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uolpnporlul 1."I \
rr su.raned.rotew aql loau.os Euutrr.rexl 'Iooq slql lo Japeal aq1 1o uedsa11l aql $nf u1
ilge:rpeLp pa6ueqr a^pq ll|^\ ]soul lsa)Pld )lurpu,tp ureurar pue paloof pue paleaq 'uall?l
)up uesrl 'passa"rdulor uaaq lo pal-p!/n ts eq sueaf,g 'a:eds pue,aulp ur a:e;.lns laueld
rql uo sa6ueq: lngaruod 1o sjole)lpu! tsrP q)rqrv\ ]o llB 'azls lo 'ssbulua!)up 'aluPpunqe
pue
ssauqru sa1:ads ur ln))o suraued 'Aep luosa.rd aq1 o1 o6e slear{ tro suollllu, ulo4
ssiiqe daap ar+ oi smollells aql u.roJl '!ts4ual uPB)o ol flseo) tuol] 's:1do.r1 aq1 oi salod
,ql u.tol] luapl^a oslP are 'aroqs aql le snol^qo os 'su.rs1ue6,ro 1o suraued 'd;a1 ;o lsarc;
)l sJtsaJ le;o) pue sureld luauJlpas ol sabuel uplunoul easlapun u.ro.r; '[llerpeulelp
ahueqr ade:suearo aql ]o asusdxa spr^ aql 'll Ual la^au sleu.rlue lo (el^qd) sdnorb
ofeu"r lsol 's:ea,{ 1o suollltru }o spaJpunq ro} }uatlluorl^ua qql ot paulluo) pauleu,laJ
tup sueo)o aU] ur ue6aq a1t1 :ssa5 pue suealo $eA aql ul sl quea uo alll ro] ajeds lsoyrl
soo6
eql ul slJ,ltsIlsd
v< t0\J I ra]deqf
fiztvli
ffi; ],ir1':,11::P,'r':':**'.::1ry.II
;l:'i.''-'"':::'iJ''.,.':, ' Funhermore even a simple climb up a mountain can reveal altitudir
. pushing'contineirts apart, aad eventually disappears down beneath
- .-, . 1-: ,' ,1" . . , . , conrinental ..ust, forming deep oceanic trenches" Considering its changes and, though few have experienced it, most people understa
volume, the vast maiority (c.99"/,1of ttre earth's habitat is marinc and that polar regions are deserts compared to the generally species-ri
;iff"T""J:i:ffi,lfJilt"
'" p.ohasized nv ri," r"a tirt ;. i, where most maior rypes of animal (phyla) evolved and continue to live tropics. In contrast, the water column and wide ocean basins might
-##;;;;;;;;;; exclusively. Most of rhe vast water column and seabed (benthic) habitat envisaged as fairly monotonous, uniform ecosystems. However, rht
_
nw remains unobserved by human eyes. New species are still routinely are mary features that puncfuate them abruptly or gradually into ma
- dei*ibd, but rhat entirely
ctasses or phyla (the highest found in deep-sea samples and even some of the larger animals on earth different environments. Changes in time, topography, chemistry, a
ta,(onomic levels of animal types) which live th".r, ..g**outh SharkS and giant Squid, have never been oceanography allow for the development of patterns acros$ a wide ran
- hlave been discovered in iust the of scales in time and space, and form the subject o{ this fust chapter a;
seen alive in their natural habitat.
last couple of decades' are themes thar reoccur throughout this book.
Standing at the edge of a forest looking out over a prairie, lake, or into
rhe tree canopy, it is easy to see how fragmented the land can be-
1.1.1 Zonation
Patterns in the marine environment are often beyond our immedia
h 1995 an enlire new phylum of tiny animals, the Cycliophora, was reported from a perception; we cannot always see them. Often, patterns are only reveal,
discovery two years earlier. As it is only 350pm in size and superficially resembles indi- through sampling and subsequent interpretation. The number and si
viduals of several other phyla of small animals (Gastfotricha. Rotifera, and Entopfocta), it of samples collected and the rype of equipment used will have a pr
could be considere.d unsurpdsing that such animals are stilt being discovered. Surpris- '
found effect on wh::: . .rrd. Strong differences in opinion exist abo
ingly, though, the single species lSymbion pandard livel on the mouthparts of
'lVe,phrops even the most ba$ic ir-:, ,rre biotic panerns, each defined by eviden,
norvegiats -a very common, well studied and widely consumed species, often refered
from a discrete $e[ of samples. In many respects, p4ttern$ in the sr
to as scampi. 5. fundora, or the 'Pandora', attaches to its host using a sucker and
resemble those on land, for exarnple at a large scale along gradien
suspension feeds.on particles in the water, a small parasitic male (whose sole purpose
seems to be for breeding) is also shown attached in the picture below'
of solar radiation (latitudinal gradients), altitudinal {which in tl
marine environment is depth, thus barhymetry), from the coast to ocen
continent centre$ and from young to old areas (e.g. from the mir
Atlantic ridge (new) to the far eastern or western Atlanric sea-flot
(oldest)). In the shallowe$t parts of the sea there are plenty of plac,
where the type or nature of the organisms changes over tens of cent
metres or metres, which we term zonation (Chapter 5). In warm tropic:
waters, the type and dominance of corals changes quickly with dept
(Chapter 10). In polar seas the abundance and richness of marine lit
alters equally sharply in response to decreasing physical disturbance h
floating icebergs that scour the seabed {Chapter t1). In certain rypes <
environment, such as estuaries (Chapter 4)' rapid changes in biologicr
constitueflts occur along the length of dre estuary in response to a suite r
changing environmental variables; parterff rhat are repeated at virnrall
all latirudes. Zonation is most apparent where the land meets the sea an
it is here that it has been studied in most detail {Fig. 1.1a}. Zonation is nc
iust driven by tolerance to physical conditions (though this is mor
important at the high shore level). Connell's (1951)work with barnacle
demonstrated L:hat interspecifi c compedtion and differential predatio
prbssures strongly in{luence the locatlon where species survive. In temper
are regions across [he globe different species and colours of algae an,
lichens indicate the gradient of immersion on the lower and upper region
of the shore. Shore zonation is equally apparert in some muddy shores ir
feeding (plantotrophic), or have their own yolk sacs (lecithotrophic) and There are many features thar partition the ocean environment, and t
behaviour. The water column, like the land and seabed, has many sharply posirions of continents, islands, and subsurface marine mountain chaj
changing physical features that effecdvbly form environmental barriers form obvious physical barriers. In the open oceau, differences in wa'
that constrain rhe organisms that live there {Chapters 2, 3, and 6). densiry, curreflts, and fronts provide the conditions that lead to distil
lryater masses. Although current direction and velocity of water mas
and patterns of wind across Ehe globe are complex, there are gener
1.1.2 Oceanography large-scale pamerns fhat occur, most obviously in water flow. Most de
On a larger scale the water column has a strong pattern of zonarion rhat tseabed) water is derived from the Southern Oceau. Cooled dcnse wat
occurs across irs full depth range, similar ro that found in the Iittoral zone. (termed Antarctic Bottorn'W'ater, AAB!() sinks in the Southern Oce
Most importanrly in the top few to 200 metres of the water column rhere is and flows away from Antarctica well into the northern hemisphe
enough light (during the day) for primary producers to photosynthesize, abyssal regions" There may be several currents that occur at differe
termed the euphotic zone (Chaprers 2 and 6). As a result, the top L00 to depths between the deep AABII and surface water. For exarnple, co
200 m ofopen ocean water and 1 to 50 m ofcoastal wateris a very different northern water llows southwards in the mid regiou of the water colun
environment to that found below. Of course, even within the euphoric
zone there is a stroog gradient of lighc intcnsity and wavelength with
depth. Beyond a depth of 1000 m (most of the world's ocean volume) the
ocean is effectively lightless, with a few srnall-scale exceptions, such as the
bioluminescence produced by ba*eria found in the light organs of deep sea
biota. Light striking the surface of the marine environment also imparts
heat, thus the surface layers are the warmest and hence have lower densiry
than the cold water beneath- Globaily the temperarure of deep water is
relatively uniform at iust a few degrees Celsius, in conrrast with shallower
water temperatures that fiuctuate with latitudp and sea$on. In the polar '5
regions, surface water is near freezing point at -1'85 'C in the winter, and
just positive in summer. However, the water in polar regions is well rnixed
as dense cold water sinks and wave height and wind speed are greatest at a =
latitude of 5O-60" (Fig. 1.2). Moving away frorn the polar regions, with a
few exceptions such as regions of upwelling, the global ocean is stradfied
Lotilude (degrax)
into a warm upper layer, a rapidlysooling zone, and a lower cold zone.
a Thermocline! are a vertiqal
zone of rapid temperature
The nature of thermoclines changes fqom place to place but in general the
change in the water columri. therrnoclines in the tropics ,r. *orJ'[hrn 10"C warmer than rhose in
temperate regions and they ,re permlnent rather than seasonal- Such
stratification is imponant as organisrns require nutrients and minerals as
well as light and respiratory gases- Thus away from Permanently mixed
areas such aE polar seas or upwelling zones, the surface layers of the sea
also have strong changes in molecules used or produced by organisms,
such as nitrates. Locally other gradients' e.8. halocliries (satinity) or
pycnoclines (oxygen) rnay also stratify water layers. The salinity of sea- =
water is rypically about 35 psu. Generally changes in saliniry rhat occur
with either larirude or longirude across oceans are small, but are a Iittle
greater in the tropice (due to increased evaporation of surface water) and
=
a psu - pract;cal salinity uniB. lower close to continents and in the arctic due to the influence ofgreater
(see 4.1.4.). fresh water run-off (Chapter 4).
loitude {da$reer}
1: pATrEfiNs tNrHE MARTNE EN'IR'NMENI
@
with time scale'
Table 1.1 Example of changes in the rnarine environrnent
Time s(ale
by a
in the Atlantic. At the sea surface, the world's oceans are c[ominiited
in the northern hernisphere and anti-
scries of gyres rotating clockwise
meet in the equarorial (')
clockwise-in the sourhern hemisphere. where these
cnrrents llowing wesrwards occur (as well as smaller ?50
region strong !00
1.1-3 Climate
There is now much concern about c[imate change and rhe
extcnt to
to this phenomenon (chapter 14).
which human acriviries are linkef,
The climate of the earth, however, has been in a state of constant
change, only the magnitudc of rhe i:rrte of ch.ange has varied
rvith time
*oui envirohrnental variable associated with
G'-ig.i.:1. Ttt. familiar
.While
air or land temperatures fluctuate more
.tiior. is remperirure.
iru*".i..11y, ,"u ,urfr.. teruperature (SST) changes are more subtle due
to the huge voiume and high latent heat of water which gives it a strong
through with rime. Clobal sea level change from the Mesoz"oic to
buffering effect. !ilhen global renrperatures are high sea levels rise Fig, 1.4 Sea level change
of ice. There is great connectivity presenr (a) and the coastline of NW Europe {b) and Australasia {c) c.18 thousar
thermal expansion and melting yea.rs ago (in light green) and now (in dark green)'
between m;iny environmental variables, such that when SSTs rise,
c.rther
change concomitantly. For exarnple warmer
environmental parameters
lvater is able ro hclcl less gas and hence oxygen needed for respiration
rnuch larger scale, each ice age spreads the icecap to lowcr latitudes fror
(Chapter 1a). At present, icecaps opcur over ti'le rvso polar regions'
the poles, lhen retreats in interglacial periods' The rise and fall of sea levr
rvhicL is a relarively unusual condition in palacontological time scales-
extent' but not has clearand importa$r irnplicarions for rhe marine habitat (Fig. 1'4
Ice sheet expansions occur cvery winter (the geographical
to winter)' On a very the shailow sloping continental shelf is one of the most importar
the volume, of Antarctic ice doubles from summer
'f,"Uii^tt primary
for most organisms with 90% of the world's marine
0ennings & Kaiser 1998)' The extent of the continental shelf Methane clathrate is a type of ice that has methane (cHJ hErnd within its crystalline
.- '.: "roa,r.,ior,
ffi;;;;ii"rruiy wirh iust a 50m change.in mean stobal sea level slructure. Euild-ups oi this hydratg are probably created by gases moving along
geo
ha:
a logical fault lines whae they crystallize on mixing with cold water. Methane clathrate
--
The extent of the continental
6^;;t; ; and 7). Changes in ice e$ent'.-which occur simultaneouslv
fietf changes <ondderably with away entire long been thought to be comrnon in the outer solar system but mote recently it has been
.'. jus"a 50m change in mean *'if, ifro*. in sea level u'"1ko important' literally scraping discovered in considerable quantities below some oceanic sedirnents. This i5 importanl
t"iitrt. and thereby oPer uP new space for colonists"
grlobal sea level' 'l;;;;;tcrals forteveral reasons: first, it is A maior source of fossil fuel (m:ybe >90% of natural
gas
of climate change have been found in the Iast centurv' reserves) and Second, release of ozerlying pressure on these deposits could result
in large
1945 *d fto* 1976 tathe present' fVtft$1
n";;fitiy"f*m 1916 toremperarur may have increased by only 0'6'C
sCale cJischarges of methane through the ocean into the atmosphere. Such releases are
Iurrt ', mean surface air thought l0 have occurred rapidty, on various occasions and may have been responsible
west Antarctic air and
i" some regions, soch as the Arctic and for sudden climate warming" For example. some of the rapid increases in thE temper'
" "-r*ty,
;;il ;p;r.es, huu" **'med more than this in a decade (Iralther et al' ature of the fuuthern ocean (Fig. 1.3) seem likely to be calced by methane clalhrate
rapid
drasric and
ffi;):'fil; is not withour pfecedeur; there have been in the sea' These
changing from s.iorage in ice crystals to atmospheric gas and then having a
profound
even
.i*"**"i, the earth's so'fai" temperature before'clathrate releases from
effeqt as a greenhouse gas.
methafle
;;;;;t occurred in response to rnassive after metenrite collisions
and
i.i.# ,ft- seabed, maior volcanic events
of the Cretaceous period]
at the end
a Past climate changes occurted [r.r, * irr", rear th; Yucatan aspects of r'' .'t climate change have 1.1,4 ProductivitY
io relponse to massive methane
ir.. S"* 1.3). Other prominent
irr.r*A.a increased C02 levels and ihattges
clathrate releases {rom below rspheric ozone thick'
Unlike on land, true plants generate linle of global marine producti<
the seabed, maior volcanic events Ti- n*t of these is strongly related to rL: ' :ture changes through
{except in intertidal saltmarshes, seagrass meadows,
and mangro'
and after meteorite collisionr' ".t*
ir."i-trrppinc gnd is referreJ to as 'the greenhouse effect" Seasonal
swamps). ln cool shallow coasral warers, macroalgac such as subtidal kelp
(ozore holes) permits increased
thinning of the upper atmosphere ozone or intertidal greeq brown and red rnacroalgae, are highly productive ar
of light with their potential
n"r"irrir", of tr,. ,rttmuioiet wavelengths of water is limimd to iust a
can grow rapidly (Fig, 1.5)' The productiviry of thesea mostly deoends c
Lmaging effec* to organisms' tlV peneuation to
I Ahhough the Peneftation of
a serious issue in inicioscopic free-living single'celled algae collectively refe*ed ,
potentially
UV tight affects only the upper f.* ,i.tJ.u of the sea zurface, so although
ir does not affect the phytoplankCIn (see Chapter 2). Although phytopla:rkton are individual ('
shaliow aquatic habitats
{ew metres 0f leawater, it ha5 the ;;;*;;;i;L littoral,
and very
uggr.lrtiont of) single cells they vary in size from large (20G-20p
fuide from a
potential to adversely affect fish *ri"ti,, of the volume oi rhe global ocean environment'
climate change' there are dil-*Jt r), diatoms {microplankton)'to tiny (2-0.2prn diameter} ba'cter
laruae and other biota' ui.o.rg .lgrrri of seasonaliry in the oceans and (picoplankton) or evec smaller cells (see Chapters 2 rnd 3)' In tropical cor
varioustrtherlonger.te,*bo.ryclicaleventssuchasMilankovitchcycles ,."f iyrt*m, much of the prinearl productiviry is rssociated wirh micr,
concern Iolar activiry' tqth."" solar,flares)
and dre El Nllo
i;ilh considerable scopic algae that occur as symbionts within the tissues.of animals, such ,
."r*-,, such, as the buffi stream {which rakes warm water to florth-
and changes in phytoplankton production *rough often appears tohave a higher bioma
;;; Etrrope), resulting in colder climatic-condirions L..**. the turnover of primary productivity is so rapid'
of ENSO has been recorded for more
;;;lipr"-ttt The phenomeaon been traced back nearly 500
The rate of primary productiou varies along gradients of light ar
rhan hundred years and discrete events have
a nutrient availability. As a resulr productivity vades across the tropic
ao weli as physical and
,.ars. ENSo and other elements of climate change, temperate and polar regions, with depth, with the proximity to the coa
;;;;.;nography have a stropg influence on the magnirude of
aod other fearures such as upwellings. Pelagic productiviry is rypical
oroductivirv in th. o.*"os (below)' This has consequences for-higher of magnitude lower than for benthic system
more than an order
;;;J;;;t; ,u.h ,, fistr, mammals, and, seabirds as well as fisheries
The maloriry of benthic producdon is, however, restricted to only a tir
(Grantham et al" 2004)'
ry
ffi 1: pAriEBNs rN ?'rtE MARtItr Ei\v'tAcwiv:ENr
Thus on a plot wirh log scales on both axes, the relationship approximatr
-
to a straight line shown for example by plant numbers on Australia
islands or even {approxirnarelyi for species on contimnts. At the other en
of sparial scale a number of authors have immersed serdement paoels c
various sizes into the sea and demonstrated simila; increases of specie
@ f : PATTERNS fff lrig MARIME EIlVlkOr\]i/IENT
with area of these milliarure islands (see e.g. Jackson 1979). There are a parameter with a larger scale unir, the unit in quesdon needs ro
d.fin.d. Although we have already considered a very brief history of
t
'
numberofcausescrflspecies*arearelafionships'suchaShighernunrbersof partirioning of ihe terrestrial and marine realms into smaller areas
you encounter)' more
samples (the more samples you take the more species to t
greater ratios of sPeciati.rn units (1.2), it is appropriate ro consider rhis in more depth due
habitars, immigration and extinction rates, and ro biogeography'
less likely tn be encountered by lvoLrld-be imporrance of this issue
ro extinction. imaller islancls are
larger land rnass.
colonizers because they represent a smaller target than a
similarly rhe more isolated an islanct, tl"Ie smaller the pool of organisms 1.2.3 Biomes, o(eans, and PrCIvinces
or airborne orgarrism wo'-rld take
,hn, .r"'.np"Ule of reaching it. A srater of
using a small ser of factors, it is possible to predict a series cor
, iong tl-" to reach Bouvet Island in t6e Sout{rern Ocean' which is ,nonity types or 'biomes' on land- Using high-resoiurion CZCS images
rho,slnds of kilometres fron-r rhe nesrest land of any kind. Of
c.urse
play a maior role in determining phytoplanktotl 'greenfiess' (a proxy for the biomass of a phytoplankt'
wincls, currents, and chance
prevailing
Ll,ro*1, an analogous series of biomes can be generated for the mari
e lsland size and itolatiofl are *t l.n ofirnisms arrive, where and when, which in part cxplains why e,rvironm.nt. Thus four primary biomes are recogniz.able from chara
important determinants of
there is oit.n much 'noise' in species*area plors of sampie
data'
specie5 ri(hness" 'l-here are all sorts of islands in the marine environment varying frorn terisric patterns of phytoplanktooic algal growth, .the environment
or hydro- forcing agents {winrl, currents) and fauna at other trophic leve
enclosed coastal lagoons, sea k:chs, and fjords to seamounts lWesrerlies, (3) Polr
(chapter 8)" Thc same principals of colon' t ongtiur*fqtl98) details these as the (1)Trades, (2)
thermai venrs in deep water are all defined by the agent th
tr: r- :r:onments' However' and (4) Coastal biomes' I'hese
ization and dynamics of fauna apply these
determines the deprh of rhe mixed layer of seawater: wind acting ov
,p*.i*r**."u relarionships tend to be compl' other factors: for
large-scale distanies (Trades), local wind speed and light intensi
Jamaica contains many more specii
rst organisrns than or a complex
(\{f,sterliesi, surface recluced salinity water (Poiar},
"xn*ple,
rhe F;lkland Islands even rhough rheir land masses are of
similar size
processes (Coastal). With the exceplion of the Trades biome, all
inclu'
and the
and disrance from their a<ljacent continental land mass' Jamaica For example, rhe lolar biome occurs (
regions (carihbean cf. ,.urrrl disjunct waler masses.
Falkland Islands are situate<l in very different high lariturle) in each hemisphere and can be split into Antarctic, Nor
the di{ferences we observe in
southern Arlantic ocean), which explains Atianric, ancl North paci6c secondary associated biomes. As lriomes a
rheir species compliment. defined inrg.ly on the basis of physical oceanographic conditi.ns and
t'
exact boundaries are prone to alter wi
response oiphytoplankton, the
1.2-2 Local versus regional patterns .eoi.r,.,rl, EI.lSb, or other climatic alrerations that have even long
cycles. The extenr and shape of biomes and their subdivision into
pr
,Some regions are richer in taxa than others' Japarrese coastal waters are' on geographic positioning of continent
,inc*s i, very rnuch dependent
for exaiple, particularly rich in .lrachiopods and the Antarctic contin- features of coestiines, and oceanic circulation paf,tsrns, As coastiines
ar
e*tal sheives (p.ycnogon.ids). Overall che Indo-West
ri.h in sea spiders
lndonesia, has higher numbers even oceanic frontal zones are apprclximately fractal, a serial subdivisit
Pacific region, and in particular ardgnd
molluscs and probably marly other marine of units could be argucd as a valid c$ncept, however, this wou
of sp..iei o{ fish, corals, require progressively more and more derailed dam from which
arlimrls than elsewhere. on land rhere are typically more plant species
tl,an in other tropical regions, which are .onrrru.i rh-em- The establishment of provinces has followed simil
(per unit area) in the neotropics
or palearctic regions' Thus smali biogeochemical steps t0 rhose for the designation oI biomes {climatolol
i, ,urn more speciose than near(tic
of mixed layer depth, water rranspaiency, and surface nutrienr statul
areas, or u,r[um.s (though these are rarely measured), in species-rich 'fhe designation of biomes and provinces is ve
but ar a high resnlurion.
regions will contain niore species rhan similar sized areas in
species-poor
usefui (or examining many aspects of large-scale patterns in organis
,.[i.r.,r. So in acldition to effects such as species-area relationships, there ccology anrl evolution (Fig. 1.7). This is particularly tire case for tl
I GenerallY, there is a $rong
positive relationrhiP between
islypically a positive relationship betwcen local a*d regional diversity. invrrlg,,rtion o{ the relationship of species-grr:upings in one area to tho
local and reqional diversity, This relationship is useful as it enables prediction of changes at one in oth.is. One of the mosl fundamental areas of biogeographic researr
although to date this has been scale from observations at anorher' To date, exampl*s of this regional (or high
is rhe study of the degr'ee c:f endemism, i.e ' how many species
demonstrated best in terrestrial local species relationship principally have been from terrestrial systerns taxa) are unique to a Particular area?
systeml- (Caley 8c Schlurer 1997).Inorder to appreciate the relarionship of a local
history of areas. Isolation prevenrs taxa evolving locally and ther
spreading into other regions, thus the more extreme and longe:
the isolation the more intense the tendency to endemism. During thr
fragmentation of the supercontinent of Gondwana, many of the result
ing fragrnentr became isolatid. Some of the resulting islands, such at
Australia, Madagascar, and New Zealand, are famous for their endemit
terrestrial vertebrates. Geographically isolated islands such as Hawai:
and, on a continental scale, Australia have very high levels ofendernisn
in marine, freshwater, and terrestrial faunas. Another island, Antarctica.
becarne even more oceanographically and climatologically isolated, and
a high proportion of the species of marine phyla are endemic (Arntz et al.
1997). Antarctica's marine fauna has few genera and even fewer families
rhat are eodemic, which contrasts sharply with paiterns in the Austra'
lasian and Madagascan vertebrates. Young islandq such es Ascension
Island (tropical, mid Atlantic) or regions, such as the Arctic basin, have
few endemics, Before comparisons of endemism can be made between
Fig. 1.7 Longhurst,s (1998) biogeographic divisions of the marine environmeat.
The locations" considerations of the principles of island biogeography must
o"looi. bouidr.i"s oi biomes (-) and provinces (-) are shown. The biomes (and be remembered. Ascension Island is tiny and young, and even the arctic
(sANT,
provinccs wirhin) are Antarctic Polar {ANTA, A}LR}, Antarctic \festerlylfinds basin is small in comparison with the Southern Ocean (which covers
SSTC), etl"nti. Coasrat (BENG, BRAZ, CNRY, FKLD, GUIA,
GLIIN' NECS' NWCS)'
c.7A% of the surface of the globe). Io one sense, the use of similar-siaed
Atlonti. Polar (ARCT, BPLR, SARC), Atlantic Trade lfind
(CARB, ETRA' NAT&
areas would be ideal for comparisons of endemism, but what size would
SATL,V/TRA),Atlantic'!(/estertyVind{GFST,}{EDI,I{ADR,NAST),lndianOcean
c"r"rr tenari" AUSW, EAFR, INDE, INDW, REDS), Indien ocean Trade wind {ISSG, be meaningful and would this be consistent from place to place? In the
MONS); Paci6c Coastal (ALSK, AUSE, CALC, CAMR" CHIN, HUMB, NEVZ' marine environmeot biomes and provinces are obvior:s starting points
pEQD, PNEC' SPSG'
SLIND), Pacific Polar (BERS), PacificTrade'Wind (ARCH" NIrfG'
Vanfrat, and Pacific lfesterly.Winds (KURO, NPPR PSAG, TASM)'
Tabte t.2 Endemism in marine and terrestrial archipelagos (modi6ed from Myers 1997).
Shallow water amphipods and vascular plants are used as exaraple taxa from the rwo
1.2,4 Endemism realrns. For each localiry the island size, total number of species, and percentage of
endemics is givcn. The one polar localiry, South Gcorgia, is shown in italics.
At a global scale all species areen{emic,'at a whole ocean scale many
taxa lre likely to be endemic but decrease at smaller spatial scales. Land area x l0o krn: Shallow arnphipods Ierrestrial planB
Typically endemism is used in the cbptext of countrie$, a;chipelagos,
or-islands and seas but also berweJin different sorts of organisms
No. species %'endemics No. species 7o endemicj
(e.g" while all but one of the marine lizards in the Galapagos Islands are
Ne{ Caledonia 150 172 70 2740 95
less than 20"/, at the fish are endemicl' As with most aspects
"rrde*ic, Nw Zealand ?680 fi3 66 161 I 8I
of biogeography and biodiversiry, refresrial data is considerably more Madagascar 5877 3r4 7 10000 80
advanfed-compared with rhat for the marine environment, and the s0 4l 50
fiii 183 1528
values for many marine invertebrate taxa are simply rough estimates. N 4422 34
Japan 3800 300
Table 1.2 shows the similariry in endemism {except in polar archipelagos) 80 51 33 ?01 41
Galapagos
of rwo of rhe befter-known marine.and terrestrial taxa at example islands' 24 33 70 ?o
Trist n da Cunha 1.0
care is needed in the interpretation of such data, animals with long lived 32 623 ul
I Animals with long-lived
planktonic larvae (e.g. crabs] are likely, to have lower rates of endemism
Society b. 6.4 3.|
locnl ecosystern fr,,nction (Ch-apter 5). Hence the prtvalence of rare spec
1.3 Bioeliversity rhat share sirnilar fulctional roles may explain, t0 an extent, the resilier
of marine ecosystems lo environmenml and anthropogenic influences'
1.3.1 What is it?
a resrilt, species-poor systems are expected to be those thar are mr
Since the Convention on Biological
Diversiry (CBD) held at the city of vulnerable to external forcing factors, as the eliminarion of one or n
[f" l. j-*f.o, Brazil in L992, ehe at:breviated term biodiversity has species may have a proPortionateiy much greater effect on ecosyst{
L..orn" ,r, o.c.pt"d term' lnterpretations of what rhe term means vary' functioniug'
to eflcomPass
lur.ft*.riu*ty iirefers to rhe li{e on rhe planet, and is used 'Ihe realizarion that che functional diversity of benthic assemblag
main threads of
.*.ino ancl living organisms (Box 1.5). Ther:e ar:e three
may differ considerably fron'r the diversity ascerrained from t
and habitats)' organ- qr.rantification of s^:" -ate species (e-g. species richness) has led to t
t *iiu.rri,y, .cologic,t (e'g' biomes, ecosystems'
and genetic (populations' 'rniques that describe community srruc[ure a
i**rf l*.g. kingdoms, phyla, and species)' development of
injiuiau^'it, anJ genes)' Sorne genes are
of these cornponents' such as
composirion. Ta.r .- diversity (D) and taxonomic distiuctness (f
t BiodiverritY comPrises or raxonomic ranks are
ecological, organismal, and ;;;,;i;.,;tt, *liil. others sr-rch as populations
probably the unit of mosr
,re p.,rposed to be more sensitive to variatiop i1 environmenfal strt
genetic comPonents' aiin.rft ," clefine *ith precision' Species is than rractitional cliversity inclices as rhey use information derived frc
reference and ipecies richness one of the rnnst comrnonly
used
;;r;;- the hierarchical taxonomic tree upon which species identities are bas
of qLr*nrifying diversity' As with any taxonomic rank (or (Chapter 14). Taxonomic diversity is defined as the average path leng
"r.ifr"a,
i,ra."a Ui.al"ersity),'th" term'species' does not
have a single definition'
between every pair of individual organisms identi8ed from within
to generate yolrng'
An interbreeding group of organisms thar is unable sample. Incliviciuals derived from the same genus have a shorcer avera
viable, wirh any orher species is a path length than individuals within the same family' Taxonon
*i;.1, u.. themielves reproductively
suitable definition. Ho*"ve., it is sorrrervhat
difncult to demonstrate tlre distinctness is ctefined as the average path length between every pair
iatterforfossilspecieslThr"rsthetermmorpho.specieshasbeenwidely indivicluals ignoring those between individuols of the same specir
*.a (rnor. with observable clistinct structural differences, e'g. the coloul These inclices circumvent two problems rhat rrormally confound dett
and shape of a gastropocl shell) - !u1
genetic str"rdies have demonstrated ticrn of more slbtle responses to stless by standard diversity indices su
many cryptic spccies.
,[r^t .o,r..," morpl.,o-.p*cies actually comprise
.Eu.nsmall(0.1m2)sanrplcsofspecicsassemblagesinthemrrine as Shaunon-'Wiener (H') and Margalef's (d). For example, in stress
environments such as those subjected to severc organic pollution, t
.nrrironrr,*r,, can contain aflything frbm ] to 10000 individuals repres- dominant species include polychaetes in t'he Capitelta capitata compl
.".i,gf'.*1to150species.Inmostcasestherewiilbefrom20to50 of species and the gews Oltbryotrocbh. This group contains relat
species that have a similar trophic role or function. Llence despite r
presence of a high nnmber of species from within this group, t
and
,diverstty,,
essentially Iefefs to
,the weighting given to these species is down-weighted compared with a le
This term, a combination of the ',vords,triological,
used the deiinition
,the Variability p.rturbed assemblage comprising only a few individurls of the
variety of }i{e, (5ee Gaslon & Spicer 2004). The CBD
anrong living otganisrns irom all scurces including, iflter alia, terrestrial, marine and other .lo*ely reiated species bur grearer numbers of taxonomically less relnt,
aquati erosystems and the ecological cornplexes of which
they are part' this includer species (Fig. 1.S). In addition, these indices incorporate a mr'rltivaria
diversiiywitfrinspecies,berweenspeciesantlofecosystems'Gdston&Spicer(2004) cornpolle]lr by retaining the informarion derived frorn species identi
good general one if 'living' is whe..as standard inclices give no weighting to species identity such th
sugg"sted that, amonglt the many definitions' this is a
,rrered to,living and all those that have ever
lived'tb take into account past forms (the rwo assemblages with an entirely diflerenr species composition cou
vast maioritY of life)' have the same index of diversiry.
i, :a;r,'pr'rs ii'r pte'vreirrT x EllvlsollMlf(r
rhe same scale would be far roo small to assess their wading-bird or I
predators that are far less abundant and rnuch more mobile (Kai
5. Ptrylum 2003). So the time or sParial scale to be used t0 measure lriodivers
has to be specifically geared to the ecosystem, communiry, trophic lel
4.00r:
and lifespan of organisms, and even the ryPe of sampling apparatus a
protocol (Chaprer 14). Given such differences in the method of me
urement, comparison of biodiversity values in time and spnce is I
3. 0rder
2. tomily
straightforward. Certainly when making or evaluating comparisons tl
have treen made, it is vital to consider the imp[ications and bias due
l.6cnus scale effects.
0.Spetier
O EF Ill,
Sonrple B
1.3.2 Biodiversity through time
Sonple A
(oltuloling 0' By measuring biodiversity of particular types of habitars or Particu
Somple A
Somple I groups of organisms, it is easy to see that biodiversity is not static
Bt 0
rE!{XYl time or space. The numtrer of lower taxonomic unirs (species, gene
A 1 2 2
-fair.il'"' etc) has increased many-fold over the approxirnately 570*6
B I I 2 I 'ars for which we have a fossil record of macro-organis
( I L:
I L
(oI
1000
The KT mass extinction refers to thB sudden decline of organisms ot the boundary of t
Cretaceous and Tertiary periods. ln the KT rnass extinctio'r (K is for Kreriie, meani
'chalk' in German, which describ*s the chalky sediment layer from that time; I is
Tertiary, the next geologic period). all land animals over about 25 kg went extinct, as t
many smaller orgnnisms. ln tolal 1 I % of marine families were lost (which compar
with olher rnass extinctions. except the Permian*Triassi( in which 52% of mari
farnilies went extinct). The KT mass extinction rs famously linked rruith seyeral even
0E first. it was approximalely at this time that a iarge rneteorite collided with earth in t
-!0{ Caribbean near the Yucatan Peninsula. Second, the remaining dinouurs perished
B0r thi$ tirne.
a (,)
polychaetes (C) crawled on these muds -fo dare only a few rnarine data sets cover entire hemispheres
phyla. such as the priapulan Ofio'h {8) and equator.
i"g"tin, *itl, strange forms like Haltucigenia l)l' Wiwaxia (E) and Eurgessia (D' other even iust multiple oceans. Even the most robust data sets cover only r
@wl
0 100 200 rm {00 tq} E0 I00 m0 900 l00lt 0 lw ?m l0 {p 5m6m 700 8$ 9ml0$1100. 0
tt,
is so poo4y studied;
following reasoos: most of the ocean\ environrnent
in but there are few
;;;;";e.g. nematodes, may be veiy rich preyiously species
well-smdied
i**oro*irt Io describe the species; aod some
cryptic whose identi
taxa appear t'o be a complex of many sPecies'
.t I .
,::::.. .j ._'; ,.;:'.''
\
1,4 Abundance and Size l
6
! B
it $
g
:
!
40
f,
ol&?oom4@
taxa are supeficially similar, both have a shell composed of two
valves
Eivolt grnom
and they are benthic and suspension feeding animals' The brachiopods
were in;ially more abundanr and diverse than bivalves, but this trend Fig. 1.14 Patterns o( and berween, brachiopod and bivalve (mollusc) tichness.
was reversei following the biggest of the mass extinctions, at
the end- (a) Numbers of families with rime over the last 600 miilion years, (b) numbers o{ gcne
Brachiopods are preseotly most abundant.and over rhe same period, and (c) gcneric richness of brachiopodr vs rhat of bivalves befc
Permian (Fig. 1.14a,b).
and after thc end-Permiaa extinction event Dam from Gould 8c Calloway (L980).
speciose in ihe deep sea and potar regions, rwo of the few
areas where
The photograph illustrates abundant fossil spiriferid brachiopods.
bir*lu*, are not dominant. More than a hundred years ago the match
1; PATTFBNS ilu ?H MA&iNE
rfiiVt$OiltuI6Ni
i3.
patterns of richness these two T a
tharcomperition explains the a O e
""rriUtiiry a a t
-"i.*p.ai*n animals is still
io-o..oriin* debated-
regions: habitats' Io o
a a a.
has been investigated in marry dif{erent
ancl anrong anrl within a wicle
uJriety of rnarine organisms' Relativeiy few
I
a a
a
n
a ta
t. Y
.t
;;;;;-;atial and tn*ono*it po"*'nt lrave emcrgied'
one of the few
Eo a a t
a '49
UO
Eatterns that has eme rgcC
ls tl"'at of sh'rllc'n encrusring (bryozcan)
much more hierarchical at high laritude
ccm-
(polar . C eT
ffi;;t...i.,;r. ,vpitirtv hierarchv in polar
;;;;t iF,t 1'15)' ihis t"u" th't the d'r.i,ance wins virtuaily all
regions is more pronottnced:
the top cornpetitor
'fhe next highest ranked competitor
;;".;;;r.r, wirh any other species' dominant' and so on down
wins all encourlters except th;se
with the most (,oled h flei! grn(k id
;;';;;*-;ith., growrh or maximum size are usually con- r Other major isperts of ecology also chanqe drastically along time and sp
"tlt""d
shorter life history'
'l'trere have been many explanltions " nts. Thi5 (hapter discussed the evidence for jrst a few. including h
i",t"a.J ty th.i, (sr"rch as redr"rction of predotion . :ion appears to be more hierarchicai at the poles than at lower latitudes i
;ffi;; "J,r*,-i. shor"rlcl increase inbutsizerhere been little consensus of
1.i'
ti. . rhanqes with the taronomic relatedness of competitors.
-, i".t.rtitt feeding' efficiency)' rhan inhas rhe formation of colonial
Y:.
r Success can be rneasured in many ways; some orga nism type: have become succes:
;i.";;;;;;rr;c" of Jwar'sm orher
, Thui colonies of organirms
ecologically (abundance, ubiquity and evolutionarily (great geological age, mi
size' without any One effective metirod of escapirrg surface area to volume
can increase in
taxa), though not always at the same time. Many others have had some ecologica
surface area ot "r.*','#r"trr- feeding surfaces is coloniality' In virtuaily
.on.rrrrinm on respiratory or
changc in the
evolutianary success, whilst some may always have been rare - but was this thror"
or feeding 5urlaces' rnany corals),
resPiratorY
hecause the modules
rematn ;ii;;r;;;f .otoni,rt raxa (c.g. many. ascidians, bryozoans, solitarylu,icary
design or chance?
nrodules are smaller thsn similar
species,
fixed in size-
ir*-."i"*"r in size'
r Finally, evidence to date suggests the iargest organisms that ever lived are alive lod
[u, ,t-,. colony may be unlimited but the largest iand animals died tens of millions oi yearsago and the largest.inse
.fhecrrrrerrtinteresringlobalbiodiversitymeansthatthereisintense
lived even earlier. Giqantism and dwarfisrn oc(urs in (artain environrnents even nl
i",",*'intherelativeextinctionra|e$ofsmallversuslargebodied (deep sea and polar regions). The evolution of body size b complex and depends
In recent time the extinction ol of the largc vertebrates
;;;t. Tl"y far less
many factors.
,.;.* .*"i"-.ts and islands coincided with human inhabiration
,r"trro*naboursieerelatedcxtinctieinsfurtherhackintime'most
based
a]'rthc end-Permiarr' Prer{ictions
notably in the lar:gest extincrion .$ FURTHER READING
larger mamtn:rls have higher
;';;;.*il, alone l.,ave s,ggesred'ttl,at
lines q:f evidence susSesr thar adaptive Gaston and Spicer (?004) provide an excellent treatment ol biodiversity and currr
;;;;; various
;rrbabilities. This set the scene
topical issues, while McSlrel (1996) provides a detailed treatment of trends betvve
at small hody size'
frt."lrf""ritftts in {arlnas occr:rrecl melazoan complexity and evolution.
well as the nurnber of species at the stafi of
i.tl^nra irlr.n',t' in size as
. Gaston, K. J. and .1. l. Spicer 2A04. Eiodiversifl: An intraduclian (2nd edition).
,t. ir*frti*n and a(ter the various Mesozoic extiflctions' Blacknell Science, Oxford.
. McShea, D. W. 1 996. Mpl:zoar) cornplexity and evolution; ls there a trend? fL?/ufij
5A'. 477*92.
A CHAPTER SUMMARY
For an introduclion to the wider implicaUons of b,:dy dze, see LaBarbera (1986) for o,
.Themarineonvilonmenta(CountsIar99%oftheEanh,slivingspaceanditiswhere of the best revieuvs of this subject.
neariyallmalortypesofanimal(phyla)evolved'Mostr:fthe*animalsrenrainendemic
(unique)totiremarineenvironmenl.Ma5sextinctionsandradiatisnsofspeciesaf{ec1ed r LaBarbera, IV. 1986. The evolution and ecology of hody size. ln D.M. Raup and
marineorganiStlsmosidramatically'Asrnarineorganismstendtobebetterpreserved D. .Jablonski (eds), Patterns and Prorcsses in the History ol Life. Springer-Verlag,
our perception of past liie comes frorn marine fossils'
tlran thosl on land. most of Eerlin, pp.69-98.