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Pollen-limited reproduction in blue oak: Implications for wind pollination in fragmented populations

Human activities are fragmenting forests and woodlands worldwide, but the impact of reduced tree population densities on pollen transfer in wind-pollinated trees is poorly understood. In a 4-year study, we evaluated relationships among stand density, pollen availability, and seed production in a thinned and fragmented population of blue oak (Quercus douglasii). Geographic coordinates were established and flowering interval determined for 100 contiguous trees. The number of neighboring trees within 60 m that released pollen during each tree's flowering period was calculated and relationships with acorn production explored using multiple regression. We evaluated the effects of female flower production, average temperature, and relative humidity during the pollination period, and number of pollen-producing neighbors on individual trees' acorn production. All factors except temperature were significant in at least one of the years of our study, but the combination of factors influencing acorn production varied among years. In 1996, a year of large acorn crop size, acorn production was significantly positively associated with number of neighboring pollen producers and density of female flowers. In 1997, 1998, and 1999, many trees produced few or no acorns, and significant associations between number of pollen-producing neighbors and acorn production were only apparent among moderately to highly reproductive trees. Acorn production by these reproductive trees in 1997 was significantly positively associated with number of neighboring pollen producers and significantly negatively associated with average relative humidity during the pollination period. In 1998, no analysis was possible, because too few trees produced a moderate to large acorn crop. Only density of female flowers was significantly associated with acorn production of moderately to highly reproductive trees in 1999. The effect of spatial scale was also investigated by conducting analyses with pollen producers counted in radii ranging from 30 m to 80 m. The association between number of pollen-producing neighbors and acorn production was strongest when neighborhood sizes of 60 m or larger were considered. Our results suggest that fragmentation and thinning of blue oak woodlands may reduce pollen availability and limit reproduction in this wind-pollinated species.

Oecologia

Linking biodiversity to ecosystem function: Implications for conservation ecology

We evaluate the empirical and theoretical support for the hypothesis that a large proportion of native species richness is required to maximize ecosystem stability and sustain function. This assessment is important for conservation strategies because sustenance of ecosystem functions has been used as an argument for the conservation of species. If ecosystem functions are sustained at relatively low species richness, then arguing for the conservation of ecosystem function, no matter how important in its own right, does not strongly argue for the conservation of species. Additionally, for this to be a strong conservation argument the link between species diversity and ecosystem functions of value to the human community must be clear. We review the empirical literature to quantify the support for two hypotheses: (1) species richness is positively correlated with ecosystem function, and (2) ecosystem functions do not saturate at low species richness relative to the observed or experimental diversity. Few empirical studies demonstrate improved function at high levels of species richness. Second, we analyze recent theoretical models in order to estimate the level of species richness required to maintain ecosystem function. Again we find that, within a single trophic level, most mathematical models predict saturation of ecosystem function at a low proportion of local species richness. We also analyze a theoretical model linking species number to ecosystem stability. This model predicts that species richness beyond the first few species does not typically increase ecosystem stability. One reason that high species richness may not contribute significantly to function or stability is that most communities are characterized by strong dominance such that a few species provide the vast majority of the community biomass. Rapid turnover of species may rescue the concept that diversity leads to maximum function and stability. The role of turnover in ecosystem function and stability has not been investigated. Despite the recent rush to embrace the linkage between biodiversity and ecosystem function, we find little support for the hypothesis that there is a strong dependence of ecosystem function on the full complement of diversity within sites. Given this observation, the conservation community should take a cautious view of endorsing this linkage as a model to promote conservation goals.

Oecologia

The influence of vines on an oligohaline marsh community: Results of a removal and fertilization study

The effects of competitive suppression by vines on the non-vine plant community have received little attention in temperate habitats. This study investigated the impact vines have on their herbaceous hosts in a wetland community at two soil fertility levels. Plots in an oligohaline marsh were treated in a 2 x 2 factorial design with vine removal and fertilization over two growing seasons. There was no significant interaction between removal and fertilization treatments on any of the measured variables. Vine removal initially caused an increase in light penetration through the canopy, but by the end of the study, plots with vines removed had less light due to a 25% increase in biomass by the plants released from competition with vines. For plots with vines removed, species richness was higher during a brief period in the spring of the second year, but by the end of the study, richness in removal plots decreased relative to controls. Fertilization caused a 40% increase in biomass overall, although only two species, Sagittaria lancifolia L. and Polygonum punctatum Ell., showed dramatic increases. Despite fertilization causing a 40% decrease in light penetration to the ground, no change in species richness was observed. Overall, these results show that vine cover in this wetland suppresses non-vine species and reduces community biomass. Removal of vines increased biomass of non-vine dominants but resulted in only an ephemeral change in species richness. Fertilization did not increase the effects of vines on the non-vine community.

Oecologia

Study design and interpretation of mammalian carnivore density estimates

Ecological theory and wildlife management often depend on reliable comparison and interpretation of population density estimates. A synthesis of 1,772 mammalian carnivore population estimates (713 unique to reference, species, site, and size of study area) from 74 species revealed global patterns among aspects of study and interpretive design that undermine the reliability and usefulness of density comparisons. The spatial extent of the study area could explain most of the variation in density, probably because study areas are typically delineated around population clusters. We related the scale-defined density estimates (regression residuals) to 28 other variables measured from the published literature, but none provided convincing biological explanation of the variation in density. Many aspects of study and interpretive design were possibly ill-suited to identifying the factor(s) influencing density. Study attributes and findings were reported inconsistently, and were subject to ideological motivations. Descriptions of vegetation were most difficult to relate to density. More intensive sampling and estimation methods produced above-average density estimates, but the differences were slight and the evidence lacking for concluding whether these more intensive methods were also more accurate. The first underlying factor extracted from principle-components analysis described the growing recognition of population declines and range reductions among large-bodied carnivores, which has also influenced study design. Another factor described an increasing trend for density to be compared and extrapolated to larger areas, but without adjusting for the effect of scale. To understand the factors influencing carnivore distribution and abundance, sampling and reporting methods (e.g., site description with maps) will need to represent the available pool of species, locations, and ecological conditions at larger-than-conventional spatial and temporal scales.

Oecologia

The effect of canopy cover and seasonal change on host plant quality for the endangered Karner blue butterfly (Lycaeidesmelissasamuelis)

Larvae of the Karner blue butterfly, Lycaeidesmelissasamuelis , feed solely on wild lupine, Lupinusperennis , from the emergence to summer senescence of the plant. Wild lupine is most abundant in open areas but Karner blue females oviposit more frequently on lupines growing in moderate shade. Can differences in lupine quality between open and shaded areas help explain this disparity in resource use? Furthermore, many lupines are senescent before the second larval brood completes development. How does lupine senescence affect larval growth? We addressed these questions by measuring growth rates of larvae fed lupines of different phenological stages and lupines growing under different shade conditions. The habitat conditions under which lupines grew and plant phenological stage did not generally affect final larval or pupal weight but did significantly affect duration of the larval period. Duration was shortest for larvae fed leaves from flowering lupines and was negatively correlated with leaf nitrogen concentration. Ovipositing in areas of moderate shade should increase second-brood larval exposure to flowering lupines. In addition, larval growth was significantly faster on shade-grown lupines that were in seed than on similar sun-grown lupines. These are possible advantages of the higher-than-expected oviposition rate on shade-grown lupines. Given the canopy-related trade-off between lupine abundance and quality, maintenance of canopy heterogeneity is an important conservation management goal. Larvae were also fed leaves growing in poor soil conditions and leaves with mildew infection. These and other feeding treatments that we anticipated would inhibit larval growth often did not. In particular, ant-tended larvae exhibited the highest weight gain per amount of lupine eaten and a relatively fast growth rate. This represents an advantage of ant tending to Karner blue larvae.

Indiana

C4 photosynthetic modifications in the evolutionary transition from land to water in aquatic grasses

Cladistic analysis supports the conclusion that the Orcuttieae tribe of C 4 grasses reflect evolution from a terrestrial ancestry into seasonal pools. All nine species in the tribe exhibit adaptations to the aquatic environment, evident in the structural characteristics of the juvenile foliage, which persist submerged for 1–3 months prior to metamorphosis to the terrestrial foliage. Aquatic leaves of the least derived or basal genus Neostapfia have few morphological and anatomical characteristics specialized to the aquatic environment and have retained full expression of the C 4 pathway, including Kranz anatomy. Orcuttia species have many derived characteristics and are more specialized to the aquatic environment. These latter species germinate earlier in the season and persist in the submerged stage longer than Neostapfia and evidence from the literature indicates length of submergence is positively correlated with fitness components. Aquatic leaves of Orcuttia species lack Kranz or PCR bundle sheath anatomy, yet 14 C-pulse chase studies indicate >95% malate + aspartate as the initial products of photosynthesis and these products turn over rapidly to phosphorylated sugars, indicating a tight coupling of the C 4 and C 3 cycles. Presence of the C 4 pathway is further supported by enzymological data. Contemporary dogma that Kranz anatomy is a sine qua non for operation of the C 4 pathway is contradicted by the patterns in Orcuttia ; however, it is unknown whether the pathway acts as a CO 2 concentrating mechanism in these aquatic plants.

California

Effects of flooding, salinity and herbivory on coastal plant communities, Louisiana, United States

Flooding and salinity stress are predicted to increase in coastal Louisiana as relative sea level rise (RSLR) continues in the Gulf of Mexico region. Although wetland plant species are adapted to these stressors, questions persist as to how marshes may respond to changed abiotic variables caused by RSLR, and how herbivory by native and non-native mammals may affect this response. The effects of altered flooding and salinity on coastal marsh communities were examined in two field experiments that simultaneously manipulated herbivore pressure. Marsh sods subjected to increased or decreased flooding (by lowering or raising sods, respectively), and increased or decreased salinity (by reciprocally transplanting sods between a brackish and fresh marsh), were monitored inside and outside mammalian herbivore exclosures for three growing seasons. Increased flooding stress reduced species numbers and biomass; alleviating flooding stress did not significantly alter species numbers while community biomass increased. Increased salinity reduced species numbers and biomass, more so if herbivores were present. Decreasing salinity had an unexpected effect: herbivores selectively consumed plants transplanted from the higher-salinity site. In plots protected from herbivory, decreased salinity had little effect on species numbers or biomass, but community composition changed. Overall, herbivore pressure further reduced species richness and biomass under conditions of increased flooding and increased salinity, supporting other findings that coastal marsh species can tolerate increasingly stressful conditions unless another factor, e.g., herbivory, is also present. Also, species dropped out of more stressful treatments much faster than they were added when stresses were alleviated, likely due to restrictions on dispersal. The rate at which plant communities will shift as a result of changed abiotic variables will determine if marshes remain viable when subjected to RSLR.

Oecologia

Role of brown bears ( Ursus arctos ) in the flow of marine nitrogen into a terrestrial ecosystem

We quantified the amount, spatial distribution, and importance of salmon ( Oncorhynchus spp.)-derived nitrogen (N) by brown bears ( Ursus arctos ) on the Kenai Peninsula, Alaska. We tested and confirmed the hypothesis that the stable isotope signature (&delta; 15 N) of N in foliage of white spruce ( Picea glauca ) was inversely proportional to the distance from salmon-spawning streams ( r =&ndash;0.99 and P <0.05 in two separate watersheds). Locations of radio-collared brown bears, relative to their distance from a stream, were highly correlated with &delta; 15 N depletion of foliage across the same gradient ( r =&ndash;0.98 and &ndash;0.96 and P <0.05 in the same two separate watersheds). Mean rates of redistribution of salmon-derived N by adult female brown bears were 37.2&plusmn;2.9 kg/year per bear (range 23.1&ndash;56.3), of which 96% (35.7&plusmn;2.7 kg/year per bear) was excreted in urine, 3% (1.1&plusmn;0.1 kg/year per bear) was excreted in feces, and <1% (0.3&plusmn; 0.1 kg/year per bear) was retained in the body. On an area basis, salmon-N redistribution rates were as high as 5.1&plusmn;0.7 mg/m 2 per year per bear within 500 m of the stream but dropped off greatly with increasing distance. We estimated that 15.5&ndash;17.8% of the total N in spruce foliage within 500 m of the stream was derived from salmon. Of that, bears had distributed 83&ndash;84%. Thus, brown bears can be an important vector of salmon-derived N into riparian ecosystems, but their effects are highly variable spatially and a function of bear density.

Alaska

The effect of community composition on persistence of prey with their predators in an assemblage of pond-breeding amphibians

We examined whether the species composition of a community influences the persistence of larval Ambystoma maculatum in assemblages composed of two larger intraguild predators (A. opacum and A. jeffersonianum) and an alternative prey species (tadpoles of Rana sylvatica). We predicted a priori that A. maculatum would have higher survival in more diverse communities containing alternative species of prey and top predators (A. opacum), the latter of which may lower the abundance of intermediate predators (A. jeffersonianum) via intraguild predation. In a factorial experiment, we manipulated the presence of larval A. opacum, A. jeffersonianum, and R. sylvatica in replicated artificial ponds containing larval A. maculatum. The presence of all three species significantly depressed biomass production in A. maculatum: biomass was highest in ponds lacking the other species and was zero in ponds initially containing all four species. Tadpoles severely reduced the growth of filamentous algae in the ponds. This, in turn, may have affected the abundance of some herbivorous prey of larval salamanders, although this possibility was not tested. The presence of congeneric predators severely restricted the presence in the water column of larval A. maculatum, which otherwise exhibited significant diel patterns of activity in the absence of predators. Together, the presence of tadpoles and a predator-mediated reduction in activity patterns may have limited foraging opportunities for A. maculatum, thus exacerbating the direct impact of predation on survival in this species. These results suggest that diverse assemblages consisting of these particular species may actually inhibit, rather than promote, inclusion of A. maculatum in some communities of pond-breeding amphibians.

Oecologia

Association of amphibians with attenuation of ultraviolet-b radiation in montane ponds

Ambient ultraviolet-b (UV-B) radiation (280–320 nm) has increased at north-temperate latitudes in the last two decades. UV-B can be detrimental to amphibians, and amphibians have shown declines in some areas during this same period. We documented the distribution of amphibians and salmonids in 42 remote, subalpine and alpine ponds in Olympic National Park, Washington, United States. We inferred relative exposure of amphibian habitats to UV-B by estimating the transmission of 305- and 320-nm radiation in pond water. We found breeding Ambystoma gracile , A. macrodactylum and Rana cascadae at 33%, 31%, and 45% of the study sites, respectively. Most R. cascadae bred in fishless shallow ponds with relatively low transmission of UV-B. The relationship with UV-B exposure remained marginally significant even after the presence of fish was included in the model. At 50 cm water depth, there was a 55% reduction in incident 305-nm radiation at sites where breeding populations of R. cascadae were detected compared to other sites. We did not detect associations between UV-B transmission and A. gracile or A. macrodactylum . Our field surveys do not provide evidence for decline of R. cascadae in Olympic National Park as has been documented in Northern California, but are consistent with the hypothesis that the spatial distribution of R. cascadae breeding sites is influenced by exposure to UV-B. Substrate or pond depth could also be related to the distribution of R. cascadae in Olympic National Park.

Oecologia

The influence of climatic variability on local population dynamics of Cercidium microphyllum (foothill paloverde)

This study investigated correlations among climatic variability, population age structure, and seedling survival of a dominant Sonoran Desert tree, Cercidium microphyllum (foothill paloverde), at Tucson, Arizona, USA. A major goal was to determine whether wet years promote seedling establishment and thereby determine population structure. Plant age was estimated from basal circumference for a sample of 980 living and dead trees in twelve 0.5-ha plots. Ages ranged from 1 to 181 years. Age frequency distribution showed that the population is in decline. Most (51.2%) of the 814 living trees were 40–80 years old; only 6.5% were younger than 20 years. The average age of the 166 dead trees was 78 years. Fifty-nine percent of dead trees were aged 60–100 years. Survival of newly emerged seedlings was monitored for 7 years in a 557-m 2 permanent plot. Mean survival in the 1st year of life was 1.7%. Only 2 of 1,008 seedlings lived longer than 1 year. Length of survival was not correlated with rainfall. Residual regeneration, an index of the difference between predicted and observed cohort size, showed that regeneration was high during the first half of the twentieth century and poor after the mid-1950s. Trends in regeneration did not reflect interannual variation in seasonal temperature or rain before 1950, that is, in the years before urban warming. Taken together, the seedling study and the regeneration analysis suggest that local population dynamics reflect biotic factors to such an extent that population age structure might not always be a reliable clue to past climatic influences.

Oecologia

Monitoring intensity and patterns of off-highway vehicle (OHV) use in remote areas of the western USA

The continued growth of off-highway vehicle (OHV) activities – demonstrated by the dramatic increase in OHV sales, number of users, and areas experiencing OHV use – has elevated concerns about their ecological effects, the impacts on wildlife, and the sustainability of OHV use on secondary and tertiary road networks. Conflicts between visitors and wildlife are raising concerns about system resiliency and sustainable management. In order to quantify the spatial and temporal impacts of OHV use it is imperative to know about the timing and patterns of vehicle use. This study tested and used multiple vehicle-counter types to study vehicular OHV use patterns and volume throughout a mountainous road network in western Colorado. OHV counts were analyzed by time of day, day of week, season, and year. While daily use peaked within a two to three hour range for all sites, the overall volume of use varied among sites on an annual basis. The data also showed that there are at least two distinct patterns of OHV use: one dominated by a majority of use on weekends, and the other with continuous use throughout the week. This project provided important, but rarely captured, metrics about patterns of OHV use in a remote, mountainous region of Colorado. The techniques described here can provide land managers with a quantitative evaluation of OHV use across the landscape, an essential foundation for travel management planning. They also provide researchers with robust tools to further investigate the impacts of OHV use.

Oecologia Australis

Comparison of neotropical winter bird populations in isolated patches versus extensive forest

Wintering birds were captured with mist nets at 12 pairs of forested sites in the New World tropics in 1984 and 1985 to compare populations in small isolated woodlands (generally 5-50 ha) with those in extensive forests (> 1,000 ha). Net-hours of effort were similar in large and small sites, as were total birds captured and banded, but species composition was very different. Members of the Todidae, Dendrocolaptidae, Formicariidae and Thraupinae were significantly more common in extensive forest than in small isolated tracts, indicating that these birds are especially vulnerable to effects of forest fragmentation. However, in winter many species of North American migrants, even species that are restricted to extensive forest during the breeding season, were just as common in small forest fragments as in extensive forest. A high percentage of the North American migrants banded in January 1984 (40 to 50 % for some species) were recaptured in 1985.

Acta Oecologica Oecologia Generalis

Improving atlas methodology

We are studying a sample of Maryland (2 %) and New Hampshire (4 %) Atlas blocks and a small sample in Maine. These three States used different sampling methods and block sizes. We compare sampling techniques, roadside with off-road coverage, our coverage with that of the volunteers, and different methods of quantifying Atlas results. The 7 1/2' (12-km) blocks used in the Maine Atlas are satisfactory for coarse mapping, but are too large to enable changes to be detected in the future. Most states are subdividing the standard 7 1/2' maps into six 5-km blocks. The random 1/6 sample of 5-km blocks used in New Hampshire, Vermont (published 1985), and many other states has the advantage of permitting detection of some changes in the future, but the disadvantage of leaving important habitats unsampled. The Maryland system of atlasing all 1,200 5-km blocks and covering one out of each six by quarterblocks (2 1/2-km) is far superior if enough observers can be found. A good compromise, not yet attempted, would be to Atlas a 1/6 random sample of 5-km blocks and also one other carefully selected (non-random) block on the same 7 1/2' map--the block that would include the best sample of habitats or elevations not in the random block. In our sample the second block raised the percentage of birds found from 86% of the birds recorded in the 7 1/2' quadrangle to 93%. It was helpful to list the expected species in each block and to revise this list annually. We estimate that 90-100 species could be found with intensive effort in most Maryland blocks; perhaps 95-105 in New Hampshire. It was also helpful to know which species were under-sampled so we could make a special effort to search for these. A total of 75 species per block (or 75% of the expected species in blocks with very restricted habitat diversity) is considered a practical and adequate goal in these States. When fewer than 60 species are found per block, a high proportion of the rarer species are missed, as well as some of the common ones. Similarity indices based on fewer than 60 species per block reflect coverage rather than habitat affinities. Atlas blocks that are ecologically similar should have similarity indices (S) of at least 0.80 to be considered adequately covered. S = 2C/(A + B), where C is the number of species in common and A and B are species totals for each of the two blocks being compared. A series of 15 13-minutes roadside stops yielded more species than 15 off-road stops, but off-road stops always had some species not detected at the roadside stops. A series of timed stops is an excellent way to map relative abundance if the stops are standardized with respect to time of day and weather, and the counts are made by observers of comparable ability. Efforts to estimate Atlas block populations in powers of 10 (as in the French Atlas) have not gained acceptance in U.S.A. Most observers feel unqualified to make estimates. An efficient way to Atlas a block is to make at least 3 early morning visits to 15 or more specific stops. Arrive in the block early enough to check for nocturnal species on at least two days; and after completing the specific stops, search the block for other species and for confirmations.

Acta Oecologica Oecologia Generalis

Temporal variability of local abundance, sex ratio and activity in the Sardinian chalk hill blue butterfly

When capturing and marking of individuals is possible, the application of newly developed capture-recapture models can remove several sources of bias in the estimation of population parameters such as local abundance and sex ratio. For example, observation of distorted sex ratios in counts or captures can reflect either different abundances of the sexes or different sex-specific capture probabilities, and capture-recapture models can help distinguish between these two possibilities. Robust design models and a model selection procedure based on information-theoretic methods were applied to study the local population structure of the endemic Sardinian chalk hill blue butterfly, Polyommatus coridon gennargenti. Seasonal variations of abundance, plus daily and weather-related variations of active populations of males and females were investigated. Evidence was found of protandry and male pioneering of the breeding space. Temporary emigration probability, which describes the proportion of the population not exposed to capture (e.g. absent from the study area) during the sampling process, was estimated, differed between sexes, and was related to temperature, a factor known to influence animal activity. The correlation between temporary emigration and average daily temperature suggested interpreting temporary emigration as inactivity of animals. Robust design models were used successfully to provide a detailed description of the population structure and activity in this butterfly and are recommended for studies of local abundance and animal activity in the field.

Oecologia