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**H. RE¸ ˙T AKÇAKAYA* AND JONATHAN L. ATWOOD† SI
**

*Applied Biomathematics, 100 North Country Road, Setauket, NY 11733, U.S.A., email ramas@gramercy.ios.com †Manomet Observatory for Conservation Sciences, P.O. Box 1770, Manomet, MA 02345, U.S.A.

Abstract: We present an analysis of the metapopulation dynamics of the federally threatened coastal California Gnatcatcher (Polioptila c. californica) for an approximately 850 km2 region of Orange County, California. We developed and validated a habitat suitability model for this species using data on topography, vegetation, and locations of gnatcatcher pair observations. Using this habitat model, we calculated the spatial structure of the metapopulation, including size and location of habitat patches and the distances among them. We used data based on field studies to estimate parameters such as survival, fecundity, dispersal, and catastrophes, and combined these parameters with the spatial structure to build a stage-structured, stochastic, spatiallyexplicit metapopulation model. The model predicted a fast decline and high risk of population extinction with most combinations of parameters. Results were most sensitive to density-dependent effects, the probability of weather-related catastrophes, adult survival, and adult fecundity. Based on data used in the model, the greatest difference in results was given when the simulation’s time horizon was only a few decades, suggesting that modeling based on longer or shorter time horizons may underestimate the effects of alternative management actions. Modelo de Metapoblación Basado en el Hábitat de Polioptila c. Californica Resumen: Presentamos un análisis de la dinámica metapoblacional de Polioptila c. californica, especie amenazada de extinción de las costas de California, para una región de aproximadamente 850 km2 del condado de Orange, California. Describimos y validamos un modelo de ajuste para esta especie utilizando datos de topografia, vegetación y sitios de observación de la especia. Utilizando este modelo de hábitat, estimamos la estructura espacial de la metapoblación incluyendo tamaño y localización de parches de hábitat así como las distancias entre ellos. Utilizamos datos basados en estudios de campo para estimar parámetros tales como la supervivencia, fecundidad, dispersión y catastrofes, también se combinaron estos parámetros con la estructura espacial con la finalidad de construir un modelo metapoblacional estructurado por pasos, estocástico y espaciálmente explícito. El modelo predijo una declinación rápida y un alto riego de extinción de la población con la combinación de la mayoría de los parámetros. Los resultados fueron mas sensitivos a efectos densodependientes, la probabilidad de catastrofes ocasionadas por el clima, la supervivencia de adultos y la fecundidad de los adultos. Basados en los datos utilizados en este modelo, las diferencias mayores en los resultados se produjeron cuando el horizonte de tiempo de las simulaciones fue de solo unas cuantas décadas, sugiriendo que el modelado basado en horizontes de tiempo largos o cortos pueden desestimar los efectos de acciones alternativas de manejo.

Introduction

The California Gnatcatcher’s northernmost subspecies (Polioptila c. californica) has declined due to extensive

Paper submitted May 15, 1996; revised manuscript accepted October 10, 1996.

agricultural and urban development of coastal sage scrub, the species’ primary habitat type in southern California and northwestern Baja California (Atwood 1993). Listed in 1993 as threatened under the U.S. Endangered Species Act (ESA), protection of the gnatcatcher and its habitat has become a major focus in the inaugural application of the State of California’s Natural Community Conserva-

422

Conservation Biology, Pages 422–434 Volume 11, No. 2, April 1997

and shape of habitat patches in which [sub]populations of the metapopulation exist).e. for reviews see Kingston 1995 and Boyce 1996).e. 1995. and forest categories). Based on these maps. such as carrying capacities. aimed at delineating core tracts of essential natural habitat while allowing economic development in areas of lower biological significance. We used the population viability analysis program RAMAS/GIS.e. vegetation. we used ranges such as 20% to 100% for parameters based on crude estimates and ranges such as 5% to 10% for parameters based on more reliable data. 0 for north). and (3) to analyze the sensitivity of the viability of this species to uncertainties in model parameters. estimating lower and upper bounds.. We analyzed the viability of a California Gnatcatcher metapopulation in central and coastal Orange County. Habitat and Census Data The habitat variables and census data that formed the basis of our analysis were provided by the GIAS lab of Southern California Edison in the form of digital raster (grid) maps with a resolution of 100 m. the values for the slope map (SLP) are in percentages (i. These maps described topography. size. we combined the original 20 subassociations of coastal sage scrub into a single. Vegetation maps. We used these ranges to estimate upper and lower bounds on the estimated viability of the species.g. 0 for flat areas. Conservation Biology Volume 11. 90 for east and west.. The NCCP plans that were recently prepared for central and coastal Orange County did not incorporate these results.. woodland. (1995. Because participation in the NCCP program was formally recognized as an alternative to the requirements for incidental “take” permits as authorized under Section 10(a) of the ESA. with values ranging from 0 m2 (no coastal sage scrub) to 10. 2. stage-structured. with each grid cell being represented by the value at the cell’s central point. April 1997 . in part because our analysis was not completed until after the planning process was well underway. each cell in the coastal sage scrub (CSS) data layer was assigned the value of a 9-cell (3 3) moving average of these area estimates. The values for elevation map (named ELV) are elevation in meters above sea level. slope. and grasslands. we created three data layers describing distance of each cell to (1) the nearest cell with at least 10% cover of trees or other woody vegetation (DTR). The total amount of coastal sage scrub was calculated for each map cell. which is designed to link landscape data from a geographic information system with a metapopulation model (Akçakaya 1995. to identify the location. ranging from 0 to 180. We used data on the current distribution of the species’ habitat to find the spatial structure of the metapopulation (i. in addition to a “medium” estimate). for another application of the program see Akçakaya et al. Because of uncertainties regarding accuracy of the raw vegetation data. stochastic model of the California Gnatcatcher metapopulation based on habitat suitability maps we developed and on demographic data from Atwood et al. and the values for aspect (ASP) are in units of degrees from north (e. and gnatcatcher distribution in approximately 853 km2 of central and coastal Orange County. 100 for a 45 incline).000 m2 (full cover). woody vegetation (combining chaparral. This voluntary. unpublished manuscript). regional land-use planning process. We also used the Jones and Stokes (1993) vegetation data to create maps of wetland vegetation (including riparian habitat).Akçakaya & Atwood California Gnatcatcher Metapopulation 423 tion Planning (NCCP) program (Atwood & Noss 1994). were converted from ARC/Info coverages to grid (raster) format. Our aims were (1) to develop a habitat suitability model for California Gnatcatcher. generalized map. (2) the nearest cell of grassland (DGR). most NCCP efforts in southern California have revolved around planning decisions intended to conserve gnatcatchers and their habitat. We accounted for uncertainties arising from measurement errors and lack of data by making three estimates of each parameter (i. (2) to demonstrate how this model can be linked to a metapopulation model for viability analysis. Because field experience suggested that the Jones and Stokes (1993) classification of various coastal sage scrub subassociations might include a relatively high degree of subjective interpretation and because a preliminary analysis failed to detect any consistent patterns regarding presence or absence of gnatcatchers in different subassociations. as well as the rate of dispersal between patches and the degree of similarity of environmental fluctuations that different populations experience. has been described as a potential model for resolving conflicts between conservation and economic concerns (Reid & Murphy 1995). In addition to spatial structure. and (3) the nearest The Model We developed a spatially-explicit. 180 for south. the amount of year-to-year variability in vital rates. No. it is our hope that these results will contribute to future NCCP planning efforts elsewhere in southern California. as well as provide a basis for long-term habitat management within the NCCP coastal sage scrub reserve system in central and coastal Orange County. initial abundances and vital rates of each population. originally prepared by Jones and Stokes (1993) from color aerial photographs. In most cases the estimation of a range is quite arbitrary. we incorporated parameters related to demography. and aspect. Nonetheless.. Topographical data layers included elevation.

or residential development. which was then used to calculate an index of habitat suitability for each cell.. Carrying Capacities and Initial Abundances The program allows the calculation of carrying capacities (K) based on the total habitat value of each patch (i. (2) Absent localities were not placed within 300 m of other randomly-placed points because too much clustering would lead to redundancy and not add information to the analysis. even if they were rendered not significant with the addition of an interaction term. No. We excluded patches with K 5 because only larger populations would have at least one pair of adults at stable stage distribution.. (3) Absent localities were not placed in cells dominated by agricultural fields.. within the neighborhood distance of each other) that have HS values higher than or equal to the threshold HS and labels them as patches. We used a stepwise approach with a significance level of p 0. We selected these “absent” localities randomly. the program finds clusters or groups of nearby cells (i. We used 1/4. only those cells that had a habitat value of 0.5 (i..69 ( 0. which were estimated to be 4. the sum of habitat values of all cells that are included in a patch). We estimated this proportion based on the ratio of the total number of observed pair locations (for both 1992 and 1994. We used total habitat value instead of the total area as the argument because the latter includes areas with low habitat value. All of these habitat types are clearly not suitable gnatcatcher habitat. coastal sage scrub dominated by Artemisia californica generally occurs in closer proximity to grassland areas. we randomly added 273 (50% of the number of cells where pairs were observed) points intended to represent locations unsuitable for gnatcatchers. For example. which we used for validation purposes. (1) “Absent” localities were not placed within 300 m of cells where pairs or single individuals were observed. In surveys of California gnatcatchers (conducted in Orange County in 1991–1992 and 1994). The sum of habitat values in a patch is also related to the area of the patch. Habitat Suitability Map and Patch Structure We used logistic regression (SAS Institute 1990) to calculate a habitat suitability function.69 ha on average in Orange County (Bontrager 1991). 50%) as the threshold HS. unpublished manuscript). We believe that variables describing distance to woody vegetation or grassland provide a more objective characterization of coastal sage scrub subassociations than do the classifications used by Jones and Stokes (1993). rural development. for example. it is likely that cells surrounding an observation point would support suitable habitat even though no observations were obtained in these cells. water. Cells where one or more pairs were recorded were assigned a value of 1 (present) (n 547). After all variables were tested. but excluding the 1994 locations within 150 m of 1992 locations) (503) to the number of pairs (629) predicted by the above calculation of the total carrying capacity.g. but with the following constraints. and their inclusion would not add information to the analysis.55 for the threshold HS parameter and a range of 250 m to 350 m for the neighborhood distance parameter (Table 1).05 for adding and removing variables. 325 pairs and 80 single individuals were found in 1991–1992.5 or above were considered when habitat patches were analyzed. the foraging distance of the species. Given these two parameters.. which corresponds to the assumption that any two suitable locations within about 300 m of each other are in the same habitat patch. We used 0. In a separate data layer. We truncated distance values greater than 2 km for these data layers. Threshold HS is the minimum habitat suitability (HS) value below which the habitat is not suitable for reproduction and/or survival. Because the species’ territory size may be larger than one 1 ha (Atwood et al.424 California Gnatcatcher Metapopulation Akçakaya & Atwood cell of wetland or riparian vegetation (DWT). because distributional surveys did not certainly document gnatcatcher absence within the study area. We specified the initial number of individuals in each patch as a fixed proportion of the carrying capacity of that patch. We used a neighborhood distance of 3 cells. we started adding interaction terms (and removing those that became non-significant).213) as a scaling constant in calculating K of each patch by multiplying it with the total habitat value in each patch. and 238 pairs and 80 single individuals were located in 1994. we consider it likely that many observations of single birds occurred in suitable breeding habitat and that areas near these points should not be used to characterize unsuitable habitat. we also assigned a value of 1 (present) to all cells (n 129) where single birds were observed.e.e. April 1997 . We used 20% of this parameter as the upper and lower limits. The link between the habitat map and the metapopulation model was characterized by two parameters. Logistic regression is indicated in cases where the response (dependent) variable is binary (e. We estimated carrying capacities based on territory sizes. or 80% Conservation Biology Volume 11. These parameters are used by a patch-recognition algorithm that delineates patches on the habitat map. so that territories in areas with low habitat value are assumed to be larger. 0 or 1). Finally. 2. Similarly.e. each patch had a population of 503/629. Neighborhood distance is used to identify nearby cells that belong to the same patch and may represent. whereas subassociations dominated by Salvia mellifera often occurs near chaparral or oak woodland (and distant from grassland). Thus. but weighted by the amount of habitat in each cell. We analyzed the sensitivity of extinction risk to these two parameters by using a range of 0. In this phase we retained all previously-added single variables.45 to 0.

a number for re-sighting is more reasonable than. b S represents the standard deviations given in Table 2.50 300 0.4975 0. as well as in coastal Orange County (Erickson & Pluff. and M is maternity or fertility (number of fledglings per breeder). c See text. Mock. This may be a result of the exceptionally wet and cold winter of 1994–1995.28 4 120 This parameter is multiplied with the total habitat suitability value of the patch to calculate carrying capacity and initial abundance in the patch. Sj is survival rate of juveniles. In parameterizing this stage-structured model.5376 0. and for the upper limit we used 100% (assuming some of the singles represent pairs). or both of the breeders are nesting for the first time (i. were juveniles last year).. To calculate upper and lower bound on vital rates. Parameter estimate Parameter Habitat-demography link Threshold HS Neighborhood distance (m) Carrying capacity (K) multiplier a Initial abundance (as % of K) Mean dispersal distance (km) Stage matrix Juvenile survival rate (Sj ) Juvenile fecundity (Fj) Adult survival rate (Sa ) Adult fecundity (Fa ) Density dependence Density dependence type Allee effects (local threshold as % of K) Density-dependent dispersal Stochasticity Standard deviations b Correlation of fluctuations (b) Catastrophe (weather) probability Number of fires Metapopulation extinction threshold a Low Medium High 0. Adult fecundity (Fa) is equal to Sa · M. say. (3) all adults breed (so that the proportion of last year’s adults who are breeders this year is simply the survival rate of adults). unpublished manuscript. (2) the population is censused immediately after each breeding season (a post-reproductive census. Stage Structure We modeled the dynamics within each patch with a stage-structured. and Sj based on data from the California Gnatcatcher population on the Palos Verdes peninsula (Atwood et al. say. Low.4650 0. For this reason.. we assumed the populations have a stable distribution of individuals to stages at the start of the simulation. and high estimates of parameters used in the model of the California Gnatcatcher metapopulation.55 350 0. we assumed a measurement error of 1 individual in the recovery (re-sighting) of banded individuals. Bontrager et al.5874 0. The data for 1994–1995 suggests a sharp decline. If the census is actually made. 10% of the vital rate itself because the latter does not account for differences in sam- Conservation Biology Volume 11. the survival rates for 1994–1995 would be an exception rather than the rule. page 426. which is what was observed on the Palos Verdes Peninsula (Atwood et al. We estimated PJB. The two elements in the first row of the matrix are fecundities. PJB is proportion of last year’s juveniles that are breeders this year. then the re-sighting might be an overestimate (if a bird actually dies before the “correct” time). unpublished manuscript).14 2 60 1. medium.5 0.5 son (a “birth-pulse” population. The rationale for these limits is based on the assumption of birth-pulse population. Chambers Group and LSA Associates.e. Caswell 1989). unpublished manuscript).170 52 2. stochastic matrix model with two stages (juveniles and adults).9876 ceiling c 0 none 2 weak contest c 4 strong 0. 1994–1995.256 100 3. of its carrying capacity at the beginning of the simulation.2 S 100 0. One hypothesis is that gnatcatchers..Akçakaya & Atwood California Gnatcatcher Metapopulation 425 Table 1. Caswell 1989). 2.7975 0. April 1997 . No. and (5) the stage matrix is the same in all populations. Sa. because of their small body size and high metabolic rates. If this is correct.. for details.45 250 0.4901 0. The parameters are estimated assuming that the census is made exactly at the same time each year.g. (4) the maternity rate (number of fledglings per breeder) is the same whether none.0 0. and juvenile fecundity (Fj) is equal to PJB · M. The data set represented three transitions: 1993–1994.07 1 30 S 30 0.3441 0. With these assumptions.5300 0. For the lower limit we used 52% (based on only the 1992 pairs). There were no comparable data on survival from Orange County.3275 0. 1995).8 S 10 0.8899 0. At a practical level. and M based on data from Orange County (Woehler. 0.3606 0. and 1995–1996. the stage matrix is PJB ⋅ M S a ⋅ M Sj Sa . unpublished manuscript). one.213 80 3. For all simulations. unpublished manuscript. we assumed (1) all reproduction in the population takes place in a relatively short breeding sea- where Sa is survival rate of adults. we used the average of 1993–1994 and 1995–1996 transitions to estimate the average stage matrix and the 1994–1995 data as a basis for modeling catastrophes (Table 2). have higher mortality under such weather conditions (e. one day earlier than this exact time.. unpublished data).

This model allows the populations to fluctuate independent of the population size (N ). We used the range 0. The number 1 (instead of 2) is used as a minimum. (1995. such as potential conversion of coastal sage scrub to grassland (Anderson 1991).1180 0.8899 SD for (93. No. Based on this we assumed two fires. unpublished manuscript).9b 0. to estimate because it is not clear which aspect of the weather makes the crucial difference. and adult fecundity (Fa Sa · M).. we focused on the minimum temperatures in October through December. We also modeled density dependence with a contest model (which uses the Beverton-Holt equation) with the assumption that Sj c PJB c Fj Sa c Fa 1. such as those that may have characterized the winter of 1994–1995.1558c 0.2986 0. Meade Consulting. In addition. thus the required standard deviation is one among years. In this model. (1995) c Atwood et al. 5 in NCCP Habitat Conservation Plan prepared by R. Density Dependence and Allee Effects For most simulations we assumed a ceiling-type density dependence model for each population and used the carrying capacities calculated based on habitat data as population ceilings. proportion juveniles that become breeders (PJB).7766 0.14.2937 0. Environmental and Demographic Stochasticity We modeled environmental stochasticity by sampling the set of vital rates used to project the dynamics of each population from random (lognormal) distributions with means taken from the mean stage matrix and standard deviations taken from a “standard deviations matrix. Two fires have occurred within the study area since 1948 (Fig.0225 0. Keeley 1982). above). the vital rates are again sampled from the average stage matrix (estimated based on the average of 1993–1994 and 1995–1996 transitions) with the standard deviations given above.4375 1. Stage matrix parameters: juvenile survival rate (Sj). frequency of average of the three monthly minimum temperatures less than or equal to that in 1994 was about 14% in the last 98 years.95) 0. corresponding to about 48% of the initial abunTable 2.8840 2. we believe they capture most of its short and mediumterm effects on gnatcatcher populations.7 0. Another type of catastrophe with direct impact on gnatcatcher populations may be extreme weather conditions. 2.2102 0. and in 1993 one occurred in the area identified as the largest patch.5376 0. The population then remains at this level until a population fluctuation takes it below the ceiling.28 to evaluate the sensitivity of results to this assumption. until the population reaches the ceiling.07–0. so we did not include this transition in estimating the standard deviations.95) 0.5200 0.0873 0. We analyzed the sensitivity of results to fire by running simulations with only one fire (in the largest patch). Table 2). The resulting ranges are listed in Table 1. J. In 1982 a fire was located in the area identified here as the second largest patch.1757 0. We used the 1994–1995 transition to model catastrophes. Catastrophes One type of catastrophe that may affect species living in coastal sage scrub is fire. unpublished manuscript) b a Conservation Biology Volume 11. We assumed that the effect of such a catastrophe is a sharp decline in vital rates beyond the normal year-toyear fluctuations (see Environmental and Demographic Stochasticity. and with two additional fires affecting the largest and second largest patches in years 25 and 14.3a 0. After a catastrophe year. we used the demographic parameters estimated from 1994–1995 to model such catastrophes.4314c 0. April 1997 . if not impossible. but even this gave quite large uncertainties at the risk level. Inc.2568 0.426 California Gnatcatcher Metapopulation Akçakaya & Atwood ple size for different vital rates.1892c 0. The frequency of such extreme population fluctuations caused by weather conditions is difficult. et al. Year 1993 1994 1995 M a dance in the modeled population occupying the largest patch. respectively. Thus we chose the annual catastrophe probability of 0.4750c 0. according to the stage matrix and the standard deviations matrix.2390 0.0909c 0. We assumed that each fire will decrease K by 48% and K will increase to its original level in 10 years in each case. one affecting the second largest patch in year 34 and the other affecting the largest patch in year 45 in the 50-year simulations of our model. A statistical estimation is possible only after several years of vital rate estimates. We incorporated demographic stochasticity by sampling number of survivors from a binomial distribution and number of offspring from a Poisson distribution (Akçakaya 1991). Although these assumptions ignore long-term affects of fires. In the absence of such information. 1995). especially in areas where the habitat occurs in close proximity to human population centers (Westman 1982. juvenile fecundity (Fj PJB · M). In the two weather stations closest to the study area (Tustin and Newport).3441 0.” The sampling was done at each time step (in this case each year).0059 Woehler (unpublished manuscript) Average of M from Woehler (unpublished manuscript) and Bontrager. We estimated these standard deviations based on the variability of survival rates and fecundities from the two transitions (1993–1994 and 1995–1996. adult survival rate (Sa).8958 Average for (93. we incorporated demographic stochasticity in dispersal. The 1993 fire destroyed habitat occupied by approximately 127 gnatcatcher pairs (Atwood et al.4975 0.4118c 0. Coastal sage scrub is frequently subject to fire.

distance-dependent spatial autocorrelation among rainfall and used this relationship between rainfall correlation and distance as a basis for setting correlation among population dynamics. No. and a and b are model parameters. (1994) found strong. In the current model we incorporated Allee effects by specifying a local extinction threshold for each population. In our model dispersal refers to the movement of birds among habitat patches. We assumed adults have negligible dispersal among populations (they can disperse within the same patch) and used dispersal parameters discussed above only for the juvenile stage. d is the distance (km) between the centers of these two populations.0 and 3. the above equation was modified to reflect a maximum dispersal distance of 15 km. By considering a population to be extinct once it reaches or falls below its threshold. 2%. Allee effects. In addition. We used b values of 100 (high correlation). we used the fitted function to model minimum dispersal. The relatively small study area (with the maximum distance between any two patches about 40 km) and the apparent dependence of gnatcatcher biology to weather conditions suggests that weather patterns such those seen in southern California (affecting substantial areas) impose a correlation structure on the metapopulation dynamics. We set the local thresholds at 0%. who report the number of juveniles that dispersed different distances.1). 30 (medium correlation). d is the distance (km). In our model M is the dispersal rate. The parameter b is the average dispersal distance. LaHaye et al. We divided the number of dispersing juvenile birds in each distance class by the total sample size (32) to obtain proportion dispersing. which may cause a reduction in vital rates when population size becomes very small. The function is C exp( d/b). 60.Akçakaya & Atwood California Gnatcatcher Metapopulation 427 each population grows with a rate of 15% when N is low and is stable with 0% growth when N K. The function gives the correlation between the vital rates of two populations separated by the indicated distance. In addition. The patch then remains unoccupied. Correlation-distance functions [y exp( x / b)] used in the model. and 4% of the carrying capacity of the patch. (un- published manuscript). the abundance in the population. are not well-studied for this species. We modeled density dependence in dispersal for each population such that the dispersal rate was directly proportional to population size. 2%. and set the average dispersal distance to 3. We incorporated demographic stochasticity in dispersal among populations by sampling the number of dispersers from a binomial distribution with sample size equal to the number of juveniles in the source population and probability equal to the dispersal rate based on distance.4 and b 2. We used the mid-point of each distance class (in km) as the independent variable (d) in the exponential model M a exp( d/b). where C is the coefficient of correlation between the vital rates of two populations. the model assumes the population to be extinct by setting the abundance to zero.5 km for medium and high dispersal. and used this as the dependent variable. April 1997 . and dispersal rate (proportion dispersing from target population to source population) may depend on the distance between source and target populations. The asymmetry of this measure of distance allows for more realistic modeling of dispersal between a large and a small patch. We modeled two levels of Figure 1. Because of the relative isolation and small size of the Palos Verdes peninsula where these data were collected. we specified a metapopulation extinction threshold of 30. The distance metric we used in this study is one between the center of the source population to the edge of the target population. Once any population falls to or below its local threshold. 2. In a study of the southern California metapopulation of the Spotted Owl. and 10 (low correlation). In our model we used three different correlation-distance functions to set the correlation of vital rates among populations (Fig. Conservation Biology Volume 11. unless it is colonized by dispersers from another patch. the model need not accurately predict the dynamics of the populations at these low abundance levels.5 km (Fig. and b is a parameter that describes how fast the correlation declines with increased distance between populations. and 4% of the total initial metapopulation size) and calculated the viability results in terms of falling below this threshold. and whether the bird is a juvenile or an adult. The model was fitted with a 0. Metapopulation Dynamics: Correlation-distance Function and Dispersal Parameters related to dynamics at the metapopulation level include the interdependence of environmental fluctuations among populations and patterns of dispersal. 2). or 120 individuals (about 1%. We estimated the distance dependence of dispersal based on data for 1993 and 1994 from Atwood et al.

the proportion dispersing is lower in proportion to the ratio of N/K. as well as the number of dispersers were always integer numbers. Slope. K is the carrying capacity. At each time step the number of juveniles and adults at each population were projected using a set of vital rates drawn from a random (lognormal) distribution. whereas the elements of the stage matrix within a population were perfectly correlated. and each replication projected the abundance of the each population for 50 time steps (years).14 to 1. We used three measures to express the predicted viability of the metapopulation: (1) median time to fall below the metapopulation extinction threshold. used to model density-dependent dispersal for each population. 3).00. The frequency distribution of the habitat suitability values of locations with gnatcatcher observations is given in Fig.5).86. we ran three simulations for each parameter.30. When dispersal is density-independent (“none” on the graph). The goodness of fit statistics given at the end of the table show that the regression is highly significant.428 California Gnatcatcher Metapopulation Akçakaya & Atwood density dependence in dispersal. M is the total rate of dispersal from the population to all other populations. The curve is the function y 0. with an average of 0. The sampled stage matrices of different populations were correlated according to one of three sets of correlations. and the probability column shows that regression coefficients are also statistically significant. Conservation Biology Volume 11. the dispersal rate is M regardless of abundance. given by the dispersal distance function. The mean values of these vital rates (survival and fecundity) were taken from the stage matrix.4 · exp( x/2. and the standard deviation of vital rates are those given by the standard deviation matrix. April 1997 . To analyze the sensitivity of model results to each parameter. 2. The frequency distribution of habitat suitability values in the landscape is given in Fig. Only 24% has habitat values of 0. Figure 2. 4. Each simulation consisted of 10. These calculations followed the algorithm described by Akçakaya (1995). in addition to densityindependent dispersal (Fig. The parameters M and K are different for each population. and (3) risk of falling below the threshold anytime within 50 years. Total rate of dispersal as a function of population size. when the population size (N) is lower than the carrying capacity (K). Analysis and Viability Measures Used The analysis of the dynamics of the California Gnatcatcher metapopulation with the model described above consisted of a series of simulations. Data are from Atwood et al. parameter and the medium estimates of all the other parameters (Table 1). The locations where gnatcatcher pairs were observed have habitat values ranging from 0. aspect. (2) risk of falling below the metapopulation threshold anytime within 20 years. 5a. and distance from wetlands were non-significant. Proportion of dispersing juveniles as a function of distance. and the average habitat suitability is about 0. No. (unpublished manuscript) for 1993 and 1994. using the lower.5 or above. The number of individuals in each stage of each population.5 or above.000 replications. Results Habitat Suitability Map The results of the logistic regression analysis are summarized in Table 3. About 95% of the observations are in locations with habitat values of 0. medium. and upper estimates of that Figure 3. Under strong densitydependent dispersal.

One method of validation is to separate the current observations in two and use only one set for estimation and the other set for testing. 5b) show the frequency distribution of the habitat suitability values of locations with gnatcatcher pairs in the south.4068 · 10 0.544 with 7 df (p 0.783 with 7 df (p 0.6781 · 10 2. The habitat function estimated from the northern half successfully predicted the observations in the southern half. We separated the observation locations and random points into two sets. We made two additional types of validation.8326 9. which gave a slightly different equation from the one estimated with Figure 5.8223 4 3 3 3 7 7 6 SE 1. Frequency distribution of the habitat suitability (HS) values of locations where California Gnatcatcher pairs were observed in all of the study area (a) and in the southern part of the study area (b).2153 · 10 2.4385 10. dividing the map roughly in half with all coastal observations in the southern half. they can be used to validate the model. aspect. Except for the small sample size (because only half the data are used). coefficient 6.0001 0.0017 0. We used this equation to predict the habitat suitability values of observed pair locations in the southern half of the data set. based on the habitat suitability function estimated with the northern half of the data set.5797 4 3 3 3 7 7 6 Wald 2 p 0.7118 20.6755 · 10 1. 2.8806 Goodness of fit (chi-square for covariates): Log likelihood statistic 523.5879 · 10 1.5 or more.0004 0. No.3759 · 10 5.5961 · 10 0.2860 · 10 8.0001) Score statistic 374. 1-ha cells in the study area.333.0017 16.1813 9. Slope.0001 0. The HS values in (b) are predicted using data only from the northern half of the data set. and thus are used as one of three validations of the habitat function.0002 0. distance from grassland (DGR) and distance from trees (DLR). this analysis does not provide an independent validation of the habitat function because these are the observations we used in the statistical analysis. elevation (ELV). Because these observations were not used in the estimation of the function.8438 · 10 1.6093 · 10 0.0001) *The habitat variables are coastal sage scrub (CSS). the results are similar to those in the previous figure: a large proportion (90%) of the observations are in cells with predicted habitat values of 0.8427 12. Validation of the Habitat Function Despite the highly significant fit of the data.3337 · 10 4. Conservation Biology Volume 11. and distance from wetlands were non-significant. Results of logistic regression for habitat function of California Gnatcatcher. and the obvious difference between the distribution of habitat suitability (HS) values at all locations and at locations with gnatcatcher observations. The results of this validation (Fig. with a habitat function that was estimated by regression of presence-absence data on habitat variables. with all inland observations in the northern half.8863 · 10 18. We re-estimated the habitat function with the northern half of the data set.1762 13.1444 25. The HS values were calculated for each of 85.9122 · 10 2. Frequency distribution of habitat suitability (HS) values for California Gnatcatchers in the study area. April 1997 .Akçakaya & Atwood California Gnatcatcher Metapopulation 429 Table 3. the whole data set.7494 · 10 8.* Variable* CSS ELV DGR DTR CSS · ELV CSS · DTR ELV · DTR constant Regr.0010 0.0001 0. We used the function estimated with the location of pairs to pre- Figure 4.

and about 76% in 50 years.7 Area as percentage of patches 49. No. and density dependence in dispersal. In other words.00 Figure 6. The results showed moderate sensitivity to seven other parameters: juvenile survival.03 4. or 553 adult pairs. and Allee effects that describe density effects at low abundances).430 California Gnatcatcher Metapopulation Akçakaya & Atwood dict the HS values of locations of single birds. and the habitat function based on 1992 pairs successfully predicted the observations of 1994 pairs. and areas of the patches identified by the model with medium values of parameters. The 2 largest patches made up about 84% of the total area of all patches (Table 4. white outlines are the outer borders of patches. Not all cells within the patch boundaries are suitable at the threshold level of 0. With medium estimates of all variables. Carrying capacities (K).0 68. The total carrying capacity was 3357 birds.3 3. the risk of falling below the metapopulation threshold of 60 individuals was about 19% in 20 years Table 4.28 0. 2.21 1. the patches have “holes” in them. The median time to fall below the threshold was about 34 years.4 202. All five parameters caused more than 10 years difference in median time to extinction and/or a difference of more than 0. threshold habitat suitability. Shades of gray represent habitat suitability (the darker the color.92 4.2 in risk of extinction (falling below the metapopulation threshold) in 50 years.1 2.7 0.8 8.82 33.99 0.2 8.6 0. Rank 1 2 3 4 5 6 7 8 9 10 11 12 13 Total K 1721 1130 133 124 72 39 36 32 32 13 11 7 7 3357 Init. metapopulation extinction threshold. 6). abund. These sensitivity results are taken from cumulative time to decline curves (as in Fig. as discussed above. dispersal distance.4 0. standard deviation of fluctuations in vital rates. initial abundances. carrying capacities.33 2. 6). The total initial abundance was 2952 birds.5 or more. and the habitat values in these areas are not used in the calculation of carrying capacities (K) or initial abundances (N0 ) reported in Table 4. We used a function estimated with the location of pairs in 1992 to predict the HS values of locations of pairs in 1994. and parameters related to density dependence (the type of density dependence. Such areas are not counted in the area of patches.1 2.35 0.21 100. correlation among population fluctuations.0 0.5 2.19 0.05 0. The results were most sensitive to the probability of weather-related catastrophes. and the thin.10 1. the model predicted a fast decline and a high risk of extinction of the gnatcatcher populations. or (at stable distribution) 629 adult pairs. The results were not sensitive to neighborhood distance. the black outline is the border of the study area. the higher the suitability).8 4. adult survival. Conservation Biology Volume 11.5 or more. initial abundances. April 1997 . About 82% of the pair locations in 1994 were predicted to have a HS value of 0. Fig. juvenile fecundity. as represented by the lighter gray regions within patches (Fig.51 1. Patch structure of the model superimposed on the habitat suitability map of California Gnatcatcher in Orange County. and the number of fires. The habitat function estimated with observations of pairs successfully predicted the observations of singles. 7). About 89% of the single locations were predicted to have a HS value of 0. 1514 994 117 109 63 34 32 28 28 11 10 6 6 2952 Area (km2) 101. and also very sensitive to adult fecundity. The sensitivities of these three results to each parameter are listed in Table 5.5. Patch Structure Given the habitat map and the (medium) parameter estimates described above the program found 13 habitat patches (clusters of suitable cells within the neighborhood distance of each other). Viability With most parameter combinations.

5% 21.2% 45. We incorporated natural variation (resulting from temporal fluctuations in environmental factors) in the form of randomly distributed vital rates (survivals and fecundities) and as natural catastrophes.9% 15.8% 84.5% Max.3 0.9% 15.0857 0.5% 16. One of potential improvements to the model involves expanding it to include the populations of California Gnatcatcher in other areas.3 7.3% Prob. 2.2 7. The sensitivity of results to catastrophe probability is not surprising.0160 0. in addition to a “medium” estimate. Number of fires Metapopulation threshold Effect a abs b 8.g. diff.1605 0.1278 0.0837 0. of decline in 20 years abs b 0.0% 14. April 1997 .1 6.0656 0.0.1204 perc c 44.0402 0. we used a wide range for this parameter.3% 23.Akçakaya & Atwood California Gnatcatcher Metapopulation 431 Table 5.145.2 7. Indicates whether high parameter value resulted in higher ( ) or lower ( ) viability.1452 0. For example.1627 0.4% 6.1056 0.0382 0. excluding some of the areas now included as habitat in the patches. c Difference as a percentage of the result with the medium value of the parameter. Most parameters are not precisely known because of measurement errors and lack of data.7% 55. In such cases. The declines predicted by the model also explains the insensitivity of metapopulation viability to fires.7% 37. We also modeled demographic stochasticity in reproduction. the function may be narrowed.5% 30.2% 30.7 years.5 11.8 years) with the medium estimate of all parameters. and dispersal.9% 4. and declines and extinctions occurred mostly due to catastrophic declines in the vital rates.4% 22.2% 76. Because we did not have enough data to correlate vital rates with weather-related variables. We modeled fires as causing a decrease in carrying capacity (K).0 5.2 perc c 25. but the populations were most of time well below K and thus were not affected by a decline in K caused by fires.0119 0.1021 perc c 21. the major weakness of this function is the lack of data on absences (locations where gnatcatchers are absent).1119 0.1119 0.5% 21.4% 8.2% 20. d Year at which the difference in risk of decline (with the high and low values of the parameter) was the largest.8% 4.0812 0.2332 0. or 77% of the risk (0. This might have caused an overestimation of habitat quality in certain locations. ) predicted by the stage matrix (with medium values of vital rates) was above 1.0914 0.0502 0. higher adult fecundity resulted in higher viability. including adult fecundity.9% 2.8 8.189) with the medium estimate.5% 43.4% 96. We used these types of natural variation (envi- ronmental and demographic) to express the model results in probabilistic terms (e. In other words.4% 1. as well as year-to-year environmental fluctuations and Allee effects. We used these ranges to estimate upper and lower bounds on the estimated viability of the species.8% 8.3% 181. which forced us to use random locations.7 3..0719 0.1617 0.9% 48. One of the specific aims of this analysis was to demonstrate how uncertainty and variability can be incorporated in models for determining the threat a species faces.0875 0.5% 40.1205 0. The difference in risk (of falling below the metapopulation threshold of 60 individuals within 20 years) between the low and high adult fecundity was about 0.1452 0.6% 42. A surprising result was the negative (but weak) effect Conservation Biology Volume 11.0334 0. estimating lower and upper bounds. The deterministic.5% Prob. we made three estimates of the parameter.2393 0.7% 9.6% 26. However. and the difference in median time to fall below the threshold between high and low estimates of adult fecundity was 14. We thus incorporated uncertainty that results from lack of knowledge in the form of parameter ranges.5% of the median time (33.5 7.1% 31.2% 21. If such data become available.7 10. The coastal sage scrub in the study area may be connected to similar habitat in southern Orange county and elsewhere.8% 21.0771 0.2% 34. Discussion The results of three types of validation of the habitat suitability function suggest the function is a good estimator of the quality of habitat in occupied locations.8% 19.1% 13.6% 21.9% 43.7 14. of decline in 50 years abs b 0. risk of decline).9% 24. (year)d 41 35 35 26 41 42 38 38 40 47 34 36 33 33 32 43 31 a Effect on metapopulation viability. No.3% 63.1662 0. b Absolute difference between the results of the two models with high and low values of the parameter.7342 0.9% 38. long-term growth rate (eigenvalue.1% 8.2% 19.4 14.6546 0.0604 0. survival.2% 345.0307 0. we used natural variability to estimate risks and uncertainties to estimate the uncertainty of these risk estimates. This corresponded to 43. Median time to decline (yr) Parameter Threshold HS Neighborhood distance Carrying capacity Initial abundance Dispersal distance Juvenile survival Juvenile fecundity Adult survival Adult fecundity Density dependence Allee effects (local threshold) Density dependent dispersal Standard deviations Correlation Catastrophe (weather) prob.1% 14.8 2.2% 34.7% 46.7 3.8 61. Another limitation of the habitat map is its geographic coverage.2888 0. Sensitivity of model results to parameters: difference in median time to decline and probability of decline between models with high and low values of each parameter.

. which can be explained by a type of source-sink dynamics. However.432 California Gnatcatcher Metapopulation Akçakaya & Atwood Figure 7. Specifically. When catastrophes were evenly distributed. Sensitivity of risk of decline to adult fecundity (Fa ). the difference in risk of decline in 20 years (B) and 50 years (C). Two of the model parameters. Increased dispersal distance meant a larger number of dispersers going from the more stable larger populations to these smaller and more extinction-prone populations. this result should not be interpreted in terms of reserve design because we also assumed there was no dispersal to the patches we deleted from the model (i. The arrows show four sensitivity results: the difference between median time to decline (A). The results of these simulations showed that when the smaller populations were excluded. The curves show the probability of falling below the metapopulation threshold as a function of time.e. which caused this effect. whether one wants to minimize Type I or Type II error). The vertical line marks the metapopulation extinction threshold of 60 individuals. 2. April 1997 . which in this case occurred around year 40.. of dispersal distance on viability. Conservation Biology Volume 11. even if their vital rates are the same as the larger populations. Threshold habitat suitability reflects a compromise between making sure that no suitable habitat is left out and making sure that unsuitable habitat is not included in patches. no guarantee that gnatcatchers would not attempt to disperse to these areas whether or not they are part of a reserve system. threshold habitat suitability and metapopulation extinction threshold. The other group includes density dependence parameters. No. increased dispersal caused a decrease in extinction risks. The sensitivity of results suggests that the results should not be interpreted in absolute terms. One group is the vital rates (especially fecundity) and the frequency and amount of change in vital rates caused by catastrophes. with the ninth largest and fifth largest populations. and the maximum difference in risk of decline (D). Such a data set may also help link the vital rates to local habitat suitability. To demonstrate that the smaller populations indeed act as sink populations. Although all populations in our model had the same vital rates (hence the same ). Detailed data on vital rates may also help estimate these parameters. have necessarily arbitrary values. The vertical dashed lines show the median time to decline. including Allee effects. so do these parameters. Increased dispersal had a similar effect of increasing extinction risk in another model: Akçakaya and Baur (1996) concluded that when some populations were modeled as being subject to more severe catastrophes than others. eliminating one of the stronger assumptions of our model (that vital rates are the same for all populations). the extinction threshold depends on whether it is more important not to overestimate or not to underestimate extinction risks. We made this comparison only to demonstrate that small populations can act as sink populations. Just as the level of statistical significance depends on priorities (e. The rationale for selecting any value for these thresholds is similar to the one used in deciding a probability level for statistical significance. There is. and 5 populations.g. 9. Similarly. The results point to a need to better estimate two groups of ecological parameters. the risk of decline to any level was lower (Fig. it would be inappropriate to use the results of this Figure 8. by simply deleting the smaller populations. it created a type of source-sink dynamics. we ran two other simulations. of course. 8). dispersal was only to the patches that remained in the model). for example by comparing fecundities in regions with different density of gnatcatchers. Risk of decline (by 80% to 100% from the initial metapopulation abundance) predicted by models with 13. the smaller populations were more prone to extinction as a result of demographic stochasticity. It is therefore positive that the results were only moderately sensitive to these parameters.

H. Cactus Wrens. Interface between ecology and land development in California. Pearce. Once a model is developed. R. Note Added in Proof Various data in this paper are from submitted. Pimm and two anonymous reviewers made helpful comments and suggestions. F. The consequences of these parameter sets can be estimated by the model in terms of the viability of the species and then used to rank alternative management actions. L. 1995. Bontrager. manuscripts. Manomet Observatory for Conservation Sciences.’s involvement was financially supported by the trustees of Manomet Observatory for Conservation Sciences. L. home range and breeding biology of the California Gnatcatcher in south Orange County. Lebben for his help in organizing the GIS data. New York. 1996. S. M.. but all assumptions may still predict that a particular management option ranks better than its alternatives in terms of the viability of the species. H. C. Baur. Luttrell.. 1991. L.S. J. R. H. California. No. As more demographic and ecological data accumulate. This would make such a relative ranking a much more reliable prediction than a prediction of exactly what the population size would be 50 years from now. 1995 breeding biology of the California Gnatcatcher in the San Joaquin Hills. Ginzburg. Pages 73–87 in A. A method for simulating demographic stochasticity. Despite this uncertainty. Bontrager and E. RAMAS/GIS: Linking landscape data with population viability analysis (ver 2. R. Chapman and Hall. and M. H. Basel. Southern California Edison. it is possible that relative rankings of management options will not be as sensitive. Even if results of a model are sensitive to various model parameters. Habitat requirements. it will become possible to make assessments with longer time horizons. Birkhaeuser Verlag.. and C. 1991. Akçakaya. A. Habitat preferences of the California Gnatcatcher in San Diego County. London. R. Downer for his help in the initial phase of the research. California. 1991. 1993. A.0t).Akçakaya & Atwood California Gnatcatcher Metapopulation 433 model to conclude that gnatcatcher populations in Central/Coastal Orange County are either threatened by extinction or secure from such a threat. Ferson. H. April 1997 .A. and H. In other words. 1995. T. Atwood. 1994. J. Loeschcke. R.. R. it can be improved with such analyses carried out at least once a year to incorporate new data collected during that year. 1991. The second use of such a model is to rank management options in terms of their predicted effect on the viability of target species. J. Carlsbad. but as yet unpublished. M. Orange County. Massachusetts. R. Quarterly Review of Biology 71:167–168. and to evaluate the importance of different parameters. Akçakaya. California. Atwood. and conservation of coastal sage scrub on the Palos Verdes Peninsula: progress report no. S. M. thesis. and D. Gnatcatchers and development: a “train wreck” avoided? Illahee 10:123–120. Applied Biomathematics. Oecologia 105:475–483. In principle. Atwood.. The cumulative time to quasiextinction results showed that the greatest difference between the results given by different parameter combinations occurred around 30 to 40 years. Risk assessment in conservation biology. R.. Akçakaya. Linking landscape data with population viability analysis: management options for the helmeted honeyeater. Pages 149–169 in J. Ecological Modelling 54:133–136. J. Akçakaya. D. Biological Conservation 73:169–176. and the Superpark Project. H. E. Southern California Academy of Sciences. U. D. Palmer for making this collaboration possible. we believe the model can have practical use in two areas. R. and R. M. California Gnatcatchers. The sensitivity analysis described is one potential use of modeling in the conservation and management of the coastal sage scrub community. K. Boyce. Literature Cited Akçakaya. Noss. Reynolds. Gorospe. McCarthy. Effects of population subdivision and catastrophes on the persistence of a land snail metapopulation. J. 1995. L. D. Acknowledgments This study was funded by Southern California Edison. Kamada. all possible management actions can be represented as changes in habitat suitability or demographic parameters (including their variances and the initial values of variables such as abundances in each stage and each population). California. and J. L. California gnatcatchers and coastal sage scrub: the biological basis for endangered species listing. R. The uncertainty of model results depends on the time horizon of the simulation. Conservation Biology Volume 11. Results of population viability models such as this one are less reliable if interpreted as absolute predictions than if interpreted as relative to other sets of assumptions or scenarios. This suggests that the results are most sensitive to changes in parameters when the time horizon is only a few decades. 1993. editor. Burgman. 1995. Setauket. Akçakaya. Longer or shorter time horizons may underestimate the effects of management actions because the model results may not be as sensitive to changes in parameters (as a function of the simulated effect of the management action). the specific predictions of the model may change as a function of parameters and assumptions. San Diego. These papers are available from the authors. Seitz and V. A. Anderson. San Diego State University. R. There is simply too much uncertainty about most of the parameters to predict with confidence what the population size will be in 50 years or what the risk of extinction might be. Almanza provided valuable discussion of gnatcatcher biology in Orange County. Manomet. 2 (1994). This might be an appropriate time horizon if the model is used in the future to compare alternative management options. Santa Margarita Company. We thank J. and L. to prioritize conservation measures. E. Fish and Wildlife Service. J. Review of RAMAS/GIS. Species conservation: a population-biological approach. 2. Los Angeles. A. Unpublished report. and B. it gives information about which parameters need to be estimated more carefully. Keeley. Bontrager.. editors. 1996. Ecological risk analysis for single and multiple populations. Overbey.

Inc. LaHaye. Central and coastal subregion. SAS user’s guide: statistics. S. T. E.R. 3rd edition. Report. Valuable modeling tool: RAMAS/GIS: Linking landscape data with population viability analysis. Gutiérrez. Massachusetts. BioScience Supplement 1995 (Science and Biodiversity Policy):84–90. W. C. North Carolina. Jones and Stokes Associates. and interpretation. SAS Institute. E. General Technical Report PSW-58. Akçakaya. Reid. Conrad and W. natural community conservation plan/habitat conservation plan (NCCP/HCP) Draft. U. Providing a regional context for local conservation action. Berkeley. U. Proceedings of the symposium on environmental consequences of fire and fuel management in mediterranean ecosystems. U. 1982. 1989. J. General technical report PSW-58. T. Keeley. Spotted owl metapopulation dynamics in southern California. 1995. 1994. 1982. R. 2. Sunderland. H. SAS Institute. editors. D. Cary. Matrix population models: construction. Pacific Southwest Forest and Range Experiment Station. Kingston. No. and H. J. Westman. editors.S.S. Forest Service. Sinauer Associates. Distribution of lightning and man-caused wildfires in California. Pages 91–98. Coastal sage succession. Oechel. California Department of Fish and Game. J. Oechel. 1993.. Inc. Conrad and W. Santa Ana. April 1997 . Orange County Environmental Management Agency. Forest Service. Murphy. Conservation Biology Volume 11. Meade Consulting. 1995. Journal of Animal Ecology 63:775–785. analysis. in C. version 6. S.S.. California. E. W. Conservation Biology 9:966–968. 1990. C. Fish and Wildlife Service. 1995. E. Orange County Environmental Management Agency. Proceedings of the symposium on dynamics and management of Mediterranean-type ecosystems. Pages 431–437. and D. California. Berkeley California. in C.434 California Gnatcatcher Metapopulation Akçakaya & Atwood Caswell. Methods used to survey the vegetation of Orange County parks and open areas and The Irvine Company property. Pacific Southwest Forest and Range Experiment Station. R.

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