Fish species richness in spring‐fed ponds: effects of habitat size versus isolation in temporally variable environments

1 Environmental fluctuations can hypothetically promote greater species diversity, but this has rarely been experimentally tested in plant communities. We predicted that high temporal variability in soil moisture would promote high diversity in a community of recruiting seedlings. Using a glasshouse experiment designed to imitate field conditions in species-rich limestone pavement habitats, we examined the effects of temporal variability and total quantity of water provided on seedling species richness, evenness and composition within a single growing season. 2 Temporal variability was altered by varying the frequency at which water was added while keeping the total quantity of water constant. Low temporal variability was created by high frequencies of water addition. High variability was associated with more extreme fluctuations between droughted and flooded conditions. Each of three variability treatments was carried out using three total quantities of water added to create a gradient in total soil water availability. 3 Species richness was greatest in higher quantity and lower variability treatments. Higher total quantities of water added buffered the negative effects of higher temporal variability, and lower temporal variability buffered the negative effects of lower quantity. Species richness was negatively correlated with temporal variability in soil moisture content both in the glasshouse and in a field study. Evenness was negatively correlated with richness. Low richness, high evenness communities had low densities and were prone to extinction. Thus, neither richness nor evenness was maintained by higher temporal variabilities within a growing season. 4 Lower species richness under more temporally variable conditions was attributable to both random extinctions due to reductions in total community density and species-specific differences in mortality. Final species relative abundances differed between water treatments, but there were no separate effects of variability in soil moisture content or total quantity of water added. Upland species were more abundant in treatments where drought was more prevalent, either due to low total quantities of water or high temporal variability in soil moisture. Spatial heterogeneity in moisture supply, or interannual fluctuations between dry and wet years, may thus contribute to coexistence of drought-tolerant and flood-tolerant species in the field.

We tested the hypothesis that higher temporal variability in water supply will promote higher species richness of germinating and surviving seedlings using assemblages of 70 species of herbaceous plants from limestone pavement habitats. In a two-factor greenhouse experiment, doubling the total volume of water added led to greater germination (measured as number of germinated seeds and species) and establishment (survival and biomass) but the effects of temporal variability depended on the response variable considered. Low pulse frequencies of water addition with total volume added held constant resulted in greater temporal variability in soil moisture concentration that in turn promoted higher density and richness of germinated seedlings. Low pulse frequencies caused an eight-fold greater mortality in the low total volume treatment and biomass production to decline by one-third in the high total volume treatment. The effects of increasing temporal variability in water supply during recruitment stages can thus be opposite on different components of plant fitness and may also depend on total resource quantity. While greater species richness in more temporally variable soil moisture conditions was attributable to sampling effects rather than species-specific responses to the water treatments, species relative abundances did vary significantly with temporal variability. Changes in the amplitude or frequency of resource fluctuations may alter recruitment patterns, and could have severe and relatively rapid effects on community structure in unproductive ecosystems.

Relationships between local annual immigration and extinction rates of plant species and total species richness were determined from long-term data in permanent plots in tallgrass and shortgrass prairies in Kansas. Combining plots resulted in higher equilibrium numbers of species as predicted from immigration and extinction rates. Immigration and extinction rates also increased with scale. Extinction rates are higher because the regional scale supports more rare species which, in turn, have high probabilities of extinction. We also tested the hypotheses that extinction rates would be higher on burned versus unburned grasslands, and that immigration rates would be higher on grazed versus ungrazed grasslands. Extinction rates were positively correlated with the number of species at a site, and this relationship was not altered by burning or grazing. Immigration rates were variable, but were sometimes positively correlated with growing season precipitation. Immigration rates decreased in years sites were burned. Therefore, after fire, the number of species going locally extinct was still dependent on earlier species richness, but the number of species added to the site was reduced. Variances in immigration and extinction rates were high, therefore, confident predictions regarding the effects of burning or grazing regimes on species richness could not be made. Variance in rates of immigration and extinction results in a range of values within which the equlibrium number of species fluctuates randomly.

Stream landscapes are highly variable in space and time and, like terrestrial landscapes, the resources they contain are patchily distributed. Organisms may disperse among patches to fulfill life-history requirements, but biotic and abiotic factors may limit patch or locality occupancy. Thus, the dynamics of immigration and extinction determine, in part, the local structure of assemblages. We sampled fishes and stream habitat at 12 localities for two years (96 samples) to examine the deterministic nature of immigration and extinction processes in stream fish assemblages. Mean immigration rates for assemblages were highest at large stream localities, where the pool of potential immigrants was largest. Mean extinction rates were highest where variability in the flow regime was high, though local refugia appeared to modify the extinction process at one locality. Significant nested subset patterns in species composition occurred over time for 7 of the 12 localities. The strength of the nesting was associated with mean immigration and extinction rates. Higher extinction rates corresponded to stronger nestedness, whereas higher immigration rates were associated with weaker nestedness. Across all species, both immigration and extinction rates were strongly associated with mean abundance. Species with high local abundances had higher immigration rates and lower extinction rates than did species with low local abundances. There were no significant associations between trophic guild or body size and immigration and extinction rate. This work supports the hypothesis that immigration and extinction rates for assemblages are predictable along environmental gradients, and that species are less prone to local extinction and more prone to colonize areas when they maintain high local abundances. The extinction process in local assemblages can be a highly ordered event leading to strong nested subset patterns, but immigration appears to be more stochastic.

Precision Agriculture (PA) recognizes and manages intra-field spatial variability to increase profitability and reduced environmental impact. Site Specific Crop Management (SSCM), a form of PA, subdivides a cropping field into uniformly manageable zones, based on quantitative measurement of yield limiting factors. In Mediterranean environments, the spatial and temporal yield variability of rain-fed cropping system is strongly influenced by the spatial variability of Plant Available Water-holding Capacity (PAWC) and its strong interaction with temporally variable seasonal rainfall. The successful adoption of SSCM depends on the understanding of both spatial and temporal variabilities in cropping fields. Remote sensing phenological metrics provide information about the biophysical growth conditions of crops across fields. In this paper, we examine the potential of phenological metrics to assess the spatial and temporal crop yield variability across a wheat cropping field at Minnipa, South Australia. The Minnipa field was classified into three management zones using prolonged observations including soil assessment and multiple year yield data. The main analytical steps followed in this study were: calculation of the phenological metrics using time series NDVI data from Moderate Resolution Imaging Spectroscope (MODIS) for 15 years (2001-2015); producing spatial trend and temporal variability maps of phenological metrics; and finally, assessment of association between the spatial patterns and temporal variability of the metrics with management zones of the cropping field. The spatial trend of the seasonal peak NDVI metric showed significant association with the management zone pattern. In terms of temporal variability, Time-integrated NDVI (TINDVI) showed higher variability in the “good” zone compared with the “poor” zone. This indicates that the magnitude of the seasonal peak is more sensitive to soil related factors across the field, whereas TINDVI is more sensitive to seasonal variability. The interpretation of the association between phenological metrics and the management zone site conditions was discussed in relation to soil-climate interaction. The results demonstrate the potential of the phenological metrics to assess the spatial and temporal variability across cropping fields and to understand the soil-climate interaction. The approach presented in this paper provides a pathway to utilize phenological metrics for precision agricultural management application.

Temporal Environmental Variability Drives the Evolution of Cooperative Breeding in Birds

In dry areas, natural and artificial ponds experience frequent water level fluctuation, affecting conditions for some aquatic and amphibiotic taxa. Water beetles, bugs, and dragonflies make up much of pond diversity, and are responsive to changes in environmental conditions. Using a drought-prone pondscape within the Greater Cape Floristic Region biodiversity hotspot, we determine (1) the relative extent to which species richness, abundance, and composition are affected by pond water level fluctuation, (2) the effects of environmental variables and vegetation characteristics relative to fluctuating water levels, and (3) make recommendations to improve pondscape conservation. We found that the degree of fluctuation had a significant effect on beetle species richness, but had no significant effect on the other focal taxa. Water temperature, pH, and conductivity, and vegetation cover and composition were drivers of aquatic insect species richness, abundances, and assemblage structures. Habitat heterogeneity supported rich aquatic insect assemblages. We recommend that a range of ponds with various degrees of water level fluctuation should be maintained, along with naturally diverse marginal vegetation. Such a dynamic pondscape can contribute greatly towards maintenance of local and regional aquatic insect diversity in drought-prone regions and should be considered as a main focus in conservation efforts.

Black hornbills (Anthracoceros malayanus) appear to be the principal long-distance seed dispersers of Aglaia sp. (Meliaceae) at Pasoh Forest Reserve in Peninsular Malaysia. The squirrel Callosciurus prevostii removed some of the large seeds at least as far as adjacent crowns and sometimes dropped them after consuming only the orange, oily sarcotesta. It chased other squirrel species, which are probably seed predators, and hornbills out of the fruiting crown. Seeds on the ground beneath the parent crown were removed more rapidly than those farther away by rodent and (possibly) phasianid seed predators. Sitophilus sp. (Curculionidae) was also an Aglaia seed predator or parasite. Aglaia juveniles grew slowly under closed canopy, and small ones (<50 cm in height) had only 9.7 percent mortality per year during a 4-year period. Seedling survival was positively size dependent. For unknown reasons, small seedlings near the parent had a higher death rate than those at a greater distance; this was apparently a long-term pattern, because large seedlings and saplings did not occur within 10 and 35 m, respectively, of the parent tree's base. For this species an advantage of seed dispersal is avoidance of disproportionate seed and seedling mortality near the parent. Although Aglaia grows much faster in gaps than under closed canopy, it may require several episodes of growth in successive gaps before becoming reproductively mature. IN THEIR REVIEW OF SEED DISPERSAL, Howe and Smallwood (1982) remarked that little is known about its ecological and evolutionary advantages to the plant. Here we combine observations on seed dispersal and predation and juvenile mortality and spatial distribution to show how dispersal helps the tropical tree Aglaia sp. (Meliaceae) avoid seed and seedling mortality that is greater near the parent than farther away. Webb et al. (1967), Janzen (1970), and Connell (1971) suggested that such a mortality pattern, generated by seed predators, herbivores, or allelopathy, would promote the maintenance of high tree species richness in tropical forests. More recently, Connell (1978, 1979) has withdrawn his support for this hypothesis because field studies have shown that seed and seedling mortality is not invariably higher near conspecific adults. However, Clark and Clark's (1984) review of 24 data sets on mostly neotropical, woody plants showed that most evidence indicates either densityor distance-dependence in progeny mortality, as originally predicted by Janzen and Connell. Hubbell (1979, 1980) challenged the Janzen-Connell hypothesis on empirical and theoretical grounds, daiming the latter to demonstrate that spacing between conspecific adults could contribute little to the maintenance of high species richness. His theoretical analysis has been questioned by Becker et al. (1985), who consider the issue unresolved for nonequilibrium

Large disturbances can cause rapid degradation of coral reef communities, but what baseline changes in species assemblages occur on undisturbed reefs through time? We surveyed live coral cover, reef fish abundance and fish species richness in 1997 and again in 2007 on 47 fringing patch reefs of varying size and depth at Mersa Bareika, Ras Mohammed National Park, Egypt. No major human or natural disturbance event occurred between these two survey periods in this remote protected area. In the absence of large disturbances, we found that live coral cover, reef fish abundance and fish species richness did not differ in 1997 compared to 2007. Fish abundance and species richness on patches was largely related to the presence of shelters (caves and/or holes), live coral cover and patch size (volume). The presence of the ectoparasite-eating cleaner wrasse, Labroides dimidiatus, was also positively related to fish species richness. Our results underscore the importance of physical reef characteristics, such as patch size and shelter availability, in addition to biotic characteristics, such as live coral cover and cleaner wrasse abundance, in supporting reef fish species richness and abundance through time in a relatively undisturbed and understudied region.

AimMacArthur and Wilson's theory of island biogeography was revolutionary, and also inspired the more recent unified neutral theory of biodiversity and biogeography. The unified neutral theory has the potential to make predictions about island biogeography that are not well studied. Here we aim to unify the two theories by using an ecological neutral model to study immigration and extinction rates on islands – the cornerstone of MacArthur and Wilson's theory.MethodsWe conduct simulations of a spatially implicit neutral model and measure species abundances, immigration rates and extinction rates. We study the behaviour of the model at dynamic equilibrium and on approach to dynamic equilibrium both from volcanic origin (low initial diversity) and from land bridge origin (high initial diversity). We extend the model to study the effects of clustered immigration and to explicitly account for the distinction between immigration and colonization.ResultsOur model, in accord with the simplest version of MacArthur and Wilson's theory, predicts linear immigration and extinction rates as functions of species richness at dynamic equilibrium. In contrast, the approach to dynamic equilibrium produces rich and unexpected behaviour where immigration and extinction rates are non‐monotonic functions of species richness, at odds with other theory. Once examined, however, this behaviour makes biological sense and results from the influence of the species abundance distribution over immigration and extinction rates. The turnover predicted by our first model appears high, but can be lowered to realistic levels with an alternative model of clustered immigration or by accounting for the difference between the immigration of a new species and its true colonization of the island.Main conclusionsMacArthur and Wilson's theory of island biogeography and ecological neutral theory are different, but there are strong similarities in their assumptions and predictions that should not be overlooked when evaluating them. Our results highlight the importance of species abundances as indicators of immigration and extinction rates; species richness alone is insufficient. In particular, extinction rate and species abundances are unavoidably linked, as rarity usually precedes extinction.

Dictated by limited resource availability for land acquisition, a central question in conservation biology is the ability of areas of different size to maintain species diversity. The selected reserves should not only be species rich at the moment, but should also maintain species diversity in the long run. We used two sets of data on vascular plant species in boreal lakes collected in 1933/34 and 1996 to test the relationships between lake area and the extinction, immigration and turnover rates of the species. Moreover, we investigated, whether the number of species in 1933/34 or water connection between lakes was related to extinction, immigration and turnover rates of species. We found that lake area or shoreline length was not correlated with immigration or turnover rate. But extinction rate was slightly negatively correlated with shoreline length. The original number of species was positively related to the number of species extinctions and to the absolute turnover rate in the lakes, which indicates that species richness does not create stability in these communities. Species number was not correlated with immigration rate. Upstream water connections in the lakes did not affect immigration, extinction or turnover rates. We conclude that length of the shoreline is a better measure of suitable area for water plants than the lake area, and that because the correlation between shoreline length and extinction rate was slight, also small lakes can be valuable for conservation.

Unsaturated Fish Communities in African Rivers

AimIsland biogeography theory states that species richness increases with habitat diversity and decreases with isolation from source pools. However, ecological theory must incorporate effects of human activity to explain contemporary patterns of biodiversity. We contemporized island biogeography theory by conceptualizing island trajectories of how species richness changes over time with accelerating land development and economic trade, which increase extinction and immigration rates, respectively. With this contemporized theory, we then articulate and empirically assess expected relationships of native, introduced and total species richness with natural and anthropogenic metrics of habitat diversity and isolation from source pools.LocationGreater Caribbean region.Time periodDatabase finalized in 2020.MethodsWe built a database of 1,042 native and introduced reptiles and amphibians (herps) for 840 Caribbean islands. For each island, we calculated natural and anthropogenic metrics of island habitat diversity and isolation from source pools and used linear model averaging to assess the expected relationships under the contemporized theory for 15 major herp clades.ResultsNatural habitat diversity metrics exhibited positive relationships with native and introduced species richness, strengthening total species richness–area relationships across herp clades. Geographic isolation exhibited negative relationships with native and positive relationships with introduced species richness, weakening total species richness–isolation relationships. Economic area, based on developed land, and economic isolation, based on maritime trade, exhibited negative relationships with native species richness, but positive and negative relationships, respectively, with introduced species richness. Total species richness relationships with these two anthropogenic metrics were strongest in clades with many introduced species.Main conclusionsA contemporized island biogeography theory that includes the effects of land development and economic trade on species extinction and immigration explained current Caribbean herp species richness patterns. As human activity continues to accelerate, the contemporized theory we articulate here will increasingly predict island biogeography of the Anthropocene.

Dispersal processes, such as immigration and extinction rates, and habitat properties play a crucial role in determining species composition and nestedness patterns within communities. In the current global scenario of changing environmental conditions and habitat fragmentation, information on the role of natural dispersal mechanisms and disturbance factors are especially important for understanding dynamics of species composition changes in stream ecosystems. We investigated spatial and temporal patterns of nestedness of fish communities in tropical stream systems of central India and the relationship of various dispersal factors (immigration–extinction rates and their variability) with habitat properties (habitat size and heterogeneity) and anthropogenic disturbances. We tested predictions of the classical model by Schlosser on immigration–extinction dynamics along longitudinal gradients in stream community composition for these streams. The results revealed significant patterns of nestedness for all study sites. Sites exposed to varying degrees of disturbance (induced by human activities) showed lower nestedness than undisturbed ones. Immigration rates did not show strong relations with nestedness but extinction rates were significantly related (negatively) to nestedness. In addition, disturbance played an important role in determining immigration rates and variability in immigration rates. Stream characteristics, such as habitat-size and habitat-variability gradients, were not statistically significant predictors of immigration and extinction rates. Our results demonstrate the influence of local anthropogenic disturbances on dispersal dynamics of species. A reduction in availability of suitable habitats could lower immigrations.

Aim To test for control of vascular plant species richness in the riparian corridor by exploring three contrasting (although not mutually exclusive) hypotheses: (1) longitudinal patterns in riparian plant species richness are governed by local, river‐related processes independent of the regional species richness, (2) riparian plant species richness is controlled by dispersal along the river (longitudinal control), and (3) the variation in riparian plant species richness mirrors variation in regional richness (lateral control).Location The riparian zones of the free‐flowing Vindel River and its surrounding river valley, northern Sweden.Methods We used data from three surveys, undertaken at 10‐year intervals, of riparian reaches (200‐m stretches of riverbank) spanning the entire river. In addition, we surveyed species richness of vascular plants in the uplands adjacent to the river in 3.75‐km2 large plots along the same regional gradient. We explored the relationship between riparian and upland flora, and various environmental variables. We also evaluated temporal variation in downstream patterns of the riparian flora.Results Our results suggest that local species richness in boreal rivers is mainly a result of local, river‐related processes and dispersal along the corridor. The strongest correlation between species richness and the environment was a negative one between species number and soil pH, but pH varied within a narrow range. We did not find evidence for a correlation between species richness on regional and local scales. We found that the local patterns of species richness for naturally occurring vascular plants were temporally variable, probably in response to large‐scale disturbance caused by extreme floods. Most previous studies have found a unimodal pattern of species richness with peaks in the middle reaches of a river. In contrast, on two of three occasions corresponding to major flooding events, we found that the distribution of species richness of naturally occurring vascular plants resembled that of regional diversity: a monotonic decrease from headwater to coast. We also found high floristic similarity between the riparian corridor and the surrounding landscape.Main conclusions These results suggest that local processes control patterns of riparian species richness, but that species composition is also highly dependent on the regional species pool. We argue that inter‐annual variation in flood disturbance is probably the most important factor producing temporal variability of longitudinal species richness patterns.