Predicting species responses to climate change: demography and climate microrefugia in California valley oak (Quercus lobata)

Anticipating species movement under climate change is a major focus in conservation. Bioclimate models are one of the few predictive tools for adaptation planning, but are limited in accounting for (i) climatic tolerances in preadult life stages that are potentially more vulnerable to warming; and (ii) local‐scale movement and use of climatic refugia as an alternative or complement to large‐scale changes in distribution. To assess whether these shortfalls can be addressed with field demographic data, we used California valley oak (Quercus lobata Nee), a long‐lived species with juvenile life stages known to be sensitive to climate. We hypothesized that the valley oak bioclimate model, based on adults, would overpredict the species' ability to remain in the projected persisting area, due to higher climate vulnerability of young life stages; and underpredict the potential for the species to remain in the projected contracting area in local‐scale refugia. We assessed the bioclimate model projections against actual demographic patterns in natural populations. We found that saplings were more constricted around surface water than adults in the projected contracting area. We also found that the climate envelope for saplings is narrower than that for adults. Saplings disappeared at a summer maximum temperature 3 °C below that associated with adults. Our findings indicate that rather than a complete shift northward and upward, as predicted by the species bioclimate model, valley oaks are more likely to experience constriction around water bodies, and eventual disappearance from areas exceeding a threshold of maximum temperature. Ours is the first study we know of to examine the importance of discrete life stage climate sensitivities in determining bioclimate modeling inputs, and to identify current climate change‐related constriction of a species around microrefugia. Our findings illustrate that targeted biological fieldwork can be central to understanding climate change‐related movement for long‐lived, sessile species.     

Population ecology and spatial synchrony in the abundance of leaf gall wasps within and among populations of valley oak (Quercus lobata)

What factors drive population variability through space and time? Here we assess patterns of abundance of seven species of gall wasps in three genera occurring on the leaves of valley oaks (Quercus lobata) at 10 sites throughout this species' statewide range in California, from 2000 to 2006. Our primary goals were to understand the factors driving variability in gall abundance and to assess the extent of spatial synchrony in gall wasp communities at both large and small geographic scales. On the large, statewide scale, there was significant site-to-site variation in gall abundance, driven in all cases primarily by differences in mean maximum seasonal temperatures, and lesser year-to-year variation. In contrast, on the small, local scale, differences were more pronounced from year to year than from tree to tree, and were to some extent correlated with differences in acorn production, suggesting an interaction with the reproductive effort of hosts. Significant spatial synchrony was detected, particularly at the statewide scale, but in no case did synchrony decline significantly with distance, despite sites being up to 741 km apart. Variation in spatial synchrony was correlated with a number of exogenous factors, including seasonal weather conditions, the acorn crop at the statewide scale and soil phosphorus availability at the local scale; however, most variation in spatial synchrony in our analyses remained unexplained.     

种子大小和干旱胁迫对辽东栎幼苗生长和生理特性的影响

在温室内遮阴条件下,设置80%、60%、40%和20%田间持水量(对照、轻度、中度和重度干旱)4个处理,研究种子大小和干旱胁迫对盆栽辽东栎幼苗生长和生理特性的影响。结果表明:大种子(3.05±0.38 g)幼苗的单株叶面积、总干质量和根冠比在所有处理均显著大于小种子(1.46±0.27 g)幼苗,前者的株高、基径、叶片数、比叶面积、相对生长率和净同化率等生长参数在轻度、中度和重度干旱处理均不同程度大于后者。大种子幼苗叶片过氧化物酶(POD)、过氧化氢酶(CAT)和超氧化物歧化酶(SOD)活性均高于小种子幼苗,前者叶片丙二醛(MDA)、可溶性蛋白、游离脯氨酸含量和叶绿素总量在部分干旱处理显著大于后者。除根冠比外,其他生长参数均随干旱胁迫增强逐渐减小,重度干旱处理大、小种子幼苗总干质量分别比对照降低19.4%和20.0%。POD、CAT和SOD活性均随干旱胁迫增强先升后降,在中度干旱处理,大、小种子幼苗POD活性分别显著高于对照126.7%和142.1%,CAT活性分别显著高于对照170.0%和151.9%。在重度干旱处理,大、小种子幼苗MDA含量分别显著高于对照86.5%和68.9%。可...     

The effects of seed size, cotyledon reserves, and herbivory on seedling survival and growth in Quercus rugosa and Q. laurina (Fagaceae)

In a greenhouse experiment, seedling survival of two oak species (Quercus rugosa and Q. laurina) was greatly affected by the excision of cotyledons 1 mo after germination, with a greater impact on Q. laurina. The effect of seed size was also significant for both species, with a positive correlation between seed mass and survival and growth. The effect of cotyledon excision on seedling growth persisted throughout the first growing season in Q. rugosa and was not analyzed for Q. laurina due to the low number of seedlings that survived cotyledon excision. Seed size significantly affected seedling height, diameter, leaf area, and biomass at 6 mo. Seed size and cotyledon retention affected the ability of Q. rugosa to recover from herbivory, as both factors had a significant effect on relative growth rates after aerial biomass removal. The results show that seedlings originating from large seeds can better endure loss of cotyledons and aerial biomass and thus are better equipped to confront stress early in their lives.     

辽东栎幼苗生长对种子大小和子叶去除处理的响应

该文在温室内盆播条件下,设置不去除子叶(对照)、去除1/4子叶(轻度子叶去除)、1/2子叶(中度子叶去除)和3/4子叶(重度子叶去除)4个处理,研究了种子大小和子叶去除处理对辽东栎幼苗生长及其形态学可塑性的影响.结果表明:(1)大种子幼苗株高、基茎、叶片数、单株叶面积、总干质量和根冠比在所有处理均大于小种子幼苗,除株高...     

沙埋和种子大小对柠条锦鸡儿种子萌发、出苗和幼苗生长的影响

研究了沙埋深度和种子大小对内蒙古毛乌素沙地植被群落中占优势的柠条锦鸡儿(Caragana korshinskii Kom.)种子萌发、出苗、幼苗存活和生长的影响.结果表明,沙埋深度显著影响柠条锦鸡儿的种子萌发率、休眠率、出苗率、幼苗存活率及生物量.在0.5-2cm的浅层沙埋下,种子萌发率、出苗率、幼苗存活率及生物量最高,休眠率最低;沙埋深度≥4 cm时,柠条锦鸡儿的种子萌发率、出苗率、幼苗存活率及生物量随着沙埋深度增加显著降低,而休眠率却显著升高;沙埋深度≥12 cm时,柠条锦鸡儿种子不能够出苗,幼苗也不能够存活.种子大小对柠条锦鸡儿种子萌发率没有显著影响,但对出苗率、幼苗存活率及生物量影响显著.在各个沙埋深度下,不同大小的柠条锦鸡儿种子间的萌发率没有显著差异.当沙埋深度≤6 cm时,不同大小的柠条锦鸡儿种子在同一沙埋深度下的出苗率间没有显著差异;但当沙埋深度≥8 cm时,在同一沙埋深度下,大种子的出苗率显著高于中种子和小种子的出苗率,而中种子和小种子出苗率间没有显著差异.0.5-10 cm的沙埋深度中,除6 cm和8 cm深度下中种子和小种子萌发幼苗的生物量间没有显著差异外,其余深度下,大种子萌发的幼苗的存活率及生物量显著高于同一沙埋深度下中种子萌发的幼苗的存活率及生物量,后者又显著高于小种子萌发的幼苗的存活率及生物量.可能正是种子萌发对沙埋环境的忍耐或响应能力以及种子的多态性提高了柠条锦鸡儿在毛乌素沙地的适合度,为其在流动或半流动沙丘环境中成功定居并形成优势群落奠定了基础.     

杭州石荠苧的传粉、小坚果散布和邻群大小

作者在杭州葛岭对杭州石荠苧自然居群的传粉和小坚果散布进行了详细的观察和实验。结果显示 ,以 2 6 0个传粉昆虫的飞行距离推算的花粉绝对散布方差为 0 .0 82 m2 ,798个小坚果的亲子散布方差为0 .1587m2。在上述散布特点和异交率为 0 .92 5条件下的邻群面积为 2 .2 33m 2 ,在有效密度为 4 7.7株 / m 2的有效密度下的邻群大小为 10 6 .5株。杭州石荠以异交为主 ,非常有限的基因散布导致邻群面积很小 ,居群容易发生遗传分化。     

间伐强度对秦岭南坡栓皮栎种子库及实生苗生长的影响

2010年8-11月和2011年8月,以秦岭南坡商洛地区近自然经营的不同间伐强度(30％、20％和10％)下的栓皮栎纯林为对象,分析了间伐4～5年后栓皮栎种子库的种子数量,实生苗的存活率和生长状况,及其与环境因子的关系.结果表明:与对照相比,间伐后第4年,间伐10％、20％和30％强度下种子库中的种子总数分别增加了2.0％、21.1％和34.8％,翌年1年生实生苗数量分别增加了28.5％、99.7％和122.2％;间伐5年后,间伐10％、20％和30％样地内实生苗的存活率分别提高了25.5％、235.7％和480.0％.间伐使林地的光照、温度、土壤水分和养分等环境因子发生变化,促进了各年龄段实生苗的高度、新梢生长量及叶面积指数的增长,且随间伐强度的增加,3项生长指标均增加.在郁闭度≥0.85的栓皮栎林,间伐强度30％(郁闭度保留0.6)最有利于栓皮栎种子萌芽和实生苗生长.     

Heterogeneity in recruitment and habitat patterns of valley oak (Quercus lobata Née) at the site and landscape scale in the Santa Monica Mountains, California, USA

Valley oak, Quercus lobata Née, has been a focus of conservation attention due to concerns about regeneration failure, habitat fragmentation and modification, and potential range contraction associated with climate change. A drawback of previous studies of valley oak stand structure and the regeneration problem has been their emphasis on one site or spatial and temporal scale. Generalization about valley oak dynamics from such studies is problematic because demographic parameters are not likely to be constant across sites, but heavily influenced by site-specific conditions. This study examines site-to-site variability across three locations in the Santa Monica Mountains National Recreation Area, California, USA. Variability in measures of stand structure and spatial pattern was examined across sites and within the landscape context. Estimates of expected landscape-scale evenness were derived by repeated random sampling of the data set and used as benchmarks for comparison with site-scale measures. Saplings were more abundant than expected at all three sites with sapling:adult ratios ranging from 0.5 to >1.0. Size-class structure suggested past recruitment problems at two sites, but recruitment at one site appears to have been steady for some time. Spatial distribution of adult stems at each site roughly corresponded to the estimated landscape pattern, but sapling establishment has shifted to north-to-east aspect hillsides and riparian-adjacent positions. In hilltop and swale habitats, adult valley oak are senescing without sapling recruitment. Shifting habitat-recruitment associations may lead to increasingly uneven distribution of stems across the landscape, altering landscape patterns and ecological processes.     

Growth of Valley Oak (<Emphasis Type="Italic">Quercus Lobata Nee</Emphasis>) in Four Floodplain Environments in the Central Valley of California

Although there have been numerous studies of California upland oak regeneration and growth there has been no research investigating oak sapling growth in riparian environments. This study looks at the growth response of young valley oak (Quercus lobata Nee), a dominant late successional riparian species in California, across environmental variation on a floodplain. Growth was measured over the course of three years at four different sites. The sites were chosen to represent the range of successional stages and surface age. Growth was significantly higher on younger, unforested sites. There was no difference in growth rate in the two forested sites (early successional cottonwood willow forest and late successional mature oak forest). Herbivory was highest in the cottonwood willow forest, where density of young oaks was also highest. The impact of flooding was measured on the youngest floodplain surface, an open floodplain restoration site where acorns were planted the year our study began. There was a significant negative impact of flooding on sapling growth in all but the first year of growth. Taken together these results suggest that existing forest trees and flooding both inhibit the growth of valley oaks on the floodplain, and that restoration in open sites may be more successful than restoration under an existing canopy.     

Mature male parr contribution to the effective size of an anadromous Atlantic salmon (Salmo salar) population over 30 years

We describe temporal changes in the genetic composition of a small anadromous Atlantic salmon (Salmo salar) population from South Newfoundland, an area where salmon populations are considered threatened (COSEWIC 2010). We examined the genetic variability (13 microsatellite loci) in 869 out‐migrating smolt and post‐spawning kelt samples, collected from 1985 to 2011 for a total of 22 annual collections and a 30 year span of assigned cohorts. We estimated the annual effective number of breeders (N_b) and the generational effective population size (N_e) through genetic methods and demographically using the adult sex ratio. Comparisons between genetic and demographic estimates show that the adult spawners inadequately explain the observed N_e estimates, suggesting that mature male parr are significantly increasing N_b and N_e over the study period. Spawning as parr appears to be a viable and important strategy in the near absence of adult males.     

Seasonal changes in pollinator activity influence pollen dispersal and seed production of the alpine shrub Rhododendron aureum (Ericaceae)

In alpine ecosystems, microscale variation in snowmelt timing often causes different flowering phenology of the same plant species and seasonal changes in pollinator activity. We compared the variations in insect visitation, pollen dispersal, mating patterns, and sexual reproduction of Rhododendron aureum early and late in the flowering season using five microsatellites. Insects visiting the flowers were rare early in the flowering season (mid-June), when major pollinators were bumblebee queens and flies. In contrast, frequent visitations by bumblebee workers were observed late in the season (late July). Two-generation analysis of pollen pool structure demonstrated that quality of pollen-mediated gene flow was more diverse late in the season in parallel with the high pollinator activity. The effective number of pollen donors per fruit (N(ep)) increased late in the season (N(ep) = 2.2-2.7 early, 3.4-4.4 late). However, both the outcrossing rate (t(m)) and seed-set ratio per fruit were smaller late in the season (t(m) = 0.89 and 0.71, seed-set ratio = 0.52 and 0.18, early and late in the season, respectively). In addition, biparental inbreeding occurred only late in the season. We conclude that R. aureum shows contrasting patterns of pollen movement and seed production between early and late season: in early season, seed production can be high but genetically less diverse and, during late season, be reduced, possibly due to higher inbreeding and inbreeding depression, but have greater genetic diversity. Thus, more pollinator activity does not always mean more pollen movement.     

Accounting for missing data in the estimation of contemporary genetic effective population size (Ne)

Theoretical models are often applied to population genetic data sets without fully considering the effect of missing data. Researchers can deal with missing data by removing individuals that have failed to yield genotypes and/or by removing loci that have failed to yield allelic determinations, but despite their best efforts, most data sets still contain some missing data. As a consequence, realized sample size differs among loci, and this poses a problem for unbiased methods that must explicitly account for random sampling error. One commonly used solution for the calculation of contemporary effective population size (N_e) is to calculate the effective sample size as an unweighted mean or harmonic mean across loci. This is not ideal because it fails to account for the fact that loci with different numbers of alleles have different information content. Here we consider this problem for genetic estimators of contemporary effective population size (N_e). To evaluate bias and precision of several statistical approaches for dealing with missing data, we simulated populations with known N_e and various degrees of missing data. Across all scenarios, one method of correcting for missing data (fixed‐inverse variance‐weighted harmonic mean) consistently performed the best for both single‐sample and two‐sample (temporal) methods of estimating N_e and outperformed some methods currently in widespread use. The approach adopted here may be a starting point to adjust other population genetics methods that include per‐locus sample size components.     

Interannual variation in effective number of breeders and estimation of effective population size in long‐lived iteroparous lake sturgeon (Acipenser fulvescens)

Quantifying interannual variation in effective adult breeding number (N_b) and relationships between N_b, effective population size (N_e), adult census size (N) and population demographic characteristics are important to predict genetic changes in populations of conservation concern. Such relationships are rarely available for long‐lived iteroparous species like lake sturgeon (Acipenser fulvescens). We estimated annual N_b and generational N_e using genotypes from 12 microsatellite loci for lake sturgeon adults (n = 796) captured during ten spawning seasons and offspring (n = 3925) collected during larval dispersal in a closed population over 8 years. Inbreeding and variance N_b estimated using mean and variance in individual reproductive success derived from genetically identified parentage and using linkage disequilibrium (LD) were similar within and among years (interannual range of N_b across estimators: 41–205). Variance in reproductive success and unequal sex ratios reduced N_b relative to N on average 36.8% and 16.3%, respectively. Interannual variation in N_b/N ratios (0.27–0.86) resulted from stable N and low standardized variance in reproductive success due to high proportions of adults breeding and the species' polygamous mating system, despite a 40‐fold difference in annual larval production across years (437–16 417). Results indicated environmental conditions and features of the species' reproductive ecology interact to affect demographic parameters and N_b/N. Estimates of N_e based on three single‐sample estimators, including LD, approximate Bayesian computation and sibship assignment, were similar to annual estimates of N_b. Findings have important implications concerning applications of genetic monitoring in conservation planning for lake sturgeon and other species with similar life histories and mating systems.     

Temporal genetic variation of mitochondrial DNA and the female effective population size of red drum (Sciaenops ocellatus) in the northern Gulf of Mexico

We studied genetic drift of mitochondrial DNA (mtDNA) haplotype frequencies in a natural population of red drum (Sciaenops ocellatus) from the northern Gulf of Mexico (Gulf). The amount of genetic drift observed across temporally adjacent year classes (1986–89) was used to estimate variance effective (female) population size (N_ef). N_ef was estimated to be 14 308 and the ratio of female effective size to adult female census size was approximately 0.004, which is among the lowest value reported for vertebrate animals. Low effective size relative to census size among red drum in the northern Gulf may result from yearly fluctuations in the number of breeding females, high variance in female reproductive success, or both. Despite low genetic effective size relative to census size, the genetic effective population size of red drum in the northern Gulf appears sufficiently large to preclude potentially deleterious effects of inbreeding.     

Effect of pollen load size and source (self, outcross) on seed and fruit production in highbush blueberry cv. 'Bluecrop' (VACCINIUM CORYMBOSUM; Ericaceae)

Reproductive fitness of a plant is ultimately determined by both number and quality of seed offspring. This is determined by sexual selection of pollen microspores and ovules during pollination and fertilization. These processes may include pollen competition and seed abortion, which reduce the number of microspores and ovules available for final seed production. Thus, even an excess of pollen microspores to ovules does not result in fertile seeds equal to ovule number. We investigated pollen requirements of highbush blueberry (Vaccinium corymbosum cultivar 'Bluecrop') for maximal seed production and how fertile seed number translates into fruit quality, since fruit quality would ultimately determine the dispersal of its offspring. We demonstrate that individual blueberry flowers with a mean of 106 ovules reach their maximum fruit set and mass and minimum time to ripen when 125 outcross pollen tetrads pollinate a flower, compared to 10 or 25. Three hundred tetrads resulted in the increase of fertile seeds, but did not result in a further increase of fruit mass or fruit set, or decrease in time to ripen. We also examined the effect of pure and mixed loads of self and outcross pollen (25 and 125 tetrads), and found no differences in fertile seed number, fruit mass, or percentage fruit set when pollen loads were either 25 self or outcross pollen tetrads, although number of days to ripen was significantly shorter by 8 d with 25 outcross tetrads. When the pollen load of 125 tetrads consisted of self or a 50:50 mixture of self and outcross pollen, fruit mass, days to ripen, and percentage fruit set were not different from loads of 125 outcross pollen. In addition, a pollen load of 25 outcross tetrads resulted in fertile seed number and fruit quality in between that of 25 self, and 125 self, 125 mixed, or 125 outcross tetrads. Large, small, and flat seed types were identified, and only large seeds (length = 1.7 mm) were fertile. These results improve our understanding of pollen load size and source requirements of a crop plant and the limits to pollen transfer when translated to fruit growth.     

The annual number of breeding adults and the effective population size of syntopic newts (Triturus cristatus, T. marmoratus)

Pond-breeding amphibians are deme-structured organisms with a population genetic structure particularly susceptible to demographic threats. We estimated the effective number of breeding adults (Nb) and the effective population size (Ne) of the European urodele amphibians Triturus cristatus (the crested newt) and T. marmoratus (the marbled newt), using temporal shifts in microsatellite allele frequencies. Eight microsatellite loci isolated from a T. cristatus library were used, five of which proved polymorphic in T. marmoratus, albeit with high frequencies of null alleles at two loci. Three ponds in western France were sampled, situated 4-10 kilometers apart and inhabited by both species. Parent-offspring cohort comparisons were used to measure Nb; samples collected at time intervals of nine or 12 years, respectively, were used to measure Ne. The adult population census size (N) was determined by mark-recapture techniques. With one exception, genetic distances (FST) between temporal samples were lower than among populations. Nb ranged between 10.6 and 101.8 individuals, Ne ranged between 9.6 and 13.4 individuals. For the pond where both parameters were available, Nb/N (overall range: 0.10-0.19) was marginally larger than Ne/N (overall range: 0.09-0.16), which is reflected in the temporal stability of N. In line with the observed differences in reproductive life-histories between the species, Nb/N ratios for newts were about one order of magnitude higher than for the anuran amphibian Bufo bufo. Despite of the colonization of the study area by T. cristatus only some decades ago, no significant genetic bottleneck could be detected. Our findings give rise to concerns about the long-term demographic viability of amphibian populations in situations typical for European landscapes.     

Exploring the structure of genetic variation and the influences of demography on effective population size in the gag grouper Mycteroperca microlepi (Goode & Bean)

Using 11 microsatellite markers, genetic analyses of three successive year-classes of gag Mycteroperca microlepis juveniles across the north-eastern Gulf of Mexico revealed a lack of spatial structure and very little temporal variation between year-classes. These results are consistent with long-term effective population sizes on the order of 30 000 adults. The importance of reproductive-style and sex-ratio variation is discussed as an important influence on long-term effective sizes.