Responses of a Key Intertidal Predator to Varying Recruitment of Its Prey
Carlos Robles, Robin Sherwood. Stephens, Mario Alvarado
Ecology, Volume 76, Issue 2 (Mar., 1995), 565-579.
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RESPONSES OF A KEY INTERTIDAL PREDATOR TO
VARYING RECRUITMENT OF ITS PREY’
Caatos ROBLES, ROBIN SHERWGOD-STEPHENS,? AND MARIO ALVARADO
olows Department, California State Cntvertin, Lox Angeles, Califone 90032 11SA
Abstract How a predator resgonds to fluctuating prey abundances determines its ca
pacity Co regulate the prey’s population levels. Uslike many terrestrial and a few marine
arthropods, Key intertidal invertebrate predstors, ¢.@, sea stars and whelks, have been
portrayed 25 licking rapid behavioral responses that would permit them to control episodes
‘of massive prey zeerultment. Que field experiments demensiated that intexidal sea stars,
Phaster ochraceus, rapidly change densities and alter diet in response 0 var Ying recrattment
ff their prey, the mussels Motilur spp. Densities of Pisaster foraging at high tide were
positively correlated with an index of recruitment of Afyilus califarnianus. The sea stars
Aggregated in episodes of massive mussel recruitment and dispersed as local abucances
‘of juvenile mussels declined. Agaregation and dispersal were reproduced in field experi-
ents by adding or removing masses af Ist-ye Mytilus pp. to naturally occurring mussel
beds
‘Over the 7 yr ofthe study, the lower margins of the mussel beds, a distributional limit
set at least im Some circumstances by the sea stars, remained relatively constant, despite
significant year-co-yeat variation in mussel recruitment rates. We speculate dhat aggregation
acts in concert with previously hiypochesized long-term responses to stabilize dhe lower
distributional limite oF the mussels.
Key nord: ggrenative response; foraging: intertidal gonaton: Mytilus; Pisaster; population
regulation: predaion: predator response: sab)
Intaopuction
Predacors respond in diverse ways as prey sbun-
dances change. Individual, predators respond by shift-
ing diets (eg., Tinbergen 1960, Murdoch 1969, Rusen
et al. 1972), altering foraging behavior (e.g.. Banks
1997, Royama 1970, Curia 1976), changing. growth
tates (e.g., Swingle and Smith 7949, Dillard and Nov-
inget 1975, Paine 1976), and ultimately, thriving oF
perishing (e.g, Adams and DeAngelis (987). Oceur-
ring Caroughout the population, these individual actions
constitute predator population tesponses to changing
prey population structure, As examples, shrews aggre
sate in local concontrations of saw fly larvae (Holling
1959), and young-of-the-year largerouth bass grow
and successfully overwinter, or remain small and die,
depending upon order of magnitude ucwations in the
availability of their forage fsh (Adams anc DeAngelis
1987)
‘The types of responses in 2 predator’s repercnice are
importent because they may determine whether and
hhow the predator regulates the prey population (Hol
1959, Hassel and May 1974, Murdach and Oaten 1975,
Hassell 1978, Murdoch and Bence 1987, Hassel and
May 1990). Responses based on behavioral mecia
nisms, ¢.g, functional and aggregative tesponses (re~
viewed for marine species by Valiela 1984), may permit
“Meacuserige received 16 July 1995; revised 6 June 1894;
accepted 21 fuse 198
“Present adcress: Department of Biology, Univesity af
Pennsylviain, Phisdelpss, Pemneylvania (3100-80L8 USA
predation to “track” changes i prey abundance with
comparatively little delay. Type 3 sigmoidal curve}
functional responses have the potential to generate sa
bilizing mortality, but they appear to be elatively rare,
at least among, invertebrate taxa (Waliela 1984, Mur
doch and Bence 1987). Aggregative responses concen
trace predators in local areas, and may play an important
sole where prey populacons occur as local ufestations
‘oF “patches” Consequently, theoreteal teatments
have focussed an the circumstances in which predator
(oF parasitoid aggregations may stabilize prey popula
toons (eg, Massel and May 1974, 1990, Murdach and
‘Oaten 1975, 1989, Hasse] 1978, Murdoch etal. 1992),
In contrast to responses based on behavior, changes
in predatar reproduction or maxtality rates cnomeriest
responses, Holling 1959, Valiela 1984) or changes in
body size or species With indeterminate gro%t, a type
‘of developmental response (Murdoch [97f}, fallow
‘comparatively long time courses. Suck slow responses
are thought (9 have limited potential to effect stabiliz-
ing regulation because the resulting time lags generated
instability in model predator-prey systems (discussion
im Hassel 1978, Hastings (984, Murdoch end Bence
1987),
Experimental confirmation of aggregative responses
in marine benshic communities 1s limited to a single
taxa’ brachyuran crabs. Aggregation by shell-crushing
crabs produced density-depencent mortality of mfaunal
bivalves whose densities were manipulated in intertidal
Goulding and Hay (984) and subtidal (Hines et al
1990, Mansour and Lipcius £991} arenas. However, che 386
mast intensively stodied benthic predators, whelks and
sea stars, appear comparatively sluggish, and are some
times portrayed as lacking an effective aggregation re~
sponse (e.g, Palumbi and Freed 1988, bar see Pair.
‘weather 1987 for an anesdocal account of aggregation
bby whelks following manipulations of their ptey)
‘The ease af the sea star Pisaster ochraceus deserves
attention because i functions as 2 keystone predator,
at least in some circumstances, limiting the, abundance
of the mussel Myclus californianus, the apparent cox
petitive dominant on exposed eastern Pacific shores
(Paine (96, 1974, 1976). Pisaster is intolerant of pro~
longed exposure, aad the impact of its foraging dimin-
ishes cowards higher shore levels, which remain ex-
posed longer during che tidal oyele. Above some critical
shore level, mussel receusts survive well, accumulating
as dense beds of adults. At lower levels the sea stars
usually lf al mussel recruits, but oceasionally ME
cealiforsiantes may survive long enough t© gro t6 ine
‘vulnerable sizes, one instance of size-limited predation
(Paine 1976)
How does Pisarter respond wo differences in the
availability of mussels within the tone ofits foraging?
Paine (1976} suggests thac the sea star's recruitment 1s
rate of infrequent and that movements alangshore of
adult sea scars are negligible. Observing many separate
sites, he noted an inverse eelationship between sea star
densities and their mean body masses, suggesting that
indeterminate geowth allows this predator t@ adjust
body size to its density and the level of prey avaulabulity
chartcteristic of a given site. Stich adjustments con-
stiute 2 fong-term developmental response because
booth the size and numaber of prey killed inerease with
the sea star's size
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