AbsLracL We dlscuss LheoreLlcal aspecLs of Lhe broader foodweb research

agenda parLlcularly Lhe background and varlous approaches used for

modellng foodweb dynamlcs ln absLracL sysLems wlLh more Lhan Lwo Laxa
Much of Lhls Lype of modellng has orlenLed lLself around Lhe classlc and
endurlng complexlLysLablllLy debaLe ln ecology especlally Lhose aspecLs
whlch relaLe Lo Lhe LheoreLlcal and assoclaLed emplrlcal foodweb research
lnLo Lhe relaLlonshlps beLween ecosysLem complexlLy ofLen characLerlzed as
number of llnks and/or number of specles ln a communlLy and varlous
aspecLs of ecosysLem sLablllLy SLablllLy" ln ecology ls a caLchall Lerm LhaL
has been varlously deflned Lo reflecL aspecLs of populaLlon and/or sysLem
equlllbrlum perslsLence reslllence reslsLance and robusLness ln some
cases sLablllLy refers Lo Lhe ouLcome of lnLernal dynamlcs whlle ln oLher
cases lL reflecLs Lhe response of a populaLlon or sysLem Lo a
perLurbaLlon 1he very noLlon of whaL ls a plauslble deflnlLlon of ecosysLem
sLablllLy has drlven some of Lhe fundamenLal shlfLs ln modellng
meLhodology LhaL wlll be descrlbed

lnLroducLlon
undersLandlng Lhe sLrucLure and dynamlcs of ecologlcal neLworks ls crlLlcal
for undersLandlng Lhe perslsLence and sLablllLy of ecosysLems ueLermlnlng
Lhe lnLerplay among neLwork sLrucLure neLwork dynamlcs and varlous
aspecLs of sLablllLy such as perslsLence robusLness and reslllence ln
complex realworld" neLworks ls one of Lhe greaLesL currenL challenges ln
Lhe naLural and soclal sclences and lL represenLs an exclLlng and
dramaLlcally expandlng area of crossdlsclpllnary lnqulry (SLrogaLz 2001)
WlLhln ecology foodweb research represenLs a long LradlLlon of boLh
emplrlcal and LheoreLlcal neLwork analysls (eg LlLon 1927 Llndeman 1942
MacArLhur 1933 alne 1966 May 1973) All aspecLs of ecologlcal neLwork
research have lncreaslng relevance (McCann 2000) for a world faclng
blodlverslLy and hablLaL loss lnvaslve specles cllmaLe change and oLher
anLhropogenlc facLors LhaL are resulLlng ln Lhe drasLlc reorganlzaLlon of
many ecosysLems and ln some cases may lead Lo Lhe collapse of ecosysLems
and Lhe vlLal underappreclaLed servlces Lhey provlde human socleLy (ually
1997) MosL LheoreLlcal sLudles of Lrophlc dynamlcs have focused narrowly
on predaLorprey or paraslLehosL lnLeracLlons and have Lhus lgnored
neLwork sLrucLure ln naLural ecosysLems such lnLeracLlon dyads are
embedded ln dlverse complex food webs where many addlLlonal Laxa and
Lhelr dlrecL and lndlrecL effecLs can play lmporLanL roles for boLh Lhe
sLablllLy of focal specles as well as Lhe sLablllLy of Lhe broader communlLy
Movlng beyond Lhe one or Lwospecles populaLlon dynamlcs modellng
paradlgm oLher research
has expanded Lhe focus Lo lnclude 3 Lo 8 specles explorlng dynamlcs ln
sllghLly more complex sysLems Powever Lhese lnLeracLlon modules sLlll
presenL a drasLlc slmpllflcaLlon of Lhe dlverslLy and sLrucLure of naLural
ecosysLems CLher approaches have focused on hlgher dlverslLy ecologlcal
neLworks buL have elLher lefL ouL dynamlcs alLogeLher (eg neLwork
Lopology and carbonbudgeL sLudles) or lgnored neLwork sLrucLure ln order
Lo conducL analyLlcally LracLable dynamlcal analyses 1he naLure of
modellng requlres [udlclous slmpllflcaLlons (?odzls and lnnes 1992) buL such
cholces can leave emplrlcal ecologlsLs susplclous and LheoreLlcal ecologlsLs
perplexed boLh wonderlng how Lo brldge Lhe gap beLween slmple
maLhemaLlcal models and complex naLural sysLems AL worsL LheoreLlclans
and emplrlclsLs end up lgnorlng or derldlng each oLher's work dlsmlsslng lL
as lrrelevanL for belng Loo absLracL or Loo parLlcular never Lhe Lwaln shall
meeL
ln Lhls paper we focus on LheoreLlcal aspecLs of Lhe broader foodweb
research agenda parLlcularly Lhe background and varlous approaches used
for modellng foodweb dynamlcs ln absLracL sysLems wlLh more Lhan Lwo
Laxa Much of Lhls Lype of modellng has orlenLed lLself around Lhe classlc
(May 1972) and endurlng (McCann 2000) complexlLysLablllLy debaLe
especlally Lhose aspecLs whlch relaLe Lo Lhe LheoreLlcal and
assoclaLed emplrlcal foodweb research lnLo Lhe relaLlonshlps beLween
ecosysLem complexlLy ofLen characLerlzed as number of llnks and/or
number of specles ln a communlLy and varlous aspecLs of ecosysLem
sLablllLy SLablllLy" ls a caLchall Lerm LhaL has been varlously deflned Lo
reflecL aspecLs of populaLlon and/or sysLem equlllbrlum perslsLence
reslllence reslsLance and robusLness (see McCann 2000 for some broad
buL noL exhausLlve sLablllLy deflnlLlons) ln some cases sLablllLy refers Lo Lhe
ouLcome of lnLernal dynamlcs whlle ln oLher cases lL reflecLs Lhe response of
a populaLlon or sysLem Lo a perLurbaLlon 1he very noLlon of whaL ls a
plauslble deflnlLlon of ecosysLem sLablllLy has drlven some of Lhe
fundamenLal shlfLs ln modellng meLhodology LhaL wlll be descrlbed ln
followlng secLlons We do noL address Lhe more emplrlcally and
experlmenLally based dlverslLysLablllLy" debaLe of recenL years (eg
naeem eL al 1994 PecLor eL al 1999 1llman eL al 2001) whlch mosLly
focuses on Lhe relaLlonshlp beLween planL specles rlchness and prlmary
producLlvlLy 1hls chapLer dlscusses how Lhree approaches Lo dynamlcal
modellng of ecologlcal neLworks Lry Lo sLrlke a balance beLween slmpllfylng
and embodylng aspecLs of Lhe
complexlLy of naLural sysLems Lo galn a beLLer undersLandlng of aspecLs of
ecosysLem sLablllLy ln parLlcular Lhe degree Lo whlch Lhe dlfferenL
approaches lncorporaLe dlverslLy
neLwork sLrucLure nonllnear dynamlcs and emplrlcally measurable
parameLers wlll be recurrlng Lhemes We do noL address spaLlal
heLerogenelLy meLapopulaLlon dynamlcs
ageclass sLrucLure envlronmenLal varlablllLy or sLolchlomeLry lncluslon of
Lhose lmporLanL facLors ln dynamlcal foodweb models may well alLer many
of our noLlons abouL Lhe lnLerplay beLween ecosysLem complexlLy and
sLablllLy Whlle we do noL
expllclLly revlew sLudles focused on foodweb sLrucLure or Lopology (eg
Cohen eL al 1990 Wllllams and MarLlnez 2000 uunne eL al 2002a
Carlaschelll eL al 2003) we do
refer Lo Lhem Lo Lhe degree LhaL Lhey have lnfluenced dynamlcal
approaches A fundamenLal message of Lhls chapLer ls LhaL Lhe lnslghLs from
sLudles on Lhe neLwork sLrucLure of complex food webs need Lo be merged
wlLh dynamlcal approaches lf we are Lo make serlous headway regardlng
lssues of Lhe sLablllLy of complex neLworks and Lhe poLenLlal for uslng
dynamlcal models ln a conservaLlon and managemenL conLexL
We focus on one end of a specLrum of modellng (Polllng 1966) Lhe use of
absLracL models Lo elucldaLe general quallLaLlve lnslghLs abouL ecosysLem
sLrucLure dynamlcs and sLablllLy We do noL dlscuss more expllclLly applled
approaches for slmulaLlng parLlcular ecosysLems a sLraLegy whlch has been
used for many aquaLlc sysLems (eg massbalance carbonbudgeL models
such as LcoaLh/LcoSlm ChrlsLensen and WalLers 2004) Such approaches
are presenLed elsewhere ln Lhls book (ChrlsLlan eL al Morrls eL al) 1hese
researchers have ofLen referred Lo Lhelr work as neLwork analysls"
Powever ln our vlew neLwork analysls" ls a very broad Lerm LhaL
encompasses all Lypes of research LhaL LreaL a sysLem as a neLwork of nodes
and edges" or llnks regardless of how Lhose nodes and llnks are deflned or
Lhelr relaLlonshlps are analyzed (SLrogaLz 2001) ln ecology Lhere has ofLen
been a Lenslon beLween research focused on absLracL models versus LhaL
focused on applled slmulaLlon models of parLlcular ecosysLems We shall
see LhaL as absLracL models seek Lo represenL greaLer ecologlcal
complexlLy Lhey Lake on some of Lhe characLerlsLlcs of applled slmulaLlon
modellng hopefully narrowlng Lhe gap beLween LheoreLlcal and emplrlcal
work by carefully lnLegraLlng characLerlsLlcs and goals of Lhe Lwo modellng
exLremes ln Lhls splrlL we wlll end by revlewlng a recenL appllcaLlon of a
mulLlspecles nonllnear dynamlcal foodweb model Lo Lhe lssue of how
culllng a Lop predaLor ls llkely Lo affecL Lhe hake flshery yleld ln Lhe 8enguela
ecosysLem (?odzls 1998 2000 2001)

8ackground
ln Lhe flrsL half of Lhe 20Lh cenLury many ecologlsLs belleved LhaL naLural
communlLles develop lnLo sLable sysLems Lhrough successlonal dynamlcs
AspecLs of Lhls bellef developed lnLo Lhe noLlon LhaL complex communlLles
are more sLable Lhan slmple ones Cdum (1933) MacArLhur (1933) LlLon
(1938) and oLhers clLed an array of emplrlcal evldence supporLlng Lhls
hypoLhesls Some popular examples lncluded Lhe vulnerablllLy of
agrlculLural monoculLures Lo calamlLles ln conLrasL Lo Lhe apparenL sLablllLy
of dlverse Lroplcal ralnforesLs and Lhe hlgher frequency of lnvaslons ln
slmple lsland communlLles compared Lo more complex malnland
communlLles lL was LhoughL LhaL a communlLy comprlsed of specles wlLh
mulLlple consumers would have fewer lnvaslons and pesL ouLbreaks Lhan
communlLles of specles wlLh fewer consumers 1hls
was sLaLed ln a general LheoreLlcal way by MacArLhur (1933) who
hypoLheslzed LhaL a large number of paLhs Lhrough each specles ls
necessary Lo reduce Lhe effecLs of overpopulaLlon of one specles"
MacArLhur concluded LhaL sLablllLy lncreases as Lhe number of llnks
lncreases" and LhaL sLablllLy ls easler Lo achleve ln more dlverse assemblages
of specles Lhus llnklng communlLy sLablllLy wlLh boLh lncreased Lrophlc llnks
and lncreased numbers of specles CLher Lypes of LheoreLlcal conslderaLlons
emerged Lo supporL Lhe poslLlve complexlLysLablllLy relaLlonshlp lor
example LlLon (1938) argued LhaL slmple predaLorprey models reveal Lhelr
lack of sLablllLy ln Lhe
osclllaLory behavlor Lhey exhlblL alLhough he falled Lo compare Lhem Lo
mulLlspecles models (May 1973) 1he noLlon LhaL complexlLy begeLs
sLablllLy" whlch already had greaL lnLulLlve appeal as well as Lhe welghL of
hlsLory behlnd lL was Lhus accorded a gloss of LheoreLlcal rlgor ln Lhe mld
1930's 8y Lhe laLe 30's lL Look on Lhe paLlna of convenLlonal wlsdom and aL
Llmes was elevaLed Lo Lhe sLaLus of ecologlcal Lheorem or
formal proof" (PuLchlnson 1939) Whlle some subsequenL emplrlcal
lnvesLlgaLlons ralsed quesLlons abouL Lhe concluslveness of Lhe relaLlonshlp
(eg PalrsLon eL al 1968) lL wasn'L unLll Lhe early 70's LhaL Lhe noLlon LhaL
complexlLy lmplles sLablllLy as a LheoreLlcal generallLy was expllclLly and
rlgorously challenged by Lhe analyLlcal work of 8oberL May (1972 1973) a
physlclsL by Lralnlng and ecologlsL by lncllnaLlon May's 1972 paper and
hls1973 book SLablllLy and ComplexlLy ln Model LcosysLems" (reprlnLed ln a
2nd
edlLlon ln 1974 wlLh an added reface AfLerLhoughLs and
8lbllography mosL recenLly reprlnLed ln an 8Lh edlLlon ln 2001 as a
rlnceLon Landmarks ln 8lology" volume) were enormously lmporLanL on
several fronLs MeLhodologlcally
May lnLroduced many ecologlsLs Lo a muchexpanded reperLolre of
maLhemaLlcal Lools beyond LoLkavolLerra predaLorprey models and also
seL Lhe sLage for ma[or ecologlcal conLrlbuLlons Lo Lhe developmenL of
Lheorles of deLermlnlsLlc chaos (eg May 1974 1976) AnalyLlcally May
used local sLablllLy analyses of randomly assembled
communlLy maLrlces Lo maLhemaLlcally demonsLraLe LhaL neLwork sLablllLy
decreases wlLh complexlLy ln parLlcular he found LhaL more dlverse
sysLems compared Lo less dlverse
sysLems wlll Lend Lo sharply LranslLlon from sLable Lo unsLable behavlor as
Lhe number of specles Lhe connecLance (a measure of llnk rlchnessLhe
probablllLy LhaL any Lwo
specles wlll lnLeracL wlLh each oLher) or Lhe average lnLeracLlon sLrengLh
lncrease beyond a crlLlcal value We descrlbe hls analyses ln more deLall ln
Lhe nexL secLlon May (1973) also polnLed ouL LhaL emplrlcal evldence ls noL
concluslve wlLh regard Lo a parLlcular complexlLysLablllLy relaLlonshlp (eg
Lhere are a number of hlghly sLable producLlve and slmple naLural
ecosysLems such as easL coasL SparLlna alLernlflora marshes complex
conLlnenLal communlLles have suffered Lhe ravages of pesLs such as Lhe
Cypsy MoLh eLc) May's analyLlcal resulLs and hls concluslon LhaL ln
general maLhemaLlcal models of mulLlspecles communlLles complexlLy
Lends Lo begeL lnsLablllLy" (p 74 2001 edlLlon) Lurned earller ecologlcal
lnLulLlon" on lLs head and lnsLlgaLed a dramaLlc shlfL and refocuslng of
LheoreLlcal ecology Pls resulLs lefL many emplrlcal ecologlsLs wonderlng
how Lhe asLonlshlng dlverslLy and complexlLy Lhey observed ln naLural
communlLles could perslsL even Lhough May hlmself lnslsLed Lhere was no
paradox As May puL lL ln shorL
Lhere ls no comforLable Lheorem assurlng LhaL lncreaslng dlverslLy and
complexlLy begeL enhanced communlLy sLablllLy raLher as a maLhemaLlcal
generallLy Lhe opposlLe ls Lrue
1he Lask Lherefore ls Lo elucldaLe Lhe devlous sLraLegles whlch make for
sLablllLy ln endurlng naLural sysLems" (p 174 2001 edlLlon) MosL
subsequenL work relaLed Lo food webs was devoLed Lo flndlng neLwork
sLrucLures specles' sLraLegles and dynamlcal characLerlsLlcs LhaL would
allow complex communlLles Lo perslsL ln some cases Lhe research was
underLaken Lo corroboraLe May's flndlngs and oLher research soughL Lo
reanlmaLe whaL many now felL Lo be Lhe ghosL of poslLlve complexlLy
sLablllLy pasL WhaLever Lhe moLlvaLlon lmporLanL pleces of Lhls ongolng
research agenda lnclude
examlnlng Lhe role of omnlvory Lhe number of Lrophlc levels weak
lnLeracLlons beLween specles adapLlve foraglng of consumers and complex
neLwork sLrucLure on varlous
measures of ecosysLem sLablllLy Powever subsLanLlally dlfferlng
meLhodologles sLablllLy deflnlLlons and a varleLy of someLlmes apparenLly
confllcLlng resulLs have somewhaL muddled Lhe orlglnal quesLlon of wheLher
complexlLy begeLs sLablllLy lnsLead Lhe quesLlon ls becomlng more usefully
recasL along Lhe llnes of May's noLlon of devlous sLraLegles" whaL are Lhe
general characLerlsLlcs of complex ecologlcal neLworks LhaL allow for
promoLe and resulL from Lhe sLablllLy and perslsLence of naLural
ecosysLems Local SLablllLy Analyses of CommunlLy MaLrlces ln hls semlnal
sLudles on food web sLablllLy May (1972 1973) measured local or
nelghborhood sLablllLy ln Lhls analysls lL ls assumed LhaL Lhe communlLy
resLs aL an equlllbrlum polnL where all populaLlons have consLanL values
(llg 1a) 1he sLablllLy of Lhls equlllbrlum ls LesLed wlLh small perLurbaLlons
lf all specles reLurn Lo Lhe equlllbrlum monoLonlcally or by damped
osclllaLlons lL ls sLable (llg 1a) ln conLrasL lf Lhe
populaLlon denslLles evolve away from Lhe equlllbrlum denslLles
monoLonlcally or osclllaLory Lhey are unsLable (llg 1b) neuLral sLablllLy
represenLs a Lhlrd posslblllLy ln
whlch Lhe perLurbaLlon nelLher grows nor decays as Lhe populaLlon denslLles
elLher reach an alLernaLlve equlllbrlum or osclllaLe wlLh a consLanL
ampllLude ln llmlL cycles (llg 1b)
ln a communlLy of n specles Lhls approach ls based on an n x n !acoblan
communlLy maLrlx of specles lnLeracLlon coefflclenLs LhaL descrlbe Lhe
lmpacL of each specles l on Lhe growLh of each specles [ aL equlllbrlum
populaLlon denslLles ln food webs Lhese coefflclenLs may be poslLlve (le l
ls eaLen by [) zero (no lnLeracLlon) or negaLlve (le l eaLs [) 1o descrlbe
lnLeracLlons by slngle coefflclenLs lL ls necessary Lo
assume llnear lnLeracLlons 1he n elgenvalues of Lhe communlLy maLrlx
characLerlze lLs Lemporal behavlor (see May 1973 for a deLalled descrlpLlon
how Lhese elgenvalues are
calculaLed) lor every populaLlon poslLlve real parLs of Lhe elgenvalues
lndlcaLe perLurbaLlon growLh negaLlve real parLs lndlcaLe perLurbaLlon
decay and zero real parLs lndlcaLe neuLral sLablllLy Accordlngly lf any of
Lhe elgenvalues has a poslLlve real parL Lhe sysLem wlll be unsLable le aL
leasL one of Lhe specles does noL reLurn Lo Lhe equlllbrlum SLablllLy may be
measured elLher as a blnary varlable reLurn Lo equlllbrlum
or noL or as a meLrlc varlable ln form of Lhe reLurn Llme needed by Lhe
populaLlon denslLles Lo seLLle back Lo Lhe equlllbrlum May (1972 1973)
used communlLy maLrlces ln whlch specles were randomly llnked wlLh
random lnLeracLlon sLrengLh Lo show LhaL Lhe local sLablllLy decreases wlLh
complexlLy (measured as connecLance) dlverslLy and average lnLeracLlon
sLrengLh among Lhe specles 1he use of such random communlLy maLrlces
has aLLracLed much crlLlclsm lL was shown Lo be exLremely unllkely LhaL any
of Lhese random communlLles even remoLely resembles ecosysLems wlLh a
mlnlmum of ecologlcally reallsm such as
conLalnlng aL leasL one prlmary producer a llmlLed number of Lrophlc levels
and no consumers eaLlng resources LhaL are Lwo or more Lrophlc levels
hlgher (Lawlor 1978) 1he nonrandomness of ecosysLem sLrucLure has been
demonsLraLed ln deLall by more recenL foodweb Lopology sLudles (eg
Wllllams and MarLlnez 2000 uunne eL al 2002a
2004 Morrls eL al Lhls volume) Accordlngly subsequenL work added more
sLrucLural reallsm Lo Lhose random communlLy maLrlces by lncludlng
emplrlcal paLLerns of food web sLrucLure and lnLeracLlon sLrengLh
dlsLrlbuLlons varylng connecLance ln 10 specles food webs ue Angells
(1973) LesLed May's
(1972) resulLs wlLhln a more reallsLlc model envlronmenL and found Lhree
condlLlons for poslLlve complexlLysLablllLy relaLlonshlps (1) Lhe blomass
asslmllaLed by consumers ls a
small fracLlon (30) of Lhe amounL of blomass removed from Lhe resource
populaLlon (2) Lhe consumers are sub[ecL Lo sLrong selfdampenlng (up Lo
20 Llmes Lhe maxlmum growLh raLe of prlmary producers) whlch resLrlcLs
Lhelr populaLlon growLh and (3) a blas Lowards boLLomup conLrol of Lhe
lnLeracLlons (le resource blomass has a sLronger lnfluence on Lhe
lnLeracLlon sLrengLh values Lhan consumer blomass) whlch also means LhaL
consumers have very llLLle lnfluence on Lhe dynamlcs of Lhelr resources 1he
reallsm of Lhese condlLlons however ls quesLlonable as asslmllaLlon
efflclencles for carnlvores should generally be hlgher (83 ?odzls lnnes
1992) and such exLreme self dampenlng or low lnfluence of consumers on
resources appear unllkely neverLheless for a long Llme Lhe resulLs of ue
Angells (1973) remalned Lhe only maLhemaLlcal models showlng poslLlve
complexlLysLablllLy relaLlonshlps ln a more sLrucLural approach lmm and
LawLon (1977 1978) used communlLy
maLrlces of four lnLeracLlng specles 1hey sysLemaLlcally varled food chaln
lengLh and lnLroduced omnlvory by leLLlng one of Lhe specles consume
resources of mulLlple Lrophlc
levels 1hey found LhaL Lhe sLablllLy of Lhe communlLy maLrlces decreases
wlLh Lhe number of Lrophlc levels 1he resulLs for omnlvory however
varled as omnlvory decreases sLablllLy ln Lerms of Lhe fracLlon of sLable
models LhaL reLurn Lo equlllbrlum buL aL Lhe same Llme lncreases sLablllLy of
Lhese sLable models by decreaslng Lhelr reLurn Llme neverLheless Lhey
concluded LhaL Lhe number of Lrophlc levels ln naLural communlLles ls
llmlLed by Lhe sLablllLy of populaLlon dynamlcs (lmm LawLon 1977) and
Lhey predlcLed LhaL omnlvory should be a rare phenomenon ln naLural food
webs (lmm LawLon 1978) ln parLlcular Lhe laLLer predlcLlon was noL
found Lo be conslsLenL wlLh emplrlcal daLa as more deLalled food webs were
complled (eg MarLlnez 1991 olls 1991) ln a furLher approach Lo analyze
Lhe relaLlonshlp beLween foodweb sLrucLure and dynamlcs ?odzls (1981)
showed LhaL emplrlcal communlLy maLrlces correspondlng Lo 40 real food
webs are more sLable Lhan randomly assembled communlLy maLrlces 1hls
clearly lndlcaLed LhaL naLural food webs are sLrucLured ln a way LhaL
sLablllzes Lhe lnLernal populaLlon dynamlcs buL Lhe quesLlon remalned open
as Lo whlch parLlcular sLrucLural
properLles were lmporLanL lnLeresLlngly Moore and PunL (1988) showed
LhaL ln Lhose 40 food webs specles are blocked lnLo smaller subwebs of hlgh
connecLance wlLh only a few llnks connecLlng Lhe subwebs 8ased on
analyses of May (1972) Lhey argued LhaL such comparLmenLallzaLlon may
lncrease neLwork sLablllLy and Lhus be responslble for Lhe
comparaLlvely hlgh sLablllLy of emplrlcal food webs
Lvldence for clusLerlng or comparLmenLallzaLlon ln more deLalled naLural
food webs has been suggesLlve buL scanL krause eL al (2003) uslng food
web neLwork daLa
wlLh unwelghLed llnks as well as wlLh llnks welghLed by lnLeracLlon
frequency carbon flow or lnLeracLlon sLrengLh ldenLlfled slgnlflcanL
comparLmenLs ln 3 of 3 complex food
webs Powever ouL of 17 verslons of Lhe 3 food webs examlned only 6
verslons revealed slgnlflcanL comparLmenLallzaLlon Clrvan eL al (2002)
found ecologlcally
meanlngful comparLmenLs based on unwelghLed Lopology for a Chesapeake
8ay food web (8alrd ulanowlcz 1989) buL had dlfflculLy ldenLlfylng
comparLmenLs ln oLher more hlghly resolved food webs (Clrvan pers com)
1opologlcal analysls of 16 dlverse wellresolved food webs showed LhaL Lhelr
clusLerlng coefflclenL (WaLLs SLrogaLz 1998) ls
low ln comparlson Lo oLher Lypes of neLworks (uunne eL al 2002a) 1he low
clusLerlng coefflclenLs of food webs ls ln parL be aLLrlbuLable Lo Lhelr hlgh
connecLance (llnks/specles 2) compared Lo oLher realworld neLworks
(uunne eL al 2002a) ln effecL Lhere ls such a hlgh concenLraLlon of llnks ln
mosL food webs LhaL Lhe posslblllLy of comparLmenLallzaLlon aL leasL from a
purely Lopologlcal perspecLlve ls drowned ouL ln
addlLlon Lo belng based on unwelghLed llnks whlch may obscure
comparLmenLallzaLlon Lhe clusLerlng coefflclenL mlsses comparLmenLs of
nonomnlvorous specles based on a
slngle basal specles ln sum Lhe lssue of wheLher and how food webs are
comparLmenLallzed and Lhe lmpacLs of comparLmenLallzaLlon on sLablllLy
remalns an open quesLlon LhaL requlres more research LxLendlng Lhe
approach of addlng emplrlcal reallsm Lo local sLablllLy analyses de
8ulLer eL al (1993) parameLerlzed Lhe general communlLy maLrlx model wlLh
emplrlcal foodweb sLrucLures and lnLeracLlon sLrengLhs among Lhe specles
ln Lhelr emplrlcal daLa Lhey found a paLLern of sLrong Lopdown effecLs of
consumers on Lhelr resources aL lower Lrophlc levels ln food webs and
sLrong boLLomup effecLs of resources on Lhelr consumers aL hlgher Lrophlc
levels Addlng emplrlcal lnLeracLlon sLrengLh paLLerns Lo Lhe communlLy
maLrlces lncreased Lhelr local sLablllLy ln comparlson Lo maLrlces wlLh
random lnLeracLlon sLrengLh values MosL lmporLanLly Lhese resulLs
demonsLraLed LhaL naLural foodweb sLrucLures as well as Lhe dlsLrlbuLlon of
lnLeracLlon sLrengLhs wlLhln Lhose sLrucLures conLrlbuLe Lo an lncreased
local sLablllLy of Lhe correspondlng communlLy maLrlces neuLel eL al (2002)
explalned Lhls flndlng wlLh resulLs LhaL showed LhaL weak lnLeracLlons are
concenLraLed ln long loops ln Lhelr analysls a loop ls a paLhway of
lnLeracLlons from a cerLaln specles Lhrough Lhe web back Lo Lhe same
specles wlLhouL vlslLlng oLher specles more Lhan once 1hey measured loop
welghL as Lhe geomeLrlc mean of Lhe lnLeracLlon sLrengLhs ln Lhe loop and
showed LhaL loop welghL decreases wlLh loop lengLh Agaln
when applled Lo Lhe communlLy maLrlx Lhls emplrlcally documenLed paLLern
of lnLeracLlon sLrengLh dlsLrlbuLlons lncreased lLs local sLablllLy ln
comparlson Lo random
lnLeracLlon sLrengLh dlsLrlbuLlons 1hus over Lhe lasL 30 years sLudles of
Lhe local sLablllLy of communlLy maLrlces have conLrlbuLed Lo our
undersLandlng of Lhe relaLlonshlp beLween complexlLy and sLablllLy ue
Angells (1973) lndlcaLed some speclflc condlLlons for poslLlve
complexlLysLablllLy relaLlonshlps whereas none of Lhe subsequenL sLudles
has lnverLed Lhe negaLlve complexlLysLablllLy relaLlonshlp demonsLraLed by
May (1972) neverLheless oLher sLudles conslsLenLly suggesLed LhaL local
sLablllLy lncreases when ecologlcal reallsm ls added Lo Lhe communlLy
maLrlx ln Lhls sense sLablllLy ls galned by a llmlLed number of Lrophlc levels
(lmm LawLon 1977) naLural foodweb sLrucLures (?odzls 1981
ulanowlcz 2002) and lnLeracLlon sLrengLh dlsLrlbuLlons (de 8ulLer eL al 1993
neuLel eL al 2002) Powever Lo galn maLhemaLlcal LracLablllLy Lhe
communlLy maLrlx approach wlLh
or wlLhouL Lhe addlLlon of more plauslble ecologlcal sLrucLure makes
several assumpLlons LhaL may resLrlcL Lhe generallLy of lLs resulLs llrsL
communlLy equlllbrlum ls assumed whlch excludes chaoLlc populaLlon
dynamlcs LhaL may characLerlze many naLural sysLems (llg 1c PasLlngs eL
al 1993) Second only small perLurbaLlons are LesLed Larger perLurbaLlons
mlghL yleld alLernaLlve equlllbrla Cnly addresslng local or nelghborhood
sLablllLy does noL necessarlly predlcL global sLablllLy alLhough Lhey can
colnclde 1hlrd llnear lnLeracLlon coefflclenLs are only a rough
approxlmaLlon of specles' dynamlcs LhaL wlll only hold for arblLrarlly small
varlances ln populaLlon denslLles lor any larger populaLlon varlances non
llnear specles' lnLeracLlons wlll be poorly approxlmaLed by
llnear coefflclenLs 1hls assumpLlon of small populaLlon varlances however
ls llkely Lo hold once Lhe analyses are resLrlcLed Lo sllghLly perLurbed
populaLlons aL equlllbrlum
Powever LhaL case ls a raLher small sllce of a much broader ecologlcal ple
nonLlnear nonLqulllbrlum opulaLlon SLablllLy ln loodWeb Modules 1o
overcome Lhe llmlLaLlons of Lhe equlllbrlum assumpLlon and llnear
lnLeracLlons lncluded ln communlLy maLrlx analyses an alLernaLlve approach
Lo analyzlng foodweb sLablllLy emerged ln Lhe early 1990s (eg PasLlngs
owell 1991 ?odzls lnnes 1992 McCann ?odzls 1994ab) Clvlng up
Lhose assumpLlons prohlblLed Lhe use of communlLy maLrlces lnsLead
Lhese sLudles deflned less analyLlcally LracLable buL more ecologlcally
plauslble models of nonllnear specles' lnLeracLlons Lo sLudy Lhe populaLlon
dynamlcs ln speclespoor foodweb modules such as Lhreespecles food
chalns ln effecL one aspecL of ecologlcal plauslblllLy was Lraded for
anoLher nonllnear nonequlllbrlum dynamlcs were embraced buL dlverslLy
was losL MosL of Lhese nonllnear dynamlcal sLudles have used an
allomeLrlc bloenergeLlc consumerresource model of specles
blomass evoluLlon LhaL developed by ?odzls lnnes (1992) lL deflnes for
every specles how much blomass ls losL due Lo meLabollsm or belng
consumed and how much blomass
ls galned due Lo prlmary producLlon or feedlng on oLher specles ln Lhls
approach prlmary producer growLh ls based on loglsLlc growLh equaLlons
and consumer growLh ls based on nonllnear funcLlonal responses of
consumpLlon Lo resource denslLles (Skalskl Cllllam 2001) AnoLher aspecL
of ecologlcal plauslblllLy ls added Lhrough Lhe basls of mosL of lLs parameLers
on specles bodyslze and meLabollc caLegorles (le endoLherm ecLoLherm
verLebraLe lnverLebraLe prlmary producer) Speclfylng Lhls bloenergeLlc
model for every specles ln a food web ylelds a seL of ordlnary dlfferenLlal
equaLlons (CuLs) 1he numerlcal lnLegraLlon of Lhe CuLs resulLs ln Llme
serles of specles' blomass evoluLlon LhaL may be used Lo characLerlze
populaLlon dynamlcs (eg sLable
equlllbrla llmlL cycles chaoLlc cycles llg 1) lurLhermore lf specles
exLlncLlon ls deflned as falllng below a Lhreshold of blomass denslLy (eg 10
30) Lhe perslsLence of Lhe populaLlons ln Lhe food web can be sLudled 1hls
approach led Lo Lhe ldenLlflcaLlon of perslsLenL chaos ln Lhree specles food
chalns where Lhe populaLlon mlnlma are bounded
well away from zero (PasLlngs owell 1991 McCann ?odzls 1994a llg
1c) Powever on a longer Llme scale perslsLenL chaos may Lransform lnLo
LranslenL chaos due
Lo sudden caLasLrophlc populaLlon crashes (McCann ?odzls 1994b)
1ogeLher Lhese resulLs demonsLraLe LhaL specles and communlLy
perslsLence ls noL resLrlcLed Lo equlllbrlum dynamlcs (llg 1ab) wlLh Lhe
poLenLlal for chaoLlc populaLlon dynamlcs (llg 1c) LhaL may or may noL be
perslsLenL over shorL and long Llme scales Lxplorlng anoLher aspecL of
ecologlcal complexlLy McCann PasLlngs (1997)
lnvesLlgaLed Lhe lmpacL of omnlvory on dynamlcs by alLerlng Lhe neLwork
sLrucLure so LhaL Lhe Lop specles of Lhe food chaln consumed boLh Lhe
herblvorous and basal specles 1hey found LhaL Lhls Lype of omnlvory
sLablllzes populaLlon dynamlcs by elLher ellmlnaLlng chaoLlc dynamlcs or
boundlng Lhelr mlnlma away from zero ln conLrasL Lo Lhe earller
communlLy maLrlx work of lmm and LawLon (1978) Lhls suggesLs LhaL
omnlvory ls a sLablllzlng feaLure of complex neLworks lL has Lo be
cauLloned however LhaL Lhese resulLs were obLalned under Lhe assumpLlon
LhaL Lhe omnlvore has a hlgher preference for Lhe lnLermedlaLe specles Lhan
for Lhe basal specles whlch rarely appears ln naLural food webs (Wllllams
MarLlnez 2004a) lurLhermore Lhe consequences of omnlvory ln a
comparable slmple experlmenLal sysLem are hlghly dependenL on nuLrlenL
enrlchmenL slnce coexlsLence of boLh consumers ls resLrlcLed Lo
lnLermedlaLe nuLrlenL saLuraLlons (ulehl lelssel 2001) ln an exLended
sLudy of foodweb modules McCann eL al (1998) showed LhaL weak
lnLeracLlons mlghL generally dampen populaLlon osclllaLlons and Lhus
sLablllze
dynamlcs ln ecologlcal neLworks 1hey suggesLed LhaL such weak
lnLeracLlons among specles mlghL be Lhe key for undersLandlng Lhe
exlsLence of complex communlLles lf
mosL lnLeracLlons ln such communlLles are weak osL eL al (2000)
demonsLraLed LhaL a seL of Lwo Lhreespecles food chalns llnked by Lhe same
Lop consumer can be sLablllzed
when Lhe Lop consumer can swlLch lLs preferences beLween Lhe Lwo
lnLermedlaLe specles Such preference swlLchlng appears llkely whenever
consumers mlgraLe beLween dlfferenL
hablLaL comparLmenLs such as Lhe pelaglc and llLLoral food webs of lakes
(osL eL al 2000) 1hls suggesLs a poLenLlally lmporLanL role of spaLlal
paLLerns on foodweb sLablllLy
LhaL remalns Lo be elucldaLed LxLendlng Lhe slze of Lhe food web modules
lussmann and Peber (2002) demonsLraLed LhaL Lhe frequency of chaoLlc
dynamlcs lncreases wlLh Lhe
number of Lrophlc levels buL decreases wlLh oLher sLrucLural properLles LhaL
cause hlgher food web complexlLy lnLeresLlngly Lhese resulLs lndlcaLe LhaL
dependenL on whlch
process domlnaLes ln a parLlcular food web populaLlon sLablllLy mlghL
lncrease or decrease wlLh food web complexlLy
Cverall Lhe numerlcal lnLegraLlon of nonllnear populaLlon dynamlc models
of foodweb modules has added a greaL deal Lo our undersLandlng of
dynamlcal sLablllLy of mulLlspecles sysLems Whlle noL as slmple absLracL
and analyLlcally Lldy as communlLy maLrlx analyses Lhese models appear Lo
capLure more falLhfully some cenLrally lmporLanL aspecLs of naLural
ecologlcal communlLlesaspecLs whlch appear very relevanL Lo lssues
concernlng complexlLy and sLablllLy MosL lmporLanLly Lhls modellng
approach has clarlfled LhaL nonequlllbrlum populaLlon dynamlcs do noL
prevenL specles' perslsLence (PasLlngs owell 1991 McCann ?odzls
1994a) lurLher resulLs suggesL how omnlvory (McCann PasLlngs 1997
ulehl lelssel 2001) weak lnLeracLlons (McCann eL al 1998) and spaLlal
processes (osL eL al 2000) may sLablllze food webs 8elng resLrlcLed Lo
speclespoor food web modules however none of Lhese sLudles can
adequaLely explore or LesL complexlLysLablllLy relaLlonshlps ln dlverse
neLworks LhaL reflecL more of Lhe complexlLy of naLural sysLems Whlle lL
has been suggesLed LhaL resulLs from lnLeracLlon module sLudles can be
generallzed Lo more dlverse complex neLworks LhaL noLlon ls based on Lwo
poLenLlally problemaLlc assumpLlons (1) populaLlon
dynamlcs respond slmllarly Lo lndependenL varlables ln foodweb modules
and complex neLworks and (2) populaLlon sLablllLy colncldes wlLh
communlLy sLablllLy lL has been
shown LhaL specles' perslsLence does noL colnclde wlLh populaLlon equlllbrla
(McCann ?odzls 1994a) and LhaL Lhe frequency of chaoLlc dynamlcs ls
lnfluenced by foodweb
complexlLy (lussmann Peber 2002) l Specles erslsLence ln Complex
lood Webs More recenLly Lhe numerlcal lnLegraLlon approach has been
exLended Lo complex
foodweb models (Wllllams MarLlnez 2004a ms) ln Lhls approach
bloenergeLlc models are applled Lo a dlverse assemblage of specles ln
complex Lrophlc neLworks 1he
sLrucLure of Lhese complex neLworks ls deflned by a seL of slmple Lopologlcal
models random cascade or nlche model food webs (Wllllams MarLlnez
2000) 1he nlche
model has been found Lo successfully predlcL Lhe general sLrucLure of
complex food webs whlle Lhe cascade model (Cohen eL al 1990) predlcLs
some aspecLs and Lhe random model predlcLs almosL no aspecLs of food
web sLrucLure (Wllllams MarLlnez 2000 uunne eL al 2004) uynamlcs
based on Lhe ?odzls and lnnes (1992) approach are run on each of Lhe Lhree
Lypes of foodweb sLrucLures and Lhe ensulng shorL and longLerm
populaLlon dynamlcs are lnvesLlgaLed 1he expllclL lnLegraLlon of complex
neLwork sLrucLure wlLh nonllnear nonequlllbrlum bloenergeLlc modellng
allows for Lhe exploraLlon and LesLlng of complexlLysLablllLy relaLlonshlps ln
more plauslbly complex food webs whlle reLalnlng cerLaln aspecLs of
modellng slmpllclLy 1he Lopology models whlch are analyLlcally solvable
(eg Camacho eL al 2002) use Lwo readlly esLlmaLed parameLers
specles rlchness (8rose eL al 2003a) and connecLance (MarLlnez eL al 1999)
and a few slmple rules for asslgnlng llnks 1he dynamlcs model (Wllllams
and MarLlnez 2004b)
modlfled and generallzed from ?odzls and lnnes (1992) whlle noL
analyLlcally LracLable ls deflned by a narrow seL of parameLers LhaL can be
measured relaLlvely readlly (eg
allomeLrlc daLa) slmple rules governlng blomass galn and loss and
lnLeracLlon dynamlcs LhaL have been sLudled exLenslvely ln emplrlcal
seLLlngs (eg funcLlonal response) As descrlbed ln Lhe prevlous secLlon Lhe
bloenergeLlc dynamlcs model has Lhe addlLlonal beneflL of havlng been
sLudled exLenslvely ln a slmple module seLLlng ln Lhese lnLegraLed sLrucLure
dynamlcs neLworks global sLablllLy can be measured as perslsLence le Lhe
fracLlon of specles wlLh perslsLenL dynamlcs (whlch lncludes equlllbrlum
llmlL cycle and chaoLlc dynamlcs) uslng such foodweb models Wllllams
and MarLlnez (ms) found LhaL perslsLence decreases llnearly for
communlLles of 13 Lo 30 specles and connecLance values beLween 003 and
03 Lhus quallLaLlvely repllcaLlng Lhe
resulLs of May (1972) ln a compleLely dlfferenL model envlronmenL Also
Lhey found
LhaL food webs wlLh random Lopology dlsplay very low perslsLence whlle
addlng some
neLwork sLrucLure ln Lhe form of Lhe cascade model lncreases perslsLence by
an order of
magnlLude use of Lhe nlche model sLrucLure lmproves perslsLence yeL
agaln over Lhe
cascade model (Wllllams MarLlnez ms) Clven LhaL Lhls colncldes wlLh a
decrease ln
emplrlcal adequacy of Lhe sLrucLural models Lhe resulLs sLrongly suggesL
LhaL Lhe complex
neLwork sLrucLure of naLural communlLles provldes a hlgher probablllLy of
specles
perslsLence lnLeresLlngly Lhe sLrucLure of Lhe perslsLenL food webs LhaL
remalned afLer
specles exLlncLlons was more conslsLenL wlLh emplrlcal daLa Lhan Lhe
orlglnal sLrucLural
models even ln Lhe case of Lhe nlche model (Wllllams MarLlnez ms)
1hese resulLs suggesL a LlghL relaLlonshlp beLween foodweb sLrucLure
dynamlcs and sLablllLy slnce
neLwork sLrucLure lnfluences Lhe populaLlon dynamlcs LhaL ln Lurn
deLermlne perslsLenL
foodweb sLrucLures ln addlLlon Lo Lhe lmporLanL role of complex neLwork
sLrucLure for
perslsLence Wllllams and MarLlnez (2004b ms) showed LhaL small
varlaLlons ln Lhe
funcLlonal response of consumpLlon Lo resource and/or predaLor denslLy can
have
dramaLlc effecLs on sLablllzlng Lhe dynamlcs of parLlcular specles as well as
overall
specles perslsLence Cne lmporLanL sLablllzlng varlaLlon ln funcLlonal
response ls Lhe
sllghL relaxaLlon of consumpLlon aL low resource denslLles rlor dynamlcal
models have
Lyplcally assumed 1ype ll funcLlonal responses (Polllng 1939) whlch lack Lhls
Lype of
feedlng relaxaLlon or have looked aL sLablllzlng aspecLs of sLrong verslons of
relaxaLlonLype responses (eg sLrong 1ype lll) ln model sysLems wlLh only
Lwo specles (Murdoch
eL al 1973 Passell 1978 ?odzls lnnes 1992) 1he relaxaLlon of feedlng aL
low
resource denslLy ls evocaLlve of a varleLy of welldocumenLed ecologlcal
mechanlsms
(Skalskl Cllllam 2001) lncludlng predaLor lnLerference prey swlLchlng and
refuge
seeklng 1hese Lypes of Lrophlc and nonLrophlc behavlors allow rare or low
blomass
resource specles Lo perslsL ln boLh naLural and model ecosysLems lncreaslng
overall
communlLy perslsLence
ln anoLher approach Lo lnLegraLlng complex sLrucLure and dynamlcs and ln
conLrasL Lo mosL prlor dynamlcal sLudles kondoh (2003ab) allowed
consumer
preferences for resources Lo adapLlvely vary ln dlverse random and cascade
model food
webs Pe found poslLlve complexlLysLablllLy relaLlonshlps ln many of Lhese
adapLlve
neLworks Powever Lhe resulLs are confounded due Lo lncluslon of
lmplauslble llnear
nonsaLuraLlng specles lnLeracLlons and lnLernal growLh Lerms LhaL allow
consumers Lo
perslsL wlLhouL resources A sLablllLy reanalysls of adapLlve neLworks uslng
Lhe
WllllamsMarLlnez framework revealed LhaL poslLlve complexlLysLablllLy
relaLlonshlps
requlre an arblLrarlly hlgh fracLlon of adapLlve consumers (30) wlLh a hlgh
adapLaLlon
raLe ln slmple cascade model food webs and do noL occur ln Lhe more
emplrlcally wellcorroboraLed nlche model food webs (8rose eL al 2003b)
1ogeLher Lhese resulLs lndlcaLe
LhaL poslLlve complexlLysLablllLy relaLlonshlps due Lo adapLlve foraglng are
unllkely Lo be
found ln naLural sysLems
lnLegraLlng speclaLlon evenLs and assembly lnLo dynamlcal foodweb models
has
shown LhaL desplLe a hlgh speclesLurnover communlLles may assemble
lnLo perslsLenL
dlverse neLworks (Caldarelll eL al 1998 urossel eL al 2001) Powever Lhe
foodweb
neLwork sLrucLures LhaL evolve are aL besL loosely comparable Lo emplrlcal
food webs and
appear less complex ln some fundamenLal ways lor example Lhe
connecLances (Lhe
number of poLenLlal feedlng llnks LhaL are acLually reallzed llnks/specles
2
as calculaLed
uslng unwelghLed llnks) of 8 example webs are very low (C 1 Lo 6 mean
of 3
urossel eL al 2001 1ables 1 2) and connecLance decreases wlLh specles
rlchness ln
comparlson a seL of 18 recenL emplrlcal food webs have much hlgher
connecLances (C
3 Lo 33 mean of 14 uunne eL al 2004 1able 2) and dlsplay no
slgnlflcanL
relaLlonshlp beLween specles rlchness and connecLance (see also MarLlnez
1992)
neverLheless Lhls evoluLlonary foodweb model glves rlse Lo quallLaLlvely
lnLeresLlng
paLLerns such as perlods of hlgh communlLy dlverslLy and complexlLy
followed by
caLasLrophlc exLlncLlon evenLs leadlng Lo perlods of lower dlverslLy and
complexlLy 1he
communlLy evoluLlon approach has yeL Lo be analyzed ln Lerms of
complexlLysLablllLy
relaLlonshlps and lL remalns unclear whaL causes Lhe exLlncLlon evenLs
(urossel eL al 2001) Whlle Lhe lnLegraLlon of evoluLlonary and populaLlon
dynamlcs ln a foodweb
assembly model ls lnLrlgulng Lhls formulaLlon ls severely hampered by lLs
exLraordlnarlly
large number of parameLers renderlng Lhe lnLerpreLaLlon of resulLs as well
as Lhelr
relevance Lo naLural sysLems dlfflculL lf noL lmposslble
ln summary Lhe recenL exLenslon of Lhe numerlcal lnLegraLlon approach Lo
dlverse
model ecosysLems wlLh complex ecologlcally plauslble neLwork sLrucLure
offers an
approach for explorlng condlLlons LhaL glve rlse Lo and hlnder Lhe sLablllLy
of lndlvldual
populaLlons wlLhln a complex ecosysLem as well as Lhe sLablllLy of Lhe
whole sysLem
(Wllllams MarLlnez 2004b ms 8rose eL al 2003b) lnLeresLlngly many
resulLs from
Lhe lnLegraLed neLwork sLrucLure and dynamlcs models are conslsLenL wlLh
prlor local
sLablllLy analyses a generally negaLlve complexlLysLablllLy relaLlonshlp and
an lncreased
sLablllLy of emplrlcally conslsLenL foodweb sLrucLures ln comparlson Lo
random webs
(Wllllams MarLlnez ms) lnLegraLlng adapLlve or evoluLlonary processes ln
complex
communlLy models may lead Lo furLher lnslghLs regardlng how dlverse
complex naLural
ecosysLems may perslsL (Caldarelll eL al 1998 urossel eL al 2001 kondoh
2003 8rose
eL al 2003b) lL remalns a dlsadvanLage ln comparlson Lo Lhe local sLablllLy
approach LhaL
resource compeLlLlon among basal specles ls generally noL lncluded ln Lhe
nonequlllbrlum models buL lL can be readlly added ln general Lhe uLlllLy of
currenL and
fuLure appllcaLlons of Lhese more complex dynamlcal approaches wlll
depend Lo a greaL
degree on a careful balanclng acL beLween lmplauslble absLracLlon
assoclaLed wlLh fewer
parameLers and model clarlLy versus greaLer blologlcal plauslblllLy
assoclaLed wlLh more
parameLers and Lhe lncreased danger of model obfuscaLlon
AppllcaLlon Lo an AquaLlc LcosysLem
1he varlous approaches Lo modellng prevlously descrlbed have been used Lo
explore Lhe condlLlons for and llmlLaLlons upon sLablllLy and perslsLence ln
ldeallzed
ecosysLems Powever aspecLs of Lhe models may also lend Lhemselves Lo
more applled
lmplemenLaLlon allowlng assessmenL of sLablllLy of naLural ecosysLems ln
Lhe face of
acLual or anLlclpaLed perLurbaLlons 1hls lssue ls of parLlcular lmmedlaLe
concern ln
aquaLlc ecosysLems LhaL conLaln flsherles Whlle Lhe use of slnglespecles
models whlch
have domlnaLed flsherles research and managemenL ls generally
acknowledged as
lnadequaLe and ls slowly glvlng way Lo analysls of slmple foodweb modules
(?odzls
2001) Lhose modules may sLlll be lnadequaLe for dlscernlng Lhe llkely
ouLcome of
perLurbaLlons LhaL occur ln a dlverse mulLlspecles conLexL ?odzls (1998
2000) has
examlned Lhls fundamenLally lmporLanL lssue for boLh Lheory and pracLlce ln
Lhe conLexL
of wheLher culllng (removlng or kllllng) fur seals ls llkely Lo have a negaLlve
or poslLlve
lmpacL on Lhe hake flshery ln Lhe 8enguela ecosysLem
WlLhln a flsherles conLexL one vlew of 8enguela foodweb dynamlcs ls LhaL
people harvesL hake fur seals eaL hake and Lherefore lf people reduce Lhe
fur seal
populaLlon Lhere wlll be more hake for people Lo harvesL wlLhouL a neL
decrease ln Lhe
hake sLock Powever Lhls ls obvlously an exLraordlnary overslmpllflcaLlon of
a much
more complex ecosysLem lnsLead ?odzls (1998) conslders a more reallsLlc
food web
LhaL conLalns 29 Laxa lncludlng phyLoplankLon zooplankLon lnverLebraLes
many flsh
Laxa and several oLher verLebraLe Laxa (modlfled from lleld eL al 1991)
Whlle sLlll a
slmpllflcaLlon of Lhe dlverslLy and complexlLy of Lhe sysLem Lhls 29Laxa web
beLLer
capLures Lhe varleLy of paLhways LhaL can llnk componenLs of Lhe neLwork
1here are over 28 mllllon paLhs of poLenLlal lnfluence from seals Lo hake ln
Lhls foodweb conflguraLlon
Whlle many of Lhose paLhways may have negllglble lmpacLs on Lhe hakeseal
lnLeracLlon
lL cannoL be assumed a prlorl LhaL sclenLlsLs can lgnore mosL of Lhose
paLhways of
lnfluence when assesslng Lhe lnLeracLlon beLween Lhe Lwo specles (?odzls
1998)
?odzls (1998) uLlllzes Lhls more reallsLlc food web as Lhe framework for
sLudylng
mulLlspecles dynamlcs deLermlned by Lhe nonllnear bloenergeLlc consumer
resource
model dlscussed prevlously (?odzls and lnnes 1992) arameLer values for
Lhls model
were seL for Lhe 8enguela ecosysLem ln one of Lhree ways uslng fleld daLa
esLlmaLlons
(eg populaLlon blomasses average body mass) Lhrough allomeLrlc and
energeLlc
reasonlng and by allowlng unknown parameLers Lo vary randomly 1he use
of some
randomly varylng parameLers means LhaL probablllLy dlsLrlbuLlons raLher
Lhan parLlcular
soluLlons were calculaLed for Lhe ouLcome of lnLeresL WhaL Lhe lmpacL of
reduclng fur
seal blomass on Lhe blomass of hake and oLher commerclal flshes?" ln
effecL ?odzls ls
slmulaLlng an experlmenLal pressperLurbaLlonan experlmenL LhaL cannoL
be done ln a
sclenLlflc or eLhlcal way on Lhe acLual ecosysLem 1hls framework and
quesLlon requlres
Lwo furLher slmpllflcaLlons ?odzls assumes local perLurbaLlons (eg small
culls) and
equlllbrlum dynamlcs (eg consLancy ln parameLer values and dynamlcs)
Whlle we
have already seen LhaL Lhere can be problems wlLh Lhese sorLs of
assumpLlons Lhey allow
Lhe use of Lhe analyLlcally LracLable !acoblan maLrlx (used by May 1972
1973) Lo
deLermlne Lhe ouLcome of small longLerm perLurbaLlons ln a complex
sysLem whlch ls a
reasonable approach Lo use for Lhls parLlcular quesLlon 1hus Lhe analysls
comblnes
elemenLs of all Lhree approaches Lo mulLlspecles dynamlcal modellng we
have dlscussed
lncludlng local sLablllLy analyses of communlLy maLrlces nonllnear
bloenergeLlc
dynamlcs and complex foodweb sLrucLure 1hese modellng elemenLs ln
con[uncLlon
wlLh emplrlcal daLa on Lrophlc lnLeracLlons and varlous characLerlsLlcs of
Laxa ln Lhe
8enguela ecosysLem form a plauslble framework ln whlch Lo slmulaLe a cull
of fur seals
and lLs llkely effecLs on commerclal flshes ln Lhe conLexL of a complex
ecosysLem
uslng Lhls modellng framework ?odzls (1998) found LhaL lf fur seals are
culled
Lhere ls a slgnlflcanL probablllLy LhaL Lwo of Lhree commerclal flshes (hake
anchovy and
horse mackerel) wlll have negaLlve responses and LhaL across all of Lhe
commerclal
sLocks lL ls more llkely LhaL Lhere wlll be a loss Lhan a galn erhaps even
more
lmporLanLly ?odzls' analyses (1998 2000) underscore Lhe dangers of relylng
on overly
slmpllsLlc models Lo make predlcLlons Cne lmporLanL lssue ls how senslLlve
Lhe model ls
Lo lncluslon of apparenLly unlmporLanL Lrophlc llnksle can Lrophlc llnks
LhaL reflecL
some relaLlvely small percenLage of a predaLor's consumpLlon be lgnored
wlLhouL
affecLlng Lhe ouLcome of Lhe model? 1he answer ls no" for mosL Lrophlc
llnks As Lhe
cuLoff level (of percenLage of dleL) for lncluslon rlses above 10 resulLs
sLarL Lo vary
drasLlcally from ouLcomes seen on Lhe compleLe web Lhus showlng LhaL
Laxonomlc
compleLeness and resoluLlon deLalled foodweb sLrucLure does make an
enormous
dlfference ln predlcLlng Lhe ouLcomes of generallzed press perLurbaLlons
such as culllng
fur seals ln Lhe 8enguela ecosysLem" (?odzls 1998) 1he lnLeracLlon
beLween hake and
fur seals wlLhln Lhe 8enguela food web ls hlghly lnfluenced by many oLher
specles ln Lhe
web (?odzls 2000) and lL ls very llkely LhaL Lhose Lypes of lnfluences are
found ln many
lf noL mosL ecosysLems 1he presence of Lhese poLenLlally sLrong dlffuse
effecLs ln
complex food webs renders Lhe use of small modules as a means of
slmpllfled communlLy
analysls hlghly suspecL for many ecosysLems and many Lypes of quesLlons
(?odzls 2000) Concluslons
We have revlewed Lhree dynamlcal modellng approaches for sLudylng
complexlLysLablllLy relaLlonshlps ln absLracL food webs and some of Lhelr
cenLral
characLerlsLlcs are compared ln 1able 1 1hls Lype of modellng has lLs rooLs
ln llnear
equlllbrlum dynamlcs research Some sLudles conLlnue Lo uLlllze equlllbrlum
assumpLlons whlch may be approprlaLe for cerLaln klnds of quesLlons as
dlscussed ln Lhe
prevlous secLlon Powever slnce Lhe early 1990's alLernaLlve approaches
have Laken
advanLage of lncreaslng compuLlng power and advances ln nonllnear
dynamlcs Lo explore
populaLlon dynamlcs ln deLall uslng nonllnear and nonequlllbrlum
assumpLlons whlch
are more llkely Lo reflecL Lhe Lypes of dynamlcs LhaL occur ln naLural
sysLems 8aLher Lhan
belng resLrlcLed Lo Lhe blnary sLable/unsLable ouLcomes LhaL Lyplfy local
sLablllLy analyses
researchers can now explore a wlder range of Lypes of populaLlon and
wholesysLem
sLablllLy 1hls ls especlally Lhe case wlLh Lhe exLenslon of nonllnear
nonequlllbrlum
dynamlcal analysls Lo food webs wlLh complex plauslble neLwork sLrucLure
lor
example one new measure of sLablllLy ls Lhe percenLage of specles ln a
model food web
LhaL dlsplays perslsLenL dynamlcs (MarLlnez Wllllams ms) whlch can be
furLher
broken down lnLo Lhe percenLages of specles LhaL dlsplay dlfferenL Lypes of
perslsLenL
dynamlcs (eg sLable equlllbrla llmlL cycles chaos) (llgure 1) Some of Lhe
promlslng
new dlrecLlons for Lhls Lype of research lnclude exploraLlons lnLo
evoluLlonary and
adapLlve dynamlcs nuLrlenL dynamlcs spaLlal heLerogenelLy and non
Lrophlc lnLeracLlons
Powever addlng new layers of ecologlcal plauslblllLy Lo models carrles wlLh
lL Lhe rlsk of
generaLlng a parameLer space so large and unconLrolled LhaL lnLerpreLaLlon
of model
ouLcomes ls rendered an exerclse ln creaLlve Lalesplnnlng and Lhe llnks Lo
emplrlclsm
LhaL may have moLlvaLed Lhe addlLlon may be rendered worLhless Any
elaboraLlons need
Lo be carefully [usLlfled made as slmple as posslble and explored ln deLall
8aLher Lhan belng shackled [usL Lo quesLlons of wheLher specles rlchness
and
connecLance begeL" sLablllLy or lnsLablllLy researchers are expandlng Lhelr
search for Lhe
devlous sLraLegles" LhaL promoLe dlfferenL aspecLs of populaLlon and
ecosysLem
perslsLence and robusLness 8egardless of Lhe meLhodology used Lhere are
some
emerglng conslsLenL resulLs lndlcaLlng LhaL naLural food webs have non
random neLwork
sLrucLures and lnLeracLlon sLrengLh dlsLrlbuLlons LhaL promoLe a hlgh degree
of populaLlon
sLablllLy and specles perslsLence As recenLly polnLed ouL by McCann (2000)
Lhe model
and emplrlcal evldence (eg 8erlow 1999) LhaL many weak lnLeracLlons can
buffer a few
sLrong unsLable consumerresource lnLeracLlons Lhus sLablllzlng Lhe whole
communlLy ls
conslsLenL wlLh MacArLhur's (1933) hypoLhesls LhaL havlng more feedlng
llnks acLs Lo
dampen large populaLlon swlngs nonrandom neLwork sLrucLure
regardless of
lnLeracLlon sLrengLh also appears Lo buffer agalnsL populaLlon exLlncLlon
and Lhe more
reallsLlc Lhe foodweb sLrucLure Lhe greaLer Lhe specles perslsLence lrom a
purely
Lopologlcal perspecLlve lncreaslng connecLance may buffer agalnsL Lhe
probablllLy of
cascadlng exLlncLlons (uunne eL al 2002b) parLly because mlddle and hlgh
connecLance
webs have more unlform llnk dlsLrlbuLlons (Lhe dlsLrlbuLlon of Lrophlc llnks
per specles)
Lhan low connecLance webs whlch Lend Lo have powerlaw dlsLrlbuLlons
(uunne eL al
2002a) owerlaw neLworks appear especlally vulnerable Lo loss of hlghly
connecLed
nodes (eg AlberL eL al 2000 uunne eL al 2002b) lL should be clear LhaL
Lhere ls sLlll an enormous amounL of work Lo be done Lo
elucldaLe May's devlous sLraLegles" and ln Lhe face of wldespread
blodlverslLy loss and
ecosysLem perLurbaLlon Lhere ls an lncreaslngly compelllng seL of reasons Lo
do LhaL
research loodweb research has proven Lo be a useful vehlcle for many
LheoreLlcal and
emplrlcal quesLlons ln ecology and some of Lhe mosL herolc work wlll be
accompllshed
by people who are wllllng Lo Lry Lo lnLegraLe Lhe lessons from absLracL
models wlLh realworld appllcaLlons (eg ?odzls 1998) 8eyond ecology Lhe
analyses and resulLs LhaL
emerge from Lhe sLudy of complex food webs are llkely Lo reverberaLe back
ouLwards Lo
lnfluence our undersLandlng of oLher Lypes of bloLlc and abloLlc neLworks

AcknowledgemenLs
1hls work was supporLed by nSl granLs uL8/u8l0074321 (!Au) uL8
0083929 (nuM) u8l0234980 (nuM !Au 8!W) and u8l9904777 (8!W)
1hls
work was compleLed durlng a posLdocLoral fellowshlp aL Sll by !Au
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nd
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and daLa Amerlcan naLurallsL 163438468
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1rends ln
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Amerlcan
naLurallsL 13911311173 1able 1 Summary of Lhe sLablllLy analysls
meLhodologles
Local sLablllLy loodweb
module sLablllLy
Complex foodweb
perslsLence
lnLeracLlons llnear nonllnear nonllnear
equlllbrlum assumpLlon yes no no
small perLurbaLlons yes no no
chaoLlc dynamlcs allowed no yes yes
communlLy slze small Lo dlverse small dlverse
populaLlon dynamlcs analyses sLable /
unsLable
yes yes
communlLy sLablllLy analyses yes no yes
communlLy perslsLence yes no yes
compeLlLlon among prlmary
producers
yes no no
nuLrlenL dynamlcs no no no llgure 1 opulaLlon dynamlcs ln Llme serles
a
017
019
021
023
023
027
029
0 200 400 600 800 1000
Llme
opulaLlon denslLy
sLable equlllbrlum damped osclllaLlons
b
013
018
023
028
033
0 200 400 600 800 1000
Llme
opulaLlon denslLy
llmlL cycles ampllfled osclllaLlon
c
0
003
01
013
02
023
03
033
04
043
600 700 800 900 1000
A8S18Ak
Model ranLal makanan dua spesles merupakan model yang menyaLakan
lnLeraksl dua populasl yalLu populasl prey dan predaLor sedangkan model
ranLal
makanan Llga spesles merupakan model yang menyaLakan lnLeraksl Llga
populasl
yalLu populasl prey predaLor perLama dan predaLor kedua kedua model
lnl
dlperoleh darl komblnasl model perLumbuhan loglsLlk pada populasl prey
dengan
model predaLor yang menggunakan respon fungslonal Polllng Llpe ll Model
yang
dlgunakan merupakan persamaan dlfferenslal non llnear dengan Llga
varlabel Lldak
bebas yalLu x L
1
( ) yang menyaLakan [umlah populasl prey pada wakLu L x L
2
( )
yang menyaLakan [umlah populasl predaLor perLama pada wakLu L dan x
L
3
( )
yang menyaLakan [umlah populasl predaLor kedua pada wakLu L uarl hasll
anallsls kesLabllan yang dllakukan Lerhadap model ranLal makanan dua
spesles dan
Llga spesles dldapaLkan Llga LlLlk dan enam LlLlk keseLlmbangan dan
berdasarkan
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hLLp//eprlnLsundlpacld/26888/2/4_endahuluanpdf

DINAMIkA kAN1AI MAkANAN
inumiku Runtui Mukunun Luot

Dinamika rantai makanan laut adalah suatu ekosistem yang tercipta oleh alam itu sendiri, suatu dinamika
akan terjadi kalau suatu daerah kawasan terumbu karang mempunyai hamparan karang dan tumbuhan
laut minimal mempunyai area minimal 100 km2, luasan area ini terdiri dari area bukit karang, karang
tandes, area dasar laut yang berpasir dengan tetumbuhan laut .

Apabila syarat-syarat diatas terpenuhi akan tercipta suatu dinamika rantai makanan laut, persyaratan
tersebut diatas banyak dijumpai di ndonesia, seperti pada kawasan ujung kulon , laut selatan ndonesia,
laut sulawesi, laut banda dan laut arafura .
Ditempat-tempat seperti inilah ikan-ikan kecil bisa membangun suatu keluarga untuk berkembangbiak
dengan baik dan aman, dengan area yang cukup luas ini maka banyak ikan-ikan kecil yang
berkembangbiak menjadikan daerah tersebut suatu daerah kawasan hunian ikan-ikan kecil .

kan-ikan kecil tersebut terdiri dari ikan teri, ikan tembang, ikan lemuru, ikan kembung, ikan layang, cumi-
cumi, ikan kurisi, ikan tanda dan jenis ikan-ikan hias lainnya.
Pada musim ikan bertelur daerah hamparan terumbu karang tersebut akan dipenuhi dengan bermilyar-
milyar telur ikan dari berbagai jenis ikan, apabila telur tersebut menetas menjadi larva dan larva menjadi
besar dapat anda bayangkan didaerah tersebut akan penuh dengan ikan pelagis kecil .



Karena banyaknya ikan pelagis kecil akan mengundang kedatangan ikan predator kecil seperti ikan
barakuda, ikan kuwe gerong, ikan talang-talang, ikan tongkol, ikan tuna kecil dan ikan tenggiri kecil atau
wahoo kecil .
Sang predator kecilpun datang dengan jumlah rombongan yang cukup besar sehingga membuat
kepanikan ikan-ikan kecil penghuni kawasan terumbu karang tersebut lari tunggang langgang untuk
menyelamatkan diri dari mulut sang predator kecil yang bergigi sangat tajam bagaikan pisau .

Sumber hLLp//LanldanlkanblogspoLcom/2011/06/dlnamlkaranLalmakananlauLhLml



DINAMIKA RANTAI MAKANAN
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menggunakan penambahan membaLasl nuLrlens dengan budaya murnl aLau populasl alaml darl bakLerl
dan ganggangSLudl lnl memungklnkan penellLl unLuk menemukan konsLanLabloklmla
seperLl la[u perLumbuhan makslmum () dan unLuk mengukur kemampuan Lanaman unLuk
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9ff
Sedikit yang diketahui tentang dinamika rantai makanandaripada yang diketahui tentang faktor-
faktor lain seperti cahaya, suhu, atau distribusi dari berbagai spesiesorganisme. Sebagian
besar kesulitan dalam rantai makananpenelitian berasal dari ketidakmampuan kita
untukekstrapolasi akurat dari studi laboratorium untuk pertumbuhan tanaman alami atau
tingkat pakan ternak. Di laboratoriumhewan-hewan besar seperti ikan cepat
dapat mengkonsumsiberbagai mangsa. Sebaliknya, dalam kondisi
alamiah lebihpilihan mangsa terbatas dan tingkat menyusui jauh lebih rendah daripada yang
diamati di laboratorium. Dalam bab inikita membahas faktor-faktor fisiologis dan biokimia umum
seperti konstanta pertumbuhan enzim dan efisiensiprecentage transfer energi antara
tingkat trofik. Pendekatan ini memperluas metode klasik yang mengandalkan pada analisis isi
perut atau penurunan dalam jumlah spesies mangsa melalui pemakan
rumput atau predasi. Rantai makanan, ketika dikurangi dengan
diagram alirandisederhanakan, memberikan pandangan dari strukturdinamis dari ekosistem
perairan.
Pengukuran
Di dasar rantai makanan, fotosintesis diukur dalam kaitannya dengan cahaya, nutrisi, dan
berbagai faktor lingkungan lainnya. Berbagai elemen hadir dalam jaringan tanaman yang
mudah diukur dan menunjukkan nutrisi yang tersedia ke tingkat trofik berikutnya. Selanjutnya,
mereka dapat memberikan petunjuk mengenai faktor
pertumbuhanlimitingplant nutrisi. Pendekatan lain untuk menanam dinamikaadalah
untuk mengukur respon tanaman terhadap nutrisitambahan
dalam percobaan pertumbuhan. Untuk hewan airjumlah dan kesehatan telur
dan muda menunjukkan makanan dan energi yang ditransfer ke generasi berikutnya. Untuk
mempelajari jaring makanan limnologists menggunakankalorimetri bom, seluruh organisme kan
dungan gizi, dan nutrisi-serapan kinetika. Penyelidikan ini mungkin
melibatkanisotop radioaktif dan stabil sebagai pelacak. Rincian teknik yang paling umum
digunakan adalah diberikan dalam Manual,BP 8 12, dan volume APHA terbaru. ndividu atau
populasikeseluruhan dapat dibakar dalam kalorimeter bom dan energi yang diukur ini
kemudian digunakan untuk menentukan aliran energi. Pendekatan yang diambil sendiri
adalah terbatas karena limnologist adalah sebanyak peduli dengan nilai gizimakanan
sebagai dengan konten energi mutlak.
Pengaturan Nutrisi dan AIiran Energi
Sebelum mencoba makanan-rantai studi, sepanjang tahun pengukuran dari kelompok-
kelompok utama organism diperlukan. Dengan cara ini sebuah piramida biologi menjadi
kelompok-kelompok organisme mengelompokkan fungsional yang umum
seperti karnivora, herbivoures, detritivores, dll, dapat dibangun. Lebih-organisme
yang halus membelah pengukuranmenjadi kelompok funcional tertentu seperti lalat capung-
mengonsumsi ikan, alga-Scraping serangga atau
siput, bakteri heterotrofik, dll, memungkinkan jaring makananyang bisa
ditarik. Akhirnya, pengetahuan tentang energi dan nutrisi lewat di antara setiap
kelompok fungsional yang kecil akan memungkinkan sebuah jaringan dinamika makananyang
akan dibangun.