长江口中华鲟自然保护区潮间带鱼类群落结构

为深入了解鱼类对潮间带水域的利用,2010年5月、8月、11月在长江口中华鲟自然保护区潮间带水域进行了插网调查。结果表明:调查期间共出现28种鱼类,隶属于10目16科,鲤形目种类最多,其次为鲈形目;群落优势种为中国花鲈(Lateolabrax maculates)、鮻(Liza haematocheila)和窄体舌鳎(Cynoglossus gracilis);长江口中华鲟自然保护区潮间带鱼类群落可以分为河口定居鱼类、海洋偶见鱼类、降海洄游鱼类、溯河洄游鱼类、海洋洄游鱼类和淡水鱼类6种生态类群,其中河口定居鱼类和海洋洄游鱼类占据明显优势。对调查水域鱼类群落进行聚类分析,结果表明秋季和春夏季差异显著,鮻、中国花鲈、斑尾刺虾虎鱼(Acanthogobius ommaturus)是对季节差异贡献最大的3种鱼类。主成分分析结果表明,夏季鱼类群落主要受到海洋洄游鱼类的影响,而秋季主要受到河口定居鱼类的影响,这2种鱼类的幼鱼在不同季节利用潮间带水域索饵肥育可能是造成季节差异的主要原因。     

Spatial and seasonal patterns in freshwater ichthyofaunal communities of a tropical high island in Fiji

We surveyed freshwater ichthyofaunal communities in streams of Vanua Levu, Fiji, under a range of land cover to assess differential, seasonal effects on fish abundance and diversity. We collected fish from 32 families, 19 genera and 87 species, representing approximately 50% of the known Fijian freshwater and estuarine fish fauna. Position in reach was the strongest overall factor influencing fish abundance and diversity, particularly in the larger, steeper catchments. However, fish communities exhibited strong seasonal specificity with over half (55%) of species observed in only one season. There were greater numbers of estuarine and marine migrants and fishes with poor swimming ability in the dry season, with more schooling species, large predators and fish that prefer muddy benthos in the wet. In the more pristine catchments of Kubulau District, higher species abundance and diversity were observed in the wet season and were associated with significantly greater flow, pH and dissolved oxygen. We observed the opposite pattern for fish diversity and abundance from the more degraded catchments of Macuata Province. These results suggest that the wet season is having a net positive effect on habitable space for fishes in Kubulau and a net negative effect in Macuata, as species may be lost due to increased runoff from heavily cleared and cultivated catchments. Integrated water resource management across the full range of habitats utilized by Fiji’s freshwater fishes is recommended in order to maintain species diversity and abundance.     

Spatial and seasonal distribution patterns of fish assemblages in the Río Champotón, southeastern Mexico

Río Champotón in the Yucatán Peninsula is within the Usumacinta Province, the most diverse in fish fauna in Mexico, and is part of the Mesoamerican hotspot, with high endemism and exceptional habitat loss. The spatial and seasonal variability of its fish fauna and their relation to physical habitat characteristics were studied, finding 53 taxa. Canonical correspondence analysis shows salinity, distance from the river mouth, and substratum type differentiate an estuarine and a freshwater region. Fish fauna replacement was observed, with taxa exclusive to each region: 26 estuarine and 27 freshwater species. Six freshwater species enter the estuarine region when the high river flow depletes salinity. Estuary showed greater seasonal differences in specific richness than the freshwater region, which have sites with higher local specific richness. Despite its small drainage, Río Champotón has a highly diverse fish fauna and some areas are affected by human activity and the hurricane season. Its conservation is of critical importance.     

Fish abundance and species richness across an estuarine–freshwater ecosystem in the Neotropics

We investigated the distributional patterns of shallow-water fish and their environmental correlates along a broad spatial scale encompassing estuarine and freshwater ecosystems. Marine-vagrant species were restricted to the sites under the influence of salinity intrusion, whereas estuarine-related and freshwater guilds were found along the entire estuarine–freshwater gradient. Primary- and secondary-freshwater fish guilds had the most widespread spatial distribution and comprised a major fraction of the total abundance and species richness. Abiotic factors correlated with fish abundance and composition along two main environmental axes, one related with variation in salinity, water transparency, and sediment granulometry and the other with the slope gradient. Species richness was significantly higher at the natural channel linking the estuarine- and freshwater-ecosystem, which probably was due to: (a) a steeper slope that favored the confluence of fish from the littoral (<2 m) and limnetic (>2 m) zones and (b) the sporadic inflow of saltwater that carried into this region several marine-related species. Although estuarine–freshwater ecotones are known to support few species, mainly salinity tolerant, our results suggest that habitat features and seasonal fish movement associated with salinity intrusion could lead to more diverse fish assemblages in this transitional zone.     

The composition and community structure of the ichthyofauna of the upper Scheldt estuary: synthesis of a 10-year data collection (1991–2001)

Fish collections on the cooling water filter screens of the Doel power plant were used to analyse temporal trends in estuarine fish populations of the upper Scheldt estuary, Belgium. Between 1991 and 2001, 62 species shared the estuary, with 15 species present at any one time. Identified were 23 marine stragglers, 16 freshwater species and four diadromous species, but their contribution to the total catch was insignificant. In terms of numbers, marine estuarine opportunists (17 species) mainly occurring as juveniles and estuarine residents (two species) were the dominant life cycle categories. Six fish species contributed to >97% of the total catch: three Pomatoschistus species [P. minutus (32%), P. microps (30%) and P. lozanoi (6%)], two Clupeidae [Clupea harengus (16%) and Sprattus sprattus (8%)] and Syngnathus rostellatus (5%). Abundance of almost all species in the estuary was highly seasonal and characterized by pronounced abundance peaks. Ordination of the data showed that variability in the temporal structure of estuarine fish community was first determined by differences in annual recruitment and then by predictable, cyclical patterns of species abundance.     

The icthyofauna of the Mhlathuze coastal lakes: some preliminary results

The fish faunas of the four Mhlathuze coastal lakes and the lower river comprise a diverse assortment of over fifty marine, estuarine and freshwater species. Three freshwater species are endemic to KwaZulu-Natal and nine estuarine species are endemic to southern Africa. Five species are of conservation significance. Species numbers in Lakes Mzingazi and Cubhu are similar historically and both lakes served as secondary nursery habitats for estuarine associated fishes. This role has been impacted by the construction of weirs at their outlets which prevent successful recruitment of estuarine species, especially during drought years when lake water levels are low. The fish faunas of Lakes Nsezi and Mangeza are depauperate and lack marine or estuarine components. In order that these systems fulfil their potential function as secondary nursery habitats to many estuarine fish species, minimum lake water levels must be set to ensure sufficient outflows at proposed fish ladders during critical spawning and migration times.     

Spatiotemporal Variation in Shallow-Water Freshwater Fish Distribution and Abundance in a Large Subtropical Coastal Lagoon

Patos Lagoon is located off the southern Brazilian coast and represents one of the largest coastal lagoons in the world. We estimated hydrological and physicochemical conditions associated with spatial variation in the abundance and diversity of freshwater fishes along the lagoon, and inter-annual variability in abundances of freshwater fishes occurring in its estuarine zone. During our study, the region experienced two periods of average rainfall and two periods with above-average rainfall. The characids Astyanax eigenmaniorum and Oligosarcus jenynsii and the siluriform Parapimelodus nigribarbis were the most abundant freshwater fishes in the estuary during wet periods when water levels were higher and salinity was lower. Increases in abundance of these species in the estuarine area, all of which members of primary-division freshwater families, apparently were associated with pulses of reproduction and passive transport from freshwater habitats located near middle and upper lagoon reaches. Abundance of species from secondary freshwater families, such as poeciliids and cichlids, were less correlated with hydrological conditions, and their patterns of occurrence in the estuary suggest active migration from nearby freshwater habitats draining into this area. Findings indicate that freshwater discharge in the basin and expansion/retraction of freshwaters in the middle-upper lagoon determined patterns of freshwater fish abundance and species richness in the estuarine zone.     

The structure and components of European estuarine fish assemblages

This paper discusses the structure of fish assemblages using information from 17 European estuarine areas (in the British Isles, Portugal, Belgium, France, the Netherlands, Germany, Norway and Spain). Binary (presence/absence) and quantitative data for each assemblage have been used to assess the assemblage structure according to taxonomy (i.e. species identity). Following this, a total of 29 functional guilds were created in order to describe the use made of an estuarine area for each taxon encountered: feeding preferences, reproduction type, substratum preferences (for bottom dwelling fish) and position within the water column (vertical preference guild). The paper focuses on the designation and determination of the proportions of the guild members of the fish assemblage within each estuary. Ecological guilds within the assemblage include estuarine residents, marine juvenile migrants, catadromous and anadromous migrants, marine seasonal users, and freshwater and marine adventitious species. Feeding guilds include detritivores, planktonic feeders, infaunal croppers and sediment ingesters, piscivores, and active predators of mobile crustaceans. Reproduction guilds include planktonic and demersal spawners and those using brood-protection. The substratum preference indicates the proportions of sand, mud, rock and vegetation dwellers, and the vertical preference denotes benthic, demersal or pelagic species. The analysis has allowed both the estuaries to be grouped according to taxonomic and guild similarity and the characterisation of a typical European estuarine fish assemblage. Within the limits posed by differing sampling methods, times of sampling and survey rationale, there is a high similarity between estuaries. The data indicate common patterns of estuarine usage irrespective of the differences between the estuaries although such patterns cannot be interpreted fully given the incomplete knowledge of their physical and anthropogenic characteristics.     

Coping with the cold: energy storage strategies for surviving winter in freshwater fish

For many ectothermic animals, the acquisition, storage and depletion of lipids is integral to successfully coping with reduced metabolic rates and activity levels associated with cold, winter periods. In fish, lipids are crucial for overwinter survival and successful reproduction. The timing and magnitude of seasonal lipid storage should therefore vary predictably among fish with different thermal preferences and spawn times. Small‐ and large‐bodied fish should also face different constraints associated with season that influence lipid cycling. However, much work to date has been species‐ and location‐specific and a general conceptual model for the seasonal energy budgets of freshwater fish is lacking. Here, we conducted a comprehensive literature review of seasonal lipid levels in freshwater fishes. We predicted that warm and cool water species would be more likely to demonstrate peak lipid levels during warm months than cold water species, and expected a larger magnitude of annual lipid cycling in warm and cool water compared to cold water fish. We also expected dampened lipid cycling in larger fish due to their lower mass‐specific metabolic rates. Observed patterns in the timing and magnitude of lipid storage contradicted our prediction because lipid cycling was widespread across species, despite thermal guild, with peak lipid levels commonly occurring during warmer months, even in cold water fish. For body size effects, larger bodied fish species had dampened seasonal lipid cycling, as predicted. We developed a conceptual framework describing how the ‘scope’ for variation in annual lipid cycling changes with body size both among and within species in order to guide future work. Together, our findings suggest that energy acquired during warm months is broadly important for overwinter survival and reproduction in fishes, and provide a new perspective on the differential constraints and physiological responses to seasonality among freshwater fish. Improving our understanding of these dynamics is especially pressing given that a changing global climate is anticipated to alter existing seasonal signals.     

Recruitment and growth of two small‐bodied resident fish species (Gobiidae and Atherinidae) in oligohaline,seasonally open lagoons

Spatio‐temporal recruitment patterns, growth and survival of the Swan River goby Pseudogobius olorum and western hardyhead Leptatherina wallacei are described from two small, coastal lagoons on the south coast of Western Australia. In these lagoons, estuarine salinity dynamics were relatively stable over much of the autumn–spring period when freshwater inputs from rivers were reduced and there was no oceanic connection. Preflexion and flexion stages of both fish species contributed strongly to population size structure in downstream reaches, whereas upstream reaches were dominated by postflexion larvae and juvenile stages. Spawning of both species was protracted and largely asynchronous, although the episodic presence of stronger preflexion and flexion cohorts suggested some synchronized spawning had occurred. Comparison with estuarine conditions over this period provided evidence that synchronized spawning may be related to temperature and salinity variations from a combination of freshwater inputs and periods of marine exchange. Uninterrupted growth and the progression of cohorts through to juvenile stages were consistent with the generally stable estuarine conditions. Larval and juvenile stages of both species were also tolerant of abrupt changes in salinity and temperature, which occurred due to a non‐seasonal oceanic connection. These findings were consistent with the euryhaline nature of adults of both species.     

Changes of fish assemblages after construction of an estuary barrage in the lower Nakdong River,South Korea

In the Nakdong River, an estuary barrage was constructed in 1987, and it divided the freshwater and the seawater, resulting in a change in the ecosystem. To estimate the impact of barrage construction on fish assemblages, we evaluated 20 years of monitoring data before and after the construction of the estuary barrage, and evaluated the role of fishways. The barrage construction generated entire changes of fish assemblages. After construction, the number of fish species dropped sharply, and 36 species disappeared. Conversely, 18 species appeared at this site, including eight freshwater species, seven of which were exotic or translocated species. Barrage construction affected freshwater fish more severely than it did estuarine and marine species because of the existence of an estuarine environment below the barrage. We did not detect any evidence of recovery of fish assemblages. A total of 31 species were collected at fishways, and the number of individuals collected at each fishway was positively correlated with the amount of discharge from the estuary barrage, mean daily tide level, and water temperature. Migratory fish using a fishway had obvious occurrence periods. Therefore, the efficiency of fishway use can be increased if an appropriate management plan is prepared and implemented.     

Shock,Stress or Signal? Implications of Freshwater Flows for a Top-Level Estuarine Predator

Physicochemical variability in estuarine systems plays an important role in estuarine processes and in the lifecycles of estuarine organisms. In particular, seasonality of freshwater inflow to estuaries may be important in various aspects of fish lifecycles. This study aimed to further understand these relationships by studying the movements of a top-level estuarine predator in response to physicochemical variability in a large, temperate south-east Australian estuary (Shoalhaven River). Mulloway (Argyrosomus japonicus, 47–89 cm total length) were surgically implanted with acoustic transmitters, and their movements and migrations monitored over two years via fixed-position VR2W acoustic receivers configured in a linear array along the length of the estuary. The study period included a high degree of abiotic variability, with multiple pulses (exponentially high flows over a short period of time) in fresh water to the estuary, as well as broader seasonal variation in flow, temperature and conductivity. The relative deviation of fish from their modal location in the estuary was affected primarily by changes in conductivity, and smaller fish (n = 4) tended to deviate much further downstream from their modal position in the estuary than larger fish (n = 8). High-flow events which coincided with warmer temperatures tended to drive mature fish down the estuary and potentially provided a spawning signal to stimulate aggregation of adults near the estuary mouth; however, this relationship requires further investigation. These findings indicate that pulse and press effects of freshwater inflow and associated physicochemical variability play a role in the movements of mulloway, and that seasonality of large freshwater flows may be important in spawning. The possible implications of river regulation and the extraction of freshwater for consumptive uses on estuarine fishes are discussed.     

Distribution and abundance of fishes in the San Francisco Bay estuary between 1980 and 1982

In 1980 a long-term study of the fishery resources of the San Francisco Bay estuary was initiated in an effort to delineate the importance of freshwater inflow to fish and invertebrate abundance and distribution in the bay. An analysis of the trawl data collected between January 1980 and December 1982 illustrates the influence of the timing and magnitude of freshwater inflows on fish fistribution and abundance in this estuary from the perspective of monthly, seasonal and annual time scales. Normally found in the delta, Suisun Bay and San Pablo Bay during periods of increased salinity, pelagic species moved downstream after the two peak flows studied, while demersal species usually found in Central San Francisco Bay moved upstream. Such upstream movements may be due in part to transport by strong density-driven currents.Timing and magnitude of monthly catches of some species varied on a seasonal cycle coincident with variations of freshwater inflow. Most species, especially the marine species, showed no consistent cycle of monthly catches. In the wet years of 1980 and 1982 the distributions of freshwater, estuarine and anadromous species were extended downstream into San Pablo, Central and South San Francisco Bays and some marine species, including the flatfish, were more abundant in the upstream areas. In the dry year of 1981 when bay salinities were higher, few marine species extended their distributions upstream into San Pablo and Suisun Bays. Jacksmelt was the only fish of the 15 most abundant species with its peak abundance in 1981. Most marine species were more abundant in the San Francisco Bay estuary in the wet years.     

Composition and intra-annual variation of the macroinfauna in the estuarine zone of the Pando Stream (Uruguay).

We studied the temporal patterns of macroinfaunal distribution in the estuarine portion of Pando Stream, Uruguay. We found a very low number of species and a high dominance of the polychaete Heteromastus similis. There were clear seasonal patterns in abundance with almost all species peaking in autumn, coinciding with the periods of higher salinity. Minimal values were found in winter and spring when salinity was at a minimum. We suggest that the seasonal pattern is a consequence of the combined effects of recruitment, and of high osmotic stress and disturbance during the periods of freshwater discharge that are common in winter and spring.     

Water use patterns of estuarine vegetation in a tidal creek system

Water availability is a key determinant of the zonation patterns in estuarine vegetation, but water availability and the use of different water sources over space and time are not well understood. We have determined the seasonal water use patterns of riparian vegetation over an estuarine ecotone. Our aim was to investigate how the water use patterns of estuarine vegetation respond to variations in the availability of tidal creek water and rain-derived freshwater. The levels of natural stable isotopes of oxygen and hydrogen were assessed in the stem of the mangrove Avicennia marina (tall and scrub growth forms), Casuarina glauca and Melaleuca quinquenervia that were distributed along transects from river/creek-front towards inland habitats. The isotopic composition of plant tissues and the potential water sources were assessed in both the wet season, when freshwater from rainfall is present, and the dry season, when mangrove trees are expected to be more dependent on tidal water, and when Casuarina and Melaleuca are expected to be dependent on groundwater. Our results indicate that rainwater during the wet season contributes significantly to estuarine vegetation, even to creek-side mangroves which are inundated by tidal creek water daily, and that estuarine vegetation depends primarily on freshwater throughout the year. In contrast, high intertidal scrub mangroves were found to use the greatest proportion of tidal creek water, supplemented by groundwater in the dry season. Contrary to prediction, inland trees C. glauca and M. quinquenervia were found also to rely predominantly on rainwater—even in the dry season. The results of this study reveal a high level of complexity in vegetation water use in estuarine settings.     

Lipid and fatty acid composition during embryo and larval development of puye Galaxias maculatus Jenyns, 1842, obtained from estuarine, freshwater and cultured populations

Galaxias maculatus eggs and larvae obtained from broodfish captured either in an estuarine or a freshwater environment, as well as from cultured broodstock were analysed to compare their lipid and fatty acid profiles. Results showed a lower lipid content in embryos and larvae from estuarine populations than those from fresh water, denoting the influence of environmental conditions. The n-3:n-6 ratio was higher in eggs from estuarine and cultured populations, being in the range of marine fishes, whereas for eggs from freshwater fish was lower and typical of freshwater fishes. The polyunsaturated fatty acids (PUFA), particularly docosahexaenoic acid (22:6n-3) and eicosapentaenoic acid (20:5n-3), were higher in eggs and larvae of broodstock coming from culture or estuarine environments than in those from fresh water. Moreover, these fatty acids markedly increased after hatching in larvae coming from estuarine populations, suggesting the effect of the environment on fatty acid profiles to physiologically prepare the larvae to adapt to higher salinity conditions. Linoleic acid (18:2n-6) content was higher in fresh water fish and its reduction during embryo and larval development was accompanied by a significant increase of arachidonic acid (20:4n-6), which was not observed in embryos or larvae from broodstock fish from estuary or aquaculture origin. Both environment and diet of broodstock fish affected lipid and fatty acid composition of G. maculatus embryo and larvae as well as their changes during development.     

Temporal abundance patterns and growth of juvenile herring and sprat from the Thames estuary 1977–1992

Most herring Clupea harengus and sprat Sprattus sprattus sampled from West Thurrock power station intake screens, middle Thames estuary, between 1977 and 1992 were age-0 and followed regular patterns of seasonal occurrence. Juvenile herring entered the estuary in July, peaked in abundance November to March, and then declined. Juvenile sprat first appeared in September and peaked in abundance in January. Neither species was abundant in summer samples. While in the estuary, herring and sprat increased in length an average of 4·0 and 0·33 cm respectively. Abundance of both species was significantly affected by temperature, temporal trend, shoaling behaviour and seasonal variables, and of herring by suspended solids. Interactions between environmental variates did not influence the abundance of either species. As estuarine clupeids were influenced by a complex set of events within and outside the estuary, estuarine monitoring studies alone will not be sufficient for understanding the changes in estuarine fish communities resulting from future human activity.     

The influence of freshwater flows on two estuarine resident fish species show differential sensitivity to the impacts of drought,flood and climate change

Estuaries are particularly susceptible to climate change and drought resulting in atypical changes to freshwater flows. How such changes in flow impact on the ecology of estuarine fishes may depend on how a species moves in response to changing flow conditions. Acoustic telemetry was used to interpret fine-scale movements of two co-inhabiting estuarine fish species, black bream, Acanthopagrus butcheri and estuary perch, Macquaria colonorum in relation to freshwater flows, season and moon phase. We found black bream to be highly mobile, regularly travelling the length of the estuary and into the neighbouring estuaries. In contrast, estuary perch had particular home ranges and made occasional, upstream or downstream movements. Possibly influenced by freshwater flows, estuary perch moved at greater rates in the Tambo compared to fish in the Mitchell. Black bream resided in the upper estuary during winter and spring and the lower estuary during summer and autumn, whereas estuary perch remained in the upper estuary throughout the year, with occasional downstream movements in winter and spring. This study revealed 1) significantly large increases in freshwater flows result in mass downstream movements in both species, 2) fish moved upstream during full moons and 3) there are contrasting spatio-temporal patterns in movement between species. The results from this study highlight that estuarine fishes are likely to show differential sensitivity to the impacts of drought and climate change and illustrate how acoustic telemetry methods can be used to determine the environmental needs of fishes and help efforts to conserve and manage estuaries worldwide.     

Dynamics of the shallow-water fish assemblage of the Patos Lagoon estuary (Brazil) during cold and warm ENSO episodes

In southern Brazil, cold ( La Niña ) and warm ( El Niño ) episodes of the El Niño Southern Oscillation (ENSO) phenomenon cause drought and high rainfall, respectively. The low precipitation and freshwater outflow associated with La Niña during 1995–1996 were associated with an increase in the abundance of marine species in the Patos Lagoon estuary. During the 1997–1998 El Niño , high precipitation and river discharge were associated with low abundance of marine species in the estuary. ANOVA results showed a higher abundance during La Niña than El Niño for estuarine resident (RES) and estuarine dependent (DEP) fishes. During La Niña catch per unit of effort (CPUE) of RES increased from the marine to estuarine area, but during El Niño CPUE increased at the marine area and diminished during summer and autumn in some estuarine sites. DEP fishes had an opposite abundance pattern. During La Niña , these fishes were abundant at the coastal marine area and along some estuarine sites, but during El Niño , CPUE remained almost the same at the marine area but dropped along some estuarine sites. These different abundance patterns for dominant fish groups yielded a positive interaction between stations and climatic events. With higher river discharge and the consequent decline of dominant euryhaline fishes, such as Mugil platanus and Atherinella brasiliensis , freshwater species increased in abundance and richness in the shallow waters of the stuary. The ENSO phenomenon influences precipitation and estuarine salinity in southern Brazil and thereby seems to have a strong influence on recruitment, immigration, and emigration dynamics of fish species living within and adjacent to estuarine habitats.     

Spatial structure of fish assemblages in a tropical estuarine lagoon: combining multivariate and geostatistical techniques

The spatial structure and seasonal changes of estuarine fish assemblages in the Ciénaga Grande de Santa Marta (CGSM) were analysed based on four seasonal comprehensive surveys conducted in 1993-1994 and 1997. Geostatistical and multivariate techniques were used to: (a) determine seasonal changes in spatial distribution of the species richness, and (b) identify assemblages of estuarine fish and their relation to abiotic factors. Potential biotic interactions affecting the assemblage structure were also explored. A total of 11075 individuals representing 39 species were collected, with Eugerres plumieri, Diapterus rhombeus, Micropogonias furnieri, Mugil incilis, Cathorops spixii, Elops saurus and Anchovia clupeoides as dominant species between seasons. Spatial distribution of fish richness differed between rainy and dry seasons in each year, whereas species mapping showed spatial patchiness in 1993-1994 and gradients during 1997. Strong evidence of species saturation was found in all seasons, suggesting biotic interactions limiting species richness at a threshold density of ca. 50 ind./5000 m². Marine, marine-estuarine and freshwater species were classified in each season according to their capability to cope with salinity fluctuations. Associations defined by functional feeding guilds were also identified. Empirical and statistic evidence showed that fish assemblages differed between seasons within each year, and each assemblage was always dominated by a small number of species, notably E. plumieri in both years 1993-1994 and 1997. Between-season differences in fish assemblage structure in the CGSM seem to be driven by abiotic factors; however, evidence of species saturation could suggest the existence of density-dependent factors operating together.