Dispersal per recruit: an efficient method for assessing sustainability in marine reserve networks.

Marine reserves are an increasingly important tool for the management of marine ecosystems around the world. However, the effects of proposed marine reserve configurations on sustainability and yield of populations are typically not estimated because of the computational intensity of direct simulation and uncertainty in larval dispersal and density-dependent recruitment. Here we develop a method for efficiently assessing a marine reserve configuration for persistence and yield of a population with sedentary adults and dispersing larvae. The method extends the familiar sustainability criteria of individual replacement for single populations based on eggs-per-recruit (EPR) to spatially distributed populations with sedentary adults, a dispersing larval phase, and limited carrying capacity in the settlement-recruit relationship. We refer to this approach as dispersal-per-recruit (DPR). In some cases, a single DPR calculation, based on the assumption that post-settlement habitat is saturated (i.e., at maximum recruitment), is sufficient to determine population persistence, while in other cases further iterative calculations are required. These additional calculations reach an equilibrium more rapidly than a full simulation of age- or size-structured populations. From the DPR result, fishery yield can be computed from yield-per-recruit (YPR) at each point. We assess the utility of DPR calculations by applying them to single reserves, uniformly distributed systems of reserves, and randomly sized and spaced systems of reserves on a linear coastline. We find that for low levels of EPR in fished areas (e.g., 10% or less of the natural, unfished EPR when post-settlement habitats are saturated by 35% of natural settlement), a single DPR calculation is sufficient to determine persistence of the population. We also show that, in uniform systems of reserves with finite reserve size, maximal fisheries yield occurs when the density of reserves is such that all post-settlement habitat is nearly saturated with settlers. Finally, we demonstrate the application of this approach to a realistic proposed marine reserve configuration.     

Seascape-dependent subtidal-intertidal trophic linkages

In this study, we test in southern New Zealand a conceptual model of food web linkage that is seascape dependent, which can explain some of the variability in rocky shore community structure among sites and coasts. Using a comparative-experimental approach at local and distant sites we demonstrate that mobile subtidal predators (fish and crabs) can exert strong predation pressure on small mussels in the low tidal zone, but only in sites where the seascape includes subtidal reefs. On intertidal benches with adjacent subtidal reefs (+SR), 60-100% of small (5-15 mm) transplanted mussels were removed within a day from experimental tiles on the low shore when unprotected from predation, compared to fully caged controls that had approximately 100% survival over several months. In partial cages that exclude fish but not crabs, survivorship was intermediate. In contrast, on benches without subtidal reefs (-SR) 40-100% of mussels survived for months, even if unprotected. This difference is expressed in lower cover (0-60%) of mussels on rocks at +SR benches compared to -SR benches (70-99%). The central to northern west coast of the South Island is composed mostly of -SR benches, and predation on small mussels there was low and similar to the -SR benches on the east coast, whereas the +SR benches on the east coast had much greater predation. This contrasts to other studies in New Zealand that examined only predation on larger mussels by seastars and concluded that predation is strong on the west coast and weak on the east coast. Excluding large predators from low-shore areas with new recruits for a year in one +SR site showed longer-term predation effects on their abundance and cover. Short-term sampling at the east coast sites showed that mussel settlement was greater in -SR compared to +SR sites, providing some evidence that seascapes may also affect settlement. Overall, predation depended on the local seascape and ultimately affected community structure via suppression of effective recruitment rates. This study emphasizes the importance of predation on early life stages of basal species and the influence of seascapes on top-down interactions between subtidal predators and their intertidal prey.     

A trait-based test for habitat filtering: convex hull volume

Community assembly theory suggests that two processes affect the distribution of trait values within communities: competition and habitat filtering. Within a local community, competition leads to ecological differentiation of coexisting species, while habitat filtering reduces the spread of trait values, reflecting shared ecological tolerances. Many statistical tests for the effects of competition exist in the literature, but measures of habitat filtering are less well-developed. Here, we present convex hull volume, a construct from computational geometry, which provides an n-dimensional measure of the volume of trait space occupied by species in a community. Combined with ecological null models, this measure offers a useful test for habitat filtering. We use convex hull volume and a null model to analyze California woody-plant trait and community data. Our results show that observed plant communities occupy less trait space than expected from random assembly, a result consistent with habitat filtering.     

Importance of competition mechanisms in successive invasions by polyphagous tephritids in La Reunion

Understanding the strength and modes of interspecific interactions between introduced and resident species (native or previously introduced) is necessary to predict invasion success. We evaluated different mechanisms of interspecific competition among four species of polyphagous fruit flies (Diptera: Tephritidae) from the island of La Reunion: one endemic species, Ceratitis catoirii, and three exotic species, C. capitata, C. rosa, and Bactrocera zonata, that have successively invaded the island. Larval competition experiments, i.e., co-infestations of the same fruit, and behavioral interference experiments measuring the ability of one female to displace another from a fruit, were performed among all pairs of the four species. We observed asymmetric and hierarchical interactions among species in both larval and adult interference competition. In agreement with the hypothesis that invasion is competition-limited, the competitive hierarchy coincided with the temporal sequence of establishment on the island, i.e., each newly established species tended to be competitively dominant over previously established ones.     

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.     

Contrasting structure and composition of the understory in species-rich tropical rain forests

In large samples of trees > or = 1 cm dbh (more than 1 million trees and 3000 species), in six lowland tropical forests on three continents, we assigned species with >30 individuals to one of six classes of stature at maturity (SAM). We then compared the proportional representation of understory trees (1-2 cm dbh) among these classes. The understory of the three Asian sites was predominantly composed of the saplings of large-canopy trees whereas the African and American sites were more richly stocked with trees of the smaller SAM classes. Differences in class representation were related to taxonomic families that were present exclusively in one continent or another. Families found in the Asian plots but not in the American plot (e.g., Dipterocarpaceae, Fagaceae) were predominantly species of the largest SAM classes, whereas families exclusive to the American plots (e.g., Melastomataceae sensu stricto, Piperaceae, and Malvaceae [Bombacacoidea]) were predominantly species of small classes. The African plot was similar to Asia in the absence of those American families rich in understory species, while similar to America in lacking the Asian families rich in canopy species. The numerous understory species of Africa were chiefly derived from families shared with Asia and/or America. The ratio of saplings (1-2 cm dbh) to conspecific canopy trees (>40 cm dbh) was lower in American plots than in the Asian plots. Possible explanations for these differences include phenology, moisture and soil fertility regimes, phyletic constraints, and the role of early successional plants in forest development. These results demonstrate that tropical forests that appear similar in tree number, basal area, and the family taxonomy of canopy trees nonetheless differ in ecological structure in ways that may impact the ecology of pollinators, dispersers, and herbivores and might reflect fundamental differences in canopy tree regeneration.     

Carbon sequestration in California agriculture, 1980-2000.

To better understand agricultural carbon fluxes in California, USA, we estimated changes in soil carbon and woody material between 1980 and 2000 on 3.6 x 10(6) ha of farmland in California. Combining the CASA (Carnegie-Ames-Stanford Approach) model with data on harvest indices and yields, we calculated net primary production, woody production in orchard and vineyard crops, and soil carbon. Over the 21-yr period, two trends resulted in carbon sequestration. Yields increased an average of 20%, corresponding to greater plant biomass and more carbon returned to the soils. Also, orchards and vineyards increased in area from 0.7 x 10(6) ha to 1.0 x 10(6) ha, displacing field crops and sequestering woody carbon. Our model estimates that California's agriculture sequestered an average of 19 g C x m(-2) x yr(-1). Sequestration was lowest in non-rice annual cropland, which sequestered 9 g C x m(-2) x yr(-1) of soil carbon, and highest on land that switched from annual cropland to perennial cropland. Land that switched from annual crops to vineyards sequestered 68 g C x m(-2) x yr(-1), and land that switched from annual crops to orchards sequestered 85 g C x m(-2) x yr(-1). Rice fields, because of a reduction in field burning, sequestered 55 g C x m(-2) x yr(-1) in the 1990s. Over the 21 years, California's 3.6 x 10(6) ha of agricultural land sequestered 11.0 Tg C within soils and 3.5 Tg C in woody biomass, for a total of 14.5 Tg C statewide. This is equal to 0.7% of the state's total fossil fuel emissions over the same time period. If California's agriculture adopted conservation tillage, changed management of almond and walnut prunings, and used all of its orchard and vineyard waste wood in the biomass power plants in the state, California's agriculture could offset up to 1.6% of the fossil fuel emissions in the state.     

Swim speeds of free-ranging great cormorants

Information about foraging speeds is particularly valuable when the impact of a predator species upon a community of prey has to be defined, as in the case of great cormorants. We measured the swim speed of 12 (six males and six females) free-ranging great cormorants Phalacrocorax carbo, foraging off the Greenland coast during the summer of 2003, using miniaturized data-loggers. Although mean body mass of males was 27% greater than that of females, and mean swim speed of males were 29–57% higher than that of females during foraging phases (but not descent phases) of dives, these differences in speeds were not significant due to high variances. Birds descended to the mean maximum depth of 4.7 m at an average speed of 1.6±0.5 m s⁻¹, a speed similar to that measured in captive cormorants in previous studies. Although bursts of up to 4 m s⁻¹ were recorded, speed usually decreased during the deepest (foraging) phase of dives, being on average 0.8±0.6 m s⁻¹. Speeds measured here should be taken with caution, because the large propeller loggers used to measure speed directly decreased descent speeds by up to 0.5 m s⁻¹ when compared to smaller depth-only loggers. Cormorants in Greenland seem to combine two searching strategies, one requiring low speed to scan the water column or benthos, and one requiring high speed to pursue prey. These two strategies depend on the two main habitats of their prey: pelagic or demersal.     

Residency and depth movements of a coastal group of Atlantic cod (Gadus morhua L.)

Atlantic cod stocks are subdivided into geographically and biologically discrete populations; some being locally resident, others migratory. The reasons for the variation in migration pattern and the consequences it may have for population structuring is poorly understood. We studied a group of cod from the coastal waters of the Shetland Isles in the northern North Sea. During the spawning season, electronic tags that record depth and temperature over time were implanted into 133 individuals and the fish released within a few kilometres of where they were captured. Thirty-nine cod have been recaptured up to 608 days later, throughout the year and, in all but two cases, within 15 km of their release site. Geolocation methods based on temperature and bathymetry also suggested that the cod had a limited home range, remaining resident year-round in coastal waters. The cod were deeper during the winter and moved to shallower water in late spring and summer. They showed diel, fortnightly and monthly cycles in depth movement that varied much within and between individuals over the season. Residency and the individual variability in vertical movement may reflect a combination of locally complex depth strata, variable seabed substrate and the wide range of seasonally available prey resources.