Fertility Life Table of <Emphasis Type="Italic">Tetranychus palmarum</Emphasis> Flechtmann &amp; Noronha (Acari: Tetranychidae) in Oil Palm

Some species of spider mites belonging to the Tetranychidae family are known to associate with oil palm (Elaeis guineensis Jacq. – Arecaceae). The occurrence of Tetranychus palmarum Flechtmann & Noronha (Acari, Tetranychidae) was verified on oil palm seedlings under greenhouse conditions in the State of Pará in Northern Brazil. Plants with colonies of T. palmarum presented yellowish spots on leaflets and leaves with chlorosis. The objective of this study was to access the biology and fertility life table of T. palmarum in E. guineensis leaves. The experiment was conducted under four constant temperatures, 22, 25, 28, and 31°C, at 70 ± 10% RH under a 12:12 LD photoperiod. The duration of the egg-to-adult period was 18.4 and 9.8 days, at 22 and 31°C, respectively. The parameters of the fertility life table showed that 28°C is most suitable for the development and reproduction of T. palmarum, with higher values for reproductive parameters (R _o , r _m , and λ) and lower values for duplicating the population (TD). Therefore, it is apparent that the best temperature conditions for the development of T. palmarum are found in the warmer regions of Brazil, such as those observed in northern Brazil.     

Morphological variation of <Emphasis Type="Italic">Melanargia russiae</Emphasis> (Esper, 1783) (Lepidoptera,Satyridae) from the main part of the range and in case of its expansion to the north under climate change conditions

A climate-related shift in the range boundaries of the western–central Eurasian subboreal species Melanargia russiae in the Ural region from the northern forest-steppe zone to pine–birch forests was found. Morphological variation of M. russiae wings from the boundary northern populations and populations from the main part of the range in the Urals were studied. The results of a complex analysis of the morphological traits (size, wing shape, and wing pattern eyespots) confirm the hypothesis that the local population formed in the south of Sverdlovsk oblast and contradict the hypothesis of the migratory origin of M. russiae imago.     

Population demographics and trade-offs to reproduction of an invasive and noninvasive species of <Emphasis Type="Italic">Rubus</Emphasis>

Do trade-offs between growth and reproduction differ between an invasive and noninvasive plant species and how do such trade-offs relate to population demographics? To help address these questions, we compared demographics for an invasive plant species, Rubus discolor, with a noninvasive congener, R. ursinus, in several populations of varying density. Removal of floral buds from reproductive canes increased the size of juvenile canes that arose from clonal sprouting in R. ursinus, suggesting a trade-off between current reproduction and growth. Removal of floral buds had no effect on growth of R. discolor. R. ursinus displayed trade-offs between reproduction (sexual and vegetative) and future growth based on negative correlations between leaf area production and both clonal sprouting and seedling production during the previous year. R. discolor did not exhibit these trade-offs. Both species had high population growth rates in low-density populations, but exhibited little or no growth in high-density populations. A life table response experiment was used to determine the underlying cause for the effect of density on population growth. For R. ursinus, lack of population growth in high-density populations was due primarily to increased mortality of clonally sprouting canes, while for R. discolor, it was due to decreased clonal cane production. Elasticity analysis revealed that clonal growth was more important than sexual reproduction for population growth of both species. However, elasticity values for sexual reproduction in R. discolor were greater in high- than low-density populations. This suggests an increased reliance on sexual reproduction in populations that had reached stable sizes, which could increase the capacity of R. discolor to disperse to new sites. Elasticity analyses were also used to simulate the efficacy of various control strategies for R. discolor. Control of this species could be attained by reducing clonal production within existing populations while reducing seed production to limit establishment of new populations.     

Microhabitat mosaics are key to the survival of an endangered ground beetle (<Emphasis Type="Italic">Carabus nitens</Emphasis>) in its post-industrial refugia

Biota dependant on early seral stages or frequently disturbed habitats belong to the most rapidly declining components of European biodiversity. This is also the case for Carabus nitens, which is threatened across Western and Central Europe. We studied one of the last remaining populations of this ground beetle in the Czech Republic, which inhabits post-extraction peat bogs. In line with findings from previous studies, we show that C. nitens prefers patches characterized by higher light intensity and lower vegetation cover. Abundance of females was positively correlated with the cover of plant species requiring higher temperature. In addition, we demonstrate its preference for periodically moist, but not wet or inundated plots, suggesting that the transition between dry heathland and wet peat bog might be the optimal habitat for this species. This hypothesis is further supported by results showing a positive correlation between the abundance of C. nitens and vegetation cover comprising of a mix of species typical for heathland, peat bog, and boreal habitats. Our results show that C. nitens mobility is comparable to other large wingless carabids. The maximum covered distance was ~?500 m in a month. To ensure the survival of this population, sites of recent peat extraction should be spared from reclamation and afforestation. In contrast, active measures should be taken to facilitate nutrient removal, disturbance of vegetation cover, and the creation of depressions with a humid microclimate. These actions will create a mosaic of heath, bog, and bare ground, which seems to be the preferred habitat of C. nitens at our study site.     

Patterns of microbial food webs in Mediterranean shallow lakes with contrasting nutrient levels and predation pressures

Hygraula nitens is a New Zealand native moth with aquatic larvae that feed on submerged aquatic plants. The larvae have been mainly observed using native Potamogeton and Myriophyllum species as a food source, although some studies reported larvae feeding on the alien macrophytes Hydrilla verticillata, Lagarosiphon major and Ceratophyllum demersum. Experimental mesocosm studies showed larvae had a major effect on H. verticillata, C. demersum, L. major, Elodea canadensis and Egeria densa. In both no choice and choice experiments H. nitens larvae showed a clear preference for and the highest consumption of C. demersum, while the native macrophyte Myriophyllum triphyllum ranked fourth out of five alien and two native plant species, indicating a preference of the larvae for alien macrophytes. Additional choice experiments using C. demersum, sampled from different waters in NZ, illustrated that there was a clear difference in H. nitens preference for plants based on their source. However although C. demersum had the lowest leaf dry matter content (LDMC) compared with the other macrophytes, neither the LDMC nor leaf carbon, nitrogen, phosphorus or total phenolic contents alone could explain the preferences of H. nitens, and we conclude that food choice is based on a combination of these and/or additional factors.     

Geographic patterns of genetic variation in nuclear and chloroplast genomes of two related oaks (<Emphasis Type="Italic">Quercus aliena</Emphasis> and <Emphasis Type="Italic">Q. serrata</Emphasis>) in Japan: implications for seed and seedling transfer

In this study, we assessed geographic patterns of genetic variations in nuclear and chloroplast genomes of two related native oaks in Japan, Quercus aliena and Q. serrata, in order to facilitate development of genetic guidelines for transfer of planting stocks for each species. A total of 12 populations of Q. aliena and 44 populations of Q. serrata were analyzed in this study. Genotyping of nuclear microsatellites in Q. aliena was done with only nine populations (n = 212) due to limited numbers of individuals in two populations, while all 12 populations (n = 89) were used in sequencing chloroplast DNA (cpDNA). In Q. serrata, 43 populations (n = 1032) were genotyped by nuclear microsatellite markers, while cpDNA of 44 populations (n = 350) was sequenced. As anticipated, geographic patterns detected in the variations of Q. aliena’s nuclear genome and its chloroplast haplotype distribution clearly distinguished northern and southern groups of populations. However, those of Q. serrata were inconsistent. The geographic distribution of its chloroplast haplotypes tends to show the predicted differentiation between northern and southern lineages, but geographic signals in the genetic structure of its nuclear microsatellites are weak. Therefore, treating northern and southern regions of Japan as genetically distinct transferrable zones for planting stocks is highly warranted for Q. aliena. For Q. serrata, the strong NE-SW geographic structure of cpDNA should be considered.     

Life History of <Emphasis Type="Italic">Cochliomyia macellaria</Emphasis> (Fabricius, 1775) (Diptera,Calliphoridae), a Blowfly of Medical and Forensic Importance

The life history traits of blow fly Cochliomyia macellaria (Fabricius, 1775) was studied under semi-controlled laboratory conditions at 29.14°C temperature, 72.53% relative humidity, and 12-h photoperiod. The raw data were analyzed based on the age-stage, two-sex life table, considering the development rates among individuals of both sexes. Cochliomyia macellaria survival rate was 0.43 (♂) and 0.40 (♀), while life expectancy was 17.9 (♂) and 20.9 (♀) days, for adult males and females, respectively. The total fecundity was 681.15 eggs/female, with an average of 3.65 batches/female and 199 eggs/batch. The intrinsic rate of increase (r) was 0.327 days^?1, the finite rate of population increase (λ) was 3.35 days^?1, the mean generation time (T) was 17.15 days, and the net reproduction rate (R ₀ ) was 272.46 offspring/individual. The population parameters found here corroborates that C. macellaria population act as a r selected species under laboratory conditions. Additionally, development data and accumulated degree days (ADD) for each stage of C. macellaria are provided and its implications for the forensic use are discussed.     

<Emphasis Type="Italic">Quinua</Emphasis> and Relatives (<Emphasis Type="Italic">Chenopodium</Emphasis> sect.<Emphasis Type="Italic">Chenopodium</Emphasis> subsect.<Emphasis Type="Italic">Celluloid</Emphasis>)

Traditionally viewed as an Andean grain crop,Chenopodium quinoa Willd. includes domesticated populations that are not Andean, and Andean populations that are not domesticated. Comparative analysis of leaf morphology and allozyme frequencies have demonstrated that Andean populations, both domesticated(quinua) and free-living(ajara), represent an exceptionally homogeneous unit that is well differentiated from allied domesticates of coastal Chile(quingua) and freeliving populations of the Argentine lowlands(C. hircinum). This pattern of relationships indicates that Andean populations represent a monophyletic crop/weed system that has possibly developed through cyclic differentiation (natural vs. human selection) and introgressive hybridization. Relative levels of variation suggest that this complex originated in the southern Andes, possibly from wild types allied withC. hircinum, with subsequent dispersal north to Colombia and south to the Chilean coast. Coastal populations were apparently isolated from post-dispersal differentiation and homogenization that occurred in the Andes. Other data point toward a center of origin in the northern Andes with secondary centers of genetic diversity subsequently developing in the southern Andes and the plains of Argentina. Comparative linkage of South American taxa, all tetraploid, with North American tetraploids of the subsection will eventually clarify this problem. While the possibility of a direct phyletic connection betweenC. quinoa and the Mexican domesticate(C. berlandieri subsp. nuttalliae,) cannot be excluded, available evidence indicates that the latter represents an autonomous lineage that is associated with the basal tetraploid, C. b. subsp.berlandieri, through var.sinuatum, whereas South American taxa show possible affinities to either var. zschackei or var.berlandieri. An extinct domesticate of eastern North America,C. b. subsp.jonesianum, represents either another instance of independent domestication, possibly from subsp. b. var.zschackei, or a northeastern outlier of subsp.nuttalliae.     

Intraspecific Variation in Mitogenomes of Five <Emphasis Type="Italic">Crassostrea</Emphasis> Species Provides Insight into Oyster Diversification and Speciation

A large number of Crassostrea oysters are found in Asia-Pacific. While analyses of interspecific variation have helped to establish historical relationships among these species, studies on intraspecific variation are necessary to understand their recent evolutionary history and current forces driving population biology. We resequenced 18 and analyzed 31 mitogenomes of five Crassostrea species from China: Crassostrea gigas, Crassostrea angulata, Crassostrea sikamea, Crassostrea ariakensis, and Crassostrea hongkongensis. Our analysis finds abundant insertions, deletions, and single-nucleotide polymorphisms in all species. Intraspecific variation varies greatly among species with polymorphic sites ranging from 54 to 293 and nucleotide diversity ranging from 0.00106 to 0.00683. In all measurements, C. hongkongensis that has the narrowest geographic distribution exhibits the least sequence diversity; C. ariakensis that has the widest distribution shows the highest diversity, and species with intermediate distribution show intermediate levels of diversity. Low sequence diversity in C. hongkongensis may reflect recent bottlenecks that are probably exacerbated by human transplantation. High diversity in C. ariakensis is likely due to divergence of northern and southern China populations that have been separated without gene flow. The significant differences in mitogenome diversity suggest that the five sister species of Crassostrea have experienced different evolutionary forces since their divergence. The recent divergence of two C. ariakensis populations and the C. gigas/angulata species complex provides evidence for continued diversification and speciation of Crassostrea species along China’s coast, which are shaped by unknown mechanisms in a north–south divide.     

Effect of salinity on vegetative growth and sexual reproduction of algae from the genus <Emphasis Type="Italic">Ardissonea</Emphasis> de Notaris (Bacillariophyta)

Effect of salinity on vegetative growth and sexual reproduction was investigated in laboratory cultures of a benthic diatom Ardissonea crystallina (C. Agardh) Grunow from the Black Sea and vegetative growth of Ardissonea sp. from the coast of Martinique, the Atlantic Ocean. Typical water salinity in the places where the populations inhabit differs two times. It was shown that the clones of both populations have a broad tolerance and great ability to adapt to changes in salinity. Cells of different size (i.e., in different stages of the life cycle) responded differently to desalination. Salinities optimal for vegetative growth of these species were not the same. Ardissonea from the coast of Martinique was more sensitive to low levels of salinity as compared with A. crystallina from the Black Sea. It was surprising that salinity optimal for vegetative growth and sexual reproduction of the Black Sea species did not coincide with the real salinity level of the Black Sea. Position of physiological optimums indicates an oceanic (or Mediterranean) origin of the Black Sea population of A. crystallina.     

Mitochondrial Evidence of Refugial Distribution of the Pygmy Field Mouse <Emphasis Type="Italic">Sylvaemus uralensis</Emphasis> Pall. (Rodentia,Muridae) in the Northwestern Caucasus

Variation of the 838-bp fragment of the mitochondrial cytb gene was analyzed in Sylvaemus uralensis from the northern macroslope of the Western Caucasus. On the basis of two fixed nonsynonymous substitutions, cytb sequences of the population sample studied can be considered as a distinct Lago-Naki haplogroup, which is clustered in the European cytb lineage. As estimated on the basis of the known rate of substitutions per third codon position in S. sylvaticus, the population must have been isolated for all or a part of the last glaciation period (10000 to 100000 years ago). The observed differentiation of cytb haplotypes is indicative of the refugial distribution of S. uralensis in the northern macroslope of the Western Caucasus, as well as of a secondary contact between the Caucasian and the Russian Plain populations during the Holocene.     

Otolith variation in Pacific herring (<Emphasis Type="Italic">Clupea pallasii</Emphasis>) reflects mitogenomic variation rather than the subspecies classification

Pacific herring (Clupea pallasii) is divided into three subspecies: two in northeast Europe and one in the north Pacific Ocean. Genetic studies have indicated that the populations in northeast Europe have derived from the northwest Pacific herring recently, or during the last 10–15 kyr, and that they are distinct from the population in the northeast Pacific. In addition, hybridization between the Pacific herring and the Atlantic herring has been documented. Otolith variation has been considered to be largely affected by environmental variation, but here we evaluate whether the genetic differentiation is reflected in otolith shape differences. A clear difference in otolith shape was observed between the genetically differentiated herring species Clupea harengus from the Atlantic and C. pallasii. The otolith shape of C. p. suworowi in the Barents Sea was different from the shape of C. pallasii in northern Norway and C. p. pallasii from the Pacific. Populations of C. p. pallasii, sampled east and west of the Alaska Peninsula, which belong to two genetically different clades of the C. p. pallasii in the Pacific Ocean, show a clear difference in otolith shape. C. p. suworowi and the local C. pallasii peripheral population in Balsfjord in northern Norway are more similar to the northwest Pacific herring (C. p. pallasii) than to the northeast Pacific herring (C. p. pallasii), both genetically and in otolith shape. The Balsfjord population, known to be influenced by introgression of mtDNA from the Atlantic herring, does not show any sign of admixture in otolith shape between the two species. A revised classification, considering the observed genetic and morphological evidence, should rather group the northwest Pacific herring in the Bering Sea together with the European populations of C. pallasii than with the northeast Pacific herring in the Gulf of Alaska.     

Polymorphism of the human <Emphasis Type="Italic">MDR1</Emphasis> gene in Siberian and Central Asian populations

The multidrug resistance gene MDR1 (ABCB1) codes for a P-glycoprotein that acts as an ATP-dependent transporter and is involved in removing drugs, xenobiotics, and peptides from the cell. MDR1 is expressed in the brain, kidneys, liver, and gastrointestinal tract. The P-glycoprotein is thought to play a role in individual resistance to xenobiotics and infections. Several polymorphisms of MDR1 are associated with the level of its expression and resistance to various neurodegenerative and gastrointestinal diseases. The allele and haplotype frequencies, genetic differentiation, and linkage disequilibrium for five MDR1 single nucleotide polymorphisms (3435C/T, 2677G/T/A, 1236C/T, +139C/T, and ?1G/A) were studied in the Russian, Tuvinian, and northern and southern Kyrgyz populations. Significant genetic differences were observed between Russians and northern Kyrgyz and between Tuvinians and northern Kyrgyz. The linkage disequilibrium pattern was characterized by high population specificity.     

Reproduction and development in some species of the weakly electric genus <Emphasis Type="Italic">Campylomormyrus</Emphasis> (Mormyridae,Teleostei)

Reproduction in captivity of four species of the mormyrid genus Campylomormyrus was investigated. Cyclical reproduction was provoked by changing water conductivity (C) alone: decreasing C led to gonadal recrudescence, an increase induced gonad regression. Data on the reproduction and development of three species are presented. All three species are indeterminate fractional spawners. Spawning intervals ranged from 6 to 66 days in C. rhynchophorus, 10–75 days in C. tshokwe, and 18 days in C. compressirostris (calculated values). Fecundities (eggs per fractional spawning) ranged from 70 to 1570 eggs in C. rhynchophorus, 100–1192 in C. tshokwe, and 38–246 in C. compressirostris. Spawnings/ovipositions occurred during the second half of the night; no parental care was observed; no special spawning substrates were necessary. C. compressirostris successfully spawned in breeding groups, C. rhynchophorus as pair. Agonistic behavior in the C. tshokwe pair forced us to divide the breeding tank; therefore, only ovipositions occurred. However, injection of an artificial GnRH hormone allowed us to obtain ripe eggs and sperm and to perform successful artificial reproduction. All three species produce yolky, slightly sticky eggs. Egg diameter ranges from 2.3–3.0 mm. Hatching occurred on day 3, feeding started on day 11. Transition from larval to juvenile stage occurred at around 20 mm total length (TL). At this size C. rhynchophorus developed a higher body than the two other species and differences between the species in the melanin pigmentation of the unpaired fins occurred. Between 32 and 35 mm TL the upper and lower jaws started to elongate.     

A review of the history of research and control of <Emphasis Type="Italic">Rhipicephalus</Emphasis> (<Emphasis Type="Italic">Boophilus</Emphasis>) <Emphasis Type="Italic">microplus</Emphasis>, babesiosis and anaplasmosis in Uruguay

The effect of five constant temperatures (16, 20, 24, 28 and 32 °C) on the development, survival and reproduction of Tetranychus cinnabarinus (Boisduval) [=?Tetranychus urticae Koch (red form)] fed on cassava leaves was examined in the laboratory at 85% relative humidity. Development time of various immature stages decreased with increasing temperature, with total egg-to-adult development time varying from 27.7 to 6.7 days. The lower thermal threshold for development was 10.8 °C and the thermal constant from egg to adult was 142.4 degree-days. Pre- and post-oviposition period and female longevity all decreased as temperature increased. The longest oviposition period was observed at 20 °C with 20.4 days. Under different temperatures, mated females laid, on average, 1.0, 2.9, 4.7, 4.7 and 4.9 eggs per day, respectively. The maximum fecundity (81.5 eggs per female) was at 28 °C and the intrinsic rate of increase (r _m ) was highest (0.25) at 32 °C. The results of this study indicate that T. cinnabarinus population could increase rapidly when cassava leaves serve as a food source. At the appropriate temperature T. cinnabarinus could seriously threaten growth of cassava.     

Life history differences and latitudinal variation in recruitment in two species of arctic-boreal perciform fishes,the fish doctor <Emphasis Type="Italic">Gymnelus viridis</Emphasis> and the Arctic shanny <Emphasis Type="Italic">Stichaeus punctatus</Emphasis>

Growth and recruitment were examined in two arctic-boreal, shallow water marine perciform species: the fish doctor Gymnelus viridis (Zoarcidae) and the Arctic shanny Stichaeus punctatus (Stichaeidae). G. viridis ranges in the Canadian Arctic from northern Hudson Bay to northern Ellesmere Island, whereas S. punctatus has a more southerly range from Nova Scotia coastal waters to northern Hudson Bay. At Nuvuk Islands in northeastern Hudson Bay, where the two species are sympatric in shallow water, they had comparable juvenile growth rates but temporal variation in 0+ recruitment was substantially greater in S. punctatus. This difference may stem from their contrasting early life histories. S. punctatus spawns large numbers of small demersal eggs that hatch into pelagic larvae, whereas G. viridis spawns small numbers of large demersal eggs that hatch directly into demersal juveniles with no pelagic stage, suggesting that recruitment in G. viridis should be less sensitive to yearly variation in the onset of the ice-free period in Hudson Bay and the subsequent pulse of pelagic invertebrate production. Relative to Nuvuk, recruitment variation in G. viridis was found to be greater at Resolute, Cornwallis Island, close to its northern range limit, whereas recruitment variation in S. punctatus was not evident in Newfoundland, closer to its southern range limit.     

Variability of quantitative morphological traits of mosquito larvae in some species of the <Emphasis Type="Italic">Culex pipiens</Emphasis> complex (Diptera,Culicidae)

The variability of 13 larval morphological characteristics in the urban mosquito, Culex pipiens f. molestus and Culex torrentium, was investigated. These species, belonging to the Culex pipiens complex, are characterized by rapid ecological and evolutionary processes following the human-induced environmental changes. A comparison of 4 geographical larval populations of C. pipiens f. molestus and Culex torrentium revealed different rates of divergence of different morphological traits. In the older form in terms of evolutionary development, C. torrentium, the two local populations differed in the structures of the trophic morphological complex characterized by a higher functional trophic load, while the structures with a lower functional load did not differ between the populations. On the contrary, the populations of the younger form, C. pipiens f. molestus, which had recently colonized new urban sites, differed in the structures with a low functional trophic load (some structures of siphon and abdominal segment IX), whereas structures of the trophic morphological complex were similar in different populations. The possible mechanisms of these morphological transformations and their role in the evolution of mosquito larvae are discussed.     

Experimental hybridization and an evaluation of the fertility of some forms of the house mouse supraspecies complex <Emphasis Type="Italic">Mus musculus</Emphasis> (Rodentia,Muridae)

The degree of development of the mechanisms of postcopulatory isolation was evaluated on the basis of experimental hybridization of representatives of three subspecies of M. musculus (M. m. musculus, M. m. wagneri, and M. m. gansuensis) and remote populations of the subspecies M. m. musculus. Experimental crosses between the different subspecies and populations indicated the presence of initial stages of postcopulatory reproductive isolation between some forms of house mice. In a number of crosses conducted between different populations and subspecies of M. musculus, asymmetry was observed. In one variant of mating, M. m. musculus (male) × M. m. wagneri (female), a reduced intensity of breeding and nonviability of pups were observed. A decrease in the intensity of reproduction was found in all variants of crosses that used male M. m. musculus from the city of Ishim. These data are assumed to confirm the previous assumption about the hybrid origin of mice inhabiting that city. The results confirm a significant level of divergence of the subspecies M. m. musculus and M. m. wagneri. Thus, initial stages both of post- and precopulatory isolation mechanisms between M. m. wagneri and M. m. musculus were shown.     

Population structure and distribution of deep-water cardinal fish <Emphasis Type="Italic">Epigonus occidentalis</Emphasis> (Epigonidae) and <Emphasis Type="Italic">Epigonus pandionis</Emphasis> (Epigonidae) in the Colombian Caribbean Sea

Cardinal fish are commercially valuable species. In the Colombian Caribbean Sea E. occidentalis and E. pandionis occurred in 37and 35% of tows during a research trawl survey (>200 m), respectively and the biology of these vulnerable species is unknown. The aim of this work is to describe the spatial distribution patterns of biomass, size structure and morphometric relationships of E. occidentalis and E. pandionis in deep waters of the Colombian Caribbean Sea. The samples were collected by trawling in depths between 200 and 550 m. No statistical differences were found in size by sexes in both species. In female and male E. occidentalis the growth was isometric, while in female and male E. pandionis it was positive allometric. E. occidentalis occurred mainly in the northern zone of Colombian Caribbean Sea, with highest aggregations off Santa Marta and Riohacha. E. pandionis was distributed in the northern area between Santa Marta and Riohacha and in the southern area between Cartagena and Morrosquillo Gulf. However, before the exploitation of these species is considered, further research is required to determine basic life history traits, such as growth, reproduction, recruitment, and mortality.