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. P.O. 119891 NeuriJ/ 34 Costanu. R., Sklar. F.H. and Day. I.W. University Press Computing: Theory and Pr3Ctice. Chapman 119901 BioScience 40. 91-107 , 13 Costanza. R. and Maxwell. T. feo/. & Hall l5 Haslett. I.H.ll'l901 Trends £co/, Eve'. 5. Mode/. lin pressl H Hogeweg. P. and Hesper. B. (19901 214-218 14 Hard. R.M.11990) Parallel Math. Comput. Mode/. 13. 83-90 36 Coulson, R.N. et al. (1990) 111 \ Supercomputing in SJMD Architectures. 25 Huston. M.• DeAngelis. D. and Post. W. Ouantitative Methods in LiJndscape eRe Press II981U BioScience 38. 682~9O Ecology {Ecological Studies. Vol. 85/ 15 Hillis, W.O. (19851 The Connection 26 Hara. T. (19831 Trends Ecof, Evo/. 3. (Tumer, M.G. alld Gardner. R.H .• eds). Machine, MIT Press 129-133 pp. 153-112. Springer-Verlag I: 16 Toffoli. T. and Margulis, N.119871 27 DeAngelis. D.L. t 19881 Ern/. Model. 4), 31 Taylor. CE.. Jefferson. D.R.. Turner. 5.R Cellular AUfOmata Machines: A New 57-73 and Go)dman. 5.E.1I9881 in Art,-fbiJl Life Envlronmenr for Modellf",. MIT Press JS Olivieri. I.. Couvet. D. and Gouyon. P-H. ISat1ta Fe Institute Studies in the Sciences 17 Hubennan. B.A.• ed.11988) The EcoJoffY 11990) Trends Ecol. Evol. 5. 207-210 of Complexity. Vol. 6J (Lane:ton. CG.. ed.l. I' of ComplJt.)!i... ~ Elsevier 20 "','nl',,'~, "I:- 1''''1(,1 Crowth"nd " .... :c-- ..,,,- ., '··~n"· >~•••\. 1," ,;, ' ....,·'''.''11 ,.~ !~1 , fir.. l:i ,-_ £eol. £>01. i. 36-1" )0 u,.. "v",".~. 11\1...01 Oi~05 ,;. ~2-1.'1 UI,"-_'.', ;. '.. ". i •. ,;: .ed.i. 20 Hogeweg. P. (19881 App/. Math. Compul. )1 U1.ano",'''. R.C t 19&'IJ CeDI. Mood. 4). 1->;'. :.,:-}io. AJJj,,,n·'.V,,~I,,y 27.81-100 45--5-6 )i Hogeweg. P.and Hesper. B.1198Sl :11 Tamayo. P. and Hartman. H. (Iml in ), P;ll:l'l'. H.t!, 119Ml in Anifiei;:,! Life ). Tlicor. Ciol. II J. J 11-3Jv
Arctic Biogeography: The Pcli'cldvx t."~ ,...~ ~ -~,J T(I<" ,,:),:.:< ["'::{I:·' f,liol.: a'l.1 f:~r
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&. Atl temperate in character and prob tinuity for at least 60 million years, The sudden appearance of a P" ably contained a large number of until the late Pliocene (3,0-3.5 flood of Pacific mollusks in Iceland on species which later become a million years agol'}·'·, The dosing of about 35 mimon years ago suggests sp. portion of our present algal 'cos this conne<:tion pennanently ended that the Bering Land Bridge was PI, mopolitan' groupo, Subsequently, in the exchange of truly bathyal or breached in the late Pliocene'). The th, the late Cretaceous, the Arctic abyssal fauna between the North opening of this shallow-water pass~ of OCean developed two large seaway Pacific and the primeval Arctic age resulted in migrations of biota an. conneaions, one with the subtropi· OCean. The effectiveness of the through an ke-free Arctic OCean. cal Gulf of Mexico (Western Interior shallow (70 m) Bering Strait as a This resulted in a high-latitude biota QUi SeawayI and the second with the bathymetric barrier to dispersion of maximum diversity as reflected in ,'" tropical Tethys Sea ITurgai Straitl'J, is strongly reflected in the ab a fauna and flora composed of I Arctic species. with phylogenetic sence of bathyal Pacific-boreoarctic Arctic and North Atlantic forms that wh J 7 affinities with warm-water taxa may polychaetes , ostracods' , and other were strongly supplemented by ag' have originated during this period Pacific deep-water benthic fauna'l·" migrants from the Pacific6- IO , The na' as subtropical migrants along these in the arctic basin. The Isolation of spectacular eastward dispersal of pe '84 2 TREE vol. 7, no. 6, June 1992 reviews
00",
c j , Lopfev ._. ) 0i;/Q. ('-"~~(Jr-
r? NSf
cos! ~ ... Norwegian Sib~rian SQ. Seo ' ...... - .- " ~.-. ' .. .. . "p " ... . + -- ' + .. .. ···, -~·X
v v ALASKA
fiCo l. Contemporary surface and subsurface circu!iltion in the ArctiC OCean"A''''. !'lSI: New Sibcri..n Isl,lnds, f)L: FfanL losef Land: CAP: Canadian Arthlpelaeo lQueen Elizabeth Islandst: 81: Bathurst Island. Map adapted from Ref. 35.
North Pacific species occurred in a approximately equal duration of tinental shelf was accomplished by cold boreal but totally ice-free Arctic 10000-20000 years !D. Hopkins. the few survivors of the repeated Ocean; a perennial ice cover did not pers. commun.l. The last major glacial episodes that retreated into develop until about 0.7-2.0 million glaciation (Wisconsin or Wurrnl the North Atlantic or North Pacific 7 years agol • This large invasion may reached its maximum extent ap or took refuge in unglaciated shelf be attributed to both favorable proximately 18000 years ago; sig areas of the East Siberian and eastward~flowinglongshore currents nificant deglaciation did not begin Beaufort Seas or deeper bathyal along the coasts of northern Alaska until 14000 years ago and ended by waters of the Eurasian sector'·-·". It and canada and the development 6000 years agol!. Reconstruction of is likely that the benthic seaweeds. of passages through the canadian full glacial conditions included a which depend heavily on stable Archipelago into the western North sea level drop of at least 85 m19.lO, hard substrate for attachment. were Atlantic'6 (Fig. II. However the exposing vast areas of the arctic eliminated from the Arctic during process occurred. it appears that continental shelf'6.2'l IFig. 21. Major glacial episodes. retreating into the only a small percentage of benthic ice sheets covered much of North North Atlanticl2. Their dependence species emerged from repeated America. Greenland. Iceland. Scan on light for photosynthetic carbon Pleistocene glacial periods. It is dinavia. the Barents Sea and the production would have further com these survivors thatfonn a large part Kara Sea. Only the continental promised their ability to survive of our contemporary arctic faunal shelves of northern Alaska (the either in shallow water or at depth. and noral assemblages. Chukchi and Beaufort Seas) and the especially follOWing the develop East Siberian Sea remained lall':ely ment of a perennially ice-covered Ouaternary glaciation: consequ~nces for unglaciated yet almost entirely ArcticQcean by 0.7-2.0 million years arctk biota emergent2'lJ' . ago. During the Quaternary Period, Such severe conditions occurred which began about 1.8 million years many times during the Quaternary. Dtllt'lopmtnt of a post.P/tisloCLllt ago. the Earth's climate alter and there is a consensus that shelf btlltnil fauna nated between numerous glacial faunal assemblages were nearly Nesis" attributes the occur periods and wanner intervals of eradicated u.s. Reinvasion of the con- rence of stenobathic. shallow-water '85 TREE vol. 7, no. 6, June 7992 TR x················ .):m:::::::::·::::::.:::::::::::.: ...... - . Eo ...... , . n B, sp aff
fa' Eo no thi th, Th wa tal Its th, sh he
p, ne
,. lliJ Glaciated Areas Wi' f; [] Sea-Ice ,.
...... ;:1
~', :"'-." .,<.... "Sil Fig. 2. Geography of the Arctic during the height 01 the Wisconsin or Wiirm Glaciation,·J,,". Moull!ain glaciers 'lIe omitted <0 "'H. lsi marine fonns in the Eastern or Cenainly one of the most dis The Atlantic character of Western of Pacific Arctic (the area between the tinguishing features of the Arctic Arctic fauna is influenced by North New Siberian Islands and Bathurst OCean biota from a biogeographical Atlantic water carried into the "A< Island in the Canadian Archipelagol perspective must be the different Barents Sea by the Norwegian a, versus the presence of eurybathlc. distribution of species with Pacific Current and waters carried past P;; deeper sublittoral species in the or Atlantic affinities. As noted by Spitzbergen and into the central fa, JJ Western or Atlantic Arctic. to geo· Ekman , these distribution pat Arctic by strong boundary currentsJ\ 51 graphic differences in the con terns fonned the basis of two. (Fig. II. By contrast. the Eastern b. ditions of the Quaternary glaciation. extreme viewpoints. First. that the (PacifiCI Arctic is characterized by a p, This hypothesis is based on evi present arctic fauna is entirely higher frequency of fauna with d, dence that the entire shelf of the derived from post-Pleistocene At Pacific rather than Atlantic affinities. Atlantic Arctic (from Greenland east lantic populations. and secondly. The Pacific chardcter of the Eastern into the Norwegian. Barents and that the arctic fauna is exclusively of Arctic fauna is probably related to Kara Seas. see Fig. 21 was covered Pacific origin. Although the majority Pacific water flOWing northward with glacial ice that descended consensus is that the arctic fauna is through the Bering Strait. al J deep into the ocean !, These ice of diverse boreal origin (involving though this volume is one-half shelves would have precluded the both Atlantic and Pacific forms). the to one-fifth the volume of At existence of a shallow-water fauna. influence of Pacific migrations to the lantic water entering the Arctic In contrast, the sublittoral zone of arctic shelf in post·glacial times Is OCean tK. Aagaard, pers. commun.l. the Pacific Arctic. although emerg made very dearby the large number Nonetheless, animal species with ent. was not glaciated'·. The benthic of Pacific elements that-characterize Pacific affinities are widespread in sublittoral fauna of the Pacific Arctic the faunal assemblage in the Chuk the Eastern ArcticM.Jt>, and in wastherefore not forced intodeeper chi, East Siberian and Beaufort Seas some taxonomic groups Pacific water, but contin ued to evolve in of the Eastern (Pacificl ArcticU.l<. forms predominate over Atlantic J7 shallow waters of Widely fluctuating In the Atlantic Arctic, there exists elements - l9• Analysis of recent temperature and salinity conditions an expectedly higher frequency of collections from the nearshore characteristic of the arctic nearshore both animal and algal spe<:ies with littoral zone 10-50 ml reveals that environment,a. Atlantic rather than Pacific affinities. benthic faunal assemblages in the 186 TREE vol. 7, no. 6, June 1992 reviews --_ .. ~--
Table 3. Dislrlbutloo of benthic Ilttonl{0--50 IIIlI marlae fauna of the Arctic Ck:ean~ Region (nl Biogeographic group 1%)
Atlantic Pilcific Boreill Eastern Arctic (particularly the Arctic boreal boreill ilrctic ilnd Chukchi. East Siberian and Alaskan endemics arctic arctic cosmopolitiln Beaufort seas) are dominated by Chukchi (150) 9 6 27 58 species ofPacific rather than Atlantic Beaufort lAlaskan and 8 13 15 64 affinity ITable 3). canadian) f37l) The high frequency of Pacific Alaskan f2061 8 6 17 69 faunal elements in the Beaufort and Cilnadian 12661 8 17 10 65 East Siberian seas is related to the northward flow of Padfic waters Cilnadian Archipelago 11581 5 20 9 66 that enter the Bering Strait and Barents (186) 14 15 8 63 then bifurcate in the Chukchi sean. lapleY 11521 12 19 10 59 This intrusion is composed of two Ea!>t Siberian" 9 5 16 70 water masses. the Alaskan Coas -Oata al'lCl biogeographic affinities derived from sources listed in Table 1. tal water and Bering Sea water.c. "Number of species not available. Its eastward flow is restricted to the outer Beaufort sea continental shelf I> 50 m) and is opposite to both the anticyclonic Beaufort sea waters into these areas', the veg Tlit tfftd or pliysiCJd factors Oft' td9llf gyre»··l and the net westward flow etation (especially along the Barents di~ptfSQl ill If" Padfic Arcrie of surface water that corresponds to C0.:.3t) jj o\erwhelll.....;·, L0r temperatures in the seaweed-rich combination of these physical fac recolonization in the nOM polar s, Aleutian Islands range between _)0 tors has probably played an import basin since the last glaciation and th and 5<>C compared to -1.BoC to Pc ant role in slowing the invasion of their requirements for hard sub te in the Alaskan Beaufort Seau.~. North Pacific algal species into the strate in areas characterized by I. The rich diversity of the Antarctic Arctic. at least in comparison to the frequent ice scour. Secondly. the ;" seaweeds. which includes some benthic fauna. Unlike the benthic presence of Pacific elements in the b, North Pacific representatives (D. algae. many arctic marine invert Arctic OCean biota raises the p, Moe, pelS. commun.). also suggests ebrates are not restricted to hard paramount question of the origin of that cold temperature tolerance is substrates in the shallow, ice the arctic biotic assemblage. not a factor limiting northward dis scoured euphotic zone of the arctic Zoogeographers appear unani '" persal of North Pacific species. shelf. mous in their assertions that the especially those growing along the contemporary fauna is neither n~~_~ .o\!euti.1.n shelf :::! the Serl", andent iI.e. a Mesozoic rdid nor Sea. biogeopaphiul perspective well establisned. and that the One possible explanation for the Delineation of the boundaries present fauna is a young coloniZing lack of Pacific floral elements in the of the Arctic Region is a sub biocoenosist.u.•. l't.4&. On the otner Eastern Arctic may be related to a ject of continuing and spirited de hand. phyco!ogistsmaintain that our combination of factors that includes bate. mo<:t recently sUmm<1riled by current NNth "'bnlie.. Nnl1h Pacific ._. -. ,. " , .. intcequency of hard substrate along ul thes~ b,)l.HH.....oi .. 5 i; imp,Jl~a:.t. species must be refined to assess StilrUord Univcrslty Press ]I Macpherson. E.l 19711 ftlad Mus.. Qn. the impact of global changes in 11 joy. JA and Cl.lIrk. D.L 119m PtlbI. Bioi. Oce.,nogt: 3. 10t9 MiaopakofltoioD 23.129-154 It Fedyakov. V.V. ilnd Naumov. A.O.119&91 temperature. insolation and circu [I Ne5/s. K.N.(19331 Soviet I. AUf. BiaL 9. in The.4lrtfc SeilS IHennan. Y~ ed.l. pp. lation on the biosphere. as reflected 235-243 )ol-324. Van Nostrand Reinhold in the time-averaged response of I' Dumam,I.W. and MacNeil. F.s.119671In 40 Mountain. D.C. I 1974J In The COiISl and benthic animals and algae to such The Beri~ Land Brid#"" lHopkins. D..M_ Shelfalehe Beaufort Sea lReed.I.e. and cdJ. pp_ )26-349. Stanford University Press Sater. I.E.. edsl. pp. 27--42. Arctic Institute perturbations. 20 Eldhofm. O. and Thicde. I. 119&11 01 North Amertcao PafCOf~r. Pakoclfmaco'. Pa~. )0. 41 (olony. R. and Thomdike. AS. 119341 -.-~ 243-259 I. Geop/lys. Res. 119. 10(2)...1(1629 I ;am especially gl1lteflJl to R.T. W[ke and 21 Barry. RC.119891In The Arctic: Sfras 42 Barnes. P.W. and Reimnitz. E_1I97411n D.M. Hopkinsfor their hlghlyconstruetiveand IHennan. Y.• ed.!. pp. 1-46. Van Nostrand The Coast and Sheffof the Beaufort Sfrd thorough criticisms. which helped Improve Reinhold lReed.I.e. and S.IIter.I.E~ edsl. pp. this review considerably. S.V. Schonberl and 22 Savin. S.M .. DouII.llS. R.C...nd Stehli. 09-476. Arctic Inslltute of North America R.T. Wlice provided Invaluable assistance F.C.1197S1 Bull. C~. Soc:". Am. 36. 4J Mohr.I.L. Willmovsky. N.I. and Dawson. In the compilation and organization of the 149'9-1510 E.Y.Il9571 AJrtlc 10. 45-52 ta~onomlc literature. I also thank K.l\apard 2J Berggren. W.A. and SChnitker. D. 119831 44 Dunton. K.H .. Relmnin. E. and for his review and F. Bacon-Dayton for his in Suucwre and Development of the Schonbcl'l. S.V. t 19821 Arctic 35. 465-484 Inspirational support. This work was sup C~en/anfi--&otldndRidge lBott. H.P. et a/.• 4S Smith. S.L .lind Vidal. I. 1[9841 J. Mar. ported in p;art by NSF irant DPP-900060S. the cds). pp. 495-548, Plenum Press Res. 42, 221-257 National Acildemy 01 Sciences and BP u· 24 Hoell. C. van den 119841 HefBoUnder 4Il Holthe. T. 119711ISiJrsia 63. J91-1911 ploratlOn IAlaskal. Inc. This article Is contri Mu~~nters3&. 221-251 47 Rozycki. O. 119871 Polish Polar Res. 8, bution number 1132 from The University of 25 Stam. W.T.. Bot. P.V.M .. Bocle·80s. SA. 107-120 T('l..1s .\~,Jrine Science Institute. ' ..... " ...... 1.):·1...... : ,~ .• ..:~., tlod·. e. {I~.),;l H..:,plJndi!r MeereSlJnlr:rs J2. :!, 1-207 i; H (Sles. I.A. and Steinberg. P.O. [19831 ••. :•._~'. .• ~""~". L II i"lOlin ReferencleS r ·__ ~.Jlogy IJ.I9-lO J ••,I'., .. ,Jl:raphyofrheM.)rln(> I Colikov. A.N. and Sc.iIIfato. OA 119Mlin 2i (L"ke. D_L 119901 in The lvaicOceiln -" _':J<, 01 :.'>c '\l"rh All:Jntic ICarbary. 01. The Arctic~;u; lHerman. Y_ ed.l. pp. Rt"gion IS"'eeney.1 F~ ed.!. pp. n-62. .:",.1 S-,,,.t!. C Il . <'d~l. pp_ 187-2().1. 251-2&0. Van NOW'ancl Reinhold Ce.>logical Sodety 01 America S;l' ~r,;of··:,;ot' ..: 2 Brtccs.I.C-119111 Af4Jrine ZOO¥e