Dramatic declines in mussel bed community diversity: response to climate change?

Mussel beds along the wave-exposed coast of the eastern North Pacific Ocean serve as an important habitat, harboring a high diversity of species. A comparison of California mussel bed community diversity in 2002 to historical data (1960s to 1970s) revealed large declines (mean loss 58.9%), including some declines >141 species (approximately 80% loss). Concurrent work revealed inconsistent changes in mussel populations (biomass and bed thickness) along the California coast, suggesting that diversity declines may be related to large-scale processes rather than local habitat destruction. Potential factors causing declines in mussel community diversity are discussed, with regional climate change associated with the Pacific Decadal Oscillation and climate change induced alterations of ecological interactions and biological processes suggested as likely causes. Although extensive literature has predicted the potential effects of climate change on global diversity, this study is one of the few examples of declines attributed to climate change.     

The effects of experimental bait collection and trampling on a<Emphasis Type="Italic"> Mytilus californianus</Emphasis> mussel bed in southern California

Rocky shores in southern California are heavily visited by humans. At sites used by recreational fishers, the effects of foot traffic combined with the collection of mussels for bait may reduce mussel cover and create mussel-free gaps. To test this hypothesis, the effects of trampling and bait-removal on mussel populations were experimentally examined. Plots in a mussel bed were subjected to monthly combinations of trampling (0, 150, or 300 steps) and simulated bait-removal (0 or 2 removed mussels). Although the experiment was done during a period of high natural disturbance associated with the 1997–1998 ENSO, plots receiving treatments experienced significantly greater reductions in mussel cover, mass, and density than controls. These results indicate that visitor foot traffic and bait-removal by fishers can significantly reduce mussel cover, density, biomass, and sizes.Communicated by P.W. Sammarco, Chauvin     

Conditions for coexistence of freshwater mussel species via partitioning of fish host resources

Riverine freshwater mussel species can be found in highly diverse communities where many similar species coexist. Mussel species potentially compete for food and space as adults, and for fish host resources during the larval (glochidial) stage. Resource partitioning at the larval stage may promote coexistence. A model of resource utilization was developed for two mussel species and analyzed to determine conditions for coexistence. Mussel species were predicted to coexist when they differed in terms of their success in contacting different fish host species; very similar strategies offered limited possibilities for coexistence. Differences in the mussel species’ maximum infestation loads on the fish hosts that coincided with differences in their fish host contact success promoted coexistence. Mussel species with a given set of trade-offs in fish host use were predicted to coexist only for a subset of relative fish host abundances, so a shift in relative fish host abundances could result in the loss of a mussel species. An understanding of the conditions for freshwater mussel species coexistence can help explain high mussel diversity in rivers and guide ongoing conservation activities.     

Comparison of mussel bed community between two intertidal mytilids Septifier virgatus and Hormomya mutabilis

Mussel bed community structure of two intertidal mytilids, Septifer virgatus (Wiegmann) and Hormomya mutabilis (Gould), whose beds were contiguous vertically on a rocky intertidal shore (Wakayama Prefecture, Japan), was compared between 1982 and 1983. In the upper S. virgatus bed, crustaceans and bivalves were dominant in terms of both number of individuals and biomass. There were three barnacle species representative of the epizoans, two isopods and one amphipod as mobile fauna, and two bivalves as infauna. The lower H. mutabilis bed supported virtually no epizoans or mobile fauna. Infaunal free-ranging polychaetes and sipunculids were dominant in terms of both number of individuals and biomass. The H. mutabilis bed contained a much greater amount of sediment than did the S. virgatus bed, and the interstices among individual H. mutabilis and among their byssal threads were filled with sediment. The biomass of six of nine species dominant in the S. virgatus bed was negatively correlated with the amount of sediment. Recruitment of these faunal assemblages into artificial mussel clumps was examined in mussel enclosure experiments, and a negative sediment effect in H. mutabilis clumps was detected for one isopod and one limpet species. Factors causing the differences between two mussel bed communities are discussed, focusing on the effects of sediment.     

Influence of wave exposure on South African mussel beds and their associated infaunal communities

This study investigates effects of wave exposure on beds of the mussel Mytilus galloprovincialis and the infaunal communities associated with them. Nine sites of varying wave exposure were sampled near Groenrivier, on the west coast of South Africa. Wave exposure was measured using a Palumbi device, and exposure ranged from 7.17×10³ to 18.49×10³ N m^–2. Percentage mussel cover at each site was measured using 50×50 cm quadrats, and three 10×10 cm samples were removed from the mussel beds in the mid-intertidal zones of each site for examination of infauna. Percentage mussel cover, mean mussel length, mussel biomass and mussel bed depth all peaked at intermediate exposures, declining towards both the most sheltered and most exposed sites. Infaunal species diversity and richness both showed the reverse trend, peaking at the most sheltered and most exposed sites, and declining at intermediate exposures. Neither infaunal abundance, nor biomass, was significantly correlated to wave exposure, although abundance was highest at sheltered sites and biomass greatest at the most exposed site, which was dominated by limpets and large robust polychaetes.Communicated by J.P. Thorpe, Port Erin     

Wave disturbance overwhelms top-down and bottom-up control of primary production in California kelp forests

We took advantage of regional differences in environmental forcing and consumer abundance to examine the relative importance of nutrient availability (bottom-up), grazing pressure (top-down), and storm waves (disturbance) in determining the standing biomass and net primary production (NPP) of the giant kelp Macrocystis pyrifera in central and southern California. Using a nine-year data set collected from 17 sites we show that, despite high densities of sea urchin grazers and prolonged periods of low nutrient availability in southern California, NPP by giant kelp was twice that of central California where nutrient concentrations were consistently high and sea urchins were nearly absent due to predation by sea otters. Waves associated with winter storms were consistently higher in central California, and the loss of kelp biomass to winter wave disturbance was on average twice that of southern California. These observations suggest that the more intense wave disturbance in central California limited NPP by giant kelp under otherwise favorable conditions. Regional patterns of interannual variation in NPP were similar to those of wave disturbance in that year-to-year variation in disturbance and NPP were both greater in southern California. Our findings provide strong evidence that regional differences in wave disturbance overwhelmed those of nutrient supply and grazing intensity to determine NPP by giant kelp. The important role of disturbance in controlling NPP revealed by our study is likely not unique to giant kelp forests, as vegetation dynamics in many systems are dominated by post-disturbance succession with climax communities being relatively uncommon. The effects of disturbance frequency may be easier to detect in giant kelp because it is fast growing and relatively short lived, with cycles of disturbance and recovery occurring on time scales of years. Much longer data sets (decades to centuries) will likely be needed to properly evaluate the role of disturbance relative to other processes in determining patterns of NPP in other systems.     

Decline of a Native Mussel Masked by Sibling Species Invasion

Abstract: The European blue mussel (  Mytilus galloprovincialis ) has extensively invaded southern California, whereas a native species, M. trossulus , is abundant in northern California and further north. In this study, a portion of the 16S ribosomal RNA gene was sequenced from mussels collected in southern and central California in the nineteenth century. I aligned these sequences with 22 sequences representing modern M. galloprovincialis , M. edulis , and M. trossulus . Phylogenetic analysis using parsimony identified the nineteenth-century mussels as M. trossulus . Because mussel populations have been continuously present in this region since the time these mussels were collected, these data indicate that M. trossulus were present during the initial invasion of M. galloprovincialis , but, due to the morphological similarity of the two species, declined without notice.     

Regional extinction of a conspicuous dorid nudibranch (Mollusca: Gastropoda) in California

Felimare californiensis (=Hypselodoris californiensis) was once common throughout the Southern California Bight (SCB) and California Channel Islands. This well-known shallow-water nudibranch, which specializes on dysideid sponges, has persisted for decades in Mexico, but in California disappeared from its entire range by 1984. Since reappearing in 2003, it has been found only at Santa Catalina Island, plus sightings of single individuals in 2011 at Santa Cruz Island and San Diego. The decline of F. californiensis in California was documented using published historical records, museum collections, unpublished field accounts, and images posted online. The loss of this emblematic species is unique among Californian nudibranchs, including (1) its congener Felimare porterae (=Mexichromis porterae), with which it appears to overlap in diet, and (2) opisthobranch species with similar historical geographic ranges and mode of development. The decline in F. californiensis is not predicted by warming trends and climate variation over the past 40 years, including the strong El Niño events of 1983 and 1998. Coastal pollution from the large human population in southern California may have impacted Dysidea amblia, the primary reported prey of F. californiensis. Historical overcollecting of the nudibranch and habitat loss through the development of major ports likely also contributed to its decline. Sightings since 2003 are consistent with a nascent recovery, as elements of water quality have improved in the SCB in recent decades.     

Nucleic acids and ATP to assess microbial biomass and activity in a marine biosedimentary system

 Sedimentary DNA, RNA and ATP concentrations were measured at six stations on a tidal flat in the German Wadden Sea from April to December 1995 to determine factors controlling microbial abundance and activity. Two stations (1 and 2) were located in a mussel bed (Mytilus edulis) in the middle of the flat, and the other four were arranged in a line following ebb tide direction. DNA and ATP concentrations were converted into carbon equivalents using literature coefficients and considered to represent total microbial biomass and viable microbial biomass, respectively. The difference between C_DNA and C_ATP was considered as detrital biomass. C_DNA and detrital C_DNA biomass both increased from spring to summer and decreased in autumn to reach lowest values in December. The most prominent seasonal increase was observed at the mussel bed stations and two adjacent stations. C_ATP biomass peaked in early spring, especially at the mussel bed stations, and showed a secondary peak in summer. Regression analysis yielded a highly significant relation between the silt content of the sediment and C_DNA and C_ATP. However, C_DNA and C_ATP biomass are not correlated if silt is taken into account. The relation between silt and microbial biomass in combination with the decrease of the silt content with distance from the mussel bed indicate the importance of mussel beds for the microbial activity on the tidal flat, most likely through deposition of feces and pseudofeces. The low silt content and microbial biomass in winter are attributed to resuspension events. The zero intercepts of the regressions between DNA and ATP or RNA suggest that extracellular DNA is absent in our samples. Hence, detrital C_DNA biomass in our samples probably consists of intact but dormant cells. Compared to other marine habitats the percentage of detrital C_DNA biomass on the tidal flat is quite high, i.e. up to >95% of the total C_DNA. However some doubt is expressed about the validity of factors used to convert ATP and DNA to carbon. The proportion of active biomass, expressed by the ratio C_ATP/C_DNA being the complement of the detrital C_DNA percentage shows a steady decrease with time and with diminishing distance towards the mussel beds. The role of a mussel bed as a biosedimentary system influencing spatial and temporal trends in specific activity is discussed. Received: 26 January 2000 / Accepted: 3 July 2000     

Density-dependent biodiversity effects on physical habitat modification by freshwater bivalves

Several decades of research have shown that biodiversity affects ecosystem processes associated with resource capture and the production of biomass within trophic levels. Although there are good reasons to expect that biodiversity influences non-trophic ecosystem processes, such as the physical creation or modification of habitat, studies investigating the role of biodiversity on physical processes are scarce. Here we report the results of a study using artificial streams to test the influence of freshwater mussel biodiversity on gravel erosion during high flows while manipulating mussel abundance. Mussel species vary in traits that should influence their effects on erosion, such as size, shell morphology, and burrowing behavior. We found that mussel species richness was associated with an increase in erosion at both low and high densities. Planned contrasts showed that the erosion observed in species mixtures was purely additive at low density, indicating that erosion in a species polyculture could routinely be predicted by the performance of monocultures. However, at high density certain combinations of species showed nonadditive effects on erosion, suggesting that organism abundance can fundamentally alter biodiversity effects. Although this may have been a result of altered species interactions at high density, our study design cannot confirm this.     

Islands at bay: rising seas,eroding islands,and waterbird habitat loss in Chesapeake Bay (USA)

Like many resources in the Chesapeake Bay region of the U.S., many waterbird nesting populations have suffered over the past three to four decades. In this study, historic information for the entire Bay and recent results from the Tangier Sound region were evaluated to illustrate patterns of island erosion and habitat loss for 19 breeding species of waterbirds. Aerial imagery and field data collected in the nesting season were the primary sources of data. From 1993/1994 to 2007/2008, a group of 15 islands in Tangier Sound, Virginia were reduced by 21% in area, as most of their small dunes and associated vegetation and forest cover were lost to increased washovers. Concurrently, nesting American black ducks (Anas rubripes) declined by 66% , wading birds (herons-egrets) by 51%, gulls by 72%, common terns (Sterna hirundo) by 96% and black skimmers (Rynchops niger) by about 70% in this complex. The declines noted at the larger Bay-wide scale suggest that this study area maybe symptomatic of a systemic limitation of nesting habitat for these species. The island losses noted in the Chesapeake have also been noted in other Atlantic U.S. coastal states. Stabilization and/or restoration of at least some of the rapidly eroding islands at key coastal areas are critical to help sustain waterbird communities.     

Remarkable Amphibian Biomass and Abundance in an Isolated Wetland: Implications for Wetland Conservation

Abstract:  Despite the continuing loss of wetland habitats and associated declines in amphibian populations, attempts to translate wetland losses into measurable losses to ecosystems have been lacking. We estimated the potential productivity from the amphibian community that would be compromised by the loss of a single isolated wetland that has been protected from most industrial, agricultural, and urban impacts for the past 54 years. We used a continuous drift fence at Ellenton Bay, a 10-ha freshwater wetland on the Savannah River Site, near Aiken, South Carolina (U.S.A.), to sample all amphibians for 1 year following a prolonged drought. Despite intensive agricultural use of the land surrounding Ellenton Bay prior to 1951, we documented 24 species and remarkably high numbers and biomass of juvenile amphibians (>360,000 individuals; >1,400 kg) produced during one breeding season. Anurans (17 species) were more abundant than salamanders (7 species), comprising 96.4% of individual captures. Most (95.9%) of the amphibian biomass came from 232095 individuals of a single species of anuran (southern leopard frog [Rana sphenocephala ]). Our results revealed the resilience of an amphibian community to natural stressors and historical habitat alteration and the potential magnitude of biomass and energy transfer from isolated wetlands to surrounding terrestrial habitat. We attributed the postdrought success of amphibians to a combination of adult longevity (often >5 years), a reduction in predator abundance, and an abundance of larval food resources. Likewise, the increase of forest cover around Ellenton Bay from <20% in 1951 to >60% in 2001 probably contributed to the long-term persistence of amphibians at this site. Our findings provide an optimistic counterpoint to the issue of the global decline of biological diversity by demonstrating that conservation efforts can mitigate historical habitat degradation.     

Predator-prey relationships between the rock lobster Jasus lalandii and the mussel Aulacomya ater at Robben Island on the Cape West Coast of Africa

The natural diet and mode of feeding of the rock lobster Jasus lalandii (H. Milne Edwards) was determined in a rock-lobster sanctuary near Cape Town, South Africa. Field observations were tested and confirmed by means of aquarium studies. Rock lobsters feed mainly upon ribbed mussels Aulacomya ater (Molina), which comprise the largest component of the sessile benthic fauna. Mussel remains were found as the major constituent in 97% of the rock-lobster stomachs examined. The density of rock lobsters averaged 8,100 per hectare (0.81 m^-2), while mussel biomass averaged more than 5 kg (wet whole weight) m^-2 within the same depth range (12 to 30 m). More than 80% of the mussel biomass comprised large individuals between 60 and 90 mm in length. Large rock lobsters (mainly males) were capable of feeding on all sizes of mussels, although many of these were inaccessible to predation. Smaller rock lobsters became progressively more limited in the size range of mussels which they could crack open and consume. Competition between rock lobsters for small mussels appeared to be intense, as mussels of suitable size for feeding were generally in short supply to most of the rock-lobster population. Hence, feeding and growth rates of rock lobsters are likely to be affected by the relative population densities of predator and prey. Growth rates of rock lobsters could be limited by food supplies even in areas where mussel biomass is comparatively large.     

Fire-climate history and landscape patterns of high burn severity areas on the California southern and central coast

There is mounting evidence that fire size and severity have been growing on the central and southern California coastal landscape over the past several decades. Landsat satellite data was analyzed for the 20 largest fires on the Central California coast since 1984 to determine the relationships between climate/weather conditions at the time of ignition and the size of high burn severity (HBS) areas. The study also examined the relationship between area burned and landscape patterns of HBS coverage, including patch size, edge complexity, perimeter-to-area ratio, and aggregation metrics. Results showed that climate conditions at the time of ignitions have been significant controllers of the total area of HBS and the complexity of HBS patches on the fire landscape. As maximum air temperatures for the month of ignition approached 40^o C, the percentage of HBS to total area burned frequently exceeded 20%. The percentage of HBS to total area burned also exceed 20% when the precipitation total recorded during the previous 12 months was less than 25% of the annual average precipitation. Landscape analysis results showed that, as the total area burned in fires on the Central California coast grows, the edge lengths and areas of HBS patches also grows at a rapid rate. At the same time, the perimeter-to-area ratio of HBS patches decreases gradually and the HBS patches become more aggregated as total burned area grows.     

A spatial gradient in the potential reproductive output of the sea mussel <Emphasis Type="Italic">Mytilus californianus</Emphasis>

Spatial variation in reproductive output from different populations within a region could have important consequences for recruitment, and cascading effects on populations and communities of marine species, but is rarely examined over meso-scales (i.e., tens to hundreds of kilometers). In this study, reproduction in the dominant mid-intertidal mussel, Mytilus californianus, was examined from sites spanning Point Conception, California over a 6-month period (March–August 2000). There was a dramatic geographic pattern in the relationship between size and potential reproductive output that was qualitatively similar across all 6 months sampled. Increases in allocation to reproductive tissue with increasing body size occurred at all sites, but the slope nearly doubled at sites south of Point Conception compared to northern sites. The spatial variation in size-specific reproductive output, coupled with additional spatial gradients in mussel density and size distributions, combined to increase total reproductive output by over eightfold at southern relative to northern sites. This study highlights the need to explicitly examine spatial patterns of reproductive output at these meso-scales, in order to better understand connectivity and source–sink dynamics in marine systems.     

Local and meso-scale patterns of recruitment and abundance of two intertidal crab species that compete for refuges

Interference competition for limited habitat or refuges is known to produce density-dependent mortality and generate patterns of micro-habitat distribution. While in mobile species the outcome of interference at a local scale can usually be determined from differences in body size and behavior, the population-level consequences of such interactions vary depending on rates of settlement and recruitment at a site, which are not directly correlated to local reproductive success. Previous experimental studies in central Chile demonstrated that interference competition for refuges is the primary factor driving microhabitat segregation between the predatory crabs Acanthocyclus gayi and Acanthocyclus hassleri, with the latter species monopolizing galleries inside mussel beds and excluding A. gayi to rock crevices. Between April 2001 and March 2006 we quantified monthly recruitment rates in artificial collectors at 17 sites over 900 km of the central coast of Chile. Results show that recruitment rates of A. hassleri are almost two orders of magnitude lower than those of A. gayi, and that they are tightly and positively correlated among sites across the region, suggesting that at scales of kilometers larval stages of these species are affected by similar oceanographic processes. Total crab densities per site were also positively correlated between species and strongly associated to mussel cover, with overall low crab densities at all sites where mussel cover was lower than about 60%. At all sites with mussel cover >60%, the ratio of A. gayi to A. hassleri density progressively decreased from recruits (2.6) to juveniles (0.5) to adults (0.04), overcoming initial differences in recruitment rates. The relative success of the inferior competitor at sites with low mussel cover does not appear to provide a potential mechanism favoring regional coexistence through dispersal to other sites (“mass effects”), because their densities were lower than at sites of high mussel cover. Yet, at many sites of low mussel cover the dominant competitor is virtually absent, allowing A. gayi to attain larger population sizes at the scale of the region. Thus, the factors limiting the dominant competitor from successfully utilizing other microhabitats seem to be the most critical factor in promoting both local and regional coexistence between these species.     

Importance of heterotrophic planktonic communities in a mussel culture environment: the Grande Entrée lagoon,Magdalen Islands (Québec,Canada)

Mussel culture in coastal environments relies on the availability of food of sufficient quality and quantity. Both to determine this availability and to examine impacts that this aquaculture practice may have on the environment, it is important to have good knowledge of the type of plankton communities present in aquaculture sites. It is usually thought that phytoplankton make up the bulk of mussel diet in many of these sites. Here we show that the Grande-Entrée lagoon [Magdalen Islands, Gulf of St Lawrence (GSL), Canada], where commercial mussel culture has been on-going since 1980, differs from this pattern. Heterotrophic protists dominate for most of the summer-early fall season (apart from short diatom bursts), with a high average biomass of 160 mg C m⁻³. The dominance of small-sized phytoplankton cells (notably green algae), low nutrient concentrations (e.g. 0.3 μM NO₃⁻ on average) and high biomass of heterotrophic protists (mostly naked ciliates and tintinnids) all point to the importance of the microbial food web in this shallow marine environment. Sustained cultivation of suspended mussels in the lagoon suggests that these heterotrophic protists could be an important source of food for the mussels, supplementing the small amount of phytoplankton present.     

A methodological framework for integrating computational fluid dynamics and ecological models applied to juvenile freshwater mussel dispersal in the Upper Mississippi River

Interdisciplinary research in hydraulics and ecology for river management and restoration must integrate processes that occur over a wide range of spatial and temporal scales, which presents a challenge to ecohydraulics modelers. Computational fluid dynamics (CFD) models are being more widely used to determine flow fields for ecohydraulics applications. In the Upper Mississippi River (UMR), the mussel dynamics model was developed as a tool for management and conservation of freshwater mussels (Unionidae), which are benthic organisms, imperiled in North America, that are inextricably linked with the hydraulics of river flow. We updated the juvenile dispersal component of the mussel dynamics model by using stochastic Lagrangian particle tracking in a three dimensional flow field output from CFD models of reaches in the UMR. We developed a methodological framework to integrate hydrodynamic data with the mussel dynamics model, and we demonstrate the use of the juvenile dispersal model employed within the methodological framework in two reaches of the UMR. The method was used to test the hypothesis that impoundment affects the relationship of some hydraulic parameters with juvenile settling distribution. Simulation results were consistent with this hypothesis, and the relationships of bed shear stress and Froude number with juvenile settling were altered by impoundment most likely through effects on local hydraulics. The methodological framework is robust, integrates Eulerian and Lagrangian reference frameworks, and incorporates processes over a wide range of temporal and spatial scales, from watershed scale hydrologic processes (decades), to reach scale (km) processes that occur over hours or days, and turbulent processes on spatial scales of meter to millimeter and times scales of seconds. The methods are presently being used to assess the impacts of pre- and early post-settlement processes on mussel distributions, including the effects of bed shear stress, and the sensitivity of the location of the host fish when juveniles excyst, on juvenile settling distribution.     

Wave exposure effects on population structure and recruitment in the mussel <Emphasis Type="Italic">Perna perna</Emphasis> suggest regulation primarily through availability of recruits and food,not space

Recruitment and population structure of Perna perna in low shore mussel beds were investigated over 15 months at six sites along the south coast of South Africa. Initial, subjective classification of sites as wave exposed or wave sheltered (three of each) was confirmed using the dissolution of cement blocks to measure average water flux and dynamometers for maximum wave force. Recruitment occurred throughout the year, but recruit (1–5 mm) densities were significantly higher from January to April 1996 on both shore types. Recruit densities were positively correlated with adult (>15 mm) densities for both shore types (P < 0.05) but the correlations were extremely weak (r ² < 0.06 in each case). In areas with 100% cover, adult size (mean and maximum lengths) was greater on exposed sites, but density showed the reverse and was negatively correlated with maximum wave strength (r = −0.84). Despite differences in adult densities and sizes, biomass, which is a product of the two, showed no significant difference between the two shore types (ANOVA P > 0.05). Thus wave exposure dramatically affects density, recruitment and mussel size, but not recruitment timing or biomass where there is 100% cover, and mediates a three-way interaction among food supply, larval supply and intraspecific competition for space. In contrast to shores with saturation recruitment, mussel biomass here appears to be limited by recruit supply and constraints of food, especially on sheltered shores, while density is regulated through intraspecific competition for space primarily on exposed shores and at small spatial scales.     

Possible Environmental Factors Underlying Amphibian Decline in Eastern Puerto Rico: Analysis of U.S. Government Data Archives

Abstract: The past three decades have seen major declines in populations of several species of amphibians at high elevations in eastern Puerto Rico, a region unique in the humid tropics because of the degree of environmental monitoring that has taken place through the efforts of U.S. government agencies. I examined changes in environmental conditions by examining time-series data sets that extend back at least into the 1980s, a period when frog populations were declining. The data include forest cover; annual mean, minimum, and maximum daily temperature; annual rainfall; rain and stream chemistry; and atmospheric-dust transport. I examined satellite imagery and air-chemistry samples from a single National Aeronautics and Space Administration aircraft flight across the Caribbean showing patches of pollutants, described as thin sheets or lenses, in the lower troposphere. The main source of these pollutants appeared to be fires from land clearing and deforestation, primarily in Africa. Some pollutant concentrations were high and, in the case of ozone, approached health limits set for urban air. Urban pollution impinging on Puerto Rico, dust generation from Africa ( potential soil pathogens), and tropical forest burning ( gaseous pollutants) have all increased during the last three decades, overlapping the timing of amphibian declines in eastern Puerto Rico. None of the data sets pointed directly to changes so extreme that they might be considered a direct lethal cause of amphibian declines in Puerto Rico. More experimental research is required to link any of these environmental factors to this problem.