Body cooling and the diving capabilities of muskrats (Ondatra zibethicus): a test of the adaptive hypothermia hypothesis

We tested the hypothesis that immersion hypothermia enhances the diving capabilities of adult and juvenile muskrats by reducing rates of oxygen consumption (V O2). Declines in abdominal body temperature (T(b)) comparable to those observed in nature (0.5-3.5 degrees C) were induced by pre-chilling animals in 6 degrees C water. Pre-chilling did not reduce diving V O2 of any animal tested in 10 degrees C or 30 degrees C water, irrespective of the nature of the dive. Most behavioural indices of dive performance, including average and cumulative dive times, were unaffected by T(b) reduction in adults, but depressed in hypothermic juveniles (200-400 g). Hypothermia reduced diving heart rate only on short (<25s) dives (16% reduction, P=0.01), but did not affect the temporal onset of diving bradycardia. Post-immersion V O2 was higher for pre-chilled than for normothermic muskrats, but the difference became insignificant on longer (>90 s) dives. Our findings suggest that the mild hypothermia experienced by muskrats in nature has minimal effect on diving and post-immersion metabolic costs, and thus has little impact on the dive performance of this northern semi-aquatic mammal.     

Population regulation of Serengeti Wildebeeest: a test of the food hypothesis

Summary The food hypothesis proposes that density dependent mortality regulates populations through food shortage. For Serengeti wildebeest, we found an empirical relationship between dry season adult mortality rate, density and food supply. This relationship predicted that: (1) the population would stabilize between 1.0 and 1.5 million animals, (2) dry season mortality would be density dependent and sufficient to account for the levelling off of this large ungulate population. Recent observations have tested and confirmed these predictions.     

A test of the hydraulic limitation hypothesis in fast-growing Eucalyptus saligna

The hydraulic limitation hypothesis proposes that (1) reduced growth in taller trees is caused by decreased photosynthesis resulting from a decrease in hydraulic conductance promoted by a longer root‐to‐leaf flow path, and (2) this mechanism reduces stand productivity after canopy closure. This hypothesis was tested by comparing the physiology of 7 m (1 year) and 26 m (5 year) Eucalyptus saligna plantations where above‐ground productivity for the 26 m trees was approximately 69% of that for the 7 m trees, and water and nutrients were not limiting. The study compared whole tree physiology [water flux (Q_l), average crown conductance (G_T), crown hydraulic conductance per unit leaf area (K_L), carbon isotope discrimination (δ¹³C)] and leaf physiology under light saturation (leaf water potential at the canopy top (Ψ_LEAF), photosynthetic capacity (A_max), and photosynthesis (A) and stomatal conductance (g_s). K_L was 50% lower in the taller trees, but whole tree Q_l and G_T were the same for the 7 m and 26 m trees. Photosynthetic capacity was the same for leaves at the canopy top, but δ¹³C was ?1.8‰ lower for the 26 m trees. A and g_s were either lower in the taller trees or equal, depending on sampling date. The taller trees maintained 0.8 MPa lower Ψ_LEAF during the day and had 2.6‐times higher sapwood area per unit leaf area; these factors compensated for the effects of increased height and gravitational potential in the taller trees to maintain higher G_T. The hydraulic limitation hypothesis (as originally stated) failed to explain the sharp decline in net primary productivity after canopy closure in this study. The effects of increased height appear to be a universal hydraulic problem for trees, but compensation mitigated these effects and maintained Q_l and G_T in the present study. Compensation may induce other problems (such as lower Ψ_LEAF or higher respiratory costs) that could reduce carbon gain or shift carbon allocation, and future studies of hydraulic limitation should consider compensation and associated carbon costs. In this study, the combination of similar G_T and lower δ¹³C for the 26 m trees suggests that total crown photosynthesis was lower for the 26 m trees, perhaps a result of the lower Ψ_LEAF.     

Variation in scope for growth: a test of food limitation among intertidal mussels

Wellington Harbour supports large populations of the mussels Aulacomya maoriana, Mytilus galloprovincialis and Perna canaliculus that are almost entirely absent from nearby coastal locations in Cook Strait. We calculated scope for growth (SFG) using ambient Cook Strait water over a broad temporal scale and a broad range of seston conditions to determine if negative SFG explains this phenomenon. Although all three mussel species had positive mean SFG values, variation in SFG was high and negative values often occurred: A. maoriana 19.1 J g⁻¹ h⁻¹, 43% of mussels showed negative SFG; M. galloprovincialis 1.26, 52% negative SFG; P. canaliculus 45.6, 27% negative SFG. Negative SFG was most often due to negative absorption efficiency caused by metabolic faecal loss that is characteristic of mussels feeding in environments with low seston quality. From our ecophysiology data we constructed a model to estimate SFG based on physiological responses to the narrow range of seston conditions typical of Cook Strait (Model One), and a model to estimate SFG based on physiological responses of mussels to the broad range of seston conditions typical of Wellington Harbour and Cook Strait (Model Two). We used seston data collected over an 18-month period from sites in Wellington Harbour and Cook Strait to derive 159 estimates of species-specific mussel SFG from both models. Both models produced higher estimates of SFG for mussels in the Harbour compared with those at Cook Strait sites. This was consistent with elevated particulate concentrations in the Harbour than at Cook Strait sites, and in agreement with previous studies. For Cook Strait mussels, both models produced negative estimates of net energy balance for long periods of time (several months), whereas for Harbour mussels negative SFG estimates were generally short in duration. We conclude that our short-term laboratory-based determinations of SFG and our long-term bioenergetics modelling estimates do not conclusively support the hypothesis of food limitation for three coexisting taxa of mussels in the intertidal region of Cook Strait, New Zealand. Handling editor: P. Viaroli     

Density-dependent habitat selection in muskrats: a test of the ideal free distribution model

Summary Two predictions of the ideal free distribution model, a null hypothesis of habitat selection, were examined using free-ranging muskrats. We rejected the prediction that the proportion of the animals found in each of five habitats was independent of population size. Data on over-winter occupancy of muskrat dwellings tend also to refute the prediction of equal fitness reward among habitats. Habitat type and water-level had a profound effect on the suitability of a site for settlement. We concluded that the observed pattern of muskrat distribution followed more closely an ideal despotic distribution where some individuals benefited from a higher fitness because of resource monopolization. Current theories of density-dependent habitat selection, which assume an ideal free distribution, would not apply to muskrats and possibly to many other mammal species.     

Below-ground bud banks increase along a precipitation gradient of the North American Great Plains: a test of the meristem limitation hypothesis

In perennial grasslands, the below-ground population of meristems (bud bank) plays a fundamental role in plant population dynamics. Here, we tested the 'meristem limitation hypothesis' prediction - that bud banks increase along an increasing precipitation/productivity gradient in North American grasslands - and assessed the seasonal dynamics of bud banks. We sampled bud and stem populations quarterly at six sites across a 1100 km gradient in central North America. Bud banks increased with average annual precipitation, which explained 80% of the variability between the sites. In addition, seasonal changes in grass bud banks were surprisingly similar across a 2.5-fold range in precipitation and a 4-fold range of productivity: densities peaked in March, decreased in June and increased slightly in September. Increasing meristem limitation may constrain vegetation responses to inter-annual changes in resources. An important consequence of this is that biomes with large bud banks may be the most responsive to environmental change. If meristem limitation represents an important constraint on productivity responses to environmental variability, then bud banks must be considered in developing predictive models for grassland responses to environmental change.     

Migrants in tropical bird communities: the balanced breeding limitation hypothesis

Explanations for the ecological integration of migratory and non-migratory (resident) insectivorous birds in the tropics have been complicated by the paradox that arthropod abundances are low when bird abundances reach their annual peak. The breeding currency hypothesis and the nest predation hypothesis both account for this paradox by postulating that residents are held below the non-breeding season carrying capacity, which frees resources available for migratory insectivores. The breeding currency hypothesis suggests residents are limited by food suitable for nestlings, whereas the nest predation hypothesis emphasizes the primacy of high rates of nest predation. However, theoretical arguments suggest that food availability and predation risk interact strongly to limit breeding birds. We use graphical analyses to extend the breeding currency hypothesis to incorporate effects of nest predation. This yields a more synthetic and realistic model for the integration of migrant and resident insectivores in the tropics – the balanced breeding limitation hypothesis.     

Environmental rugosity, body size and access to food: a test of the size-grain hypothesis in tropical litter ants

In terrestrial walking organisms, long legs help to decrease the cost of running, allowing animals to step over environmental interstices rather than walking through them. However, long legs can complicate the infiltration of these interstices, which may contain food sources and refugia. Since the number of environmental interstices perceived by an organism (rugosity) increases as it body size decreases (size-grain hypothesis, SGH), natural selection should favor proportionally smaller legs with decreasing body size. Recent work demonstrated that ants fit this hypothesis. We experimentally tested the assumption of the SGH that small ants, which have proportionally smaller legs than larger ants, are more successful in exploring environmental interstices because they can easily penetrate them. We examined the ability of tropical litter ant species with different body sizes to access food baits in 'landscapes' (=plots) with different levels of rugosity and food exposure. In the first experiment, three levels of landscape rugosity were defined by manipulating the density of leaf litter placed on the ground plots: a) plain landscape: no litter fall, b) intermediate rugosity (∼0.5 kg of litter fall) and c) high rugosity (∼1 kg). In a second experiment, food baits were in plain landscapes, exposed or covered by leaf litter. The body lengths of ants that first accessed food baits ranged from 1.5 to 12 mm. Ants that first reached food baits in the most rugose landscapes were ∼40% smaller than ants that first found baits in plain landscapes. Smaller ants were also the first to access covered food. The application of a phylogenetic comparative method suggested the same patterns. We conclude that these results support the size grain hypothesis. Environmental rugosity might have operated as a selective force to shape the morphological characteristics of litter ant species.     

Response of the copper butterfly Lycaena tityrus to increased leaf nitrogen in natural food plants: evidence against the nitrogen limitation hypothesis

This study examined the effects of increased leaf N in natural food plants on oviposition, preimaginal survival, growth, and adult size of the butterfly Lycaena tityrus. Female butterflies did not discriminate between leaves of high and low N content. In accordance with previous studies, we found higher growth rates and concomitantly decreased development times at a high N level. However, because of high pupal (and larval) mortality (overall 73.0%) as well as a reduction in adult size (by ca. 8%) this was, overall, not beneficial to the butterflies. Thus, our results were not consistent with the broad interspecific trend that insect herbivore performance is positively correlated with leaf N. These findings undermine the general applicability of the N limitation hypothesis. As the detrimental effects were largely confined to the pupal and adult stages, results obtained from the larval phase only may not yield reliable results and must therefore be interpreted with caution. If negative effects of N enrichment are found more frequently in declining species inhabiting nutrient poor grassland, this will have major implications for the conservation of these species. Received: 30 November 1999 / Accepted: 14 February 2000     

Resource polyphenism increases species richness: a test of the hypothesis

A major goal of evolutionary biology is to identify the causes of diversification and to ascertain why some evolutionary lineages are especially diverse. Evolutionary biologists have long speculated that polyphenism—where a single genome produces alternative phenotypes in response to different environmental stimuli—facilitates speciation, especially when these alternative phenotypes differ in resource or habitat use, i.e. resource polyphenism. Here, we present a series of replicated sister-group comparisons showing that fishes and amphibian clades in which resource polyphenism has evolved are more species rich, and have broader geographical ranges, than closely related clades lacking resource polyphenism. Resource polyphenism may promote diversification by facilitating each of the different stages of the speciation process (isolation, divergence, reproductive isolation) and/or by reducing a lineage''s risk of extinction. Generally, resource polyphenism may play a key role in fostering diversity, and species in which resource polyphenism has evolved may be predisposed to diversify.     

Movement of modulator in maize: a test of an hypothesis

A model of Modulator movement (Greenblatt 1968) from one chromosomal site to another requires that all movements produce potential twin mutations within the affected cell lineage. From this model the prediction would be that (1) untwinned red and untwinned light-variegated sectors within the pericarp of medium-variegated maize must occur in equal frequency even though partners become lost, during ear morphogenesis, to the final pericarp tissue, and that (2) among the backcross progeny of a homozygous P(rr)Mp individual mated with P(wr)/P(wr) pollen, red offspring and light-variegated offspring would occur in equal frequency. Both expectations have been realized and herein reported.     

Condition and size of damselflies: a field study of food limitation

Summary Based on evidence from field manipulations, several authors have recently suggested that interference competition among larval odonates reduces individual growth rates and biomass by reducing foraging rates. This study was designed to test the effects of food shortage on condition (relative mass per unit head width) of larval Ischnura verticalis (Odonata: Coenagrionidae) under laboratory conditions and to use these results to estimate the degree of food shortage of larvae under naturally occurring field conditions. In the laboratory, there were marked differences in condition of larvae fed diets ranging from ad libitum feeding with worms to ad libitum feeding with Daphnia 1 day out of every 8. Condition of larvae collected from May through October from 17 different sites in southern Ontario indicated that, for most of the year, larvae had conditions similar to those fed ad libitum with Daphnia in the laboratory. There was no evidence that larval condition was related to population density. Condition of larvae in most sites during July was similar to that of larvae fed poor diets in the laboratory. It is unlikely that the low conditions were due to competition as there were no correlations with density across sites and population densities during July were at their lowest. Adult head widths showed a seasonal decline from mid June to the end of the flight season. There was no evidence that head widths were related to population density although there was some evidence that head widths of males were positively related to larval condition. My results do not support the hypothesis that competition is important in affecting foraging rates and subsequent development of larvae. Contrasts between my results and other studies may stem from difficulties with the interpretation of field experiments, that densities in my study may have been low due to fish predation, and/or that I. verticalis larvae are slow moving relative to other larvae and thus less likely to interact.     

Quantifying food limitation of arthropod predators in the field

A method for quantifying food limitation of arthropod predators in the field is presented and applied to species of ground beetles (Carabidae) and sheet-web spiders (Linyphiidae) from a cereal field. Food limitation is expressed quantitatively as accumulated hunger (=starvation) by transforming 24-h food consumption at 20°C of animals newly collected in the field into days of starvation at 20°C. This is done by means of a reference curve relating 24-h food intake at 20°C to starvation periods (days) at 20°C. Such a reference curve was obtained for the carabid beetle Agonum dorsale in the laboratory. For other species the reference curve was modified with species-specific data. The procedure makes it possible to compare the feeding conditions of different species populations that are part of the same community. Hunger levels in the field for females of A.␣dorsale were equivalent to c. 15 days of starvation in early spring, c. 5 days in June, increasing to c. 10 days in late summer. Two ground beetles occurring during summer also showed increasing hunger levels from June to July, probably the result of a dry summer. Two spider species experienced a hunger level in the range of 4–8 days of starvation. Received: 15 August 1997 / Accepted: 22 January 1998     

Evolution of clutch size in cavity-excavating birds: the nest site limitation hypothesis revisited

There are two major competing hypotheses for variation in clutch size among cavity-nesting species. The nest site limitation hypothesis postulates that nesting opportunities are more limited for weak excavators, which consequently invest more in each breeding attempt by laying larger clutches. Alternatively, clutch size may be determined by diet; the clutch sizes of strong excavators may be smaller because they are able to specialize on a more seasonally stable prey. We built a conceptual model that integrated hypotheses for interspecific variation in clutch size and tested it with comparative data on life-history traits of woodpeckers (Picidae) and nuthatches (Sittidae). In most analyses, diet explained more variation in clutch size among species than did propensity to excavate. Migratory status was positively associated with clutch size but was difficult to distinguish from diet since resident species consumed more bark beetles (a prey available in winter) and had smaller clutches than migratory species. The literature suggests that cavities are not limited in natural, old-growth forests. Although our data do not rule out nest site limitation, we conclude that annual stability of food resources has a larger impact on the evolution of clutch sizes in excavators than does limitation of nest sites.