Tree-Dependent Lichens and Beetles as Indicators in Conservation Forests

We tested whether the conspicuous lichen Lobaria pulmonaria indicates the number of tree-dependent, red-listed species in a hemiboreal forest in southern Sweden. In 18 naturally regenerated, mainly old deciduous forest plots considered to be of high or very high conservation value, the number of red-listed tree lichens or wood beetles was not positively correlated with the area of the forest stands studied (8–56 ha). The 8 stands with L. pulmonaria had about nine (median) red-listed lichens, but 10 stands without L. pulmonaria had only about one such species, a highly significant difference. Similarly, the variation between stands in the number of red-listed wood beetles was considerable, but it was very weakly correlated with the number of red-listed lichens. The number of red-listed wood beetles dependent on dead trees was not different in areas with or without Lobaria pulmonaria , but the number of red-listed wood beetles dependent on hollow trees was higher in stands with Lobaria pulmonaria (median of seven species) than in those without (three species) the lichen. Stands with this lichen species also contained significantly more of other types of lichens that have been proposed as indicators of forest continuity according to three lists. We stress the need for identification of species that could serve as indicators of different types of forest continuity and identify some organism groups that may indicate the different types.     

Community assembly in epiphytic lichens in early stages of colonization

Colonization studies may function as natural experiments and have the potential of addressing important questions about community assembly. We studied colonization for a guild of epiphytic lichens in a former treeless heathland area of 170 km2 in southwest Norway. We investigated if epiphytic lichen species richness and composition on aspen (Populus tremula) trees corresponded to a random draw of lichen individuals from the regional species pool. We compared lichen communities of isolated young (55-120 yr) and old (140-200 yr) forest patches in the heathland area to those of aspen forest in an adjacent reference area that has been forested for a long time. All thalli (lichen bodies) of 32 selected lichen species on trunks of aspen were recorded in 35 aspen sites. When data for each site category (young, old, and reference) were pooled, we found the species richness by rarefaction to be similar for reference sites and old sites, but significantly lower for young sites. The depauperated species richness of young sites was accompanied by a skew in species composition and absence of several species that were common in the reference sites. In contrast, genetic variation screened with neutral microsatellite markers in the lichen species Lobaria pulmonaria showed no significant differences between site categories. Our null hypothesis of a neutral species assembly in young sites corresponding to a random draw from the regional species pool was rejected, whereas an alternative hypothesis based on differences in colonization capacity among species was supported. The results indicate that for the habitat configuration in the heathland area (isolated patches constituting < 0.4% of the area) lichen communities may need a colonization time of 100-150 yr for species richness to level off, but given enough time, isolation will not affect species richness. We suggest that this contradiction to expectations from classical island equilibrium theory results from low extinction rates.     

Microsite-limited recruitment controls fern colonization of post-agricultural forests

Assessing the relative roles of dispersal limitation and environmental effects in population dynamics and community assembly is fundamental to understanding patterns of species distribution and diversity. In forests growing on abandoned agricultural lands, both legacies of vegetation disturbance and changes in the abiotic environment shape the diversity and composition of recovering communities. Here I specify how interactions among historical, environmental, and biological factors influence species distributions, focusing on three fern species with contrasting distributions across forests of different history in central New York, USA: Dryopteris carthusiana, Dryopteris intermedia, and Polystichum acrostichoides. Using population surveys, spore-trap and spore-bank studies, and a three-year field experiment, I compare demographic rates among species and between forest types to determine which life history stages limit colonization and which traits explain species distributions. Adult plants of all three species were larger and more likely to produce spores in post-agricultural forests than in adjacent, uncleared stands. Though lower population densities led to fewer spores in post-agricultural soils, spore availability still exceeded recruitment by four to five orders of magnitude. Sowing additional spores had relatively little effect, while microhabitat conditions had the greatest impact on establishment rates. Given similar microsites, the two forest types had equal rates of establishment, but some forest-floor features preferentially occupied by juvenile plants were less frequent in post-agricultural stands. The availability of suitable sites for establishment, created by small-scale heterogeneity on forest floors, thus limits both the growth of fern populations and the colonization of new habitats. In fact, reduced microtopographic variation in post-agricultural forests may represent a greater hindrance to plant establishment than changes in mean environmental conditions. Among the three fern species, establishment rates differed as species distributions would predict, with the strongest colonizer consistently having the highest rates and the slowest colonizer the lowest. Rather than random or trait-mediated dispersal, the different distributions of these species reflect life history traits that determine establishment rates and thus colonization ability. This case study demonstrates that ecological interactions based on the unique life histories of individual species can override dispersal in determining species distributions.     

Hunting Increases Dispersal Limitation in the Tree Carapa procera, a Nontimber Forest Product

Abstract:  The sustainability of seed extraction from natural populations has been questioned recently. Increased recruitment failure under intense seed harvesting suggests that seed extraction intensifies source limitation. Nevertheless, areas where more seeds are collected tend to also have more intense hunting of seed-dispersing animals. We studied whether such hunting, by limiting disperser activity, could cause quantitative dispersal limitation, especially for large crops and for crops in years of high seed abundance. In each of four Carapa procera (Meliaceae) populations in French Guiana and Surinam, two with hunting and two without, we compared seed fate for individual trees varying in crop size in years of high and low population-level seed abundance. Carapa seeds are a nontimber forest product and depend on dispersal by scatter-hoarding rodents for survival and seedling establishment. Hunting negatively affected the proportion of seeds dispersed and caused greater numbers of seeds to germinate or be infested by moths below parent trees, where they would likely die. Hunting of seed-dispersing animals disproportionally affected large seed crops, but we found no additional effect of population-level seed abundance on dispersal rates. Consistently lower rates of seed dispersal, especially for large seed crops, may translate to lower levels of seedling recruitment under hunting. Our results therefore suggest that the subsistence hunting that usually accompanies seed collection is at the cost of seed dispersal and may contribute to recruitment failure of these nontimber forest products. Seed extraction from natural populations may affect seedling recruitment less if accompanied by measures adequately incorporating and protecting seed dispersers.     

Epiphyte metapopulation dynamics are explained by species traits, connectivity, and patch dynamics

The colonization-extinction dynamics of many species are affected by the dynamics of their patches. For increasing our understanding of the metapopulation dynamics of sessile species confined to dynamic patches, we fitted a Bayesian incidence function model extended for dynamic landscapes to snapshot data on five epiphytic lichens among 2083 mapped oaks (dynamic patches). We estimate the age at which trees become suitable patches for different species, which defines their niche breadth (number of suitable trees). We show that the colonization rates were generally low, but increased with increasing connectivity in accordance with metapopulation theory. The rates were related to species traits, and we show, for the first time, that they are higher for species with wide niches and small dispersal propagules than for species with narrow niches or large propagules. We also show frequent long-distance dispersal in epiphytes by quantifying the relative importance of local dispersal and background deposition of dispersal propagules. Local stochastic extinctions from intact trees were negligible in all study species, and thus, the extinction rate is set by the rate of patch destruction (tree fall). These findings mean that epiphyte metapopulations may have slow colonization-extinction dynamics that are explained by connectivity, species traits, and patch dynamics.     

Effects of Forest Fragmentation on Seed Dispersal and Seedling Establishment in Ornithochorous Trees

Abstract: Habitat fragmentation increases seed dispersal limitation across the landscape and may also affect subsequent demographic stages such as seedling establishment. Thus, the development of adequate plans for forest restoration requires an understanding of mechanisms by which fragmentation hampers seed delivery to deforested areas and knowledge of how fragmentation affects the relationship between seed‐deposition patterns and seedling establishment. We evaluated the dispersal and recruitment of two bird‐dispersed, fleshy‐fruited tree species (Crataegus monogyna and Ilex aquifolium) in fragmented secondary forests of northern Spain. Forest fragmentation reduced the probability of seed deposition for both trees because of decreased availability of woody perches and fruit‐rich neighborhoods for seed dispersers, rather than because of reductions in tree cover by itself. The effects of fragmentation went beyond effects on the dispersal stage in Crataegus because seedling establishment was proportional to the quantities of bird‐dispersed seeds arriving at microsites. In contrast, postdispersal mortality in Ilex was so high that it obscured the seed‐to‐seedling transition. These results suggest that the effects of fragmentation are not necessarily consistent across stages of recruitment across species. Habitat management seeking to overcome barriers to forest recovery must include the preservation, and even the planting, of fleshy‐fruited trees in the unforested matrix as a measure to encourage frugivorous birds to enter into open and degraded areas. An integrative management strategy should also explicitly consider seed‐survival expectancies at microhabitats to preserve plant‐population dynamics and community structure in fragmented landscapes.     

On the relative contributions of wind vs. animals to seed dispersal of four Sierra Nevada pines

Selective pressures that influence the form of seed dispersal syndromes are poorly understood. Morphology of plant propagules is often used to infer the means of dispersal, but morphology can be misleading. Several species of pines, for example, have winged seeds adapted for wind dispersal but owe much of their establishment to scatter-hoarding animals. Here the relative importance of wind vs. animal dispersal is assessed for four species of pines of the eastern Sierra Nevada that have winged seeds but differed in seed size: lodgepole pine (Pinus contorta murrayana, 8 mg); ponderosa pine (Pinus ponderosa ponderosa, 56 mg); Jeffrey pine (Pinus jeffreyi, 160 mg); and sugar pine (Pinus lambertiana, 231 mg). Pre-dispersal seed mortality eliminated much of the ponderosa pine seed crop (66%), but had much less effect on Jeffrey pine (32% of seeds destroyed), lodgepole pine (29%), and sugar pine (7%). When cones opened most filled seeds were dispersed by wind. Animals removed > 99% of wind-dispersed Jeffrey and sugar pine seeds from the ground within 60 days, but animals gathered only 93% of lodgepole pine seeds and 38% of ponderosa pine seeds during the same period. Animals gathered and scatter hoarded radioactively labeled ponderosa, Jeffrey, and sugar pine seeds, making a total of 2103 caches over three years of study. Only three lodgepole pine caches were found. Caches typically contained 1-4 seeds buried 5-20 mm deep, depths suitable for seedling emergence. Although Jeffrey and sugar pine seeds are initially wind dispersed, nearly all seedlings arise from animal caches. Lodgepole pine is almost exclusively wind dispersed, with animals acting as seed predators. Animals treated ponderosa pine in an intermediate fashion. Two-phased dispersal of large, winged pine seeds appears adaptive; initial wind dispersal helps to minimize pre-dispersal seed mortality whereas scatter hoarding by animals places seeds in sites with a higher probability of seedling establishment.     

Recreational Portage Trails as Corridors Facilitating Non-Native Plant Invasions of the Boundary Waters Canoe Area Wilderness (U.S.A.)

Abstract:  Wilderness areas are protected and valued in part for recreation; recreational use, however, can negatively impact these areas. In particular, recreational use can facilitate transport of non-native propagules and create open sites for establishment of non-native species. We examined the role of recreational portage trails in the introduction and establishment of non-native plants into the Boundary Waters Canoe Area Wilderness of northern Minnesota (U.S.A.). On 20 portages, we sampled non-native plant richness and cover at four distances (0, 10, 25, and 50 m) from trails. Non-native richness and cover were not related to distance from wilderness entry point. Non-native richness and cover were, however, negatively related to distance from trails. All six non-native species we observed were either directly on or within 1 m of trails. These results suggest that recreational trails act as corridors facilitating invasions of non-native plants into wilderness areas. It remains unclear, however, whether these effects are caused by dispersal of propagules, creation of bare ground, or changes in the native plant community.     

林木外生菌根真菌彩色豆马勃巢式PCR检测

为探讨林木外生菌根真菌的分子检测方法,利用真菌通用引物ITS1-F/ITS4-B扩增了外生菌根真菌彩色豆马勃的核糖体DNA内转录间隔区并进行了序列测定.通过序列比较,设计了一对彩色豆马勃特异性引物PtF/PtR.利用该对特异性引物与ITS1-F/ITS4-B组合进行巢式PCR,能从供试的彩色豆马勃10个菌株中特异性地扩增出1条347 bp的条带,而供试的其他6个参比菌株未出现扩增产物.经分析,该巢式PCR检测技术的灵敏度可达到10 fg的DNA,是常规PCR检测的1 000倍.利用该技术从马尾松苗木菌根中检测到目的外生菌根真菌.这表明采用本研究设计的特异性引物,利用巢式PCR技术可以灵敏、准确地从林木外生菌根中检测出彩色豆马勃.     

Introduced Birds and the Fate of Hawaiian Rainforests

Abstract:  The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.     

Dispersal, competition, and shifting patterns of diversity in a degraded oak savanna

Diversity is a balance between processes that add and limit species (e.g., dispersal vs. competition), but reconciling their contributions remains a challenge. Recruit-ment-based models predict that dispersal barriers are most limiting for diversity, while competition-based models predict that dispersal matters only when competition is minimized. Testing these models is difficult because their influence varies with scale and site productivity. In a degraded oak savanna, we used plot-level (seed additions, burning) and site-level (proportions of regional functional groups found locally) analyses in areas with variable soil depth to examine how dispersal and competition influence diversity. At the plot level, added species persisted where they were formerly absent, but few established naturally despite fire-induced resource enrichment and nearby populations, revealing the importance of dispersal limitation for diversity. This result did not vary with soil depth or standing crop. Although competition could not prevent establishment in unburned plots, it significantly lowered survival, indicating that resource limitations exacerbate dispersal inefficiencies. At the site level, the concordance between regional and local diversity for native species was associated with soil depth heterogeneity, not dispersal or competition. This suggests that persistence is determined primarily by the influence of the environment on population demographics. Given that the formation of new populations is unlikely, those remaining appear to be confined to optimal habitat where they resist competitive or stochastic displacement, possibly explaining why species loss is rare despite substantial habitat loss and invasion. For exotics, there was no relationship between diversity and soil depth heterogeneity. Annuals with presumed dispersal capabilities were significantly overrepresented in all sites while perennial forbs, the largest regional functional group, were significantly underrepresented. We interpret the native-exotic discrepancies as reflecting the recent arrival of exotics (150 years ago), suggesting that local establishment occurs slowly even for species with regional prevalence. The accumulation lag may be explained by the need for founder populations to be demographically stable; otherwise persistence requires continual immigration favoring overrepresentation by dispersers. Our findings support the view that dispersal limitation restricts diversity within plant communities, but suggests that the impacts of environment on demographic performance ultimately determine the pattern and rate of community assembly.     

鼎湖山锥栗种子扩散过程中死亡因素分析

研究了锥栗(Castanopsischinensis)种子扩散过程中死亡的主要因素。实验分为无处理组、鸟类取食组、排除鸟类取食组、排除哺乳动物组和种子袋法。结果表明,扩散前种子主要受栗实象甲(Curculiodavidi)取食和病原体感染,其取食率在鼎湖山3种林型之间显示了较大差异。排除实验的研究表明,锥栗种子在扩散后承受啮齿类、鸟类等动物取食和病原体感染致死的巨大压力,其中无覆盖处理实验组的种子取食平均比例高达93.3%,鸟类对种子取食率约为80%,排除鸟类实验组中的种子平均取食率为92.4%,非哺乳动物对种子的取食在3个林型有差异,但不是显著性的。因此,巨大的种子死亡或丧失是锥栗自然更新率非常低的主要原因。     

Role of Propagule Size in the Success of Incipient Colonies of the Invasive Argentine Ant

Abstract: Factors that contribute to the successful establishment of invasive species are often poorly understood. Propagule size is considered a key determinant of establishment success, but experimental tests of its importance are rare. We used experimental colonies of the invasive Argentine ant (   Linepithema humile ) that differed both in worker and queen number to test how these attributes influence the survivorship and growth of incipient colonies. All propagules without workers experienced queen mortality, in contrast to only 6% of propagules with workers. In small propagules (10–1,000 workers), brood production increased with worker number but not queen number. In contrast, per capita measures of colony growth decreased with worker number over these colony sizes. In larger propagules ( 1,000–11,000 workers), brood production also increased with increasing worker number, but per capita brood production appeared independent of colony size. Our results suggest that queens need workers to establish successfully but that propagules with as few as 10 workers can grow quickly. Given the requirements for propagule success in Argentine ants, it is not surprising how easily they spread via human commerce.     

Ecological responses to variation in seasonal snow cover

Seasonal snow is among the most important factors governing the ecology of many terrestrial ecosystems, but rising global temperatures are changing snow regimes and driving widespread declines in the depth and duration of snow cover. Loss of the insulating snow layer will fundamentally change the environment. Understanding how individuals, populations, and communities respond to different snow conditions is thus essential for predicting and managing future ecosystem change. We synthesized 365 studies that examined ecological responses to variation in winter snow conditions. This research encompasses a broad range of methods (experimental manipulations, measurement of natural snow gradients, and long-term monitoring), locations (35 countries), study organisms (plants, mammals, arthropods, birds, fish, lichen, and fungi), and response measures. Earlier snowmelt was consistently associated with advanced spring phenology in plants, mammals, and arthropods. Reduced snow depth often increased mortality or physical injury in plants, although there were few clear effects on animals. Neither snow depth nor snowmelt timing had clear or consistent directional effects on body size of animals or biomass of plants. However, because 96% of studies were from the northern hemisphere, the generality of these trends across ecosystems and localities is also unclear. We identified substantial research gaps for several taxonomic groups and response types; research on wintertime responses was notably scarce. Future research should prioritize examination of the mechanisms underlying responses to changing snow conditions and the consequences of those responses for seasonally snow-covered ecosystems.     

Dispersal ecology of deadwood organisms and connectivity conservation

Limited knowledge of dispersal for most organisms hampers effective connectivity conservation in fragmented landscapes. In forest ecosystems, deadwood‐dependent organisms (i.e., saproxylics) are negatively affected by forest management and degradation globally. We reviewed empirically established dispersal ecology of saproxylic insects and fungi. We focused on direct studies (e.g., mark‐recapture, radiotelemetry), field experiments, and population genetic analyses. We found 2 somewhat opposite results. Based on direct methods and experiments, dispersal is limited to within a few kilometers, whereas genetic studies showed little genetic structure over tens of kilometers, which indicates long‐distance dispersal. The extent of direct dispersal studies and field experiments was small and thus these studies could not have detected long‐distance dispersal. Particularly for fungi, more studies at management‐relevant scales (1–10 km) are needed. Genetic researchers used outdated markers, investigated few loci, and faced the inherent difficulties of inferring dispersal from genetic population structure. Although there were systematic and species‐specific differences in dispersal ability (fungi are better dispersers than insects), it seems that for both groups colonization and establishment, not dispersal per se, are limiting their occurrence at management‐relevant scales. Because most studies were on forest landscapes in Europe, particularly the boreal region, more data are needed from nonforested landscapes in which fragmentation effects are likely to be more pronounced. Given the potential for long‐distance dispersal and the logical necessity of habitat area being a more fundamental landscape attribute than the spatial arrangement of habitat patches (i.e., connectivity sensu strict), retaining high‐quality deadwood habitat is more important for saproxylic insects and fungi than explicit connectivity conservation in many cases.     

Analyzing the spatial structure of a Sri Lankan tree species with multiple scales of clustering

Clustering at multiple critical scales may be common for plants since many different factors and processes may cause clustering. This is especially true for tropical rain forests for which theories explaining species coexistence and community structure rest heavily on spatial patterns. We used point pattern analysis to analyze the spatial structure of Shorea congestiflora, a dominant species in a 25-ha forest dynamics plot in a rain forest at Sinharaja World Heritage Site (Sri Lanka), which apparently shows clustering at several scales. We developed cluster processes incorporating two critical scales of clustering for exploring the spatial structure of S. congestiflora and interpret it in relation to factors such as competition, dispersal limitation, recruitment limitation, and Janzen-Connell effects. All size classes showed consistent large-scale clustering with a cluster radius of approximately 25 m. Inside the larger clusters, small-scale clusters with a radius of 8 m were evident for recruits and saplings, weak for intermediates, and disappeared for adults. The pattern of all trees could be divided into two independent patterns: a random pattern (nearest neighbor distance > 8 m) comprising approximately 12% of the trees and a nested double-cluster pattern. This finding suggests two independent recruitment and/or seed dispersal mechanisms. Saplings were several times as abundant as recruits and may accumulate several recruit generations. Recruits were only weakly associated with adults and occupied about half of the large-scale clusters, but saplings almost all. This is consistent with recruitment limitation. For approximately 70% (95%) of all juveniles the nearest adult was less than 26 m away (53 m), suggesting a dispersal limitation that may also be related to the critical large-scale clustering. Our example illustrates the manner in which the use of a specific and complex null hypothesis of spatial structure in point pattern analysis can help us better understand the biology of a species and generate specific hypotheses to be further investigated in the field.     

Quantifying the importance of local niche-based and stochastic processes to tropical tree community assembly

Although niche-based and stochastic processes, including dispersal limitation and demographic stochasticity, can each contribute to community assembly, it is difficult to quantify the relative importance of each process in natural vegetation. Here, we extend Shipley's maxent model (Community Assembly by Trait Selection, CATS) for the prediction of relative abundances to incorporate both trait-based filtering and dispersal limitation from the larger landscape and develop a statistical decomposition of the proportions of the total information content of relative abundances in local communities that are attributable to trait-based filtering, dispersal limitation, and demographic stochasticity. We apply the method to tree communities in a mature, species-rich, tropical forest in French Guiana at 1-, 0.25- and 0.04-ha scales. Trait data consisted of species' means of 17 functional traits measured over both the entire meta-community and separately in each of nine 1-ha plots. Trait means calculated separately for each site always gave better predictions. There was clear evidence of trait-based filtering at all spatial scales. Trait-based filtering was the most important process at the 1-ha scale (34%), whereas demographic stochasticity was the most important at smaller scales (37-53%). Dispersal limitation from the meta-community was less important and approximately constant across scales (-9%), and there was also an unresolved association between site-specific traits and meta-community relative abundances. Our method allows one to quantify the relative importance of local niche-based and meta-community processes and demographic stochasticity during community assembly across spatial and temporal scales.     

Establishment, persistence, and introgression of entomopathogenic nematodes in a forest ecosystem.

Entomopathogenic nematodes (EPN) are currently marketed worldwide for use in inundative biological control, where the applied natural enemy population (rather than its offspring) is expected to reduce insect numbers. Unlike classical biological control, in inundative control natural enemy establishment is not crucial in order to achieve pest suppression. Field trials in Irish forestry provided the opportunity to test predictions regarding the establishment of two exotic (Steinernema carpocapsae and Heterorhabditis megidis) and two indigenous (Steinernema feltiae and Heterorhabditis downesi) species. Nematodes were inundatively applied to pine stumps to control populations of pine weevil, Hylobius abietis, on three clearcut sites, and their persistence and spread monitored for up to five years. All species were recovered three years after application but only S. feltiae was recovered in years 4 and 5. Limited horizontal dispersal to 20 cm (but not 100 cm) was observed, but the majority of nematodes were recovered close to the area of application. Steinernema feltiae was also recovered from nearby stumps to which it had not been applied, indicating possible phoretic dispersal by weevils or other stump-associated fauna. EPN were not recovered from stumps outside the treated area, suggesting that such dispersal is quite localized. Two strains of S. feltiae (Irish and exotic) were applied. Amplified fragment length polymorphism (AFLP) analysis on 11 populations isolated from soil four years later showed that all had a much closer affinity to the applied Irish strain, suggesting persistence of this genotype and extinction of the exotic one. Some strains were clustered close together, and this is interpreted in the light of possible population genetic scenarios. The findings from the field study confirm predictions based on background knowledge of the species and demonstrate the importance of medium-term studies, as a 3-year study would have overestimated the risk of establishment of exotic species. Short-term persistence and spread of S. carpocapsae, S. feltiae, and H. downesi was also studied in pine forest mesocosms. Similar trends to field results, such as limited horizontal dispersal, even vertical distribution, and more abundant recovery of S. feltiae than of other species, point to the utility of mesocosm studies as a predictive tool.     

The interplay between environmental conditions and allee effects during the recovery of stressed zooplankton communities

Many important ecological phenomena depend on the success or failure of small introduced populations. Several factors are thought to influence the fate of small populations, including resource and habitat availability, dispersal levels, interspecific interactions, mate limitation, and demographic stochasticity. Recent field studies suggest that Allee effects resulting from mate limitation can prevent the reestablishment of sexual zooplankton species following a disturbance. In this study, we explore the interplay between Allee effects and local environmental conditions in determining the population growth and establishment of two acid-sensitive zooplankton species that have been impacted by regional anthropogenic acidification. We conducted a factorial design field experiment to test the impact of pH and initial organism densities on the per capita population growth (r) of the sexual copepod Epischura lacustris and the seasonally parthenogenetic cladoceran Daphnia mendotae. In addition, we conducted computer simulations using r values obtained from our experiments to determine the probability of extinction for small populations of acid-sensitive colonists that are in the process of colonizing recovering lakes. The results of our field experiment demonstrated that local environmental conditions can moderate the impacts of Allee effects for E. lacustris: Populations introduced at low densities had a significantly lower r at pH 6 than at pH 7. In contrast, r did not differ between pH 6 and 7 environments when E. lacustris populations were introduced at high densities. D. mendotae was affected by pH levels, but not by initial organism densities. Results from our population growth simulations indicated that E. lacustris populations introduced at low densities to pH 6 conditions had a higher probability of extinction than those introduced at low densities to a pH 7 environment. Our study indicates that environmental conditions and mate limitation can interact to determine the fate of small populations of sexually reproducing zooplankton species. If a more rapid recovery of acid-damaged zooplankton communities is desired, augmentation of dispersal levels may be needed during the early phases of pH recovery in order to increase the probability of establishment for mate-limited zooplankton species.     

Cryptic species of Clavelina (Ascidiacea) in two different habitats: harbours and rocky littoral zones in the northwestern Mediterranean

Marinas and harbours provide ideal sites for the study of population genetics of marine invertebrates with restricted dispersal capabilities. They combine a confinement effect, particular ecological conditions (pollution, turbidity), and the possibility of high gene flow through ship-borne propagules, which greatly increases the natural dispersal capability of sexual and asexual propagules in many species with short-lived larvae. We studied the genetic structure of populations of the ascidian Clavelina lepadiformis living inside and outside harbours in the north-western Mediterranean. A 500-bp segment of the cytochrome c oxidase subunit I (COI) mitochondrial gene was sequenced in three populations from inside harbours (interior form) and in three populations from the rocky littoral (exterior form). Two congeneric Mediterranean species, Clavelina sp. and C. dellavallei, were used for comparison. We found that the interior and exterior forms of C. lepadiformis belong to two distinct clades, with a genetic divergence of 5%. Gene-flow values among these forms were insignificant. The lack of gene flow and the genetic divergence suggest that the interior and exterior forms of C. lepadiformis are in fact cryptic species rather than differentiated populations of the same species. Levels of gene flow were higher among interior habitats than among exterior habitats, a pattern likely maintained by genetic exchange through ships. We discuss the possible origins of the present-day distribution of these cryptic species. We contend that the study of species living both inside and outside these particular habitats will reveal more instances of genetic discontinuities allowing local adaptations.