Effects of human disturbance on reproductive success and population viability of Serapias cordigera (Orchidaceae)

Fragmentation of habitats by roads, railroads, fields, buildings and other human activities can affect population size, pollination success and fruit production, especially in plants showing pollinator limitation, such as Mediterranean orchids. In this study, we investigated the effect of human activity on the population dynamics and reproductive success of the orchid Serapias cordigera. Three anthropic and three natural populations were monitored over 14 years (1999–2012), classifying individuals into five stage classes and evaluating reproductive success. Population growth rates differed between anthropic and natural populations. Our results demonstrated that small anthropic populations have lower population viability compared with large natural populations. The proportion of flowering plants, the number of reproductive plants and the percentage of fruits were significantly lower in anthropic than in natural populations. This strong decline in fruit production in populations in urban areas may reflect lower pollination attraction and higher inbreeding in small than in natural populations. Calculation of extinction probabilities showed that the anthropic populations will drop below the survival threshold of 15–20 years. This study highlights that continued monitoring is needed to improve information on population viability and for appropriate conservation management. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 408–420.     

Outcomes of reciprocal invasions between genetically diverse and genetically uniform populations of <Emphasis Type="Italic">Daphnia obtusa</Emphasis> (Kurz)

Ecological theory predicts that genetic variation produced by sexual reproduction results in niche diversification and provides a competitive advantage both to facilitate invasion into genetically uniform asexual populations and to withstand invasion by asexual competitors. We tested the hypothesis that a large group of diverse clones of Daphnia obtusa has greater competitive advantage when invading into genetically uniform populations of this species than a smaller group with inherently less genetic diversity. We compared competitive outcomes to those of genetically uniform groups of small and large size invading into genetically diverse populations. Genetically diverse invaders of initially large group size increased their representation by more than those of initially small size; in contrast, genetically uniform invaders of initially large group size diminished on average by more than those of initially small size. These results demonstrate an advantage to the genetic variation produced by sexual reproduction, both in invasion and resisting invasion, which we attribute to competitive release experienced by individuals in genetically diverse populations.     

Variation in reproductive modes and population genetic structures of a monocarpic perennial herb,Cardiocrinum cordatum,in relation to habitat fragmentation

Reproductive systems are recognized as having a profound influence on the extent and structure of genetic variation in plant populations. To investigate the spatiotemporal variation in the reproductive modes (sexual and vegetative reproduction) and population genetic structures of a monocarpic perennial herb, Cardiocrinum cordatum (liliaceae), we selected a variety of habitats (e.g. large forested area including primeval forest, small fragmented secondary forest, and so on) around Sapporo City, Japan. We conducted breeding experiments, monitored the fate and growth of marked individuals for 3 years, and also analyzed the spatiotemporal genetic variation of flowering plants within the populations using allozyme variation. Plants emasculated prior to anthesis produced mature fruits in all populations examined. However, seed production was significantly lower in the small fragmented populations, possibly because of the low availability of pollinators and subsequent pollen limitation. In these fragmented populations, the mature flowering plants tended to be more dependent on vegetative reproduction for their recruitment, because they can only produce flowers once in their lifetime. Genetic diversity using samples from mature flowering plants in each population was lower in the small fragmented populations than in the large populations. In addition, although genotypic compositions in the fragmented populations were more or less similar during the 3 years of the study, the dominant genotypes changed temporally and spatially every year in the large populations. The present study demonstrated that the reproductive features of C. cordatum can be altered in various environmental conditions, such as habitat fragmentation, and these changes considerably affected the population genetic structures and vice versa.     

The ontogeny of flowering and sex expression in divergent populations of Lomatium grayi

Summary Populations of Lomatium grayi (Umbelliferae), an andromonoecious, perennial herb, differe in growth rates, flowering frequency, and survivorship. Effects of these different life histories on the ontogeny of sex expression were analyzed for plants from two populations grown from seedlings in a common garden and monitored for six years. Plants from Smoot Hill, Washington grew faster, had a higher probability of flowering at each age and size after the first year of growth, and a higher probability of flowering repeatedly among years than did plants from Clarkston, Washington. The proportion of plants producing some hermaphroditic flowers increased with plant size in both populations. Smoot Hill plants, however, were more likely to begin flowering as small, staminate plants than Clarkston plants. Clarkston plants did not begin flowering until they were older and larger, and most of these plants produced some hermaphroditic flowers when they began reproduction. The positive association between production of hermaphroditic flowers and both plant size and age was consistent with the hypothesis that hermaphroditic flowers are more costly to produce than staminate flowers.Although the populations did not differ in the total number of flowers per plant produced at any age or size, Smoot Hill plants consistently produced a lower percentage of hermaphroditic flowers than Clarkston plants at larger sizes and later ages. Consequently, selection for faster growth rates and higher flowering frequency at small sizes and early ages may have favored the more staminate-biased sex ratios in the Smoot Hill population.     

Synergistic effects of an extreme weather event and habitat fragmentation on a specialised insect herbivore

Habitat fragmentation is considered to be one of the main causes of population decline and species extinction worldwide. Furthermore, habitat fragmentation can decrease the ability of populations to resist and to recover from environmental disturbances such as extreme weather events, which are expected to occur at an increasing rate as a result of climate change. In this study, we investigated how calcareous grassland fragmentation affected the impact of the climatically extreme summer of 2003 on egg deposition rates, population size variation and survival of the blue butterfly Cupido minimus, a specialist herbivore of Anthyllis vulneraria. Immediately after the 2003 summer heat wave, populations of the host plant declined in size; this was paralleled with decreases in population size of the herbivore and altered egg deposition rates. In 2006 at the end of the monitoring period, however, most A. vulneraria populations had recovered and only one population went extinct. In contrast, several butterfly populations had gone extinct between 2003 and 2006. Extinction probability was significantly related to initial population size, with small populations having a higher risk of extinction than large populations. These results support the prediction that species of higher trophic levels are more susceptible to extinction due to habitat fragmentation and severe disturbances.     

EFFECTS OF HERBIVORE SIZE AND HUNGER LEVEL ON PERIPHYTON COMMUNITIES1

The effects of herbivore size and hunger level were tested on lotic periphyton community structure and ash-free dry mass (AFDM). My hypotheses were 1) that small herbivores would remove more periphyton per unit biomass than large herbivores of the same species because of energetic demands and 2) that within the same size class, starved herbivores would remove more periphyton than nonstarved herbivores. The herbivore used was the prosobranch snail Elimia clavaeformis Lea. Seven treatments were employed: 1) no snails (control); 2) small “starved” snails; 3) large “starved” snails; 4) small and large “starved” snails; 5) small fed snails; 6) large fed snails; and 7) small and large fed snails. Although snails removed significant amounts of periphyton AFDM relative to controls, neither snail size nor degree of starvation had a significant effect on loss of total AFDM. Small snails removed significantly more erect forms of Stigeoclonium tenue (C. A. Ag.) Kütz. than large snails, but snail size had no other significant effect. Starved snails removed significantly more Cocconeis placentula Ehr. than fed snails, suggesting that after the more susceptible growth forms (e.g. erect Stigeoclonium) were removed, Cocconeis cells became more vulnerable to grazing by hungry snails. When small and large snails were combined in chambers, large snails gained weight over time, whereas small snails lost weight. These results suggest that intraspecific competition may occur within populations of Elimia.     

Population viability analysis of Saxifraga cotyledon,a perennial plant with semelparous rosettes

The aim with this study was to analyse the population dynamics ofSaxifraga cotyledon, a rare, long lived herb, withsemelparous rosettes. In Sweden, S. cotyledon grows infragmented habitats at high altitude and is classified as vulnerable accordingto the IUCN system. From a five year demographical study we estimatedpopulationgrowth rates and extinction risks for one small and one large subpopulation. Inthe small subpopulation deterministic matrix simulations showed largevariation,with two transitions projecting negative and two projecting positive populationgrowth. The large subpopulation also showed large variation, but all yearlytransitions projected positive population growth. In both subpopulationssurvival and growth contributed more than twice as much to population growthrates than did sexual reproduction, vegetative reproduction and the seed bankall taken together. In stochastic simulations the maximum likelihood growthestimator waslarger than 1 for both subpopulations. None of the two subpopulations sufferfrom high extinction rates and although the effect of demographic stochasticityincrease extinction risk in small populations it is enough with 70 individualsfor a viable population of S. cotyledon. Hence, for thestudied population of S. cotyledon, rareness per se is nota good indicator of vulnerability.     

Population dynamics of the perennial Plantago media in semi‐natural grasslands

Abstract. In this study we investigated population dynamics of a perennial forb, Plantago media, in semi‐natural grasslands in southeastern Sweden. Plantago media is a rather common inhabitant of grasslands in this region, but it has been shown to experience dispersal limitations both among managed grassland sites and at potential dispersal routes along road verges. The demography of P. media was studied in 22 permanent plots at two sites over five years. A seed sowing experiment was also performed at each site. The life‐cycle of P. media includes seedlings, juveniles, small vegetative rosettes, large vegetative rosettes and flowering rosettes. The population growth rates (λ‐values) were negative for both populations, with one exception: the first year transition matrix at one site. The projected life span for individuals surviving from seedlings to flowering rosettes varied between 19.6 yr and 227.8 yr. Elasticity analysis showed that remaining in the large rosette stages, both vegetative and flowering, was the most important factor influencing population growth rate. LTRE analysis gave similar results, indicating that spatial and temporal variation have similar effects on the population growth rate. The expected time to extinction in populations with an initial size of between 100 and 1000 individuals varied between 60 and 200 yr. The seed sowing experiment showed that seedling emergence was enhanced by both seed addition and disturbance, suggesting that recruitment is limited by seed and microsite availability. Recruitment was not found to have much impact on population growth rate. However, long‐term population maintenance may depend on occasional small‐scale disturbances that enhance pulses of recruitment. Fragmented landscape is likely to effect plant populations including rare and endangered species as well as more common species which have limited dispersal mechanisms.     

Changes in the status of tortoise populations in Greece 1984–2001

Three species of tortoise (Testudinidae) occur in Greece (Testudo hermanni, T. graeca and the endemic T. marginata), all of which have been listed as rare or vulnerable. This paper describes the current situation of 75 populations that were last observed in the 1970s and 1980s and described in 1989. Twenty-nine populations had declined in density and/or status (significantly more than had improved), 10 of which were functionally extinct. A particularly notable loss was a dense population of T. marginata at Gytheion in the Peloponnese due to a widespread fire. Declining populations were significantly associated with identified high threat in 1989 and with close proximity to human settlement. There were no significant effects of tortoise species, area of site, characteristics of surrounding areas or original density on these changes. Declines were less associated with agriculture than predicted in 1989, reflecting changing economic conditions in Greece in the 1990s. Continuing threats to tortoise habitats make their long-term future appear bleak outside of protected areas. The risk of extinction from stochastic variation in small populations was also assessed, using the VORTEX program. This risk was much lower than that from habitat loss; 60–96% of populations of 100 tortoises would survive for 1000 years, depending on adult body size, compared to only 0–0.02% of habitats. The high survival potential of small populations would facilitate conservation of the high intraspecific diversity of tortoises in Greece.     

Identifying a surrogate metric for monitoring the population status of a secretive habitat specialist,the heath skink Liopholis multiscutata,in south‐eastern Australia

Threatened species that exist in small isolated populations are vulnerable to extinction processes, so effectively monitoring the trajectory of such populations will help determine the most appropriate management actions to combat extinction threats. In this study, we aimed to track the population status of the fossorial heath skink Liopholis multiscutata that is listed as threatened in Victoria, south‐eastern Australia, and exists there as a few small and highly disjunct populations, by using an appropriate surrogate population monitoring metric. This secretive lizard is a habitat specialist, is highly localised in Victoria and lives in warrens in semi‐arid heathland or mallee on large dunes. Survey data, which included every warren and their constituent burrows, as well as an assessment of whether each burrow was ‘active’, were collected for the four known Victorian populations in 2007 and annually during 2014–2018 inclusive. We compared five population indices per monitoring site: number of active warrens (NAW), number of active burrows (NAB), population area for 80% of active warrens (PA80), percentage of warrens that were active (PAW) and average number of active burrows per active warren. The heath skink currently occurs in small populations (8–46 active warrens) and these populations have typically declined over recent years. NAW was the most robust metric; NAB and PA80 did not reveal strong temporal trends. PAW indicated that inactive warrens and burrows persist less than a year and hence may provide information about recent (within months) population changes. It is imperative to establish a material link between the effective monitoring of small, vulnerable populations and the implementation of management actions that benefit such populations. Here, NAW could be used as a long‐term monitoring tool to provide an estimate of the minimum population size of the heath skink at a site. Its use would also ensure continuity in monitoring approaches for the Victorian populations.     

Demographic consequences of drought in the herbaceous perennial Cryptantha flava: effects of density,associations with shrubs,and plant size

The demographic consequences of a severe drought year were examined for two experimental plantings of the herbaceous desert perennial Cryptantha flava(Boraginaceae) in northeastern Utah, United States. A total of 6680 nutlets were planted individually or in clusters of four both under shrubs and in open microhabitats within two natural populations. Survival, growth, and flowering as a function of density and microhabitat were followed for 7 years, including 1 year when precipitation just before and during the growing season was 74.5% below normal. The design permitted assessment of how intraspecific density and shrub cover affect demographic response to drought. Mortality increased and flowering decreased dramatically during drought but neither varied with density or between shrub and open microhabitats. For plants growing under shrubs, survival (at Site 1) and growth (at Site 2) varied with shrub species. Average aboveground plant size also decreased during drought. Population size hierarchies were rearranged because larger plants lost leaf rosettes while many smaller plants grew. Density and microhabitat affected plant performance in non-drought years but more often at Site 1 than at Site 2. Individuals growing alone often were more likely to flower and/or produced more inflorescences when they did flower than did individuals growing with at least one other plant. However, for 2 years, survival rates at Site 1 were higher for plants growing in clumps than for single individuals. Shrubs also had mixed effects on plant performance. In some years, survival was higher under shrubs, but at Site 1 plants in the open often were more likely to flower and/or produced more inflorescences. Thus despite severe demographic consequences of drought, the study provided no evidence that intraspecific competition, interference by shrubs, or facilitation by shrubs increases under limited soil water.     

VARIATION IN INDIVIDUAL FLOWERING TIME AND REPRODUCTIVE SUCCESS OF AGALINIS STRICTIFOLIA (SCROPHULARIACEAE)

Field studies on two populations of Agalinis strictifolia were conducted over a 3-year period to investigate the relationship between flowering time of individuals and plant size, flowering duration, flower and fruit production, fruit predation, and growth rate. Seasonal patterns of pollinator visitation were compared with those of individual flowering time, flower density, percent fruit production, and mean seeds/fruit. In general, early and middle flowering individuals (as determined by either first flowering date or peak flowering) were larger, flowered longer, and produced more flowers and fruits than late flowering individuals. Early and middle flowering individuals (based on first flowering date) also grew faster than late flowering individuals. Although early and middle flowering individuals produced more fruits, fruit predators did not damage a disproportionate number of fruits compared to late flowering individuals. Patterns of bee visitation showed no association with seasonal patterns of flower density, percent fruit production, mean seed/fruit, or individual flowering time. In populations of A. strictifolia, it would seem that biotic or environmental determinants of growth rate (hence size and reproductive success) may be more important in generating variation in individual flowering time than patterns of pollinator visitation or fruit predation.     

Sexual reproductive biology of the endangered Japanese red maple (<Emphasis Type="Italic">Acer pycnanthum</Emphasis>)

Japanese red maple, Acer pycnanthum, is an endangered wetland species having dioecious sex expression. Prior studies conclude that land development has caused severe loss of habitat, and seedling regeneration is rare. However, information is lacking on sexual reproduction, which is the first critical stage of regeneration. My research objective was to describe flowering and fruiting characteristics of Japanese red maple, focusing on tree sizes at the onset of flowering, factors affecting flowering regularity and fruit abundance, periodicity of fruit production, and sex ratio. In a study of a young population, Japanese red maple initiated flowering at a small size under 100% light intensity. Initiation of flowering occurred at relatively lower height in males than females, and males flowered more regularly than females. A survey of 1,106 clones from 46 natural populations revealed that male clones were significantly more abundant than female clones. In a 4-year study of mature populations, nearly all clones flowered regularly, and fruit abundance of dominant females was typically high. Fruit abundance at the population level also remained high, although fruit abundance of individual female clones varied year by year. Therefore, sexual reproductive ability is highly vigorous. Sex ratio exhibited a major deviation from a 1:1 ratio in populations with a small number of clones. In conservation planning, we should prioritize avoidance of an unbalanced sex ratio by increasing population sizes. In addition, populations having many clones >40 cm dbh with large, well-lighted crowns have a high potential of regular and abundant seed production.     

The reproductive assurance benefit of selfing: importance of flower size and population size

Autonomous selfing can provide reproductive assurance (RA) for flowering plants that are unattractive to pollinators or in environments that are pollen limited. Pollen limitation may result from the breakdown of once-continuous habitat into smaller, more isolated patches (habitat fragmentation) if fragmentation negatively impacts pollinator populations. Here we quantify the levels of pollen limitation and RA among large and small populations of Collinsia parviflora, a wildflower with inter-population variation in flower size. We found that none of the populations were pollen limited, as pollen-supplemented and intact flowers did not differ in seed production. There was a significant effect of flower size on RA; intact flowers (can self) produced significantly more seeds than emasculated flowers (require pollen delivery) in small-flowered plants but not large-flowered plants. Population size nested within flower size did not significantly affect RA, but there was a large difference between our two replicate populations for large-flowered, small populations and small-flowered, large populations that appears related to a more variable pollination environment under these conditions. In fact, levels of RA were strongly negatively correlated with rates of pollinator visitation, whereby infrequent visitation by pollinators yielded high levels of RA via autonomous selfing, but there was no benefit of autonomous selfing when visitation rates were high. These results suggest that autonomous selfing may be adaptive in fragmented habitats or other ecological circumstances that affect pollinator visitation rates.     

Inbreeding and extinction: The effect of environmental stress and lineage

Human activities are simultaneously decreasing the size of wildlife populations (causing inbreeding) and increasing the level of stress that wildlife populations must face. Inbreeding reduces population fitness and increases extinction risk. However, very little information on the impact of stressful environments on extinction risk under inbreeding is available. We evaluated the impact of full sib inbreeding on extinction risk, using Drosophila melanogaster, in a benign and three stressful environments. The three stressful environments involved the addition to the medium of copper sulfate, methanol or alternating copper sulfate and methanol. There were 128 replicate populations for each of the four treatments. Under inbreeding, extinction rates were significantly higher in all three stressful environments compared with the benign environment. The percent extinct at generation eight (F = 0.826) for the four treatments were: 62.5% in the benign environment, 75.8%in the copper sulfate environment, 82.8% in the methanol environment, and 83.6% in the variable stress environment. However, the extinction rate in the variable stress environment did not differ significantly from the constant stress environments. Highly significant differences, among lineages, in extinction risk were detected. The results of this study indicate that wild populations are more vulnerable to inbreeding than indicated by extrapolation from captive environments.     

Flowering phenology of <Emphasis Type="Italic">Ulex europaeus</Emphasis>: ecological consequences of variation within and among populations

Reproductive phenology of gorse (Ulex europaeus L., Genisteae, Fabaceae) is unusual in that the onset and duration of flowering vary greatly among individuals within populations: some plants initiate flowering in autumn or winter and continue flowering through spring, others initiate flowering in early spring. To understand the origin of this diversity and its ecological consequences, we investigated flowering phenology of randomly sampled individuals from five different natural populations in Brittany (France). Reproductive success was evaluated for individuals with contrasting flowering patterns, from 16 natural populations. Flower production, pod production, seed production and seed predation were estimated. Plants initiating flowering in spring produced larger numbers of flowers and pods over a shorter period than plants flowering from winter to spring, which produced few flowers and pods at a time but over a longer period. Pod production of long-flowering plants did not differ significantly between winter and spring, but their pods were more intensively attacked by seed predators in spring than in winter. We discuss our results in relation to biotic and abiotic parameters. We postulate that long-flowering can be interpreted as a bet-hedging strategy, spreading the risk of pod failure (rotting or freezing) in winter and of seed predation in spring.     

Extinction of populations due to inbreeding depression with demographic disturbances

The process of population extinction due to inbreeding depression with constant demographic disturbances every generation is analysed using a population genetic and demographic model. The demographic disturbances introduced into the model represent loss of population size that is induced by any kind of human activities, e.g. through hunting and destruction of habitats. The genetic heterozygosity among recessive deleterious genes and the population size are assumed to be in equilibrium before the demographic disturbances start. The effects of deleterious mutations are represented by decreases in the growth rate and carrying capacity of a population. Numerical simulations indicate rapid extinction due to synergistic interaction between inbreeding depression and declining population size for realistic ranges of per-locus mutation rate, equilibrium population size, intrinsic rate of population growth, and strength of demographic disturbances. Large populations at equilibrium are more liable to extinction when disturbed due to inbreeding depression than small populations. This is a consequence of the fact that large populations maintain more recessive deleterious mutations than small populations. The rapid extinction predicted in the present study indicates the importance of the demographic history of a population in relation to extinction due to inbreeding depression.     

Genetic and phylogenetic consequences of island biogeography

Abstract.— Island biogeography theory predicts that the number of species on an island should increase with island size and decrease with island distance to the mainland. These predictions are generally well supported in comparative and experimental studies. These ecological, equilibrium predictions arise as a result of colonization and extinction processes. Because colonization and extinction are also important processes in evolution, we develop methods to test evolutionary predictions of island biogeography. We derive a population genetic model of island biogeography that incorporates island colonization, migration of individuals from the mainland, and extinction of island populations. The model provides a means of estimating the rates of migration and extinction from population genetic data. This model predicts that within an island population the distribution of genetic divergences with respect to the mainland source population should be bimodal, with much of the divergence dating to the colonization event. Across islands, this model predicts that populations on large islands should be on average more genetically divergent from mainland source populations than those on small islands. Likewise, populations on distant islands should be more divergent than those on close islands. Published observations of a larger proportion of endemic species on large and distant islands support these predictions.     

Evolutionary ecology of mast-seeding in temperate and tropical oaks (Quercus spp.)

Mast-seeding is the synchronous production of large seed crops within a population or community of species every two or more years. This paper addresses three non-mutually exclusive hypotheses explaining the evolution of mast-seeding in temperate tree species, especially the genus Quercus: (1) mast-seeding is a consequence of mast-flowering which evolves to increased pollination efficiency in mast-flowering years; (2) mast-seeding has evolved as an anti-predator adaptation by which large seed crops during mast years satiate the seed predators and allow survival of some of the seeds; (3) selection on seed size by habitat can indirectly affect the evolution of masting if trees with large seeds require more time to accumulate reserves to mature those seeds. I find support for the pollination hypothesis in several wind-pollinated temperate tree species but not oaks. However, oaks show evidence favoring the predation and seed size hypotheses. I then develop a model to illustrate the relationships among the three hypotheses in their effects on the evolution of masting. Finally, using data from herbaria and Floras, the influence of selection via flowering, fruiting, and seed size in the evolution of masting in tropical oaks is discussed. I conclude that the need for a supra-annual cue to synchronize flowering and fruiting as well as the larger seed size found in many tropical oak species should contribute to the evolution of masting to a greater extent than seen among temperate oaks.