Optimizing conservation in species-specific agricultural landscapes

Recovery of grassland birds in agricultural landscapes is a global imperative. Agricultural landscapes are complex, and the value of resource patches may vary substantially among species. The spatial extent at which landscape features affect populations (i.e., scale of effect) may also differ among species. There is a need for regional-scale conservation planning that considers landscape-scale and species-specific responses of grassland birds to environmental change. We developed a spatially explicit approach to optimizing grassland conservation in the context of species-specific landscapes and prioritization of species recovery and applied it to a conservation program in Kentucky (USA). We used a hierarchical distance-sampling model with an embedded scale of effect predictor to estimate the relationship between landscape structure and abundance of eastern meadowlarks (Sturnella magna), field sparrows (Spizella pusilla), and northern bobwhites (Colinus virginianus). We used a novel spatially explicit optimization procedure rooted in multi-attribute utility theory to design alternative conservation strategies (e.g., prioritize only northern bobwhite recovery or assign equal weight to each species’ recovery). Eastern meadowlarks and field sparrows were more likely to respond to landscape-scale resource patch adjacencies than landscape-scale patch densities. Northern bobwhite responded to both landscape-scale resource patch adjacencies and densities and responded strongly to increased grassland density. Effects of landscape features on local abundance decreased as distance increased and had negligible influence at 0.8 km for eastern meadowlarks (0.7–1.2 km 95% Bayesian credibility intervals [BCI]), 2.5 km for field sparrows (1.5–5.8 km 95% BCI), and 8.4 km for bobwhite (6.4–26 km 95% BCI). Northern bobwhites were predicted to benefit greatly from future grassland conservation regardless of conservation priorities, but eastern meadowlark and field sparrow were not. Our results suggest similar species can respond differently to broad-scale conservation practices because of species-specific, distance-dependent relationships with landscape structure. Our framework is quantitative, conceptually simple, customizable, and predictive and can be used to optimize conservation in heterogeneous ecosystems while considering landscape-scale processes and explicit prioritization of species recovery.     

Sexual selection and cuckoldry in a monogamous songbird: implications for sexual selection theory

Sexual selection is generally assumed to be weaker in monogamous than in polygynous animals. Recently, though, extra-pair fertilizations have been hailed as an important force in generating variance in reproductive success among males in socially monogamous species, thereby increasing the intensity of sexual selection. To see if extra-pair copulations contribute to variance in male reproductive success in the house finch (Carpodacus mexicanus), we used DNA fingerprinting to determine the paternity of chicks from 35 nests. This species is a socially monogamous passerine in which plumage brightness serves as a sexually selected indicator of male quality. Out of 119, nestlings 10 (8.3%) were fathered by a male other than the attending male, but cuckoldry occurred randomly with respect to the plumage colouration, size, or age of the attending male. Thus extra-pair fertilizations do not generate variance in male reproductive success with respect to plumage colour. On the other hand, a strongly male-biased sex ratio and asynchronous breeding by females may generate substantial variance in male reproductive success and could explain the evolution of ornamental colouration.     

Decoupling fragmentation from habitat loss for spiders in patchy agricultural landscapes

Habitat loss reduces species diversity, but the effect of habitat fragmentation on number of species is less clear because fragmentation generally accompanies loss of habitat. We compared four methods that aim to decouple the effects of fragmentation from the effects of habitat loss. Two methods are based on species-area relations, one on Fisher's alpha index of diversity, and one on plots of cumulative number of species detected against cumulative area sampled. We used these methods to analyze the species diversity of spiders in 2, 3.2 × 4 km agricultural landscapes in Southern Judea Lowlands, Israel. Spider diversity increased as fragmentation increased with all four methods, probably not because of the additive within-patch processes, such as edge effect and heterogeneity. The positive relation between fragmentation and species diversity might reflect that most species can disperse through the fields during the wheat-growing season. We suggest that if a given area was designated for the conservation of spiders in Southern Judea Lowlands, Israel, a set of several small patches may maximize species diversity over time.     

Comparing geographic boundaries in songbird demography data with vegetation boundaries: a new approach to evaluating habitat quality

To plan for the habitat needs of forest songbirds of conservation concern, managers need to understand how spatial heterogeneity in forest conditions influences habitat quality. I used difference boundary detection (wombling) and spatially constrained clustering to delineate boundaries in various combinations of four forest vegetation variables (understory height, understory density, percent deciduous vs. conifer understory, and percent canopy closure) in two Michigan northern hardwood forests. My goal was to identify vegetation boundaries that corresponded with boundaries in an understory-dependent songbird’s distribution, and with boundaries in demographic measures for this songbird that indicate habitat quality (e.g., occupancy by older vs. yearling males, reproductive success). Both forests were actively-managed, mature stands: The first site (78 ha) was heavily deer-browsed (HB), with many browse-resistant conifers in the understory, and the second (62 ha) was less-browsed (LB), with deciduous-dominated understory. I compared the vegetation difference and cluster boundaries to difference boundaries based on 6 years of distribution and demographic data for black-throated blue warblers (Dendroica caerulescens). At the HB site, warbler boundaries overlapped strongly with vegetation boundaries that included all four variables, and clustering effectively divided the habitat into areas with different warbler occupancy and demographic characteristics. At the LB site, warbler distribution showed high overlap with difference and cluster boundaries based on just the height and density of understory vegetation, and cluster boundaries again effectively partitioned the study area into sites that varied in habitat quality. Thus, geographic boundary analysis is likely to be a useful tool for identifying key vegetation variables for management, and for delineating clusters (habitat patches) within sites that capture differences in habitat quality.

Grassy field margins as potential corridors for butterflies in agricultural landscapes: A simulation study

Over the last decades, agricultural intensification has caused a dramatic reduction of grassy habitats. This habitat loss has had a strong negative effect on many meadow-living insect populations, including butterflies. As a part of the cross-compliance measures of the Common Agricultural Policy of the European Union, subsidies for creation and maintenance of grassy field margins (GFM) have been launched. Among other environmental issues, they may serve as corridors for movement of various meadow-living species between individual meadows. Their role as corridors has, however, not yet been demonstrated at the landscape scale and their characteristics that most significantly increase landscape connectivity are unknown. Empirical data for such studies are missing, as the GFM subsidies were launched only 3 years ago. One possibility to get some predictions of their outcomes is provided by simulation models. Here we present our simulation results, using an extension of the model developed by Kindlmann et al. (2004) for the Meadow Brown butterfly, Maniola jurtina. The extension includes the probability to cross a boundary (Conradt and Roper, 2006) that negatively influences dispersal rates but increases sensitivity to the corridor effect. Our simulations show that GFMs increase the dispersal rates between habitat patches and we predict the optimal combinations of width and number of GFMs in the landscape. This way we provide a decision-making tool for increasing landscape connectivity for M. jurtina and similar species. Although our simulations are based on a particular species, they may be generalized because this species shows dispersal rates that are typical of butterfly metapopulations (Conradt et al., 2000), and a potentially widespread dispersal kernel (i.e. “foray search”) that has been reported in a wide variety of species (see Conradt et al., 2003 for a review).     

A comparison of land-sharing and land-sparing strategies for plant richness conservation in agricultural landscapes

Strategies for conserving plant diversity in agroecosystems generally focus on either expanding land area in non-crop habitat or enhancing diversity within crop fields through changes in within-field management practices. In this study, we compare effects on landscape-scale species richness from such land-sharing or land-sparing strategies. We collected data in arable field, grassland, pasture, and forest habitat types (1.6 ha sampled per habitat type) across a 100-km2 region of farmland in Lancaster County, Pennsylvania, USA. We fitted species-area relationships (SARs) for each habitat type and then combined extrapolations from the curves with estimates of community overlap to estimate richness in a 314.5-ha landscape. We then modified these baseline estimates by adjusting parameters in the SAR models to compare potential effects of land-sharing and land-sparing conservation practices on landscape richness. We found that species richness of the habitat types showed a strong inverse relationship to the relative land area of each type in the region, with 89 species in arable fields (66.5% of total land area), 153 in pastures (6.7%), 196 in forests (5.2%), and 213 in grasslands (2.9%). Relative to the baseline scenario, major changes in the richness of arable fields produced gains in landscape-scale richness comparable to a conversion of 3.1% of arable field area into grassland habitat. Sensitivity analysis of our model indicated that relative gains from land sparing would be greatest in landscapes with a low amount of non-crop habitat in the baseline scenario, but that in more complex landscapes land sharing would provide greater gains. These results indicate that the majority of plant species in agroecosystems are found in small fragments of non-crop habitat and suggest that, especially in landscapes with little non-crop habitat, richness can be more readily conserved through land-sparing approaches.     

Reptile and arboreal marsupial response to replanted vegetation in agricultural landscapes.

We report reptile and arboreal marsupial responses to vegetation planting and remnant native vegetation in agricultural landscapes in southeastern Australia. We used a hierarchical survey to select 23 landscapes that varied in the amounts of remnant native vegetation and planted native vegetation. We selected two farms within each landscape. In landscapes with plantings, we selected one farm with and one farm without plantings. We surveyed arboreal marsupials and reptiles on four sites on each farm that encompassed four vegetation types (plantings 7-20 years old, old-growth woodland, naturally occurring seedling regrowth woodland, and coppice [i.e., multistemmed] regrowth woodland). Reptiles and arboreal marsupials were less likely to occur on farms and in landscapes with comparatively large areas of plantings. Such farms and landscapes had less native vegetation, fewer paddock trees, and less woody debris within those areas of natural vegetation. The relatively large area of planting on these farms was insufficient to overcome the lack of these key structural attributes. Old-growth woodland, coppice regrowth, seedling regrowth, and planted areas had different habitat values for different reptiles and arboreal marsupials. We conclude that, although plantings may improve habitat conditions for some taxa, they may not effectively offset the negative effects of native vegetation clearing for all species, especially those reliant on old-growth woodland. Restoring suitable habitat for such species may take decades to centuries.     

Simulated predator extinctions: predator identity affects survival and recruitment of oysters

The rate of species loss is increasing at a global scale, and human-induced extinctions are biased toward predator species. We examined the effects of predator extinctions on a foundation species, the eastern oyster (Crassostrea virginica). We performed a factorial experiment manipulating the presence and abundance of three of the most common predatory crabs, the blue crab (Callinectes sapidus), stone crab (Menippe mercenaria), and mud crab (Panopeus herbstii) in estuaries in the eastern United States. We tested the effects of species richness and identity of predators on juvenile oyster survival, oyster recruitment, and organic matter content of sediment. We also manipulated the density of each of the predators and controlled for the loss of biomass of species by maintaining a constant mass of predators in one set of treatments and simultaneously using an additive design. This design allowed us to test the density dependence of our results and test for functional compensation by other species. The identity of predator species, but not richness, affected oyster populations. The loss of blue crabs, alone or in combination with either of the other species, affected the survival rate of juvenile oysters. Blue crabs and stone crabs both affected oyster recruitment and sediment organic matter negatively. Mud crabs at higher than ambient densities, however, could fulfill some of the functions of blue and stone crabs, suggesting a level of ecological redundancy. Importantly, the strong effects of blue crabs in all processes measured no longer occurred when individuals were present at higher-than-ambient densities. Their role as dominant predator is, therefore, dependent on their density within the system and the density of other species within their guild (e.g., mud crabs). Our findings support the hypothesis that the effects of species loss at higher trophic levels are determined by predator identity and are subject to complex intraguild interactions that are largely density dependent. Understanding the role of biodiversity in ecosystem functioning or addressing practical concerns, such as loss of predators owing to overharvesting, remains complicated because accurate predictions require detailed knowledge of the system and should be drawn from sound experimental evidence, not based on observations or generalized models.     

Neutral modelling of agricultural landscapes by tessellation methods—Application for gene flow simulation

Neutral landscape models are not frequently used in the agronomical domain, whereas they would be very useful for studying given agro-ecological or physical processes. Contrary to ecological neutral landscape models, agricultural models have to represent and manage geometrical patches and thus should rely on tessellation methods. We present a three steps approach that aimed at simulating such landscapes. Firstly, we characterized the geometry of three real field patterns; secondly, we generated simulated field patterns with two tessellation methods attempting to control the value of some of the observed characteristics and, thirdly, we evaluated the simulated field patterns. For this evaluation, we considered that good simulated field patterns should capture characteristics of real landscapes that are important for the targeted agro-ecological process. Real landscapes and landscapes simulated using either a Voronoi or a rectangular tessellation were thus compared when used as input data within a gene flow model. The results showed that neither tessellation method captured field shapes correctly, thus leading to over or (small) under estimation of gene flow. The Voronoi tessellation, though, performed better than the rectangular tessellation. Possible research directions are proposed to improve the simulated patterns, including the use of post-processing, the control of cell orientation or the implementation of other tessellation techniques.     

Fire history and tree recruitment in the Colorado Front Range upper montane zone: implications for forest restoration

Forests experiencing moderate- or mixed-severity fire regimes are presumed to be widespread across the western United States, but few studies have characterized these complex disturbance regimes and their effects on contemporary forest structure. Restoration of pre-fire-suppression open-forest structure to reduce the risk of uncharacteristic stand-replacing fires is a guiding principle in forest management policy, but identifying which forests are clear candidates for restoration remains a challenge. We conducted dendroecological reconstructions of fire history and stand structure at 40 sites in the upper montane zone of the Colorado Front Range (2400-2800 m), sampled in proportion to the distribution of forest types in that zone (50% dominated by ponderosa pine, 28% by lodgepole pine, 12% by aspen, 10% by Douglas-fir). We characterized past fire severity based on remnant criteria at each site in order to assess the effect of fire history on tree establishment patterns, and we also evaluated the influence of fire suppression and climate. We found that 62% of the sites experienced predominantly moderate-severity fire, 38% burned at high severity, and no sites burned exclusively at low severity. The proportion of total tree and sapling establishment was significantly different among equal time periods based on a chi-square test, with highest tree and sapling establishment during the pre-fire-suppression period (1835-1919). Superposed epoch analysis revealed that fires burned during years of extreme drought (95% CI). The major pulse of tree establishment in the upper montane zone occurred during a multidecadal period of extreme drought conditions in the Colorado Front Range (1850-1889), during which 53% of the fires from the 1750-1989 period burned. In the upper montane zone of the Colorado Front Range, historical evidence suggests that these forests are resilient to prolonged periods of severe drought and associated severe fires.     

Spatiotemporal variation in survival rates: implications for population dynamics of yellow-bellied marmots

Spatiotemporal variation in age-specific survival rates can profoundly influence population dynamics, but few studies of vertebrates have thoroughly investigated both spatial and temporal variability in age-specific survival rates. We used 28 years (1976-2003) of capture-mark-recapture (CMR) data from 17 locations to parameterize an age-structured Cormack-Jolly-Seber model, and investigated spatial and temporal variation in age-specific annual survival rates of yellow-bellied marmots (Marmota flaviventris). Survival rates varied both spatially and temporally, with survival of younger animals exhibiting the highest degree of variation. Juvenile survival rates varied from 0.52 +/- 0.05 to 0.78 +/- 0.10 among sites and from 0.15 +/- 0.14 to 0.89 +/- 0.06 over time. Adult survival rates varied from 0.62 +/- 0.09 to 0.80 +/- 0.03 among sites, but did not vary significantly over time. We used reverse-time CMR models to estimate the realized population growth rate (lamda), and to investigate the influence of the observed variation in age-specific survival rates on lamda. The realized growth rate of the population closely covaried with, and was significantly influenced by, spatiotemporal variation in juvenile survival rate. High variability in juvenile survival rates over space and time clearly influenced the dynamics of our study population and is also likely to be an important determinant of the spatiotemporal variation in the population dynamics of other mammals with similar life history characteristics.     

Patterns of animal diversity in different forms of tree cover in agricultural landscapes.

As tropical regions are converted to agriculture, conservation of biodiversity will depend not only on the maintenance of protected forest areas, but also on the scope for conservation within the agricultural matrix in which they are embedded. Tree cover typically retained in agricultural landscapes in the neotropics may provide resources and habitats for animals, but little is known about the extent to which it contributes to conservation of animal species. Here, we explore the animal diversity associated with different forms of tree cover for birds, bats, butterflies, and dung beetles in a pastoral landscape in Nicaragua. We measured species richness and abundance of these four animal taxa in riparian and secondary forest, forest fallows, live fences, and pastures with high and low tree cover. We recorded over 20,000 individuals of 189 species including 14 endangered bird species. Mean abundance and species richness of birds and bats, but not dung beetles or butterflies, were significantly different among forms of tree cover. Species richness of bats and birds was positively correlated with tree species richness. While the greatest numbers of bird species were associated with riparian and secondary forest, forest fallows, and pastures with >15% tree cover, the greatest numbers of bat species were found in live fences and riparian forest. Species assemblages of all animal taxa were different among tree cover types, so that maintaining a diversity of forms of tree cover led to conservation of more animal species in the landscape as a whole. Overall, the findings indicate that retaining tree cover within agricultural landscapes can help conserve animal diversity, but that conservation efforts need to target forms of tree cover that conserve the taxa that are of interest locally. Preventing the degradation of remaining forest fragments is a priority, but encouraging farmers to maintain tree cover in pastures and along boundaries may also make an important contribution to animal conservation.     

Effects of incubation temperature on hatchling pine snakes: implications for survival

Incubation temperature in ectothermic vertebrates affects incubation periods, and in some reptiles it affects sex ratios and behavior. I present evidence that incubation temperature affects emergence and post-hatching behavior of pine snakes (Pituophis melanoleucus) that could influence survival in the weeks before hibernation. Hatchlings incubated at low temperatures remained in the nest longer, had fewer alternate nest openings, and fewer underground tunnels to hide in than did hatchlings from warmer temperatures. These conditions could render hatchlings from low-temperature nests more vulnerable to predation because, if a nest is opened, they are not inside tunnels where they would be protected. Hatchlings from nests incubated at low temperatures took longer to find shade during a thermoregulation test, and were less likely to move about in search of other cover than were those from higher-incubation-temperature artificial nests. Similarly, hatchlings from nests with low incubation temperatures were less responsive to a predatory stimulus and had a longer latency to strike than other hatchlings. Taken together, hatchlings from nests with low incubation temperatures might be less able to avoid predators and find shade than those from nests incubated at higher temperatures, and thus could be expected to have lower survival in nature. Received: 21 July 1997 / Accepted after revision: 15 February 1998     

Community recruitment: Settlement and juvenile survival of seven co-occurring species of sessile marine invertebrates

Recruitment of a group of co-occurring sessile invertebrates (a serpulid polychaete, an oyster, a bryozoan, and several colonial and one solitary ascidian) that encrust floating docks in Pearl Harbor, Hawaii, was studied in August 1985. Daily photographs were taken of acrylic settling plates over a 14 d period, and daily settlement and juvenile mortality were measured. Settlement rates were compared between species and days, while juvenile mortalities were compared between species, days, juvenile ages and densities of juveniles on the plates. Species differed in abundance of both settlers and juveniles, and in rate and pattern of juvenile mortality. Settlement intensity varied between days for some species. Significant juvenile mortality occurred during the 14 d for most species; it appeared to be caused by fish predation. For two species, mortality varied with juvenile age, with older individuals suffering higher mortality. Mortality was density-dependent for some species, increasing with numbers of juveniles on the plates. Mortality patterns varied even within the colonial species. Assessing larval settlement and juvenile mortality for a single species or type of organism in a community thus may not indicate that similar patterns exist for co-occurring species.     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

Spatiotemporal variability in recruitment around Iriomote Island, Ryukyu Archipelago, Japan: implications for dispersal of spawning corals

Spatiotemporal recruitment patterns of scleractinian corals were investigated around Iriomote Island, Ryukyu Archipelago, Japan, in relation to adult coral cover in 2005 and 2006. Although almost all corals were broadcasting spawners, the relationship between recruitment and adult coral cover differed among coral families (Acroporidae, Poritidae, and Pocilloporidae), likely due to differences in embryonic development time. For spawning pocilloporid corals, whose larvae develop relatively more rapidly, recruitment was higher at sites where adult coral cover was higher. In contrast, recruitment was not related to adult coral cover in acroporid and poritid corals, whose embryonic development times were relatively slow. Moreover, recruitment of acroporid corals varied between years, and recruitment was greater at leeward compared to windward reefs for a few days after spawning. These results suggest that embryonic development time and wind-driven surface currents affect larval dispersal and subsequent recruitment patterns at a local scale. Based on embryonic development time, some spawning corals are more likely to have higher rates of self-seeding than others. Our results predict that among spawning corals, local populations of acroporid and poritid corals, whose larvae potentially disperse over long distances and recruit in neighboring reefs, are more resilient to local disturbances than those of pocilloporid corals, whose recruitment relies upon local stock.     

Dynamics and driving forces of agricultural landscapes in Southern Ethiopia – a case study of the Chencha and Arbaminch areas

Agricultural landscapes in Ethiopia have undergone unprecedented changes. The direction of change, however, is unsustainable as manifested in land degradation, biodiversity loss, and low agricultural productivity. The objective of this study is to examine the patterns and trends of agricultural landscape development and responses of the local people within the framework of the dynamics of demography, socioeconomic conditions, politics, and natural resources in the Chencha and Arbaminch areas, Southern Ethiopia, during the last century. Information on cultivated and grazing land areas was acquired by satellite image interpretation. Interviews and group discussions provided important information on agricultural land use systems. A review and an analysis of secondary sources and documents of past studies were also used for trend analysis as a baseline and a supplement to oral history. The results show that cultivated land was expanded by 39% from 1973 until 2006, but per capita farming land holdings decreased enormously. In the same period of time, grassland shrank by 69% thus causing a significant decrease in livestock. Cultivated land scarcity can mostly be related to demographic pressure, which was exacerbated by government policy, land tenure, and the nature of subsistence agriculture. The farmers, however, were resourceful and developed skills over millennia to cope with the problems associated with population density and scarce resources. However, these traditional land use activities and land management practices have been deteriorating recently. Land use planners and environmental managers should take local knowledge and innovation into account in order to make sound decisions for the future.     

Fragmentation profiles for real and simulated landscapes

When a natural landscape is represented by a series of categorical raster maps of varying resolution, a multiresolution characterization of spatial pattern can be obtained in which entropy is computed at each resolution conditional on the next coarser resolution. The series of entropy values is plotted as a function of resolution, resulting in a multiresolution profile of fragmentation pattern in the landscape. If a categorical raster map is available at a single resolution only, a series of degraded maps at increasingly coarser resolutions is generated and the fragmentation profile is computed for this series. An algorithm has been developed for obtaining the profile directly from the single resolution map without having to generate and store the coarser resolution maps. A hierarchical stochastic model is described for simulating categorical raster maps and the fragmentation profile of the generating process is obtained in terms of the model parameters. These process profiles provide benchmarks for assessing empirical profiles obtained from raster maps of actual landscapes. Methods of the paper are applied to several watersheds of Pennsylvania using landcover maps derived from satellite imagery. These examples indicate that characteristic landscape types induce characteristic features in their fragmentation profiles.     

The invasive gastropod Crepidula fornicata: reproduction and recruitment in the intertidal at its northernmost range in Wales, UK, and implications for its secondary spread

The establishment and spread of a non-native species in an introduced range depends to a large extent on the performance of the species under the prevailing environmental conditions. The spawning, larval and spatfall periods of the invasive gastropod Crepidula fornicata were monitored in the intertidal zone at its northernmost range in Wales, UK, between February 2010 and January 2011. The duration of the reproductive season was similar to that recorded from more southerly European populations. Spawning and larval release occurred throughout most of the year even at low seawater temperatures of <7 °C, but benthic recruitment was observed over a much shorter period at seawater temperatures >16 °C. Recruitment was low and likely controlled by post-settlement mortality. These observations suggest that C. fornicata’s northwards spread in Welsh waters will not be limited by seawater temperature negatively affecting reproduction, but by processes acting after larval release. These data show the importance of incorporating settlement and post-settlement processes into studies on recruitment success when aiming to predict the potential spread of a potentially harmful invader such as C. fornicata.     

Effects of anaerobiosis on root metabolism of Zostera marina (eelgrass): implications for survival in reducing sediments

The temperate seagrass Zostera marina L. typically grows in highly reducing sediments. Photosynthesis-mediated O₂ supplied to below-ground tissues sustains aerobic respiration during photosynthetic periods. Roots, however, experience daily periods of anoxia and/or hypoxia at night and under conditions that reduce photosynthesis. Rhizosphere cores of Z. marina were collected in August 1984 from Great Harbor, Massachusetts, USA. We examined short-term anaerobic metabolism of [U-¹⁴C]sucrose in excised roots and roots of intact plants. Under anaerobic conditions roots showed appreciable labeling of CO₂, ethanol and lactate, and slight labeling of alanine and other metabolites. Over 95% of the ¹⁴C-ethanol was recovered in the root exudate. Release of other metabolites from the roots was minimal. Ethanol was also released from hypoxic/anoxic roots of intact plants and none of this ethanol was transported to the shoot under any experimental conditions. Loss of ethanol from roots prevented tissue levels of this phytotoxin from increasing during anaerobiosis despite increased synthesis of ethanol. Anaerobic metabolism of [U-¹⁴C]glutamate in excised roots led to appreciable labelling of -aminobutyrate, which was known to accumulate in eelgrass roots. Roots recovered to fully aerobic metabolism within 4 h after re-establishment of aerobic conditions. The contributions of these root metabolic responses to the ability of Z. marina to grow in reducing marine sediments are related to light-regulated interactions of shoots and roots that likely dictate depth penetration, distribution and ecological success of eelgrass.