Demography when history matters: construction and analysis of second-order matrix population models

History matters when individual prior conditions contain important information about the fate of individuals. We present a general framework for demographic models which incorporates the effects of history on population dynamics. The framework incorporates prior condition into the i-state variable and includes an algorithm for constructing the population projection matrix from information on current state dynamics as a function of prior condition. Three biologically motivated classes of prior condition are included: prior stages, linear functions of current and prior stages, and equivalence classes of prior stages. Taking advantage of the matrix formulation of the model, we show how to calculate sensitivity and elasticity of any demographic outcome. Prior condition effects are a source of inter-individual variation in vital rates, i.e., individual heterogeneity. As an example, we construct and analyze a second-order model of Lathyrus vernus, a long-lived herb. We present population growth rate, the stable population distribution, the reproductive value vector, and the elasticity of λ to changes in the second-order transition rates. We quantify the contribution of prior conditions to the total heterogeneity in the stable population of Lathyrus using the entropy of the stable distribution.     

Gutsy genetics: identification of digested piscine prey items in the stomach contents of sympatric native and introduced warmwater catfishes via DNA barcoding

A major focus of ecology is understanding trophic relationships and energy flows in natural systems, associated food web dynamics and changes in food webs due to introduced species. Predator-prey interactions are often assessed by examining stomach contents. However, partially digested remains may be difficult to accurately identify by traditional visual analysis. Here we evaluate the effectiveness of DNA barcoding to identify digested piscine prey remains in invasive Blue Catfish Ictalurus furcatus, non-native, but established Channel Catfish Ictalurus punctatus and native White Catfish Ameiurus catus from Chesapeake Bay, USA. Stomach contents were examined and piscine prey items were scored as lightly digested, moderately digested or severely digested. A 652 base pair region of the cytochrome c oxidase subunit I (COI-5P) mitochondrial DNA gene was sequenced for each prey item. Edited barcode sequences were compared to locally-caught and validated reference sequences in BOLD (Barcode of Life Database). A large majority of prey items were sufficiently digested to limit morphological identification (9.4 % to species and an additional 12.1 % to family). However, overall barcoding success was high (90.3 %) with little difference among the digestion classifications. Combining morphological and genetic identifications, we classified 91.6 % of fish prey items to species. Twenty-three fish species were identified, including species undergoing active restoration efforts (e.g., Alosa spp.) and commercially important species, e.g., Striped Bass Morone saxatilis, White Perch Morone americana, American Eel Anguilla rostrata and Menhaden Brevoortia tyrannus. We found DNA barcoding highly successful at identifying all but the most heavily degraded prey items and to be an efficient and effective method for obtaining diet information to strengthen the resolution of trophic analyses including diet comparisons among sympatric native and non-native predators.     

The effects of polymorphisms in <Emphasis Type="Italic">growth hormone</Emphasis> and <Emphasis Type="Italic">growth hormone receptor</Emphasis> genes on production and reproduction traits in Aberdeen-Angus cattle (<Emphasis Type="Italic">Bos taurus</Emphasis> L., 1758)

The study was aimed to analyze the relation between individual genotypes and allelic variants of SNPs g.2141C>G of growth hormone gene, g.914T>A and g.257A>G of growth hormone receptor gene with growth and reproduction traits and to evaluate the populationgenetic structure in Aberdeen-Angus cattle (Bos taurus L., 1758) sample of Eastern Ukraine according SNPs studied. Allele C of SNP g.2141C>G has a positive correlation with birth weight, body stature, bigger rump, udder and total exterior evaluation score, shorter calving interval and better calve birth weight and negative correlation with calve average daily gain. Allele T of SNP g.914T>A has positive correlation with the muscle and udder size; live weight in each age, average daily gain, weight and average daily gain of calves born conform to the principle AA>TT≥TA. SNP g.257A>G showed a positive correlation for G allele with muscle size. The population is in equilibrium for SNPs g.2141C>G and g.257A>G, and in disequilibrium for SNP g.914T>A. The analysis showed no linkage disequilibrium between SNPs g.914T>A and g.257A>G. Inbreeding coefficient F_ST in Aberdeen-Angus group studied was 16.1%.     

Fullerene transformations as analogues of radiolarian skeleton microevolution

The analogy in the structures of fullerenes (i.e., polyhedral molecules with only the pentagonal and hexagonal facets) and radiolarian skeletons (Heliosphaera inermis, H. tenuissima, H. actinota, H. echinoides, H. elegans; Circogonia dodecahedra; Haliomma capillaceum, Ethmosphaera siphonophora Hkl., etc.) is briefly considered. Modern chemical methods allow fullerenes to be changed to desirable forms, e.g., the most symmetrical and, therefore, stable ones. The computer algorithms (e.g., SW-transformations) divide the variety of fullerenes-isomers into the classes of equivalency. The hypothesis is stated that the transformations of fullerenes simulate the radiolarian skeleton microevolution, while the classes of equivalency are of taxonomic significance.     

Genetic divergence within the monotypic tree genus <Emphasis Type="Italic">Platycarya</Emphasis> (Juglandaceae) and its implications for species’ past dynamics in subtropical China

Subtropical East Asia harbours a large plant diversity that is often attributed to allopatric speciation in this topographically complex region characterized by a relative climate stability. Here, we use observations of Platycarya, a widespread subtropical Asian tree genus, to explore the consequences of past climate stability on species’ evolutionary history in subtropical China. This genus has a controversial taxonomy: while it is now prevailingly treated as monotypic, two species have been originally described, Platycarya strobilacea and P. longipes. Previous information from species distribution models, fossil pollen data and genetic data based on chloroplast DNA (cpDNA) were integrated with newly obtained genetic data from the two putative species. We used both cpDNA (psbA-trnH and trnL-F intergenic spacers, including a partial trnL gene sequence) and nuclear markers. The latter included sequences of the internal transcribed spacer region (ITS1–5.8S–ITS2) of the nuclear ribosomal DNA and random genomic single nucleotide polymorphisms. Using these nuclear genetic markers, we found interspecific genetic divergence fitting with the ‘two species’ scenario and geographically structured intraspecific variation. Using cpDNA markers, we also found geographically structured intraspecific variation. Despite deep inter- and intraspecific genetic divergence, we detected genetic admixture in southwest China. Overall, our findings of genetic divergence within Platycarya support the hypothesis of allopatric speciation. However, episodes of population interconnection were identified, at least in southwest China, suggesting that the genus has had a dynamic population history.     

Phylogenetic study of clavicipitaceous fungi using acetaldehyde dehydrogenase gene sequences

Aciculosporium and Heteroepichloë (Clavicipitaceae) are characteristic bambusicolous fungi in east Asia. In this study, we examined their intergeneric relationships based on the ALDH1-1 gene, which encodes a member of the aldehyde dehydrogenase family. In the clavicipitaceous fungi examined in this study, the nucleotide sequence of the third exon of ALDH1-1 (Exon-3) is 889 bp in length and has no insertion/deletion. A phylogenetic tree based on Exon-3 indicated that the clavicipitaceous fungi could be divided into two large groups: Cordyceps, Nomuraea, and Ustilaginoidea species formed a paraphyletic group, and the other grass biotrophic species formed a monophyletic group. This monophyletic group was further divided into three groups with high bootstrap support: i.e., species with Neotyphodium anamorphs (e.g., Epichloë), species with Ephelis anamorphs (e.g., Heteroepichloë), and Aciculosporium-Claviceps species. We discuss the relationships among Aciculosporium, Heteroepichloë, and other clavicipitaceous fungi.     

Glutathione improves survival of cryopreserved embryogenic calli of <Emphasis Type="Italic">Agapanthus praecox</Emphasis> subsp. <Emphasis Type="Italic">orientalis</Emphasis>

The effect of supplementation of reduced glutathione (GSH) to cryoprotectant solution on the generation of reactive oxygen species (ROS) (e.g., H₂O₂, OH^·, and O ₂ ^·? ) and antioxidants (e.g., SOD, POD, CAT, AsA, and GSH), as well as membrane lipid peroxidation (i.e., MDA content) mitigation in cryopreserving of embryogenic calli (EC) of Agapanthus praecox subsp. orientalis was investigated. The vitrification-based cryopreservation method was used in this study. The addition of GSH at a final concentration of 0.08 mM to the cryoprotectant solution has significantly improved cryotolerance of A. praecox EC. The EC post-thaw survival rate increased by 68.34 % using the cryoprotectant solution containing 0.08 mM GSH as compared to the control (GSH-free). EC treated with GSH displayed the reduction in  OH^· generation activity and the contents of H₂O₂ and MDA, as well as enhancement in the inhibition of O ₂ ^·? generation and the antioxidant activity. Treatment with exogenous GSH also increased endogenous AsA and GSH contents after dehydration step. Expression of stress-responsive genes, e.g., peroxidase (POD), peroxiredoxin, ascorbate peroxidase (APX), monodehydroascorbate reductase (MDHAR), and glutathione peroxidase (GPX), was also increased during cryopreservation processes. The expression of DAD1 (Defender against apoptotic cell death) was elevated, while cell death-related protease SBT was suppressed. These results demonstrated that the addition of GSH to cryoprotectant solution affects the ROS level and could effectively improve survival of A. praecox EC through enhancing antioxidant enzyme activities and decreasing cell death.     

Predation on crown-of-thorns starfish larvae by damselfishes

Examining the functional response of predators can provide insight into the role of predation in structuring prey populations and ecological communities. This study explored feeding behaviour and functional responses of planktivorous damselfishes when offered captive reared larvae of crown-of-thorns starfish, Acanthaster sp., with the aim of determining whether these predators could ever play a role in moderating outbreaks of Acanthaster sp. We examined predatory behaviour of 11 species of planktivorous damselfish, testing: (1) the relationship between predator size and predation rate, both within and among fish species; (2) consumption rates on larvae of Acanthaster sp. versus larvae of a common, co-occurring coral reef asteroid Linckia laevigata; (3) maximal feeding rates upon both Acanthaster sp. and L. laevigata; and (4) functional responses of planktivorous fishes to increasing densities of Acanthaster sp. Consumption rates of crown-of-thorns larvae by damselfishes were independent of predator size; however, there was a significant negative relationship between predator size and consumption rate of L. laevigata, when pooling across all predatory species. Some damselfishes, including Acanthochromis polyacanthus and Amblyglyphidodon curacao, consumed larval Acanthaster sp. at a greater rate than for L. laevigata. Most predatory species (all except A. curacao and Pomacentrus amboinensis) exhibited a Type II functional response whereby the increasing feeding rate decelerated with increasing prey density. In addition to revealing that a wide range of planktivorous fishes can prey upon larvae of Acanthaster sp., these data suggest that planktivorous damselfishes may have the capacity to buffer against population fluctuations of Acanthaster sp. Importantly, predators with Type II functional responses often contribute to stability of prey populations, though planktivorous fishes may be swamped by an abnormally high influx of larvae, potentially contributing to the characteristic population fluctuations of Acanthaster sp.     

The influence of temperature on life history traits of <Emphasis Type="Italic">Diadegma mollipla</Emphasis> (Hymenoptera: Ichneumonidae), an African parasitoid of the diamondback moth, <Emphasis Type="Italic">Plutella xylostella</Emphasis> (Lepidoptera: Plutellidae)

The use of Diadegma mollipla (Holmgren) as a biocontrol agent for diamondback moth (DBM)—a notorious pest of crucifers—can potentially be improved with increased scientific knowledge of the D. mollipla’s life history. This study investigated the influence of temperature on the life history traits of D. mollipla parasitizing DBM under laboratory conditions with 20 female D. mollipla for each of five constant temperatures i.e., 15, 20, 25, 30 and 35 °C. Data analysis using curvilinear regression models showed that D. mollipla’s highest intrinsic and finite rates of population increase, net reproductive rate, survival of immatures, daily fertility, shortest immature development and doubling times were achieved between 26 and 30 °C. Thus the results of this study provide evidence to that D. mollipla does not thrive well outside the 26–30 °C temperature range. These results provide baseline information for guiding the use of D. mollipla in the biological control of the destructive DBM pest.     

Gamete-associated flavobacteria of the oviparous Chinook salmon (<Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>) in lakes Michigan and Huron,North America

Flavobacterial diseases, caused by multiple members of the Family Flavobacteriaceae, elicit serious losses in wild and farmed fish around the world. Flavobacteria are known to be transmitted horizontally; however, vertical transmission has been suspected but proven only for one fish-pathogenic flavobacterial species (e.g., Flavobacterium psychrophilum). Herein, we report on the isolation and molecular identification of multiple Flavobacterium and Chryseobacterium taxa from the ovarian fluid and eggs of feral Great Lakes Chinook salmon (Oncorhynchus tshawytscha). Identified egg- and ovarian fluid-associated flavobacteria were either well-known flavobacterial fish pathogens (e.g., F. psychrophilum and F. columnare), most similar to emerging fish-associated flavobacteria (e.g., F. spartansii, F. tructae, F. piscis, C. piscium, C. scophthalmum), or were distinct from all other described Chryseobacterium and Flavobacterium spp., as determined by phylogenetic analyses using neighbor-joining, Bayesian, and Maximum Likelihood methodologies. The gamete-associated flavobacteria fell into three groups (e.g., those that were recovered from the ovarian fluid but not eggs; those that were recovered from the ovarian fluid and eggs; and those that were recovered from eggs but not ovarian fluid), a portion of which were recovered from eggs that were surface disinfected with iodophor at the commonly used dose and duration for egg disinfection. Some gamete-associated flavobacteria were also found in renal, splenic, and neurological tissues. Systemic polymicrobial infections comprised of F. psychrophilum and F. columnare were also detected at nearly an 11% prevalence. This study highlights the potential role that sexual products of female Great Lakes Chinook salmon may play in the transmission of fish-associated flavobacteria.     

Meta-analysis reveals asymmetric reduction in the genetic diversity of introduced populations of exotic insects

Factors promoting the invasion success of introduced populations have been receiving increased attention in studies of biological invasions. Previous reports have indicated that successful invasions may be attributable to reduced genetic diversity in the invasive species. However, there is large variation in the magnitude and direction of the impact of exotic species that have remained unexplained. Here, we present a structured meta-analysis of papers investigating the genetic diversity of native and introduced populations of exotic insects using nuclear microsatellites and mitochondrial DNA sequences. The results indicate that invasion by exotic insects had an overall reducing effect on the genetic diversity of the invading population, with nonzero effect sizes for the number of alleles (NA), observed heterozygosity (Ho), expected heterozygosity (He) and nucleotide diversity (Nd). However, when analyzing different orders (e.g., Lepidoptera, Hemiptera), the effect sizes of NA, Ho and Nd in Lepidoptera were found to bracket zero, as did the effect size of He in Hemiptera. These results suggest an asymmetric reduction in the genetic diversity of introduced populations of exotic insects, indicating diverse mechanisms underlying their successful invasion.     

New Zealand Eocene,Oligocene and Miocene Macrofossil and Pollen Records and Modern Plant Distributions in the Southern Hemisphere

The modern New Zealand flora has a relatively low number of families and genera in relation to land area, but well-preserved macrofossils and pollen from three sites in southern New Zealand suggest that the floras in Eocene, Oligocene and Miocene times were much more diverse at the generic level. At Pikopiko, Southland, a late Eocene in situ forest with fern understory was dominated by conifers, Casuarinaceae, Lauraceae, Nothofagus, Proteaceae, and mesothermal angiosperms including palms (aff. Calamus), Sapindaceae: Cupaniae and Picrodendraceae. At Newvale Mine, Southland, a leaf bed within a thick lignite seam represents leaf fossils preserved in a late Oligocene oligotrophic bog. This site demonstrates that Agathis, Dacrycarpus, Dacrydium, Halocarpus, Microcachrys, Podocarpus and Phyllocladus coexisted with diverse angiosperms including Nothofagus, Gymnostoma, Cunoniaceae, Ericaceae, Sapindaceae and several Proteaceae. Pollen data add Meliaceae, Myrtaceae, Onagraceae and Rubiaceae to the flora. At Foulden Maar, Otago, mummified leaves and flowers, including several with in situ pollen, demonstrate the existence of a diverse flora surrounding an Early Miocene lake. This site contains numerous monocot macrofossils including Astelia, Cordyline, Ripogonum and Typha, as well as the oldest fossils known for Orchidaceae and Luzuriagaceae. This flora was dominated by Lauraceae with affinities to Cryptocarya and Litsea, but other families include Araliaceae, Cunoniaceae, Elaeocarpaceae, Euphorbiaceae sensu lato, Menispermaceae, Myrsinaceae, Myrtaceae, Onagraceae, Proteaceae and Sterculiaceae. Many ferns, conifers, and Nothofagus are from lineages with Gondwanan ancestors, whereas other taxa show links to Australia (e.g., Gyrostemonaceae), New Caledonia (e.g., Beauprea) and South America (e.g., Luzuriaga, Fuchsia). Many of these taxa are now extinct in New Zealand, and therefore indicate much wider biogeographic ranges for many families and genera in the past.     

Taxonomic composition of bacteria associated with cultivated mollusks <Emphasis Type="Italic">Crassostrea lugubris</Emphasis> and <Emphasis Type="Italic">Perna viridis</Emphasis> and with the water of the Gulf of Nha Trang lagoon,Vietnam

One hundred and four strains of heterotrophic bacteria have been isolated and characterized from two species of bivalve mollusks cultivated in the Gulf of Nha Trang (Vietnam) and from the water of a mariculture farm. The isolates have been identified on the basis of morphological, physiological, biochemical, and chemotaxonomic properties, as well as by the content of G+C bases in DNA. In the microflora of mollusks, Vibrio alginolyticus was predominant; the pathogenic species V. harveyi and V. splendidus were found as well. Staphylococci and bacilli occupied the second place in abundance after vibrios. In addition, coryneforms and enterobacteria, as well as Pseudomonas spp. and Pseudoalteromonas spp., were revealed. The composition of the water microflora was more diverse as compared with the microflora of mollusks. In the water, Bacillus spp., Vibrio spp., and Pseudomonas spp. were predominant. Brevibacterium spp. and other coryneform bacteria, as well as enterobacteria, occurred in significant amounts. In addition, Pseudoalteromonas spp., Marinococcus sp., Halobacillus sp., Shewanella sp., Sulfitobacter sp., and bacteria of the CFB cluster were noticed. The presence of pathogenic and conditionally pathogenic bacterial species in the water and mollusks is probably the reason for the high death rate of cultivated animals at the mariculture farm.     

Definition of unit internal (physiological) time

We provide a definition of the unit internal (physiological) time based on metabolism. If q(t) is specific rate of metabolism, i.e. the amount of energy (oxygen) consumed by unit of active mass per physical time unit, the unit of physiological time τ(t) is defined as physical time, during which unit of active mass consumes one unit of energy: τ(t) = 1/q(t). The dimension of unit physiological time is the same as that of unit physical time and its value depends on q(t). Therefore, the unit physiological time τ(t) is a variable value, while the internal time is unequal relative to the physical time. The more internal time units τ, i.e., elementary acts of energy consumption, fit in the unit physical time t, the longer is the unit physical time for the unit active mass relative to the internal time unit, i.e., the physical time is seemingly slowed down. And, on the contrary, the less elementary acts of energy consumption take place during unit physical time, the shorter seems unit t, i.e. physical time is seemingly accelerated. Unequal course of the internal time is determined by the curve of specific metabolism q(t) during the life under specific conditions and, hence, internal time is individual. It has been questioned that the total (during lifetime) specific metabolism, often called Rubner constant, can serve as specie specific characteristic.     

DNA barcoding significantly improves resolution of invasive lionfish diet in the Northern Gulf of Mexico

Invasive Indo-Pacific red lionfish (Pterois volitans) have become well-established residents within reef communities across the western Atlantic Ocean where they pose substantial threats to native fish communities and reef ecosystems. Species-specific identification of prey is necessary to elucidate predator–prey interactions, but can be challenging with traditional visual identification methods given prey are often highly digested, thus not identifiable visually. To supplement visual diet analysis of lionfish (n = 934) sampled in the northern Gulf of Mexico, we applied DNA barcoding to identify otherwise unidentifiable fish prey (n = 696) via amplification of the cytochrome c oxidase subunit I (COI) of the mitochondrial genome. Barcoding nearly doubled the number of identifiable fish prey, thereby greatly enhancing our ability to describe lionfish diet. Thirty-three fish prey species were identified via barcoding, twenty-four of which were not previously detected by traditional methods. Some exploited reef fishes were newly reported (e.g., red snapper, Lutjanus campechanus) or found to constitute higher proportions of lionfish diet than previously reported (e.g., vermilion snapper, Rhomboplites aurorubens). Barcoding added a significant amount of new dietary information, and we observed the highest prey diversity reported to date for invasive lionfish. Potential cannibalism on juveniles also was identified via DNA barcoding, with the highest incidence corresponding to high lionfish densities, thus suggesting density-dependent prey demand may have driven this response. Overall, DNA barcoding greatly enhanced our ability to describe invasive lionfish diet in this study, suggesting that even studies with relatively large diet sample sizes could benefit from barcoding analysis.     

Olenekian brachiopods from the Kamenushka River basin,South Primorye: New data on the brachiopod recovery after the end-Permian mass extinction

The brachiopods (e.g., Heterelasma sp. nov., Uniplicatorhynchia sp. nov., Quadratirhynchia sp., and Sphriganaria sp.) discovered from the Olenekian deposits of the Kamenushka River basin and Russky Island, South Primorye, allow refining the information on the stratigraphic range of the relevant families and recovery of the biota after the Late Permian mass extinction. A new terebratulid genus, Bittnerithyris gen. nov., and a new rhynchonellid species, Piarorhynchella tazawai sp. nov., are established.     

Identification of the di-<Emphasis Type="Italic">n</Emphasis>-butyl phthalate-biodegrading strains and the biodegradation pathway in strain LMB-1

DNA isolated from a greenhouse soil (Nanjing, Jiangsu Province, China) was suitable for PCR amplification of gene segment coding for the 16S rRNA. Diverse PCR products were characterized by cloning and sequencing, and analysis of bacterial colonies showed the presence over 26 phyla. The most bacteria belonged to Proteobacteria, Actinobacteria, Gemmatimonadetes, Acidobacteria and Planctomycetes. Furthermore, after the enrichment procedure of DBP-degrading microorganisms, 4 strains were isolated from the soil sample with di-n-butyl phthalate (DBP) biodegradability, and they were identified to be Rhizobium sp., Streptomyces sp., Pseudomonas sp. and Acinetobacter sp. Analysis of the degradation products by LC-MS led to identification of metabolites of DBP in strain LMB-1 (identified as Rhizobium sp.) which suggests that DBP was degraded through β-oxidation, demethylation, de-esterification and cleavage of aromatic ring.     

A PCR-based analysis of plant DNA reveals the feeding preferences of <Emphasis Type="Italic">Apolygus lucorum</Emphasis> (Heteroptera: Miridae)

The mirid bug Apolygus lucorum (Meyer-Dür) (Heteroptera: Miridae) is a severe pest of cotton and other crops in China. The feeding preferences of this pest are unclear due to its frequent movement among different host plants and the inconspicuous signs of its feeding. Here, we present results of a field trial that used direct observation of bug densities and a PCR-based molecular detection assay to detect plant DNA in bugs to explore relationships between A. lucorum population abundance and its feeding preference between two host plants, Humulus scandens (Loureiro) Merrill and Medicago sativa L. The field-plot samples showed that A. lucorum adults generally prefer flowering host plants. Its density was significantly higher on flowering H. scandens than on seedlings of M. sativa, and a similarly higher bug density was observed on flowering M. sativa than on seedlings of H. scandens. In the laboratory, we designed two pairs of species-specific primers targeting the trnL-F region for H. scandens and M. sativa, respectively. The detectability of plant DNA generally decreased with time post-feeding, and the half-life of plant DNA detection (DS₅₀) in the gut was estimated as 6.26 h for H. scandens and 3.79 h for M. sativa with significant differences between each other. In mirid bugs exposed to seedlings of H. scandens and flowering M. sativa, the detection rate of M. sativa DNA was significantly higher than that of H. scandens. Meanwhile, in mirid bugs exposed to seedlings of M. sativa and flowering H. scandens, a significantly higher detection rate of H. scandens DNA was found. We developed a useful tool to detect the remaining plant food species specifically from the gut of A. lucorum in the current study. We provided direct evidence of its feeding preference between H. scandens and M. sativa at different growth stages, which strongly supported a positive correlation between population abundance and feeding preference of A. lucorum on different plants under field conditions. The findings provide new insights into the understanding of A. lucorum’s feeding preference, and are helpful for developing the strategies to control this pest.     

The fall and rise of the Icelandic Arctic fox (<Emphasis Type="Italic">Vulpes lagopus</Emphasis>): a 50-year demographic study on a non-cyclic Arctic fox population

In territorial species, observed density dependence is often manifest in lowered reproductive output at high population density where individuals have fewer resources or are forced to inhabit low-quality territories. The Arctic fox (Vulpes lagopus) in Iceland is territorial throughout the year and feeds mostly on birds, since lemmings are absent from the country. Thus, the population does not exhibit short-term population cycles that are evident in most of the species’ geographical range. The population has, however, gone through a major long-term fluctuation in population size. Because of the stability in hunting effort and reliable hunting records since 1958, the total number of adult foxes killed annually can be used as an index of population size (N _t ). An index of carrying capacity (K) from population growth data for five separate time blocks during 1958–2007 revealed considerable variation in K and allowed a novel definition of population density in terms of K, or N _t /K. Correlation analysis suggested that the reproductive rate was largely determined by the proportion of territorial foxes in the population. Variation in litter size and cub mortality was, on the other hand, related to climatic variation. Thus, Arctic foxes in Iceland engage in typical contest competition but can adapt their territory sizes in response to both temporal and spatial variation in carrying capacity, resulting in surprisingly little variation in litter size.