INVESTIGATING THE BIASES IN THE USE OF HARD PREY REMAINS TO IDENTIFY DIET COMPOSITION USING ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA) IN CAPTIVE FEEDING TRIALS

The analysis of pinniped scats has been used to quantify their diet, using prey remains to identify species and to estimate the numbers and sizes of prey consumed. There are, however, potential biases involved with scat analysis and, for this method to be used successfully, these biases need to be quantified. Thirty-six Antarctic fur seals ( Arctocephalus gazella ) were fed meals of exclusively either fish, squid, or krill and their scats were collected and analyzed. The major sources of error in the analysis of prey remains from scats were the differential erosion and passage rate of items in relation to their size. However, using simple correction functions, such as those which model otolith erosion, it is possible to reduce these errors. Using plastic beads as dietary markers showed recovery rates were negatively related to their size. Larger squid beaks had lower recovery rates than smaller beaks, but there was no size-related bias in the recovery rates of krill carapaces. Only 33% of the squid beaks and 27% of the otoliths originally fed were recovered from the scats. Recovery rates were greater for squid (77%) and fish (50%) eye lenses and these structures gave a better estimate of numbers eaten. Differences found between experimentally derived and published regression equations (used to calculate prey sizes eaten from prey remains) highlights the need for regression equations based on local prey characteristics, if these are to be used with any success to describe the prey eaten.     

Fish diet of the Antarctic fur seal,Arctocephalus gazella,in summer,at Stranger Point,King George Island,South Shetland Islands

 In February 1992, 34 faecal samples from non-breeding male Antarctic fur seals, Arctocephalus gazella, were collected at Stranger Point, King George Island, South Shetlands. Fish constituted an important part of the diet, occurring in 90% of those scats containing prey remains. From 1162 otoliths found in the faeces, 1086 were identified to at least family level. Myctophids and nototheniids represented together almost 90% of the fish eaten. The dominant species were Gymnoscopelus nicholsi, Pleuragramma antarcticum and Electrona antarctica, contributing 33.3%, 30.8% and 12.0% of the otoliths respectively. The standard length of these three species was estimated from otoliths with little or no signs of erosion. This study showed that fur seals fed mainly on pelagic fish species that are often associated with krill. These findings are corroborated by fur seal diving patterns. Received: 7 October 1994/Accepted: 5 October 1995     

A captive feeding study with the Pacific harbor seal (Phoca vitulina richardii): Implications for scat analysis

Seven prey species ( n _total > 2,700) were fed to seven captive male Pacific harbor seals ( Phoca vitulina richardii ) in 177 experimental meals to quantify biases associated with scat analysis and current consumption models. Hard parts from an individual meal were recovered in an average of 3.8 ± 1.8 scats (range 1–10; mean ± SD). Overall, 57.7 ± 33.2% of otoliths and 89.5 ± 15.5% of squid beaks were recovered. Recovery rates varied, and prey with smaller, fragile otoliths were recovered in lesser quantities than prey with larger, robust otoliths. Recovery rates of all prey except pink salmon were improved by a mean of 31.7% when all diagnostic structures were included in estimates. Estimated recovery of pink salmon was 9.5 times that fed seals based on the all-structure technique. Mean length reduction of recovered otoliths was 20.4 ± 10.1%. Correction factors calculated from average length reduction improved length estimates for all fish species. Grade-specific length correction factors (gLCFs) reduced variability in all of the estimates and significantly improved estimates of prey with highly eroded otoliths including Pacific hake and shortbelly rockfish. The Biomass Reconstruction (BR) model accurately predicted biomass consumption within 4% of known consumption, whereas estimates based on frequency of occurrence were inaccurate.     

DIET OF WEDDELL SEALS LEPTONYCHOTES WEDDELLII AT VESTKAPP, EASTERN WEDDELL SEA (ANTARCTICA), IN RELATION TO LOCAL FOOD SUPPLY

Thirteen Weddell seals ( Leptonychotes weddellii ) were collected at Vestkapp, eastern Weddell Sea coast, in austral spring 1986. All stomachs contained partially digested food. The mean wet weight of stomach contents was 7.5 kg, 3.3% of the. mean body weight of the collected seals. Twelve fish species and three cephalopod species were identified from 372 left otoliths and 25 lower beaks, representing 58.4% of 679 total prey items obtained. Composition by number of total prey was: Chionodraco myersi (15.8%), Trematomus eulepidotus (10.0%), Pagetopsis maculatus (9.7%), Racovitzia glacialis (9.6%) and Cryodraco antarcticus (4.1%). Otoliths of the seven other fish species and beaks of the three cephalopod species together represented 9.1% of total prey numbers. The pooled wet weights calculated from 13 prey species (regressions for two octopod species were not available) amounted to 43.5 kg food mass and represented 44.7% of the combined food mass in all stomachs. Composition by mass was: C. myersi (44.5%), T. eulepidotus (19.8%), squid Psychroteuthis glacialis (8.5%), P. maculates (7.9%), C. antarcticus (7.1%) and R. glacialis (6.2%). The remaining 7 fish species together represented 5.8% by mass. Temporal variation in food availability was apparent. Midwater fish Pleuragramma antarcticum was the staple food of Weddell seals from the same area during the 1985 summer, whereas it was absent in the samples taken in spring 1986. Estimates of fish biomass from net hauls demonstrate a highly variable availability of pelagic food resources for top predators in the Vestkapp area.     

Seasonal and regional variations in the feeding habits of the harbour seal, Phoca vitulina, in the Skagerrak and the Kattegat

Studies of the feeding of harbour seals have been carried out at the Tjärnö Marine Biological Laboratory since 1977. The studies are based on fish otoliths found in faeces at seal haulouts. The present paper compares feeding habitats at two different localities. Three families of fish, gadoids, pleuronectids and clupeoids were predominant in the seals' diet at a rocky shore habitat. Pleuronectids made up 75% of the diet at a sandy shore habitat. Temporal variations in feeding habits are also examined. The results indicate that harbour seals are opportunists in their choice of prey species, but some locally abundant species do not appear in their diet.     

Improved estimates of digestion correction factors and passage rates for harbor seal (Phoca vitulina) prey in the northeast Atlantic

Diet composition in pinnipeds is widely estimated using hard prey remains recovered from feces. To estimate the size and number of prey represented in fecal samples accurately, digestion correction factors (DCFs) must be applied to measurements and counts of fish otoliths and cephalopod beaks. In this study, 101 whole prey feeding trials were conducted with six harbor seals (Phoca vitulina) and 18 prey species. We derived species‐ and grade‐specific estimates of digestion coefficients (DCs) and species‐specific recovery rates (RRs) to account for partial and complete digestion, respectively. Greater than 98% of otoliths were passed within three days of consumption. RRs were smallest for Atlantic salmon smolts (RR = 0.306, SE = 0.031) and increasingly larger for sandeels (RR = 0.494, SE = 0.017), flatfish (RR = 0.789, SE = 0.033), and large gadoids (RR = 0.944, SE = 0.034). Species‐specific otolith width DCs were smallest for Trisopterus species (DC = 1.14, SE = 0.015) and increasingly larger for flatfish (DC = 1.27, SE = 0.045), large gadoids (DC = 1.32, SE = 0.067) and sandeels (DC = 1.57, SE = 0.035). RRs were similar to those from gray seals (Halichoerus grypus), but harbor seal species‐ and grade‐specific DCs were generally smaller. Differences in partial and complete digestion rates among prey species and between seal species highlight the importance of applying DCFs when reconstructing diet.     

Diet composition and biomass consumption of harbour seals in Telemark and Aust-Agder,Norwegian Skagerrak

ABSTRACT

To explore ecosystem dynamics and functions it is vital to obtain knowledge on predator–prey relations. Harbour seals are piscivorous predators that can come into conflict with fisheries. Recently, as the Skagerrak and Kattegat population of harbour seals has increased, claims have emerged that seals are depleting coastal cod populations. The diet of harbour seals in Norwegian Skagerrak was investigated based on otolith identification from scats. The overall seal diet included 20 different fish species/groups. The most important prey (combined index Q _i ) were haddock/pollack/saithe (32.7%), genus Trisopterus (Norway pout/poor cod/bib, 12.5%), plaice (12.4%) and herring (10.0%). Plaice also had the largest biomass (24.1%). Gadoids and pleuronectids comprised 88.6% of the diet (combined index Q _i ) and 87.1% in terms of biomass. Cod constituted 0.7% (combined index Q _i ) of the overall diet and 2.3% in terms of biomass. Fish length estimates showed that seals generally prefer small fish below minimum allowed landing size. Estimated total amount of fish consumed was 315 tons per year and was dominated by non-commercial species. Annual cod consumption was an estimated 7.1 tons, representing 5% of annual cod landings, suggesting that competition between local fisheries and harbour seals is limited.     

DIETS OF BAIRD'S BEAKED WHALES, BERARDIUS BAIRDII, IN THE SOUTHERN SEA OF OKHOTSK AND OFF THE PACIFIC COAST OF HONSHU, JAPAN

Stomach contents were analyzed from 127 Baird's beaked whales, Berardius bairdii , taken in coastal waters of Japan. During late July-August of 1985–1987, 1989, and 1991, 107 samples were collected from off the Pacific coast of Honshu. An additional 20 samples were collected from whales taken in the southern Sea of Okhotsk during late August—September of 1988 and 1989. Prey identification using fish otoliths and cephalopod beaks revealed the whales fed primarily on deep-water gadiform fishes and cephalopods in both regions. Prey species diversity and the percentage of cephalopods and fish differed between the two regions. Off the Pacific coast of Honshu the whales fed primarily on benthopelagic fishes (81.8%) and only 18.0% on cephalopods. Eight species of fish representing two families, the codlings (Moridae) and the grenadiers (Macrouridae), collectively made up 81.3% of rhe rotal. Thirty species of cephalopods representing 14 families made up 12.7%. In the southern Sea of Okhotsk, cephalopods accounted for 87.1% of stomach contents. The families Gonatidae and Cranchiidae were the predominant cephalopod prey, accounting for 86.7% of the diet. Gadiform fish accounted for only 12.9% of the diet. Longfin codling, Laemonema longipes , was the dominant fish prey in both regions. Depth distribution of the two commonly consumed fish off the Pacific coast of Honshu indicate the whales in this region fed primarily at depths ranging from 800 to 1,200 m.     

Otolith size and location in digestive tracts of northern fur seals (Callorhinus ursinus): Implications for dietary interpretations

Walleye pollock (Theragra chalcogramma) otoliths (n= 2,706) recovered from stomachs, small intestines, and colons of 43 northern fur seals (Callorhinus ursinus) were evaluated for size and wear by location in the digestive tract. Pollock fork length was regressed on otolith length after correction for erosion, and age was estimated from the calculated body size. Age‐1+ pollock otoliths (≥6.3‐mm length) were concentrated in stomachs while age‐0 otoliths (≤6.2‐mm length) were concentrated in colons. Less than 10% of otoliths were found in the small intestines. Pollock age decreased with progression along seal gastrointestinal tracts. Otolith quality increased along gastrointestinal tracts in numbers ≥20, which was typical of age‐0 otoliths recovered from colons. Otolith distribution by age and quality along gastrointestinal tracts suggests that small (≤12 cm) schooling prey are consumed in large volume and passed as a bolus rapidly through the digestive tract before significant erosion of bony remains occurs; while larger prey are eaten in smaller volume and subjected to otolith erosion due to longer retention in the stomach. Our results illustrate the importance of multiple sampling strategies to comprehensively represent prey size in pinniped diet.     

Response of

The fish prey consumed by New Zealand fur seals (Arctocephalus forsteri) was investigated by analysis of faeces collected between February and August at sites on the east and west coasts of South Island, New Zealand. Twelve species were identified from otoliths recovered from faeces. Lanternfish (Symbolophorus sp. and Lampanyctodes hectoris), the most frequent fish prey, comprised 79% of all otoliths, followed by anchovy (Engraulis australis) at 12%, ahuru (Auchenoceros punctatus) with 3.9%, and hoki (Macruronus novaezelandiae) at 3.7%. Of these species only hoki is commercially important. Regional and seasonal differences in the proportions of species were evident, and the results are compared with those from previous studies.     

Morphology and taxonomic significance of the otoliths of some bathypelagic Anguilloidei and Saccopharyngoidei from the Sargasso Sea

The sagittal otoliths of seven anguilliform species belonging to the families Nemichthyidae, Serrivomeridae and Eurypharyngidae are described and illustrated, and a key for their identification is provided. Although the leptocephali of the various fish species treated here have similar otoliths, some species develop specific otolith characteristics during the adult stage. The shape as well as the ratio of fish length to otolith length ratio in adults may serve as a taxonomic aid. Since the sagittae of these fishes can be used for specific identification, the digested remains of prey in the stomachs and guts of predators can be identified.     

东太平洋公海茎柔鱼饵料中鱼类和头足类组成初步分析

茎柔鱼(Dosidicus gigas)广泛分布于东太平洋海域,在海洋生态系统中扮演着重要角色。本研究根据2019年6—12月我国鱿钓生产船在东太平洋公海采集的62尾茎柔鱼样本,对其胃含物残留的4131个耳石和75个角质颚样本进行鉴定分析。结果表明: 茎柔鱼主要摄食鱼类10种、头足类4种。从摄食饵料的出现频率和数量百分比来看,主要摄食的鱼类饵料为荧串光鱼、朗明灯鱼和墨西哥尾灯鱼,荧串光鱼在不同胴长组的茎柔鱼胃内均有出现,是茎柔鱼最重要的饵料;摄食的头足类主要有茎柔鱼、鸢乌贼和爪乌贼。随着胴长的增大,茎柔鱼摄食的饵料种类不断增加,头足类和灯笼鱼饵料比例增加,且摄食的饵料尺寸增大,摄食营养结构发生改变,摄食营养水平随之提高。研究结果可为评估不同饵料生物在茎柔鱼食性转变过程中的食物贡献提供基础资料。     

The cephalopod prey of the Weddell seal, <Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>, a biological sampler of the Antarctic marine ecosystem

Weddell seals, Leptonychotes weddellii, are important apex predators in the food web of the Antarctic marine ecosystem. However, detailed information on their trophic relationships with cephalopods is scarce. Moreover, cephalopods play a key role in the marine environment, but knowledge of their feeding habits is limited by lack of data. Here, we have combined the use of this seal as a biological sampler together with measurements of the stable isotopic signature of the beaks of their cephalopod prey. Thus, the aims of the present study were: (1) to examine in detail the cephalopod portion of the diet of Weddell seals by means of scat analysis and (2) to assess the habitat use and trophic level of the different cephalopod prey taxa identified. From January to February 2009, a total of 48 faecal droppings were collected at Hope Bay, Antarctic Peninsula. Cephalopods were mainly represented by beaks (n = 83) which were identified to the lowest possible taxonomic level. Furthermore, subsamples of beaks were separated for further isotopic analysis. Relative abundance of stable isotopes of carbon (δ¹³C) and nitrogen (δ¹⁵N) was determined by continuous-flow isotope-ratio mass spectrometry. Cephalopods were represented uniquely by octopods of the subfamily Eledoninae. Pareledone turqueti was the dominant prey species followed by the papillated Pareledone species group and Adelieledone polymorpha. We conclude that Weddell seals preyed primarily on benthic prey resources. Furthermore, the relatively similar δ¹³C and δ¹⁵N values in beaks of the three octopod prey taxa suggest that these share the same type of habitat and occupy similar trophic level positions.     

Predation on cephalopods by Weddell seals, <Emphasis Type="Italic">Leptonychotes weddellii</Emphasis>, at Hope Bay,Antarctic Peninsula

Cephalopods play a key role in marine environments as food resources for top predators such as marine mammals and seabirds. However, detailed information on their trophic relationships with Antarctic seals is scarce. The aim of the present study was to examine the cephalopod portion of the diet of adult and subadult Weddell seals, Leptonychotes weddellii, at Hope Bay, Antarctic Peninsula, through the analysis of scats collected during three consecutive summers (2003, 2004 and 2005). Cephalopods occurred in almost 45% of the 217 samples collected during the whole period of study. A total of 662 beaks (358 upper and 304 lower) were removed from scats containing cephalopod remains (n = 93). Octopods were largely dominant in comparison with teuthoids constituting in numerical abundance over 95% of the cephalopod prey. The octopod Pareledone turqueti was the most frequent and dominant prey species representing, respectively, 57.9 and 71.1% in numbers and biomass of cephalopods consumed. Species belonging to the group of papillated Pareledone were second in importance. The results were compared with information from previous dietary studies of L. weddellii at other localities of Antarctica. Based on the examination of the cephalopod prey taxa identified in this study, it is suggested that during the study period Weddell seal individuals foraged mainly on benthic prey resources close to the coast, in inshore waters where octopods were dominant in comparison with pelagic squid.     

FEEDING METHOD AFFECTS OTOLITH DIGESTION IN CAPTIVE GRAY SEALS: IMPLICATIONS FOR DIET COMPOSITION ESTIMATION

Analysis of prey structures from scats has become widely used for assessing the diet of free-ranging seals. Captive feeding experiments have played an important role in the development of scat analysis but methodology has varied. We investigated whether using an experimental otolith-carrier species or in situ otolith experimental prey to feed haddock, plaice, and sand eel otoliths to two captive gray seals affected the amount of otolith digestion, and therefore our ability to estimate fish size and diet composition. Otolith recovery rates varied among the three prey species, but were not affected by feeding method. Otoliths were more digested in carrier experiments; resulting digestion coefficients were greater than those from in situ experiments. As a result, while the lengths of whole fish fed experimentally could be accurately predicted by applying in situ digestion coefficients, the lengths of these fish could not be accurately predicted using carrier digestion coefficients. Consequently, the estimated proportion by weight of each species in the diet can change markedly when carrier, as opposed to in situ , digestion coefficients are used. The implications for diet composition estimation are significant.     

STOMACH CONTENTS OF FALSE KILLER WHALES (PSEUDORCA CRASSIDENS) STRANDED ON THE COASTS OF THE STRAIT OF MAGELLAN,TIERRA DEL FUEGO

We examined the stomachs of 25 false killer whales collected from a mass stranding of 181 animals along both coasts of the Strait of Magellan, Chile, in March 1989. The 21 stomachs (nine males and 12 females), with food remains contained 11 prey species (nine cephalopods and two fishes) with a total of 442 individuals. Except for one case, food remains were meager, indicating that the animals had not eaten for some time or through stress had vomited on the beach. Eleven of the 21 animals had mud (often with squid beaks) in the esophagus and first stomach. The prey were identified employing squid beaks, fish otoliths and bones, and their wet weights were estimated using regressions between hard parts and known weights of species. The most important prey were the oceanic and neritic-oceanic squids, Martialia byadesi and Illex argentinus, followed by the neritic fish, Macruronus magellanicus. Of less importance were the oceanic squid, Todaroes fillipovae, the oceanic and epipelagic octopus, Ocytboe sp., and the oceanic squid, Moroteuthis ingens. The rest of the prey were poorly represented and included four oceanic squids and one neritic fish. The prey species of these animals were subantarctic, with two antarctic species, abundant over the Patagonian shelf and adjacent oceanic waters around Tierra del Fuego.     

Predation on cephalopods by Antarctic fur seals, Arctocephalus gazella, at two localities of the Scotia Arc, Antarctica

The cephalopod remains from 206 Antarctic fur seal (Arctocephalus gazella) scats collected at Mossman Peninsula, South Orkney Islands (n?=?105) and at Stranger Point, South Shetland Islands (n?=?101) contained 148 beaks (57 lower and 91 upper). The lower beaks were sorted and measured. Identification of 33 of the lower beaks that were collected at Mossman Peninsula revealed two squid species, Brachioteuthis ?picta (n?=?26) and Psychroteuthisglacialis (n?=?7), with lower rostral lengths (LRL) of 2.0–3.5?mm, and 1.0–2.5?mm, respectively. Identification of 15 of the lower beaks collected at Stranger Point revealed the same squid species, with the LRL of B. ?picta ranging from 3.0–3.4?mm (n?=?9), and that of P. glacialis from 2.0 to 3.5?mm (n?=?6). Estimated squid sizes and wet masses indicate that Antarctic fur seals feed on the small sub-adult squid which inhabit the surface layers. We have compared the squid diet estimated for the seals with that reported for its congeners in lower latitudes and other Antarctic seals, and conclude that cephalopods do not form an important food resource for Antarctic fur seals.     

Fish prey of Antarctic fur seals Arctocephalus gazella at Ile de Croy,Kerguelen

 The composition of Antarctic fur seal prey was assessed through analysis of scats collected in March 1994 on Ile de Croy, Iles Nuageuses. Fish remains predominated in samples, occurring in 95% of droppings. A total of 968 otoliths allowed the identification of 16 fish species. Myctophid fish (12 species) dominated the diet both by number (94% of the otoliths) and by fish reconstituted mass (76%). Three fish species constituted together 87% of the reconstituted mass: the myctophids Gymnoscopelus nicholsi (52%) and G. piabilis (12%), and the channichthyid Champsocephalus gunnari (23%). Prey distribution suggests that during late summer seals forage in upper slope waters in the northeast of the Kerguelen Archipelago. Received:1 March 1996/Accepted: 20 May 1996     

The diet of the subantarctic fur seal Arctocephalus tropicalis at Marion Island

Scats of subantarctic fur seals Arctocephalus tropicalis at Marion Island were collected from 1996 to 2,000, to examine temporal variability in the diet, factors affecting the variability and how the diet differed from that of the Antarctic fur seal A. gazella in the same period. For A. tropicalis, 19 prey species, of which 18 were fish and one a cephalopod, were identified in 213 scats. Fish were the main prey, occurring in 98.1 % of scats, whereas the cephalopod was present in only 1.4 % of scats. Amongst fish species, Myctophidae were most abundant, with Gymnoscopelus piabilis, G. fraseri and Electrona carlsbergi being the commonest prey items. Other fish families present in the diet in small numbers were Channichthyidae, Paralepididae, Nototheniidae, Microstomatidae and Notosudidae. Fish eaten ranged in size from Protomyctophum bolini and Krefftichthys anderssoni of standard length (SL) 25 mm to a single Dissostichus eleginoides of SL 249 mm. Differences in the diet existed between summer and winter. However, prey type accounted for most variability in the diet. In previous studies based on scats, a dominance of fish in the diet of A. tropicalis was also found at Possession Island (Iles Crozet), Amsterdam Island and Macquarie Island, but the dominant prey species differed between the various localities, also suggesting that prey availability is a major determinant of diet. At Marion Island, from 1996 to 2000 the diet of A. gazella comprised similar prey to that of A. tropicalis, but the proportional contribution of prey types differed in instances.     

Diet estimation based on an integrated mixed prey feeding experiment using Arctocephalus seals

Food web models depend on identifying which taxa are eaten and in what proportion they are consumed. Arctocephalus seals are generalist foragers and are an ongoing focus of Southern Hemisphere marine ecosystem research. This is the first feeding experiment to use Arctocephalus spp. to assess the utility of hard part scat analysis for diet estimation, based on mixed prey diets integrated over several days. Recovery rates of otoliths were extremely low for all taxa (0-9%). Although we could not collect scats produced during a 90 min period each day, during which the seals had access to a large pool, this result could not be attributed to otolith robustness, pinniped species or class, activity level, meal size or frequency, or fat content of the diet. We conclude that the unusually low recovery rates in this study may be due to unaccounted scats produced during 90 min of each day, if they contained otolith numbers an order of magnitude greater than all otoliths retrieved from scats produced during the other 22.5 h of each day, and/or may be related to the digestive processing of a mixed prey diet. Our study demonstrates the inadequacy of using otoliths in field collected scats for diet estimation due to the high level of unexplained variability of otolith occurrence in scats. We also identify two new potential sources of this variability. These are variability in numbers of otoliths per scat depending on activity level when a scat is excreted, and variability in recovery rates of otoliths as a function of the complexity of the diet.