osp:ervuno sarrour llenpar8 pue perlrJol $ }sruf, ,Iueef,o suone:)ol

asoqt rV 'sa8pu urrro-P1u Io sa8uer ulelunoul BeslaPun ag] '(q '(po

oplolq salrauropl arenbs 3o sputsnoql Jo suar uuds rrql sulseq ParaAof,
.rur*tprr e3:e1 aqr grtrn\ tsrrtuos ,leqs Pruaunuor aql o] dn sestr doats
rq1 'stusrue8ro r(uru.r dq qldap gr lnoq3norql parepdod sI luetuuorlluo
lesorsu?urrparrqt sql 'daap ut 666.p {1"leeu st {aru;rns
s.t{lreo eql
p %TS pue) retrqeil ,ru?a)o aq] ,o lsow Jleqs Ieruaulruos arll ss un'rou{
Jaap sartau psrpunq rrra; u un[ seas /rollrqs ]o ulro; arll sa{El lral
-slsora ruznbp slqr sluauuuo) aI{} ol asol} 'a)trms sll Jo %0/< Jo^ol
ruB;)o aqt pue :aueld enlq B sI guea eql lad 'lueuruo:tAua lelrlsalral aql
:{r],ra TBIIIIuEJ rsour ,(lluanbasuo, PuB satlads Suraq-pue1 E a'B sueunl{

uolpnporlul 1."I \

'quea uo rll+o uollnlo^a pue sso))ns

)ururalap leqt sassarotd aql olul 1q6rsu1 6uo4s e sea16 ,$olorq Jlaql pup surstueEro

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

,,\uvl il,uns u3ldvH)

Iuaui usj l^u:l su!JEk1tr

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
behaviour. The water column, like the land and seabed, has many sharply
changing physical features that effecdvbly form environmental barriers
that constrain rhe organisms that live there {Chapters 2, 3, and 6).
1.1.2 Oceanography
On a larger scale the water column has a strong pattern of zonarion rhat
occurs across irs full depth range, similar ro that found in the Iittoral zone.
Most importanrly in the top few to 200 metres of the water column rhere is
There are many features thar partition the ocean environment, and t
posirions of continents, islands, and subsurface marine mountain chaj
form obvious physical barriers. In the open oceau, differences in wa'
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
large-scale pamerns fhat occur, most obviously in water flow. Most de
tseabed) water is derived from the Southern Oceau. Cooled dcnse wat
(termed Antarctic Bottorn'W'ater, AAB!() sinks in the Southern Oce
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

a Thermocline! are a vertiqal
zone of rapid temperature
change in the water columri.
a psu - pract;cal salinity uniB.
(see 4.1.4.).
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.
The nature of thermoclines changes fqom place to place but in general the
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
=
lower close to continents and in the arctic due to the influence ofgreater
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

Hou15 High to low tides :i,

Days Spring to neap tides

Months 5eason.1l temperaiure/salinity/currents

E0
Years El Ni6o 5outhern Osciilation

Decad5 Climate warming, i(e sheet ral.eat q

4
lcB ages, Milankovilch cyclel, :ea level fluctuations,
Cenluries to millennia
sea floor spreading o
,.40

iVillionr of yeor ogo

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

colrnt..curr.nts flowing eastwards)' The current sy$tem of the

Southem 150

flows around the m

Ocean is dominated by the circumpolar current' which
100
50

continent in a clockwise direction' Surface currents *re Parricularly 0

I Surface currents are particular{y

i*po.ron, in mixing the water layers, which is a powerful agent that
import3nt in mixing the water
layers due to the frictional stress
.or.,,.ru stratification, at least in rhe shallower layers o{ the water
column. As well as these iarge and small-scale spatial valiations,
the
that occurs between two moving (Table 1'1)'
marine environrnent changes over a variety of tirne scales
bodies of water.

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

.- '.:
a logical fault lines whae they crystallize on mixing with cold water. Methane clathrate
--
The extent of the continental
fietf changes <ondderably with
.'. jus"a 50m change in mean
grlobal sea level'
a Past climate changes occurted
io relponse to massive methane
clathrate releases {rom below
the seabed, maior volcanic events
and after meteorite collisionr'
I Ahhough the Peneftation of
UV tight affects only the upper
{ew metres 0f leawater, it ha5 the
potential to adversely affect fish
laruae and other biota'
'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:
6^;;t; ; and 7). Changes in ice e$ent'.-which occur simultaneouslv
away entire long been thought to be comrnon in the outer solar system but mote recently it has been
*'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"
'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
[r.r, * irr", rear th; Yucatan aspects of r'' .'t climate change have 1.1,4 ProductivitY
ir.. S"* 1.3). Other prominent
irr.r*A.a increased C02 levels and ihattges
rspheric ozone thick'
Unlike on land, true plants generate linle of global marine producti<
Ti- n*t of these is strongly related to rL: ' :ture changes through
{except in intertidal saltmarshes, seagrass meadows,
and mangro'
".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
a serious issue in inicioscopic free-living single'celled algae collectively refe*ed ,
potentially
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
;;;*;;;i;L littoral,
and very
uggr.lrtiont of) single cells they vary in size from large (20G-20p
fuide from a
*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
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 ,
ilrh"., Oscillarion (ENSO). EN3Q has been rhe subject of
maior in 7972' 19821t983 and the lchupt r 10). Primary producrivity in the ma:irte environment vari
;;;ttfi. Jircorsion after evtqnts "orrl,
or"r r.uur*l orders of magnitude from grams of carbon m-2 d-1 to iust ter
iplot. An ENso t*"t'"\ around raised equatorial Pacific
of mg carbon m-2 d-1. Toral annual global phytoplankton primary pr'
""ar- "ot"t
seasurfacetemperatures,raisedcoastalrainfallandfloodinginthe ductiviry is patchy in space but is rypically highmt around conrinent
global climatic
*.*rn Americas but is also associated with knock-on shelyes and lowest in the centre of ocean gyres {Fig. 1'5} (Chapter ?
anomalies,Oneexampleofsuchananomalyistheweakeningofoceau secondary production (mainly by zooplanktoo Fig. 1.5) is linked directly
,
."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

proporrion of rhe global rnarine environment on the continental shelf

are con-
lCnrpr". 7). The lowest levels of total annual productivity
..rrrrrr..l around the centre of oceanic gyre$ such as the Pacific Ocean
basins north anrl south of the equator. seasonally high productivity is

@ "sm [-**l zm.mo ffi.-:1;E rm.zso [-J

Fig, 1.5 Pamerns of annual phytoplankton producrivity in rhe global oceanic
enyironment expressed as gCrn-2year-t lAdapred from B.:rnes Er Hughes 1999).

associated with much of the shalkrrv polar or high temperare coastline e

high annual prodlrctivily is particularly norable acloss the tropics of mr
of the coastal Indo-!7est Paci6c. Patterns of high and low prodtlcti!
have a largely predicrable distribution and are likely to be a major car
in rhe way other marine organisms are arranged globally. The proc
of prirnary production is dealt with in detail in Chapters 2 and 3.
This first chapter examinss large-scale patrerns in the distribution
marine organisrns. The emerging discipline of macro-ecology has gro'
to quantify, analyse, and explain the parterns of organism groups
large scales (Brown 1995). Biogeography is rhe science of geographi,
and l'risrorical relationships berween and among different organis
(taxa), and is a natural starting point for rhe examinarion of large-sc'
prrt*'rnr in the marine environmert. Later chapters consider patet
and processes that occur in specific systems.

Fig. 1.5 Macro- anri nricro- phyroplankton and zooplarrkton' Examples of

ml.ro- pri*ory prcducers are subtidal kelp (a), intertidal red and brown macrophytes
(b) and a u.rgrri, meadow (c). Examples of micro- primary producers are a diatom
(e) aad a radiolarian (f)- Foraminiferans are protoctist
1.2 Biogeography
iCocane;s) (dl, a silico-flagellare
micro-consumers (g). Example zooplankton {h) includes a iellyfish, a polychaete worm' Biogeography explores the lisribution of species, how groupings
a euphausid crustaiean and a ctenophore
(comb i*tly) and various invertebrate larvae (ii.
such species form distinct ecosystems and their geographical limi
phoiogrrphs, (a, b ao4 c) David Barnes, (d and e) Sandra ll(ilks, Bricish Anarctic Survey,
(f) Clalre Allen, British Antarcric Survey, (g) Mark lfilliarns, {h) Simon I}rockington,
Individual species (and in some cases higher taxa) each occur across
(i) David Bowden' certain geographic range. Under experimental conditions, many speci

a At the edge of their
geographical range, rpecies are
much more susceptible to local
extinctionJ that oc(ur with
changing cnvironmental
(onditions.
can endure environmental conditions (e.g- temperature, salinity, sedi-
menrarion, and pressure) far in excess of the conditions in which their
populations naturally are found. Theie are many reasons why species
are not found across tle entire range oftheir physiological tolerance. At
the edge of their geographical range, species are much more su$ceptible
ro local exlirctio$ rhat occur with changing environmental coadirions,
hence the presence of a species along a gradient may simply rellect the
time since the last local extinction event. At a larger scale, there is
evidence in some groups of animals that the geographic ranges of
species alter along major gradients, and ultimately influence biodiversity
(Box 1.4). Species and higher taxa tend to form characterisric groupings
by regions around the globe, as well as in particular habitats. The major
theme of biogeography has been to'identify and characterize the
geographic groupings of species and the biogeochemical conditions that
make them differ (Longhurst 1998).
Each species occurs across a certain range of latitude. Roy et al. (1998) compared
the number of gastropod species with latitude along the Pacific (blue line) and Atlantic
(red line) coasts of North America (iosert plot). 'they then investigated whether these
species+ichnes patterns could be explained by specks at lower latitude having
narrower htitudinal ranges (Rapoport's rule). They found, however. no obviour link
and furtherrnore latitudinal ranges of many tropical gastropod rnolluscs were higher
than those at high latitudes. Roy et al.'s (1 998) studywas the mostserious testof Rapoport't
rule and oneofthestrongestdatasetsforlatitudinal rangesof speciesforany marine group.
:E
Biogeography has been of inrerest ro both terrestrial and mar
biologists for more than a century since early boanists began to ma
vegetation sEuctures to regional climate rypes. A succession
pioneering botanists classified groupings of vqetation structures
climare into a series of life zones, biomes or ecoregions according
environmental characterisrics such as precipitation, temperarure, :
humidity. Similar advances were nor easy in tle marine environm
where mismatches in differenr aspcts of biogeographic knowlet
occurred even in the most accessible $ysrems. Ir the last few deca<j
underwater scientific instrumentation, advances in marine sampl.
platforms (ships and autonemous underwater vehicles), and m
recently analysis of high qualiry ocea*ographic data (Coastal Zc
Colour Scanner, CZCS) from rhe NIMBUS satellite have revolutionir
our understanding of ocean surface biogeochemis[y. ln rerms of mari
climate, the global marine enyironment has been considered to consisr
three broad domains: polar, temperate, and tropical. In contrast,
term$ of their fauna the early zooiogists divided rhe globe inro d bro
realms: Nearctic (Norrh America) and the sirnilar Palearctic (rrr.
of Eurasia and Northwest Africa), Neotropical {Central and Sou
America), Ethiopian (Africa), Oriental, and Australian. Boundaries a:
subdivisions have gone through many subsequenr refinements. Althoq
designed around terresrrial bioiogy, rheories of island biogeograp.
represent a good inrroduction ro maoy of the inportant facror$ th
determine the distriburions of organisms.
1.2.1 lsland biogeography
The global rerrestrial environmenr is formed of e series of islands th
vary in size, age, isolation, nature, and history. The fauna of islan,
refiects these factors. The larger, older, and closer to a continenr i
island is, the greater the number of taxa present. MacArrhur ar
'!0ilson's (1.967) theories
of islaud biogeography explained many r
these concepts. \[hether marine or terrestrial, animals or plants, tl
number of species {S) {or higher taxa) increases proportionally to t}
area of the island (,4) according ro the Arrhenius relatiouship:
Log S : c * z ( log A), where the values of the constanrs are
c=c.0.3andz=c.0.16.
Thus on a plot wirh log scales on both axes, the relationship approximatr
-
to a straight line shown for example by plant numbers on Australia
islands or even {approxirnarelyi for species on contimnts. At the other en
of sparial scale a number of authors have immersed serdement paoels c
various sizes into the sea and demonstrated simila; increases of specie
@ f : PATTERNS fff lrig MARIME EIlVlkOr\]i/IENT
e lsland size and itolatiofl are
important determinants of
specie5 ri(hness"
I GenerallY, there is a $rong
positive relationrhiP between
local and reqional diversity,
although to date this has been
demonstrated best in terrestrial
systeml-
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
*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
there is oit.n much 'noise' in species*area plors of sampie
data'
'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
islypically a positive relationship betwcen local a*d regional diversity. invrrlg,,rtion o{ the relationship of species-grr:upings in one area to tho
This relationship is useful as it enables prediction of changes at one in oth.is. One of the mosl fundamental areas of biogeographic researr
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
local species relationship principally have been from terrestrial systerns taxa) are unique to a Particular area?
(Caley 8c Schlurer 1997).Inorder to appreciate the relarionship of a local

Fig. 1.7 Longhurst,s (1998) biogeographic divisions of the marine environmeat.
The
o"looi. bouidr.i"s oi biomes (-) and provinces (-) are shown. The biomes (and
(sANT,
provinccs wirhin) are Antarctic Polar {ANTA, A}LR}, Antarctic \festerlylfinds
SSTC), etl"nti. Coasrat (BENG, BRAZ, CNRY, FKLD, GUIA,
GLIIN' NECS' NWCS)'
Atlonti. Polar (ARCT, BPLR, SARC), Atlantic Trade lfind
(CARB, ETRA' NAT&
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,
MONS); Paci6c Coastal (ALSK, AUSE, CALC, CAMR" CHIN, HUMB, NEVZ'
pEQD, PNEC' SPSG'
SLIND), Pacific Polar (BERS), PacificTrade'Wind (ARCH" NIrfG'
Vanfrat, and Pacific lfesterly.Winds (KURO, NPPR PSAG, TASM)'
1.2,4 Endemism
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.
Typically endemism is used in the cbptext of countrie$, a;chipelagos,
or-islands and seas but also berweJin different sorts of organisms
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
locations" considerations of the principles of island biogeography must
be remembered. Ascension Island is tiny and young, and even the arctic
basin is small in comparison with the Southern Ocean (which covers
c.7A% of the surface of the globe). Io one sense, the use of similar-siaed
areas would be ideal for comparisons of endemism, but what size would
be meaningful and would this be consistent from place to place? In the
marine environmeot biomes and provinces are obvior:s starting points
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
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.
Land area x l0o krn: Shallow arnphipods Ierrestrial planB
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.|

planktonic larvae have lower rates Eermuda 0.5 51 1? 165 I

ihan thor" wirh only limircd oppornrnirie$ for dispersal, such a$ animais
of endemign than those with lreland 820 263 1210 <l
limited dirPersal, such as animalS
that brood rheir young ie.g. amphipods). Nevemheless, levels and Par- South Georgia 36 151 35 25 4
drat brood their Young. terns of endemism provide a powerful insight into rhe evolutionary
Hffi 1: pArrEElJ5 tN r!-tE lfiAflII'iE
Erivli{o*iliE'rir
W they are often geograpliically species within the sample, but a few species will accounr
for rnosr
but so are the coastai zones of islands as rhe inclividuals. This means rhar.many different species within a saml
variable"
j degree of endemism is often a most important
disdnct. The .-
l nI rnnsrrveiion are likely to have similar functional roles within the assemblage' T
-L^
nulri.uturty in ihe ^^--;r-.^rinn
considerarion of conservarion issues.
iss*es. I-Iowever.
I-Iowever,
raises rhe ecological question of 'what is the purpose of so mu
;;;;;it";t between distinct regions are often based on eirher species duplicarion of similar ecological roles within an assemhlage?' Perhr
richness or diversicy per unit area' more intriguing is rhe notion rhar some species are redundant such tl
their. extirpation fronr a defined area would have no consequences
J

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

). This represents less than 5Y" of living specics past a

,l
0 2 6 (1'u.
t present. l:lowever, we can only estimate the number of species rhat rn
F
have existed at any one rime, as for many the fossil record is poor
Totol Poth length [P) 33 9l
absenc. For every ore species for which we have fossil evidcnce, thr
n-r-f"'rii'i.u;''0 21 2l
may be a thousand whose existence remains unknown. To complic:
157 433 mamers furtlrer, che duration of the existence of a species can vary fr<
D'=P/L
approximately 1" million to t0 miilion years" From the limired evider
Fig'1.8 Anexarnple ofthemethodofcalculatingD.fortwodifferentsamples;A,rypical
The hierarchical linkages are we have of fossil taxa (Fig. 1".9), the overall increase in numbers
site artd B, typical of an unpoliuted area'
"i, ,air,"J path l"ngtti each level' The species (A' B' C' D' E' F' G' \{' x' Y' z) animal families has been unevefl> punctuated by a series of sudd
ti"r^|. *rtf,
the
" to as extinctions, and the larg
catculared. D- is rhe decreases. These decreases are referred
;;; ;;;; #* the p-ath lengths betwecn each speciessre
totrl paih length divided.bv the total number o[ linkages' occurrcnces as mass extinctions, which themseives have interesti
;;;;;"
parterns in time. The rate of speciation also appears to have be
biodiversity..is the scale at variable and maior radiations have generally occurred 'soon' after t
A principal issue wirh rhe quantiHcation of
There ale vast numbers occurrence of exrinctions. o.ne theory to explain this sudden radiation
which measurement or sampling is uhderteken.
in time and space varies over mafly 'barrel filling', a term used to refer to the rapid dilersificariorr of biota
oirtual., of biodiversiry whose scale
ro rhe rypes of organisms under inyesr- fill the many empty niches fotlowing an extinction event' The Partt
.rJ.., rnagnitude according shown in ligure 1.9a shows the trend in rhc nurnber of animal famil
"rF;t i sp"cific community' there is a standard method ior
;;;;. Samples are repeat- across tirne and uses data pooled from all animal {arnilies- Such gene
Jdmating how representative sampling has been'
new frorn each successive sample figures mask the different patterns that occur for tach of the compont
;;;rJ;,
-pr.".a ,nd the number of species
!7hen species animal groupings. For example, trilobites (Fig. 1-9b) were one
i, against rhe cumulative number of samples.
an asymPtote)' the more abundant and species-rich benthic animal types found
nGU*r, nJ lor.g", increase (the curve reaches
the
(though.this does make a number of Palacozoic seas, but it seems that rione survived the biggest m:
;;lt;g l, gaugid as sufficiert 'fhe extinction that occttrred at the end of the Permian period' The panel
;;;"*|;"''t| {ttit i' known as a specics accumulation curve'
of increase and decrease speciation differed with both the taxonon
are appropriate for sampling different
,."Uiri" tt that different scales \flhile a 1m2 sample would be level considered and between different specific taxa. For example,
organisms, communltles and ecosystems'
the richness and abundance of tiny in(auna, the bryozoan orders Cheilostomatida and Cyclosromatida, contrasts
iol n. ,rnit ro
turg* assess
"
1: PATTEfiNS r$ THE
MASIilE;NVlRONtulgi{T

(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 (,)

mid-Cambrian rocks there has treen a sequential series of dorninance

higher taxa, such a:i trilobites and brachiopods in tlre Palaenzoic
l0
ammernites and rnarine reptiles in the Mesozoic. Since the Creracec
rhe cheilostome bryozoans (Fig. 1.9c,d), gastropod molluscs, and tele
0 0G
"?00 fish have dramatically radiated in richness and disparity, a pattern a
Iimo (nrillions of Ymn ugo) reflected in the radiation of land mammals. Some taxa, such as
corals and sponge$, havc been speciose and abundant rhroughout rl
with time (miilionsof yea*)' (a) Overall rumbers
Fig. 1.9 Changes in taxon richness fossil record to the present day, Other groups such as the priapu
families' aad
.t' tiifossil triL:bites, an extinct phylurn' (c) numbers of bryozoan
f
worms, exh.ibit remarkably few morpl.rological changes over nea
"i'fr*,f genera' The blue line codes for the order Cheilostomatida
iiirr*fr"r, of bryoz'oan Cyclosromntida. Plot (a)red*wn fromJablonski (1999] 600 million years,.and remain a minor group in terms of disparity a
*.lrd rhe red tin" for the orrlcr
(d) redrarvn {rbn: sepk,:ski er rrl. (2000). Phorog*ph: David
Barnes.
il ;i;" [i ;^d richness. Based on evidence from only the Cambrian period, or even t
from rhe whole Palaeozoic, ir would be difficult to predict which of
natterflscanbeseenbothamongandwithinrepresenlarivesofthese
t"ra".t. higher raxa would be successful today or ro predict which phyla mi1
genus level patterns contrast $trongly'
arrtiretlrlore family and come to donrinate in 50-100 million years time. While palaeonrologJ
,r.r*r^rr, in rclation to the KT mass.extinction 55 million years ago
are primarily interested in the pactern of species rhrough time, for b
irig. t.1., cl; Box 1'6)' logisrs rhat work with present-day taxa, pa*erns in space have attract
'- (init* <ruerall putt..r,s of lower
taxonomic levels indicate a subst&ntial
most attention and debate.
case.for the higher taxonomic
in.r"nr",,vidl time, this may nor bd'\the
Gorild p.pularized the work of several
i.r.ii *.L as phylum^ Stephen J. f*ssiliferous ar"Id ancient out-
that investigated
,.i"r.,rlrru a particuiarly
1.3.3 Biodiversity in space
.,"n.rrockcalledtheBurgessShale(G<luld1989).Tlrefossilorganisms
,."|^f.a in these rocks were not overly speciose but rhey were
very
How many species are alive in the global rnarine environment toda
liri"r.nr from each other. Most of the piryla known today
are repres-
ln roral about 2 million $pecies of organism have lreen described to di
thete were nlany oddities
-.,r.J i" early Cambrian Iocks but in additiorr from all habitats, of which most have been terrescrial insecrs or plan
phyla and resuited in many
,t,*, .outa nor be assigned easily [o existing Terrestri;ll organisnrs and habitats have been much bett*r samplt
function. The nr"rmber .f
.rlir-*irirr.rpretarions of hody form and but many trelieve rhar they will still dominate species glnbal spec
,,*,0..,*n'inthisear:lyfaunahasattractedconsiderabledebate,but richness, even when marine hahitats are better studied. Nevertheless,
shale unprecedented diversiryof body types is important fo remember that the seas are much richer in
I Organisms in the Burgess ,u-.r.'i, unrioubtecliy an almost high
were subied to early different organisms are 'lumped' into existing taxonomic levels (e.g. Phyla and Classes)" Ar present, providing estir
{orm iJ;p*i,yt. Whether
rnitiflterPretations of body new taxa are two extremes of approaches taken by
ates of the roral nurnber of species in the ocean is problernaric f<rr t
function and tontinue t0 *,r,.p, or 'split' into Marella (Box 1"7) in
and the clominance of the enigmatic
qenerate debate'
i""u"o*io$. Since
 debate exists as to the cause of such a cline (trend) or whether it is r
or partly an artefact of confounding factors (e.g.land area)' Dara frr
Maretta \A\ an enigmatic species t'longing
to an unknown phylum was the most the marine environment is even patchier, but similar debates rage o
commoninBurgessShaletocks(Cambrianmudstone)'Eartyrepresentativesofextant rhe global latitudinal pattfrn of increasing species richness towards '

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

were stalked. such as Dinornischus iG) or were

hrge swimming predators
strange fornrs
strange shrimp) was originally
rorthfrn hemisphere and, even then only the American or Atlantic coar
ls Anom/aca ris lH). Anomlacans (whose name means
such
- wtrich were thought to be One of the best-known species-laritude data series is that reported
described frorn iust the curled front-most appendages
Roy er al. {1998) for North American gastropod$. Casropod spec
,f',rirpr.rttcircularmouthandmainbodywerealsodescribedasseparaterpeciesuntil
one larqer organism' Other common richness is considerably depressed at high latitudes (Fig. 1.10a) I
all three fossils were realized as belonging to
equatorial values are also significantly lower than those at iatimdes fr<
animalsatthistime(andwellbeforethisperiodthroughtopresentday)werecnidarians
such as the iellyfish (l) and the sponges
(J)' 15 ro 30 oN. North Adantic data for bryozoans and sea-weeds a
decrease in richness towards the North Pole, but studies elsewhr
suggest that Atlantic taxoo richness levels are flot representative a

@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'

fication will require extensive genetic studies- Along a spatial scale

;;;;;;r, biodiversiry is essentially considered at Pqint {a diversitv}'

of
W ]0
f,abit.t lp diversiry; or regioqal (y diversity) levels' Discussions r0 200 t00 {00 50{l 600 7m 800 91x} t[s ll00 lzl]0 0 15

deal with 7 level

ii
;;;i futt".nt of biodiversiry almost invariablv the gpecies Fig. 1.10 Latitudinal and longitudinal pamems of raxon richncss' Numbers of: (a) spec
dui. fo, ,bou, *o centuries scientists have considered
of gasuopod molluscs, (b) sPecies of decapod crustaceans' (c) species of sgbellid
ro the tropics' This is
richness of organisms to increase from the poles polychaeie", (d) species of bivalve molluscs, and {e) genera of hermatypic corals' Plo
environment for
;;;il by"rnbur, data sets {rom the rerrestrial
x
iedrawn from data in Roy et al' (19981, Boschi (2000), Giangraade & Licciano (200
pii"rt, ior..ts, birds, reptiles, and mammals' However' considerable
Crame (2000) and Stehli 8c !flelts (1971) respecrively'
@ 1: PATIEFNs
lhi fHE MAR'SIE ET'T\':AONMENT

thus wide-scale generaiizations should

not be based on such data alone' 1.3.4 Biodiversity and ecosystem functioning
prrr"rn, crustacean (e'g' crabs and lobsters) richness The species rhat constitute ecosystems can be assigned to function,
^h;;;;;*of coasial decapoclacross the Americas (Fig' 1'10b)' and demon- types, srrch as pioneer encru$ting suspension feeders (e.g' some pol'
been described
$trate a strong decrease in richness
lowards the Arctic and a weaker chaetes and bryozoans), competitively dominant encrusring sr-rspensic
,l*.r""u. ro*urdu higher southern latiludes' feeders (e.g. some sponge$ and ascidians), benthic zooplankton feeder
-"i;;;r]; (ilig. 1.10c)
truly glJbai data series are rhose for the bivalves (e.g. some anemones), deposit feeders (e.g" echiuran worms), scavenger
corals (Fig' 1'10d)' Both (e"g. amphipocls), mobile carnivores, and orhers. The concept of bi<
forille reef-building (hermatypic)
gradients' but iongituciinai diversity can also be used in the conrexc of the numbcr of function;
I Part of the Problenr o{ ; ;;;- gioups show clistinct iatitudinalboth pattems radiate out from
^r,a-.rri.r,nr.,
discerning gradienis
in cliversity i5
;;J;;;t:*ear .c1t'atly strong' Basically groups within a system. Thus the functional diversiry of an ecosystem
the patchy coverage
of malor .hotspot, (in contrasr to terresrrial planrs, which higher if the number of funcrional groups is higher and/or the nLrmber c
groups ln the ilil;:;[;";h.
animal and Piant
,.-li.fr*, in rhe Americ"" t'opit';' Clarke {1992) suggested rhat while members of each functional group is higher. Higher funcrional diversit
maiine environment' evidence for a siruilar should nrean that ecosystems are more robust fo change and enviror
;;;;; a clear northern cline in species richness,
hemisphere was at best inconclusive' and mental stress. This is currenrly a hot topic of research, which seeks t
;;;;;.;-i, the sr:utherndara ..in[o.."s rhe existe'ce of tl"re northern answer the quesrion 'if the number of species in an ecosystem decrease
;;;i;, so roday. New
southern decrease for cloes this affect its abilicy to respond to change?'iSee also Chapters J
trend, but while some sruclies have supported .a
other studies demonstrate
clecapod crustaceans anti bivalvi molluscs'- 7 and 14.)
' pole' The
*r,'r*rrr"o"d molluscs increa$e in ri . towards the
' rnbers of some marine
Some species have been described as 'keystone' ro their comrnunitie
(Chaprers 5 and 7). Such species when experirnentally removed can hav
;;;il o.eun h,u disproportionatell
notably the Pytnogo"u, Polych'' --i
(Fig' i'5i top right)' majnr ecological consequences for the other species within the syster
;, of a glotral pa$ern -in
fru.Liopoa*, and Bryo'oa' Dtbot* on causes (Pace et al. 1999). Few species, even those considered as 'keystone', hav

the sea would seem premature given

uncertainries surrounding the been demonstrated ro have a pivotal role in their comniuniry. Snm
hemisphere. one.such simple hypotheses have been generated, however, to predict change ir
evidence fo, consistent" pafferns in
the southern
-",r,"*r",forbothtirelatituclinalandlongitudinalpatrernsofrichness ecosystem functionaliry with species number (i.e. whar happens fol
a The Southern Ocean has intheseacefltresonseabasinage'whichmighrexpiainwhylatitudinal of lowing removal of any random species). Examples of experimenta
disproPortionatelY high
numbers arctic (ir is still in rhe process diversity-ecosysrem function studies in Iow richnes (estuarine) system
PolYchaeta'
rrends are srrong rolvard, th" /o.rng
of Pycnogona,
heing invadecl) bur weak tcwards cllder
Antarctica; are given in Chapter 4. The redundancy hypothesis predicts that ar
BrachioPada, and BrYozoa' latirudinal cline in species richness is also increasing number of species increases ecosystem funcrionality Propor
ln the deep sea, a northern
o{ a decrease in richness tionally less as the number of species rises ro a point when furthe
evident bnt th.r" have also been suggestions
ir"- rir. shallows down into deep watbrs. (e'g' offforNorway)
Support this came from additions have no net effect (a curve to asymptote). If this is correct
has casr species could be lost from species-rich ecosystems without any loss o
ffi riyttrf samples, bur ofsampling',elsewhere2001)' Very shallow faunas functions. This is termed the functional redundancy of species. Suppor
,t"r;; ;. the generalitv "1tt"'l'4Cray age between 14000 to
;;. ;;; (maior tha"ges occurred ht the.lastconstant for such an idea comes from the response of many coral reefs tr
recluire reinva,sions and hcrbivore losses. In the Caril:bean, losses of herbivorous, manatees
100b0 y;rs ago) as '"f leutl changes
to the shelf edges turtles and fish have occurred but maior systern deterioration
i...*r,... n,axi*n have repeatedly bullilozed cr:lonists onlJ
quite ybung as rhe flow of occurred when the last big herbivore populations {sea urchins) crashe(
ii;frrot-t 7). The d*tp-"' fauna
also rnust be
is-relatively recent (older than and resulted in overgrorvth of coral by algae' Other hypotheses sugges
O Ttre oldest and hence ri(hert
llyg"r-lra., cold warer from Antarctica
t0 shallowshelfenvironments,butonlytensofmillionsofyearsold).Thus rhat any species addition or subtraction changes rhe level of ecosysten
fauna might be expected
margin o{ the rheoldestandrichesrfaunamightlreexpectedatshelfedgespr:csentlyat funcrions (rivet hypothesis) or thar funcdonaliry is driven by specier
occur at the
, a.p,f, of about 200 to 300 m- The richness of large (mega)benthos
and inter;rctions rather than the species per se (Box 1.8). All of thesr
continental shelf'
and away from shelf edge depths. Any bathy- hyporheses suggest that biodiversity is intricarely linked to ecosysten
fiJ';n...rr*, rowards
*.t,i..lin.isr.rnlikelytobeassharpordirectionalasdrealtitudinal srability- Intrinsic to the idea of ecosystem functionality, ecosyster
richness clincs in the terrestrial environment' trophic and food-web $tructure, are the numbers of individuals of eacl
species present within the system. The latrer also exhil'it large'scale
patterns in time and space.

e Low diversity habitats are
typified bY high ahrndances of
single sPecies, which are
themselves highlY variable in
space and time.
fuactions increase with species richness but
At low levels of species richness, ecosy:tem
species nur*ber according to the
,t higl, t*"ls ecosystem functions stop increasing with
(decrease rkhness from 32 species A to 28
rivet-theory. 5o if 4 species are removed
would sugEest measurable decreases in ecosystem func-
,o*i., ,) ihe rivet hypothesis
suggests that an experimenter would find no
ti'rns, *her"rs the redundancy theory
change in ecosystem funcilon'
.t I .
,::::.. .j ._'; ,.;:'.''
\
1,4 Abundance and Size l
t.4.1 Scale, time, and sPace
$ome are not' Abundant animals
Some animal$ are very abundant and
rhan
,r." very small, iuch that billions would weiSh less -a :iJtgle
"fr"" of some of rhe rare larger ones. Many low diversiry habitats
i.ivia"rr
,r. irrrr.ra characrerized by huge numbers of a worllt$
few species (e'g'
in marine Age fueor!
UrarrA;, snails on intertidal mudflars, capitellid
of
The abundance
ffi;*; or sea cucumbers in deep ocean trenches)'
in time and space. Thus
abundance of a taxon has so far only been considered as an aid
;;;;;t;;;is very dynamic at iertain scales quantifying diversity, but many large-ecale patterffi in the sea occur n'
one year and almost
ii i, porl*rt. for one ,p*titt to be super-abundant berween but within taxa. On land, seasonal changes in the abundance '
rhousands of
;bJ;;i" next, whilJ it is not uncommon toanncounteradiacent sample' The
many species are obviouq and in the sea this is most clearly seen
individuals in one sample and hardly ary in terms of seasonal blooms of phytoplankton. Annual changes in d
 't: PATTERN5 lN;HE M'qRlNE
=Ftvl:loilFrEi\'ir
ffi
ll lnterdecadal cytlic flu(tuationt
in the abundance of inuertebrates
such as starfishes and sea urchins
have a strong influence on the
ecology of the systems in which
they o(cilr.
I of taxa as well as species richness. changes in the abundance levels o[
Patterns of abundance are
often decouPled frorn Patterns of
species richness.
abundance of organisms are ofren due to differential surYival
of year-
classes (cohorts) of young thar *te related to environmenta[
conditions
(e.g. temperature), abundance of predators and comPetirors' There
are a
number tf [ong-r"r* data sets that demonstrate major inter-annual
Long-
ciifferences in dre abundance of animals across the world's oceans.
rerm dara sets of fisheries landings demonstrate the cyclical nature of
'good' and 'bad' year-classes in various fish species such as salmon'
.ld, *nd herring (FiS. i.11). Cyclical pafterns have also becn observed
at interclecadal scales or iinked rc ENSO events, such as sea stars
reefs) or remperate North
{Crown-of-th orr$ Acafithttster oL1 cora}
(chapters 7
American sea urchins (srrozgy lacentrotusinrnacroalgal beds)
and 10).
.we
can gain a much longer-tirne scale perspective by reference to
the fossil record that enables us to compare patterns in the abundance
Pafierns
species form the parterns of mass extinction in the fossil record.
of alrun,lrnce ar. oft.n decoupled from patterns of species richness' Two
examples rhat illustrate this are the present-day Anrarctic shallow ben-
rhic fauna anrl in the fossil record (Mesozoic to present-day) of cycio;
groups
stome bryozoans. in terms of the numbers of taxa, many benthic
of animals f<lund in the Antarcric are dominated by those that brood
their young (e.g. sea urchins or gastropod molluscs, see Fig' 1'12)'
Ho*.u"r, ih* *ot, abundant species in these grolrPs are broadcast- rhar one of the fundamental changes drar occurred in camlrrian ocea
spawners {animals that release their gametes directiy into the
rvater c.550 * million yfars ago \tras the initi*rion of competition driven
coiu*rr;. Stuclying the fossil record, lr{cKinney er al' (i998) found that the rapid increase in the abundance and diversity of marine organisn
the proportion of global bryozoan genera and species in the order Investigation of past evidence for competition is problemaric
Cycltstcmarida, has steadiiy decreased relarive to the o{der Cheilosto- McKinney {1995} measured the oulcomes of spatial interference cot
matida over the last 100 million years (Fig' 1'13a)' Prior to the cnd- petition belween cyclostome and cheilostome blyozoans preserved
Cretaceous mass extinction (KT boundary), the san'te pattern
was iossils (Fis. i.13c). Cheilostomes consistently out-competed the cyclt
evident in abr.rndance, that is cheilqstomes became progressively domin- tomes in c,66u/, ol encouoters. This suggests that one of these Sroups
ant in terms of bryozoan abundan\,g. Horvever, imciediately after the bryozoans hrrs had a clear, but not increasing, cDmpetitive advantal
KT bclundary, rhe cyclostomes becam'e very abundant again (FiS. 1.13b). Both of these organism rypes are extaflI (alive in Present day), hen
This recovery lasred only a few miliion years, but is a clear illustration Barnes and Dick (2000) were able to look for similar patterns across t
that historical abundance and richness can lre quitc separate. Both worlcl, but found no such constancy in the outcome of competiti'
cyclostomes and cheiiostomes are similar ryPes of orgaoisms
(encrusting, (FiS. 1.13c). There are two possible explanations of this discrepan<
benthic suspension feeders), which occur in the same habitats. com- greater variation in competition in modern assemblages or constancy
peririon for resources (space in che case of bryozoans) is thought to be our.o*. within rhe rype of environment thar preserves well in the fos
ih. *nin driving {orce rhat underpins major temporal patterns that are record (warm shallow continental shelf)' The latrer seems more Iikt
repeated in many co-occurring organism grouPs' and as such makes it clifficult to interpret the implication of McKinne;
(1995) long-terrn palrern of compctition, as the dominance of cheil<
tomes would then depend on the proportion o[ warnr shallow sh'
1.4.2 Competition compared to other suitable habitats, which are unknown.
Competition for resources can take many forms (Chapter 5) and can Perhaps the most famous example of inferred competition is th
drive Iarge-scale biogeographic and biodiversity patterns. Many consider between (articulate) brachiopods and bivalve molluscs- Both of the
 th)
of the two pafferns was held-uP as a 'classic' examPle of competit.
driving rhe gradual replacement of one group of organi$m$ by anor,
$uperior competitor. Modern bivalves and brachiopods comPste
e- resources in some environments, notably in NEw Zealand fiords.
3,s
60
these situarions bivalves seem to be generally superior but the outcon
9S
Ee {0 6-g ,t0
of competition are complicated by predation and other factors.jGould
EE =g
Calloway (1980) reanalysed the fossil data and found that, except at
:16
v.6
20 !:.6 n
end-Permian exdncrioa, there was no regativc reladonship betwt
OL
-I50 the species richness of the rwo groups (Fig. 1'14c)' Furthermore, d
actually {ound a weak Positive correlatior\ so that there were mr

6
! B
it $
g
:
!
40

f,

----:ifr--{o .60 '10 "10

Iimr (millionr Yeorl

and (b} relative

fig. 1-13 Cyclostome and cheilostome biyOzoans: {a) raxonomic richness
years,\end lines plotted from data in McKintey
*irnd"n.. or", the last 140 mitlion
Arrow shows timing o{KT mJhs cxincdon, das6ed bludline is species
tfgSAt"
"irf. rti condnuous red lin" is genus ouoibtt' Photograpbs show competition for
.r*U.r
spacebetweendretwogroups'McKinney's(1995)datafromfossilassernblages{blue
(c). competition in
ci..les) sugg"rt cheilosromes win c.66yo ol comperitive encounrers
iirr", (whire circleg shows much more variabiliry, data &om Barnes & Dick
^".riurrg.s
(2000). Photographs: David Barnes'

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

bivalve species' The positive rela- (ol

brachiopod species when there more The
;;;t; ;";, as the authors acknorvledged' probablv not incalrsal' a a e. e

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

hierarchy' If it *'ere not for.the influence

of disturbance'
;;;^;;;ri;r".e availaF:le space' In this
;;;;;;;t;;;oulcl ultinratelv monopolize all the 1'2) and ice-scour' is
Ioronomir divergontr of ompelitots

;;;;;tr;rbance, th'o''gh wave action.(Fis' has been described for

Fig. 1.16 competition asymmetry with taxon divergence in shallow seas. Propofiior
*otrr^ri f", rh. muinttnJ"ce of biodiversity as coral reefs'
sparial competrtors demoustraring mutual aggression decreases with increasing
sorne low-latitude communities'
In conlrast' on Jamaican ,el*,"dn"s, in remperare bryozoans (red fitled circles)(a). scded by Nei's genetic ind
species pairs
ru; found that in many intera.ctions between
Oiiyl (0.273 = within family, 0.54: wirhin genus and 0.96 = within species), the relations
A number of studies hlve reported similar seems rohus!: ir aiso holds rrue for Antarctic bryozoans (white circles) and temperar
there was no dellnitive winner' asciclians {squares}. (b} Plot modified from Barnes (2003),
I At hiqh latitudes, diversity is
for amons corals or sponges' In these
disturbante ;;:iil;; ;;.n*pttitio''t space
to each
seem to be able to exist in close proximity
maintained bY

whereas lower latitudes biological

siruations maily specles resources acquisition' In addition, very differenr species compete for t
biological interactions
int6racti0n5. o,t". nna biodiversity is m*i*tained thro*gh same resources, for exan:ple. humans' seals, and seabirds compete {
;;;r;;^ ty pnysic'il disturbance' 'lvlost investigations of competition
sand eels (Ammodytes spp.)" Barnes (2003) examined how the inte nsity
closely related competitors' often arnong
species
fr-r" fr""r, l"r**.n -makes sense' as similar comperirion varied rvith relatedness in tlvo grotrps of marine organisr
genus' Inruitively this
belonging to the sarne methods of {e.g. bryozoans and ascidians) in two regions of matching latitude
(nort
it
;;;;ffi iir.ty ,o hnue'th" n-rost similar requiremenrs and
'1. western Europe and the Antarctic Peninsula). The outcornes of compe
'i rion were as asymmetrical between organisms from differenr kingdor
(e.g. algae and ascidians) as they were between animals of differe
*.
families (Fig. 1.16). This.study showed that intraspecific comperition (tv
comperitors of the same species) was less likely ro result in aggressi
compedtion for space than between congenerics (rwo species from t
same genus), which in tiun were less likeiy than those in the sar:'re famj
but from clifferent genera. Furthermore, the more distantly related the rv
cornperitors the more uneven the outcome of competition. Trends in tl
ourcomes of cornpetition, abundance, and richness could be considered
pa$erns of success. This means that ir is possible to measure the success
different taxa.
Lctilrde (degreed

marine comqe1ili11 wlrh latitude' Data are how

1.4.3 Success
1.15 Changes in rhe structure of
'' "'-
Fig.
' ate irl the norrhern (Q) and
n, 'o".fring orclers' of bryozoan comlnunities To measure success, we firsl need a standardized definition of rhis terr
(a), scor"i using Tanaka & Nrnda.kurnar's (199a) r;rnsitiviry
southern hemispheres
with addirional d:rta' In evolutionary terms) insects are considered a succesdful taxon becau
;;;il. Plo, r*drnuuu from Barnes {2002)
 ;;' . .-' .[ri tnu. raJieted to a high.level of,.*,.i::
',,.-.;:li. ' ]l:-1t::;.*l"t:Xl]
b* consideied (genus, phvlum), as could
.-: '-,.. .' ffiil", ;;;ic levets.o,rfu

animals on earth' and are therefore

to described as the most numerous
Iered
seen already' large-scale
I.oNematodesareconsideredto ::-:,;*r;1."..,r...ssfu1'.Aswehaveseenalready, large-sc: trends
.ooria.r.d.to fra '-".t*J"f" As we have
, be ,successful.o*""*,i"Jitr'"'
-'- and the abundance of organisms' but these
-n, imong the most.r-uroui exist both in uit"J'-'itr'ness
do not necessarilv coincide
(Fie. 1.13), There are, however, many more
, animalson earth, show strong trends in space and
potential *t";;:t ;; tuttt"i"l'ittt
L ' i*tty-*tic gradient' elhiloderms-,tf::::11.1t:j:
' time: Along
'
particular) o..t" n *''il' *ia"t,aepth range than other taxa
and
"t'o*'
dominate the deep sea (Ctrapter 8)'
lu*"ri."tiy
Crustaceans, *"li;; othei phyla occur in marine' fresh- Fig, 1.17 Complexiry in suture lines of Jurassic ammonites. Four ammonite species v
'nd $o
'omt
an analysis of occupancy of the availpble surure line boundaries coloured to illustrate evolving compta'<ity lrom simple (left)
water, and ,.r'*'*iJ
'f'tems' progressively more complexly folded ones (right), Photograph; David Barnes.
habimrsinthese'y-..*'*i*dmecouldbeconsideredasananalysisof
of advanced
success is the evolution
proxy of ;;;;;;;. A frequently ,r..a pro*y of
o A frequentty used
of i;;;", o, .o*pt*Ji'y' Organism evolution has produced progressively
sucees, is the evolution example, smaller rhan brachiopods, which are all smaller than mari
;;;;;;;;*"sorcomp,exi,vlh{::lm-BJJf reptiles. However, coflrra${ing representatives of some taxa span mc
":'T#};ffi.:il,':,-:*ffi.1"-"f"i
JJ*ii"*ri"r in the'dtutiop**"t of complexiry
but' most
is debated' than four orders of magniude in size (e"g. for molluscs, while ma
simple in structure' At this snails are iust a few millimetres in length, giant squid can measure neal
;.gr"t;;- (f,tttti' 'na protoctists).remainmore complex organisms are 30 m). Other animal types, such as sponges and colonial forms, do n
whether
#;;;; should perhaps consider have definite limits to body size, much like many fungi, lichens, ar
moresuccessfuflfn-*a',yexamplesrhisdependsonfhetirnescaleunder mosses, There are strong trends in time and space for organism si:
' an{ abundant groups in.today's
consideration' Sonre of th* *o'" 'peciose In the Precambrian (c.570 million years ago), all known animals we
were relatively unspecialized in
seas (e.g' molluscs and tt'stacians) fairly small. !flhile in the Mesozoic, we find the largest known oce,
seas' Trilobites and brachiopods'
in contra$t' were more spe-
ir*briu'n 'predator' (Lioplewodon). Nevertheless the largest organisms ever
cializedandforconsiderableperiodsofevolutionarytirneweremore have occurred are found in today's seas (fin (Balaenopteraphysatwil ar
;;;i"r. *bond'nt (see Fig' 1'10)' though not since the ?alaeczoic' blue {.8. musculus) whales). Arnazingly, two of the iargest living anima
On smaller tU"' tiiif h'gt) ;** scales'-increased
"nd complexity has made
and speciose but more prore to the megamouth shark lMegachasma pelagiasJ and the giant squ
some animal types more abundant (Arcbiteuthis dux), are yet to be seen alive in their natural habitr
is the.evolution of a
t,'go
extinction on
'i'n* "'lt'' { good example
that the shelled Itrithin taxa, ma$y types of animals (mostly those with calcareo
group of tut*,o'J *oU*t'
(an\-onircs) resembled
years, families skeletons) have tended to get larger with time. It rnay be that this reflet
cephal<lpodNa wtilu,ir,wdny,s seas'
bve, t"ns of millions of
evolution away from small rather than towards large size and is :
compex ixi*:l,li:*im:x1r"ffiiff ::H:"1ffi-1ffi.'H:[|;'1ffi anefact o( increased specialization. The increasingly complex amm
. Ammon,es w,h
their shells became the ammonites with nites also tended m become larger (Fig. 1.13). For nearly all animals, r
iutrru lin", in |I|::.;il;-;;;; the mass exthctions werc moslly
increase in volume will change their surface area to volume ratio. This
which illustrares that seiection pressures for
more speciose and abundant evolu-
u il ffi;;ii;;r;, crucial to functioas such as gas exchange and feeding. There has ber
l[H::H,11fr;,TfiJff"* ff#:;;;;;i;;':;i '*::Ti::]:.:fl,::Y*]]ll]]i'evorution
trend with time' change in body size'
rnuch discussion (and little generally accepted conclusion) on wheth
e$indions' also illustrated u"itht' maior (1) there have been more opportunities for new roles (empry niches) f,
small or large animals; (2) rates of evolution of body size vary wir
1.4.4 Size population size or reproductive mode; and (3) oxygen, temperature (
ecology' physiology' and other famors were responsible for the within-fauna size variability set
Bodv size has a key role in the function' in modern faunas. Ior example, amphipods, isopods, pycnogona, ar
organisms" conrrasrs.in animal size surround
I
us,
I :;"t;;.;;;aiil*r all nematode worms are' for some other taxa in rhe deep sea and polar regions can achieve very lar;
,""", "* simply betwe*"-o'g^ti*m rypes:
 r: P,*.-iTEP(fl6 ii\'i
tiit lnAP'lil El':Vltl0$lilti'iT
ro as porar giganrism). c)n careful anarysis of
r New evidence, such as sateliite iraagery, has revealed that the oceans and sea:
. l,t size (s.merimes referred more compartmentaliged Lhan r^/e once lhought" Broad\, marine life seems spat
Chapelle E< Peck (1999) found
e The link to oxyqell
exPlains rmnhioocl sizcs frotn around ttre world' divided into twelve major biogeograpttic biomes, each with dilferent pattern
uf nor only marine, bur,also freshwarer faunas showed a
Iioun,ir* (e g' in <Jragonflies) ilffi;;r; 'I'hey tested rhis hypctthesis by
product,vity and their awn specrlir biotas.
'd,]n"n ,n. Carbroniierous Period
strong. correlation to o)cygen' larer.
r tt has long been ls6Lrgnis0d thst terrestrial organisms exhibit distinct trend
i
1
]r"i""iu.-otoituric
oxygen levels
t"r"r:t*-it"* amplripod size in a high-altitude lake (l'ake Titicaca)'
endemism, diversity. abundance, size and other characterswith physical environn're
were higher'
;il;".;;d;*ed"tl"ir predictions (they were mainlv very small)- characteristics. lntense work on nrarine organisms during the last couple of deca
I ""'D_;;;;_; in siz.e rend ro occur in rerresuial island
as iilcrcases has revealed some similarities and striking drfferences with such patterns on land.
as well
occur relarive to rnainland ccunter-
[";;;. ;;i, rately does nc chan6letypes (rrrost famously l;rrge rnamrnals
o with more extreme and longer isolation of geographic reqions, the tendency tow;

ir.rt O, iiar,ci', ce't'in animal

become dwarfs while others (such as
endemism is greater. Continental shelf rnargins around irlands, such as Hawaii o
big as Antarctica, (an (ont6in the rnajority of species fourd nowhere else. How m
suchas carnivores or mammoths)
size' For the sea' we krrow of far fewer size depends very much on the scale at which we look ior such species.
,-a"*rf tend to increase in lim.itation gradients. There is
il;;; Ju" to irot^tion or along resource . Biodiversity is much discussed but often poorly understood. lt can be measured u:
a variety o{ methods at various taxononric levels (though usually species). There is r
evidencerr{rlecreasesinsizeforsomeradiolaria(Fig.1.3f},butcon.
seem to be smaller good evidence that richness (the number of taxa) has changed through trme
;.;;i;,r;;";ses itt c'thers' Many estuarine anirnals 4), a'y.conclu-
i

;;;;h-;. fully marine equivalent-species iChapter but

differs with tatitude, longitude and other factors.

;;';;;*-;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.