 49.60 Zeppelin et al. 2004 Aleutian Is. (western stock) 1998-2000 

Pleurogrammus monopterygius P. dalli 1.95 13.10 1 Crawford 1981 N Pacific 1978-1979 

Leptocottus armatus P. vitulina 1.55 11.00 85 4.00 21.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

Leptocottus armatus P. vitulina 1.55 14.03 72 5.00 25.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 

Myoxocephalus scorpius H. grypus 2.08 24.60 1 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Myoxocephalus scorpius H. grypus 2.08 n.a.** 15.00 25.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Myoxocephalus scorpius P. hispida 1.30 11.14 35 8.00 18.00 Weslawski et al. 1994 Svalbard, Greenland Sea 1985-1987 Cottidae P. groenlandica 1.70 12.00 7 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Cottidae P. largha 1.75 10.00 19 3.30 16.50 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

Cyclopterus lumpus H. grypus 2.08 15.50 0.50 15 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Cyclopterus lumpus H. grypus 2.08 24.90 26 15.00 29.20 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Decapterus macrosoma S. attenuata 2.05 20.06 55 16.11 22.20 Wang et al. 2003 Taiwan 1994-1995 

Trachurus capensis A. p. pusillus 0.85 10.80 66 David 1987 Benguela, South Africa 1974-1985 

Trachurus capensis A. p. pusillus 0.85 16.30 449 David 1987 Benguela, South Africa 1974-1985 Trachurus capensis A. p. pusillus 0.85 14.80 3542 David 1987 Benguela, South Africa 1974-1985 

Trachurus capensis A. p. pusillus 2.05 27.15 56 Castley et al. 1991 Benguela, South Africa 1976-1990 

Trachurus capensis C. heavisidii 1.57 17.10 2.40 23 10.70 22.20 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 Trachurus capensis D. delphis 2.08 16.20 6.20 134 4.60 34.30 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 Trachurus capensis G. griseus 2.79 32.60 1 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Trachurus capensis L. obscurus 1.71 9.90 3.00 723 4.20 24.90 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Trachurus capensis T. truncatus 2.72 20.10 1.10 8 18.10 21.30 Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Trachurus declivis A. forsteri 1.40 31.00 5.00 145 Lake 1997 Tasmania 1992-1993 

Trachurus declivis A. p. doriferus 1.70 21.20 2.80 48 16.00 31.00 Gales and Pemberton 1994 Tasmania 1984-1986 

Trachurus symmetricus Z. californianus 1.80 13.10 12 10.90 16.20 Melin 2002 San Miguel Island, NE Pacific 1993, 1996 

Trachurus sp. A. forsteri 1.40 3 80.00 90.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 

Trachurus sp. A. forsteri 1.40 10 50.00 60.00 Fea et al. 1999 Otago Peninsula, New Zealand 1993-1994 Trachurus sp. M. monachus 2.40 30.00 50 Sergeant et al. 1979 Madeira, North Atlantic 1957 

Trachurus sp. P. hookeri 2.00 38.00 47 27.00 45.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Emmelichthys nitidus A. forsteri 1.40 16.00 4.00 315 Lake 1997 Tasmania 1992-1993 

Emmelichthys nitidus A. p. doriferus 1.70 17.00 1.20 194 14.00 21.00 Gales and Pemberton 1994 Tasmania 1984-1986 Eucinostomus argenteus S. fluviatilis 1.78 11.91 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Orthopristis ruber S. fluviatilis 1.78 3.73 0.73 2 3.21 4.24 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Pomadasys corvinaeformis S. fluviatilis 1.78 9.06 1.98 7 4.92 10.80 de Oliveira Santos et al. 2002 Brazil 1995-1998 Argyrozona argyrozona A. p. pusillus 0.85 16.10 2 David 1987 Benguela, South Africa 1974-1985 

Argyrozona argyrozona A. p. pusillus 2.05 37.03 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

Cheimerius nufar A. p. pusillus 2.05 11.75 1 Castley et al. 1991 Benguela, South Africa 1976-1990 

Pagellus natalensis A. p. pusillus 2.05 22.87 7 Castley et al. 1991 Benguela, South Africa 1976-1990 Pagrus africanus M. monachus 2.40 35.00 1 Sergeant et al. 1979 Madeira, North Atlantic 1957 

Pterogymnus laniarius A. p. pusillus 0.85 20.00 15 David 1987 Benguela, South Africa 1974-1985 

Pterogymnus laniarius A. p. pusillus 2.05 28.36 15 Castley et al. 1991 Benguela, South Africa 1976-1990 

Argyrosomus hololepidotus A. p. pusillus 2.05 34.68 2 Castley et al. 1991 Benguela, South Africa 1976-1990 Cynoscion jamaicensis S. fluviatilis 1.78 9.35 5.16 2 5.70 13.00 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Cynoscion leiarchus S. fluviatilis 1.78 24.80 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 Genyonemus lineatus P. vitulina 1.55 22.20 40 13.00 29.00 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 Isopisthus parvipinnis S. fluviatilis 1.78 13.98 2.73 56 10.24 19.81 de Oliveira Santos et al. 2002 Brazil 1995-1998 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

265

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Larimus breviceps S. fluviatilis 1.78 10.51 1.60 6 8.71 12.98 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Macrodon ancylodon S. fluviatilis 1.78 28.10 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 Micropogonias furnieri S. fluviatilis 1.78 16.23 7.16 11 5.68 33.08 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Paralonchurus brasiliensis S. fluviatilis 1.78 14.93 3.96 57 3.93 20.81 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Stellifer brasiliensis S. fluviatilis 1.78 10.61 1.82 52 6.09 15.54 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Stellifer rastrifer S. fluviatilis 1.78 6.19 0.99 657 3.48 10.84 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Mugil cephalus A. p. pusillus 0.85 13.80 38 David 1987 Benguela, South Africa 1974-1985 

Mugil spp. S. fluviatilis 1.78 14.76 0.35 2 14.51 15.00 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Cymatogaster aggregata P. vitulina 1.55 8.50 31 6.50 11.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

Notolabrus sp. P. hookeri 2.00 23.00 21 9.00 40.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Tautogolabrus adspersus H. grypus 2.08 25.40 1 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Odax pullus P. hookeri 2.00 31.00 36 22.00 49.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Lycodes sp. P. largha 1.75 27.30 6 22.30 30.90 Bukhtiyarov et al. 1984 Bering Sea 1976-1978 

Lycodopsis pacifica P. phocoena 1.65 10.43 1786 8.00 11.00 Walker et al. 1998 Washington and British Columbia 1990-1997 

Lycodopsis pacifica P. dalli 1.95 9.50 10175 8.00 10.50 Walker et al. 1998 Washington and British Columbia 1990-1997 

Zoarces americanus H. grypus 2.08 53.30 12 45.70 61.00 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Anarhichas lupus H. grypus 2.08 29.60 3 22.90 35.60 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Anarhichas lupus H. grypus 2.08 n.a.** 20.00 60.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Anarrhichthys ocellatus P. vitulina 200.00 1 Baldridge and Rogers 1991 Point Lobos State Reserve, CA 1986 

Dissostichus eleginoides A. gazella 1.25 45.40 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Dissostichus eleginoides H. planifrons 6.30 35.23 16.34 3 Slip et al. 1995 Heard Island, Indian Ocean 1992 

Gobionotothen acuta A. gazella 0.75 7.32 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Gobionotothen gibberifrons A. gazella 1.25 31.60 11.70 55 5.90 54.00 North 1996 South Georgia Island, S Atlantic 1983 

Gobionotothen gibberifrons A. gazella 1.50 42.10 49 23.00 59.00 Reid 1995 South Georgia Island, S Atlantic 1992 

Gobionotothen gibberifrons A. gazella 1.50 43.22 46 21.00 59.00 Reid 1995 South Georgia Island, S Atlantic 1993 Gobionotothen gibberifrons A. gazella 1.00 37.00 1 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 Gobionotothen marionensis A. gazella 1.25 11.60 2.59 4 9.70 15.30 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Lepidonotothen larseni A. gazella 1.25 14.70 3.30 91 7.80 22.00 North 1996 South Georgia Island, S Atlantic 1983 

Lepidonotothen larseni A. gazella 0.75 16.20 142 10.50 23.50 Reid and Arnould 1996 South Georgia Island, S Atlantic 1994 

Lepidonotothen larseni A. gazella 0.75 14.04 110 10.50 23.50 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

Lepidonotothen larseni A. gazella 0.75 18.55 65 13.50 25.50 Reid and Arnould 1996 South Georgia Island, S Atlantic 1993 

Lepidonotothen larseni A. gazella 1.50 16.79 193 4.00 26.00 Reid 1995 South Georgia Island, S Atlantic 1992 

Lepidonotothen larseni A. gazella 1.50 20.30 187 11.00 28.00 Reid 1995 South Georgia Island, S Atlantic 1993 

Lepidonotothen squamifrons A. gazella 0.75 3.53 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Notothenia squamifrons A. gazella 1.25 11.40 4.96 12 3.60 22.40 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Paranotothenia magellanica A. gazella 1.25 14.00 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Paranotothenia magellanica A. tropicalis 1.50 17.90 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Pleuragramma antarcticum A. gazella 1.00 14.26 1.24 136 11.59 17.80 Daneri 1996 South Shetland Islands, S Atlantic 1992 Pleuragramma antarcticum A. gazella 1.00 13.87 33 11.90 16.88 Daneri and Carlini 1999 South Shetland Islands, S Atlantic 1993-1994 

Pleuragramma antarcticum A. gazella 1.00 15.00 1.60 6 12.90 17.00 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

Pleuragramma antarcticum H. leptonyx 2.80 13.76 2.43 226 8.80 17.50 Green and Williams 1986 Davis Station, Antarctica 1984 Pleuragramma antarcticum L. weddelii 3.00 16.24 80 12.00 21.00 Plötz et al. 1991 Weddell Sea, Antarctica 1998 

Pleuragramma antarcticum L. weddelii 3.00 15.41 1296 5.00 21.50 Plötz 1986 Weddell Sea, Antarctica 1983, 1985 

Pleuragramma antarcticum L. weddelii 3.00 16.05 1034 6.00 22.00 Plötz 1986 Weddell Sea, Antarctica 1983, 1985 

Pleuragramma antarcticum L. weddelii 3.00 11.64 2.33 26 5.20 19.10 Green and Burton 1987 Mawson, Antarctica 1983-1985 

Pleuragramma antarcticum L. weddelii 3.00 15.03 1.69 169 7.20 21.00 Green and Burton 1987 Davis Station, Antarctica 1983-1985 

Pleuragramma antarcticum L. weddelii 3.00 14.97 2.33 330 5.20 25.00 Green and Burton 1987 McMurdo Sound, Antarctica 1983-1985 Pleuragramma antarcticum L. weddelii 3.00 14.07 4.27 168 5.07 25.91 Burns et al. 1998 McMurdo Sound, Antarctica 1989-1993 Pleuragramma antarcticum L. carcinophagus 2.40 15.50 4 12.20 18.40 Green and Williams 1986 Davis Station, Antarctica 1984 

Pleuragramma antarcticum M. leonina 3.45 13.15 4.14 16 9.06 22.62 Daneri and Carlini 2002 South Shetland Islands, S Atlantic 1993-2000 

Pleuragramma antarcticum O. rossii 1.85 15.10 9 11.20 18.50 Skinner and Klages 1994 King Haakon VII Sea, Antarctica 1980-1982 

Trematomus eulepidotus L. weddelii 2.00 § 18.70 68 14.40 23.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Trematomus lepidorhinus L. weddelii 2.00 § 14.90 12 12.30 19.30 Plötz et al. 1991 Weddell Sea, Antarctica 1986 Trematomus nicolai L. weddelii 2.00 § 18.90 3 16.70 20.00 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Trematomus spp. L. weddelii 3.00 23.34 7.89 6 12.73 34.10 Burns et al. 1998 McMurdo Sound, Antarctica 1989-1993 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

266

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Artedidraco loennbergi L. weddelii 2.00 8.70 2 8.50 8.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 Dolloidraco longedorsalis L. weddelii 2.00 § 9.40 5 7.70 9.60 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Histiodraco velifer L. weddelii 2.00 9.85 2 8.90 10.80 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Gerlachea australis L. weddelii 2.00 § 20.60 7 19.00 22.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Gymnodraco acuticeps L. weddelii 2.00 § 24.60 7 18.00 29.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Racovitzia glacialis L. weddelii 2.00 § 19.10 65 12.20 22.90 Plötz et al. 1991 Weddell Sea, Antarctica 1986 Chaenocephalus aceratus A. gazella 1.25 49.30 4.50 7 41.40 53.40 North 1996 South Georgia Island, S Atlantic 1983 

Champsocephalus gunnari A. gazella 0.75 21.50 7 18.49 22.94 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Champsocephalus gunnari A. gazella 1.25 21.09 530 12.00 34.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

Champsocephalus gunnari A. gazella 1.50 20.80 1263 18.75 23.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 

Champsocephalus gunnari A. gazella 1.50 22.42 877 18.25 27.75 Green et al. 1997 Heard Island, Indian Ocean 1993-1993 

Champsocephalus gunnari A. gazella 0.75 19.29 9.80 161 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Champsocephalus gunnari M. leonina 3.45 30.11 6 Green and Burton 1993 Heard Island, Indian Ocean 1988 

Champsocephalus gunnari A. gazella 1.25 23.80 8.60 201 10.00 46.00 North 1996 South Georgia Island, S Atlantic 1983 

Champsocephalus gunnari A. gazella 0.75 31.88 328 15.00 45.00 Reid and Arnould 1996 South Georgia Island, S Atlantic 1994 

Champsocephalus gunnari A. gazella 0.75 24.27 158 15.00 47.00 Reid and Arnould 1996 South Georgia Island, S Atlantic 1993 

Champsocephalus gunnari A. gazella 0.75 38.11 18 27.00 49.00 Reid and Arnould 1996 South Georgia Island, S Atlantic 1991 

Champsocephalus gunnari A. gazella 1.50 21.72 1522 12.00 44.00 Reid 1995 South Georgia Island, S Atlantic 1992 

Champsocephalus gunnari A. gazella 1.50 25.09 660 16.00 46.00 Reid 1995 South Georgia Island, S Atlantic 1993 

Champsocephalus gunnari A. tropicalis 1.50 14.10 1 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 Channichthys rhinoceratus A. gazella 1.25 32.52 101 20.00 44.00 Green et al. 1989 Heard Island, Indian Ocean 1987-1988 

Channichthys rhinoceratus M. leonina 3.45 37.83 4 Green and Burton 1993 Heard Island, Indian Ocean 1988 

Chionodraco myersi L. weddelii 2.00 § 27.30 107 20.10 34.40 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Chionodraco rastrospinosus A. gazella 1.00 15.10 2.20 2 13.60 16.70 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 

Cryodraco antarcticus A. gazella 1.00 36.60 2.00 3 35.30 38.90 Casaux et al. 1998 South Shetland Islands, S Atlantic 1996-1997 Cryodraco antarcticus L. weddelii 2.00 § 27.00 28 20.80 32.40 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Pagetopsis maculatus L. weddelii 2.00 § 16.90 66 13.30 20.50 Plötz et al. 1991 Weddell Sea, Antarctica 1986 

Pseudochaenichthys georgianus A. gazella 1.25 28.80 2.40 26 24.10 34.80 North 1996 South Georgia Island, S Atlantic 1983 

Ammodytes dubius H. grypus 2.08 19.10 SE 0.47 139 5.00 25.00 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 Ammodytes hexapterus P. vitulina 1.55 9.50 621 8.00 13.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 Ammodytes hexapterus P. phocoena 1.65 15.70 2 Walker et al. 1998 Washington and British Columbia 1990-1997 

Ammodytes hexapterus P. dalli 1.95 10.90 61 9.30 13.30 Walker et al. 1998 Washington and British Columbia 1990-1997 

Ammodytes personatus P. dalli 1.89 19.44 277 12.75 22.21 Ohizumi et al. 2000 Sea of Okhotsk & Sea of Japan 1988-1996 

Ammodytes sp. H. grypus 2.08 18.10 62 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Ammodytes sp. H. grypus 2.08 n.a.** 15.00 35.00 Hauksson and Bogason 1997 Iceland 1992-1993 

Ammodytes spp. H. grypus 2.08 16.72 324 5.00 31.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Ammodytes spp. H. grypus 2.08 17.50 1878 5.00 43.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Ammodytes spp. H. grypus 2.08 16.29 418 3.00 43.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Ammodytes spp. H. grypus 2.08 16.82 1220 3.00 45.00 Hammond et al. 1994b Hebrides Islands, NE Atlantic 1985 

Ammodytes spp. H. grypus 2.08 13.93 6752 3.00 25.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Ammodytes spp. H. grypus 2.08 13.96 5943 1.00 25.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Ammodytes spp. H. grypus 2.08 16.67 1536 3.00 31.00 Hammond et al. 1994a Orkney Island, NE Atlantic 1985 

Ammodytes spp. H. grypus 2.08 15.00 876 5.00 30.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

Gymnammodytes capensis A. p. pusillus 0.85 25.00 12 David 1987 Benguela, South Africa 1974-1985 

Gymnammodytes capensis A. p. pusillus 0.85 25.00 25 David 1987 Benguela, South Africa 1974-1985 

Ammodytidae H. grypus 2.08 16.37 2341 1.50 34.50 Prime and Hammond 1990 Donna Nook, SW North Sea 1985 

Ammodytidae P. phocoena 1.65 13.10 6.20 13 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

Clinus sp. A. p. pusillus 0.85 11.30 7 David 1987 Benguela, South Africa 1974-1985 

Callionymidae A. p. pusillus 0.85 21.50 86 David 1987 Benguela, South Africa 1974-1985 

Acanthogobius flavimanus P. vitulina 1.55 9.20 2.30 1021 3.50 20.20 Torok 1994 San Francisco Bay, NE Pacific 1991-1992 

Pomatoschistus spp. P. vitulina 1.55 3.84 641 Behrends 1982 Waddensea, North Sea 1975-1981 

Sufflogobius bibarbatus A. p. pusillus 0.85 6.10 64 David 1987 Benguela, South Africa 1974-1985 

Sufflogobius bibarbatus A. p. pusillus 0.85 6.90 48078 David 1987 Benguela, South Africa 1974-1985 

Sufflogobius bibarbatus C. heavisidii 1.57 5.30 2.00 n.a.** Sekiguchi et al. 1992 Benguela, South Africa 1969-1990 

Gobiidae P. phocoena 1.65 4.50 2.00 79 1.50 9.00 Borjesson et al. 2003 Kattegat Sea and Skagerrak Sea 1989-1996 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 4. Raw food habits data, sorted phylogenetically by prey species. See Appendix 1 for common and full scientific names of predator species. 

267

Predator Mean prey range range 

Prey species Predator species length (m) length (cm) SD N low high Reference Locality Dates 

Paradiplospinus gracilis A. gazella 0.75 3.97 1 Cherel et al. 1997 Iles Kerguelen, Indian Ocean 1994 

Paradiplospinus gracilis A. tropicalis 1.50 21.50 2.19 2 19.90 23.00 Klages and Bester 1998 Marion Island, Indian Ocean 1989-1995 

Thyrsites atun A. p. pusillus 0.85 80.00 1 David 1987 Benguela, South Africa 1974-1985 

Thyrsites atun A. p. pusillus 0.85 80.00 2 David 1987 Benguela, South Africa 1974-1985 

Thyrsites atun A. p. pusillus 0.85 80.00 10 David 1987 Benguela, South Africa 1974-1985 

Thyrsites atun P. hookeri 2.00 77.00 14 49.00 86.00 Lalas 1997 Otago Peninsula, New Zealand 1991-1992 

Lepidopus caudatus A. p. pusillus 0.85 75.00 1 David 1987 Benguela, South Africa 1974-1985 Lepidopus caudatus A. p. pusillus 0.85 75.00 5 David 1987 Benguela, South Africa 1974-1985 Trichiurus lepturus S. fluviatilis 1.78 18.29 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 

Trichiurus lepturus S. attenuata 2.05 55.50 74 49.62 65.65 Wang et al. 2003 Taiwan 1994-1995 

Auxis thazard S. attenuata 2.05 21.15 2.29 8 19 26 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Scomber australasicus S. attenuata 2.05 2.76 29 16.39 33.27 Wang et al. 2003 Taiwan 1994-1995 

Scomber japonicus A. p. pusillus 0.85 32.50 6 David 1987 Benguela, South Africa 1974-1985 

Scomber japonicus A. p. pusillus 0.85 24.70 15 David 1987 Benguela, South Africa 1974-1985 

Scomber japonicus S. attenuata 2.05 30.77 15 28.00 33.50 Wang et al. 2003 Taiwan 1994-1995 Scomber scombrus G. melas 5.90 35.80 45 26.50 42.10 Gannon et al. 1997a NW Atlantic 1989-1991 Scomber scombrus Globicephala sp. 6.00 36.32 22 Overholtz and Waring 1991 Mid-Atlantic Bight, USA 1989 

Scomber scombrus H. grypus 2.08 32.60 24 25.00 47.90 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Scomber scombrus H. grypus 2.08 20.90 8 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

Scomber scombrus P. phocoena 1.65 37.70 1.60 10 35.00 40.00 Fontaine et al. 1994 Gulf of St. Lawrence, NW Atlantic 1989 Scomber scombrus P. phocoena 1.65 29.60 Recchia and Read 1989 Bay of Fundy, NW Atlantic 1985-1987 

Scomber scombrus P. phocoena 1.65 22.40 5.30 15 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Icichthys australis A. gazella 0.75 31.81 17 Lea et al. 2002 Iles Kerguelen, Indian Ocean 1998-2000 

Seriolella brama A. p. doriferus 1.70 16.70 1.50 15 14.00 19.00 Gales and Pemberton 1994 Tasmania 1984-1986 

Cubiceps baxteri S. attenuata 2.05 14.09 0.49 3 13.61 14.58 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 

Cubiceps pauciradiatus L. hosei 2.33 11.30 55 8.50 14.20 Dolar et al. 2003 Sulu Sea, Philippines n.a. 

Cubiceps pauciradiatus S. attenuata 2.05 10.91 0.98 233 7.46 12.98 Robertson and Chivers 1997 E Tropical Pacific 1989-1991 Cubiceps pauciradiatus S. longirostris 1.70 11.89 9 11.00 13.40 Dolar et al. 2003 Sulu Sea, Philippines n.a. Peprilus triacanthus P. phocoena 1.65 9.70 1.20 38 Gannon et al. 1998 Gulf of Maine, NW Atlantic 1989-1994 

Scophthalmus aquosus P. groenlandica 1.70 23.50 1 Beck et al. 1993 Gulf of St. Lawrence, NW Atlantic 1988-1990 

Citharichthys sordidus P. vitulina 1.55 6.00 74 4.00 21.50 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

Citharichthys sordidus P. phocoena 1.65 15.20 10 12.50 17.60 Walker et al. 1998 Washington and British Columbia 1990-1997 

Citharichthys stigmaeus P. vitulina 1.55 6.50 29 5.00 10.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

Paralichthys orbignyanus S. fluviatilis 1.78 9.31 1 de Oliveira Santos et al. 2002 Brazil 1995-1998 Glyptocephalus cynoglossus H. grypus 2.08 31.30 1 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Glyptocephalus zachirus P. vitulina 1.55 16.50 113 5.00 28.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

Hippoglossoides platessoides H. grypus 2.08 34.40 5.20 6 Murie and Lavigne 1992 Gulf of St. Lawrence, NW Atlantic 1983 

Hippoglossoides platessoides H. grypus 2.08 24.30 269 11.10 46.80 Benoit and Bowen 1990 Gulf of St. Lawrence, NW Atlantic 1982-1987 

Hippoglossoides platessoides H. grypus 2.08 14.70 9 Bowen et al. 1993 Scotian Shelf, NW Atlantic 1988-1990 

Hippoglossoides platessoides H. grypus 2.08 25.91 23 15.00 40.00 Bowen and Harrison 1994 Scotian Shelf, NW Atlantic 1991-1993 

Isopsetta isolepis P. vitulina 1.55 18.00 10 7.00 26.00 Brown and Mate 1983 Netarts Bay, NE Pacific 1978-1980 

SE 5.05 

SE 1.44 SE 0.17 

SE 0.33 SE 1.56 

§ = median prey size; n.a.** = sample size not available, but probably large. 
Appendix 5. Pairings of food habits and commercial catch data. See Appendix 3 for full scientific names of prey species. See Appendix 1 for common and full scientific names of predator species. See Appendix 7 for key to numbered citations. 

268

Food Commercial 

Prey Predator Prey Range, Range, habits Food habits Comm. Range, Range, catch Commercial 

species species mean N low high citation Food habits locality dates mean N low high citations Commercial catch locality catch dates 

A. pectoralis A. p. pusillus 27.90 10 24 Benguela, South Africa 1976-1990 32.1 n.a.* 23.00 46.00 9 Benguela (ICSEAF Divs. 2.1 and 2.2) 1978-1984 

A. pseudoharengus H. grypus 22.60 10 79 Gulf of St. Lawrence, NW Atlantic 1983 27.14 3M 22.00 33.00 3, 4, 5 Gulf of St. Lawrence (Margaree and Miramichi Rivers) 1983 C. capensis A. p. pusillus 15.00 10 36 Benguela, South Africa 1974-1985 39.4 624 7.50 67.50 75 Benguela, Agulhas Bank Aug 1995 - Jan 1997 

C. gunnari A. gazella 21.09 530 12.00 34.00 54 Heard Island, Indian Ocean 1987-1988 27.3 18060 12.00 40.00 38 Kerguelen Island, Indian Ocean (NE zone) 1987-1988 C. gunnari A. gazella 22.42 877 18.25 27.75 55 Heard Island, Indian Ocean 1993-1993 23.6 1140 18.00 29.00 39 Kerguelen Island, Indian Ocean (outer shelf) 1993 

C. gunnari A. gazella 20.80 1263 18.75 23.75 55 Heard Island, Indian Ocean 1993-1993 23.6 1140 18.00 29.00 39 Kerguelen Island, Indian Ocean (outer shelf) 1993 

C. gunnari A. gazella 23.80 201 10.00 46.00 82 South Georgia Island, S Atlantic 1983 33.2 82974 20.00 47.50 63 South Georgia Island, S Atlantic 

C. harengus H. grypus 24.90 111 20.00 37.30 13 Gulf of St. Lawrence, NW Atlantic 1982-1987 32.0 456 25.00 36.00 2 Gulf of St. Lawrence (NAFO Div. 4T) 1984 

C. harengus H. grypus 25.40 48 79 Gulf of St. Lawrence, NW Atlantic 1983 32.0 456 25.00 36.00 2 Gulf of St. Lawrence (NAFO Div. 4T) 1984 

C. harengus H. grypus 34.50 160 5.00 40.00 19 Scotian Shelf, NW Atlantic 1988-1990 32.8 164M 27.50 39.00 26 Gulf of St. Lawrence (NAFO Div. 4T) 1987 C. harengus P. phocoena 33.70 47 29.00 38.00 46 Gulf of St. Lawrence, NW Atlantic 1989 32.8 164M 27.50 39.00 26 Gulf of St. Lawrence (NAFO Div. 4T) 1987 

C. harengus P. phocoena 25.40 507 18.00 33.00 52 Gulf of Maine, NW Atlantic 1989-1994 32.8 164M 27.50 39.00 26 Gulf of St. Lawrence (NAFO Div. 4T) 1987 

C. harengus P. phocoena 26.50 136 92 Bay of Fundy, NW Atlantic 1985-1987 32.8 164M 27.50 39.00 26 Gulf of St. Lawrence (NAFO Div. 4T) 1987 C. harengus P. vitulina 25.70 84 15.00 35.00 18 Nova Scotia, Canada 1988-1992 32.8 164M 27.50 39.00 26 Gulf of St. Lawrence (NAFO Div. 4T) 1987 

C. harengus P. vitulina 22.00 197 10.00 35.00 18 Bay of Fundy, NW Atlantic 1988-1992 32.8 164M 27.50 39.00 26 Gulf of St. Lawrence (NAFO Div. 4T) 1987 

E. encrasicolus A. p. pusillus 12.70 1799 36 Benguela, South Africa 1974-1985 8.27 n.a.* 4.00 13.00 103 Benguela, west coast 1986 

E. encrasicolus A. p. pusillus 9.90 2911 36 Benguela, South Africa 1974-1985 8.27 n.a.* 4.00 13.00 103 Benguela, west coast 1986 

E. encrasicolus A. p. pusillus 9.40 15871 36 Benguela, South Africa 1974-1985 8.27 n.a.* 4.00 13.00 103 Benguela, west coast 1986 

E. encrasicolus D. delphis 11.20 413 5.30 18.10 103 Benguela, South Africa 1969-1990 8.27 n.a.* 4.00 13.00 103 Benguela, west coast 1986 E. encrasicolus L. obscurus 9.10 188 4.40 15.80 103 Benguela, South Africa 1969-1990 8.27 n.a.* 4.00 13.00 103 Benguela, west coast 1986 

E. superba A. gazella 5.13 301 3.90 6.10 33 South Georgia Island, S Atlantic 1972-1977 4.1 n.a.* 2.00 6.00 104 South Georgia Island, S Atlantic 1975-1980 

E. superba A. gazella 5.54 324 3.80 6.60 96 South Georgia Island, S Atlantic 1986 4.6 n.a.* 2.80 6.00 104 South Orkney and Antarctic Peninsula 1985-1986 

E. whiteheadi A. p. pusillus 13.20 32 36 Benguela, South Africa 1974-1985 15.94 n.a.* 8.00 23.00 103 Benguela, west coast 1986 

E. whiteheadi A. p. pusillus 14.70 17 36 Benguela, South Africa 1974-1985 15.94 n.a.* 8.00 23.00 103 Benguela, west coast 1986 

E. whiteheadi D. delphis 16.20 171 8.50 23.60 103 Benguela, South Africa 1969-1990 15.94 n.a.* 8.00 23.00 103 Benguela, west coast 1986 

G. capensis A. p. pusillus 45.90 25 36 Benguela, South Africa 1974-1985 59.0 108 46.00 108.00 57 Benguela (ICSEAF Divs. 1.3 and 1.4) 1967-1976 

G. morhua H. grypus 28.20 269 6.70 79.70 13 Gulf of St. Lawrence, NW Atlantic 1982-1987 48.9 138K 20.00 90.00 27 Gulf of St. Lawrence (NAFO Div. 4T) 1984 

G. morhua H. grypus 17.30 101 10.00 40.00 79 Gulf of St. Lawrence, NW Atlantic 1983 48.9 138K 20.00 90.00 27 Gulf of St. Lawrence (NAFO Div. 4T) 1984 

G. morhua H. grypus 18.80 81 0.00 35.00 17 Scotian Shelf, NW Atlantic 1991-1993 45.0 134K 23.00 105.00 41 Scotian Shelf (NAFO Div. 4VsW) 1983-1987 

G. morhua H. grypus 25.80 115 10.00 55.00 19 Scotian Shelf, NW Atlantic 1988-1990 45.0 134K 23.00 105.00 41 Scotian Shelf (NAFO Div. 4VsW) 1983-1987 

G. morhua P. vitulina 19.00 46 10.00 35.00 18 Nova Scotia, Canada 1988-1992 45.0 134K 23.00 105.00 41 Scotian Shelf (NAFO Div. 4VsW) 1983-1987 

H. platessoides H. grypus 24.30 77 11.10 46.80 13 Gulf of St. Lawrence, NW Atlantic 1982-1987 33.2 71K 15.00 64.00 27 Gulf of St. Lawrence (NAFO Div. 4T) 1985 

H. platessoides H. grypus 25.91 23 15.00 40.00 17 Scotian Shelf, NW Atlantic 1991-1993 33.2 71K 15.00 64.00 27 Gulf of St. Lawrence (NAFO Div. 4T) 1985 

J. lalandii A. p. pusillus 4.18 89 1.00 7.00 36 Benguela, South Africa 1981 7.91 4332 5.00 11.50 100 Benguela, Olifants River to St. Helena Bay Oct 1996, Mar 1999 J. lalandii A. p. pusillus 5.20 38 1.00 7.00 36 Benguela, South Africa 1984 7.91 4332 5.00 11.50 100 Benguela, Olifants River to St. Helena Bay Oct 1996, Mar 1999 J. lalandii A. p. pusillus 2.50 253 1.00 6.00 36 Benguela, South Africa 1985 7.91 4332 5.00 11.50 100 Benguela, Olifants River to St. Helena Bay Oct 1996, Mar 1999 L. hectoris A. p. pusillus 4.80 595 36 Benguela, South Africa 1974-1985 5.2 737 3.00 7.00 30 Benguela (mostly ICSEAF Div. 1.6) 1968-1976 

L. hectoris A. p. pusillus 5.00 1849 36 Benguela, South Africa 1974-1985 5.2 737 3.00 7.00 30 Benguela (mostly ICSEAF Div. 1.6) 1968-1976 

L. opalescens G. macrorhynchus 16.20 609 8.70 23.20 107 S California Bight, NE Pacific 1969-1977 14.01 n.a.* 40 Monterey Bay, CA 1974 

L. opalescens Z. californianus 5.2 164 4.2 6.7 76 San Miguel Island, NE Pacific 1993, 1996 12.7 6200 5.8 18.5 67 Monterey Bay, CA 1989-1992, 1994 L. v. reynaudii A. p. pusillus 16.06 183 24 Benguela, South Africa 1976-1990 14.8 1000 5.00 33.00 103 Benguela (ICSEAF Div. 2.1) 1986 L. v. reynaudii A. p. pusillus 11.50 51 7.10 18.90 68 Benguela, South Africa 1974-1985 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii A. p. pusillus 15.60 596 4.10 32.40 68 Benguela, South Africa 1974-1985 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii C. heavisidii 17.40 11 9.30 24.40 103 Benguela, South Africa 1969-1990 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii D. delphis 17.30 322 2.50 34.50 103 Benguela, South Africa 1969-1990 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii G. griseus 13.00 67 2.10 32.90 103 Benguela, South Africa 1975-1990 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii K. breviceps 11.60 21 5.30 20.90 103 Benguela, South Africa 1975-1990 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii K. sima 7.10 11 5.30 8.90 103 Benguela, South Africa 1975-1988 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii L. obscurus 12.90 23 1.70 27.00 103 Benguela, South Africa 1969-1990 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii S. coeruleoalba 9.30 183 2.40 22.30 103 Benguela, South Africa 1975-1989 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

L. v. reynaudii T. truncatus 22.70 12 20.20 26.10 103 Benguela, South Africa 1975-1989 15.52 n.a.* 5.00 35.00 103 Benguela, east and west coasts 1986 

M. capensis A. p. pusillus 28.19 64 24 Benguela, South Africa 1976-1990 31.6 480 26.00 34.00 103 Benguela (ICSEAF Div. 2.1) 1986 M. hyadesi H. planifrons 30.10 25 28 Tierra del Fuego, Argentina late 1970s? 29.7 336 19.00 35.00 53 Falkland Islands, S Atlantic May 1995 

M. hyadesi M. leonina 27.10 43 19.30 42.60 98 South Georgia Island, S Atlantic 1986, 1988-1989 29.7 336 19.00 35.00 53 Falkland Islands, S Atlantic May 1995 M. ingens G. melas 21.20 19 28 Tierra del Fuego, Argentina 1977-1982 18.86 4205 5.00 50.50 58 Patgonian Shelf, Falkland Islands 1988-1996 

M. ingens H. planifrons 67.20 13 28 Tierra del Fuego, Argentina late 1970s? 18.86 4205 5.00 50.50 58 Patgonian Shelf, Falkland Islands 1988-1996 Merluccius spp. A. p. pusillus 29.50 116 36 Benguela, South Africa 1974-1985 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 

Merluccius spp. A. p. pusillus 17.40 350 36 Benguela, South Africa 1974-1985 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 

Merluccius spp. A. p. pusillus 21.30 1177 36 Benguela, South Africa 1974-1985 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 

Merluccius spp. C. heavisidii 19.50 160 4.90 28.60 103 Benguela, South Africa 1969-1990 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 

Merluccius spp. D. delphis 18.10 203 5.10 39.30 103 Benguela, South Africa 1969-1990 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 Merluccius spp. K. breviceps 21.30 11 16.20 25.50 103 Benguela, South Africa 1975-1990 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 

Merluccius spp. K. sima 21.60 20 16.60 26.80 103 Benguela, South Africa 1975-1988 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 

Merluccius spp. L. obscurus 16.50 216 4.80 42.10 103 Benguela, South Africa 1969-1990 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 Merluccius spp. S. coeruleoalba 14.90 12 7.80 17.20 103 Benguela, South Africa 1975-1989 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 Merluccius spp. T. truncatus 28.30 15 17.10 42.30 103 Benguela, South Africa 1975-1989 27.19 90K 5.00 75.00 87, 88, 103 Benguela (ICSEAF Div. 1.6) 1984-1986 

P. laniarius A. p. pusillus 28.36 15 24 Benguela, South Africa 1976-1990 32.8 n.a.* 20.00 44.00 15 Benguela (mostly ICSEAF Div. 2.2) 1974-1975 

P. laniarius A. p. pusillus 20.00 15 36 Benguela, South Africa 1974-1985 32.8 n.a.* 20.00 44.00 15 Benguela (mostly ICSEAF Div. 2.2) 1974-1975 

P. monoptyerygius E. jubatus 32.30 1685 15.30 49.60 117 Aleutian Is. (western stock) 1998-2000 39.33 92877 28.00 50.00 117 1998-2000 Pleuronectidae A. p. pusillus 21.10 32 36 Benguela, South Africa 1974-1985 32.1 n.a.* 23.00 46.00 9 Benguela (ICSEAF Divs. 2.1 and 2.2) 1978-1984 Pleuronectidae A. p. pusillus 14.60 144 36 Benguela, South Africa 1974-1985 32.1 n.a.* 23.00 46.00 9 Benguela (ICSEAF Divs. 2.1 and 2.2) 1978-1984 

Oct 1982 - Jun 1983 

Aleutian Islands; trawl fishery 

n.a.* = sample size not available, but probably large 
Appendix 5. Pairings of food habits and commercial catch data. See Appendix 3 for full scientific names of prey species. See Appendix 1 for common and full scientific names of predator species. See Appendix 7 for key to numbered citations. 

269

Food Commercial 

Prey Predator Prey Range, Range, habits Food habits Comm. Range, Range, catch Commercial 

species species mean N low high citation Food habits locality dates mean N low high citations Commercial catch locality catch dates 

R. hippoglossoides M. monoceros 35.70 140 13.80 54.70 65 Disko Bay, Greenland 2002-2003 78.4 n.a.* 30.00 90.00 106 2002 

S. japonicus A. p. pusillus 24.70 15 36 Benguela, South Africa 1974-1985 32.2 1913 11.50 56.50 31 Benguela, west coast 1972-1974 

S. sagax A. p. pusillus 14.90 24 36 Benguela, South Africa 1974-1985 16.15 n.a.* 10.00 21.00 103 Benguela, west coast 1986 

S. sagax A. p. pusillus 19.80 134 36 Benguela, South Africa 1974-1985 16.15 n.a.* 10.00 21.00 103 Benguela, west coast 1986 

S. sagax A. p. pusillus 18.70 214 36 Benguela, South Africa 1974-1985 16.15 n.a.* 10.00 21.00 103 Benguela, west coast 1986 

S. sagax D. delphis 19.00 163 12.60 24.30 103 Benguela, South Africa 1969-1990 16.15 n.a.* 10.00 21.00 103 Benguela, west coast 1986 S. sagax L. obscurus 16.60 21 13.80 25.60 103 Benguela, South Africa 1969-1990 16.15 n.a.* 10.00 21.00 103 Benguela, west coast 1986 S. sagax P. dalli 17.89 372 12.53 22.46 84 Sea of Okhotsk & Sea of Japan 1988-1996 19.2 n.a.* 13.00 22.50 101 NW Pacific, SE of Hokkaido 1988-1992 

S. sagax P. dalli 18.01 1419 13.25 23.25 113 Sea of Okhotsk 1988-1989 18.3 n.a.* 15.00 21.00 101 NW Pacific, SE of Hokkaido 1988 

S. scombrus G. melas 35.80 45 26.50 42.10 50 NW Atlantic 1989-1991 35.2 321M 26.00 43.00 25 NW Atlantic (CAFSAC Divs. 3-6) 1986-1987 

S. scombrus Globicephala sp. 36.32 22 85 Mid-Atlantic Bight, USA 1989 35.3 228M 26.00 43.00 25 NW Atlantic (CAFSAC Divs. 5-6) 1986-1987 

S. scombrus H. grypus 32.60 24 25.00 47.90 13 Gulf of St. Lawrence, NW Atlantic 1982-1987 34.9 92M 26.00 43.00 25 NW Atlantic (CAFSAC Divs. 3-4) 1986-1987 

S. scombrus P. phocoena 37.70 10 35.00 40.00 46 Gulf of St. Lawrence, NW Atlantic 1989 34.9 92M 26.00 43.00 25 NW Atlantic (CAFSAC Divs. 3-4) 1986-1987 

S. scombrus P. phocoena 22.40 15 52 Gulf of Maine, NW Atlantic 1989-1994 35.3 228M 26.00 43.00 25 NW Atlantic (CAFSAC Divs. 5-6) 1986-1987 S. scombrus P. phocoena 29.60 n.a. 92 Bay of Fundy, NW Atlantic 1985-1987 35.3 228M 26.00 43.00 25 NW Atlantic (CAFSAC Divs. 5-6) 1986-1987 

Sebastes sp. P. groenlandica 11.40 257 78 Gulf of St. Lawrence, NW Atlantic 1983 31.32 1160 10.00 46.00 99 Gulf of St. Lawrence 1983 

T. atun A. p. pusillus 80.00 10 36 Benguela, South Africa 1974-1985 91.2 n.a.* 75.00 95.00 32 Benguela, Namibian coast (ICSEAF Divs. 1.3-1.5) 1978-1986 

T. capensis A. p. pusillus 10.80 66 36 Benguela, South Africa 1974-1985 9.5 n.a.* 4.00 16.00 103 Benguela, west coast 1986 

T. capensis A. p. pusillus 16.30 449 36 Benguela, South Africa 1974-1985 9.5 n.a.* 4.00 16.00 103 Benguela, west coast 1986 

T. capensis A. p. pusillus 14.80 3542 36 Benguela, South Africa 1974-1985 9.5 n.a.* 4.00 16.00 103 Benguela, west coast 1986 

T. chalcogramma B. acutorostrata 14.50 121 11.80 17.50 49 Bering Sea 1975 31.40 n.a.* 15.00 62.00 10 Bering Sea, eastern 1975 

T. chalcogramma C. ursinus 30.40 n.a.* 69 Bering Sea 1981-1982 39.80 n.a.* 20.00 62.00 10 Bering Sea, eastern 1981-1982 

T. chalcogramma C. ursinus 19.30 n.a.* 10.00 35.00 71 Bering Sea 1974 32.10 n.a.* 11.00 63.00 10 Bering Sea, eastern 1974 

T. chalcogramma C. ursinus 32.97 39 6.00 41.00 108 Bering Sea 1981 39.08 n.a.* 20.00 61.00 10 Bering Sea, eastern 1981 

T. chalcogramma C. ursinus 6.70 1191 3.00 42.00 108 Bering Sea 1982 40.55 n.a.* 22.00 62.00 10 Bering Sea, eastern 1982 

T. chalcogramma C. ursinus 13.03 1428 4.00 47.00 108 Bering Sea 1985 42.43 n.a.* 17.00 63.00 10 Bering Sea, eastern 1985 

T. chalcogramma E. barbatus 11.80 56 6.90 14.30 7 Bering Sea 1981 39.20 n.a.* 20.00 61.00 10 Bering Sea, eastern 1981 

T. chalcogramma E. jubatus 29.80 2030 5.60 62.90 89 Gulf of Alaska 1975-1978 41.2 n.a.* 23.40 62.30 6 Gulf of Alaksa 1977-1978 

T. chalcogramma E. jubatus 46.90 280 18.40 61.40 49 Pribilof Islands, Bering Sea 1976, 1979 33.10 n.a.* 17.00 62.00 10 Bering Sea, eastern 1976 

T. chalcogramma E. jubatus 25.20 1135 8.30 64.20 49 Bering Sea 1981 39.20 n.a.* 20.00 61.00 10 Bering Sea, eastern 1981 

T. chalcogramma E. jubatus 21.8 n.a. 10.30 51.60 69 St. Matthew Island, Bering Sea 1985 42.60 n.a.* 17.00 63.00 10 Bering Sea, eastern 1985 

T. chalcogramma E. jubatus 33.5 n.a. 20.80 44.50 69 St. Paul Island, Bering Sea 1985 42.60 n.a.* 17.00 63.00 10 Bering Sea, eastern 1985 

T. chalcogramma E. jubatus 29.9 n.a. 1.70 42.70 69 E Aleutian Islands, Gulf of Alaska 1981-1982 39.80 n.a.* 20.00 62.00 10 Bering Sea, eastern 1981-1982 T. chalcogramma E. jubatus 25.5 80 4.80 55.70 23 Kodiak Island, Gulf of Alaska 1985-1986 47.6 n.a.* 21.00 70.00 80 Gulf of Alaska, Kodiak Island region 1985-1986 

T. chalcogramma E. jubatus 42.40 909 10.00 78.10 110 SE Alaska 1994-1999 52.2 2103 110 Dixon Entrance, BC 1993-1999 

T. chalcogramma E. jubatus 39.30 666 3.70 70.80 117 Aleutian Is. (western stock) 1998-2000 49.9 92133 20.00 74.00 117 1998-2000 

T. chalcogramma P. fasciata 10.26 447 6.00 20.00 48 Bering Sea 1976-1976 34.17 n.a.* 11.00 68.00 10 Bering Sea, eastern 1976-1979 

T. chalcogramma P. fasciata 11.20 468 6.50 34.40 49 Bering Sea 1976-1978 33.90 n.a.* 12.00 68.00 10 Bering Sea, eastern 1976-1978 T. chalcogramma P. largha 10.90 21 8.00 15.00 22 Bering Sea 1976-1978 33.90 n.a.* 12.00 68.00 10 Bering Sea, eastern 1976-1978 T. chalcogramma P. largha 10.90 21 8.00 15.00 49 Bering Sea 1978 36.80 n.a.* 14.00 68.00 10 Bering Sea, eastern 1978 

T. chalcogramma P. vitulina 19.20 2180 4.20 53.20 89 Gulf of Alaska 1975-1978 41.2 n.a.* 23.40 62.30 6 Gulf of Alaksa 1977-1978 

T. chalcogramma P. vitulina 31.80 23 10.30 56.30 49 Pribilof Islands, Bering Sea 1979 34.90 n.a.* 11.00 64.00 10 Bering Sea, eastern 1979 

T. chalcogramma P. vitulina 10.60 12 8.20 12.60 49 Bering Sea 1981 39.2 n.a.* 20.00 61.00 10 Bering Sea, eastern 1981 

T. t. capensis A. p. pusillus 27.15 56 24 Benguela, South Africa 1976-1990 16 n.a.* 4.00 35.00 103 Benguela, east and west coasts 1986 T. t. capensis C. heavisidii 17.10 23 10.70 22.20 103 Benguela, South Africa 1969-1990 16 n.a.* 4.00 35.00 103 Benguela, east and west coasts 1986 

T. t. capensis D. delphis 16.20 134 4.60 34.30 103 Benguela, South Africa 1969-1990 16 n.a.* 4.00 35.00 103 Benguela, east and west coasts 1986 

T. t. capensis L. obscurus 9.90 723 4.20 24.90 103 Benguela, South Africa 1969-1990 16 n.a.* 4.00 35.00 103 Benguela, east and west coasts 1986 

Disko Bay, Greenland (inshore longline fishery) 

Aleutian Islands; trawl fishery 

n.a.* = sample size not available, but probably large 
Appendix 6. Pairings of food habits and survey data. See Appendix 3 for full scientific names of prey species. See Appendix 1 for common and full scientific names of predator species. See Appendix 7 for key to numbered citations. 

270

Food Food Food 

Prey Predator habits Range, Range, habits habits Survey Range, Range, Survey 

species species mean N low high citation Food habits locality dates mean N low high citations Locality of survey Survey dates 

A. flavimanus P. vitulina 9.20 1021 3.50 20.20 111 San Francisco Bay, NE Pacific 1991-1992 9.78 219 2 18 111 San Francisco Bay, NE Pacific 1980-1992 

B. saida D. leucas 14.54 125 9.00 21.50 45 Baffin Bay, NW Atlantic 1978-1979 9.72 172 5 22.5 20 Pond Inlet, NW Atlantic 1978-1979 

B. saida E. barbatus 13.40 272 6.90 23.60 42 Baffin Bay, NW Atlantic 1978-1979 9.72 172 5 22.5 20 Pond Inlet, NW Atlantic 1978-1979 

B. saida M. monoceros 11.30 607 4.80 21.50 43 Baffin Bay, NW Atlantic 1978-1979 9.72 172 5 22.5 20 Pond Inlet, NW Atlantic 1978-1979 

B. saida P. fasciata 15.87 92 12.00 21.00 48 Bering Sea 1976-1979 13.81 119 7 22 48 Bering Sea, northern 

B. saida P. groenlandica 13.40 648 4.80 23.60 45 Baffin Bay, NW Atlantic 1978-1979 9.72 172 5 22.5 20 Pond Inlet, NW Atlantic 1978-1979 

B. saida P. hispida 10.50 4047 4.80 23.60 44 Baffin Bay, NW Atlantic 1978-1979 9.72 172 5 22.5 20 Pond Inlet, NW Atlantic 1978-1979 

B. saida P. hispida 10.90 4847 4.90 16.10 115 Svalbard, Greenland Sea 1985-1987 8.00 115 Kongsfjorden 1988-1989 

B. saida P. largha 14.90 326 7.60 20.50 22 Bering Sea 1976-1978 14.20 121 7.00 22 22 Bering Sea 1976-1978 

C. capensis A. p. pusillus 15.00 10 36 Benguela, South Africa 1974-1985 39.80 125 28 56 64 Benguela, South Africa (ICSEAF Div. 1.3) 1988 

C. gunnari A. gazella 21.09 530 12.00 34.00 54 Heard Island, Indian Ocean 1987-1988 21.73 3762 5 38 39 Kerguelen Island, inner shelf 1989-1992 

C. gunnari A. gazella 22.42 877 18.25 27.75 55 Heard Island, Indian Ocean 1993-1993 22.23 827 5 38 39 Kerguelen Island, inner shelf 1992 

C. gunnari A. gazella 20.80 1263 18.75 23.75 55 Heard Island, Indian Ocean 1993-1993 22.23 827 5 38 39 Kerguelen Island, inner shelf 1992 

C. gunnari A. gazella 23.80 201 10.00 46.00 82 South Georgia Island, S Atlantic 1983 30.99 666 11 46 62 Elephant Island 1983 

C. gunnari A. gazella 27.80 580 14.00 48.00 94 South Georgia Island, S Atlantic 1991-1994 28.39 1665 10 49 62 Elephant Island 1996 

C. gunnari A. gazella 22.74 1920 12.00 46.00 93 South Georgia Island, S Atlantic 1992-1993 20.65 10 50 86 South Georgia Island, S Atlantic 1991 

C. harengus P. phocoena 25.40 507 18.00 33.00 52 Gulf of Maine, NW Atlantic 1989-1994 25.56 33352 8 35 83 Gulf of Maine Fall 1989-1994 

C. opilio E. barbatus 5.70 336 3.20 12.30 7 Bering Sea 1981 6.25 12.5 132.5 61 Bering Sea, eastern 1975 

E. encrasicolus A. p. pusillus 12.70 1799 36 Benguela, South Africa 1974-1985 6.60 1671 2 9.5 35 Benguela, Orange River to Olifants River 1983 

E. encrasicolus A. p. pusillus 9.90 2911 36 Benguela, South Africa 1974-1985 6.60 1671 2 9.5 35 Benguela, Orange River to Olifants River 1983 

E. encrasicolus A. p. pusillus 9.40 15871 36 Benguela, South Africa 1974-1985 6.60 1671 2 9.5 35 Benguela, Orange River to Olifants River 1983 

E. encrasicolus D. delphis 11.20 413 5.30 18.10 103 Benguela, South Africa 1969-1990 6.60 1671 2 9.5 35 Benguela, Orange River to Olifants River 1983 E. encrasicolus L. obscurus 9.10 188 4.40 15.80 103 Benguela, South Africa 1969-1990 6.60 1671 2 9.5 35 Benguela, Orange River to Olifants River 1983 

E. gracilis P. largha 16.70 131 6.20 25.10 22 Bering Sea 1976-1978 19.56 903 6.00 35 116 Norton Sound and Chukchi Seas 1976, 1979 

E. mordax P. vitulina 12.90 81 4.00 15.00 111 San Francisco Bay, NE Pacific 1991-1992 4.9 13000 3 14 111 San Francisco Bay, NE Pacific 1980-1992 E. superba A. gazella 5.13 301 3.90 6.10 33 South Georgia Island, S Atlantic 1972-1977 3.55 1471 1.8 5.2 60 South Georgia to South Shetlands 1976 E. superba A. gazella 4.45 308 3.60 6.20 34 South Georgia Island, S Atlantic 1994 4.32 2.2 6.1 95 South Georgia Island, S Atlantic 1994 

E. superba A. gazella 4.26 n.a.* 72 South Georgia Island, S Atlantic 1994-1996 3.76 n.a.* 2.2 6.1 95 South Georgia Island, S Atlantic 1994, 1996 

E. superba A. gazella 4.42 2206 2.40 5.80 93 South Georgia Island, S Atlantic 1992-1993 4.32 2.2 6.1 95 South Georgia Island, S Atlantic 1994 

E. superba A. gazella 4.67 5585 3.20 6.40 94 South Georgia Island, S Atlantic 1991-1994 4.32 2.2 6.1 95 South Georgia Island, S Atlantic 1994 

E. superba A. gazella 4.63 n.a.* 3.20 6.10 95 South Georgia Island, S Atlantic 1994-1999 4.05 n.a.* 2.1 6.3 95 South Georgia Island, S Atlantic 1994-1999 

E. superba A. gazella 5.54 324 3.80 6.60 96 South Georgia Island, S Atlantic 1986 5.35 3465 2.8 6.6 96 South Georgia Island, S Atlantic 1986 

E. superba A. gazella 5.14 388 3.50 6.50 37 South Georgia Island, S Atlantic 1982-1983 3.29 7666 1.9 5.25 104 Bransfield Strait, South Georgia 1983 E. whiteheadi A. p. pusillus 14.70 17 36 Benguela, South Africa 1974-1985 13.44 4 21 112 Benguela, west coast of South Africa 1984-1989 

E. whiteheadi A. p. pusillus 13.20 32 36 Benguela, South Africa 1974-1985 13.44 4 21 112 Benguela, west coast of South Africa 1984-1989 E. whiteheadi D. delphis 16.20 171 8.50 23.60 103 Benguela, South Africa 1969-1990 13.44 4 21 112 Benguela, west coast of South Africa 1984-1989 

E.s gracilis D. leucas 11.98 530 5.00 30.00 102 Bering Sea 1978 19.56 903 6.00 35 116 Norton Sound and Chukchi Seas 1976, 1979 G. capensis A. p. pusillus 45.90 25 36 Benguela, South Africa 1974-1985 86.90 810 26.5 134.5 59 Benguela, Cape Town to Luderitz 1986-1988 

G. gibberifrons A. gazella 31.60 55 5.90 54.00 82 South Georgia Island, S Atlantic 1983 29.53 12151 13 45 62 Elephant Island 1983, 1985 

G. gibberifrons A. gazella 42.10 95 21.00 59.00 93 South Georgia Island, S Atlantic 1992-1993 33.87 3256 18 47 62 Elephant Island 1996 G. lineatus P. vitulina 22.20 40 13.00 29.00 111 San Francisco Bay, NE Pacific 1991-1992 13.5 4171 1 30 111 San Francisco Bay, NE Pacific 1980-1992 

G. morhua H. grypus 18.80 81 0.00 35.00 17 Scotian Shelf, NW Atlantic 1991-1993 53.87 235 14 118 83 NW Atlantic (NAFO Divs. 464, 465) 1983-1987 

G. morhua H. grypus 25.80 115 10.00 55.00 19 Scotian Shelf, NW Atlantic 1988-1990 50.88 231 4 124 83 NW Atlantic (NAFO Divs. 464, 465) 1983-1987 

G. morhua P. groenlandica 29.55 133 5.00 65.00 81 Ullsford, Norway 1986, 1988 45.00 527 25 85 81 Atlantic, NE; Ullsford, North Norway 1986, 1988 

G. morhua P. phocoena 15.90 92 Bay of Fundy, NW Atlantic 1985-1987 59.34 148 22 110 83 Bay of Fundy 1985-1987 

G. morhua P. phocoena 28.10 30 16 Kattegat Sea and Skagerrak Sea 1989-1996 27.89 13 48 109 Skagerrak Sea 2000-2001 

G. morhua P. vitulina 35.20 33 5.00 50.00 18 Bay of Fundy, NW Atlantic 1988-1992 51.69 94 5 116 83 Bay of Fundy 1988-1991 

G. morhua P. vitulina 19.00 46 10.00 35.00 18 Nova Scotia, Canada 1988-1992 53.18 366 4 124 83 NW Atlantic (NAFO Divs. 464, 465) 1983-1987 G. setosus P. hispida 1.53 35 1.20 2.00 115 Svalbard, Greenland Sea 1985-1987 2.50 29 2.2 3.4 115 Kongsfjorden 1988-1989 

H. dactylopterus A. p. pusillus 23.20 13 36 Benguela, South Africa 1974-1985 12.96 1722 5 35 1 Benguela (ICSEAF Divs. 1.4 and 1.5) 1987 

H. dactylopterus A. p. pusillus 20.00 145 36 Benguela, South Africa 1974-1985 12.96 1722 5 35 1 Benguela (ICSEAF Divs. 1.4 and 1.5) 1987 

H. platessoides H. grypus 24.30 77 11.10 46.80 13 Gulf of St. Lawrence, NW Atlantic 1982-1987 27.40 5213 9 57 11 NW Atlantic (CAFSCA Div. 4T, 195-79) 1975-1979 

H. platessoides H. grypus 25.91 23 15.00 40.00 17 Scotian Shelf, NW Atlantic 1991-1993 26.55 3264 10 62 114 Scotian Shelf, NW Atlantic 1998 

I. illecebrosus H. grypus 18.50 78 10.00 25.00 19 Scotian Shelf, NW Atlantic 1988-1990 29.00 3124 13 29 83 Scotian Shelf, NW Atlantic Fall 1988-1990 

I. illecebrosus P. vitulina 17.60 139 5.00 20.00 18 Bay of Fundy, NW Atlantic 1988-1992 19.57 3034 13 29 83 Bay of Fundy, fall 1988 and 1989 Fall, 1988-1989 

I. illecebrosus P. vitulina 19.10 51 15.00 25.00 18 Nova Scotia, Canada 1988-1992 17.93 3239 9 29 83 Scotian Shelf, NW Atlantic Fall 1988-1992 

J. lalandii A. p. pusillus 5.20 38 1.00 7.00 36 Benguela, South Africa 1984 7.16 8170 4.5 11.5 100 Benguela, Olifants River to St. Helena Bay Oct 1996, Mar 1999 

J. lalandii A. p. pusillus 4.18 89 1.00 7.00 36 Benguela, South Africa 1981 7.16 8170 4.5 11.5 100 Benguela, Olifants River to St. Helena Bay Oct 1996, Mar 1999 

J. lalandii A. p. pusillus 2.50 253 1.00 6.00 36 Benguela, South Africa 1985 7.16 8170 4.5 11.5 100 Benguela, Olifants River to St. Helena Bay Oct 1996, Mar 1999 

L. armatus P. vitulina 14.03 72 5.00 25.00 111 San Francisco Bay, NE Pacific 1991-1992 11.64 704 2 20 111 San Francisco Bay, NE Pacific 1980-1992 

L. ferruginea H. grypus 20.00 14 19 Scotian Shelf, NW Atlantic 1988-1990 30.33 85 18 44 83 Scotian Shelf, NW Atlantic 1988, 1990 

L. hectoris A. p. pusillus 4.80 595 36 Benguela, South Africa 1974-1985 3.87 144 0.8 6.8 91 Benguela 1982-1983 

L. hectoris A. p. pusillus 5.00 1849 36 Benguela, South Africa 1974-1985 3.87 144 0.8 6.8 91 Benguela 1982-1983 

L. larseni A. gazella 14.70 91 7.80 22.00 82 South Georgia Island, S Atlantic 1983 10.49 11494 3 23 47 South Georgia Island, S Atlantic 1984 

L. larseni A. gazella 18.52 380 4.00 28.00 93 South Georgia Island, S Atlantic 1992-1993 14.93 2525 6 22 62 Elephant Island 1996 

L. larseni A. gazella 17.21 482 10.00 25.00 94 South Georgia Island, S Atlantic 1991-1994 14.93 2525 6 22 62 Elephant Island 1996 L. limanda P. vitulina 8.13 219 < 7.0 20.00 12 Waddensea, North Sea 1975-1981 17.82 56416 9.5 31.5 70 Deutsche Bucht, Waddensea 1983-1986 

n.a.* = sample size not available, but probably large 
Appendix 6. Pairings of food habits and survey data. See Appendix 3 for full scientific names of prey species. See Appendix 1 for common and full scientific names of predator species. See Appendix 7 for key to numbered citations. 

271

Food Food Food 

Prey Predator habits Range, Range, habits habits Survey Range, Range, Survey 

species species mean N low high citation Food habits locality dates mean N low high citations Locality of survey Survey dates 

L. pealeii G. melas 19.60 578 3.70 43.60 50 NW Atlantic 1989-1991 6.79 94000 1 38 83 Mid-Atlantic Bight, USA 1989-1991 

L. pealeii G. melas 16.07 145 5.00 42.00 51 Mid-Atlantic Bight, USA 1973-1993 6.90 ##### 1 44 83 Mid-Atlantic Bight, USA 1980-1993 

L. pealeii Globicephala sp. 13.40 68 85 Mid-Atlantic Bight, USA 1989 8.22 36000 1 33 83 Mid-Atlantic Bight, USA 1989 

L. v. reynaudii A. p. pusillus 16.06 183 24 Benguela, South Africa 1976-1990 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii A. p. pusillus 11.50 51 7.10 18.90 68 Benguela, South Africa 1974-1985 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii A. p. pusillus 15.60 596 4.10 32.40 68 Benguela, South Africa 1974-1985 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii C. heavisidii 17.40 11 9.30 24.40 103 Benguela, South Africa 1969-1990 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii D. delphis 17.30 322 2.50 34.50 103 Benguela, South Africa 1969-1990 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii G. griseus 13.00 67 2.10 32.90 103 Benguela, South Africa 1975-1990 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii K. breviceps 11.60 21 5.30 20.90 103 Benguela, South Africa 1975-1990 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii K. sima 7.10 11 5.30 8.90 103 Benguela, South Africa 1975-1988 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 L. v. reynaudii L. obscurus 12.90 23 1.70 27.00 103 Benguela, South Africa 1969-1990 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii S. coeruleoalba 9.30 183 2.40 22.30 103 Benguela, South Africa 1975-1989 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

L. v. reynaudii T. truncatus 22.70 12 20.20 26.10 103 Benguela, South Africa 1975-1989 16.13 11357 4 29 8 Benguela, Orange River to Cape Agulhas 1984-1987 

M. bilinearis H. grypus 25.67 30 15.00 35.00 17 Scotian Shelf, NW Atlantic 1991-1993 20.34 8968 3 47 83 Scotian Shelf, NW Atlantic 1991-1993 M. bilinearis H. grypus 25.40 43 10.00 45.00 19 Scotian Shelf, NW Atlantic 1988-1990 20.47 5410 4 49 83 Scotian Shelf, NW Atlantic 1988-1990 M. bilinearis P. phocoena 16.40 1605 3.00 38.00 52 Gulf of Maine, NW Atlantic 1989-1994 21.52 47949 2 54 83 Gulf of Maine Fall 1989-1994 

M. bilinearis P. phocoena 19.60 173 92 Bay of Fundy, NW Atlantic 1985-1987 20.35 723 4 49 83 Bay of Funday, fall 1985-86 Fall, 1985-1986 

M. capensis A. p. pusillus 28.19 64 24 Benguela, South Africa 1976-1990 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

M. merlangus P. phocoena 18.70 65 16 Kattegat Sea and Skagerrak Sea 1989-1996 18.49 11 31 109 Skagerrak Sea 2000-2001 

M. merlangus P. phocoena 7.50 449 16 Kattegat Sea and Skagerrak Sea 1989-1996 18.49 11 31 109 Skagerrak Sea 2000-2001 

M. muelleri A. p. pusillus 3.40 1257 36 Benguela, South Africa 1974-1985 3.65 944 1 5.2 91 Benguela 1982-1983 

M. scorpius P. hispida 11.14 35 8.00 18.00 115 Svalbard, Greenland Sea 1985-1987 8.90 130 2 28 115 Kongsfjorden 1988-1989 Merluccius spp. A. p. pusillus 29.50 116 36 Benguela, South Africa 1974-1985 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. A. p. pusillus 17.40 350 36 Benguela, South Africa 1974-1985 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. A. p. pusillus 21.30 1177 36 Benguela, South Africa 1974-1985 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. C. heavisidii 19.50 160 4.90 28.60 103 Benguela, South Africa 1969-1990 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. D. delphis 18.10 203 5.10 39.30 103 Benguela, South Africa 1969-1990 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. K. breviceps 21.30 11 16.20 25.50 103 Benguela, South Africa 1975-1990 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. K. sima 21.60 20 16.60 26.80 103 Benguela, South Africa 1975-1988 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. L. obscurus 16.50 216 4.80 42.10 103 Benguela, South Africa 1969-1990 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. S. coeruleoalba 14.90 12 7.80 17.20 103 Benguela, South Africa 1975-1989 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

Merluccius spp. T. truncatus 28.30 15 17.10 42.30 103 Benguela, South Africa 1975-1989 39.11 12780 19 71 77 Benguela (ICSEAF Divs. 1.3, 1.4, 1.5) 1987 

P. antarcticum L. weddelii 16.24 80 12.00 21.00 90 Weddell Sea, Antarctica 1998 15.47 466 3 24 90 Drescher Inlet, Weddell Sea 1988 

P. borealis P. hispida 7.47 51 4.80 9.20 115 Svalbard, Greenland Sea 1985-1987 7.00 20 2.4 7.6 115 Kongsfjorden 1988-1989 

P. choriodon A. gazella 8.56 752 7.00 11.00 94 South Georgia Island, S Atlantic 1991-1994 5.87 2378 5 10 47 South Georgia Island, S Atlantic 1986-1987 

P. georgianus A. gazella 28.80 26 24.10 34.80 82 South Georgia Island, S Atlantic 1983 15.27 706 9 51 47 South Georgia Island, S Atlantic 1984 

P. laniarius A. p. pusillus 28.36 15 24 Benguela, South Africa 1976-1990 22.47 2762 3 37 14 Benguela, Algulhas Bank May-Oct 1994 

P. laniarius A. p. pusillus 20.00 15 36 Benguela, South Africa 1974-1985 22.47 2762 3 37 14 Benguela, Algulhas Bank May-Oct 1994 P. notatus P. vitulina 21.80 73 7.00 39.00 111 San Francisco Bay, NE Pacific 1991-1992 4.79 2054 1 32 111 San Francisco Bay, NE Pacific 1980-1992 

P. platessa H. grypus 24.07 51 7.50 77.50 56 Orkney Island, NE Atlantic 1985 22.13 4542 7 46 97 North Sea Winter 1985-1986 P. platessa H. grypus 31.99 247 2.50 77.50 56 Orkney Island, NE Atlantic 1985 22.13 4542 7 46 97 North Sea Winter 1985-1986 P. platessa P. vitulina 10.00 199 12 Waddensea, North Sea 1975-1981 22.13 4542 7 46 97 North Sea Winter 1985-1986 

P. triacanthus P. phocoena 9.70 38 52 Gulf of Maine, NW Atlantic 1989-1994 11.07 19225 2 20 83 Gulf of Maine Fall 1989-1994 P. virens P. phocoena 19.50 76 52 Gulf of Maine, NW Atlantic 1989-1994 33.45 875 8 112 83 Gulf of Maine Fall 1989-1994 

R. hippoglossoides M. monoceros 35.70 140 13.80 54.70 65 Disko Bay, Greenland 2002-2003 44.27 2355 13 99 66 Central Baffin Bay and No. Davis Strait 

S. scombrus G. melas 35.80 45 26.50 42.10 50 NW Atlantic 1989-1991 25.60 11490 5 44 83 Mid-Atlantic Bight, USA 1989-1991 

S. scombrus Globicephala sp. 36.32 22 85 Mid-Atlantic Bight, USA 1989 21.04 1462 14 43 83 Mid-Atlantic Bight, USA 1989 

S. scombrus P. phocoena 22.40 15 52 Gulf of Maine, NW Atlantic 1989-1994 27.31 1353 17 43 83 Gulf of Maine Fall 1989-1994 

S. septemcarinata P. hispida 5.02 19 3.60 5.60 115 Svalbard, Greenland Sea 1985-1987 5.20 29 2 6.4 115 Kongsfjorden 1988-1989 

T. angolensis A. p. pusillus 14.60 22 3.40 29.10 68 Benguela, South Africa 1974-1985 27.20 417 15 45 1 Benguela (ICSEAF Divs. 1.4 and 1.5) 1987 

T. atun A. p. pusillus 80.00 10 36 Benguela, South Africa 1974-1985 92.90 51 70 114 64 Benguela, South Africa (ICSEAF Div. 1.3) 

T. capensis A. p. pusillus 10.80 66 36 Benguela, South Africa 1974-1985 28.90 912 20 36 64 Benguela, South Africa (ICSEAF Div. 1.3) 

T. capensis A. p. pusillus 16.30 449 36 Benguela, South Africa 1974-1985 28.90 912 20 36 64 Benguela, South Africa (ICSEAF Div. 1.3) 

T. capensis A. p. pusillus 14.80 3542 36 Benguela, South Africa 1974-1985 28.90 912 20 36 64 Benguela, South Africa (ICSEAF Div. 1.3) 

T. chalcogramma B. acutorostrata 14.50 121 11.80 17.50 49 Bering Sea 1975 25.90 10 58 10 Bering Sea, eastern 1975 

T. chalcogramma C. ursinus 19.30 n.a.* 10.00 35.00 71 Bering Sea 1974 25.90 10 58 10 Bering Sea, eastern 1975 

T. chalcogramma C. ursinus 30.40 69 Bering Sea 1981-1982 38.30 9 68 69 Bering Sea, southeast 1981-1982 

T. chalcogramma C. ursinus 32.97 39 6.00 41.00 108 Bering Sea 1981 38.90 108 Bering Sea, eastern 1981 

T. chalcogramma C. ursinus 6.70 1191 3.00 42.00 108 Bering Sea 1982 39.70 108 Bering Sea, eastern 1982 

T. chalcogramma C. ursinus 13.03 1428 4.00 47.00 108 Bering Sea 1985 44.00 108 Bering Sea, eastern 1985 

T. chalcogramma E. barbatus 11.80 56 6.90 14.30 7 Bering Sea 1981 33.40 9 68 10 Bering Sea, eastern 1981 

T. chalcogramma E. jubatus 46.90 280 18.40 61.40 49 Pribilof Islands, Bering Sea 1976, 1979 26.00 6.00 63 10 Bering Sea, eastern 1975. 1979-1980 

T. chalcogramma E. jubatus 25.20 1135 8.30 64.20 49 Bering Sea 1981 33.40 9 68 10 Bering Sea, eastern 1981 

T. chalcogramma E. jubatus 33.5 20.8 44.5 69 St. Paul Island, Bering Sea 1985 35.00 9 67 10 Bering Sea, eastern 1985 

n.a.* = sample size not available, but probably large 
Appendix 6. Pairings of food habits and survey data. See Appendix 3 for full scientific names of prey species. See Appendix 1 for common and full scientific names of predator species. See Appendix 7 for key to numbered citations. 

Food Food Food 

Prey Predator habits Range, Range, habits habits Survey Range, Range, Survey 

species species mean N low high citation Food habits locality dates mean N low high citations Locality of survey Survey dates 

T. chalcogramma E. jubatus 21.8 10.3 51.6 69 St. Matthew Island, Bering Sea 1985 35.00 9 67 10 Bering Sea, eastern 1985 

T. chalcogramma E. jubatus 29.80 2030 5.60 62.90 89 Gulf of Alaska 1975-1978 31.50 9 58 21 Gulf of Alaska T. chalcogramma E. jubatus 29.9 1.7 42.7 69 E Aleutian Islands, Gulf of Alaska 1981-1982 25.50 69 Gulf of Alaska, eastern Aleutians 1981-1982 

T. chalcogramma E. jubatus 25.5 80 4.8 55.7 23 Kodiak Island, Gulf of Alaska 1985-1986 25.14 10 57 80 Gulf of Alaska, Shelikof Strait 1985-1986 T. chalcogramma P. dalli 33.64 639 15.08 52.48 84 Sea of Okhotsk & Sea of Japan 1988-1996 43.15 894 26 58 73,74 Sea of Okhotsk 1990-1991, 1996 

T. chalcogramma P. dalli 22.61 135 6.50 60.50 113 Sea of Okhotsk 1988-1989 43.15 894 26 58 73,74 Sea of Okhotsk 1990-1991, 1996 

T. chalcogramma P. fasciata 11.20 468 6.50 34.40 49 Bering Sea 1976-1978 26.00 6.00 63 10 Bering Sea, eastern 1975. 1979-1980 

T. chalcogramma P. fasciata 10.26 447 6.00 20.00 48 Bering Sea 1976-1976 11.16 899 8 23 48 Bering Sea, northern 

T. chalcogramma P. largha 10.90 21 8.00 15.00 22 Bering Sea 1976-1978 26.00 6.00 63 10 Bering Sea, eastern 1975. 1979-1980 

T. chalcogramma P. largha 10.90 21 8.00 15.00 49 Bering Sea 1978 26.00 6.00 63 10 Bering Sea, eastern 1975. 1979-1980 

T. chalcogramma P. vitulina 10.60 12 8.20 12.60 49 Bering Sea 1981 33.40 9 68 10 Bering Sea, eastern 1981 

T. chalcogramma P. vitulina 31.80 23 10.30 56.30 49 Pribilof Islands, Bering Sea 1979 24.90 6 61 10 Bering Sea, eastern 1979 T. chalcogramma P. vitulina 19.20 2180 4.20 53.20 89 Gulf of Alaska 1975-1978 31.50 9 58 21 Gulf of Alaska 

T. inermis P. hispida 1.75 189 1.20 2.60 115 Svalbard, Greenland Sea 1985-1987 1.90 23 1.2 2.4 115 Kongsfjorden 1988-1989 

T. libellula P. hispida 1.66 279 1.40 3.40 115 Svalbard, Greenland Sea 1985-1987 2.30 21 0.2 3.8 115 Kongsfjorden 1988-1989 

T. t. capensis A. p. pusillus 27.15 56 24 Benguela, South Africa 1976-1990 28.90 912 20 36 64 Benguela, South Africa (ICSEAF Div. 1.3) 

T. t. capensis C. heavisidii 17.10 23 10.70 22.20 103 Benguela, South Africa 1969-1990 28.90 912 20 36 64 Benguela, South Africa (ICSEAF Div. 1.3) 

T. t. capensis D. delphis 16.20 134 4.60 34.30 103 Benguela, South Africa 1969-1990 28.90 912 20 36 64 Benguela, South Africa (ICSEAF Div. 1.3) 

T. t. capensis L. obscurus 9.90 723 4.20 24.90 103 Benguela, South Africa 1969-1990 28.90 912 20 36 64 Benguela, South Africa (ICSEAF Div. 1.3) Urophycis spp. P. phocoena 15.90 474 52 Gulf of Maine, NW Atlantic 1989-1994 36.62 6120 3 91 83 Gulf of Maine Fall 1989-1994 

272

n.a.* = sample size not available, but probably large 
Appendix 7. Key to numbered citations in Appendices 5 and 6. 

Citation Citation 

number Citation number Citation 

1 Abello et al. 1988 49 Frost and Lowry 1986 

2 Ahrens 1985 50 Gannon et al. 1997a 

3 Alexander and Vromans 1985 51 Gannon et al. 1997b 

4 Alexander and Vromans 1988a 52 Gannon et al. 1998 

5 Alexander and Vromans 1988b 53 Gonzales et al. 1997 

6 Alton and Deriso 1983 54 Green et al. 1989 

7 Antonelis et al. 1994b 55 Green et al. 1997 

8 Augustyn 1991 56 Hammond et al. 1994a 

9 Badenhorst 1985 57 Isarev 1985 

10 Bakkala 1989 58 Jackson et al. 1998 

11 Beacham 1982 59 Japp 1990 

12 Behrends 1982 60 Jazdzewski et al. 1978 

13 Benoit and Bowen 1990 61 Kaimmer et al. 1976 

14 Booth and Buxton 1997 62 Kock 1998 

15 Booth and Hecht 1998 63 Kock and Köster 1989 

16 Borjesson et al. 2003 64 Konchina 1989 

17 Bowen and Harrison 1994 65 Laidre and Heide-Jorgensen 2005 

18 Bowen and Harrison 1996 66 Laidre et al. 2004 

19 Bowen et al. 1993 67 Leos 1998 

20 Bradstreet et al. 1986 68 Lipinski and David 1990 

21 Brown and Rose 1983 69 Lowry et al. 1989 

22 Bukhtiyarov et al. 1984 70 Lozan 1988 

23 Calkins and Goodwin 1988 71 McAlister et al. 1976 

24 Castley et al. 1991 72 McCafferty et al. 1998 

25 

26 

Castonguay and Mercille 1988 

Chadwick and Cairns 1988 

73 

74 

McFarlane et al. 1996 

McFarlane, n.d.† 

27 Chouinard and Metuzals 1985 75 McPhail et al. 2001 

28 Clarke and Goodall 1994 76 Melin 2002 

29 Clay et al. 1985 77 Morales-Nin 1991 

30 Crawford 1980 78 Murie and Lavigne 1991 

31 Crawford et al. 1978 79 Murie and Lavigne 1992 

32 Crawford et al. 1990 80 Nelson and Nunnallee 1987 

33 Croxall and Pilcher 1984 81 Nilssen et al. 1990 

34 Croxall et al. 1999 82 North 1996 

35 Cruickshank et al. 1990 83 O'Brien, n.d.‡ 

36 David 1987 84 Ohizumi et al. 2000 

37 Doidge and Croxall 1985 85 Overholtz and Waring 1991 

38 Duhamel 1989 86 Parkes 1993, via Reid 1995 

39 Duhamel 1995 87 Payne et al. 1985 

40 Evans 1976 88 Payne et al. 1986 

41 Fanning et al. 1988 89 Pitcher 1981 

42 Finley and Evans 1983 90 Plotz et al. 2001 

43 Finley and Gibb 1982 91 Prosch 1991 

44 Finley et al. 1983 92 Recchia and Read 1989 

45 Finley et al. 1990 93 Reid 1995 

46 Fontaine et al. 1994 94 Reid and Arnould 1996 

47 Frolkina et al. 1998 95 Reid and Brierly 2001 

48 Frost and Lowry 1980 96 Reid et al. 1996 

†

 G. A. "Sandy" McFarlane, Fisheries and Oceans Canada, Pacific Region, Nanaimo, British Columbia, Canada, 

pers. commun. Length frequencies of walleye pollock (Theragra chalcogramma) from the Sea of Okhotsk. 

Unpublished data, 23 September 2003. 

‡  L. O'Brien, Northeast Fisheries Science Center, Woods Hole, Massachusetts, U.S.A., pers. commun. Fish length 

frequency data sampled during fishery independent surveys conducted by Northeast Fisheries Science Center.

 Unpublished data, 1 October 2003. 

273
Appendix 7. Key to numbered citations in Appendices 5 and 6. 

Citation 

number 

97 

98 

99 

100 

101 

102 

103 

104 

105 

106 

107 

108 

109 

110 

111 

112 

113 

114 

115 

116 

Citation 

Rijnsdorp 1989

Weslawski et al. 1994 

Wolotira 1985  

Rodhouse et al. 1992

Rubec et al. 1985

Schoeman et al. 2002

Schwartzlose et al. 1999

Seaman et al. 1982

Sekiguchi et al. 1992

Siegel 1986

Siegel et al. 1998

Simonsen and Boje 2003

Sinclair 1992

Sinclair et al. 1994

Svedang 2003

Tollit et al. 2004a 

Torok 1994

Waldron et al. 1991

Walker 1996

Walsh et al. 1992

274
RECENT TECHNICAL MEMORANDUMS
 

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153 FRITZ, L. W., and C. STINCHCOMB. 2005. Aerial, ship, and land-based surveys of Steller sea lions 

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Pacific halibut longline fisheries, 93 p. NTIS No. PB2005-107312. 

151 TESTA, J. W. (editor). 2005. Fur seal investigations, 2002-2003, 72 p. NTIS No. PB2005-104811. 

150 GULLAND, F. M. D., H. PÉREZ-CORTÉS M., J. URBÁN R., L. ROJAS-BRACHO, G. YLITALO, J. WEIR, 

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148 EILER, J. H., T. R. SPENCER, J. J. PELLA, M. M. MASUDA, and R. R. HOLDER. 2004. Distribution and 

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PB2005-100707. 

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DAVIS. 2004. Electronic monitoring of seabird interactions with trawl third-wire cables on trawl vessels ­

a pilot study, 39 p. NTIS No. PB2005-100243. 

146 WING, B. L., and D. R. BARNARD. 2004. A field guide to Alaskan corals, 67 p. NTIS No. PB2005-100429. 

145 RUGH, D. J., B. A. MAHONEY, and B. K. SMITH. 2004. Aerial surveys of beulga whales in Cook Inlet, 

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