Towards a more general species–area relationship: diversity on all islands, great and small

Aim

To demonstrate a new and more general model of the species–area relationship that builds on traditional models, but includes the provision that richness may vary independently of island area on relatively small islands (the small island effect).

Location

We analysed species–area patterns for a broad diversity of insular biotas from aquatic and terrestrial archipelagoes.

Methods

We used breakpoint or piecewise regression methods by adding an additional term (the breakpoint transformation) to traditional species–area models. The resultant, more general, species–area model has three readily interpretable, biologically relevant parameters: (1) the upper limit of the small island effect (SIE), (2) an estimate of richness for relatively small islands and (3) the slope of the species–area relationship (in semi‐log or log–log space) for relatively large islands.

Results

The SIE, albeit of varying magnitude depending on the biotas in question, appeared to be a relatively common feature of the data sets we studied. The upper limit of the SIE tended to be highest for species groups with relatively high resource requirements and low dispersal abilities, and for biotas of more isolated archipelagoes.

Main conclusions

The breakpoint species–area model can be used to test for the significance, and to explore patterns of variation in small island effects, and to estimate slopes of the species–area (semi‐log or log–log) relationship after adjusting for SIE. Moreover, the breakpoint species–area model can be expanded to investigate three fundamentally different realms of the species–area relationship: (1) small islands where species richness varies independent of area, but with idiosyncratic differences among islands and with catastrophic events such as hurricanes, (2) islands beyond the upper limit of SIE where richness varies in a more deterministic and predictable manner with island area and associated, ecological factors and (3) islands large enough to provide the internal geographical isolation (large rivers, mountains and other barriers within islands) necessary for in situ speciation.

     

Four Darwinian themes on the origin,evolution and preservation of island life

Charles Darwin’s observations and insights continue to inspire nearly all scientists who are captivated by both the marvels and the perils of island life. Here I feature four themes inspired by Darwin’s singular insights: themes that may continue to provide valuable lessons for understanding the ecological and evolutionary development of insular biotas, and for conserving the natural character and evolutionary potential of all species restricted to isolated ecosystems (natural or anthropogenic).     

The island rule and a research agenda for studying ecogeographical patterns

We are currently experiencing a resurgence of interest in ecogeographical rules, which describe general trends in morphology and related traits along geographical gradients. In order to develop a more comprehensive understanding of the generality and underlying causal mechanisms for these patterns, we recommend a new, more integrated research agenda. In particular, we recommend studies that simultaneously consider different clines in morphology, geographical ranges and diversity as intricately related phenomena; all being ecological, evolutionary and biogeographical responses of organisms to selection regimes that vary non-randomly over space and time, and among species with different ecological and evolutionary histories.     

A call for a new paradigm of island biogeography

MacArthur and Wilson’s equilibrium theory of island biogeography quickly became the paradigm of the field in the 1960s and has strongly influenced this and other disciplines of ecology and conservation biology for the past three decades. Recently, however, a growing number of ecologists have begun to question whether the theory remains a useful paradigm for modern ecology. We now have a much better appreciation for the complexity of nature and we study patterns that span a very broad range in spatial, temporal and ecological scales. At such scales, assumptions that communities are in equilibrium, that species, islands and intervening landscapes or seascapes are equivalent or homogeneous with respect to factors influencing immigration and extinction, and that in situ speciation can be overlooked become very tenuous. With this in mind, this and other papers of this special feature discuss the principal, conceptual shortcomings of the equilibrium theory and offer some modifications or alternatives to the theory that we hope will eventually lead to a more comprehensive understanding of the forces structuring insular communities.     

Isolation‐driven functional assembly of plant communities on islands

The physical and biotic environment is often considered the primary driver of functional variation in plant communities. Here, we examine the hypothesis that spatial isolation may also be an important driver of functional variation in plant communities where disturbance and dispersal limitation may prevent species from occupying all suitable habitats. To test this hypothesis, we surveyed the vascular plant composition of 30 islands in the Gulf of Maine, USA, and used available functional trait and growth form data to quantify the functional composition of these islands. We categorized species based on dispersal mode and used a landscape metric of isolation to assess the potential role of dispersal limitation as a mechanism of isolation‐driven assembly. We tested for island and species level effects on functional composition using a hierarchical Bayesian framework to better assess the causal link between isolation and functional variation. Growth form composition and the community mean value of functional traits related to growth rate, stress tolerance, and nutrient use varied significantly with island isolation. Functional traits and growth forms were significantly associated with dispersal mode, and spatial isolation was the strongest driver of primary trait variation, while island properties associated with environmental drivers in our system were not strong predictors of trait variation. Despite the species‐level association of dispersal mode and functional traits, dispersal mode only accounted for a small proportion of the overall isolation effect on community‐level trait variation. Our study suggests that spatial isolation can be a key driver of functional assembly in plant communities on islands, though the role of particular dispersal processes remains unclear.     

Species-area and species-distance relationships of terrestrial mammals in the Thousand Island Region

Summary The species-area and species-distance relationships of terrestrial mammals in the Thousand Island Region of the St. Lawrence River are totally consistent with the basic predictions of the equilibrium theory of island biogeography. The power model provides the best fit for the species-area relationship, and the z-value of 0.305 does not differ significantly from Preston's canonical value (0.26). Distance (D) is a normal determinant (Se -D ²) of mammalian richness, and 93% of the variability in richness is accounted for by island area and isolation. The high z-values and poor species-distance correlations reported in previous studies of mammalian island biogeography, rather than evidencing non-equilibrium, are predictions consistent with the equilibrium theory for distant archipelagoes or, equivalently, poor immigrators such as mammals.     

A species-based theory of insular zoogeography

• 1 I present an alternative to the equilibrium theory of island biogeography, one which is based on the premise that many of the more general patterns in insular community structure result from, not despite, nonrandom variation among species.
• 2 For the sake of simplicity, the model is limited to patterns and processes operating over scales of ecological space and time: evolution is not included in the current version of the model.
• 3 The model assumes, as did MacArthur and Wilson’s model, that insular community structure is dynamic in ecological time, but the model does not assume a balance, or equilibrium, of immigration and extinction.
• 4 The model presented here is hierarchical, phenomenological (it requires little parameterization beyond that which is directly derived from distributional data), graphical, and it includes potential feedback processes (including interspecific interactions).
• 5 The model offers an alternative explanation for a variety of patterns ranging from distributions of individual species, species–area and species–isolation relationships, to patterns of assembly of insular communities. The model also generates some new predictions and identifies some potentially important areas for future studies.