Food and Feeding Ecology of the Neritic-Slope Forager Black-Browed Albatross and Its Relationships with Commercial Fisheries in Kerguelen Waters
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MARINE ECOLOGY PROGRESS SERIES Vol. 207: 183–199, 2000 Published November 22 Mar Ecol Prog Ser Food and feeding ecology of the neritic-slope forager black-browed albatross and its relationships with commercial fisheries in Kerguelen waters Yves Cherel*, Henri Weimerskirch, Colette Trouvé Centre d’Etudes Biologiques de Chizé, UPR 1934 du Centre National de la Recherche Scientifique, 79360 Villiers-en-Bois, France ABSTRACT: Food and feeding ecology of black-browed albatrosses Diomedea melanophrys rearing chicks was studied during 2 austral summers (1994 and 1995) at the Kerguelen Islands. Dietary analy- sis and satellite tracking were used to estimate potential interactions with commercial fisheries in the area. Fish comprised 73% by fresh mass of albatross diet; other significant food items were penguins (14%) and cephalopods (10%). Twenty-one species of fish (232 individuals) were identified and included mainly nototheniid and channichthyid species. The most important were Dissostichus elegi- noides (18.3% by reconstituted mass), Channichthys rhinoceratus (16.9%), Lepidonotothen squam- ifrons (11.6%), and to a lesser extent, Bathyraja sp. (4.5%) and Notothenia cyanobrancha (4.5%). The cephalopod diet was dominated by 3 taxa, the ommastrephid squids Todarodes sp. (7.6%) and Mar- tialia hyadesi (3.6%), and the octopus Benthoctopus thielei (2.4%). Satellite tracking indicated that during trips lasting 2 to 3 d, albatrosses foraged mainly over the outer shelf and inner shelf-break of the Kerguelen Archipelago. Birds moved to northern, eastern and southern waters, but never to the western Kerguelen shelf where there was a commercial longline fishery for D. eleginoides. Interac- tions with trawlers targetting D. eleginoides and Champsocephalus gunnari were of minor impor- tance in the northern shelf. There, offal from D. eleginoides was available to the birds; fish and cephalopod bycatch were negligible. Most of the natural prey of black-browed albatrosses are pri- marily benthic and semipelagic organisms not known to occur near the surface. Since we demon- strate that most of them were not scavenged behind fishing vessels, the way albatrosses catch dem- ersal organisms remains a mystery. KEY WORDS: Cephalopods · Benthic prey · Diet · Fish · Procellariiformes · Satellite tracking Resale or republication not permitted without written consent of the publisher INTRODUCTION dental mortality (DeGange & Day 1991, Robertson & Gales 1998) and the competition for common resources Increasing exploitation of marine resources by man (Montevecchi 1993); the positive include the provision has focused attention on the management of marine of new resources through offal and discards (Blaber et ecosystems and the factors affecting them. Of major al. 1995, Garthe et al. 1996). concern are the effects of competition between marine Analysis of stomach contents of seabirds allows the apex predators and fisheries, and the conservation of determination of their main prey (Duffy & Jackson endangered species. In this context, accurate informa- 1986). Food can include some easily identified items tion on the diets and feeding ecologies of predators is associated with fisheries, such as fish and squids that essential if the effects of human activities are to be are not naturally known from the area and are used as minimized. Fishery operations affect seabirds in 2 bait on longliners (Catard et al. 2000). More often, ways, negative and positive. The negative include inci- however, the identified species occur commonly in the foraging area and it is therefore a major challenge to *E-mail: [email protected] separate the species naturally eaten from those made © Inter-Research 2000 184 Mar Ecol Prog Ser 207: 183–199, 2000 available by the fisheries. It is consequently generally Kuba 1998). We consequently made a reference collec- assumed or hypothesized that prey known to be pri- tion of otoliths and bones for all the main species marily benthic or semipelagic are items discarded dur- occurring in Kerguelen waters to help identify fishes. ing fishery operations (Croxall et al. 1995, Reid et al. We also calculated allometric equations between 1996, Freeman 1998, Imber 1999), or adults that appear otolith length (OL), mandible length (ML), standard at the surface as a result of post-spawning mortality length (SL) and fresh mass (M) to estimate the size (Croxall et al. 1988), as already suggested for squids. of albatross prey. Interactions with fisheries were The first assumption is rarely verified by working on assessed by satellite tracking of adult albatrosses the quantification of fishery waste and the correspond- (Weimerskirch et al. 1997b), by comparison of the ing importance of the discarded species in seabird diet birds’ foraging locations with fishery areas, and by at the same time (Thompson 1992). Moreover, informa- quantifying birds attending vessels and fishery waste tion is lacking on the biology of some prey, thus indu- (Cherel et al. 1996, Weimerskirch et al. 2000). cing misinterpretation. It has been shown, for example, that small juvenile southern blue whiting Micromesis- tius australis, which were assumed to be taken by alba- METHODS trosses in relation to a bottom trawl fishery at the Falk- lands (Thompson 1992), is more probably a natural Study site, birds and field collection of samples. The prey there, as recently demonstrated in New Zealand study was conducted at the southern colony of Canyon waters (Cherel et al. 1999). des Sourcils Noirs (49° 41’ S, 70° 14’ E), Jeanne d’Arc The diet of albatrosses is generally dominated by fish Peninsula, southern Kerguelen Islands, where 1000 to and cephalopods. Fish ranked first by mass in the diet 1200 pairs of black-browed albatrosses breed annually of 3, and second in the diet of 6 albatross investigated (Weimerskirch et al. 1989). so far (Cherel & Klages 1998). Considering the impor- Between 10 and 27 February 1994 and between 3 tance of fish as prey, little qualitative and quantitative and 20 February 1995, 23 nests were observed contin- data are available on the contribution of different fish uously from dawn to dusk to monitor the visits of indi- species to the diet of albatrosses. Generally, the stom- vidual adults to their chick. One adult of each pair was ach contents are digested, which precludes the identi- marked on the breast with a patch of picric acid to fication of prey by the use of any external diagnostic make identification easier at a distance. The exact features. Thus, either no, or very little fish remains times of arrival and departure of the adults were noted have been identified to the species level in previous continuously. All the chicks were weighed twice daily, studies (Prince 1980, Thomas 1982, Harrison et al. in the early morning and late evening. Whenever feed- 1983, Croxall et al. 1988, Hunter & Klages 1989, ing was observed, the chick was also weighed after the Cooper et al. 1992, Ridoux 1994, Cooper & Klages departure of the parents (see Weimerskirch et al. 1995, Reid et al. 1996, Green et al. 1998a, Imber 1999). 1997b for additional information). Consequently, depending on the species considered, A total of 114 dietary samples were taken from ran- the quantitative composition of the fish diet of alba- domly-selected chicks after a returning parent had tross is still poorly or essentially unknown (Reid et al. completed feeding them. Forty-five samples were col- 1996). lected in 1994 (37 in February and 8 in early April), and The main objective of this study was to investigate 69 samples in 1995 (47 in February and 22 in late the food and feeding ecology of the black-browed March). Other prey items and additional incomplete albatross Diomedea melanophrys, and to critically stomach contents were also taken opportunistically evaluate the importance of fishery waste and natural during these periods. Samples were obtained by up- prey in its diet at the Kerguelen Islands, southern ending chicks over a plastic bucket and massaging the Indian Ocean. Black-browed albatrosses from the stomach and throat. All samples were returned deep- study colony at Canyon des Sourcils Noirs are known frozen (–20°C) to the laboratory in France for analysis. to feed mainly on fish (Weimerskirch et al. 1988), but Dietary analysis. Each sample was thawed and no detailed information on the prey eaten is available drained, and accumulated cephalopod beaks were in the literature. Traditionally, scientists working on subsequently sorted. Beaks can persist in predator piscivorous bird diets have tended to limit their stomachs for weeks and even months, thus overem- research to the examination of otoliths only, disregard- phasizing their importance in seabird diets. Following ing skeletal remains. Various factors may affect the Cherel & Klages (1998), accumulated beaks (without final qualitative and quantitative results of this method, flesh attached) were consequently analysed separately leading to the underestimation, or ignorance of the from fresh items. These beaks were not considered presence of, some species and also to misleading when calculating the reconstituted proportion by mass results in back-calculation of fish sizes (Gosztonyi & in the diet of the different prey species. Cherel et al.: Feeding ecology of albatrosses at Kerguelen 185 Fresh remains were divided into broad prey classes satellite transmitters (8 T2038 Toyocoms and 2 Micro- (fish, cephalopods, crustaceans, penguins and others), wave 100s) weighing between 55 and 80 g (i.e. 1.5 to which were weighed to calculate their proportion by 2.2% of the birds’ body mass). The transmitters were mass in the diet. Identification of prey relied almost fitted on back feathers with adhesive tape and left on totally on the examination of otoliths and bones for the birds for 1 to 5 successive foraging trips. Between 8 fish, keratinized beaks for cephalopods, and exoskele- and 15 locations were received each day per transmit- tons for crustaceans. Special care was made to use all ter, giving a total of 1290 and 857 locations in 1994 fish hard parts recovered in stomach contents (bones, and 1995, respectively (for further information, see cartilageous elements, otoliths, scales, thorns, teeth), Weimerskirch et al.