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Recent Talks
Kinlan* and Koch, et al. 2010 (Contributed Talk)
ASLO/AGU/TOS Ocean Sciences Meeting, Portland, OR, 22-26 Feb 2010

EMPIRICAL ESTIMATION OF LARVAL CONNECTIVITY ACROSS A SPECIES RANGE FROM MULTIVARIATE NATURAL TAG-RECAPTURE DATA
Kinlan, B.P.; Koch, S.E.; Zacherl, D.C.; Warner, R.R.

Predictive understanding of marine population dynamics at regional to global scales requires quantitative estimates of connectivity across species' ranges. Yet, the difficulty of tracing individual settling larvae back to their natal populations has stymied efforts to directly estimate connectivity.  The few successes in this endeavor are limited to cases of small numbers of distinct potential source populations (e.g., estuarine or island species) with unique natural geochemical or genetic signatures. The problem is much more difficult for open coast marine species, where unsampled potential source populations inevitably outnumber the locations sampled to develop a source atlas. Here, we present a novel statistical approach that couples recruit-to-source assignment with a multivariate model of spatial correlation in natural signatures to account for the contribution of unsampled sources. Applying this technique to geochemical data for a marine gastropod, Kelletia kelletii, we produce high-resolution (10km) alongcoast probability profiles of larval origins, estimate dispersal kernels between pairs of sites, and build an empirical connectivity matrix for the entire species' range. This new assignment method outperforms discriminant function analysis and existing Bayesian assignment techniques, provides full measures of uncertainty, and produces outputs that directly translate into parameters of spatial population models. It can also incorporate available information on the relative reproductive output of potential sources (e.g., from combining adult density, fecundity, and habitat distribution), and evaluate competing hypotheses about dispersal processes via model selection.  The result is a direct link between theoretical and empirical studies of spatial marine population dynamics.
Talk Abstract Archive
Kinlan* and Koch 2009 (Contributed Talk)
WSN 90th Annual Meeting, Seaside, CA, 12-15 Nov 2009

A NOVEL STATISTICAL METHOD TO PINPOINT LARVAL ORIGINS AND ESTIMATE RANGE-WIDE CONNECTIVITY IN OPEN COAST MARINE POPULATIONS
Kinlan, B.P.; Koch, S.E.

Predictive understanding of marine population dynamics at regional to global scales requires quantitative estimates of connectivity across species' ranges. Yet, the difficulty of tracing individual settling larvae back to their natal populations has stymied efforts to directly estimate connectivity. The few successes in this endeavor are limited to cases of small numbers of distinct potential source populations (e.g., estuarine or island species) with unique natural geochemical or genetic signatures. The problem is much more difficult for open coast marine species, where unsampled potential source populations inevitably outnumber the locations sampled to develop a source atlas. Here, we present a novel statistical approach that couples recruit-to-source assignment with a multivariate model of spatial correlation in natural signatures to account for the contribution of unsampled sources. Applying this technique to geochemical data for a marine gastropod, Kelletia kelletii, we produce high-resolution (10km) alongcoast probability profiles of larval origins, estimate dispersal kernels between pairs of sites, and build an empirical connectivity matrix for the entire species' range. This new assignment method outperforms discriminant function analysis and existing Bayesian assignment techniques, provides full measures of uncertainty, and produces outputs that directly translate into parameters of spatial population models. It can also incorporate available information on the relative reproductive output of potential sources (e.g., from combining adult density, fecundity, and habitat distribution), and evaluate competing hypotheses about dispersal processes via model selection. The result is a direct link between theoretical and empirical studies of spatial marine population dynamics.
Kinlan* 2008 (Contributed Talk)
WSN 89th Annual Meeting, Vancouver, BC, Canada, 6-9 Nov 2008

QUANTIFYING THE LOCAL INFLUENCE OF UPWELLING ALONG COASTLINES
Kinlan, B.P.

Coastal upwelling drives fluxes of heat, nutrients, larvae, phytoplankton, and other dissolved and suspended matter to intertidal and shallow subtidal ecosystems on eastern boundaries of ocean basins worldwide.  Ecologists working in these systems often wish to characterize upwelling variability at local spatial scales relevant to ecological field sites (<1km).  The definition of a local upwelling index is problematic, however, because fundamental physical drivers of coastal upwelling (wind, flow, and planetary motion) operate over scales much larger than the site (10's to 1000's of km).  Upwelling-related responses observed at coastal sites are actually the local manifestation of offshore meso- to regional-scale processes, modulated by smaller-scale circulation and biophysical processes often associated with topographic and bathymetric features near the coast.  The local expression of upwelling can be damped or amplified and exhibit spatial and temporal lags compared to the offshore phenomenon.  To quantify the local influence of upwelling, I combined global 50-km satellite/model blended wind data (QuikScat/NCEP 6-hour wind velocity and wind-stress curl, 1999-2008) with digital coastlines to estimate offshore upwelling transport in the vicinity of coastal sites in Western North America.  Then, I computed lagged spatiotemporal correlations between upwelling and site-specific biophysical time series (temperature, nutrients, chlorophyll, and settlement).  This approach resulted in site-specific coefficients relating each of the local responses to upwelling with the offshore wind forcing.  I show that the topographic and bathymetric structure of the coastline can be used to estimate these coefficients, leading to a predictive framework for alongshore variability in the influence of upwelling.

Kinlan* et al. 2007 (Contributed Talk)
WSN 88th Annual Meeting, Ventura, CA, 8-11 Nov 2007

POPULATION DYNAMIC RESPONSES OF THE CALIFORNIA SPINY LOBSTER  (PANULIRUS INTERRUPTUS) TO 120 YEARS OF VARIABLE OCEAN CLIMATE, RECRUITMENT, AND FISHING PRESSURE
Kinlan, B.P.; McArdle, D.; Emery, K.; S.D. Gaines

Long historical data sets that encompass wide variation in abundance, fishing pressure and ocean conditions are rare, but invaluable for understanding population dynamic responses of exploited species to past and future changes in climate and fishing.  We used catch and effort records from the beginning of the California spiny lobster (Panulirus interruptus) fishery to the present (1888-2006), to reconstruct likely changes in abundance, size structure, growth, recruitment, and mortality of P. interruptus since the fishery’s inception.  Information from process-oriented field and laboratory studies of lobster growth, mortality, and recruitment was synthesized with historical information on lobster fishing in an hierarchical Bayesian statistical framework, which was used to estimate a size-structured state-space population model.  The results reveal a complex interplay between inter-annual and decadal fluctuations in ocean climate (ENSO and PDO), changes in fishing effort (new technologies, closures and reductions of effort during wartime), and density-dependent population dynamics, and highlight historical changes in lobster abundance and biomass that span several orders of magnitude.  We conclude by exploring the possibility of long-term feedbacks between lobster abundance and lobster habitat quality via their trophic role in the kelp forest, with implications for ecosystem-based fishery management in the Southern California Bight.
Kinlan* et al. 2006 (Contributed Talk)
WSN 87th Annual Meeting, Redmond, WA, 9-12 Nov 2006

Do observed self-recruitment rates require special behavioral and oceanographic features?
Kinlan, Brian P.; Gaines, Steven D.; Siegel, David A.

Marine life histories and fluid characteristics create the possibility for long-distance dispersal, but larval behaviors and persistent oceanographic features may reduce dispersal and enhance local recruitment.  Lack of quantitative dispersal data has fueled debate about the fraction of marine larvae that return to their natal population (self-recruitment), or emigrate to another population (export).  Decoupling of local population processes from local production has been used to argue that marine systems are open and connected at large scales. Other recent studies have highlighted evidence of behaviors and circulation patterns that restrict dispersal and question the degree to which marine populations fulfill their potential for open demography. Synthesis of these perspectives has been hindered by the lack of an appropriate null hypothesis that makes quantitative predictions about expected rates of self-recruitment in the absence of special behaviors and oceanographic features.  We review recent estimates of self-recruitment rates and compare them with predictions from a null model incorporating only larval duration, competency period, and passive transport by turbulent ocean currents. We find that in many instances passive transport alone can explain relatively high rates of self-recruitment (5-50%) at relevant spatial scales (5-100 km), implying that rates of self-recruitment previously viewed as “surprising” are in fact indistinguishable from those expected under a null passive model.
Kinlan* and Broitman 2006 (Contributed talk)
7th International Temperate Reef Symposium (7ITRS), Santa Barbara, CA, 26 June-1 July 2006

Predictability in Intertidal Ecosystems: Scale-Dependent Coupling of Coastal Geomorphology, Oceanography, and Benthic Community Structure
Kinlan, B.P.* and Broitman, B.R.

A predictive approach to coastal ecology requires quantitative description of biophysical coupling over a continuous range of scales from meters to thousands of kilometers.  Here, we introduce a new approach for examining the strength and scale of coupling between biological patterns and coastline topography, and apply this method to develop a quantitative model of spatial variation of rocky intertidal community structure in three eastern boundary current ecosystems: western North America, Chile, and South Africa.  Using data from intertidal field surveys conducted over multiple years with similar methods on three continents, we found significant correlations between topographic features and community structure at several distinct spatial scales.  The scale-dependence of these relationships was consistent with the operation of distinct biophysical processes at different scales (e.g., upwelling, wave exposure).  When combined in a multi-scale model, simple indices of coastline morphology explained a significant fraction of the variance in intertidal diversity and abundance of functional groups.  The degree of predictability (maximum percentage of variance explained by topography) varied among continents and regions within continents.  Preliminary evidence suggests that predictive power is maximized when the length-scales of coastal features closely match the characteristic scales of nearshore physical processes.  Thus, the geological history of coastal regions may interact with dominant physical forcing processes to determine the predictability of ecological patterns at large scales.
Cavanaugh* and Kinlan, et al. 2006 (Contributed Poster, presented by co-author)
ASLO/AGU/TOS Ocean Sciences Meeting, Honolulu, HI, 20-24 Feb 2006

REMOTE SENSING OF KELP FOREST DISTRIBUTION AND DYNAMICS IN THE SANTA BARBARA CHANNEL USING SPOT IMAGERY
Cavanaugh, K.; Kinlan, B.P.; Reed, D.; Siegel, D.A.

A method for the remote assessment giant kelp (Macrocystis pyrifera) canopy cover is presented for the kelp beds of the Santa Barbara Channel, California for use with multispectral data from the SPOT 5 satellite.  The algorithm identifies kelp-covered pixels by the near-infrared to green band ratio after atmospheric correction by the dark pixel method.  High values of this ratio identify pixels with a surface canopy of giant kelp.  The SPOT satellite kelp coverage determinations are validated on both pixel and bed scales using data sets that include monthly aerial estimates of biomass, available photographic surveys of kelp cover, and diver mapping of kelp bed extent. Estimates of kelp canopy area are combined with field estimates of per-unit-area kelp net primary production (NPP) to enable assessment of temporal changes in regional kelp NPP.  This method is being used to create kelp cover maps on a monthly basis in order to observe intraseasonal to interannual changes in kelp cover and to determine spatial variation in persistence.  The derived coverage will be used to observe dynamics of kelp canopies in relation to changes in sea surface temperature, wave action, terrestrial runoff, and ambient chlorophyll concentrations on monthly time scales.  Future work will include investigating relationships between other aspects of kelp bed health (e.g., canopy senescence) and reflectance.
Kinlan* et al. 2005 (Contributed Talk)
WSN 86th Annual Meeting, Seaside, CA, 17-20 Nov 2005

PREDICTABILITY IN INTERTIDAL ECOSYSTEMS: SCALE-DEPENDENT COUPLING OF COASTAL GEOMORPHOLOGY, OCEANOGRAPHY, AND BENTHIC COMMUNITY STRUCTURE
Kinlan, B.P.; B.R. Broitman; S.D. Gaines; C.A. Blanchette; E. Wieters; S.E. Lester; P.T. Raimondi

Much recent ecological research has focused on generalizing results from small-scale observations to predict large-scale patterns of distribution and abundance.  In coastal ecosystems, nearshore physical processes such as upwelling, circulation, sediment transport, and wave propagation interact with topographic features of the coastline at a range of spatial and temporal scales.  These physical processes, in turn, exert a strong influence on the structure of nearshore biological communities.  A predictive approach to large-scale coastal ecology requires a quantitative description of biophysical coupling over a continuous range of scales from tens to thousands of kilometers.  Here, we introduce a new approach for examining the strength and scale of coupling between biological patterns and coastline topography, and apply this method to develop a quantitative model of spatial variation in intertidal community structure in three eastern boundary current ecosystems: western North America, Chile, and South Africa.  Using data from intertidal field surveys conducted over multiple years with similar methods on three continents, we found significant relationships between topographic features and community structure at several distinct spatial scales.  When combined in a multi-scale model, simple indices of coastline morphology explained up to 80% of the variance in intertidal diversity and abundance of functional groups.  Moreover, the distinct spatial scales of coupling could be traced to distinct physical mechanisms (e.g., upwelling vs. wave exposure), via a combination of theoretical and numerical models and remote-sensing data.  The degree of predictability (maximum percentage of variance explained by topography) varied among continents and regions within continents.  Preliminary evidence suggests that predictive power is maximized when the length-scales of coastal features closely match the characteristic scales of nearshore physical processes.  Thus, the geological history of coastal regions may interact with dominant physical forcing processes to determine the predictability of ecological patterns at large scales.
Kinlan* et al. 2005 (Contributed Talk)
2005 TOS International Ocean Research Conference, Special Session on Connectivity in Marine Populations, UNESCO, Paris, France, 5-9 June 2005

DO OBSERVED SELF-RECRUITMENT RATES REQUIRE SPECIAL BEHAVIOURAL AND OCEANOGRAPHIC FEATURES?  USING NULL MODELS TO EVALUATE THE EVIDENCE FOR ALTERNATIVE HYPOTHESES
Kinlan, Brian P.; Gaines, Steven D.; Siegel, David A.

Life histories and fluid characteristics in the ocean create the possibility for very long distance dispersal.  At the same time, larval behaviours, life history strategies, and persistent oceanographic features provide opportunities for shorter dispersal and site-fidelity.  The wide range of possible dispersal distances and scarcity of quantitative dispersal data in the marine environment have led to a debate on the degree to which marine larvae produced in a local population are likely to return to that population (self-recruitment) or emigrate to another population (export).   Two opposing perspectives have emerged in this debate.  On one side, evidence of decoupling of local population processes from local production has been used to argue that marine systems are open and connected at large geographic scales.  On the other hand, recent studies have highlighted evidence of restricted dispersal and question the degree to which marine populations fulfill their perceived potential for open demography.  Often, restricted dispersal is attributed to particular behavioural or oceanographic features.  Synthesis of these perspectives on connectivity has been hindered by ambiguity over expected rates of self-recruitment in the absence of special behaviours and oceanographic features.  Here, we describe a simple Lagrangian model, incorporating only larval duration, competency period, and passive transport by turbulent ocean currents along an idealized coastline, that can be used to generate a quantitative “null hypothesis” for rates of self-recruitment at a particular spatial scale.  Parameterizing the model with current velocity statistics and spatial scales corresponding to empirical studies, we review recent estimates of self-recruitment rates and compare them with the null model of passive transport.  We find that passive transport alone can explain relatively high rates of self-recruitment (5-50%) at relevant spatial scales (5-100 km) in many species.  Self-recruitment rates higher than expected from a simple passive model are found in some, but not all, of the empirical studies.  Our results highlight the utility of quantitative null models in examining the weight of the evidence for more complex alternatives.
Kinlan* 2005 (Invited Poster)
Hertz Foundation Research Symposium: Catalyzing the Future, San Jose, CA, 18-20 March 2005

Scaling and forecasting the spatio-temporal dynamics of coastal ecosystems
Brian P. Kinlan

(I) Biophysical coupling. Over the past five years, I have employed a combination of satellite and aerial remote sensing, field surveys, drifters, and moored instruments (depicted at left) to characterize the spatio-temporal stucture of the northeast Pacific coastal ecosystem. The dominant energy inputs to this
system are derived from photosynthesis of phytoplankton and large macroalgae (kelps) in the shallow euphotic zone. Primary production, in
turn, is strongly coupled to the process of coastal upwelling, which varies in space and time in response to atmospheric forcing at scales ranging from
10’s to 1000’s of km and days to decades. Below, I summarize some of the approaches I have taken to examine biophysical coupling in this region, resulting in a statistical model of ecosystem structure and dynamics.
(II) Larval transport in turbulent coastal flows. The coastal ocean is a turbulent environment where larvae are advected by ambient currents while
they develop competency for their next life stage. Among the vast number of released larvae, very few successfully settle upon suitable habitat and recruit to adult life stages. Key to the predictive understanding of nearshore marine ecosystem is this source/settlement relationship for larval transport. I investigated the coupling of turbulent flows to organism biology using two approaches: a highly idealized 2D random-flight model, and a three-dimensional idealized circulation model (based upon the Regional Ocean Model System) for a coastal domain in which many Lagrangian particles are tracked as models of planktonic larvae.
(III) Ecological forecasting. The ultimate goal of this research is to enable statistical prediction of ecosystem responses to climate oscillations and global climate change in the northeast Pacific basin. This stage of my research is a work in progress. Below I describe approaches I have taken to model the effects of three types of predicted climate change: changes in ENSO frequency, sea level rise, and changes in larval connectivity due to shifts in sea surface temperature & upwelling.
Kinlan* et al. 2004 (Contributed Talk)
WSN 85th Annual Meeting, Rohnert Park, CA, 11-14 Nov 2004

THE METAPOPULATION ECOLOGY OF GIANT KELP IN SOUTHERN CALIFORNIA
Kinlan, B.P.; D.C. Reed; P.T. Raimondi; L. Washburn; B. Gaylord; P.T. Drake

Although the potential importance of the metapopulation concept to marine ecology has long been recognized, detailed examples of metapopulation dynamics in the marine environment are rare.  This is due in part to the large geographic scale at which marine population dynamics often operate, and the difficulty of characterizing rates of migration in the marine environment.   Here, we combine long-term monitoring data, oceanographic modeling, and field experiments to show that the giant kelp, Macrocystis pyrifera, exhibits structured metapopulation dynamics along a continuous 500 km stretch of coastline in southern California, and estimate extinction and colonization rate parameters as a function of oceanographic conditions, patch size, and connectivity.  A 34-year time series of monthly aerial surveys showed that patches in this region underwent frequent extinctions and recolonizations that occurred over time scales ranging from several months to as much as 13 years.  Extinction probabilities were negatively correlated with patch size, and positively correlated with degree of isolation from surrounding patches.  In contrast, recolonization probabilities were positively correlated with patch size and negatively correlated with isolation.  The vast majority of patches remained extinct for less than two years before being recolonized.  Empirical and modeled estimates of spore dispersal resembled a negative power function, with the bulk of spores landing near parent patches and the tails of dispersal extending from tens of meters to several kilometers depending on oceanographic currents and waves.  Results of analyses overlaying modeled estimates of dispersal with inter-patch distances revealed that an average patch may be completely isolated or be connected to up to five neighboring patches, depending on the oceanographic setting and the size, spore output, and spacing of patches.  Implications of the metapopulation framework for the regional population dynamics, persistence, and genetic structure of giant kelp are discussed.

Kinlan* and Siegel, et al. 2004 (Contributed Talk)
ASLO/AGU/TOS Ocean Sciences Meeting, Portland, OR, 26-30 Jan 2004

Marine Larval Dispersion and Prediction in Coastal Fisheries Science
Kinlan, B.P.; Siegel, D.A.; Gaylord, B.; Gaines, S.D.

Many coastal marine organisms are sedentary as adults and are redistributed between generations by oceanic transport of planktonic larvae. We introduce a Lagrangian description of larval transport to assess larval dispersal kernels (settlement probability distributions) for a range of ocean flows and larval settlement pre-competency/competency periods. Paths of individual planktonic larval releases are modeled statistically and, by averaging over many individuals, estimates of the larval dispersal kernel are derived. Typical dispersal scales vary from a few km to $>$400 km. Modeled dispersal kernels are well explained using only a few readily available biological and oceanographic parameters and derived scales agree well with population genetic estimates. Importantly, we note that settlement patterns resulting from larval releases made over short times (days to months) should be comprised of a small number of discrete samples taken from the long-term averaged dispersal kernel. The resulting larval dispersal patterns are quasi-random in both space and time, which has important implications for interpretation of settlement time series and management of coastal fisheries.
Kinlan* and Broitman 2004 (Contributed Talk)
ASLO/AGU/TOS Ocean Sciences Meeting, Portland, OR, 26-30 Jan 2004

A COUPLED SPATIAL PATTERN OF BENTHIC AND PELAGIC ECOSYSTEM STRUCTURE IN COASTAL UPWELLING REGIONS
Kinlan, B.P.; Broitman, B.R.

A growing body of research in the benthic environment has revealed that coastal oceanographic processes largely determine the structure of ecological communities in the nearshore.  Knowledge of the spatial pattern of coupled benthic-pelagic processes is of critical importance to coastal management issues and to our understanding of nearshore ecological systems.  Yet, rigorous characterization of the spatial scales of benthic-pelagic coupling has remained elusive.  We studied patch scales of benthic and pelagic primary productivity over a large region on the west coast of North America, between Baja California del Sur, Mexico (27°N) and Oregon, USA (42°N). Benthic patch scales were determined using the alongshore distribution of shallow subtidal kelp stands from aerial photography.  Nearshore pelagic patch scales were examined through chlorophyll-a concentration from SeaWiFS.  The spatial analysis showed a striking match between the patch scales of kelp stands and chlorophyll-a concentration. Across this large extent of coastline (~3000 km) coastal patch scales are characterized by small-scale spatial cycles of 30-60 km embedded in large 150-250 km regions.  A spatial analysis of meridional anomalies in coastline orientation across the region shows that patch scales in the benthic and pelagic patterns closely follow the spatial pattern of coastline orientation. Our results suggest the existence of nested spatial scales of variation in nearshore ocean climate with strong, predictable effects on benthic and pelagic ecosystems.  The correlated spatial patterns provide strong support for topographic forcing of nearshore oceanography as the dominant process structuring coastal ecosystems in this upwelling region.  Moreover, the existence of well-defined spatial scales and a mechanistic linkage to physical forcing provides a powerful tool for current coastal management initiatives.
Kinlan* and Broitman 2003 (Contributed Talk)
WSN 84th Annual Meeting, Long Beach, CA, 9-11 Nov 2003

A COUPLED SPATIAL PATTERN OF BENTHIC AND PELAGIC ECOSYSTEM STRUCTURE IN COASTAL UPWELLING REGIONS
Kinlan, B.P.; Broitman, B.R.

A growing body of research in the benthic environment has revealed that coastal oceanographic processes largely determine the structure of ecological communities in the nearshore.  Knowledge of the spatial pattern of coupled benthic-pelagic processes is of critical importance to coastal management issues and to our understanding of nearshore ecological systems.  Yet, rigorous characterization of the spatial scales of benthic-pelagic coupling has remained elusive.  We studied patch scales of benthic and pelagic primary productivity over a large region on the west coast of North America, between Baja California del Sur, Mexico (27ºN) and Oregon, USA (42ºN). Benthic patch scales were determined using the alongshore distribution of shallow subtidal kelp stands from aerial photography.  Nearshore pelagic patch scales were examined through chlorophyll-a concentration from SeaWiFS.  The spatial analysis showed a striking match between the patch scales of kelp stands and chlorophyll-a concentration. Across this large extent of coastline (~3000 km) coastal patch scales are characterized by small-scale spatial cycles of 30-60 km embedded in large 150-250 km regions.  A spatial analysis of meridional anomalies in coastline orientation across the region shows that patch scales in the benthic and pelagic patterns closely follow the spatial pattern of coastline orientation. Our results suggest the existence of nested spatial scales of variation in nearshore ocean climate with strong, predictable effects on benthic and pelagic ecosystems.  The correlated spatial patterns provide strong support for topographic forcing of nearshore oceanography as the dominant process structuring coastal ecosystems in this upwelling region.  Moreover, the existence of well-defined spatial scales and a mechanistic linkage to physical forcing provides a powerful tool for current coastal management initiatives.
Kinlan* and Gaines 2003 (Contributed Talk)
Eastern Pacific Ocean Conference (EPOC), Catalina, CA, 27-30 Sept 2003

CONSEQUENCES OF LIFE HISTORY AND LARVAL DURATION FOR THE SCALE OF LARVAL TRANSPORT
B. P. Kinlan and S. D. Gaines

Larval transport underlies the scale and pattern of connectivity in marine benthic communities.  Because dispersal distances of marine organisms are difficult to measure directly, a predictive understanding of marine community connectivity requires knowledge of basic relationships between life history characteristics and the scale of larval transport.  We estimated average dispersal distances for ~100 marine animals and plants from genetic isolation-by-distance relationships, and used this database to examine effects of life history variation on the scale of larval dispersal.  The most important factor influencing dispersal scale was planktonic larval duration.  To a first approximation, dispersal scale estimates were concordant with a null model incorporating only physical processes and time in the plankton.  Variation in dispersal distance among invertebrates with different developmental modes could be accounted for by variation in larval duration.  Spawning mode of fishes, however, had an effect on dispersal scale that could not be explained by larval duration alone: planktonic-spawning fishes dispersed further per day than benthic spawners, suggesting a role of behavior in limiting dispersal of advanced-stage larvae.  Adult movement (whether active or passive) swamped the effect of juvenile dispersal for many organisms with larval durations < 1-3 days.   When spread rates of invasive species were examined as an index of long-distance dispersal potential, we found evidence that rare long-distance dispersal events are critical to the ecology of otherwise short-dispersing benthic organisms.  Our results suggest an interplay between community life history patterns and spatial dynamics, with important implications for the management of multi-species marine assemblages.
Broitman* and Kinlan 2003 (Contributed Talk, presented by co-author)
Eastern Pacific Ocean Conference (EPOC), Catalina, CA, 27-30 Sept 2003

A COUPLED SPATIAL PATTERN OF BENTHIC AND PELAGIC ECOSYSTEM STRUCTURE IN COASTAL UPWELLING REGIONS
Broitman, B.R.; Kinlan, B.P.

A growing body of research in the benthic environment has revealed that coastal oceanographic processes largely determine the structure of ecological communities in the nearshore.  Knowledge of the spatial pattern of coupled benthic-pelagic processes is of critical importance to coastal management issues and to our understanding of nearshore ecological systems.  Yet, rigorous characterization of the spatial scales of benthic-pelagic coupling has remained elusive.  We studied patch scales of benthic and pelagic primary productivity over a large region on the west coast of North America, between Baja California del Sur, Mexico (27ºN) and Oregon, USA (42ºN). Benthic patch scales were determined using the alongshore distribution of shallow subtidal kelp stands from aerial photography.  Nearshore pelagic patch scales were examined through chlorophyll-a concentration from SeaWiFS.  The spatial analysis showed a striking match between the patch scales of kelp stands and chlorophyll-a concentration. Across this large extent of coastline (~3000 km) coastal patch scales are characterized by small-scale spatial cycles of 30-40 km embedded in large 150-200 km regions.  A spatial analysis of meridional anomalies in coastline orientation across the region shows that patch scales in the benthic and pelagic patterns closely follow the spatial pattern of coastline orientation. Our results suggest the existence of nested spatial scales of variation in nearshore ocean climate with strong, predictable effects on benthic and pelagic ecosystems.  The correlated spatial patterns provide strong support for topographic forcing of nearshore oceanography as the dominant process structuring coastal ecosystems in this upwelling region.  Moreover, the existence of well-defined spatial scales and a mechanistic linkage to physical forcing provides a powerful tool for current coastal management initiatives.
Kinlan* 2003 (Contributed Poster)
Eastern Pacific Ocean Conference (EPOC), Catalina, CA, 27-30 Sept 2003

PHYSICAL FORCING OF KELP FOREST COMMUNITY DYNAMICS IN THE NORTHEAST PACIFIC
B. P. Kinlan
 
The giant kelp, Macrocystis pyrifera, provides food and habitat structure for a diverse array of organisms on shallow subtidal reefs (~3-30 m) along the west coast of North America.    Regional and meso-scale physical processes exert a strong influence on the structure and dynamics of kelp forest habitats in this region, generating variation at distinct, nested spatial and temporal scales.  Organisms that live in kelp forests typically disperse, reproduce, grow, and die at different temporal and spatial scales than their biogenic habitat.  As a result, variation in the spatiotemporal dynamics of kelp forest patches may exert important effects on benthic community structure and composition.  I quantified the spatial and temporal dynamics of giant kelp, using a 34-year monthly time series of kelp canopy biomass covering ~2000 km of coastline (10º of latitude) and digital maps of annual maximum canopy cover.  Canopy biomass varied at well-defined dominant periods of 12-18 months, 3-7 y, 12-14 y and >20 y, and spatial scales ranging from mesoscale (20-50 km, 100-150 km) to regional (300-500 km).   Much of the spatial variability was attributable to variation in coastal upwelling and sediment deposition forced by coastline orientation and geomorphology.  Temporal variation was attributable to basin-scale climate forcing in the ENSO and PDO bands, with regional variation in coherence with and response to these two sources of variability.  Changes in biomass were associated with shifts in the spatial structure of the kelp habitat, as revealed by high-resolution digital canopy maps.   Long-term monitoring data from areas with contrasting kelp forest structure and dynamics suggest important impacts of meso- and large-scale physical forcing on kelp forest community structure.
Kinlan* 2003 (Contributed talk)
6th International Temperate Reef Symposium (6ITRS), Christchurch, New Zealand, 12-17 January 2003

SPATIAL AND TEMPORAL VARIATION OF KELP FOREST HABITAT STRUCTURE IN THE NORTHEAST PACIFIC
Kinlan, B.P.

Habitat-forming species such as kelps and corals often differ from associated benthic species in resource requirements, sources of disturbance, and dispersal ability.  These differences in life history can cause habitat to vary at spatial and temporal scales that differ from the “optimal” scale that would promote maximum abundance of any particular associate species.  As a result, the spatiotemporal dynamics of habitat can exert important effects on benthic community structure and composition.  To quantify the spatial and temporal dynamics of giant kelp (Macrocystis pyrifera), a key habitat-former in the NE Pacific, I analyzed a 32-year monthly time series of estimated canopy biomass covering ~1500 km of coastline (7º of latitude) and digital maps of annual maximum canopy cover.  Canopy biomass varied at dominant periods of 12 months, 3-7 y, 12-14 y and >20 y, and spatial scales ranging from mesoscale (~50 km) to regional (~250 km).  Digital canopy maps revealed that changes in biomass were associated with shifts in the spatial structure of the kelp habitat.  Patch size and inter-patch distance distributions varied with canopy biomass in a non-linear, but predictable relationship at local (0.1-10 km), mesoscale (10-100 km), and regional (100-1000 km) scales.   The spatiotemporal dynamics of kelp habitat were also related to geographic region (Central vs. Southern vs. Baja California), and long-term mean size of kelp beds.  Larger kelp beds exhibited less short-term variation and greater patch coherence than smaller beds.  Long-term monitoring data suggest these differences in habitat structure have important consequences for community dynamics.
Kinlan* and Gaines 2002 (Contributed Talk)
Larval Ecology Meetings, Vigo, Spain, 15-20 Sept 2002

CONSEQUENCES OF LIFE HISTORY AND LARVAL DURATION FOR THE SCALE OF LARVAL TRANSPORT
Kinlan, B.P.; Gaines, S.D.

Larval transport underlies the scale and pattern of connectivity in marine benthic communities.  Because dispersal distances of marine organisms are difficult to measure directly, a predictive understanding of marine community connectivity requires knowledge of basic relationships between life history characteristics and the scale of larval transport.  We estimated average dispersal distances for >100 marine animals and plants from genetic isolation-by-distance relationships, and used this database to examine effects of life history variation on scale of larval dispersal.  The most important factor influencing dispersal scale was planktonic larval duration.  Dispersal scale estimates were generally concordant with a null model incorporating only physical processes and time in the plankton.  Larval duration in turn explained variation in dispersal distance among invertebrates with different developmental modes.  Spawning mode of fishes, however, had an effect on dispersal scale that could not be explained by larval duration alone: planktonic-spawning fishes dispersed further per day than benthic spawners, suggesting a role of behavior in limiting dispersal of advanced-stage larvae.  For organisms with planktonic durations <1 day (e.g. direct-developing invertebrates and many macroalgae), genetic dispersal scale was independent of time in the plankton and strongly dependent on adult dispersal potential. While average dispersal scale is a critical determinant of population connectivity, landscape processes such as invasion and recolonization may be more dependent on the tails of a dispersal distribution.  To examine the relationship between average dispersal distance and spread rates, we compared invasion rates of marine organisms with short vs. long average dispersal.  For long-dispersers, invasion rate was comparable to average dispersal distance.  For short dispersers, however, invasion rate was highly variable and frequently several orders of magnitude larger than average dispersal distance.  Thus organisms with very different life histories and scales of average larval transport may invade marine landscapes at similar speeds. Our results suggest an interplay between community life history patterns and spatial dynamics, with important implications for the management of multi-species marine assemblages.
Kinlan* and Gaines 2002 (Contributed Talk)
Ecological Society of America Annual Meeting, Tucson, AZ, 4-9 August 2002

A COMPARATIVE ANALYSIS OF DISPERSAL SCALE IN MARINE AND TERRESTRIAL SYSTEMS
Kinlan, B.P.; Gaines, S.D.

For organisms with limited adult movement (e.g. terrestrial plants, benthic marine organisms), propagule dispersal determines the spatial scale at which populations interact with their environment, spread across landscapes, and adapt.   Differences in dispersal scale among co-occuring taxonomic or functional groups may have important implications for community structure and dynamics.  On evolutionary time scales, differences in dispersal scale may be shaped by the distribution of resources in space and time, costs and availability of dispersal mechanisms, and indirect consequences of life history trade-offs (e.g. propagule size vs. fecundity) or phylogenetic history (e.g. complex life cycles involving a larval stage).    Because marine and terrestrial environments differ in resource distribution, available dispersal mechanisms, and life history/phylogeny of sessile species, dispersal patterns may vary substantially across these systems.  However, quantitative comparisons of marine vs. terrestrial dispersal scales have been hindered by the difficulty of tracking propagules in the ocean.  We use evidence from the increase in genetic differentiation with geographic distance to derive estimates of average dispersal scale for 104 marine macroalgae, invertebrates, and fish, and compare these data to existing dispersal estimates for 1) terrestrial plants and 2) a functional group (herbivorous invertebrates) that is sedentary in the ocean, but vagile on land.  Marine algae and terrestrial plants disperse over similar scales, but sessile invertebrates and demersal fish can disperse at least 2 orders of magnitude further than the longest-dispersing terrestrial plant.  Genetic estimates suggest that herbivores typically disperse much further than their plant resources both on land and in the sea, though the ecological consequences of this disparity may vary across systems.  Differences in dispersal scale may underlie important differences in the dynamics of marine and terrestrial communities.
Kinlan* and Gaines 2002 (Contributed Poster)
43rd Annual Symposium of the British Ecological Society, "Macroecology: Concepts and Consequences", Birmingham, England, 17-19 April 2002

A COMPARATIVE ANALYSIS OF DISPERSAL SCALE IN MARINE AND TERRESTRIAL SYSTEMS
Kinlan, B.P.; Gaines, S.D.

For organisms with limited adult movement (e.g. terrestrial plants, benthic marine organisms), propagule dispersal determines the spatial scale at which populations interact with their environment, spread across landscapes, and adapt. Differences in dispersal scale among co-occuring
taxonomic or functional groups may have important implications for community structure and dynamics. We use evidence from the increase in genetic differentiation with geographic distance to derive estimates of average dispersal scale for 104 marine macroalgae, invertebrates, and fish,
and compare these data to existing dispersal estimates for 1) terrestrial plants and 2) a functional group (herbivorous invertebrates) that is sedentary in the ocean, but vagile on land. Average dispersal scales of marine algae and terrestrial plants are similar, but sessile invertebrates and
demersal fish can disperse at least 2 orders of magnitude further than the longest-dispersing terrestrial plant. Genetic estimates suggest that herbivores typically disperse further than their plant resources both on land and in the sea, though the behaviorally-mediated dispersal of terrestrial
herbivores may have very different consequences from the more obligate dispersal of marine propagules. Comparison of rates of invasive species spread in marine and terrestrial systems suggests that long-distance dispersal is more common among marine macroalgae than terrestrial
plants, despite the similarity in mean dispersal scales of these groups. Systematic differences in the scale and pattern of dispersal may underlie differences in community and landscape dynamics between land and sea.
Thornber* and Kinlan, et al. 2002 (Contributed Talk, presented by co-author)

Reproductive Ecology of the Invasive Japanese Kelp Undaria pinnatifida along the California Coast
Carol Thornber, Brian Kinlan, Michael Graham, Jay Stachowicz

    The exotic kelp Undaria pinnatifida has recently become established at several locations in California coastal waters.  Undaria is native to Japan, but in recent decades it has spread via  purposeful and accidental introductions to numerous other coastal areas around the globe.  The population biology of Undaria varies greatly among geographic locations.  For example, although Undaria is an annual kelp, it may either have clearly defined seasonal cycles of growth and senescence (Japan), or overlapping generations in which individuals are present throughout the year (New Zealand).  Some invasions of Undaria (e.g. New Zealand) have resulted in significant changes to the native flora and fauna; but it is unknown what effects Undaria may haveon California’s marine communities.  This study represents the first attempt to document the reproductive ecology and population biology of Undaria in California.  We tracked the timing and magnitude of Undaria recruitment, growth, and subsequent reproductive onset in the Santa Barbara, California harbor following the discovery of a dense, reproductive population there in April 2001.  From July to September 2001, there was limited recruitment of new Undaria sporophytes.  Although these individuals did mature and reproduce, they were much smaller than the spring 2001 cohort.  A much larger recruitment pulse was observed during January-February 2002, followed by rapid growth of individuals.  This recruitment pulse is correlated with a drop in ocean temperature, and ongoing laboratory culture experiments are exploring the effects of different water temperatures on the growth of microscopic stages of Undaria.  This research  provides insight into the potential for Undaria spread and growth into previously unoccupied habitats along the California coast, as well as information for the timing of subsequent eradication efforts.
Kinlan* 2001 (Contributed Talk)

PERSISTING IN AN UNPREDICTABLE WORLD: ARRESTED DEVELOPMENT OF MICROSCOPIC SPOROPHYTES AS A MECHANISM FOR DELAYED RECRUITMENT IN MACROCYSTIS.
Kinlan, Brian P.

To persist in patchy and variable environments, kelps employ life history strategies that extend recruitment in space and time.  Dispersal is crucial to colonization and patch dynamics, and can allow persistence through short- and long-term episodes of unfavorable conditions (e.g. winter storms, grazer outbreaks, ENSO).  Yet, the scale of environmental fluctuations often exceeds the scale of propagule dispersal.  As an alternative, many kelps survive predictable environmental fluctuations by delayed recruitment of microscopic stages regulated by endogenous clocks.  Such temporally rigid strategies, however, are not responsive to unpredictable changes in environmental conditions.  I experimentally evaluated the potential for temporary reduction in key limiting resources (light, nutrients) to directly delay recruitment of giant kelp (Macrocystis pyrifera) microscopic life history stages.  Laboratory cultures of gametophytes and embryonic sporophytes were subjected to limiting conditions of light and nutrients for one month, and then exposed to non-limiting conditions for 10 days.  Gametophytes in all treatments failed to recruit to sporophytes after 2 weeks, suggesting they are not a source of delayed recruitment in giant kelp.  Sporophytes in light-stressed treatments, however, survived and grew significantly slower than non-light-stressed controls.  When stimulated with light, light-stressed sporophytes grew an average of 5-10 times faster than unstimulated controls; speed of growth following light stimulation was dependent upon nutrient availability.  These results suggest that limiting resources can delay recruitment of embryonic giant kelp sporophytes for at least one month.  Temporally rigid mechanisms of delayed recruitment may be of limited utility to a continuously- reproducing perennial such as giant kelp; instead, flexible timing of recruitment from microscopic stages may enhance survival of episodic environmental fluctuations.