Density-Dependent Growth in Invasive Lionfish (Pterois volitans)

Direct demographic density dependence is necessary for population regulation and is a central concept in ecology, yet has not been studied in many invasive species, including any invasive marine fish. The red lionfish (Pterois volitans) is an invasive predatory marine fish that is undergoing exponential population growth throughout the tropical western Atlantic. Invasive lionfish threaten coral-reef ecosystems, but there is currently no evidence of any natural population control. Therefore, a manipulative field experiment was conducted to test for density dependence in lionfish. Juvenile lionfish densities were adjusted on small reefs and several demographic rates (growth, recruitment, immigration, and loss) were measured throughout an 8-week period. Invasive lionfish exhibited direct density dependence in individual growth rates, as lionfish grew slower at higher densities throughout the study. Individual growth in length declined linearly with increasing lionfish density, while growth in mass declined exponentially with increasing density. There was no evidence, however, for density dependence in recruitment, immigration, or loss (mortality plus emigration) of invasive lionfish. The observed density-dependent growth rates may have implications for which native species are susceptible to lionfish predation, as the size and type of prey that lionfish consume is directly related to their body size. The absence of density-dependent loss, however, contrasts with many native coral-reef fish species and suggests that for the foreseeable future manual removals may be the only effective local control of this invasion.     

Pattern, process and the evolution of meaning: species and units of selection

Many of the fundamental concepts of biology lack consensual, precise definitions. Partly, this is due to a contrast between our discrete language and the continuous character of nature. Some debates over these concepts are confounded by the use of the same terms with different specific meanings, indicating a possible need for an expanded scientific lexicon. Words have their own histories, and frequently scientific terms with a vernacular origin retain associated vestigial meanings. Even terms newly coined within science have histories and changing meanings, which can lead to confusion among debaters. Debates over concepts are further confounded when the same terms are used in different fields of biology, with distinct (even conflicting) objectives, and by biologists with different approaches and perspectives. I illustrate these issues by considering the debate over the concept of species and the unit of selection.     

Can intense predation by bears exert a depensatory effect on recruitment in a Pacific salmon population?

It has long been recognized that, as populations increase in density, ecological processes affecting growth and survival reduce per capita recruitment in the next generation. In contrast to the evidence for such “compensatory” density dependence, the alternative “depensatory” process (reduced per capita recruitment at low density) has proven more difficult to demonstrate in the field. To test for such depensation, we measured the spawner–recruit relationship over five decades for a sockeye salmon (Oncorhynchus nerka) population in Alaska breeding in high-quality, unaltered habitat. Twenty-five years of detailed estimates of predation by brown bears, Ursus arctos, revealed strong density dependence in predation rate; the bears killed ca. 80 % of the salmon in years of low salmon spawning abundance. Nevertheless, the reconstructed spawner–recruit relationship, adjusted to include salmon intercepted in the commercial fishery, provided no evidence of demographic depensation. That is, in years when few salmon returned and the great majority were killed by bears, the few that spawned were successful enough that the population remained highly productive, even when those killed by bears were included as potential spawners. We conclude that the high quality of breeding habitat at this site and the productive nature of semelparous Pacific salmon allowed this population to avoid the hypothesized depressed recruitment from depensatory processes expected at low density. The observed lack of demographic depensation is encouraging from a conservation standpoint because it implies that depleted populations may have the potential to rebound successfully given suitable spawning and rearing habitat, even in the presence of strong predation pressure.     

Effects of forest disturbance on the fitness of an endemic rodent in a biodiversity hotspot

Praomys delectorum occurs abundantly in both disturbed and intact forests in the Ukaguru Mountains within the Eastern Arc Mountains (EAM), Morogoro, Tanzania. While previous studies have reported that anthropogenic disturbances such as grazing, wood cutting, and harvesting have a positive effect on the population density of P. delectorum, the impact of habitat disturbance on its demographic traits is still unknown. We performed a capture–mark–recapture study in both disturbed and intact forests from June 2018 to February 2020 in order to investigate the effects of habitat disturbance on abundance and two demographic traits: survival and maturation of P. delectorum in the Ukaguru Mountains. We found no variation in abundance or maturation between intact and disturbed forests, but habitat type did affect survival. However, this effect was sex‐dependent since female survival was higher in disturbed forests, while male survival remained similar across the two forest types potentially due to differences in predation pressure or food availability between the two habitats. Continuous demographic monitoring of P. delectorum in EAM is necessary given that the increasing human population surrounding the landscape is leading to higher deforestation rates and expansion of the pine plantation in the forest reserve.     

Phenotypic Plasticity in Response to the Social Environment: Effects of Density and Sex Ratio on Mating Behaviour Following Ecotype Divergence

The ability to express phenotypically plastic responses to environmental cues might be adaptive in changing environments. We studied phenotypic plasticity in mating behaviour as a response to population density and adult sex ratio in a freshwater isopod (Asellus aquaticus). A. aquaticus has recently diverged into two distinct ecotypes, inhabiting different lake habitats (reed Phragmites australis and stonewort Chara tomentosa, respectively). In field surveys, we found that these habitats differ markedly in isopod population densities and adult sex ratios. These spatially and temporally demographic differences are likely to affect mating behaviour. We performed behavioural experiments using animals from both the ancestral ecotype (“reed” isopods) and from the novel ecotype (“stonewort” isopods) population. We found that neither ecotype adjusted their behaviour in response to population density. However, the reed ecotype had a higher intrinsic mating propensity across densities. In contrast to the effects of density, we found ecotype differences in plasticity in response to sex ratio. The stonewort ecotype show pronounced phenotypic plasticity in mating propensity to adult sex ratio, whereas the reed ecotype showed a more canalised behaviour with respect to this demographic factor. We suggest that the lower overall mating propensity and the phenotypic plasticity in response to sex ratio have evolved in the novel stonewort ecotype following invasion of the novel habitat. Plasticity in mating behaviour may in turn have effects on the direction and intensity of sexual selection in the stonewort habitat, which may fuel further ecotype divergence.     

Component-dependent allomorphy and paradigm accessibility: evidence from Hebrew

This study examines allomorphy in vowel selection in Hebrew. We address the formation of adjectives and passive verbs in a specific morphological pattern with stem-medial glottals. While the normative vowel that precedes the glottal is o, there are cases where it is colloquially u. We provide empirical evidence for this variation (or lack thereof), arguing that vowel selection results from the distinction between adjectives and passive verbs and, more generally, from the component of the grammar where they are derived, the lexicon and the syntax. Studies of Hebrew have shown that a has a more morphological-lexical status, as it is typical of word formation processes, while e has a phonological status as it is the default vowel in epenthesis. A previous study has also shown that a is more typical of the lexicon and e is more typical of the syntax. We propose a similar distinction between o and u; u tends to be selected in passive formation in the syntax, while o is selected for words stored in the lexicon. In addition, we claim that the occurrence of each allomorph is triggered by uniformity to other forms in different paradigms that are taken into account in word formation. The study provides further evidence to a new type of allomorphy, whose evidence for conditioning is not purely phonological or morpho-syntactic, but rather depends on the locus of word formation. It highlights the strong correlation between form, meaning and the relevant grammatical component in word formation.     

Poor horse traders: large mammals trade survival for reproduction during the process of feralization

We investigated density dependence on the demographic parameters of a population of Camargue horses (Equus caballus), individually monitored and unmanaged for eight years. We also analysed the contributions of individual demographic parameters to changes in the population growth rates. The decrease in resources caused a loss of body condition. Adult male survival was not affected, but the survival of foals and adult females decreased with increasing density. Prime-aged females maintained high reproductive performance at high density, and their survival decreased. The higher survival of adult males compared with females at high density presumably results from higher investment in reproduction by mares. The high fecundity in prime-aged females, even when at high density, may result from artificial selection for high reproductive performance, which is known to have occurred in all the major domestic ungulates. Other studies suggest that feral ungulates including cattle and sheep, as these horses, respond differently from wild ungulates to increases in density, by trading adult survival for reproduction. As a consequence, populations of feral animals should oscillate more strongly than their wild counterparts, since they should be both more invasive (as they breed faster), and more sensitive to harsh environmental conditions (as the population growth rate of long-lived species is consistently more sensitive to a given proportional change in adult survival than to the same change in any other vital rate). If this principle proves to be general, it has important implications for management of populations of feral ungulates.     

A technique for analysis of density dependence in population models

The concept of density-dependent population growth is fundamental to our understanding of how populations persist. While it is generally agreed that negative density dependence must occur at high densities, the direction of density dependence may be negative (pure negative density dependence) or positive (demographic Allee effect) at low densities. In this article, we present a technique to link the direction of density dependence to generic ecological factors. This technique involves exploiting the presence of a particular bifurcation, known as a saddle-node-transcritical interaction. We first provide a method to detect this bifurcation in a given model and then demonstrate its ecological relevance using several existing mechanistic models. With a mathematical framework in place, we are able to identify scenarios in which neither a weak Allee effect nor pure negative density dependence are possible. More generally, we find conditions on parameter values that are necessary for transitions between pure negative density dependence and demographic Allee effects to occur.     

Density dependence in blowfly populations: experimental evaluation of non‐parametric time‐series modelling

Two approaches for describing density dependence in demographic rates of stage‐structured populations are compared in this study. Time‐series data from laboratory blowfly populations (Lucilia sericata) have been analysed in a separate study, with a statistical modelling approach that incorporated density dependences as unspecified (non‐parametric) functions. In this study, we assessed density‐dependent structures by manipulating densities of larvae and adults in cohorts of blowflies and measuring the demographic rates. We here compare the density‐dependent structures revealed by the cohort experiments with those estimated by the non‐parametric model. This model estimates the demographic rates to have the following density‐dependent structures: (i) larval survival was non‐linearly density‐dependent (a ‘humped’ function), (ii) adult survival was density‐independent, and (iii) reproductive rate decreased with adult density. In the cohort experiments reported here, (i) juvenile survival exhibited a positive density dependence in low densities (facilitation), which became negative at higher densities (competition). Pupal and adult size decreased with initial larval density. (ii) Adult survival was reduced by high initial larval density, but it was independent of adult density. (iii) Reproductive rate was reduced by high initial larval density, and by high adult density in populations of large individuals (from low larval density). Hence, the results from these experiments support the non‐parametric model estimates regarding density‐dependent structures of demographic rates in the blowfly populations. The mean demographic rates, however, were apparently underestimated by the model. We conclude that non‐parametric modelling is a useful first approach for exploratory analysis of ecological time‐series data.     

The Evolution of Plains Indian Kin Terminologies:

Abstract

The striking similarity in basic terminological pattern among Plains tribes, and the absence of this pattern elsewhere in North America, is not accounted for by existing general theories of the evolution of kin terminologies. The development of this pattern is explained here in terms of a theory which abandons the assumption that terminological patterns are reflections of social structure. It is argued that the basic process in this development was the tactical or metaphoric extension of sibling terms to cross-cousins as a response to the increased importance of solidarity under the conditions of Plains life, and the subsequent incorporation of this extension into the meanings of the sibling terms. This hypothesis not only accounts for the distribution of terminological patterns in the Plains area, but illuminates the general relationship of kin terminology to social structure.     

Coming to Terms with the Concept of Moving Species Threatened by Climate Change – A Systematic Review of the Terminology and Definitions

Intentional moving of species threatened by climate change is actively being discussed as a conservation approach. The debate, empirical studies, and policy development, however, are impeded by an inconsistent articulation of the idea. The discrepancy is demonstrated by the varying use of terms, such as assisted migration, assisted colonisation, or managed relocation, and their multiple definitions. Since this conservation approach is novel, and may for instance lead to legislative changes, it is important to aim for terminological consistency. The objective of this study is to analyse the suitability of terms and definitions used when discussing the moving of organisms as a response to climate change. An extensive literature search and review of the material (868 scientific publications) was conducted for finding hitherto used terms (N = 40) and definitions (N = 75), and these were analysed for their suitability. Based on the findings, it is argued that an appropriate term for a conservation approach relating to aiding the movement of organisms harmed by climate change is assisted migration defined as follows: Assisted migration means safeguarding biological diversity through the translocation of representatives of a species or population harmed by climate change to an area outside the indigenous range of that unit where it would be predicted to move as climate changes, were it not for anthropogenic dispersal barriers or lack of time. The differences between assisted migration and other conservation translocations are also discussed. A wide adoption of the clear and distinctive term and definition provided would allow more focused research on the topic and enable consistent implementation as practitioners could have the same understanding of the concept.     

An open spatial capture–recapture model for estimating density,movement, and population dynamics from line‐transect surveys

The purpose of many wildlife population studies is to estimate density, movement, or demographic parameters. Linking these parameters to covariates, such as habitat features, provides additional ecological insight and can be used to make predictions for management purposes. Line‐transect surveys, combined with distance sampling methods, are often used to estimate density at discrete points in time, whereas capture–recapture methods are used to estimate movement and other demographic parameters. Recently, open population spatial capture–recapture models have been developed, which simultaneously estimate density and demographic parameters, but have been made available only for data collected from a fixed array of detectors and have not incorporated the effects of habitat covariates. We developed a spatial capture–recapture model that can be applied to line‐transect survey data by modeling detection probability in a manner analogous to distance sampling. We extend this model to a) estimate demographic parameters using an open population framework and b) model variation in density and space use as a function of habitat covariates. The model is illustrated using simulated data and aerial line‐transect survey data for North Atlantic right whales in the southeastern United States, which also demonstrates the ability to integrate data from multiple survey platforms and accommodate differences between strata or demographic groups. When individuals detected from line‐transect surveys can be uniquely identified, our model can be used to simultaneously make inference on factors that influence spatial and temporal variation in density, movement, and population dynamics.     

Effective population size for culturally evolving traits

Population size has long been considered an important driver of cultural diversity and complexity. Results from population genetics, however, demonstrate that in populations with complex demographic structure or mode of inheritance, it is not the census population size, N, but the effective size of a population, N_e, that determines important evolutionary parameters. Here, we examine the concept of effective population size for traits that evolve culturally, through processes of innovation and social learning. We use mathematical and computational modeling approaches to investigate how cultural N_e and levels of diversity depend on (1) the way traits are learned, (2) population connectedness, and (3) social network structure. We show that one-to-many and frequency-dependent transmission can temporally or permanently lower effective population size compared to census numbers. We caution that migration and cultural exchange can have counter-intuitive effects on N_e. Network density in random networks leaves N_e unchanged, scale-free networks tend to decrease and small-world networks tend to increase N_e compared to census numbers. For one-to-many transmission and different network structures, larger effective sizes are closely associated with higher cultural diversity. For connectedness, however, even small amounts of migration and cultural exchange result in high diversity independently of N_e. Extending previous work, our results highlight the importance of carefully defining effective population size for cultural systems and show that inferring N_e requires detailed knowledge about underlying cultural and demographic processes.     

Climate, social factors and research disturbance influence population dynamics in a declining sociable weaver metapopulation

Population trends are determined by gains through reproduction and immigration, and losses through mortality and emigration. These demographic quantities and resulting population dynamics are affected by different external and internal drivers. We examined how these demographic quantities were affected by weather, research-induced disturbance, local density, colony site and year in a metapopulation of 17 sociable weaver (Philetairus socius) colonies over 17 years of study (4 years for reproduction). Most colonies declined, but at different rates. The four demographic quantities were related to different drivers. Survival strongly varied among years and colonies and was positively related to rainfall and negatively related to extreme temperature (together explaining 30 % of variation) and disturbance (measured as number of captures conducted at a colony; 7 %). There was a trend for a positive relationship between reproduction and rainfall (50 %). Movement was mainly related to local density: individuals were more likely to emigrate from small to large colonies and from colonies that were either well below or above their long-term mean. They were more likely to immigrate into colonies that were nearby, and below their mean size. We then quantified the effects of these relationships on metapopulation dynamics using a multi-site matrix projection model. Rainfall was potentially a strong driver of metapopulation dynamics. In addition, field-work disturbance might have contributed to the decline of this metapopulation but could not explain its full magnitude. Hence, through a combination of analytical methods we were able to obtain information on the main drivers affecting dynamics in a declining metapopulation.     

The Structure of the Individual Mental Lexicon from the Standpoint of L. S. Vygotsky's Ideas

Research on the structure of an individual's mental lexicon has shown that, among the set of concepts belonging to a single semantic field, there is typically one particular concept that stands out. The processes of storage, identification, and retrieval of this concept are faster and more successful than they are for the others. For example, in the vertical hierarchy of the concept fruit—apple—antonovka [McIntosh to give a familiar U. S. example—Trans.], apple plays this type of lead role and also occupies the most prominent position in the horizontal series apple—pear—plum—cherries—orange. Concepts with such characteristics are referred to as basic, source, or primary concepts. What is the reason for their special status?     

Cascading effects of artificial light at night: resource-mediated control of herbivores in a grassland ecosystem

Artificial light at night has a wide range of biological effects on both plants and animals. Here, we review mechanisms by which artificial light at night may restructure ecological communities by modifying the interactions between species. Such mechanisms may be top-down (predator, parasite or grazer controlled), bottom-up (resource-controlled) or involve non-trophic processes, such as pollination, seed dispersal or competition. We present results from an experiment investigating both top-down and bottom-up effects of artificial light at night on the population density of pea aphids Acyrthosiphon pisum in a diverse artificial grassland community in the presence and absence of predators and under low-level light of different spectral composition. We found no evidence for top-down control of A. pisum in this system, but did find evidence for bottom-up effects mediated through the impact of light on flower head density in a leguminous food plant. These results suggest that physiological effects of light on a plant species within a diverse plant community can have detectable demographic effects on a specialist herbivore.     

Local environment and density-dependent feedbacks determine population growth in a forest herb

Linking spatial variation in environmental factors to variation in demographic rates is essential for a mechanistic understanding of the dynamics of populations. However, we still know relatively little about such links, partly because feedbacks via intraspecific density make them difficult to observe in natural populations. We conducted a detailed field study and investigated simultaneous effects of environmental factors and the intraspecific density of individuals on the demography of the herb Lathyrus vernus. In regression models of vital rates we identified effects associated with spring shade on survival and growth, while density was negatively correlated with these vital rates. Density was also negatively correlated with average individual size in the study plots, which is consistent with self-thinning. In addition, average plant sizes were larger than predicted by density in plots that were less shaded by the tree canopy, indicating an environmentally determined carrying capacity. A size-structured integral projection model based on the vital rate regressions revealed that the identified effects of shade and density were strong enough to produce differences in stable population sizes similar to those observed in the field. The results illustrate how the local environment can determine dynamics of populations and that intraspecific density may have to be more carefully considered in studies of plant demography and population viability analyses of threatened species. We conclude that demographic approaches incorporating information about both density and key environmental factors are powerful tools for understanding the processes that interact to determine population dynamics and abundances.     

Density dependence at some time and place?

There appears to be widespread acceptance that for a population to persist, some demographic parameter must be density dependent at some place or time. In this paper, we question the veracity and heuristic value of treating this statement as a general principle of ecology. We also point out that some processes that have recently been defined as density dependent are, in fact, not. Taken in its original sense, density dependence implies a change in demographic rates based on biological (generally negative) feedback. Situations exist, however, in which demographic rates change in relation to density without negative biological feedback. For example, per capita recruitment in marine populations will decrease as local population size increases even as absolute numbers of arriving larvae do not change. The failure to separate these density-related processes from true density-dependent processes affects our understanding of population regulation and of the way in which the natural world functions. Furthermore, focusing solely on density-dependent processes and their role in population regulation neglects to address numerous density-independent processes like disturbance and climatic variation that may have important impacts in determining population size. Received: 20 January 1999 / Accepted: 12 January 2000     

On microrefugia and cryptic refugia

The existence of microrefugia and/or cryptic refugia has been proposed to explain the colonization patterns of temperate continents after the Last Glacial Maximum, as well as to gain an understanding of the present genetic structure of species and their populations. The concepts ‘microrefugia’ and ‘cryptic refugia’ largely overlap but do not fully coincide, which may lead to terminological duplication and eventual semantic misconceptions. Here, after a terminological and conceptual analysis, it is proposed that most situations currently characterized as cryptic refugia can be accommodated within the definition of microrefugia, with one single exception, for which some alternative terms are suggested. Moreover, the concept and definition of cryptic refugia are considered to be more imprecise, perishable and subjective than those of microrefugia, which is the preferred term in the current context.     

On the evolution of demographic strategies in populations with equilibrium and cyclic densities

Optimal demographic strategies (life histories of l_x-m_x schedules) for populations with stable and cylicly varying densities are predicted on the basis of a density dependent population dynamics model; the cyclic density pattern—often seen in lemmings and voles —is assumed to have relatively slow density increase followed by an abrupt density crash. Analysis shows that low reproductive rates for all ages and a Type I or II survival schedule will be favored by natural selection in stable populations. High reproductive rates for all ages and a Type III survival schedule will be favored by natural selection in cyclic populations. These strategies may be regarded as the endpoints of a continuum. Features predicted to be optimal for opportunistic species are expected to be more pronounced the more hostile the environment is. Since all quantities used in formulating these predictions are operationally defined, the predicted patterns are testable. Predictions derived from the model are compared, in a general way, with available data. This comparison suggests the validity of the theory.