TREE vol. 7, no. 6, June 1992 reviews 1277 ---- 119911 Nature 353. 255-258 ArtifiCial Ufe fSiJntiJ Fe In5tirute Studie5 in ISdnta Fe Institute Studies in the Sciences 10 Menge. B.A. and Olson.....M. (1990) the Sdence5 of Complexity. Vol. 61 of Complexity. Vol. 61 (Langton. CG.. ed.l, Trends frol. CVf)/. 5. 52_57 ILangton. C.G" ed.l. pp. 105-124. pp. 63-11. Addison-Wesley 11 .... lIeo, T.r.H. and Sfarf, T.B. 11982) Addi&an-Wesley H Sklar. F.H. and COStanza. R.lllXlOI in Hierilrchy: Perspectives for EcologlCilf 12 Hogeweg. P. and Hesper. B, (1989) in Ouantitative Methods in undscape Complex.ity. Unfversity of Chicago Press Modelling and Simu/iJrion Methodofogy Ecolo£y /Ecological Studies. Vol. 851 12 O'Nelll. R.V.. DeAngelis. D.L.. Waide. I,B. IEllas. MS.. Oren. TJ. and Zeigler. B.P.. (Turner. M.G. and Gardner. R.H .. edsl. and Allen. T.F.H.119851 A Hierarchical edsl. pp. 77-92. Elsevier pp. 239-2811. Springer-Verlag COn/:ept of Ecosystems, Princeton 23 Wasserman. 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J 11-3Jv Arctic Biogeography: The Pcli'cldvx t."~ ,...~ ~ -~,J T(I<" ,,:),:.:< ["'::{I:·' f,liol.: a'l.1 f:~r<J I~ll: nf M rine vr" ill{labil Ine shallw arclic subliHorill ZOIlt' comprist' a relatiwly young marine as­ semblllge characterized by species of eilner Flora PacifiC or AI/anlic affinity and notably few endemics. The young character of near­ Ken Dunton shore arctic communilits, as wtll as thtir biogeographical composition, is largely a product 0( the Pltistount gla{iatioll. How­ Arctic Ocean marine benthic biogeographic origins of the marine ever. allalysis of mort rtetnt colftdions alld communities are distinctive in benthic fauna and flora throughout comparison btlwren IHe origins of the that they are composed of a the Arctic. benlnic fauna and flora prrstnl somt relatively young fauna comprising Previous faunistic and floristic inttresling parado:us 10 biogeographers. species of Pacific and/or Atlan­ analyses of the arctic biota were One enigma is tht low !rtqutnc!! of a/gal tic affinity and few endemicsu . based on available checklists of species wilH Pacific affinilies in 'ht Arclic, These characteristics have been animals and seaweeds from various esptciall!l in Ine Cnukcni. Beaufort and cited repeatedly in zoogeographic disparate geographic regions. More East Sibtrian Stas of tht Eastern Arctic. studies of various taxonomic groups. extensive surveys of the arctic sub­ l whiCH rtceivt direc' inputs of nOrlnlllard­ including polychaetes , seastars' littoral biota have been completed nowing Pacific waltrs. In contraS!. animal and bivalve molluscs~. The marine in recent years, butmuch ofthis data specits with Pacific affinities art Iound benthic vegetation of the Arctic remains unpublished or is located Ihroughoul If1t ntalSnOrt regions of the also contains few endemics. yet in relatively inaccessible Russian Arctic, reaCHing tntir highesl frrqutncy it appears to be characterized by literature. Consequently. the ab­ in the marginal seas bttwttn the Nell' species of predominantly Atlantic sence of data from these collections Sibtriall Islands and the Gmadian Archi­ affinity in both the Amerasian 6 and hampers biogeographic investi­ pelago. Organization of published and Eurasian7.3 sectors of the Arctic. The gations. As part of this review, and unpub/iShtd data. additional fitld colltc­ low number of endemics and the in collaboration with several sys­ lions. and Ihe useo! dadislics and mo/teu/ar variable predominance of species tematists, I gathered information DNA ItehlliqutS by systematists are Q high with Atlantic or Pacific aftin itieshave from these databases and the Rus­ priority for future research in reconstructing often been cited as evidence that sian literature ITables 1 and 21 and the ewlution of tne arclic biotic assrm&lage. the geographic distribution of the compiled regional checklists for arctic biota remains in a highly the seaweeds and benthic invert­ dynamic state and is not in ebrates. Here. I review these 'new' Ken Dunton is at the University of Texas at Austin. equilibrium6.9·lo. Even more interest· data, discuss the factors that have Marine SCience Institute. Pon: Aransas. TX 7~H3, ing but not previously noted is influenced the spread and estab­ US~ the appatent difference in the lishment of marine benthic biota 183 " . TREE vol. 7, no. 6, June J992 TI T~bIe I. SUIlUUfJ oIl11i1jor ptlbIiHed ud InptlbliHtd qWlDtJtatlYe SlIn'ey5 of the beftlilk: IUMt fallU of ttle uiillIow ndk Rlbllnam lODe siDce 1970 Regia... Depth Period of Reference raoge survey Iml the North Pacific from both arctic Alaskan Chukchi Sea 0-16 197~1977 51 and tropical waters also facilitated Alaskan Beaufort Sea 197~19BO 51 (he development of cool-water biota 0-'. that started to evolve at least 40 C"nadian Beaufort Sea 0-00 1984-1988 M. Lawrence (pe~. commun.) million years before the North Canaaian Atc.~ipel.J;o 0-55 197~1983 52 Atlantic lost its subtropical charac­ Franz Josef Land lMl8 1970 53 ter. The rich boreal taxa of the North New Siberian Islal'lds 0-32 1973 34 Pacific later became an important East Siber'ian Sea 0-15 1986 A.N. Golikov (pel"$. commun.) seed stock for the late Cenozoic (Chau... Bayl invasion or the Arctic OCean and North Atlantic during the submerg~ ence of the Bering Land Bridge". The first additions of Atlantic ldAa \from the northemmostextenston of Table 2. Summary of major published and unpllbllshtd collections a( the bt'nthk Nflne alcott of the a developing North Atlantic) to (he MCtic SlIb1ltton.1 lOne Arctic Ocean flora and fauna may Regio... Depth Oates of Reference have occurred towards the end of Iml collection the Eocene, around 40 million years Alaskan Chukchi SeOli 0-5 1976-1sao t.t.A. Dube- ago. Lis h}poth~sis is based on r Drift 1965 54 (he opening of deep-v.'ater seaways Alaska... Buufort Sea CHI 1977-1991 K. Dunto... and R. Wilce" between a cool·temperate Arctic 44 and a subtropical North Atlamic Ca"'OIIdian Alchipeltlgo 0-25 1967-1976 55 in lhe late Eocene:lf. By contrast, 1974-1983 56 Barryl' maintains that no significant NortheOlist Greenland 0-40 1907-1908 57 opening for Atlantic-Arctic water Franz Josef Ltlnd 0-38 1970 53 exchange occurred until after 27 New Siberian Islands 0-32 1973 7 million years ago. Regardless of the time disparity, the opening of East Siberian Sea 0-15 '98. 8 lChaul'l Bayl sea .....ays between the Arctic a,d North Atlantic coincided ..... ith a -Herbarium collectiol'ls strong cooling trend that dropped high-latitude surface water tem­ peratures below 1O"C by 40 mil­ lion years ago". This further en­ in the Arctic. and introduce the two seaways"'··JS, although new hanced the development of a paradox of the fauna and flora that evidence reveals that exchanges distinct marine boreal province In this analysis presents to biogeogra­ along these seaways were very the Arctic during the OligocenelJ and phers. IimitedU , The marine plants of this the evolution of a cool-temperate Nc period, Le, late Paleocene-Eocene, arctic biota of Atlantic character that 001 Evolution of the Arctlc Ocean i1lld its are postulated as the likely ances­ predominated well into the late Dc benthic biota tors of today's Alttic OCean nora of MiocenelJ.llo..H. A second major cool~ de The diverse origins and affinities Indo-Pacific affinityO. inl phase about 12 million years yeo of the arctic flora and fauna are By about the end of the ago Hate Miocene) dropped high­ be strongly related to the physical Cretaceous tBO-lOO million years latitude surface temperatures to e.. developmentof the north polar sea, ago), continental plate movements less than S"en , This temperature ak which essentially originated as a dosed the deep-water connection reduction further stimulated speci­ an, large northern embayment of the between the North Pacific and ation in variouscold-watergroupsof of North Pacific in the Mesozoicll~I), primeval Arctic Ocean'J, essentially the North Pacific, such as kelpl"l, and A" This primeval Arctic OCean was cool ending Pacific-Arctic biotic con­ subsequently, their herbivoresl&.