A test of statistical techniques for detecting density dependence in sequential censuses of animal populations

Summary Principal and reduced major axes, and Bulmer's (1975) tests have been suggested as methods for detecting the presence of density dependence in a series of population censuses that are unsuitable for analysis by alternative means e.g. by k-factor analysis. These alternative methods are tested using census data, some of which are previously unpublished, from natural populations known from independent evidence to be subject to density dependent processes. All the methods fail to detect density dependence reliably, irrespective of sample size and the dynamics of the population. We conclude that none of the methods tested is sufficiently reliable to be useful as a test of density dependence in sequential censues of animal populations.     

Ontogenetic shifts within the selfish herd: predation risk and foraging trade-offs change with age in colonial web-building spiders

We tested for the existence of density dependence in annual adult rangeland grasshopper (Orthoptera: Acrididae) data from Montana, USA (1951–1991). Statistical density dependence was, in the sense of a stochastic equilibrium or return tendency, detected in all of the grasshopper mean density time-series from the three major physiographic regions of the state, Northern Glaciated Plains, Southern Unglaciated Plains, and Western Mountains. Parameters were estimated for a model that described the stochastic equilibrium characteristics of regional mean densities. The analyses showed that rangeland grasshopper regional densities fluctuate according to gamma distribution with a mean of 6.1–6.3 grasshoppers per m². Further, when regions exhibit outbreaks, the resulting infestation period (duration of outbreak) is short, spanning only a few generations.     

Early survival in roe deer: causes and consequences of cohort variation in two contrasted populations

Time- and sex-specific summer survival of roe deer fawns was estimated using capture-mark-recapture methods in two enclosed populations living in contrasting conditions. The population of Trois Fontaines (eastern France) was roughly constant in size throughout the study period, while in Chizé (western France), the population experienced frequent summer droughts and numbers decreased continuously during the study. Early survival of fawns was low and highly variable over the years at both Chizé and Trois Fontaines, and demonstrated marked variations between cohorts that need to be taken into account when modelling roe deer population dynamics. In Trois Fontaines, fawn survival was positively correlated with early body growth and total rainfall in May and June. In Chizé, fawn survival decreased with increasing density and tended to increase with increasing rainfall in May and June and adult female body mass. These factors explained more than 75% of the variability in early survival observed in both populations. Variation between cohorts had different consequences for the two populations. At Trois Fontaines, cohort variation was limited to a numerical effect on early survival. However at Chizé, cohort variation was long-lasting and affected the phenotypic quality of survivors at later ages, and thereby future survival and breeding abilities (both numerical and quality effects). Male and female fawns had similar survival over their first summer in both populations. This result contrasts with the lower survival of young males often observed in ungulates. Two ultimate causes can be proposed to account for the low and variable survival of roe deer fawns over the first summer: the high energy expenditures incurred by does during each breeding attempt and/or the low absolute body size of newborn roe deer fawns. Received: 28 April 1997 / Accepted: 14 July 1997     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

Spatial scale and the detection of density dependence in spruce budworm outbreaks in eastern North America

Using two tests for direct density dependence and standard techniques of time series analysis, we identified density dependence in defoliation time series of the spruce budworm across its outbreak range in eastern North America over the years 1945–1988. We carried out analyses for the entire region and for grid cells of defoliation maps at five spatial scales created by aggregating the smallest grid cells. The rate of detection of direct density dependence, as assessed by two previously published methods, decreased with increasing spatial scale. Using both methods, density dependence was detected more frequently at the periphery of the outbreak range, where defoliation rate was lower. This result suggested that density-dependent regulation may be stronger in those areas. The first order autoregressive process was the basic model for defoliation dynamics overall and the most common model across spatial scales. Second-order processes were encountered much less frequently, and those commonly identified as resulting from delayed density dependence generally occurred across spatial scales at a rate expected by chance alone. Our results were similar to those of other published studies, which have found the detection of density dependence to decrease at larger spatial scales. The results also reinforced the importance of considering spatial scale when diagnosing population processes using time series of abundance for single species. Received: 26 December 1999 / Accepted: 17 March 2000     

Population and prehistory II: space-limited human populations in constant environments

We present a population model to examine the forces that determined the quality and quantity of human life in early agricultural societies where cultivable area is limited. The model is driven by the non-linear and interdependent relationships between the age distribution of a population, its behavior and technology, and the nature of its environment. The common currency in the model is the production of food, on which age-specific rates of birth and death depend. There is a single non-trivial equilibrium population at which productivity balances caloric needs. One of the most powerful controls on equilibrium hunger level is fertility control. Gains against hunger are accompanied by decreases in population size. Increasing worker productivity does increase equilibrium population size but does not improve welfare at equilibrium. As a case study we apply the model to the population of a Polynesian valley before European contact.     

Individuals,populations and the balance of nature: the question of persistence in ecology

Explaining the persistence of populations is an important quest in ecology, and is a modern manifestation of the balance of nature metaphor. Increasingly, however, ecologists see populations (and ecological systems generally) as not being in equilibrium or balance. The portrayal of ecological systems as “non-equilibrium” is seen as a strong alternative to deterministic or equilibrium ecology, but this approach fails to provide much theoretical or practical guidance, and warrants formalisation at a more fundamental level. This is available in adaptation theory, which allows population persistence to be explained as an epiphenomenon stemming from the maintenance, survival, movement and reproduction of individual organisms. These processes take place within a physicochemical and biotic environment that persists through structured annual cycles, but which is also spatiotemporally dynamic and subject to stochastic variation. The focus is thus shifted from the overproduction of offspring and the consequent density dependent population pressure thought to follow, to the adaptations and ecological circumstances that support those relatively few individuals that do survive.     

Detection and analysis of genetic variation in Hordeum spontaneum populations from Israel using RAPD markers

Randomly amplified polymorphic DNA (RAPD) markers were used to analyse genetic diversity within and between Hordeum spontaneum populations sampled from Israel. Nei's index of genetic differentiation was used to partition diversity into within and between population components. Fifty-seven per cent of the variation detected was partitioned within 10 H. spontaneum populations. Using principal component and multiple regression analysis, part of the variation detected between populations was seen to be associated with certain ecogeographical factors. Fifty-eight per cent of the distribution of the phenotypic frequencies of three RAPD phenotypes detected using a single primer in 20 H. spontaneum populations could be accounted for by four ecogeographical variables, suggesting adaptive variation at certain RAPD loci.     

Genetic experiments with model populations: Fallacies in genetic analysis performed from samples of recombinants selected for two markers

Abstract: Samples selected for two markers (one from each parent) from the progeny of a genetic cross are altered both in the genotype and in the frequency distribution as compared to the original progeny population. The consequences of these alterations were analyzed in selected samples obtained from model progeny populations of hypothetical genetic crosses. In complete progeny populations, distribution of genotypes (pattern of genotypes), sequence of markers, and frequency distribution of individual genotypes exhibit genetically intelligent relationship to each other only under a sole condition: at the correct sequence of markers. In selected samples, the singularity of the interrelationship is relaxed. Consequences of the relaxation in the genetic analysis are multiple, the most striking of which is an insinuation of circularity for the linear gene map.     

Size variation in chewing lice <Emphasis Type="Italic">Docophorulus coarctatus</Emphasis>: how host size and louse population density vary together

Chewing lice of the species Docophorulus coarctatus were extracted from museum specimens of their host, the great grey shrike Lanius excubitor, by combing feathers from 36 freshly shot birds (shot between 1962 and 1974), and samples of ten individual lice (five female, five male) were randomly collected for measurements from each bird. Female lice were bigger than males for all studied measurements (P < 0.001 in all cases), although the size of both sexes obtained from individual hosts was positively correlated. The overall size of lice (derived from a principal components analysis) was positively correlated with the overall size of the avian host, and also with the population density of lice on the individual host. We suggest that variation in louse morphology is due to differences in selection pressure exerted by each host and by intraspecific competition due to conspecifics. This is, to the best of our knowledge, the first evidence that Harrison’s rule (parasites on larger host species are often bigger than those on smaller hosts) not only works in a multispecies comparison but also within a single host–single parasite system as well.     

Genetic analysis of fish genomes and populations: allozyme variation within and among Atlantic salmon from Downeast rivers of Maine

Analysis of genetic variation within and among samples of naturally produced Atlantic salmon ( n  = 372) from 7 Maine (U.S.A.) and one Canadian river were conducted based on 54 allozyme loci. Eight of the 54 loci proved polymorphic, and estimated heterozygosities ( H _S) based on all loci ranged from 0·012 to 0·026 (mean = 0·021, s . e . = 0·002). Only one of 56 tests revealed genotypic proportions that deviated significantly from Hardy–Weinberg expectations. Genetic distances ( D ) between samples ranged from 0·002 to 0·022. No obvious association existed between genetic and geographic distances. Cluster analysis of genetic distances revealed the Dennys River sample as the most differentiated when all samples were included in the analysis, though bootstrap support of the cluster analysis was generally weak. G ‐tests revealed significant differences in allele frequencies among samples at five of the polymorphic loci, and the G ‐value summed over all loci also indicated significant differences among samples. F _ST values indicated that 3·4% of the total genetic diversity was due to variability among samples, while 96·6% was due to variability within samples. These results indicate that the Atlantic salmon analyzed in this study had levels of genetic variability and differentiation among samples comparable to native populations from other areas collected across a similar geographic range.     

Density, climate and varying return points: an analysis of long-term population fluctuations in the threatened European tree frog

Experimental research has identified many putative agents of amphibian decline, yet the population-level consequences of these agents remain unknown, owing to lack of information on compensatory density dependence in natural populations. Here, we investigate the relative importance of intrinsic (density-dependent) and extrinsic (climatic) factors impacting the dynamics of a tree frog (Hyla arborea) population over 22 years. A combination of log-linear density dependence and rainfall (with a 2-year time lag corresponding to development time) explain 75% of the variance in the rate of increase. Such fluctuations around a variable return point might be responsible for the seemingly erratic demography and disequilibrium dynamics of many amphibian populations.     

Solar radiation stress in climbing snails: behavioural and intrinsic features define the Hsp70 level in natural populations of Xeropicta derbentina (Pulmonata)

Ectotherms from sunny and hot environments need to cope with solar radiation. Mediterranean land snails of the superfamily Helicoidea feature a behavioural strategy to escape from solar radiation-induced excessive soil heating by climbing up vertical objects. The height of climbing, and also other parameters like shell colouration pattern, shell orientation, shell size, body mass, actual internal and shell surface temperature, and the interactions between those factors may be expected to modulate proteotoxic effects in snails exposed to solar radiation and, thus, their stress response. Focussing on natural populations of Xeropicta derbentina, we conducted a ‘snapshot’ field study using the individual Hsp70 level as a proxy for proteotoxic stress. In addition to correlation analyses, an IT-model selection approach based on Akaike’s Information Criterion was applied to evaluate a set of models with respect to their explanatory power and to assess the relevance of each of the above-mentioned parameters for individual stress, by model averaging and parameter estimation. The analysis revealed particular importance of the individuals’ shell size, height above ground, the shell colouration pattern and the interaction height × orientation. Our study showed that a distinct set of behavioural traits and intrinsic characters define the Hsp70 level and that environmental factors and individual features strongly interact.     

Inferring local adaptation from QST-FST comparisons: neutral genetic and quantitative trait variation in European populations of great snipe

We applied a phenotypic QST (PST) vs. FST approach to study spatial variation in selection among great snipe (Gallinago media) populations in two regions of northern Europe. Morphological divergence between regions was high despite low differentiation in selectively neutral genetic markers, whereas populations within regions showed very little neutral divergence and trait differentiation. QST > FST was robust against altering assumptions about the additive genetic proportions of variance components. The homogenizing effect of gene flow (or a short time available for neutral divergence) has apparently been effectively counterbalanced by differential natural selection, although one trait showed some evidence of being under uniform stabilizing selection. Neutral markers can hence be misleading for identifying evolutionary significant units, and adopting the PST-FST approach might therefore be valuable when common garden experiments is not an option. We discuss the statistical difficulties of documenting uniform selection as opposed to divergent selection, and the need for estimating measurement error. Instead of only comparing overall QST and FST values, we advocate the use of partial matrix permutation tests to analyse pairwise QST differences among populations, while statistically controlling for neutral differentiation.     

Comment arising from a paper by Wittmer et al.: hypothesis testing for top-down and bottom-up effects in woodland caribou population dynamics

Conservation strategies for populations of woodland caribou Rangifer tarandus caribou frequently emphasize the importance of predator–prey relationships and the availability of lichen-rich late seral forests, yet the importance of summer diet and forage availability to woodland caribou survival is poorly understood. In a recent article, Wittmer et al. (Can J Zool 83:407–418, 2005b) concluded that woodland caribou in British Columbia were declining as a consequence of increased predation that was facilitated by habitat alteration. Their conclusion is consistent with the findings of other authors who have suggested that predation is the most important proximal factor limiting woodland caribou populations (Bergerud and Elliot in Can J Zool 64:1515–1529, 1986; Edmonds in Can J Zool 66:817–826, 1988; Rettie and Messier in Can J Zool 76:251–259, 1998; Hayes et al. in Wildl Monogr 152:1–35, 2003). Wittmer et al. (Can J Zool 83:407–418, 2005b) presented three alternative, contrasting hypotheses for caribou decline that differed in terms of predicted differences in instantaneous rates of increase, pregnancy rates, causes of mortality, and seasonal vulnerability to mortality (Table 1, p 258). These authors rejected the hypotheses that food or an interaction between food and predation was responsible for observed declines in caribou populations; however, the use of pregnancy rate, mortality season and cause of mortality to contrast the alternative hypotheses is problematic. We argue here that the data employed in their study were insufficient to properly evaluate a predation-sensitive foraging hypothesis for caribou decline. Empirical data on seasonal forage availability and quality and plane of nutrition of caribou would be required to test the competing hypotheses. We suggest that methodological limitations in studies of woodland caribou population dynamics prohibit proper evaluation of the mechanism of caribou population declines and fail to elucidate potential interactions between top-down and bottom-up effects on populations. An erratum to this article can be found at     

Genetic diversity in Tunisian rosy garlic populations (Allium roseum L.) as evidenced by chloroplastic DNA analysis: Sequence variation of non-coding region and intergenic spacers

Allium roseum (Alliaceae) is an important medicinal and aromatic plant in the flora of Tunisia. It is widespread in southern Tunisia, where it is commonly used as a therapeutic agent. However, its existence is seriously threatened because of over-collection.     

Sex differences in grey seal diet reflect seasonal variation in foraging behaviour and reproductive expenditure: evidence from quantitative fatty acid signature analysis

1. Intraspecific variation in diet can be an important component of a species niche breadth. We tested the hypothesis that sex differences in seasonal foraging behaviour and energy storage of sexually size dimorphic grey seals Halichoerus grypus (Fabrisius 1971) are reflected in differences in the diet and niche breadth. Diet composition was estimated for 496 adult (226 males, 270 females) and 91 juvenile (46 males/45 females; all 6 months old) grey seals sampled between 1993 and 2000 using quantitative fatty acid signature analysis. Niche breadth and overlap were estimated using the Shannon-Weaver diversity index (H') and the Morisita-Horn index (C(H)), respectively. 2. Sand lance Ammodytes dubius (Reinhardt 1837) and redfish Sebastes sp. (Cuvier 1829) accounted for a high proportion of the diet in both sexes and age groups. However, the diets of adult males were significantly more diverse across all seasons (H': males 0.36 +/- 0.007 vs. females 0.28 +/- 0.007) and less energy dense in spring (male 5.3 +/- 0.07 kJ g(-1) vs. females 5.6 +/- 0.09 kJ g(-1)) than those of adult females. 3. Season and sex explained most of the observed variation in adult diets, but there were significant sex-season interactions. These differences were most evident during the post-breeding (spring) foraging period when energy acquisition is important to female recovery of nutrient stores needed to support pregnancy. Females selected fewer and higher quality prey species in spring than males. 4. There were no sex differences in the diets of juvenile grey seals. Although many of the species overlapped with those eaten by adults, juvenile niche breadth (H': 0.41 +/- 0.014, n = 91) was significantly broader than that of adults (H': 0.30 +/- 0.011, n = 115). Juvenile diets were also of lower energy density (5.3 +/- 0.04 kJ g(-1)) than those of adults (5.6 +/- 0.09 kJ g(-1)), suggesting less selectivity in these young and relatively na?ve predators. 5. Sex-specific seasonal changes in diet correspond to seasonal changes in diving behaviour and rate of body energy accumulation of adult males and females. Sex-specific reproductive requirements appear to be a primary factor generating the intraspecific variation in the seasonal foraging ecology of this large marine carnivore. However, sex differences in the breadth and energy content of diets also suggest the influence of body-size dimorphism as a factor shaping the diet of this species.     

Population genetic analysis of cat populations from Mexico, Colombia, Bolivia, and the Dominican Republic: Identification of different gene pools in Latin America

In this paper we identify new genetic profiles of eight Latin American cat populations. In addition, we combine data from the present study and previously published data on 70 other American and European populations to discuss (1) the points of introduction of mutant alleles for cat coat phenotypes from Europe into Latin America, (2) the heterozygosity levels at these loci in the current Latin American cat populations, (3) the level of genetic heterogeneity among Latin American cat populations, and how this compares with levels found in North American and European cat populations, and (4) how many different cat gene pools are currently present in Latin America. We also include in our purview historical records of human migrations from Europe to and within the Americas. Our analyses clearly support the view that the current genetic profiles and structuring of cat populations in Latin America can be largely explained by the historical migration patterns of humans. This work is dedicated to the memory of Dr Roy Robinson. May he rest in peace.