El Niño,volcanism, and global climate

The June 1991 eruption of Mt. Pinatubo in the Philippines produced one of the greatest volcanic aerosols in the last hundred years. The estimated net decrease of radiation may have peaked at 10% in the tropics. What was the impact of the Pinatubo aerosol on regional and global climate? Besides the expected net cooling of the average global surface temperature, correlation studies indicate that other types of climate anomalies may also be expected. These include the appearance of an El Niño event, decreased Indian monsoon rainfall, fewer tropical storms in the north Atlantic Ocean in 1991–1993, and normal to above normal winter rainfall in California in 1991/92, all of which were observed. A proposed physical mechanism for the almost-simultaneous occurrence of this constellation of climate anomalies is presented. The results of correlation studies between low-latitude volcanic aerosols and the El Niño/Southern Oscillation are presented in some detail as one example. The correlation between Indian monsoon rainfall and tropical storms in the north Atlantic Ocean is also shown and is updated for the most recent 5 years.     

Linking climate change to population cycles of hares and lynx

The classic 10‐year population cycle of snowshoe hares (Lepus americanus, Erxleben 1777) and Canada lynx (Lynx canadensis, Kerr 1792) in the boreal forests of North America has drawn much attention from both population and community ecologists worldwide; however, the ecological mechanisms driving the 10‐year cyclic dynamic pattern are not fully revealed yet. In this study, by the use of historic fur harvest data, we constructed a series of generalized additive models to study the effects of density dependence, predation, and climate (both global climate indices of North Atlantic Oscillation index (NAO), Southern Oscillation index (SOI) and northern hemispheric temperature (NHT) and local weather data including temperature, rainfall, and snow). We identified several key pathways from global and local climate to lynx with various time lags: rainfall shows a negative, and snow shows a positive effect on lynx; NHT and NAO negatively affect lynx through their positive effect on rainfall and negative effect on snow; SOI positively affects lynx through its negative effect on rainfall. Direct or delayed density dependency effects, the prey effect of hare on lynx and a 2‐year delayed negative effect of lynx on hare (defined as asymmetric predation) were found. The simulated population dynamics is well fitted to the observed long‐term fluctuations of hare and lynx populations. Through simulation, we find density dependency and asymmetric predation, only producing damped oscillation, are necessary but not sufficient factors in causing the observed 10‐year cycles; while extrinsic climate factors are important in producing and modifying the sustained cycles. Two recent population declines of lynx (1940–1955 and after 1980) were likely caused by ongoing climate warming indirectly. Our results provide an alternative explanation to the mechanism of the 10‐year cycles, and there is a need for further investigation on links between disappearance of population cycles and global warming in hare–lynx system.     

A boundary current drives synchronous growth of marine fishes across tropical and temperate latitudes

Entrainment of growth patterns of multiple species to single climatic drivers can lower ecosystem resilience and increase the risk of species extinction during stressful climatic events. However, predictions of the effects of climate change on the productivity and dynamics of marine fishes are hampered by a lack of historical data on growth patterns. We use otolith biochronologies to show that the strength of a boundary current, modulated by the El Niño‐Southern Oscillation, accounted for almost half of the shared variance in annual growth patterns of five of six species of tropical and temperate marine fishes across 23° of latitude (3000 km) in Western Australia. Stronger flow during La Niña years drove increased growth of five species, whereas weaker flow during El Niño years reduced growth. Our work is the first to link the growth patterns of multiple fishes with a single oceanographic/climate phenomenon at large spatial scales and across multiple climate zones, habitat types, trophic levels and depth ranges. Extreme La Niña and El Niño events are predicted to occur more frequently in the future and these are likely to have implications for these vulnerable ecosystems, such as a limited capacity of the marine taxa to recover from stressful climatic events.     

Changes in spring arrival of Nearctic-Neotropical migrants attributed to multiscalar climate

Climate-related changes associated with the California marine ecosystem have been documented; however, there are no studies assessing changes in terrestrial vertebrate phenology on the Pacific coast of western North America. We analyze the spring phenology of 21 Nearctic-Neotropical migratory songbird species in central and northern CA. Using observational and banding data at multiple sites, we evaluate evidence for a change in arrival timing being linked to either nonclimatic or multiscalar climatic explanations. Using correlation analysis, of the 13 species with a significant ( P <0.10) change in arrival, the arrival timing of 10 species (77%) is associated with both temperature and a large-scale climate oscillation index (El Niño Southern Oscillation, ENSO; North Atlantic Oscillation, NAO; and/or Pacific Decadal Oscillation, PDO) at least at one location. Eight of the 13 species (62%) are advancing their migratory timing. All species for which spring arrival is associated with climate at multiple locations are exhibiting changes ( n =5) and all species lacking evidence for association between migration phenology and climate ( n =3) exhibit no change. Migrants tend to arrive earlier in association with warmer temperatures, positive NAO indices, and stronger ENSO indices. Twelve species negatively correlate ( P ≤0.05) with local or regional temperature at least at one location; five species negatively correlate with ENSO. Eleven species' arrival is correlated ( P ≤0.05) with NAO; 10 are negatively associated. After an exhaustive literature search, this is apparently the first documentation of an association between NAO and migratory phenology in western North America.     

Beyond annual and seasonal averages: using temporal patterns of precipitation to predict butterfly richness across an elevational gradient

1. Ecologists often make predictions about community richness and diversity using climate variables that include seasonal precipitation totals and mean daily temperatures. While means and totals can be effective predictors to a certain extent, the complexities of faunal–climate relationships might be over‐simplified through the use of coarse‐grained variables. 2. The goal of this study was to investigate less commonly studied climate variables, including indices of intra‐annual variation in the timing and intensity of precipitation events that might be used to predict butterfly richness across an elevational gradient. Data from a long‐term, single‐observer dataset at four sites in California were examined with Bayesian model averaging and structural equation modelling. Species‐specific responses to climate were compared with community responses at each site. 3. At lower elevations, it was found that the relative importance of climate variables shifted towards temporal patterns of precipitation, including the timing of the first storm event and the annual number of precipitation events. Heterogeneity among sites was apparent in the importance of specific weather variables, and temporal trends (across years) were detected for a small number of variables. Species‐specific results paralleled those obtained from analysis of species richness, thus suggesting a commonality of response to climate across site‐specific assemblages. 4. Models were improved by inclusion of the Pacific Decadal Oscillation and El Niño‐Southern Oscillation indices, indicating that regional variables can profitably be included in faunal–climate relationship analyses. These results emphasise the need for researchers to examine climate variables beyond the most readily summarised means and totals.     

ENSO, rainfall and temperature influences on extreme population declines among African savanna ungulates

Climatic variation associated with the North Atlantic Oscillation (NAO) and El Niño‐Southern Oscillation (ENSO) has a widespread influence on the population dynamics of many organisms worldwide. While previous analyses have related the dynamics of northern ungulates to the NAO, there has been no comparable assessment for the species rich assemblages of tropical and subtropical Africa. Census records for 11 ungulate species in South Africa's Kruger National Park over 1977–96 reveal severe population declines by seven species, which were inadequately explained by indices of ENSO or its effects on annual rainfall totals. An additional influence was an extreme reduction in dry season rainfall, concurrent with and perhaps related to a regional temperature rise, possibly a signal of global warming. Boundary fencing now restricts range shifts by such large mammals in response to climatic variation. Our models project near extirpation of three ungulate species from the park's fauna should these climatic conditions recur.     

Combined effect of ENSO and SAM on the population dynamics of the invasive yellowjacket wasp in central Chile

The population dynamics of the yellowjacket wasp (Vespula germanica Fabricus) in central Chile were analyzed for the first time. Using a simple Ricker logistic model and adding the effects of local weather variables (temperature and precipitation) and large-scale climate phenomena as El Niño Southern Oscillation (ENSO) and the Southern Annular Mode (SAM), we modeled the interannual fluctuations in nest density. The best model according to the Bayesian information criterion (BIC) included 1-year-lag negative feedback combined with the positive additive effects of ENSO and SAM. According to this model, yellowjacket nest density was favored by warm and dry winters, which probably influenced the survival of overwintering queens. Large-scale climatic variables [Southern Oscillation Index (SOI) and SAM] described the effect of exogenous factors in wasp fluctuations better than local weather variables did. Our results emphasize the usefulness of climate indices and simple theoretical-based models in insect ecological research.     

Global climate cycles and cyclones: consequences for rainfall patterns and lemur reproduction in southeastern Madagascar

Most studies that examine the influence of climatic change on flora and fauna have focused on northern latitudes; however, there is increasing recognition that tropical regions are also being affected. Despite this, regions such as Madagascar, which are rich in endemic biodiversity but may have low adaptive capacity to climatic change, are poorly represented in studies examining the effects of climate variability on biota. We investigated how El Niño Southern Oscillations (ENSO) influence precipitation patterns in the rainforest region of southeastern Madagascar (1962–2006) and then constructed models to assess the potential contribution of climatic variables on the reproductive parameters of the Milne Edward's sifaka, a threatened lemur species (Propithecus edwardsi), over a 20‐year period. The Southern Oscillation Index of sea surface temperature (SST) anomalies in the tropical Pacific was associated with precipitation patterns including wetter wet seasons during warmer phases and drier dry seasons following cooler phases. The best‐supported models of lemur fecundity (female offspring per female that survive to 1 year of age per year) included cyclone presence during gestation and ENSO phase before conception and during the first 6 months of life. Models also suggested that heavy rains during gestation may limit birth rates and that prolonged drought during female lactation may limit first year offspring survival; although these variables were given little importance for predicting overall fecundity relative to ENSO phases and cyclone presence. Our results linking lemur reproduction with climatic variability suggest that climatic changes may be an additional threat to Madagascar's unique and already endangered flora and fauna. The association between precipitation in southeastern Madagascar and SST anomalies in the tropical Pacific suggests that dynamics of wildlife populations even in tropical areas such as Madagascar can be affected by global climate cycles making them potentially vulnerable to global climate change.     

Large-Scale Temperature Patterns in Past Centuries: Implications for North American Climate Change

Recent climate reconstructions are analyzed specifically for insights into those patterns of climate variability in past centuries with greatest impact on the North American region. Regional variability, largely associated with the El Nino/Southern Oscillation (ENSO) phenomenon, the North Atlantic Oscillation (NAO), and multidecadal patterns of natural variability, are found to mask the emergence of an anthropogenic temperature signal in North America. Substantial recent temperature anomalies may however indicate a possible recent emergence of this signal in the region. Multidecadal North Atlantic variability is likely to positively reinforce any anthropogenic warming over substantial parts of North America in coming decades. The recent magnitudes of El Nino events appear to be unprecedented over the past several centuries. These recent changes, if anthropogenic in nature, may outweigh the projection of larger-scale climate change patterns onto the region in a climate change scenario. The implications of such changes for North America, however, are not yet clear. These observations suggest caution in assessing regional climate change scenarios in North America without a detailed consideration of possible anthropogenic changes in climate patterns influencing the region.     

Bird migration times, climate change, and changing population sizes

Past studies of bird migration times have shown great variation in migratory responses to climate change. We used 33 years of bird capture data (1970–2002) from Manomet, Massachusetts to examine variation in spring migration times for 32 species of North American passerines. We found that changes in first arrival dates – the unit of observation used in most studies of bird migration times – often differ dramatically from changes in the mean arrival date of the migration cohort as a whole. In our study, the earliest recorded springtime arrival date for each species occurred 0.20 days later each decade. In contrast, the mean arrival dates for birds of each species occurred 0.78 days earlier each decade. The difference in the two trends was largely explained by declining migration cohort sizes, a factor not examined in many previous studies. We found that changes in migration cohort or population sizes may account for a substantial amount of the variation in previously documented changes in migration times. After controlling for changes in migration cohort size, we found that climate variables, migration distance, and date of migration explained portions of the variation in migratory changes over time. In particular, short-distance migrants appeared to respond to changes in temperature, while mid-distance migrants responded particularly strongly to changes in the Southern Oscillation Index. The migration times of long-distance migrants tended not to change over time. Our findings suggest that previously reported changes in migration times may need to be reinterpreted to incorporate changes in migration cohort sizes.     

Primary production dynamics and climate variability: ecological consequences in semiarid Chile

Increase in rainfall variability has important consequences for organisms in arid and semiarid regions around the world. In South American and Australian deserts, the El Niño/Southern Oscillation (ENSO) phenomenon greatly influences rainfall patterns, and therefore the dynamics of plant communities. However, the field data needed to assess the effect of climate change on vegetational patterns is difficult to obtain because of the large spatial scale required for such studies. Normalized Difference Vegetation Index (NDVI) characteristics allow the use of several indexes related to vegetational structure. Due to its direct relationship with primary productivity, it is possible to obtain several measures of annual productivity. These include annual plant yield, annual maximum yield, onset of 'greening-up' and senescence phases, length of the 'green' season, vegetation peak, and therefore, the periods when more or less food is available for herbivores. After verification with ground-truth measures, we used NDVI data from two semiarid localities in north-central Chile (Fray Jorge and Aucó) to determine the relationship between rainfall patterns and vegetation cover and productivity related to El Niño phenomenon. With this information we gauge the influence of climatic processes on primary productivity in western South America, an area subject to strong climate variability. We predict significant variation in Chilean semiarid regions due to climate change, affecting mainly the extent and timing of annual growth season of vegetation, and also including a shorter and delayed greening-up season. Also, we predict that important decreases in rainfall levels will not have strong effects on primary production in these semiarid ecosystems.     

Variation in leaf litter nutrients of a Costa Rican rain forest is related to precipitation

By assessing current leaf litter nutrient dynamics, we may be able to predict responses of nutrient cycling in tropical ecosystems to future environmental change. The goal of this study was to assess whether nutrient cycling is related to seasonal variation in rainfall in a wet tropical forest. We examined leaf litter of an old-growth tropical rain forest in N.E. Costa Rica over a 4-year period to explore seasonal and inter-annual changes in leaf litter nutrient concentrations, and to evaluate potential short- and long-term drivers of variation in litter nutrient concentration, particularly that of phosphorus (P) and nitrogen (N). We also examined the temporal dynamics of calcium, potassium, and magnesium in the leaf litter. Leaf litter [P] and %N changed significantly with time, both seasonally and inter-annually. Seasonal changes in leaf litter [P] were strongly positively correlated with rainfall from the previous 2 weeks; cations, however, were inversely related to this measure of current rainfall, while %N was not related to rainfall. We propose that the positive relationship between current rainfall and leaf litter [P] is due to a response by the vegetation to an increase in nutrient availability and uptake. In contrast, given the negative relationship between current rainfall and cation concentrations, leaching from live leaf tissue is a more likely driver of short-term changes in cations. Should global climate change include altered rainfall patterns in this biome, one class of ecosystem-level responses could be significant changes in P and cation cycling.     

Effects of large-scale climatic fluctuations on survival and production of young in a Neotropical migrant songbird, the yellow warbler Dendroica petechia

Migrant songbirds are vulnerable to changes in climatic conditions on both the breeding and wintering grounds. For North American Neotropical migrants, the El Niño/Southern Oscillation (ENSO), via its effects on global precipitation and temperature, modulates the productivity of their temperate and tropical terrestrial ecosystems used during the course of their annual cycle. We evaluated how a densely nesting population of yellow warblers Dendroica petechia in a riparian forest in southern Manitoba, Canada, responded to the El Niño/Southern Oscillation (ENSO) between 1992 and 2001. Standardized mist netting was used to estimate apparent annual survival of adults and production of young. Both adult survival and the production of young were positively correlated with the Southern Oscillation Index (SOI). More specifically, values of both these demographic parameters were lowest during El Niño years and highest during La Niña years. These findings demonstrate the influence of climate on populations of Neotropical migrants in North America. The more frequent El Niños predicted to result from future global climate change could negatively affect populations of yellow warblers and other Neotropical migrants breeding in this region.     

Climate and wildfires in the North American boreal forest

The area burned in the North American boreal forest is controlled by the frequency of mid-tropospheric blocking highs that cause rapid fuel drying. Climate controls the area burned through changing the dynamics of large-scale teleconnection patterns (Pacific Decadal Oscillation/El Niño Southern Oscillation and Arctic Oscillation, PDO/ENSO and AO) that control the frequency of blocking highs over the continent at different time scales. Changes in these teleconnections may be caused by the current global warming. Thus, an increase in temperature alone need not be associated with an increase in area burned in the North American boreal forest. Since the end of the Little Ice Age, the climate has been unusually moist and variable: large fire years have occurred in unusual years, fire frequency has decreased and fire–climate relationships have occurred at interannual to decadal time scales. Prolonged and severe droughts were common in the past and were partly associated with changes in the PDO/ENSO system. Under these conditions, large fire years become common, fire frequency increases and fire–climate relationships occur at decadal to centennial time scales. A suggested return to the drier climate regimes of the past would imply major changes in the temporal dynamics of fire–climate relationships and in area burned, a reduction in the mean age of the forest, and changes in species composition of the North American boreal forest.     

Solar radiation and ENSO predict fruiting phenology patterns in a 15‐year record from Kibale National Park,Uganda

Fruiting, flowering, and leaf set patterns influence many aspects of tropical forest communities, but there are few long‐term studies examining potential drivers of these patterns, particularly in Africa. We evaluated a 15‐year dataset of tree phenology in Kibale National Park, Uganda, to identify abiotic predictors of fruit phenological patterns and discuss our findings in light of climate change. We quantified fruiting for 326 trees from 43 species and evaluated these patterns in relation to solar radiance, rainfall, and monthly temperature. We used time‐lagged variables based on seasonality in linear regression models to assess the effect of abiotic variables on the proportion of fruiting trees. Annual fruiting varied over 3.8‐fold, and inter‐annual variation in fruiting is associated with the extent of fruiting in the peak period, not variation in time of fruit set. While temperature and rainfall showed positive effects on fruiting, solar radiance in the two‐year period encompassing a given year and the previous year was the strongest predictor of fruiting. As solar irradiance was the strongest predictor of fruiting, the projected increase in rainfall associated with climate change, and coincident increase in cloud cover suggest that climate change will lead to a decrease in fruiting. ENSO in the prior 24‐month period was also significantly associated with annual ripe fruit production, and ENSO is also affected by climate change. Predicting changes in phenology demands understanding inter‐annual variation in fruit dynamics in light of potential abiotic drivers, patterns that will only emerge with long‐term data.     

El Niño Southern Oscillation influences the abundance and movements of a marine top predator in coastal waters

Large‐scale climate modes such as El Niño Southern Oscillation (ENSO) influence population dynamics in many species, including marine top predators. However, few quantitative studies have investigated the influence of large‐scale variability on resident marine top predator populations. We examined the effect of climate variability on the abundance and temporary emigration of a resident bottlenose dolphin (Tursiops aduncus) population off Bunbury, Western Australia (WA). This population has been studied intensively over six consecutive years (2007–2013), yielding a robust dataset that captures seasonal variations in both abundance and movement patterns. In WA, ENSO affects the strength of the Leeuwin Current (LC), the dominant oceanographic feature in the region. The strength and variability of the LC affects marine ecosystems and distribution of top predator prey. We investigated the relationship between dolphin abundance and ENSO, Southern Annular Mode, austral season, rainfall, sea surface salinity and sea surface temperature (SST). Linear models indicated that dolphin abundance was significantly affected by ENSO, and that the magnitude of the effect was dependent upon season. Dolphin abundance was lowest during winter 2009, when dolphins had high temporary emigration rates out of the study area. This coincided with the single El Niño event that occurred throughout the study period. Coupled with this event, there was a negative anomaly in SST and an above average rainfall. These conditions may have affected the distribution of dolphin prey, resulting in the temporary emigration of dolphins out of the study area in search of adequate prey. This study demonstrated the local effects of large‐scale climatic variations on the short‐term response of a resident, coastal delphinid species. With a projected global increase in frequency and intensity of extreme climatic events, resident marine top predators may not only have to contend with increasing coastal anthropogenic activities, but also have to adapt to large‐scale climatic changes.     

Breeding sites and population of seabirds on Admiralty Bay,King George Island,Antarctica

The monitoring of the status and distribution of seabird populations is necessary to understand their spatial and temporal responses to rapid climate changes occurring in the Western Antarctic Peninsula area. We surveyed and mapped Admiralty Bay bird communities and related them to climate variables—temperature, temperature anomaly, Antarctic Oscillation Index and El-Niño Southern Oscillation Index. We recorded 13 breeding seabird species over three seasons (2009/2010, 2010/2011 and 2011/2012) and mapped 10 of them over an area of 149.5 ha. The ice-free areas with the greatest number of species were Point Thomas, Keller Peninsula and Hennequin Point. The most abundant species was the Adelie Penguin (Pygoscelis adeliae) followed by the Chinstrap Penguin (P. antarcticus). We observed that the number of breeding pairs of Gentoo Penguins (P. papua), Chinstrap Penguins and skuas (Catharacta maccormicki and C. antarctica) are related to temperature, temperature anomaly and El-Niño Southern Oscillation Index. The size of breeding populations and their distributions have been fluctuating over the last 30 years in ice-free areas of Admiralty Bay. Most species showed a decreasing trend from 1978 to 2012, with the exception of Chinstrap Penguins, Southern Giant Petrels (Macronectes giganteus) and skuas, which seem to be stable in numbers in the last two decades. Decreases in seabird populations from the Antarctic Peninsula are widely recognized as a response to environmental change and anthropogenic influences such as tourism and building activities, thus highlighting the importance of monitoring to support mitigation measures.     

Relationships between climate and growth of Gymnocypris selincuoensis in the Tibetan Plateau

The consequences of climate change are becoming increasingly evident in the Tibetan Plateau, represented by glaciers retreating and lakes expanding, but the biological response to climate change by plateau–lake ecosystems is poorly known. In this study, we applied dendrochronology methods to develop a growth index chronology with otolith increment widths of Selincuo naked carp (Gymnocypris selincuoensis), which is an endemic species in Lake Selincuo (4530 m), and investigated the relationships between fish growth and climate variables (regional and global) in the last three decades. A correlation analysis and principle component regression analysis between regional climate factors and the growth index chronology indicated that the growth of G. selincuoensis was significantly and positively correlated with length of the growing season and temperature‐related variables, particularly during the growing season. Most of global climate variables, which are relevant to the Asian monsoon and the midlatitude westerlies, such as El Nino Southern Oscillation Index, the Arctic Oscillation, North Atlantic Oscillation, and North America Pattern, showed negative but not significant correlations with the annual growth of Selincuo naked carp. This may have resulted from the high elevation of the Tibetan Plateau and the high mountains surrounding this area. In comparison, the Pacific Decade Oscillation (PDO) negatively affected the growth of G. selincuoensis. The reason maybe that enhancement of the PDO can lead to cold conditions in this area. Taken together, the results indicate that the Tibetan Plateau fish has been affected by global climate change, particularly during the growing season, and global climate change likely has important effects on productivity of aquatic ecosystems in this area.     

Earlier Migration Timing,Decreasing Phenotypic Variation,and Biocomplexity in Multiple Salmonid Species

Climate-induced phenological shifts can influence population, evolutionary, and ecological dynamics, but our understanding of these phenomena is hampered by a lack of long-term demographic data. We use a multi-decade census of 5 salmonid species representing 14 life histories in a warming Alaskan stream to address the following key questions about climate change and phenology: How consistent are temporal patterns and drivers of phenology for similar species and alternative life histories? Are shifts in phenology associated with changes in phenotypic variation? How do phenological changes influence the availability of resource subsidies? For most salmonid species, life stages, and life histories, freshwater temperature influences migration timing – migration events are occurring earlier in time (mean = 1.7 days earlier per decade over the 3–5 decades), and the number of days over which migration events occur is decreasing (mean = 1.5 days per decade). Temporal trends in migration timing were not correlated with changes in intra-annual phenotypic variation, suggesting that these components of the phenotypic distribution have responded to environmental change independently. Despite commonalities across species and life histories, there was important biocomplexity in the form of disparate shifts in migration timing and variation in the environmental factors influencing migration timing for alternative life history strategies in the same population. Overall, adult populations have been stable during these phenotypic and environmental changes (λ ≈1.0), but the temporal availability of salmon as a resource in freshwater has decreased by nearly 30 days since 1971 due to changes in the median date of migration timing and decreases in intra-annual variation in migration timing. These novel observations advance our understanding of phenological change in response to climate warming, and indicate that climate change has influenced the ecology of salmon populations, which will have important consequences for the numerous species that depend on this resource.     

A Bloom of the Planktonic Diatom,Cerataulina pelagica,off the Coast of Northeastern New Zealand in 1983, and its Contribution to an Associated Mortality of Fish and Benthic Fauna

A bloom of Cerataulina pelagica dominated the sea area off the north-east coast of New Zealand for three months in the summer of 1982–83, and has been related to unusually calm conditions associated with an unusually low Southern Oscillation Index. Deaths of benthic shellfish during the bloom were attributed to anoxia induced by the bacterial decay of the diatom cells. Death of bony fish caught on long-lines was attributed to anoxia and the clogging of the gills with mucilage produced by the diatoms.