BirdLife South Africa – Species factsheet Macaroni Penguin Eudyptes chrysolophus 2013 Regional Red List status Vulnerable 2000 Regional Red List status Near threatened 2013 Global Red List status Vulnerable Reasons for change in regional status Measured change in population size Family Spheniscidae Species name author Brandt 1837 Population size 279 000–290 000 pairs Distribution size (AOO) N/A Regional endemic (South Africa, Lesotho and No Swaziland)

Justification Conservation status was assessed from the rate of decrease in estimates of numbers of Macaroni Penguins Eudyptes chrysolophus breeding at Marion Island against the IUCN criterion A2, because the decrease may not have ceased, the causes of decrease are not fully understood and they may not be reversible (Birdlife International 2000). The metric used was the percentage decline in three generations or 10 years (whichever is greater). The average generation for Macaroni Penguins is 11.4 years (M Taylor, BirdLife South Africa, in litt.). The estimates of numbers of Macaroni Penguins breeding at Marion Island decreased by 41% between 1976/77 and 2012/13 (see below), equivalent to a decrease of 38% in three generations, which merits a classification of Vulnerable. The modelled decrease in numbers breeding between 1994/95 and 2012/13 was 33% in 18 years (see below), which is equivalent to 63% in three generations. This suggests that the rate of decrease increased since the mid 1990s to a level that would justify a rating of Endangered. However, the decrease since the mid 1970s, a period equivalent to about three generations, has been used to assign the Red List status.

Reasons for inclusion in the assessment The species was listed as Near Threatened in the 2000 regional Red List assessment (Barnes 2000), and is regarded as globally Vulnerable (BirdLife 2013).

Taxonomic note The Royal Penguin E. schlegeli, which is found in the south-west Pacific Ocean, was previously treated as a subspecies of the Macaroni Penguin. The two species are similar, except for the amount of blue-black feathering on their cheeks and throat. Whereas these feathers are dark in macaroni penguins, they are white or pale grey in Royal Penguins, which also exhibit yellow-tinged feathers between the bill and eye. There are differences in measurements between these two penguins (Marchant and Higgins 1990, BMD, pers. obs.). The Macaroni Penguin is monotypic (Del Hoyo et al. 1992).

Identification Length 71 cm; weight 5–6 kg. Macaroni Penguins are blue-black on their backs and white ventrally. Long orange, yellow and black feather plumes extend from the forehead along the sides of the head and meet above the eyes, forming a broad, loose crest. There is an obvious pink fleshy gape. The bill (larger in males than females) is red-brown, eyes dull red, legs and feet pink. Juveniles are similarly marked except for greyer eyes, chin and throat and the lack distinct crest feathers. A raucous, loud braying call is made at breeding colonies. The species is usually silent at sea (Marchant and Higgins 1990, Del Hoyo et al. 1992).

Distribution The Macaroni Penguin has a more southerly distribution than most other crested penguins, breeding between about latitudes 45° and 65°S in the western Indian (Heard and McDonald, Kerguelen, Crozet and Prince Edward groups of islands) and South Atlantic (Bouvet, South Sandwich, South Georgia, South Orkney, South Shetland and Falkland groups of islands and in small numbers at islands off the Antarctic Peninsula) oceans. It also extends into the south-east Pacific Ocean along the coastline of southern Chile. The largest colonies are north of about 55°S (Marchant and Higgins 1990).

Population justification

In 2012/13, it was estimated that 266 971 pairs of Macaroni Penguins bred at Marion Island compared to 433 723 pairs in 1994/95 (Crawford et al. 2003, 2009, Department of Environmental Affairs, unpublished information). The decrease was best fitted by a linear regression, which suggested that the population in 2012/13 was 278 000 pairs (see below). An estimated 12 000 pairs bred at Prince Edward Island in 2008/09 (Crawford et al. 2009). Therefore, the overall population at the Prince Edward Islands is of the order of 279 000–290 000 pairs.

Trend justification From counts of small colonies and at two large colonies (Bullard Beach and Kildalkey Bay) measurements of areas occupied and densities of nests, or in 1974–1977 counts of birds moulting made on aerial photographs, it was estimated that there were 449 892 pairs of Macaroni Penguins at Marion Island in 1974–1977 (Siegfried et al. 1978), 405 084 pairs in the 1983/84 breeding season (Watkins 1987) and 433 723 pairs in 1994/95 (Crawford et al. 2003). It was then estimated that by 2008/09 numbers breeding had decreased by 32% (Crawford et al. 2009). By 2008/09, there were significant decreases in numbers breeding at the small colonies and at both of the large colonies, where densities of nests also decreased significantly (Crawford et al. 2009). In 2012/13, 266 971 pairs bred at Marion Island (see above). Therefore, between 1976/77 and 2012/13, a period of 36 years, estimates of the numbers breeding decreased by 41%. Numbers breeding at Marion Island were available for each season between 1994/95 and 2012/13 (Crawford et al. 2003, 2009, Department of Environmental Affairs, unpublished information). In this period, the overall decrease was best modelled by linear regression (n = 19, r = 0.861, p < 0.001), which estimated the population in 1994/95 to be 416 000 pairs and that in 2012/13 to be 278 000 pairs, a decrease of 33% in 19 years.

Ecology During October, Macaroni Penguins return to breed at Marion Island, in colonies which range from small numbers of birds to the two large colonies that presently each hold > 100 000 pairs and formerly held > 200 000 pairs (Crawford et al. 2003, 2009). Breeding areas usually have little or no vegetation due to erosion by birds (BirdLife International 2013). Traditional paths are used to access colonies (BMD, pers. obs.). At Marion Island, Macaroni Penguins mostly breed for the first time when four years old (Crawford et al. 2003). Males arrive for breeding several days earlier than females (Crawford et al. 2006). Both sexes undertake substantial fasts during courtship and breeding (Marchant and Higgins 1990). Pairs lay two eggs from late October to mid November, but only the larger second (B) eggs produce fledged chicks, which leave the island by the end of February (Williams 1980a, 1980b, Crawford et al. 2003). Incubation takes 34–39 days (Williams 1981). After breeding, Macaroni Penguins go to sea to fatten for about a month before returning to islands to moult, when they remain ashore for 20–30 days to replace all their feathers (Williams et al. 1977, Brown 1986). They then depart to over-wintering grounds to regain condition. At Marion Island, Macaroni Penguins feed mainly on crustaceans (euphausiids and amphipods), fish (mostly myctophids) and cephalopods (Brown and Klages 1987, Crawford et al. 2003, Pichegru et al. 2011). The

composition of prey changes as chicks develop (Brown and Klages 1987). The median trip duration of birds feeding chicks was 22.8 hours and most dives were < 10 m (Pichegru et al. 2011).

Threats Inadequate breeding success has been a factor in the decrease of Macaroni Penguins at Marion Island (Crawford et al. 2003). Overwintering conditions are thought to influence the proportions of Macaroni Penguins skipping breeding and breeding success; feeding conditions during breeding also may influence breeding success (Crawford et al. 2006). Global change may have altered the availability of prey to Macaroni Penguins (Crawford et al. 2003). In winter, Macaroni Penguins from Marion Island forage mostly to the south of the island, sometimes approaching the edge of the sea ice (J-B Thiebot et al., unpublished information). No fisheries target the main prey of Macaroni Penguins at the Prince Edward Islands, but should they be introduced they could decrease prey availability (BirdLife International 2013). Long-line fisheries at winter feeding grounds have inflicted limited incidental mortality on Macaroni Penguins (BMD, unpublished information). Recoveries of fur seals Arctocephalus spp. at the Prince Edward Islands (Hofmeyr et al. 2006, Bester et al. 2009) could increase competition for prey and lead to increased predation of penguins by seals. Increasing seal herds also could block access to breeding sites thereby inhibiting colony growth, as happened at Bouvet Island (Isaksen et al. 1997). Giant petrels Macronectes spp. and Subantarctic Skuas Catharacta antarctica and Lesser Sheathbills Chionis minor also inflict mortality (BMD, pers. obs.). In 1992, 1993 and 2004 disease, including avian cholera Pasteurella multocida, killed substantial numbers of Macaroni Penguins at Marion Island (Cooper et al. 2009). Oil spills could kill large numbers of Macaroni Penguins.

Conservation measures underway At Marion Island, there are strict guidelines in place for offloading diesel. Poultry products supplied to the over-wintering teams at Marion Island, or used during relief voyages to the island, do not contain bones and are irradiated in order to reduce the risk of introducing avian diseases. A contingency plan is in place to guide responses to outbreaks of disease at the Prince Edward Islands. No access is allowed to Prince Edward Island except for bona fide research and conservation purposes every 4–5 years. The Prince Edward Islands Marine Protected Area was proclaimed in 2013.

Research questions It will be difficult but important to understand how food availability for Macaroni Penguins fluctuates at both summer and winter feeding grounds, and to what extent the penguins compete with other predators for prey. Further information is required on the foraging distributions, especially during breeding, and on adult and immature survival and age at breeding of Macaroni Penguins at Marion Island. Levels of predation attributable to predators should be reassessed. Proposed implantation of transponder tags may assist in determining both population and demographic parameters. Further counts of Macaroni Penguins are required for Prince Edward Island.

Conservation measures proposed Monitoring of population numbers and demographic, diet and condition parameters at Marion Island should be continued. A population census at Prince Edward Island should be undertaken at intervals of about five years. Best practice guidelines for reducing risk of disease outbreaks at islands are being developed by the Agreement on the Conservation of Albatrosses and Petrels. Once available, these guidelines should replace those presently operating at the Prince Edward Islands. Overwintering teams should be given basic training in the rescue and rehabilitation of oiled penguins. The impact of any proposed fishery on the prey of Macaroni Penguins should be carefully assessed before such a fishery is allowed. Consideration should be given to declaring marine protected areas at important feeding grounds of Macaroni Penguins.

References Barnes KN (ed.) 2000. The Eskom Red Data Book of Birds of South Africa, Lesotho and Swaziland. Johannesburg: BirdLife South Africa. 169 pp. Bester MN, Ryan PG, Visagie J. 2009. Summer survey of fur seals at Prince Edward Island, southern Indian Ocean. African Journal of Marine Science 31: 451–455. Birdlife International. 2000. Threatened Birds of the World. Lynx Edicions, Barcelona. BirdLife International. 2013. Species factsheet: Eudyptes chrysolophus. Downloaded from http://www.birdlife.org on 27/05/2013. Brown CR. 1986. Feather growth, mass loss and duration of moult in Macaroni and Rockhopper Penguins. Ostrich 57: 180–184. Brown CR, Klages NT[W]. 1987. Seasonal and annual variation in the diets of Macaroni (Eudyptes chrysolophus chrysolophus) and Southern Rockhopper (E. chrysocome chrysocome) penguins at sub- Antarctic Marion Island. Journal of Zoology, London 212: 7–28. Cooper J, Crawford RJM, de Villiers MS, Dyer BM, Hofmeyr GJG, Jonker A. 2009. Disease outbreaks among penguins at sub-Antarctic Marion Island: a conservation concern. Marine Ornithology 37: 193– 196. Crawford RJM, Cooper J, Dyer BM, Greyling MD, Klages NTW, Nel DC, Nel JL, Petersen SL, Wolfaardt AC. 2003. Decrease in numbers of the Eastern Rockhopper Penguin Eudyptes chrysocome filholi at Marion Island, 1994/95–2002/03. African Journal of Marine Science 25: 487–498. Crawford RJM, Dyer BM, Cooper J, Underhill LG. 2006. Breeding numbers and success of Eudyptes penguins at Marion Island, and the influence of mass and time of arrival of adults. CCAMLR Science 13: 175–190. Crawford RJM, Whittington PA, Upfold L, Ryan PG, Petersen SL, Dyer BM, Cooper J. 2009. Recent trends in four species of penguin at the Prince Edward Islands. African Journal of Marine Science 31: 419–426.

Del Hoyo J, Elliott A, Sargatal J (eds). 1992. Handbook of the Birds of the Worls. Vol. 1. Ostrich to Ducks. Lynx Edicions, Barcelona. Hofmeyr GJG, Bester MN, Makhado AB, Pistorius PA. 2006. Population changes in Subantarctic and Antarctic Fur Seals at Marion Island. South African Journal of Wildlife Research 36: 55–68. Isaksen K, Hofmeyr GJG, Dyer BM, Naestvold A, Mehlum F, Gjertz I, Bakken V, Huyser O. 1997. Preliminary results from CEMP-monitoring of Antarctic Fur Seals, Chinstrap Penguins and Macaroni Penguins at Bouvetoya 1996/97. Report to CCAMLR WG-EMM 1997: 26 pp. Marchant S, Higgins PJ (Co-ordinators). 1990. Handbook of Australian, New Zealand and Antarctic Birds. 1. Ratites to Ducks. Melbourne, Oxford University Press. Pichegru L, Ropert-Coudert Y, Kato A, Takahashi A, Dyer BM, Ryan PG. 2011. Diving patterns of female Macaroni Penguins breeding on Marion Island, South Africa. Polar Biology: DOI 10.1007/s00300-010- 0950-5. Siegfried WR, Williams AJ, Burger AE, Berruti A. 1978. Mineral and energy contributions of eggs and selected species of seabirds to the Marion Island terrestrial ecosystem. South African Journal of Antarctic Research 8: 75–87. Watkins BP. 1987. Population sizes of King, Rockhopper and Macaroni Penguins and Wandering Albatrosses at the Prince Edward Islands and Gough Island, 1951–1986. South African Journal of Antarctic Resesearch 17: 155–162. Williams AJ. 1980a. Offspring reduction in Macaroni and Rockhopper Penguins. Auk 97: 754–759. Williams AJ. 1980b. The clutch size of Macaroni and Rockhopper Penguins. Emu 81: 87–90. Williams AJ. 1981. The laying interval and incubation period of Rockhopper and Macaroni Penguins. Ostrich 52: 226–229. Williams AJ, Siegfried WR, Burger AE, Berruti A. 1977. Body composition and energy metabolism of moulting Eudyptid penguins. Comparative Biochemistry and Physiology 56A: 27–30.

Further web sources: http://www.birdlife.org/datazone/species and http://www.adu.org.za. Assessor: Bruce Dyer and Robert Crawford Reviewers: Peter Ryan, Martin Taylor and Ross Wanless Recommended citation: pending...