Andy J. Green

MASS/LENGTH RESIDUALS: MEASURES OF BODY CONDITION OR GENERATORS OF SPURIOUS RESULTS?

In studies of animal ecology, it is fashionable to use the residuals from an ordinary least squares (OLS) linear regression of body mass against a linear measure of size (the body size indicator, BSI) as an index of body condition. These residual indices are used to study the relationship between condition and reproductive investment, survi- vorship, habitat use, and other parameters. I identify a series of key assumptions underlying the use of this method, each of which is likely to be violated in some or all studies. These assumptions are: (1) that the functional relationship between mass and BSI is linear; (2) that condition is independent of BSI length; (3) that BSI length accurately indicates struc- tural size; (4) that there is no correlation between the size of BSI relative to other structural components (i.e., shape) and the parameter against which the residuals are analyzed; (5) that BSI length is strictly independent of mass; and (6) that BSI length is not subjected to error. Violations of these assumptions place the results of some studies in question and explain the poor relationship observed between OLS residuals and more direct measures of condition. I use avian morphometric data to illustrate how OLS methods can easily lead to Type I and Type II errors through violations of assumptions (5) and (6). Significant relationships reported between OLS residual indices and parameters correlating with body size (e.g., size of sexual ornaments or egg size) are at particular risk of being spurious when the correlation coefficient between mass and BSI is low. Residual indices of condition are often likely to be more reliable when calculated with alternative methods such as nonparametric or model II regression. However, whatever the method used to produce them, residual indices are not suitable for studying the heritability of condition.

Implications of waterbird ecology for the dispersal of aquatic organisms

In this paper, we review some potential implications of waterbird ecology for their role as dispersers of aquatic plants and invertebrates. We focus particularly on internal transport (endozoochory) by the Anatidae (mainly ducks) and shorebirds, groups especially important for dispersal processes owing to their abundance, migratory habitats and diets. We conduct a literature review to assess the seasonal patterns shown by Anatidae in consumption of seeds and plankton, the interspecific patterns in such consumption (including the effects of body size, bill morphology, etc.), and differences in habitat use (e.g., shoreline vs. open water specialists) and migration patterns between species (e.g., true migrants vs. nomads). We show that many shorebirds are important consumers of seeds as well as plankton, and suggest that their role in plant dispersal has been underestimated. This review confirms that Anatidae, shorebirds and other waterbirds have great potential as dispersers of aquatic organisms, but illustrates how closely related, sympatric bird species can have very different roles in dispersal of specific aquatic organisms. Furthermore, great spatial and temporal variation is likely in dispersal patterns realized by a given bird population. We present evidence suggesting that northbound dispersal of aquatic propagules by endozoochory during spring migration is a frequent process in the northern hemisphere. Much more systematic fieldwork and reanalysis of the existing data sets (e.g., from diet studies) are needed before the relative roles of various waterbird species as dispersers can be fully assessed.

Biotic wetland connectivity—supporting a new approach for wetland policy

Wetlands are key habitats connected physically and socially with processes occurring over a much wider territory. The biotic connection through dispersal mechanisms among wetlands is of primary importance to wetland management and policies. However, traditional wetland conservation approaches are based on the preservation of isolated sites considered to be of special importance (typically owing to their importance for concentrations of migratory waterbirds). Research linking local species richness and bird migration suggests that the effect of wetland loss on regional diversity might be much larger than what would be expected from direct habitat loss. Since the biotic connection among wetlands serviced by waterbirds appears to be more efficient within a limited range, the distribution of wetlands in space is a key aspect determining wetland connectedness even in the absence of direct hydrologic links. Protected areas should thus be defined with regard to waterfowl movements and waterbird migration as functional processes contributing to aquatic species migration and local species richness. This calls for a regional approach to wetland management within a continental context. This paper aims at defining an operational view of the dispersion function of wetlands and its implication for conservation policies. For this purpose, we examined the conservation policies of the Ramsar Convention (the international treaty that protects wetlands) and the European Union (as an example of relevant continental level policy-making) from the viewpoint of bird-mediated dispersal of aquatic organisms. We propose nine specific avenues for the inclusion of bird-mediated dispersal in the policy documents examined. Non-governmental organisations and other organisations working in waterbird conservation should also recognise the importance of their policies for aquatic biodiversity at broader levels and avoid compartmentalising their conservation activities.

Invertebrate Eggs Can Fly: Evidence of Waterfowl‐Mediated Gene Flow in Aquatic Invertebrates

Waterfowl often have been assumed to disperse freshwater aquatic organisms between isolated wetlands, but no one has analyzed the impact of this transport on the population structure of aquatic organisms. For three cladocerans (Daphnia ambigua, Daphnia laevis, and Sida crystallina) and one bryozoan (Cristatella mucedo), we estimated the genetic distances between populations across North America using sequences of several mitochondrial DNA genes and genotypic frequencies at allozyme and microsatellite loci. Waterfowl movements across North America (estimated from band recovery data) explained a significant proportion of the gene flow occurring between populations across the continent for three of the four species, even after controlling for geographic distances between localities. The fourth species, S. crystallina, has propagules less likely to survive desiccation or ingestion by birds. Differences in the capacity to exploit bird‐mediated transport are likely to have important consequences for the ecology of aquatic communities and the spread of invasive species.

High dispersal capacity of a broad spectrum of aquatic invertebrates via waterbirds

Abstract.Speculation about the role of waterbirds in the dispersal of aquatic invertebrates pre-dates Darwin. However, there is a critical shortage of field studies quantifying such dispersal. We quantified the viability of aquatic invertebrates in the faeces of different waterfowl species collected in the field at different times during winter. Faeces were collected from four duck species (Northern Pintail Anas acuta, Mallard A. platyrhynchos, Shoveler A. clypeata, Eurasian Teal A. crecca) and Eurasian Coot Fulica atra in November 2004 and January 2005. We also collected soil samples from resting sites as an indicator of what may be transported on birds’ feet and plumage. Faecal and soil samples were incubated using two treatments (0.4 and 4.0 mS cm-1) to quantify the potential for dispersal between aquatic habitats of different salinities. We found that viable Nematoda, Rotifera, Copepoda, Ostracoda, Insecta (Tipulidae), and Daphnia and Moina cladocerans were transported internally by birds in the wild. We also found evidence that nematodes, rotifers, ostracods, copepods, tipulids, chironomids and hemipterans can be dispersed on birds’ feet and feathers. The overall incidence of hatching from all samples was higher in January (59.4%) than in November (11.5%). With the exception of bdelloid rotifers, we found no evidence that the potential for dispersal between two habitats would be impeded by salinity in the range tested. Our data suggest that the taxonomic range of dispersed invertebrates and the frequency of their dispersal via waterfowl has previously been underestimated.

Ecosystem services provided by waterbirds

Ecosystem services are ecosystem processes that directly or indirectly benefit human well‐being. There has been much recent literature identifying different services and the communities and species that provide them. This is a vital first step towards management and maintenance of these services. In this review, we specifically address the waterbirds, which play key functional roles in many aquatic ecosystems, including as predators, herbivores and vectors of seeds, invertebrates and nutrients, although these roles have often been overlooked. Waterbirds can maintain the diversity of other organisms, control pests, be effective bioindicators of ecological conditions, and act as sentinels of potential disease outbreaks. They also provide important provisioning (meat, feathers, eggs, etc.) and cultural services to both indigenous and westernized societies. We identify key gaps in the understanding of ecosystem services provided by waterbirds and areas for future research required to clarify their functional role in ecosystems and the services they provide. We consider how the economic value of these services could be calculated, giving some examples. Such valuation will provide powerful arguments for waterbird conservation.

PLANT PERFORMANCE ACROSS LATITUDE: THE ROLE OF PLASTICITY AND LOCAL ADAPTATION IN AN AQUATIC PLANT

Geographic variation can lead to the evolution of different local varieties within a given species, therefore influencing its distribution and genetic structure. We investigated the contribution of plasticity and local adaptation to the performance of a common aquatic plant (Potamogeton pectinatus) in contrasting climates, using reciprocal transplants at three experimental sites across a latitudinal cline in Europe. Plants from 54 genets, originally collected from 14 populations situated within four climatic regions (sub- arctic, cold temperate, mild temperate, and mediterranean) were grown in three different localities within three of these regions (cold temperate, Norway; mild temperate, The Netherlands; mediterranean, Spain). Tuber production was highest for the mild-temperate genets, irrespective of locality where the genets were grown. Selection coefficients indicated that populations at the European center of the species distribution perform better than all other populations, at all sites. However, marginal populations showed changes in life-history traits, such as compressed life cycles in the north and true perenniality in the south, that may allow them to perform better locally, at the limits of their distribution range. Our results thus suggest that local adaptation may overlap spatially with center-periphery gra- dients in performance caused by genetic factors (such as genetic drift and inbreeding in range-marginal populations).

The American brine shrimp as an exotic invasive species in the western Mediterranean

The hypersaline environments and salterns present in the western Mediterranean region (including Italy, southern France, the Iberian Peninsula and Morocco) contain autochthonous forms of the brine shrimp Artemia, with parthenogenetic diploid and tetraploid strains coexisting with the bisexual species A. salina. Introduced populations of the American brine shrimp A. franciscana have also been recorded in these Mediterranean environments since the 1980s. Based on brine shrimp cyst samples collected in these countries from 1980 until 2002, we were able to establish the present distribution of autochthonous brine shrimps and of A. franciscana, which is shown to be an expanding invasive species. The results obtained show that A. franciscana is now the dominant Artemia species in Portuguese salterns, along the French Mediterranean coast and in Cadiz bay (Spain). Co-occurrence of autochthonous (parthenogenetic) and American brine shrimp populations was observed in Morocco (Mar Chica) and France (Aigues Mortes), whereas A. franciscana was not found in Italian cyst samples. The results suggest these exotic A. franciscana populations originate as intentional or non-intentional inoculations through aquacultural (hatchery effluents) or pet market activities, and suggest that the native species can be rapidly replaced by the exotic species.

Creating a safe operating space for iconic ecosystems

Manage local stressors to promote resilience to global change Although some ecosystem responses to climate change are gradual, many ecosystems react in highly nonlinear ways. They show little response until a threshold or tipping point is reached where even a small perturbation may trigger collapse into a state from which recovery is difficult (1). Increasing evidence shows that the critical climate level for such collapse may be altered by conditions that can be managed locally. These synergies between local stressors and climate change provide potential opportunities for proactive management. Although their clarity and scale make such local approaches more conducive to action than global greenhouse gas management, crises in iconic UNESCO World Heritage sites illustrate that such stewardship is at risk of failing.

The conservation status of Moroccan wetlands with particular reference to waterbirds and to changes since 1978

Morgan made detailed descriptions of 24 major Moroccan wetlands visited in 1978, with a total area of 4529 ha (Morgan, N.C., 1982a. An ecological survey of standing waters in North West Africa: III. Site descriptions for Morocco. Biological Conservation, 24, 161–182.). We revisited these sites, and found that 25% of the wetland area had been destroyed by 1999. This loss was concentrated in wetland types of low salinity ( < 5 g/l NaCl), with a 98% loss of seasonal mesohaline sites, 41% loss of mountain lakes and 33% loss of seasonal Phragmites/Scirpus lacustris marshes. Surviving mountain lakes showed increased conductivities, suggesting reduced inflow. No loss of area of other wetland types was recorded, although degradation has occurred at all sites due to hydrological impacts, overgrazing or excessive reed-cutting, sedimentation, urban development, pollution, introduction of exotic fish and other causes. Similar threats face another 23 major Moroccan wetlands reviewed in this study. Of the 47 wetlands studied in total, only 10 have some kind of protection status. We assess the importance of these wetlands for waterbirds and aquatic submerged or floating plants. The number of plant species recorded is strongly correlated with the number of threatened waterbird species (but not the total number of waterbird species). Natural, freshwater wetlands most affected by wetland loss hold more species of aquatic plants and invertebrates, and are of great value for threatened waterbirds such as marbled teal (Marmaronetta angustrirostris), ferruginous duck (Aythya nyroca), ruddy shelduck (Tadorna ferruginea) and especially the crested coot (Fulica cristata). Most surviving natural, fresh wetlands are unprotected, and measures to conserve them are urgently required. Humanmade wetlands such as reservoirs have some value for threatened waterbirds (especially ruddy shelduck and marbled teal), but hold much lower densities of waterbirds than natural wetlands, and support fewer plant species. Thus, they do not compensate for the continuing loss of natural wetlands. # 2002 Elsevier Science Ltd. All rights reserved.