University of Groningen

Mirrors in ice van Franeker, Jan Andries

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Download date: 26-09-2021 1. INTRODUCTIONAND OUTLINE

Fulmarinepetrels the Diomedeidae(albatrosses), the Petrelsare seabirdsthat are characterisedby the (fulmarine petrels, gadfly petrels, prions and possessionof one or two homy tubes on top of the shearwaters), the Hydrobatidae (storm petrels), bill covering their nostrils. The petrels, or and the Pelecanoididae(diving petrels). There are tubenosedseabirds (: or considerable disagreements on evolutionary Tubinares)are probably an old group that relationships (phylogeny) in the tubenoses,which evolved in the southem hemisphere and that is leads to frequent changes in the systematics(the relatedto the penguins (Order: Sphenisciformes). ordering in groups and related system of names But unlike the penguins, petrels developed that should be used). Followed here are the remarkableflying skills that enablethem to utilize subdivisions and names as used by Warham in his the mostdistant parts of the world's oceans.Four extensive work on the petrels (Warham 1990, families are distinguished (Warham 1990): 1996).

ORDER FAMILY SABFAMILY/GROAP

FULMARINE PETRELS Fulmarínae fulmarine stormvosels

PRIONS (whale ) P ac hyp t i I a./HaI o baen a walvisstormvogels OTHER PETRELS Procellartidae storm-& pijlstormvogels PETRELS Procellaiífurmes stormvogelachtigen

SHEARWATERS Puírtnus/C alo ne c t ri s/P roce I I dr id pijlstormvogels

FIJLMARINE PETRELS ( Fulmarinae) Englishname mass (g) scientific name Dutch name distribution NorthernFulmar 725 Fulmarus glacialis Noordse Stormvogel Arctic SouthemFulmar 850 Fulmarus glacialoides Zuidelijke Stormvogel Antarctic Antarctic Petrel 690 Thalassoicaantarctica Antarctische Stormvogel Antarctic CapePehel 475 Daption capense KaapseStormvogel Antarctic SnowPetrel 360 Pagodroma nivea Sneeuwstormvogel Antarctic Northern 4500 Macronecteshalli Noordehjke Reuzenstormvogel Antarctic SouthemGiant Petrel 4500 Macronectesgiganteus Zuidelijke Reuzenstormvogel Antarctic

Mírrors in lce introduction and outline range c Within the family of the Procellariidae, the hemisphere , the Northern , is Petrel ( fulmarinepetrels seem a somewhatodd collection thought to be an offspring of ancestral Southern extends ofseven species,in five genera.They lack unique (Fulmarus glacialoides) having reached breedinl morphological or anatomical group-characters the north Pacifrc using the cold Humbold Current location and vary widely in size, shape coloration and in Pleistoceneglaciation periods (Voous 1949). Antarcti habits. Neverheless, they have usually been From the Pacific, Northern Fulmars have The Catr grouped as a subfamily, the Fulmarinae (Warham successfullycolonized all of the Arctic and now Pintado 1990). In recent yeaÍs mitochondrial DNA have separate and morphologically híghly breeding analyseshave confirmed the common ancestry of variable populations in the North Pacific and islands d these seven species(Nunn 1994). According to North Atlantic (Fisher 1952; van Franeker & widely d the same study, Blue Petrel Qlalobaena Wattel 1982). ZeaTand, caerulea), Prions (Pachyptila spp.) and All other fulmarines are restricted to the cold The Sor surprisingly the Kerguelen Petrel (Lugensa climate zones of the southem hemispere (Watson dishibuti, kerguelensis or Pterodroma kerguelensis) aÍe et al. t97l). enclosed close relatives that could be included under a The name of the Northern Giant Petrel non bree broader interpretation of the name fulmarines (Macronectes halli) is somewhat confusing, as it disperses (Nunn 1994). is a subantarctic southern hemisphere bird. Its The real c The sevenfulmarines in the old sensehave Petrel (Tl, a relatively large nasal tube, and most of (Pagodrot them nest in the open, and not in burrows winter seaice boundary live in c Antarctic Polar Fronl like the majority of the other Southern Procellariidae. The fulmarines aÍe also colonies il renowned for their habit of spitting the interior stomachoils in defense against enemies. nest on hi This smelly habit explains their name, mountain p which goesback to old Viking languages. Whether When Vikings settled on Iceland and otherwise, St.Kilda, they found a breeding bird that fulmarines was new to them, the Northem Fulmar ending as1 Fulmarus glacialis. Its colouration looked strategies c a bit like that of gulls, and that is why they many place named it 'ful-mar' which means as much Antarctic rr as 'foul gull' or'stinking gull' (Lockwood this Antarct res4). research el affecting to1 Cold climate specialists which coul A pronounced characteristic of all understand, fulmarine petrels is that they are adaptedto change". Wi the coldest of the world's environments. that the fulm They breed from extreme high-Arctic key role in r places in the northern hemisphere, to the have been al inland mountains of the Antarctic. No ANTARCTICA. The Antarctic Polar Front is the zone where together the warrner subantarctic water other bird group breeds beyond 80o of cold Antarctic surface water and studies in o; bounce against each other and submerge. It creates a sharp latitude on both poles. They are so highly petrels are temperature gradient all around the Antarctic, conveniently provide specialisedto the cold, that they are unable used as the natural northern border of the the Antarctic area. a Antarctic to live in subtropicaland tropical climate Politically, the Antarctic is defined as the area south of 60o of eco zones. Only over cold currents, some of latitude. The light gÍey areashows the extent of sea ice during the fulmarines occasionally venture into winter, which melts to about 20oÁof that size in mid suÍImer. Mirrors in ic warÍner climate zones during the non- Dark areas along the Antarctic coast show ice shelves, the Populationso breeding seasons. The single northern floating outer margins of the Antarctic icecap. many million

Mírrors in ice Introduction and outline range overlaps with that of the Southem Giant 1984;Woehler 1997;Croxall et al. 1984;Croxall Fulmar, is Petrel (M. giganteus) but the northern specres et al. 1995) that depend entirely on the small bits Í Southem extends into more temperate zones, whereas of ice-free land for breeding and largely or ng reached breedinglocations of the southem one include completely on the Southem Ocean marine rld Current locations along the continental coasts of environment for their food. ous 1949). Antarctica(Bourne & Warham 1966). The most striking feature of the Antarctic marine lars have The , also known as Cape Pigeon or system is the seasonal sea ice cover (Bonner & c and now Pintado Petrel (Daption capense), has a wide Walton 1985,SCAR 1993).In late winter, seaice ly highly breeding distribution from many subantarctic in the SouthernOcean grows to a surface of more acific and islandsdown to the Antarctic coast and disperses than 20 million lan2. That is more than half of the raneker & widely during winter, reaching the coastsof New Antarctic waters (38 million km2 south of the Zealand,South Africa and South America. Antarctic Polar Front) and effectively doubles the r the cold The Southem Fulmar has a true Antarctic ice surfaceof the continent itself. During summer, :e (Watson distnbution, as it does not breed out of the area the sea ice melts to a residual surface of about 4 enclosedby the Antarctic Polar Front. During the million km2. The ecological importance of the rnt Petrel non breeding seasons,part of its populations continuous waxing and waning of this huge sea rsing,as it dispersesto the north ofthe Polar Front. ice zone is probably still underestimated and : bird. Its The real cold climate championsare the Antarctic poorly understood(Legendre et al. 1992). Seaice --I Petrel (Thalassoicaantarctica) and does play a critical role in the life cycle of (Pagodroma nivea). Throughout the year they Antarctic Wltt (Euphausia superba) (Smetacekel : boundaryI live in close connection with sea-ice in the al. 1990,Hempel 1991). is thoughtto be the : PolarFront Southem Ocean, and part of their breeding key organism in the Antarctic food web, I I colonies is situated hundreds of kilometers into channeling primary production directly to all I the interior of the Antarctic continent where they birds, sealsand whales (Bonner & Walton 1985, I nest on high altitudes on nunataks, the isolated El-Sayed 1985). Therefore krill was and is the I mountainpeaks that prohude out the icecap. main study object in past (El-Sayed 1994) and I Whether by shared climatic preference or current major research programs (CCAMLR \'l otherwise, I have been fascinated by the 1991, 7997; Hofmann 1998). The krill to ice \l fulmarinessince my days of studenthood.Never linkage suggestshigh risks of global warming to rl ending astonishment over the extreme life the Antarctic ecosystem. Much of the current strategiesof these species has brought me to effort in Antarctic researchis thereforefocused on )i many placesand resulted in a job dedicatedto the effects of global climate change on Antarctic il Antarctic research. The general assignment for sea ice and the associatedecosystem (SCAR /Island this Antarctic position was to conduct "applied I 993,Goodwin 1997, El-Sayed 1998). I research enhancing understanding of factors The different speciesof fulmarine petrels,because I affecting top predator populations in Antarctica" of their breeding at the extreme limit of the which could be rephrased as "to measure, Antarctic climate zone and becauseof their strong understand,and predict effects of environmental linkage to the seasonal sea ice system, will change".With such freedom,it is not surprising undoubtedly be among the very first organismsto _l that the fulmarine petrels have continued to play a be affected by climate change. Changes in 'mirrored' key role in many of the Antarctic projects that I climate and ice conditions will be by ItlltÉrcltt have been allowed to conduct. This thesis brings variations in petrel breeding success,survival, and re where together the major results of those Antarctic shifts in distnbution ranges. By simultaneously ic water studies in order to demonstrate that fulmarine studying the different speciesof fulmarines, each a sharp petrels are not an isolated study-object, but of them with different ecological adaptationsto 'eniently provide a powerful tool in understanding the Antarctic environment, one obtains a number rtlc area. Antarcticecosystems. of sensitive and early indicators for a wide range rf 60o of e during of potential changesin the Antarctic environment snÍnmer. Mirrors in ice and the way in which ecosystemsare functioning. ves, the Populationsof Antarctic fulmarine petrels number This thesis aims to show that such a mirror many millions of individuals (e.g. Ainley et al. function is not just a theoretical assumption.

Mirrors in ice Introductionand outline Researchquestion populations and has Cape Petrel and Antarctic Ship-basedstr It would be incorrect to suggest that the Petrel among the target speciesfor its monitoring first marine environmental indicator or mirror function of progïam (CCAMLR 1991, 1997). Polarstem Stur fulmarines above has been the key research Further projects used in this thesis were not with old stud questionon which all projectsfor this thesishave sharply or exclusively focused on the research program neces sharplyfocused. question of fulmarine petrel ecology, but were methodology My original interest in the fulmarines concerned more a matter of adapting to political preferences compared to (BIOMASS: the rather academicquestion on the and or taking advantage of logistic opportunities. A zoogeography of the fulmarine petrel group. comparative study on Cape Petrels in the Antarctic Systr Having studied morphological variation of the Peninsulaarea could be included becauseof the discussedin cl Northern Fulmar in detail (van Franeker &'Wattel Dutch Antarctic Expedition l990l9l to King of the whole 1982), the hope for resolving phylogenetic George Island (Anonymous 1990). Further comparison be relationships in the fulmarines using just knowledge on Cape Petrels in that area was primary proó morphological characters was small. As achieved by supporting data analyses of a Chapter 4 con ecological characteristicscould hold additional Tsjechian researchproject. My first ship-based top predator clues on phylogeny, my first Antarctic project project was, to be honest, mainly startedto flag production da aimed at a comparative study of the ecology of Dutch presence in international coordinated Chapters 5 anr the four smaller species of Antarctic fulmarines researchefforts. The cruise was not specifically and ecology investigates th (SouthernFulmar, Antarctic-, Cape- , and Snow focused on top predator research, let alone Petrel).The comparativeaspect was best served fulmarine petrel research. However, the study the Antarctic by studying these birds simultaneously on a proved to be extremely productive and was Pachyptila sp. locationwhere they co-occurredand sharednest followed up by further projects on the German seem a bit t locationsand foraging areas. Such location was icebreaker Polarstem. Most of these weÍe However, accr found on Ardery Island and the Australian conducted as part of the Joint Global Ocean Flux also fulmarine Antarctic Division was prepared to support the Study JGOFS (IGBP & SCOR 1992) which aims demonstrateth project. Researchquestion, from a phylogenetic to quantiS'carbon fluxes in marine environments small spatial i background,for the project was thus: "What are in relation to atmospehericCO2 levels and global dynamic ocet the ecologicaladaptations that allow four related climate change.By bits and pieces,the combined makes the fir andsimilar species of petrelsto breedon the same results of studies at sea have considerably combining var location and to exploit the same environment?" improved my understanding of the marine pelagic ecolol Such a researchquestion actually requires the ecology of fulmarine petrels, crucial to the estimate of tl sametype of datacollection as what is neededfor interpretation of study results obtained on land. Chapter 7 is the environmental indicator function, with the The marine projects also provided me with a Antarctic I only difference that the latter more explicitly much broader awareness of how Antarctic top knowledge f needsto look into possible changesover time m predatorsmirror the ecosystemshidden below the research from relation to environmental variability. Being surface and how important the sea ice system is. paper is one employedto conduct applied Antarctic research, For this reason, parts of the thesis deviate from Antarctic seab the emphasis of follow up projects on Ardery the central question on the fulmarines. SCAR Bird Island shifted to the indicator aspect.This was prerequisite enhanced by the structure of the Netherlands Outline of the thesis providing advi AntArctic Program NAAP (CAO 1994) which The thesisconsists of two sections,the first part issues(SCAR' doesnot operateits own research station or ship. (Chapters2-7) dealingwith results from ship Research). This means that any project has to adapt to the basedresearch, and the secondpart (Chapters8- Land-based objectives of foreign pÍogÍams to obtain their 14) with resultsof land-basedstudies of the maps the birc logistic support. The Australian commitment to fulmarine petrels. Ardery Island the Ardery project has increasinglybeen based on Considerableproportions of my time, I have been with which it the EcosystemMonitoring Programof CCAMLR wrestling with methodologicalissues to ensureor under the ,\ (Commission on the Conservation of Antarctic search for comparability of data in larger methodologici Marine Living Resources). CCAMLR aims to frameworks. Chapters 2, 9 and large parts of sexing live fu collect information on potential or actual effects chapter 11 are attempts to clarify methods or to measurements of (l

Mirrors in lce Introduction and outline I Antarctic Ship-basedstudies. Chapters 2 and3 concernthe sexes lack distinguishing characters. Using all monitoring first marine science project, the European data from Northem Fulmars and the southern PolarsternStudy (EPOS 1988-89).My breaking fulmarines we searchedfor a method that can be were not with old study methods from the BIOMASS used in any fulmarine petrel population without re research programnecessitated an explanation of why new the need to kill birds for sexedreference material. methodology was needed, and how results Chapter 10, first , but were authored by Karel Weidinger, lreferences comparedto those collected by old methods gives and example of applicability of the method (BIOMASS= tunities. A BiologicalInvestigations of Marine and refines it for Cape Petrels from different ls in the Antarctic Systemsand Stocks). EPOS results are populations. ruseof the discussedin chapter 3 and focus on distributions The large chapter 11 is the link between marine to King of the whole top predator community and the and landbasedresearch and extensively discusses l. Further comparisonbetween their food requirements and foods and foraging ecology of the fulmarines of area was primary productivity measured in the water. Ardery Island, and comparesthe speciesin detail. rses of a Chapter4 continues on this issue by comparing Diet study methods are extensively discussedin ship-based top predator requirements with primary order to integrate our data with published ted to flag production data from more research cruises. information. Chapter 12, ftrst authoredby Sophie oordinated Chapters5 and 6 deal with pelagic distributions Creet, gives valuable comparative data on the pecifically and ecology of separate species. Chapter 5 diets of Cape Petrels on King George Island. let alone investigatesthe links between frontal physics in Chapter 13 is also related to diets, but discusses the Antarctic Polar Front (APF) and prions the non-food plastic items that are ingested by the study 'mirror' and was Pachyptilasp. The bird speciesand the area may petrels. It was included to show that the LeGerman seem a bit out of the focus of this thesis. function of petrels in our studies extends to the ese were However, according to Nunn (1994) prions are signalling and measuring of global levels of man- )ceanFlux also fulmarines, and the paper was included to made pollutants. The best example of this latter rhich aims demonstratethe mirror function of seabirdson the function is of course the completed study of Nico 'ironments small spatial and temporal scales that occur in van den Brink on monitoring organochlorine md global dynamic oceanic frontal systems. Chapter 6 pollutants (van den Brink 1997). Finally, chapter combined makes the first step to the real fulmarines, by 14 demonstrates the importance of long term combiningvarious data into a descnption of the ecological and environmental data collection in a nsiderably 'mirror' e marine pelagic ecology of the Antarctic Petrel and an spectacular example of effects of ll to the estimateof the size of a regional population. environmental changeon birds. I on land. Chapter7 is a species review paper on the The thesis concludes with a short English re with a Antarctic Petrel population, integrating summary with concluding remarks, an extensive arctic top knowledge from landbased and sea-going Dutch suïrmary and finally acknowledgements. below the researchfrom literature and my own data. This systemis. paper is one in a series on populations of References ,iate from Antarctic seabirds,compiled by members of the Ainley, D.G., O'Connor,E.F. & Boekelheide,R.J. SCAR Bird Biology Subcommittee as a 1984.The marine ecology of birdsin theRoss Sea, prerequisite to the subcommittees task in Antarctica.Ornithological Monograph s 32: l-91. providingadvise in conservationand management Anonymous 1990. The Netherlands Antarctic ExpeditionKing GeorgeIsland, Arctowski Station, issues(SCAR:Scientifi c Committee on Antarctic lrst part December1990 - February1991.. TheNetherlands Research). ship Antarctic Researchnr. IV. Ministry of Foreign Lpters8- Land-based studies. Chapter 8 describes and Affairs and the NetherlandsMarine Research the maps the bird populations in the study area on Foundation,The Hague. 33 pp.. fudery Island and on the nearby Odbert Island Bonner,W.N. & Walton,D.V/.H. (Eds.) 1985.Key havebeen with which it forms a Specially Protected Area Environments:Antarctica. ruCN, PergamonPress ensureor under the Antarctic Treaty. Chapter 9 is a Oxford.38lpp. Bourne,W.R.P.; Warham,J. 1966. Geographical in larger methodologicalpaper dealing with methods for variation in the Giant Petrels of the senus parts sexinglive fulmarine petrels on the basis of their of Macronectes..Ardea 54: 45-67. Lodsor to measurements.Many studies on petrel ecology tterature. require sex identification in the field, but the

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Mirrors in rce Inffoductionand outline