Conservation Benefits of Temperate Marine Protected Areas: Variation among Fish Species

Abstract:  Marine protected areas, and other fishery management systems that impart partial or total protection from fishing, are increasingly advocated as an essential management tool to ensure the sustainable use of marine resources. Beneficial effects for fish species are well documented for tropical and reef systems, but the effects of marine protected areas remain largely untested in temperate waters. We compared trends in sport-fishing catches of nine fish species in an area influenced by a large (500-km²) towed-fishing-gear restriction zone and in adjacent areas under conventional fishery management controls. Over the period 1973–2002 the mean reported weight of above-average-sized (trophy) fish of species with early age at maturity and limited home range was greatest within the area influenced by the fishing-gear restriction zone. The reported weight of trophy fish of species that mature early also declined less and more slowly over time within the area influenced by the fishing-gear restriction zone. Importantly, the mean reported weight of trophy fish of species that mature late and those that undertake extensive spatial movements declined at the same rate in all areas. Hence these species are likely to require protected areas >500 km² for effective protection. Our results also indicated that fish species with a localized distribution or high site fidelity may require additional protection from sport fishing to prevent declines in the number or size of fish within the local population.     

Invasion in a diversity hotspot: exotic cover and native richness in the Californian serpentine flora

Exotic species have been observed to be more prevalent in sites where the richness of native species is highest, possibly reflecting variation among sites in resources, propagule supply, heterogeneity, or disturbance. However, such a pattern leaves unclear whether natives at species-rich sites are subject to especially severe impacts from exotics as a result. We considered this question using path models in which relationships between exotic cover and native richness were evaluated in the presence of correlated environmental factors. At 109 sites on serpentine soils across California, USA, exotic cover was positively correlated with total native herbaceous richness and was negatively correlated with the richness of both serpentine-endemic and rare native herbs. However, in path models that accounted for the influences of soil chemistry, disturbance, overstory cover, and regional rainfall and elevation, we found no indication that exotic cover reduced any component of native herb richness. Rather, our results indicated similarities and differences in the conditions favoring exotic, native, endemic, and rare species. Our results suggest that, in spite of some localized impacts, exotic species are not exerting a detectable overall effect on the community richness of the unique native flora of Californian serpentine.     

Potential nitrogen constraints on soil carbon sequestration under low and elevated atmospheric CO2

The interaction between nitrogen cycling and carbon sequestration is critical in predicting the consequences of anthropogenic increases in atmospheric CO2 (hereafter, Ca). The progressive N limitation (PNL) theory predicts that carbon sequestration in plants and soils with rising Ca may be constrained by the availability of nitrogen in many ecosystems. Here we report on the interaction between C and N dynamics during a four-year field experiment in which an intact C3/C4 grassland was exposed to a gradient in Ca from 200 to 560 micromol/mol. There were strong species effects on decomposition dynamics, with C loss positively correlated and N mineralization negatively correlated with Ca for litter of the C3 forb Solanum dimidiatum, whereas decomposition of litter from the C4 grass Bothriochloa ischaemum was unresponsive to Ca. Both soil microbial biomass and soil respiration rates exhibited a nonlinear response to Ca, reaching a maximum at approximately 440 micromol/mol Ca. We found a general movement of N out of soil organic matter and into aboveground plant biomass with increased Ca. Within soils we found evidence of C loss from recalcitrant soil C fractions with narrow C:N ratios to more labile soil fractions with broader C:N ratios, potentially due to decreases in N availability. The observed reallocation of N from soil to plants over the last three years of the experiment supports the PNL theory that reductions in N availability with rising Ca could initially be overcome by a transfer of N from low C:N ratio fractions to those with higher C:N ratios. Although the transfer of N allowed plant production to increase with increasing Ca, there was no net soil C sequestration at elevated Ca, presumably because relatively stable C is being decomposed to meet microbial and plant N requirements. Ultimately, if the C gained by increased plant production is rapidly lost through decomposition, the shift in N from older soil organic matter to rapidly decomposing plant tissue may limit net C sequestration with increased plant production.     

Recruitment of Hexagenia mayfly nymphs in western Lake Erie linked to environmental variability.

After a 40-year absence caused by pollution and eutrophication, burrowing mayflies (Hexagenia spp.) recolonized western Lake Erie in the mid 1990s as water quality improved. Mayflies are an important food resource for the economically valuable yellow perch fishery and are considered to be major indicator species of the ecological condition of the lake. Since their reappearance, however, mayfly populations have suffered occasional unexplained recruitment failures. In 2002, a failure of fall recruitment followed an unusually warm summer in which western Lake Erie became temporarily stratified, resulting in low dissolved oxygen levels near the lake floor. In the present study, we examined a possible link between Hexagenia recruitment and periods of intermittent stratification for the years 1997 2002. A simple model was developed using surface temperature, wind speed, and water column data from 2003 to predict stratification. The model was then used to detect episodes of stratification in past years for which water column data are unavailable. Low or undetectable mayfly recruitment occurred in 1997 and 2002, years in which there was frequent or extended stratification between June and September. Highest mayfly reproduction in 2000 corresponded to the fewest stratified periods. These results suggest that even relatively brief periods of stratification can result in loss of larval mayfly recruitment, probably through the effects of hypoxia. A trend toward increasing frequency of hot summers in the Great Lakes region could result in recurrent loss of mayfly larvae in western Lake Erie and other shallow areas in the Great Lakes.     

Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.     

Serengeti wildebeest migratory patterns modeled from rainfall and new vegetation growth

We used evolutionary programming to model innate migratory pathways of wildebeest in the Serengeti Mara Ecosystem, Tanzania and Kenya. Wildebeest annually move from the southern short-grass plains of the Serengeti to the northern woodlands of the Mara. We used satellite images to create 12 average monthly and 180 10-day surfaces from 1998 to 2003 of percentage rainfall and new vegetation. The surfaces were combined in five additive and three multiplicative models, with the weightings on rainfall and new vegetation from 0% to 100%. Modeled wildebeest were first assigned random migration pathways. In simulated generations, animals best able to access rainfall and vegetation were retained, and they produced offspring with similar migratory pathways. Modeling proceeded until the best pathway was stable. In a learning phase, modeling continued with the ten-day images in the objective function. The additive model, influenced 25% by rainfall and 75% by vegetation growth, yielded the best agreement, with a multi-resolution comparison to observed densities yielding 76.8% of blocks in agreement (kappa = 0.32). Agreement was best for dry season and early wet season (kappa = 0.22-0.57), and poorest for the late wet season (0.04). The model suggests that new forage growth is a dominant correlate of wildebeest migration.     

Growth inhibition in Japanese medaka (Oryzias latipes) fish exposed to tetrachloroethylene

A recent study in our laboratory has demonstrated that tetrachloroethylene (TCE) is acutely toxic to Japanese medaka (Oryzias latipes) larvae with a 96 hr-LC50 of 18 (17-19) mg/mL (Spencer et al., 2002). In the present study we hypothesize that TCE exposure induces a developmental effect in Japanese medaka. Growth and age specific sensitivity of Japanese medaka larvae were studied with four age groups (7, 14, 21 and 28 days old) to determine tetrachloroethylene effects on these parameters. The medaka larvae were exposed for 96 hours in a single concentration (10 mg/mL) of TCE. The toxic endpoints evaluated were larvae weight, length, water content and protein concentration. The study revealed that exposure of medaka larvae to this sub-acute concentration of TCE significantly reduced length and weight in the treated group. The difference in growth between control and treated groups was more obvious in age versus length, than in age versus weight. The dry weight-fresh weight ratio (dw/fw) was shown to be higher in the control group. Water content in TCE-treated medaka was higher than in the control group, and younger fry had more water content than older ones. A higher protein concentration was also observed in TCE-treated medaka compared to the control group. These results indicate that TCE has a profound effect on the growth and development of Japanese medaka larvae.     

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.     

In situ experiment of ontogenetic variability in the otolith chemistry of Stegastes partitus

Otolith chemistry can be used to assess pelagic larval fish connectivity by comparing spatially variable otolith edge chemistry (corresponding to the site of collection) to otolith core chemistry (corresponding to the site of hatching). However, because the otolith’s edge and core represent different life stages, the deposition of elements may differ, thus complicating direct comparisons of edge and core chemistry to investigate connectivity. Here we present data from a field experiment in which otoliths from embryos (3 days post-fertilization) and juveniles of Stegastes partitus were collected at the same site and time, and chemically analyzed to assess whether elemental concentrations of otoliths vary ontogenetically. Separate multivariate analyses, each investigating the spatial/temporal variability in the chemistry of either embryo otoliths or the edges of juvenile otoliths, revealed significant differences, suggesting an environmental influence to the chemical signals of otoliths. A nested multivariate analysis assessing whether otolith chemistry varied with life history stage (i.e., ontogenetic variability) indicated that elemental concentrations of embryo otoliths were significantly greater than that of juvenile otolith edges. Specifically, embryo elemental concentrations of Mn, Zn, Sn, Ba, Ce, and Pb were between 2 and 163 times greater than those of the corresponding juvenile otoliths, and thus the environment was not the primary determinant of embryo otolith chemistry. Consequently, caution is warranted when interpreting environmental patterns of otolith cores, particularly when using them as a proxy for natal signatures.