Reconciling variability and optimal behaviour using multiple criteria in optimization models

A major objective in behavioural and evolutionary ecology is to understand how animals make decisions in complex environments. Examinations of animal behaviour typically use optimization models to predict the choices animals ought to make. The performance of animals under specific conditions is then compared against the predicted optimal strategy. This optimization approach has come into question because model predictions often do not match animal behaviour exactly. This has led to serious scepticism about the ability of animals to exhibit optimal behaviour in complex environments. We show that conventional approaches that compare observed animal behaviour with single optimal values may bias the way we view real-world variation in animal performance. Considerable insight into the abilities of animals to make optimal decisions can be gained by interpreting why variability in performance exists. We introduce a new theoretical framework, called multi-objective optimization, which allows us to examine decision-making in complex environments and interpret the meaning of variability in animal performance. A multi-objective approach defines the set of efficient choices animals may make in attempting to reach compromises among multiple conflicting demands. In a multi-objective framework, we may see variation in animal choices, but, unlike single-objective optimizations where there is one best solution, this variation may represent a range of adaptive compromises to conflicting objectives. An important feature of this approach is that, within the set of efficient alternatives, no choice can be considered to yield higher fitness, a priori, than any other choice. Thus, variability and optimal behaviour may be entirely consistent. We illustrate our point using selected examples from foraging theory where there is already an optimization program in place.     

Effects of low and elevated CO2 partial pressure on growth and reproduction of Arabidopsis thaliana from different elevations

Atmospheric CO₂ partial pressure may have been as low as 18 Pa during the Pleistocene and is expected to increase from 35 to 70 Pa before the end of the next century. Low CO₂ reduces the growth and reproduction of C₃ plants, whereas elevated CO₂ often increases growth and reproduction. Plants at high elevation are exposed to reduced CO₂ partial pressure and may be better adapted to the low CO₂ of the Pleistocene. We examined genotypes of Arabidopsis thaliana from different elevations for variation in growth and reproduction at the CO₂ levels of the Pleistocene, the present and the future. Genotypes exhibited limited genetic variation in the response of the production of biomass to changes in CO₂, but showed significant variation in reproductive characters. We found evidence that plants from high elevations may be better adapted to low CO₂ when considering seed number, which is an important component of fitness. Genotypes showed greater variation in the response of seed number between 35 and 20 Pa CO₂ compared to 35 and 70 Pa CO₂. We conclude that present-day C₃ annuals may have greater potential for evolution in response to the low CO₂ of the Pleistocene relative to the elevated CO₂ predicted for the future.     

Origins, establishment and evolution of new polyploid species: Senecio cambrensis and S. eboracensis in the British Isles

Two new polyploid species of Senecio have originated in the British Isles in recent times following hybridization between native S. vulgaris (2 n  = 40) and introduced S. squalidus (2 n  = 20). One of these is the allohexaploid S. cambrensis (2 n  = 60), the other is the recombinant tetraploid S. eboracensis (2 n  = 40). We review what is known about when and how each species originated, and their reproductive isolation from parents due to high selfing rates. We also review evidence that suggests S. cambrensis may have undergone rapid genome evolution since its origin, and comment on the risks of extinction to each species due to chance factors operating during the early establishment phase. The discovery of both species soon after their origin provides an unparalleled opportunity to examine two different but related forms of speciation following hybridization between the same parent species. Further detailed study of the ecology and genomics of S. cambrensis and S. eboracensis will help improve our understanding of the process of polyploid speciation in plants.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 467–474.     

A landscape perspective of surface–subsurface hydrological exchanges in river corridors

1. River corridors can be visualised as a three‐dimensional mosaic of surface–subsurface exchange patches over multiple spatial scales. Along major flow paths, surface water downwells into the sediment, travels for some distance beneath or along the stream, eventually mixes with ground water, and then returns to the stream. 2. Spatial variations in bed topography and sediment permeability result in a mosaic of patch types (e.g. gravel versus sandy patches) that differ in their hydrological exchange rate with the surface stream. Biogeochemical processes and invertebrate assemblages vary among patch types as a function of the flux of advected channel water that determines the supply of organic matter and terminal electron acceptors. 3. The overall effect of surface–subsurface hydrological exchanges on nutrient cycling and biodiversity in streams not only depends on the proportion of the different patch types, but also on the frequency distribution of patch size and shape. 4. Because nutrients are essentially produced or depleted at the downwelling end of hyporheic flow paths, reach‐scale processing rates of nutrients should be greater in stretches with many small patches (e.g. short compact gravel bars) than in stretches with only a few large patches (e.g. large gravel bars). 5. Based on data from the Rhône River, we predict that a reach with many small bars should offer more hyporheic refugia for epigean fauna than a reach containing only a few large gravel bars because benthic organisms accumulate preferentially in sediments located at the upstream and downwelling edge of bars during floods. However, large bars are more stable and may provide the only refugia during severe flood events. 6. In river floodplain systems exhibiting pronounced expansion/contraction cycles, hyporheic assemblages within newly created patches not only depend on the intrinsic characteristics of these patches but also on their life span, hydrological connection with neighbouring patches, and movement patterns of organisms. 7. Empirical and theoretical evidence illustrate how the spatial arrangement of surface–subsurface exchange patches affects heterogeneity in stream nutrient concentration, surface water temperature, and colonisation of dry reaches by invertebrates. 8. Interactions between fluvial action and geomorphic features, resulting from seasonal and episodic flow pulses, alter surface–subsurface exchange pathways and repeatedly modify the configuration of the mosaic, thereby altering the contribution of the hyporheic zone to nutrient transformation and biodiversity in river corridors.     

Breeding frequency in Grey-headed Albatrosses Thalassarche chrysostoma

Although Grey-headed Albatrosses Thalassarche chrysostoma are usually regarded as biennial breeders, taking a year off following a successful breeding attempt, a small proportion of successful birds attempt to breed annually. This proportion was higher at Marion Island (5.4%) than at Bird Island, South Georgia (1.0%), suggesting that conditions are more favourable at Marion Island. This hypothesis is supported by higher average breeding success and shorter lags following both successful and failed breeding attempts at Marion Island. Factors favouring reproduction at Marion Island may include reduced intraspecific competition (given the much smaller breeding population) and/or more predictable food supply (owing to production of meso-scale eddies associated with the Indian Ocean Ridge). Although annual breeding appeared to increase the risk of adult mortality, with several birds that attempted to breed annually found dead the following year, at least some birds greatly enhanced their reproductive output, with one male raising five chicks in five successive years. Contrary to life-history theory, there was no evidence that older birds were more likely to attempt annual breeding because of declining reproductive value.     

REVISION OF THE ARCHOSAUROMORPH REPTILE TRILOPHOSAURUS, WITH A DESCRIPTION OF THE FIRST SKULL OF TRILOPHOSAURUS JACOBSI, FROM THE UPPER TRIASSIC CHINLE GROUP, WEST TEXAS, USA

Abstract: The first abundant, well‐preserved fossils of the unusual archosauromorph reptile Trilophosaurus jacobsi Murry are from an Upper Triassic bonebed in the lowermost Trujillo Formation of the Chinle Group in Borden County, Texas. A nearly complete left side of the skull and incomplete but articulated mandible of a juvenile individual demonstrate that Trilophosaurus jacobsi Murry is referable to Trilophosaurus, so the putative procolophonid genus Chinleogomphius is a junior objective synonym of Trilophosaurus. Features of T. jacobsi that diagnose it from T. buettneri include asymmetrical, tricuspate teeth in which the central cusp is taller than the marginal cusp, polygonal in cross‐section and displaced slightly lingually; the lingual cusp is low, transversely broadened and anteroposteriorly compressed; teeth with prominent cingula along the mesial and distal margins; and paired sagittal (parasagittal) crests composed of portions of the postfrontals and the parietals, as well as a relatively broad skull roof between the supratemporal fenestrae. We also re‐illustrate and re‐describe the holotypes of both Trilophosaurus buettneri Case and T. jacobsi Murry. T. jacobsi is primarily Adamanian in age, and the lowest occurrence of T. jacobsi is stratigraphically above that of T. buettneri. Therefore, T. buettneri is an Otischalkian–Adamanian index taxon, and T. jacobsi is an Adamanian–?earliest Revueltian index taxon. Trilophosaurus teeth are readily identifiable to species and therefore identifiable as isolated fossils.     

A cochleosaurid temnospondyl amphibian from the Middle Pennsylvanian of Linton, Ohio, U.S.A.

Two temnospondyl amphibian specimens from the Middle Pennsylvanian locality of Linton, Ohio, USA are described as Adatnanterpeton ohioensis gen. et sp. nov. , a member of the family Cochleosauridae and sister-taxon to the clade comprising the genera Cochleosaurus+Chenoprosopus. A. ohioensis is a very rare component in the large Linton vertebrate assemblage and may have been a relatively terrestrial form, perhaps similar to Cochleosaurus florensis from the contemporaneous lycopsid-trunk locality at Florence, Nova Scotia. An overview of the edopoid and eryopoid adaptive radiations is presented and it is argued that they were sequential radiations, each of monophyletic origin filling the same range of niches.