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Steneck & Dethier. 1999. A Functional Group Approach to the Structure of Algal-dominated Communities. Oikos

Steneck & Dethier. 1999. A Functional Group Approach to the Structure of Algal-dominated Communities. Oikos

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OIKOS
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Copcnhagen
1994
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A
functional group
a
n
svroach
to the structure of algal-dominatedcornmunities
Robert S. Steneck 'and Megan N. Dethier 
Steneck, R. S. and Dethier. M. N. 19Y~. A functional group approach to the ,tructure of algal-dominated communties. - Oiko" 69: 476-498.
R. S. Sreneck. Depr 
of  
Oceallograplr.'" am/ Cerzrer/or Marille SllIdit's. Dar/in? Mari,,!.' Cemer. Univ.
(Jj 
.'v!aine. lI-'alpo/e.. \fE 04573. USA. - M.
IV.
DelÍli"r. Insr. .•.or EIl\'i-l'(}Illllell/a/ SllIdies and Fridav Harbor Labf}r(l(ories. U"i\'. oj \\ilshi"!!!l";. Frie/a\' H'lrbor. WA 98250. USA.
 Natura communities shauld be described in rerms simple
6~~
R "
",\f1;'óO~ .
e_:n_lu a be rs
000
and deraded enough to convéyuseful information about their structure and functionalcomponents. There exists a bread spectrum 01' ways todescribe panems. At one end 01'this specrrum. species arethe fundamental unit 01'measure. Because many factorscontribute to the distribution and abllndance of a givenspecies, it is often impossible to predict its behavior consistently. At the other end of rhe spectrum are func-tional groups which categorize species according to fea-tures such as body plan. behavior 
01'
rife history strategy.In this paper, we argue that analyzing cOIlununity pat-tems for marine algae via groupings based on functionalaspects of their morphology and anatomy provides sub-
Accepted 17 September 1993Cop}right
©
OIKOS 1994
ISSN 0030-1299
Printed in Denmark - aH rights reservecl
476
stantial insight into communiry strucrure. A functionalgroup analysis can be applied more broadly in 'pace fol'making biogeographical comparisons. :md in time foreconstrucríng paleocommllnities, than is possible at rhelevel of spec:.::s
01'
among relared higher taxa.O\'er the past three decades mosr community ecol-ogis(s. followin" the lead of Hutchinson and ~[;:¡c.-\rthur.
..... (.) ;C.O¡.dlO. .
stressed the
l/l1lqut!ness
of specles. That no
(\\'0
specle~can occupy (he same ecological niche has been a drivingaxiom stimulating interest in a variety oi topics includingcompetiríon. niche compression. ~haracrer displa~emenl.resource panitioning and species diversificatian. Ho\\'-ever. many af these cancepts have been qLles[ion~d andrecently criticized (e.g., see Sale 1977. Peters 1991. Bond
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MORPHOLOGY
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THALLUS S1ZE (•• 1
4
CORTtCATEDMACl'.OPHYTI:S
(ta'e~)
Clt.ondnu
and
Gi,artUia
nJNCTIONAL CROUPS COMPARATIVE ANATOMY
1
MlCROALGAE
(sin,l •••
U)
Cyanoboc1eria and di.""",
2
FILa.MENTOIJS ALGAE(uniseriato)
C1DdoploDr4 aN1B ••• , ••
3
roUOSE ALGAE (.inSk Uyer)
MOftOStTOII't4 
ar multiIayered
VI••••P"'T'."a
5
LEATHERYMACROPHYTES
Kelp and
Fucus
6
ARnC'.Jt.ATED CALCAREOUSALGAE
C""alJiNJ
andHtJi~d4
7
CRUSTOSE ALGAE
 Lilholluvnnion,
 PIJ$1DNV/i4and 
"Ralfsi."
CORnCA TED FOUOSE
3.5
ALGAE
 Djc1yo1a
and
Padind 
Fil!. l. Diagrammaticrepresentation ol" algalfunctional groups.Anatomical components arenot drawn to scale. butillustrate tissuedifferentiation such as between the cortex andmedullary regions of thethallus 01' a macroalga. Thespecific form of eachfunctional group illustratedis indicated in parentheses.A few representati ve gener;¡are given as examples foeach group. Numbers at len,algal group ("AG") numbers,are for quick reference(rankings are described inResults). Note that groups of a given morphology withincceased anaromicJicomplexity (e.g .. they arecorticatcd) are designated byhigher AG numbers. Thus.thinly corticated or  polysiphonous filaments become AG 2.5 andsimilarly corticated foliosefonns becorne AG
1.5.et al. 1992). A funcrional group approach, in contrast,stresses similarities among unrelated species that sharecritical organismal fearures. We discuss the overridingimportance of a small number of species attributes to thestructure of benthic marine algal communities, and notethat these attributes may be shared polyphyletically.The functional group approach, although having re-ceived little attention among community ecologists, mayhave been foreseen by MacArthur (1972) when he pre-dicted that "the future principIes of the ecology of coexis-tence will... be of the form 'for organisms of type A inenvironment of structure B, such and such relationshipwill hold'" (boldface ours). We will consider the types of marine algae that live under specitic marine environ-ments (defined below). We offer data and examplcs insupport of this approach as an altemative to other meansof studying communities. rather than as
éL
strict test of ouhypothesis. Our objective is to examine pattems
01"
algalfunctional group abundance, diversity and dominancerelative
10
extrinsic charucteristics of their environmenl.To demonstrate rhe broad applicability of the functionalgroup approach. we examined three biogeographicallydi'stinct regions: the westem North Atlantic, the eastem North Pacifico and the Caribbean as exemplified by sitesin Maine, Washington and St. Croix (U.S. Virgin Is-lands), respectively.
Definition of terms and rationale
We explore the hypothesis that observed pattems in thedistribution and abundance of life forms of algae (func-tional groups) result largely from two environmental pa-rameters: 1) productivity potential (factors that contribmeto the maximum possible rute of biomass production) and2) disturbance potential (factors responsible for the maxi-mum possible rute of biomass lost). It is important
lO
note
OtKOS 69:3 (1994)
477
Q
«;&Y:¡iLUtnm ....
.su
,-' ., _C
 
B
i
=
+
Pn, - Di' where
Fig. 2. The relati0nship between disturbance imensity and fre-qency. LOW, MEO and HIGH refer to disturbance levels.
Si
=
Biomass of an algal speeies
01'
functionaJ group (i)that accumulates over sorne period of time. i
=
thespccilic algal functional group(s) conccmed i.e .. 1-7(sce Fig. 1). The numeric designalion in Fig. I is basedon Ihe ranking of mass-specitic productivity (see Re-sults).R,
=
Rate of recruitment. which is a function of:1) lntrinsic properties of the algal species or funetionalgroup
(i)
such as the numbt:r and viability of propa-gules .2) Extrinsie properties of the environment or therecruitment potentinl of the environment (such asavailability of free space for gerrnination).Pni
=
Rate of o~t plj,wary producti)'. whieh is a fune-tion of:1) lntrinsic properties of the algal speeies or functionalgroup (i) or the "Mass-speeitie rate of production".2) Extrinsic properties of the environment or the"Productivity potemial of Ihe environmem".Di
=
Rate of herbivore-induced disturbance. which. as-suming no refuge, is a funclion of:1) lntrinsic properties of the oiga! species
01'
fünctic.'Oalgroup
(i),
which involves:Resistance to disturbam.:e (e.g .. due
10
mechanical properties sueh as roughness and morphology). anddeterrence of disturbance (e.g .. ehemistry of preyspecies affecting herbivore choice).
2)
Ex{nnsic properties of the environmem which werefer to as the: "Distllrbance potential oi the envi-ronmem". This can be measured as:Rate of disturbance (e.g., from herbimres). involv-ing:Disturbance intensity (amount of biomass lost peevent).Disturbance frequency (events per unir rime).We define the "productivity potemial of an en\ironmem"as being detennined by the extrinsie factors that ser anupper limir to the net primary producrivity possible inthat environment. A reduction in productivity potemial of the environment by this definion equals an increase instress (sensu Grime 1981). In rhe marine realm. factorsintlueneing rhe productivity potential include lighr. nutrí-ems, desiccation. freezing, and water motion (\\hich con-trols both nutrient and gas exchange: e.g., Blinks 1955.Leigh et al. 1987). Thus the productivity potential on hardsubstrata decreases in a logarithmic fashion from maxi-mum levels in lhe lower intenidal and shallo\\ subtidallOne toward minimal levels in the upper reaches of theintertidal lOne and the lower limits of the photic lOne.Evidence for these gradients have been published from avariety of locations (Nicotri 1977, Raffnelli 1979. Round1981, K"eser and Larson 1984, Underwood 1984a, b.Hardwick-Wirman 1985, Bosman et al. 1986. illustratedin Hawkins and Hartnoll 1983, Steneck et al. 1991).Grime (1981: 39) defines disturbanee to be ··the mech-anisms which limit plant biomass by cnusing its partial
01'
total destruetion·'. Disturbance has two componems: fre-quency and intensity (Reichle er al. 1975, Steneck 1988.Steneck et al. 1991). High Icvels of disturbance result
UNES OF EQUALDISTURBANCE(EQUAL BIOMASSREMOVED)
 NO VIABLE STRATEGYFOR PERSISTENCELOW
DISTURBANCE FREQUENCY
r  
E-
•...•
Cf)
z
~
E-
Z
•...•
that the productivity and disturbaripotemials of theenvironment are theoretically independem of resident plam assembiages and tllus may not directly reflect theobserved level of productivity
01'
disturbnnce in the sys-temo Independence of structuring environmemal compo-nems from organisms comprising the eommunity is es-sential (e.g., Van del' Steen and Scholten 1985, South-wood 1988) and thus we will earefully detail the imrinsicand extrinsic components of the community and theienvironment below. At this point we will uevelop thisidea exclusively for herbivore-induced disturbanees. al-though most aspeets apply equallY well ro abiotic disturb-ances.Functional groupings of algae are based on anatomicaland morphologieal characteristics (Stcneck and Watling1982, Steneek 1988, Fig. 1) that often eorrespond toecological charaeteristies (identified below). Thus theydiffer from guilds (sensu Roar 1973). whieh are basedstrictly on sim!larities in resource utilization. We consider an algal-dominated eornmunity as an assemblage of fune-tional groups. with the abundance of cach group mea-sured by its somatic biomass. Biomass is maintained by adynamic balance between the rates of constructive forcesof recruitment and net primary
01'
biomass production andthe destructive force of disturbance. The measurable re-sult of these processes depends on.both imrinsic proper-ties of the organisms and extrinsic properties of the envi-ronment. This relationship can be expressed in biomassunits for any given area as:478
0110;05
09:.1
119'M)

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