Records of vagrant fur seals (family Otariidae) in South Australia

Transcription

1 CSIRO PUBLISHING Australian Mammalogy, 2014, 36, Records of vagrant fur seals (family Otariidae) in South Australia Peter D. Shaughnessy A,C, Catherine M. Kemper A, David Stemmer A and Jane McKenzie B A South Australian Museum, rth Terrace, Adelaide, SA 5000, Australia. B 235 Robin Road, Semaphore, SA 5019, Australia. C Corresponding author. peter.shaughnessy@samuseum.sa.gov.au Abstract. Two fur seal species breed on the southern coast of Australia: the Australian fur seal (Arctocephalus pusillus doriferus) and the New Zealand fur seal (A. forsteri). Two other species are vagrants: the subantarctic fur seal (A. tropicalis) and the Antarctic fur seal (A. gazella). We document records of vagrant fur seals in South Australia from 1982 to 2012 based primarily on records from the South Australian Museum. There were 86 subantarctic fur seals: 49 specimens and 37 sightings. Most (77%) were recorded from July to October and 83% of all records were juveniles. All but two specimens were collected between July and vember. Sightings were prevalent during the same period, but there were also nine sightings during summer (December February), several of healthy-looking adults. table concentrations were near Victor Harbor, on Kangaroo Island and Eyre Peninsula. Likely sources of subantarctic fur seals seen in South Australia are Macquarie and Amsterdam Islands in the South Indian Ocean, ~2700 km south-east and 5200 km west of SA, respectively. There were two sightings of Antarctic fur seals, both of adults, on Kangaroo Island at New Zealand fur seal breeding colonies. Records of this species for continental Australia and nearby islands are infrequent. Additional keywords: Antarctic fur seals, Arctocephalus gazella, Arctocephalus tropicalis, geographic distribution, subantarctic fur seal, sightings, strandings. Received 8 vember 2013, accepted 20 February 2014, published online 10 June 2014 Introduction Three species of otariid seal (family Otariidae) breed on the southern coast of continental Australia and its nearby islands: the Australian fur seal (Arctocephalus pusillus doriferus), the New Zealand (NZ) fur seal (A. forsteri) and the Australian sea lion (Neophoca cinerea). Two other fur seal species have been recorded but do not breed there, the subantarctic fur seal (A. tropicalis) and the Antarctic fur seal (A. gazella); their presence in South Australia (SA) is documented here. Recent taxonomic revision (Committee on Taxonomy Society for Marine Mammalogy 2012) has resulted in changes to the names of three of the fur seal taxa: Arctophoca australis forsteri (NZ fur seal), Arctophoca tropicalis (subantarctic fur seal) and Arctophoca gazella (Antarctic fur seal). These names have not been widely accepted by the scientific community and therefore are not used here. The occurrence of vagrant seals on the Australian coast was summarised briefly by Shaughnessy (1999) and by National Seal Strategy Group and Stewardson (2007). The subantarctic fur seal is a well known visitor to Australian waters; for example, Gales et al. (1992) documented 37 records between 1980 and 1991 in Western Australia, SA, Victoria, Tasmania and New South Wales. Subsequent to that study, records of subantarctic fur seals in southern Australia have been documented for Western Australia (Mawson and Coughran 1999), Victoria (Warneke 1995), Tasmania (PMTMMCP 2011), Journal compilation Australian Mammal Society 2014 New South Wales (Llewellyn et al. 1994; Haynes-Lovell 1995) and Queensland (Haynes-Lovell 1995). In addition, there is a specimen in the Queensland Museum (Martin 1975) and two tagged juveniles were seen on Kangaroo Island in 1999 (C. Guinet in Wynen et al. 2000). The cream-coloured chest, throat and face of this species make it easily recognisable and it tends to attract media attention when sighted ashore. The Antarctic fur seal is more uniformly coloured and is difficult to distinguish from Australian and NZ fur seals. The only record of it on the Australian coast is the sighting of a juvenile at Kangaroo Island in early 1999 that had been tagged at Macquarie Island (M.-A. Lea, in Wynen et al. 2000). The subantarctic fur seal is widely distributed in the Southern Ocean, with breeding colonies at Gough Island in the South Atlantic Ocean, and at the Prince Edward Islands, Crozet Archipelago and Amsterdam Island in the South Indian Ocean (Goldsworthy 2008b). The nearest breeding colony to mainland Australia is at Macquarie Island, where some of them hybridise with the Antarctic fur seal and the NZ fur seal (Lancaster et al. 2006; Goldsworthy et al. 2009). The subantarctic fur seal is ashore for breeding in December and early January and produces a single pup annually. Adult females are mated ~1 week after parturition and return to the sea a day or two later. Pups are suckled for ~10 11 months, with adult females alternating between foraging at sea and nursing bouts ashore (Goldsworthy 2008b). Adult males haul out at colonies in early autumn to

2 Vagrant fur seals in South Australia Australian Mammalogy 155 moult and are rarely ashore during winter when subadults are ashore in small numbers (Kerley 1983). The Antarctic fur seal is also widely distributed in the Southern Ocean, with most breeding colonies in more southern latitudes than those of the subantarctic fur seal, although both species breed on some islands. Antarctic fur seal breeding colonies are situated at South Georgia, at islands in the Scotia Arc and Antarctic Peninsula, at Bouvetøya, and at islands in the South Indian Ocean including Prince Edward Islands, Isles Crozet, Kerguelen, as well as at Heard and Macquarie islands (Goldsworthy 2008a). Females produce a single pup annually in December and early January; pups wean at ~4 months of age, after which adult females and pups depart. Adult and subadult male Antarctic fur seals display a similar haul-out pattern to subantarctic fur seals, with maximum numbers ashore during summer (December February) and the autumn moulting peak (Page et al. 2003; Carlini et al. 2006). Both species were recognised as part of the Australian fauna based on breeding populations at Macquarie and Heard islands in the Southern Ocean (Shaughnessy 1992). In order to improve our knowledge of the subantarctic fur seal and Antarctic fur seal in Australian waters, we document their occurrence in SA and investigate their geographic, temporal and age sex variation. Materials and methods Sources of data The primary source of records of vagrant subantarctic fur seals for this study was the South Australian Museum, from the registered specimen collection and from files containing data relevant to sightings of live and dead animals that are not represented by specimens in the museum. They are referred to as specimens and sightings, respectively, and collectively as records. All specimens were dead but not all sightings were of dead animals. Nine early records duplicate those in Gales et al. (1992) for southern Australia and another four records are from Kemper et al.(2008) in a regional account for Gulf Saint Vincent and the northern coast of Kangaroo Island; they are included here for completeness. Records date from September 1982 to vember Many of the seals were observed and reported by officers of the Department of Environment, Water and Natural Resources (DEWNR and its predecessors). Other sources of records were the DEWNR fauna database of sightings, website of the Australian Marine Wildlife Research and Rescue Organisation, Adelaide (AMWRRO) and members of the public. The two sightings of Antarctic fur seals were made by observers who have worked with that species at Macquarie Island and are familiar with fur seals in SA. Care was taken to ensure that repeat sighting records of individuals were not included. In many instances a photograph or digital image of the seal was obtained (74% of records); records without photographic evidence were accepted only from experienced observers. The reliability of species identification was graded as: 1, certain, a museum specimen was available, an image was obtained that showed recognisable features, or a description of an adult or subadult was provided by a recognised observer; 2, probable, based on identification of some juvenile skulls, a poor image or an equivocal description from a recognised observer; 3, less certain. In the tables that summarise records, geographical coordinates refer to features listed in the South Australian State Gazetteer. The coordinates are based on the Gazetteer, or on positions from GPS, Google Earth and 1 : topographic maps. In the text and figures, we refer to seasons as summer beginning in December, autumn beginning in March, winter beginning in June and spring beginning in September. During necropsy of specimens (by South Australian Museum staff and associates), several external measurements were taken including weight (kg) and standard length (SL, in millimetres, nose to tail in a straight line) and axillary girth (mm). The body condition of each animal was noted, a search was made for external parasites and attempts were made to determine the cause of death. Skeletal material was prepared and has been incorporated into the South Australian Museum mammal collection. Barnacles collected from three subantarctic fur seals were identified by staff of the Marine Invertebrates section of the South Australian Museum and are lodged in that collection. Age and sex categories Age categories of subantarctic and Antarctic fur seals were based on those used by Goldsworthy and Shaughnessy (1994) supported by descriptions in Condy (1978) and Shaughnessy et al.(1988), and measurements in Goldsworthy (2008a, 2008b). They were: adult males, with well developed crest, chest and shoulders (standard length cm); subadult males, males smaller than adult males and as big as or slightly larger than adult females; adult females, mid-sized animals with a short, broad nose (standard length cm); juveniles, smaller than the other age classes. For juvenile specimens, sex was determined by the presence or absence of the penile opening. This method was also possible for some sightings; others were sexed by inspection of the vent. For larger animals, sex was determined by body size and general appearance, as indicated above. Species recognition (non-skeletal) Specific recognition of live subantarctic and Antarctic fur seals was aided by reference to detailed descriptions and photographs of animals by Condy (1978), Goldsworthy (2008a, 2008b) and Shaughnessy et al.(1988). The distinctive vocalisations of adult male subantarctic fur seals and Antarctic fur seals were used to distinguish them from NZ fur seals (Page et al. 2002). Subantarctic fur seals were recognised by their creamcoloured chest, throat and face. Their dorsal pelage is chocolate brown to black and it contrasts with a cream-coloured chest and face that extends above the eyes. This coloration distinguishes them from Australian and NZ fur seals, which have a brown to dark grey pelage. Adult and subadult male subantarctic fur seals have a crest of black fur on top of the head. Adult females are also dark on their dorsal surface with a cream-coloured chest and face, and a dark belly. Juveniles are silver-grey dorsally and laterally, with varying amounts of cream-coloured fur ventrally. Their colouring on the chest and throat was not as obvious as in older animals, but it was apparent on the face and above the eyes. Subantarctic fur seals are sometimes confused with adult female and juvenile Australian sea lions because of the pale coloration of the sea lions ventral pelage. But the dorsal pelage

3 156 Australian Mammalogy P. D. Shaughnessy et al. of the subantarctic fur seal is much darker than that of the sea lion. Juvenile subantarctic fur seals are sometimes confused with NZ fur seals; the more pointed nose of NZ fur seals distinguishes them from subantarctic fur seals, which have a relatively blunt nose. Pelage colour of Antarctic fur seals is uniform silver grey to brown and they can be difficult to distinguish from the resident NZ fur seals. Coloration of adult females is variable from silver-grey to brown dorsally and paler on the chest; their body form is svelte compared with female subantarctic fur seals (Condy 1978). Antarctic fur seals were recognised by their distinctly longer flippers in relation to body size and by their slow and awkward movement over rocks, features Condy (1978) noted when comparing them with subantarctic fur seals. Juvenile Antarctic fur seals are similar in external appearance to juvenile subantarctic fur seals; juvenile and adult female Antarctic fur seals lack the cream-coloured markings above the eye that are found in subantarctic fur seals. Species recognition (skeletal) Species identification of subantarctic fur seal specimens was based on three skull characters; their appearance was compared in skulls of subantarctic fur seals and the other fur seals that occur in SA, the NZ fur seal and Australian fur seal (Fig. 1). First, the general shape of the skull: when viewed from above or below, it is more triangular in subantarctic fur seals than in the other two species, in which it is more elongate. Second, the size of the postcanine teeth is smaller in subantarctic fur seals than in the other two species; this character was noted by Repenning et al.(1971). Third, the position of the palatine suture is usually posterior to the anterior extremity of the orbit in subantarctic fur seals. It is about level with the anterior extremity of the orbit in Australian fur seals and anterior to it in NZ fur seals. The position of the palatine suture was the most useful of these three characters for differentiating skulls of subantarctic fur seals from those of the other two fur seal species. There is variation in each of these skull characters and some specimens are difficult to identify reliably on the basis of these characters alone. Hybrids between A. tropicalis, A. gazella and A. forsteri have been reported from Macquarie Island (e.g. Lancaster et al. 2006) and it is possible that some hybrids could have been included in this study. The specific identity of some specimens was confirmed by the Evolutionary Biology Unit of the South Australian Museum using isozyme electrophoresis (Table 1). In this study, species identification was based on all available characters, including photographs or digital images. Any uncertainty is reflected in our ranking of the reliability of species identification. Body condition The condition was noted for 56 subantarctic fur seals by external inspection of the whole body, with added information from necropsies of specimens. Two categories were used, poor (which included emaciated ) and good (which included robust and excellent ). If the bony outline of an animal s skeleton was apparent under its pelage, it was classed as poor. Those with no blubber layer on the inside of the pelage were considered emaciated. If the bony outline of the skeleton was not apparent, the animal s condition was classed as good. Statistical analyses Data for the two condition categories for specimens and sighting records were set out in 22 contingency tables and compared using a Chi-square statistical test incorporating Yates correction for continuity. Statistical significance was accepted at the 5% level. Fig. 1. Skulls of three species of juvenile fur seal (NZ, subantarctic and Australian fur seal) showing the three characters used for identification: the general shape of the skull, the size of the postcanine teeth, and the position of the palatine suture relative to the anterior end of the orbit. The lines join the anterior extremities of the orbits; the arrows point to the palatine sutures. Specimens are: A. forsteri M25076, A. tropicalis M24279, A. pusillus doriferus M24750.

4 Vagrant fur seals in South Australia Australian Mammalogy 157 Table 1. Records of specimens of subantarctic fur seals in South Australia held at the South Australian Museum (SAM) The column headed Species ID refers to the reliability of the species identification (see Methods). KI, Kangaroo Island. AMWRRO, Australian Marine Wildlife Research and Rescue Organisation, Adelaide. RSPCA, Royal Society for the Prevention of Cruelty to Animals. SL, standard length Date first sighted SA Museum specimen. Species ID Locality and geographical coordinates Data Collected or reported by Comment Images date M South Australia Juvenile male data Registered at SAM, 5 v Sep M South Australia Juvenile male, SL 927 mm DEWNR 30 Aug A M Coorong, (35.07 S, E) 1988 A M Hanson Bay (36.03 S, E) 17 Sep A M C Blackfellows Caves (37.95 S, E) 1 v A M C 11 km S Salt Creek, (36.24 S, E) 30 Jul A M km ESE Goolwa (35.55 S, E) Aug A M Wheaton s Beach, D Estrees Bay, KI (36.02 S, E) 26 Jul A M C 9.5 km SE Goolwa (35.58 S, E) 22 Sep M C Western end of Goolwa Beach (35.52 S, E) 28 Oct B M C Penneshaw (35.72 S, E) 5 Oct M km SSE Robe (37.25 S, E) 3 Aug M Goolwa (35.53 S, E) 8 Jul M Victor Harbor (35.55 S, E) 1 Aug M Murat Bay (32.08 S, E) 25 Aug M Port Elliot (35.53 S, E) Juvenile, weight 10.3 kg, SL 950 mm Unknown Alive, taken to Marineland, Adelaide; died Dec. 1989; age estimated as 3 years Juvenile male, SL 915 mm DEWNR Collected during winter (Jun. Aug.) Juvenile female, S. Childs Found alive at high-water weight 6.5 kg, SL 810 mm mark, died 4 days later Juvenile male W. Head Alive, died in captivity Juvenile male, weight 11 kg, SL 840 mm, girth 506 mm Juvenile male, weight 11 kg, SL 872 mm Juvenile male, weight 9 kg, SL 840 mm D. J. Needham, Adelaide Alive, in poor condition, euthanased, fractured spine; barnacles L. australis attached to flippers and lower body DEWNR Dead, emaciated; death caused by peritonitis A. Wood, Port Elliott Alive, in poor condition; euthanased at RSPCA, Lonsdale 5d later Juvenile, weight ~15 kg J. Sorrell, DEWNR Alive; died at RSPCA, Lonsdale P. Seager, DEWNR Alive, in poor condition, Juvenile female, SL 920 mm Juvenile female, weight 16 kg, SL 970 mm Adult male J. Bracken and C. Halstead, DEWNR died at RSPCA, Lonsdale G. Watson, DEWNR Found dead on beach Juvenile female DEWNR Alive, taken to Adelaide Zoo for rehabilitation, but died Juvenile female, weight 12 kg, SL 822 Juvenile male, weight 13 kg, SL 910 mm, girth 380 mm DEWNR, Ceduna J. Sorrell, DEWNR Alive, in poor condition, died next day

5 158 Australian Mammalogy P. D. Shaughnessy et al. 9 Jul M Basham Beach, 1.8 km NE Port Elliot (35.52 S, E) 18 Sep M Fishery Bay, 6.3 km NE Cape Carnot (34.92 S, E) 24 Sep M Basham Beach, 2.0 km SW Middleton (35.53 S, E) 24 Oct B M Grange, Adelaide (34.91 S, E) 11 Dec M Victor Harbor (35.55 S, E) 2002 M South-east SA, (approx. 38 S, E) 12 Oct M km NNW South End (37.53 S, E) 26 Oct M Causeway at Inman River, Victor Harbor (35.56 S, E) 30 Oct M Port Kenny (33.18 S, E) 23 v M Talia Beach (33.33 S, E) 30 Jul M Talia Beach (33.33 S, E) 16 Aug M Little Dip Conservation Park, Robe (37.24 S, E) 10 Sep M The Granites, Point Westall (32.88 S, E) Oct M Anxious Bay (33.38 S, E) 2 Oct M Sheringa Beach (33.88 S, E) 7 Oct M Tumby Bay (34.37 S, E) 19 Oct M km NW Elliston (33.62 S, E) 23 Oct M Tod River mouth, Port Lincoln (34.59 S, E) 29 Oct B M Taperoo, Adelaide (34.81 S, E) Juvenile male, weight 11 kg, SL 910 mm, girth 220 mm J. Bracken, DEWNR Emaciated, stomach empty Juvenile male, weight 11.5 kg, SL 904 mm, girth 265 mm J. Tilley, DEWNR Dead, emaciated Juvenile male, weight 9 kg, SL 860 mm, girth 250 mm Juvenile male, weight 13.5 kg, girth 270 mm Juvenile female, weight 9.5 kg, SL 817 mm, girth 450 mm Juvenile male, weight 13 kg, SL 1010 mm, girth 448 mm Juvenile female, weight 5.5 kg, SL 755 mm, girth 405 mm Juvenile male, weight 8.5 kg, SL 860 mm J. Bracken, DEWNR Alive, died at RSPCA Lonsdale; emaciated, fluid in stomach (otherwise empty) M. Long and I. Tomo, SAM Found dead, decomposed J. Bracken, DEWNR Robust, stomach empty DEWNR, Mt Gambier Robust, collected before 26 Sep G. Jackway, DEWNR Dead on beach, in poor condition; barnacles (L. anatifera) attached to body T. Te, DEWNR Dead, emaciated Juvenile male, weight 9.5 kg, SL 870 mm, girth 430 mm Juvenile male, weight 9.5 kg, SL 930 mm, girth 430 mm Juvenile female, weight 12.5 kg, SL 935 mm, girth 260 mm Juvenile male, weight 8.3 kg, SL 852 mm Juvenile male, weight 10 kg, SL 940 mm, girth 410 mm Juvenile male, weight 12.5 kg, girth 280 mm L. Amey, DEWNR Dead, in poor condition L. Amey, DEWNR In poor condition D. Armstrong, DEWNR Dead, in poor condition J. Haywood and M. Christie, Carpenter Rocks Dead, in poor condition SA Fisheries, Streaky Bay Dead, emaciated L. Amey, DEWNR Dead, in poor condition, stomach empty Juvenile male, weight 6.9 kg L. Olsen Alive, emaciated, died a few days later at Adelaide Zoo Juvenile female, weight 7.5 kg, SL 800 mm, girth 400 mm Juvenile female, weight 8.5 kg, SL 892 mm, girth 210 mm Juvenile male, weight 7.5 kg, SL 800 mm, girth 400 mm Juvenile B. Pendlebury and R. Morcom, DEWNR RSPCA Emaciated L. Amey, DEWNR Dead, emaciated, stomach empty P. Wilkins, DEWNR Dead Dead, decomposed (continued next page)

6 Vagrant fur seals in South Australia Australian Mammalogy 159 Date first sighted SA Museum specimen. Species ID Locality and geographical coordinates 25 v M Middleton (35.52 S, E) 28 Aug M km SE Goolwa (35.54 S, E) 6 Oct M At sea, 4.5 km SSE of Rosetta Head, Victor Harbor (35.59 S, E) 17 Oct M At sea, 4.5 km SSE of Rosetta Head, Victor Harbor (35.59 S, E) 18 Oct M km SW Victor Harbor (35.56 S, E) 21 Oct M Smooth Pool, Point Westall (32.93 S, E) 26 Jun M Vivonne Bay (35.98 S, E) 25 Aug M Mount Camel Beach, 9 km SE Venus Bay PO (33.28 S, E) 30 Aug M km NNE Point Warna, (34.52 S, E) 30 Sep M Mt Camel Beach, 9.2 km SE Venus Bay PO (33.29 S, E) 6 Aug M Talia Beach (33.33 S, E) 20 Sep M Semaphore, Adelaide (34.83 S E) 6 Aug M Horseshoe Bay, Port Elliot (35.53 S, E) 10 Apr M Merdayerrah Sandpatch (31.67 S, E) A ted by Gales et al. (1992). B ted by Kemper et al. (2008). C Specific identity confirmed by SAM Evolutionary Biology Unit using isozymes. Table 1. (continued) Data Collected or reported by Comment Images Juvenile male, weight 8.5 kg, SL 854 mm Juvenile male, weight 11.5 kg, SL 950 mm, girth 460 mm Juvenile male, weight 10 kg, SL 944 mm, girth 410 mm P. Unsworth, DEWNR Washed ashore dead, emaciated S. Gilbert, DEWNR In poor condition P. Unsworth, DEWNR Dead, emaciated Juvenile female, weight 9.5 kg, SL 892 mm, girth 440 mm Member of the public and DEWNR Found dead in sea, emaciated, crustacean remains in stomach Juvenile male, weight 10.5 kg, SL 910 mm Juvenile female, weight 7 kg, SL 790 mm, girth 400 mm Adult male aged 12.5 years, weight 47.5 kg, SL 1482 mm, girth 762 mm P. Unsworth, DEWNR In poor condition, much haemorrhaging DEWNR Alive, in poor condition, died soon after D. Snowball and P. Arthur, Dead, in poor condition; DEWNR tagged Juvenile D. Armstrong, DEWNR Dead, decomposed Juvenile male, weight 8 kg, SL 855 mm, girth 355 mm Juvenile male, weight 8.5 kg, SL 890 mm, girth 416 mm Juvenile male, weight 6.0 kg, SL 770 mm, girth 395 mm Juvenile male, weight 9.3 kg, SL 755 mm, girth 485 mm T. Hall, DEWNR Alive, emaciated; died next day D. Armstrong, DEWNR Alive, emaciated, died a few hours later; blunt trauma and haemorrhaging to neck L. Amey, DEWNR Dead, emaciated; Juvenile male A. Machado, AMWRRO; in Kemper and Tomo (2011) Juvenile G. Armstrong, DEWNR barnacles (L. anatifera) on lower back A. Machado, AMWRRO Dead, robust, severe blunt trauma Nullarbor Biological Survey Alive, died at AMWRRO, Torrens Is. Skull picked up from beach

7 160 Australian Mammalogy P. D. Shaughnessy et al. Results Subantarctic fur seals Between 1982 and 2012 there were records of 86 individual subantarctic fur seals in SA (Tables 1 and 2, Fig. 2), which included 49 registered museum specimens and 37 sighting records. Each museum specimen was represented by a skull; some included postcranial material and/or skins. The reliability of species identification was graded as 1 for all but two of the specimens; all 16 specimens that included a skin were graded 1. For the sighting records there were 22, 13 and 2 animals in classes 1, 2 and 3, respectively. The geographic spread of records was from Blackfellows Cave in the east (Gales et al. 1992) to Merdayerrah Sandpatch in the west. table concentrations were in the area between Victor Harbor and Goolwa (19 records), Kangaroo Island (17) and Eyre Peninsula (23), of which 14 were from Coffin Bay westward to Point Westall, and 9 were at the southern end of Eyre Peninsula between Tumby Bay and Cape Carnot. These localities accounted for 69% of the records. Of the 49 museum specimens, 21 were dead when found, 16 were alive and nothing was reported for the other 12. Of the 16 live animals, 13 died within a few days of being sighted, two were euthanased and one (M15968) was taken to a captive facility where it lived for 2 years and 4 months. Of the 37 sighting records, four were found dead and 33 were alive. Of the latter, four were found dead soon afterwards. Age-class was recorded for 80 of the seals: most were juvenile (66 or 83%), four were subadults and 10 were adults. The month when animals were collected or sighted was noted for 82 records (Fig. 3, which excludes the pickup skull, M25874). Most (77%) were between July and October. All but two of the museum specimens were reported between July and vember. Sightings were also prevalent then, but there were nine sightings during summer, six of them in January. Of the nine summer sightings, four were of adult males, two were of subadult males, two were of juveniles and one was of an adult female. The distribution of sex and age-classes through the four seasons is available for 76 records (Fig. 4); most juveniles (95%) were recorded in winter and spring, and the sole adult femalewasseeninsummer. Sex was determined for 61 animals, of which 46 (75.4%) were male and 15 (24.6%) were female. Sex was determined for 47 of the 67 juveniles, of which 33 (70%) were male, which differs significantly from a 1 : 1 ratio (c 2 = 7.7, d.f. = 1, P = 0.006). The number of records increased with time during the study, with the 5-year period from January 2003 to December 2007 having the most records (32: Fig. 5). For the six 5-year periods between January 1982 and December 2012, the mean increase was 4.2 animals per period. The year with most records was 2005, with 13. Body condition data were available for 54 animals (Table 3). The distribution of the two categories of body condition among specimens and sighting records was heterogeneous (c 2 = 21.0, d.f. = 1, P < 0.001). Body condition was better among sighting records (75% were rated as good) than specimens (all of which were dead, 10%). Among juveniles, body condition was noted for 41 animals (Table 3). The distribution of the two categories of body condition among specimens and sighting records for juveniles was heterogeneous (c 2 = 8.16, d.f. = 1, P = 0.004). Body condition was better among sighting records (58% were rated as good) than specimens (10%). Some of the juveniles could have been in their second year. The heaviest juvenile was a female (M17423, 16 kg) that was collected in October This is heavier than the weaning weight of female pups at Macquarie (13.6: Goldsworthy 1992), Amsterdam (14.4 kg: Tollu 1974) and Marion Islands (13.5 kg: Kerley 1985), suggesting that this animal was in its second year. The other juveniles were lighter than the weaning weights of male and female pups at Amsterdam and Marion Islands, but because some of them were emaciated, it is not appropriate to estimate their age. Examination of tooth structure would provide useful information on their age and enhance our understanding of these options. One tagged subantarctic fur seal (M24981) is included in the specimens, an adult male found dead on the beach at Vivonne Bay, Kangaroo Island, on 26 June 2007 (Table 1). It was in poor condition and showed signs of trauma. It was tagged on the foreflipper with an orange tag that included details of the research institution. It had been tagged as a pup on Amsterdam Island in March 1995 (C. Guinet, in litt.) and hence was 12.5 years old. Barnacles (Lepas spp.) were attached to the pelage and/or flippers of three animals: L. australis were on a juvenile male (M16249) and L. anatifera were attached to a juvenile female (M23780) and a juvenile male (M25572). Antarctic fur seals The two records of Antarctic fur seals for SA were sightings of live animals in good condition from NZ fur seal colonies on Kangaroo Island (Table 4). They were made during summer by experienced seal biologists studying NZ fur seals. An adult female was sighted at Cape du Couedic on the south coast of Kangaroo Island on 2 February 1994 by PDS and C. Bald. It was on the seaward edge of the Nautilus Rock breeding colony in an area used by non-breeding animals. It was more svelte in appearance than the NZ fur seals, and its gait over rocks was slower and more awkward. An adult male was sighted at Cape Gantheaume on the south coast of Kangaroo Island on 14 December 2000 by JM. Its pelage coloration and vocalisations assisted in its identification. It was resting on a headland within the fur seal colony among adult and subadult male NZ fur seals in a haul-out area used primarily by males. Although the observation was made during the fur seal breeding season, no territorial behaviour was apparent in that haul-out area. The animal was judged to be an adult male because it was similar in size to the adult male NZ fur seals nearby. tags or tag rips were seen on its fore-flippers. The Antarctic fur seal was not seen again, even though JM inspected the Cape Gantheaume fur seal colony daily during the summer of Discussion Subantarctic fur seals The 86 records documented here include nine of the 10 records of Gales et al. (1992), seven specimens and two sightings. The tenth record (SA Museum M15971) was reidentified by Brunner

8 Vagrant fur seals in South Australia Australian Mammalogy 161 Table 2. Sighting records of subantarctic fur seals in South Australia from reports at the South Australian Museum (SAM) and other sources as indicated The column headed Species ID refers to the reliability of the species identification (see Methods). KI, Kangaroo Island. AMWRRO, Australian Marine Wildlife Research and Rescue Organisation, Adelaide. SARDI, SA Research and Development Institute, Adelaide. RSPCA, Royal Society for the Prevention of Cruelty to Animals. SL, standard length. Place names in quotes are unofficial Date Species ID Locality and geographical coordinates Data Collected or reported by Comment Images 6 Sep A 2 Tumby Bay (34.37 S, E) 14 Aug A 1 Middleton (35.51 S, E) 23 Aug The Gap, south of Cape Elizabeth (35.29 S, E) May and Jun Cape Gantheaume, KI (36.07 S, E) 12 Jul B 2 rmanville (35.45 S, E) 17 Dec Corny Point (34.90 S, E) 30 Aug Louth Bay (34.54 S, E) 27 Jan Cape Gantheaume, KI (36.07 S, E) 28 Jul Seal Bay, KI (35.96 S, E) 28 Aug Flaherty Beach, Yorke Peninsula (34.95 S, E) 9 Aug Tea Tree Crossing, Younghusband Peninsula (36.19 S, E) Juvenile female, weight 9.3 kg, SL 950 mm G. Abel, Marineland, Adelaide Alive on beach, died next day Juvenile J. Sorrell, DEWNR Alive, in poor condition Juvenile L. Holmes, Weetulta Alive with netting on neck, caught, released next day nearby at Balgowan; decomposed body found at The Gap several weeks later was likely same seal Adult male Shaughnessy (1994; p. 13) Seen alive by S. Troy on several occasions a few metres from an adult male A. forsteri; in good condition Juvenile male, weight 18 kg C. Cock, RSPCA, Lonsdale, Adelaide Alive, rehabilitated at RSPCA and released at Fishery Beach, Cape Jervis, 25 Jul Penguin remains in faeces. Juvenile L. Holmes, Weetulta Alive, in poor condition Juvenile, weight 9 kg T. Berden and D. Forman, DEWNR Alive, in poor condition, released in Spalding Cove, Lincoln National Park Adult male M. and S. Berris, KI Alive, in good condition at headland 1 km north-east of Cape Gantheaume, in nonbreeding part of the NZ fur seal colony Juvenile C. Groves, DEWNR Alive in Western Prohibited Juvenile G. and P. Pain, Warooka, and L. Holmes, Weetulta Juvenile CMK and V. Weisbecker, SAM Area of the Australian sea lion colony (Dennis 2005); returned to sea Alive, in good condition, returned to sea Alive, cuts on left fore-flipper, returned to sea Sketch

9 162 Australian Mammalogy P. D. Shaughnessy et al. 10 Aug Seven Mile Beach, Coffin Bay National Park (34.51 S, E) 6 Jul Berris Point, Cape Gantheaume Conservation Park, KI (36.06 S, E) 13 Aug Berris Point, Cape Gantheaume Conservation Park, KI (36.06 S, E) 6 Jan Cape Gantheaume, KI (36.07 S, E) 19 Jan Cape Gantheaume, KI (36.07 S, E) 8 Aug Poonindie, Eyre Peninsula (34.58 S, E) 17 Oct Louth Bay (34.54 S, E) 20 Oct Newland Head (35.64 S, E) 25 Oct B 2 Ethelton, Adelaide (34.85 S, E) Jan rth Neptune Is. (main island) (35.23 S, E) 13 Jul Middleton (35.51 S, E) 23 Aug Port Lincoln (34.73 S, E) 30 Jan rth Casuarina Is (36.07 S, E) 2 Feb Liguanea Island (34.99 S, E) 7 Aug St Kilda (34.75 S, E) T. Gershwitz, DEWNR Alive, in poor condition, picked up by tourists, released next day in Coffin Bay Juvenile JM Alive, in good condition Juvenile B. Page, SARDI Alive, in good condition Subadult male JM Alive, in excellent condition, possible injured left shoulder or fore-flipper; seen nearby on 14, 16, 18 Jan Juvenile female JM Alive, in excellent condition Juvenile male L. Dunning; K. Villiers, DEWNR Alive, resting on beach in good condition, no apparent injuries Juvenile male M. Holton, DEWNR Dead Juvenile male S. Thompson, DEWNR Dead, in good condition, entangled in trawl net Juvenile, weight 8.1 kg A. Machado, AMWRRO Alive, in poor condition, on a road, died at Adelaide Zoo Adult male B. Page, SARDI Holding a territory in NZ fur C. O Neil, Middleton; S. Southcombe, DEWNR seal breeding colony; in good condition. Presumably it was this animal holding a territory in same area in previous and following breeding seasons. Dead, no obvious injuries; report included white fur around eye Juvenile male P. Wilkins, DEWNR Alive, on beach in town, relocated and released Adult male PDS Alive, in NZ fur seal colony in Adult female S. Goldsworthy and B. Page, SARDI good condition; also seen 31 Jan. and 1 Feb Alive, in NZ fur seal colony at north-eastern end of island; in good condition but no pup found Juvenile S. Taylor, DEWNR Alive, returned to sea by next day (continued next page)

10 Vagrant fur seals in South Australia Australian Mammalogy 163 Date Species ID Locality and geographical coordinates 16 May Admirals Arch, Cape du Couedic, K.I (36.06 S, E) 9 Jul Elliston (33.65 S, E) 12 Dec Cape du Couedic, KI (36.06 S, E) 16 v Admirals Arch, Cape du Couedic, K.I (36.06 S, E) 4 v Sceale Bay (32.97 S, E) 25 Jan Cape Gantheaume, KI (36.07 S, E) 20 Jul Glenelg (34.98 S, E) 27 Aug Twin Rocks, Head of Bight (31.47 S, E) 31 Aug Port Price (34.26 S, E) 10 Oct Seal Slide, Cape Gantheaume Conservation Park, KI (36.03 S, E) 21 v ra Creina (37.33 S, E) A ted by Gales et al. (1992). B ted by Kemper et al. (2008). Table 2. (continued) Data Collected or reported by Comment Images B. Chuan, DEWNR Alive Juvenile I. Quinn, Eyre Peninsula NRM and C. Bell, DEWNR Alive, rope removed from neck, returned to sea in good condition Adult male C. Baxter, KI Alive, in good condition, on rocks 1 m from a nursing NZ fur seal Adult male C. Baxter, KI Alive, in good condition; thermoregulating with hindflippers in a pool Juvenile A. Payne, Baird Bay Alive, in poor condition. On Sceale Bay beach, then at township on 5 v.; presumably same seal near Talia on 30 v Subadult male S. Goldsworthy, SARDI Alive, in good condition in NZ fur seal colony Subadult male, weight 63 kg A. Machado, AMWRRO Alive, in good condition, rehabilitated at AMWRRO, Torrens Is., released at Outer Harbor Juvenile, SL ~80 cm P. Rogers, SARDI Dead on beach, fresh Juvenile female A. Machado, AMWRRO Alive, rehabilitated at AMWRRO, Torrens Is., released at Port Elliott on or after 21 Sep Adult male K. Peters and J. Hicks, DEWNR Alive, in good condition. Fur seal of similar appearance seen at D Estrees Bay next day was likely same animal Subadult male G. Jackway, DEWNR Alive, in poor condition on beach, departed several days later

11 164 Australian Mammalogy P. D. Shaughnessy et al. (a) 130 E 132 E 134 E 136 E 138 E 140 E South Australia 32 S 32 S Point Westall 34 S Eyre Peninsula 34 S Tumby Bay Coffin Bay Adelaide Cape Carnot 36 S Kangaroo Is Victor Harbor 36 S N 38 S kilometres 38 S 130 E 132 E 134 E 136 E 138 E 140 E (b) 130 E 132 E 134 E 136 E 138 E 140 E South Australia 32 S 32 S Point Westall 34 S Eyre Peninsula 34 S Tumby Bay Coffin Bay Adelaide Cape Carnot 36 S Kangaroo Is Victor Harbor 36 S N 38 S kilometres 38 S 130 E 132 E 134 E 136 E 138 E 140 E Fig. 2. Map of South Australia showing locations for records of subantarctic fur seals: (a) specimens, (b) sighting records. Locations of several records are close to each other and are not shown separately. (2004) asa. forsteri and was collected at Middleton on 18 May The temporal distribution observed in this study, with 78% of the records between July and October, is similar to that observed by Gales et al.(1992), namely 81% of their 37 records for southern Australia were in the same months. During summer there were nine sightings of live subantarctic fur seals but no specimens. The temporal distribution of live animals may have been biased because researchers frequented NZ fur seal colonies during summer rather than at other times of the year. However, JM undertook field work at the NZ fur seal colony at Cape Gantheaume through most months for several years during the 2000s and recorded live subantarctic fur seals only during summer. The frequency of subantarctic fur seal records in SA appears to have increased slightly from nine records between August 1987 and August 1991 (Gales et al. 1992) at 2.3 records per year, compared with 86 records from September 1982 to vember 2012, at 2.9 records per year. This is likely to have resulted from the increased population size of this species in the South Indian Ocean, at St Paul and Amsterdam Islands and also Crozet Archipelago (Guinet et al. 1994). Other contributing factors are likely to be an increase in the reporting effort by DEWNR staff following the establishment of a stranding network (primarily for cetaceans) initiated by CMK, the increased number and awareness of people who frequent coastal SA, and from the increasing number of researchers working in NZ fur seal colonies in SA.

12 Vagrant fur seals in South Australia Australian Mammalogy Sightings Specimens Table 3. Body condition of subantarctic fur seals in South Australia. of records Jan Feb Mar Apr May Jun Jul Aug Sep Oct v Dec Fig. 3. Monthly distribution of records of subantarctic fur seals on the South Australian coast; open bars are sighting records, black bars are museum specimens.. of records Adult male Subadult male Adult female Juvenile Summer Autumn Winter Spring Fig. 4. Seasonal distribution of records of subantarctic fur seals of various age-classes on the South Australian coast; black bars are adult males, vertical hatching represents subadult males, horizontal hatching represents an adult female, open bars are juveniles.. of records Fig. 5. Records of subantarctic fur seals on the South Australian coast in 5-year intervals. Poor or emaciated Condition Good robust or excellent Total All records Specimens Sighting records Juveniles Specimens Sighting records The most frequent age-class among the records of subantarctic fur seals in SA was juveniles (83%), and almost allofthem(95%)occurredinwinterandspring.asimilar seasonal distribution for southern Australia was noted by Gales et al. (1992). Subantarctic fur seal pups wean between early September and late October (Kerley 1983), although some could be weaned by mid-august (Bester 1981). Juveniles that come ashore in southern Australia in winter could be pups that weaned early, as proposed by Gales et al. (1992) who interpreted the influx as a post-weaning dispersal coupled with assistance from the eastward-flowing West Wind Drift. During winter, they would be assisted by the eastward-flowing Leeuwin Current along the southern coast of Australia that averages up to cm s 1 (Middleton and Bye 2007). This preponderance of juveniles ashore in southern Australia contrasts with the situation in South Africa in the 1970s (Shaughnessy and Ross 1980) where far more adult males (60%) were recorded than juveniles (27%). The preponderance of males among the records (76%) is partly because nearly all adult and subadult animals recorded were male (13 of 14). These are easily recognised because of their striking colour pattern, whereas adult females are more likely to be confused with resident NZ fur seals and hence overlooked. Among juveniles, the high percentage of males (71%) reflects the usual pattern of dispersal in mammals by which males tend to disperse more than females (Greenwood 1980), which is seen in otariid seals (Page et al. 2003; Carlini et al. 2006). Barnacles are typically observed on animals that have recently returned ashore after extended periods at sea, and they drop off or are groomed off after the seal has been ashore for several days. Lepas australis has been reported on adult female subantarctic fur seals at Gough Island (Setsaas and Bester 2006) and on adult female Antarctic fur seals at South Georgia and Macquarie Island (Bonner 1968; Shaughnessy et al. 1988). L. australis is distributed in cold temperate to Antarctic waters in the Southern Hemisphere (Foster 1978). The juvenile male subantarctic fur seal observed with L. australis in this study is thus likely to have been in southern waters that are colder than those of SA. On the other hand, L. anatifera is distributed in tropical and temperate seas, indicating that the two juvenile subantarctic fur seals bearing this species were likely to have been in local waters for some time before going ashore. Antarctic fur seals The two records of Antarctic fur seals reported here appear to be the only ones for continental Australia and its nearby islands,

13 166 Australian Mammalogy P. D. Shaughnessy et al. other than a juvenile seen at Kangaroo Island that had been tagged at Macquarie Island (M.-A. Lea, in Wynen et al. 2000). There are two confusing records for New South Wales (Llewellyn et al. 1994) that refer to Antarctic fur seal (Arctocephalus tropicalis). We assume that these refer to subantarctic fur seals, as did Shaughnessy (1999). The paucity of records of Antarctic fur seals in waters of continental Australia (compared with records of subantarctic fur seals) is not surprising because the former species breeds on islands south of, or close to, the Polar Front, whereas the subantarctic fur seal breeds further north on islands north of the Polar Front (Goldsworthy et al. 2009). Furthermore, the Antarctic fur seal is similar in appearance to the resident NZ fur seal and is likely to be overlooked, whereas subantarctic fur seals are of striking appearance and easier to identify. Possible sources of animals Subantarctic fur seals are well known for their dispersing ability. Tagged individuals from Marion Island have been recorded at Heard Island (Bester 1989) and tagged individuals from Amsterdam Island have been sighted at Macquarie Island (Wynen et al. 2000; Goldsworthy et al. 2009). Subantarctic fur seals have been sighted at several localities where other fur seal species breed, including South Georgia (Payne 1979), Juan Fernandez Archipelago (Torres and Aguayo 1984), South America (de Moura and Siciliano 2007), South Africa (Shaughnessy and Ross 1980), Heard Island (Goldsworthy and Shaughnessy 1989) and New Zealand and some of its subantarctic islands (Taylor 1990). The nearest breeding colony of subantarctic fur seals to SA is at Macquarie Island (Fig. 6), 2700 km south-east of SA. It could be the source for animals seen on coasts of southern Table 4. Sightings of Antarctic fur seals in South Australia Geographical coordinates of localities are from the South Australian State Gazetteer. The column headed Species ID refers to the reliability of the species identification (see Methods). The abbreviation KI refers to Kangaroo Island Date Species ID Locality and geographical coordinates Age sex category Reported by Comment Images 2 Feb Cape du Couedic, KI (36.06 S, E) 14 Dec Cape Gantheaume, KI (36.07 S, E) Adult female PDS and C. Bald, DEWNR; Shaughnessy (1994) Svelte, pale pelage, broad foreflippers. In the non-breeding part of the Nautilus Rock colony of NZ fur seals Adult male JM On headland1 kmnorth-east of Cape Gantheaume, in nonbreeding part of the NZ fur seal colony 60 E 90 E 120 E 150 E 30 S Australia 30 S Ile Amsterdam Kangaroo Is Iles Crozet 60 S Heard Is Macquarie Is 60 S kilometres 60 E 90 E 120 E 150 E Fig. 6. Map of portion of the Southern Ocean showing possible sources of subantarctic and Antarctic fur seals in South Australia from islands in the Southern Ocean.

14 Vagrant fur seals in South Australia Australian Mammalogy 167 Australia, in particular the two specimens with reliability of species identification of 2 that might be hybrids. However, the population size at Macquarie Island is small, with fewer than 50 adult females in 2007 (Goldsworthy et al. 2009). A likely source is Amsterdam Island in the South Indian Ocean, 5200 km west of SA, where this species breeds and the population is increasing (Guinet et al. 1994). The genetic profile of two vagrant subantarctic fur seals housed at Melbourne Zoo indicated they were from Amsterdam Island (Wynen et al. 2000). Three tagged animals from there have been seen on Kangaroo Island: an adult in 2007 (M24981) and two juveniles in 1999 (C. Guinet in Wynen et al. 2000). The next closest breeding colony to SA is at Crozet Archipelago (7000 km south-west of SA), which is another possible source. For the Antarctic fur seals sighted in SA, a possible source is also at Macquarie Island, the nearest breeding colony, where they are more abundant than subantarctic fur seals (Goldsworthy et al. 2009). Furthermore, a juvenile tagged therewassightedatkangarooisland(m-a.leainwynen et al. 2000). Another possible source is Heard Island, 5200 km south-west of SA, where they breed in larger numbers than at Macquarie (Page et al. 2003). Concluding remarks It is unlikely that either of these fur seal species will set up successful breeding colonies on the SA coast because conditions ashore and in the sea are warmer than at the islands where they breed. The south coast of Tasmania with its cooler climate and records of subantarctic fur seals (PMTMMCP 2011) is a more likely place for them to attempt to breed. Nevertheless, the presence of adult male and adult female subantarctic fur seals in SA raises the possibility of hybridisation with NZ fur seals. Because archived tissue samples are available for most of the specimens lodged at the South Australian Museum, a study could be conducted to confirm this. The records reported here have been collected opportunistically without active searching or regular monitoring. Consequently, they must underestimate the incidence of vagrant fur seals ashore in SA. With increasing population size of these species, and the increase in public interest and awareness of marine mammals, the number of sighting records of vagrant fur seals in SA is likely to increase. Acknowledgements We thank the many people who reported sightings and collected specimens of vagrant fur seals on the coast of South Australia. We are especially indebted to staff of the Department of Environment, Water and Natural Resources (DEWNR) for their assistance. We thank A. Machado of Australian Marine Wildlife Research and Rescue Organisation for reports of several animals. We thank Dr C. Guinet, Centre National de la Recherche Scientifique, France, for information about the tagged fur seal from Kangaroo Island, and D. Snowball and P. Arthur of DEWNR for collecting the animal. We thank two anonymous reviewers for helpful comments on the manuscript. The following staff of the South Australian Museum are also thanked: L. Queale for the effort she made in obtaining seal records while Collection Manager, T. Segawa for producing the maps and T. Laperousaz for identifying barnacles. References Bester, M. N. (1981). Seasonal changes in the population composition of the fur seal Arctocephalus tropicalis at Gough Island. South African Journal of Wildlife Research 11, Bester, M. N. (1989). Movements of southern elephant seals and subantarctic fur seals in relation to Marion Island. Marine Mammal Science 5, doi: /j tb00339.x Bonner, W. N. (1968). The fur seal of South Georgia. British Antarctic Survey Scientific Reports 56, Brunner, S. (2004). Fur seals and sea lions (Otariidae): identification of species and taxonomic review. Systematics and Biodiversity 1, doi: /s x Carlini, A. R., Daneri, G. A., Casaux, R., and Márquez, M. E. I. (2006). Haulout pattern of itinerant male Antarctic fur seals (Arctocephalus gazella) at Laurie Island, South Orkney Islands. Polar Research 25, doi: /j tb00029.x Committee on Taxonomy, Society for Marine Mammalogy (2012). List of marine mammal species and subspecies. Society for Marine Mammalogy. Available at: [accessed 17 December 2013]. Condy, P. R. (1978). Distribution, abundance, and annual cycle of fur seals (Arctocephalus spp.) on the Prince Edward Islands. South African Journal of Wildlife Research 8, de Moura, J. F., and Siciliano, S. (2007). Straggler subantarctic fur seals (Arctocephalus tropicalis) on the coast of Rio de Janeiro State, Brazil. The Latin American Journal of Aquatic Mammals; (LAJAM) 6, doi: /lajam00114 Dennis, T. E. (2005). Australian sea lion survey (and historical) records for South Australia. Report to the Wildlife Conservation Fund. Department for Environment and Heritage, South Australia. Foster, B. A. (1978). The marine fauna of New Zealand: barnacles (Cirripedia: Thoracica). New Zealand Oceanographic Institute Memoir 69, Gales, N. J., Coughran, D. K., and Queale, L. F. (1992). Records of subantarctic fur seals Arctocephalus tropicalis in Australia. Australian Mammalogy 15, Goldsworthy, S. D. (1992). Maternal care in three species of southern fur seal (Arctocephalus spp.). Ph.D. Thesis, Monash University, Melbourne. Goldsworthy, S. D. (2008a). Antarctic fur seal Arctocephalus gazella (Peters, 1875). In The Mammals of Australia. 3rd edn. (Eds S. Van Dyck and R. Strahan.) pp (Reed New Holland: Sydney.) Goldsworthy, S. D. (2008b). Subantarctic fur seal Arctocephalus tropicalis (Gray, 1872). In The Mammals of Australia. 3rd edn. (Eds S. Van Dyck and R. Strahan.) pp (Reed New Holland: Sydney.) Goldsworthy, S. D., and Shaughnessy, P. D. (1989). Subantarctic fur seals Arctocephalus tropicalis at Heard Island. Polar Biology 9, doi: /bf Goldsworthy, S. D., and Shaughnessy, P. D. (1994). Breeding biology and haul-out pattern of the New Zealand fur seal, Arctocephalus forsteri, at Cape Gantheaume, South Australia. Wildlife Research 21, doi: /wr Goldsworthy, S. D., McKenzie, J., Page, B., Lancaster, M. L., Shaughnessy, P. D., Wynen, L. P., Robinson, S. A., Peters, K. J., Baylis, A. M. M., and McIntosh, R. R. (2009). Fur seals at Macquarie Island: post-sealing colonisation, trends in abundance and hybridisation of three species. Polar Biology 32, doi: /s y Greenwood, P. J. (1980). Mating systems, philopatry and dispersal in birds and mammals. Animal Behaviour 28, doi: /s (80) Guinet, C., Jouventin, P., and Georges, J.-Y. (1994). Long term population changes of fur seals Arctocephalus gazella and Arctocephalus tropicalis on subantarctic (Crozet) and subtropical (St. Paul and Amsterdam) islands and their possible relationship to El Niño Southern Oscillation. Antarctic Science 6, doi: /s