Influence of Habitat Patchiness on Genetic Diversity and Spatial Structure of a Serpentine Endemic Plant

Abstract: Conservation of rare plant species often involves small, local populations dispersed within apparently suitable habitat. We used enzyme electrophoresis to study genotypic diversity in 32 populations of Calystegia collina , a self-incompatible clonal plant endemic to serpentine substrate in northern California's Coast Range. Genotypic diversity within a population was a good predictor of seed set by randomly marked flowers. Ramets from the most abundant genotypes in a population were most likely to produce flowers, but flowers from abundant genotypes were less likely to produce seed capsules than were flowers from rarer genotypes. These results are consistent with previous findings that reproductive success in C. collina is limited by the availability of compatible pollen. On small serpentine outcrops supporting only one or two populations, C. collina did not show reduced genotypic diversity or heterozygosity compared with populations on large outcrops supporting many populations. Instead, genotypic diversity and outcrop size had strong but independent influences on reproductive success. Although large serpentine outcrops contain more populations and provide better conditions for flower and seed production, significant diversity of unique genotypes clearly exists in isolated serpentine outcrops. Our findings suggest that plant conservation strategies must take into account the natural distribution of populations. The effects of habitat fragmentation on C. collina and other plant species that occur naturally in small, discrete patches may be unlike those that have been documented in more recently fragmented species.     

Testing the abundant center model using range-wide demographic surveys of two coastal dune plants

It is widely accepted that species are most abundant at the center of their geographic ranges and become progressively rarer toward range limits. Although the abundant center model (ACM) has rarely been tested with range-wide surveys, it influences much thinking about the ecology and evolution of species' distributions. We tested ACM predictions using two unrelated but ecologically similar plants, Camissonia cheiranthifolia and Abronia umbellata. We intensively sampled both throughout their one-dimensional distributions within the Pacific coastal dunes of North America, from northern Baja California, Mexico, to southern Oregon, USA. Data from > 1100 herbarium specimens indicated that these limits have been stable for at least the last 100 years. Range-wide field surveys detected C. cheiranthifolia at 87% of 124 sites and A. umbellata at 54% of 113 sites, but site occupancy did not decline significantly toward range limits for either species. Permutation analysis did not detect a significant fit of geographical variation in local density to the ACM. Mean density did not correlate negatively with mean individual performance (plant size or number of seeds/plant), probably because both species occur at low densities. Although size and seeds per plant varied widely, central populations tended to have the highest values for size only. For C. cheiranthifolia, we observed asymmetry in the pattern of variation between the northern and southern halves of the range consistent with the long-standing prediction that range limits are imposed by different ecological factors in different parts of the geographical distribution. However, these asymmetries were difficult to interpret and likely reflect evolutionary differentiation as well as plastic responses to ecological variation. Both density and seeds per plant contributed to variation in seed production per unit area. In C. cheiranthifolia only, sites with highest seed production tended to occur at the range center, as predicted by the ACM and assumed by theory proposing that range limits evolve via antagonism between natural selection and gene flow.     

Geographic structure of pollinator communities, reproductive assurance, and the evolution of self-pollination

Reproductive assurance is often invoked as an explanation for the evolution of self-fertilization in plants. However, key aspects of this hypothesis have received little empirical support. In this study, I use geographic surveys of pollinator communities along with functional studies of floral trait variation to examine the role of pollination ecology in mating system differentiation among populations and subspecies of the annual plant Clarkia xantiana. A greenhouse experiment involving 30 populations from throughout the species' range indicated that variation in two floral traits, herkogamy and protandry, was closely related to levels of autofertility and that trait variation was partitioned mainly among populations. Emasculation experiments in the field showed that autonomous selfing confers reproductive assurance by elevating fruit and seed production. Surveys of pollinator communities across the geographic range of the species revealed that bee pollinator abundance and community composition differed dramatically between populations of the outcrossing subspecies xantiana and the selfing subspecies parviflora despite their close proximity. Specialist bee pollinators of Clarkia were absent from selfing populations, but they were the most frequent visitors to outcrossing populations. Moreover, within the outcrossing subspecies xantiana, there was a close correspondence between specialist abundance and population differentiation in herkogamy, a key mating system trait. This spatial covariation arose, in part, because geographically peripheral populations had reduced herkogamy, higher autofertility, and lower pollinator abundance compared to central populations of xantiana. Finally, I detected strong spatial structure to bee communities both across the range of the species and within the outcrossing subspecies. In both cases, spatial structure was stronger for specialist bees compared to generalist bees, and pollinator communities varied in parallel with population variation in herkogamy. These results provide evidence that mating system differentiation parallels spatial variation in pollinator abundance and community composition at both broad and more restricted spatial scales, consistent with the hypothesis that pollinator abundance and reproductive assurance are important drivers of plant mating system evolution.     

Reproductive Success in Reestablished versus Natural Populations of a Threatened Grassland Daisy (Rutidosis leptorrhynchoides)

Abstract: Populations of the endangered Rutidosis leptorrhynchoides (Asteraceae) have been reestablished in conservation reserves to reduce their risk of extinction in the wild. The reproductive success (number of seeds per inflorescence, percent seed set, germinability) of five small reestablished populations (at 5–10 years after establishment) was compared to that of two large natural remnant populations from which they were derived. Initial growth rates under glasshouse conditions were compared for seedlings derived from seed from remnant and reestablished populations. Seed set per inflorescence in all five reestablished populations was equal to or greater than seed set in remnant population plants. The resulting seed was as germinable, and in some cases more germinable, than seed derived from remnant populations; the seedlings then grew as large or larger than remnant population seedlings grown under glasshouse conditions. Fitness reductions in small reestablished populations of R. leptorrhynchoides are not evident at the reproductive stage. Hence, these populations have some potential to maintain natural regeneration processes and might therefore positively contribute to the conservation of this species.     

Nonrandom, Size- and Timing-Biased Breeding in a Hatchery Population of Steelhead Trout

Abstract:  Hatcheries have been built and operated to buffer salmon and trout populations from overfishing and to compensate for habitat lost or degraded by human activities. These facilities are now so prevalent that in some cases hatchery-produced salmon outnumber salmon produced in the wild. By default, this makes them an important component in the current ecology and evolution of salmonids. Hatcheries differ from natural environments in many ways, and among the most fundamental is the necessity that humans select fish for breeding instead of allowing natural processes of mate choice and competition. We examined the mating system for steelhead trout ( Oncorhynchus mykiss ) at Forks Creek Hatchery in southwest Washington and investigated factors affecting selection of individual steelhead for spawning by the hatchery staff. Despite efforts by the staff to not spawn selectively, data on steelhead spawned over 7 years revealed selection for large adult body size and early reproductive timing and a tendency for size-assortative mating (i.e., large with large). Selection on size was related to selection on reproductive timing because early returning fish tended to be larger than those returning later. To improve the fitness of both hatchery fish destined to spawn in the wild and hatchery fish designated to be spawned in the hatchery, a better understanding of factors associated with the range of reproductive success and mate-choice mechanisms in the wild is vital. This knowledge may then be applied to artificial propagation programs designed for conservation or enhancement.     

Quantifying Plant Population Persistence in Human-Dominated Landscapes

Abstract:  We assessed population performance of rare plants across a gradient from rural to urban landscapes and evaluated 2 hypotheses central to strategic conservation planning: (1) population performance declines with increasing human dominance and (2) small populations perform poorly relative to larger ones. Assessing these hypotheses is critical to strategic conservation planning. The current conservation paradigm adheres to the well-established ecology theory that small isolated populations, particularly those in human-dominated landscapes, are the least likely to succeed over the long term. Consequently, conservation planning has strongly favored large, remote targets for protection. This shift in conservation toward ecosystem-based programs and protection of populations within large, remote systems has been at the expense of protection of the rarest of the rare species, the dominant paradigm for conservation driven by the endangered species act. Yet, avoiding conservation of small populations appears to be based more on theoretical understanding and expert opinion than empiricism. We used Natural Heritage data from California in an assessment of population performance of rare plants across a landscape with an urban-rural gradient. Population performance did not decrease in urban settings or for populations that were initially small. Our results are consistent with a pattern of few species extinctions within these landscapes over the past several decades. We conclude that these populations within compromised landscapes can contribute to overall biodiversity conservation. We further argue that conservation planning for biodiversity preservation should allocate relatively more resources to protecting urban-associated plant taxa because they may provide conservation benefit beyond simply protecting isolated populations; they may be useful in building social interest in conservation.     

Genetic Diversity, Population Size, and Fitness in Central and Peripheral Populations of a Rare Plant Lychnis viscaria

Abstract: Genetic diversity is expected to decrease in small and isolated populations as a consequence of bottlenecks, founder effects, inbreeding, and genetic drift. The genetics and ecology of the rare perennial plant Lychnis viscaria (Caryophyllaceae) were studied in both peripheral and central populations within its distribution area. We aimed to investigate the overall level of genetic diversity, its spatial distribution, and possible differences between peripheral and central populations by examining several populations with electrophoresis. Our results showed that the level of genetic diversity varied substantially among populations (  H _exp = 0.000–0.116) and that the total level of genetic diversity (mean H _exp = 0.056) was low compared to that of other species with similar life-history attributes. The peripheral populations of L. viscaria had less genetic variation (mean H _exp = 0.034) than the central ones (0.114). Analysis of genetic structure suggested limited gene flow (mean F _ST = 0.430) and high differentiation among populations, emphasizing the role of genetic drift (  N _e m = 0.33). Isolation was even higher than expected based on the physical distance among populations. We also focused on the association between population size and genetic diversity and possible effects on fitness of these factors. Population size was positively correlated with genetic diversity. Population size and genetic diversity, however, were not associated with fitness components such as germination rate, seedling mass, or seed yield. There were no differences in the measured fitness components between peripheral and central populations. Even though small and peripheral populations had lower levels of genetic variation, they were as viable as larger populations, which emphasizes their potential value for conservation.     

Genetic Structure of Fragmented Populations of the Endangered Daisy Rutidosis leptorrhynchoides

Abstract: The endangered grassland daisy Rutidosis leptorrhynchoides has been subject to severe habitat destruction and fragmentation over the past century. Using allozyme markers, we examined the genetic diversity and structure of 16 fragmented populations. The species had high genetic variation compared to other plant species, and both polymorphism and allelic richness showed strong positive relationships with log reproductive population size, reflecting a loss of rare alleles (frequency of q < 0.1) in smaller populations. Fixation coefficients were positively related to size, due either to a lack of rare homozygotes in small populations or to Wahlund effects (owing to spatial genetic structure) in large ones. Neither gene diversity nor heterozygosity was related to population size, and other population parameters such as density, spatial contagion, and isolation had no apparent effect on genetic variation. Genetic divergence among populations was low , despite a large north-to-south break in the species' current distribution. To preserve maximum genetic variation, conservation strategies should aim to maintain the five populations larger than 5000 reproductive plants, all of which occur in the north of the range, as well as the largest southern population of 626 plants at Truganina. Only one of these is currently under formal protection. High heterozygosity in smaller populations suggests that they are unlikely to be suffering from inbreeding depression and so are also valuable for conservation. Erosion of allelic richness at self-incompatibility loci, however, may limit the reproductive capacity of populations numbering less than 20 flowering plants.     

Living at the Edge: Local versus Positional Factors in the Long‐Term Population Dynamics of an Endangered Orchid

Abstract: Populations at the margin of geographic ranges of distribution have been considered more vulnerable than central ones, but recent reviews have caste doubt on this generalization. We examined the reproductive and demographic performance of a rare Euroasiatic orchid (Cypripedium calceolus) at its southwesterly range limit and compared our findings with those of previous studies of nine central populations at the center of the orchid's range. We sought to test the central‐marginal model and to evaluate factors involved in long‐term performance of forest Eurosiberian species with peripheral populations in southern European mountains. We characterized (structure, temporal fluctuations, herbivory, reproductive success, and recruitment at different habitats) four Pyrenean populations of C. calceolus of different sizes (5–3500 ramets) and monitored three of them for up to 13 years. Two quantitative stochastic models (count data and matrix models) were used to assess population trends and viability and the effect of herbivory. Contrary to expectations, and despite the negative effect of sporadic events of herbivory, the peripheral populations we studied (except the smallest one) performed similarly or better than populations occurring in central part of the species’ range in terms of reproductive success and population growth rates. Landscape changes over the last 50 years suggest that natural reforestation could be involved in the success of this plant at its southern limit. Forest expansion in the mountain regions of southern Europe may provide new opportunities for plants with geographic distributions centered mainly at higher latitudes and give some hope for their recovery in future scenarios dominated by biodiversity loss.     

Role of climate and competitors in limiting fitness across range edges of an annual plant

It is often assumed that the geographic distributions of species match their climatic tolerances, but this assumption is not frequently tested. Moreover, few studies examine the relative importance of abiotic and biotic factors for limiting species ranges. We combined multiple approaches to assess the extent to which fitness of a widespread native annual legume, Chamaecrista fasciculata, decreases at and beyond its northern and western range edges, and how this is influenced by the presence of neighbors. First, we examined plant fitness and the effect of neighbors in natural populations at different geographic range locations for three years. Fitness decreased toward the northern range edge, but not the western edge. Neighbor removal had a consistently positive effect on seedpod production across all years and sites. Second, we established experimental populations at sites within the range, and at and beyond the northern and western range edges. We tracked individual fitness and recorded seedling recruitment in the following year (a complete generation) to estimate population growth rate. Individual fitness and population growth declined to near zero beyond both range edges, indicating that C. fasciculata with its present genetic composition will not establish in these regions, given conditions currently. We also carried out a neighbor removal treatment. Consistent with the natural populations, neighbors reduced seedpod production of reproductive adults. However, neighbors also increased early-season survival, and this positive effect early in life history resulted in a net positive effect of neighbors on lifetime fitness at most range locations. Our data show that the population growth rate of C. fasciculata includes values above replacement, and populations are well adapted to conditions up to the edge of the range, whereas the severely compromised fitness at sites beyond the edge precludes immediate establishment of populations and thereby impedes adaptation to these conditions.     

Influence of growth on reproductive traits and its effect on fertility and gene diversity in a clonal seed orchard of scots pine, Pinus Sylvestris L

This study was carried out in a clonal seed orchard of scots pine (Pinus sylvestris L.), to determine the difference and interaction for reproductive and growth characters among clones and its impact on fertility variation and gene diversity Numbers of female and male strobili, and height and diameter at breast height were studied on six grafts chosen randomly in each of the 27 clones for the purpose. One-way analysis of variance revealed large differences in both reproductive and growth characters among clones. The differences were higher in growth characters than in reproductive traits. There was significant phenotypic correlation among growth and reproductive characters. So, growth characters had a greater effect on male and female fertility Estimates of total fertility variation (Sibling coefficient = 1.012), status number (26.8) and relative gene diversity (0.981) were computed. Fertility variation among clones was low, which caused a high relative population size (99% of census number). The positive phenotypic correlation between growth and reproductive characters showed that enhanced growth rate could be effective in improving fertility and gene diversity of seed orchard crop. The results of the study have implications in breeding and selection of plus tree and populations, establishment and thinning of seed orchards of the species.     

Targeted gene flow and rapid adaptation in an endangered marsupial

Targeted gene flow is an emerging conservation strategy. It involves translocating individuals with favorable genes to areas where they will have a conservation benefit. The applications for targeted gene flow are wide-ranging but include preadapting native species to the arrival of invasive species. The endangered carnivorous marsupial, the northern quoll (Dasyurus hallucatus), has declined rapidly since the introduction of the cane toad (Rhinella marina), which fatally poisons quolls that attack them. There are, however, a few remaining toad-invaded quoll populations in which the quolls survive because they know not to eat cane toads. It is this toad-smart behavior we hope to promote through targeted gene flow. For targeted gene flow to be feasible, however, toad-smart behavior must have a genetic basis. To assess this, we used a common garden experiment, comparing offspring from toad-exposed and toad-naïve parents raised in identical environments, to determine whether toad-smart behavior is heritable. Offspring from toad-exposed populations were substantially less likely to eat toads than those with toad-naïve parents. Hybrid offspring showed similar responses to quolls with 2 toad-exposed parents, indicating the trait may be dominant. Together, these results suggest a heritable trait and rapid adaptive response in a small number of toad-exposed populations. Although questions remain about outbreeding depression, our results are encouraging for targeted gene flow. It should be possible to introduce toad-smart behavior into soon to be affected quoll populations.     

Using circuit theory to model connectivity in ecology, evolution, and conservation

Connectivity among populations and habitats is important for a wide range of ecological processes. Understanding, preserving, and restoring connectivity in complex landscapes requires connectivity models and metrics that are reliable, efficient, and process based. We introduce a new class of ecological connectivity models based in electrical circuit theory. Although they have been applied in other disciplines, circuit-theoretic connectivity models are new to ecology. They offer distinct advantages over common analytic connectivity models, including a theoretical basis in random walk theory and an ability to evaluate contributions of multiple dispersal pathways. Resistance, current, and voltage calculated across graphs or raster grids can be related to ecological processes (such as individual movement and gene flow) that occur across large population networks or landscapes. Efficient algorithms can quickly solve networks with millions of nodes, or landscapes with millions of raster cells. Here we review basic circuit theory, discuss relationships between circuit and random walk theories, and describe applications in ecology, evolution, and conservation. We provide examples of how circuit models can be used to predict movement patterns and fates of random walkers in complex landscapes and to identify important habitat patches and movement corridors for conservation planning.     

Behavioral differences between Pogonomyrmex rugosus and dependent lineage (H1/H2) harvester ants

The discovery of genetic caste determination (GCD) in populations of Pogonomyrmex harvester ants raises many questions about the evolution and persistence of such populations. The genetic caste determination arises from the existence of two distinct, but mutually dependent, genetic lineages within a population. Workers always develop from a combination of the two lineages, but their sister queens develop from within-lineage matings. Maintaining genetic caste determination appears to be costly because many queen-destined eggs are wasted when a colony is not in the reproductive stage, yet these populations appear to be widespread. We investigated whether inter-lineage workers have novel traits that give GCD colonies a selective advantage in certain environments. In particular, we compared ecologically relevant behavioral characteristics of inter-lineage workers in H-lineage colonies with co-occurring normal colonies of P. rugosus. First, we measured colony defensive response toward a simulated vertebrate predator. Second, we set up direct competitive foraging and recruitment experiments between dependent lineage and P. rugosus colonies. Last, we measured individual aggressive response to foreign inter-lineage and P. rugosus workers. We found that H1/H2 inter-lineage workers explored objects on the nest more thoroughly and responded much more aggressively to simulated predator disturbance than the P. rugosus colonies. In individual encounters, H1/H2 inter-lineage and P. rugosus workers were equally aggressive toward foreign ants, but both worker types could discriminate P. rugosus from inter-lineage intruders and were more aggressive toward ants of the alternate type to themselves. When competing directly for resources, however, P. rugosus colonies consistently dominated seed piles. In summary, H1/H2 GCD colonies show distinct behavioral differences, but there is no clear ecological advantage from the traits we examined.     

Rapid Evolution in the Wild: Changes in Body Size, Life-History Traits, and Behavior in Hunted Populations of the Japanese Mamushi Snake

Abstract:  Rapid evolution caused by human exploitation of wildlife is not usually addressed in studies of the impacts of such exploitation despite its direct relevance to population persistence. Japanese mamushi ( Gloydius blomhoffii ), an endemic venomous snake of the Japanese archipelago, has been heavily hunted by humans, and many populations appear to be declining or are already extirpated. We compared local populations that have been hunted regularly with populations that have not been hunted. Mamushi in hunted populations were smaller, had fewer vertebrae, produced more and smaller offspring, had increased reproductive effort among smaller females, and in nature fled at greater distances from an approaching human and were less defensive than mamushi in unhunted populations, as predicted from life-history theory. Heritability estimates for body size, number of vertebrae, and antipredator behavior were statistically significant, and neonates from hunted sites showed the same distribution of altered characters (compared with those from unhunted sites) as adults. Thus, distribution of the divergent trait between hunted and unhunted sites appeared in part to be genetically based, which suggests rapid evolution to human predation pressures. Trait distributions in hunted populations probably deviate from naturally (as opposed to anthropogenically) selected optima and, therefore, may have long-term negative repercussions on population persistence. Because rapid evolution affects a suite of parameters that characterize exploited populations, accurate understanding of the impacts of exploitation and effective resource management and conservation can only be achieved if evolutionary consequences are considered explicitly.     

The possible origin of Zostera noltii in the Canary Islands and guidelines for restoration

Seagrasses and their habitat are declining worldwide. Zostera noltii in the Canary Islands has been drastically reduced, mainly by anthropogenic disturbance, to three small surviving patches in a single harbor in Lanzarote. A previous genetic study, using neutral microsatellite markers, revealed that these patches consist of a single clonal individual. Here, an assignment test, using microsatellite data, was used to locate the most likely population of origin from a set of possible donor populations. Our results show that the Moulay Bousselham population in northern Morocco is assigned as the most likely population of origin (88%), although the probability of being at one generation time distance is low (2.7%). This result, however, allows locating the most closely related stands that may be the most successful donor populations for future restoration based on shoot or seed transplantation.     

Extinction times in experimental populations

Predicting population extinctions is a key element of quantitative conservation biology and population ecology. Although stochastic population theories have long been used to obtain theoretical distributions of population extinction times, model-based predictions have rarely been tested. Here I report results from a quantitative analysis of extinction time in 281 experimental populations of water fleas (Daphnia magna) in variable environments. To my knowledge, this is the first quantitative estimate of the shape of the distribution of population extinction times based on extinction data for any species. The finding that the distribution of population extinction times was extraordinarily peaked is consistent with theoretical predictions for density-independent populations, but inconsistent with predictions for density-dependent populations. The tail of the extinction time distribution was not exponential. These results imply that our current theories of extinction are inadequate. Future work should focus on how demographic stochasticity scales with population size and effects of nonrandom variable environments on population growth and decline.     

Population Persistence in Florida Torreya: Comparing Modeled Projections of a Declining Coniferous Tree

Abstract: The Florida torreya (   Torreya taxifolia ) is a coniferous tree endemic to a 35-km stretch of bluffs and ravines along the east side of the Apalachicola River in northern Florida and southern Georgia. This formerly locally abundant tree has declined as a result of disease during the 1950s and is on the U.S. endangered species list. With no seed production in the wild, this species is headed toward extinction. We conducted a survey on roughly 200 trees from 1988 to 1996 and used these data to estimate the likelihood of population persistence during the next several decades. We compared a stage-class transition matrix model ( RAMAS) and an individual-based stochastic model ( TORSIM) of growth and mortality to project future populations. Given the current lack of seed production in the wild, all models predict extinction. The question of concern is the imminence of this predicted extinction. Differing predicted times to extinction would suggest different immediate management recommendations. Both models predicted an over 90% likelihood of persistence during the next 50 years. Predictions differed in that the transition matrix model was less optimistic than the individual-based model regarding persistence. Model sensitivity analysis showed that the results were robust to significant decreases in growth and sprouting probabilities. Submodels identified different persistence likelihoods in different populations. Nonetheless, unless management of the population can facilitate maturation and seed production, extinction of this species in the wild is inevitable.     

Payments for Ecosystem Services as a Framework for Community-Based Conservation in Northern Tanzania

Abstract:  Payments for ecosystem services (PES) are an increasingly promoted approach to conservation. These approaches seek to develop financial mechanisms that create economic incentives for the maintenance of ecosystems and associated biodiversity by rewarding those who are responsible for provision of ecological services. There are, however, few cases in which such schemes have been used as a strategy for conserving wildlife in developing countries and very few operational examples of such schemes of any sort in sub-Saharan Africa. In savannah ecosystems, large mammal populations generally depend on seasonal use of extensive areas and are widely declining as a result of habitat loss, overexploitation, and policies that limit local benefits from wildlife. Community-based conservation strategies seek to create local incentives for conserving wildlife, but often have limited impact as a result of persistent institutional barriers that limit local rights and economic benefits. In northern Tanzania, a consortium of tourism operators is attempting to address these challenges through an agreement with a village that possesses part of a key wildlife dispersal area outside Tarangire National Park. The operators pay the community to enforce voluntary restrictions on agricultural cultivation and permanent settlement in a defined area of land. The initiative represents a potentially cost-effective framework for community-based conservation in an ecologically important area and is helping to reconcile historically conflicting local and national interests relative to land tenure, pastoralist livelihoods, and conservation. Wider adaptation of payments for ecosystem services approaches to settings where sustaining wildlife populations depends on local stewardship may help address current challenges facing conservation outside state-protected areas in savannah ecosystems in sub-Saharan Africa and other parts of the world.     

Effects of Habitat Size and Patch Isolation on Reproductive Success of the Serpentine Morning Glory

Abstract: We examined the effects of habitat area and patch isolation on reproductive success in serpentine morning glory ( Calystegia collina [Convolvulaceae]), a primarily self-incompatible clonal plant endemic to serpentine outcrops in northern California's coast ranges. Within a 4000-km ² region, we compared the reproductive success of C. collina on 16 small (<5 ha) and 7 large ( >300 ha) outcrops. Flower and fruit production were significantly higher on large serpentine outcrops than on small outcrops. Fruit production also was positively correlated with the soil's ratio of calcium to magnesium. Successful pollination was positively affected by flower density and the number of other flowering patches within 100 m of a C. collina patch. The number of nearby flowering patches was considerably higher on large than on small outcrops. Flowers on large outcrops did not receive significantly more bee visitors than flowers on small outcrops, suggesting that pollination success is related to the quality rather than the quantity of pollen deposited. Fruit production by plants on both small and large outcrops was enhanced by the experimental addition of pollen from other patches, but not by the addition of pollen from the same patch. These findings demonstrate that the size of habitat may have strong effects on the reproductive success of locally endemic plants by enhancing opportunities for successful sexual reproduction. They also warn against the presumption that naturally patchy plant species are invulnerable to the effects of habitat fragmentation.