Seed banks of grazed and ungrazed Baltic seashore meadows

Abstract. The seed banks of three grazed and three ungrazed seashore meadows were studied on the west coast of Finland. 8486 seedlings (mean 13 669 seedlings/m²) germinated from cold-treated samples (n = 343; depth = 10 cm). Most seedlings and species were monocots and perennials. The seed bank flora included 54 dicots vs. 28 monocots and 66 perennials vs. 16 annuals. The most abundant species were Juncus gerardii, Schoenoplectus tabernaemontani, Eleocharis uniglumis, Agrostis stolonifera, Juncus bufonius and Carex nigra, which made up 73% of the seed bank. Numbers of species and seedlings differed between elevation classes. Species richness was highest in elevation class 50–70 cm. The highest seed density occurred in class 20–50 cm. A model for size and species composition of the seed bank in relation to elevation is presented. The seed bank was larger and richer in species in the ungrazed than in the grazed sites, but not so in the upper elevations and closest to the open sea. Grazing reduced the size of the seed bank of Agrostis stolonifera, A. capillaris, Calamagrostis stricta, Elymus repens, Juncus bufonius, Limosella aquatica and Schoenoplectus tabernaemontani, but increased that of J. gerardii. 32 species germinated only from ungrazed samples and 11 species only from grazed ones. Multivariate classification resulted in nine sample groups. The ordination scatter was best explained by the flooding stress variables, elevation, the distance from the water line and the number of helophyte species in samples. 75 species were found both in the seed bank and in the vegetation, but there was a significant lack of resemblance (in the Mantel test) due to over-representation of some species. Eight species occurring only in the seed bank were mainly annuals or biennials (63%); those occurring only in the established vegetation (86 species) were mainly perennials (86%).     

Nonequilibrium dynamics of sedge meadows grazed by cattle in southern Wisconsin

Equilibrium theory predicts that after disturbance, ecosystemseventually regain the structural and functional properties characteristic oftheir predisturbance condition. This study tested this idea by examining theeffects of cattle grazing and exclusion on the long-term structuralcharacteristics of sedge meadows in southern Wisconsin. To compare structuralchanges in mean percentage cover and height, repeated measures analysis wasconducted on two sedge meadows over a twenty year period from 1977 to 1997. Onesedge meadow was recovering from cattle grazing (cattle excluded in 1973) andthe other was a reference area (nearly undisturbed). Both of these study siteschanged structurally from 1977 to 1997, supporting non-equilibrium theory.Additional observations were made in a heavily and lightly grazed sedge meadowthat were surveyed in 1977. As based on the positions of subunits in anordination graph produced using Non-Metric Multidimensional Scaling (NMS), therecovery sedge meadow became less structurally similar to the grazed and moresimilar to the reference site over the 20 year study. However, from theperspective of mean maximum height in another NMS analysis, the recovery sedgemeadow became less similar to the reference site over time likely because by1997, a shrub carr of Cornus sericea had developed in therecovery sedge meadow that had been dominated by graminoids and forbs in 1977(mean maximum height: 1977 vs. 1997; 0 vs. 47 cm). Seedlings ofCornus sericea were invading the grazed sedge meadows andin the recovery sedge meadow (cattle excluded 4 years earlier) in 1977. A shrubcarr did not develop in the reference sedge meadow. Changes in the referencesite were relatively minor over this time interval; certain species eitherincreased or decreased in dominance, e.g., Carex strictaincreased in cover (1977 vs. 1997, 20 and 28 mean percentage (%) cover,respectively). A few short-term species of the recovery sedge meadow followedthe tenets of equilibrium theory. These became less common or disappeared4–9 years after cattle exclusion including Asterlanceolatus, Calamagrostis canadensis,Poa compressa, Solidago altissima andVerbena hastata. Some of these species were eaten andlikely spread by the cattle. This study suggests that the progression of sedgemeadow to shrub carr may not be an inevitable outcome of succession but insteadcan be a consequence of past cattle grazing history. Also, because the recoveryand the reference sedge both changed structurally over time, the tenets ofnon-equilibrium theory were supported by this study.     

Nest predation and semicolonial breeding in Linnets Carduelis cannabina

Capsule Nest predation in Linnets in Denmark was caused mainly by Hooded Crows Corvus corone cornix and Magpies Pica pica with no experimental evidence of antipredator aggression.

Aims To investigate the influence of nest clumping on the risk of nest predation in Linnets, a species that often breeds semi-colonially.

Methods The predation rates on clumped and randomly distributed nests were compared by the use of artificial nest experiments. Additionally, the behaviour of Linnets towards a potential nest predator was studied by presenting a mounted Magpie near the nest of breeding Linnets.

Results The artificial nest experiments showed that semi-colonial breeding was not likely to increase the risk of nest predation, and an experiment with the mounted Magpie showed that Linnets did not exhibit anti-predator aggression.

Conclusion Semi-colonial breeding, resulting in increased local nest density, did not appear to influence the risk of nest predation in Linnets. We suggest that semi-colonial breeding in Linnets is a means of pooling information on food sources.     

Nest predation and the species diversity of birds

In a recent series of papers, Thomas E. Martin has suggested that selective, density-dependent predation on nests may promote ecological diversification of birds and constrain the number of coexisting species in a community. In an experimental study, Martin has demonstrated density-dependent predation on artificial nests constructed and placed so as to mimic natural ones. Comparisons between habitats indicate that availability of suitable nest sites may influence community diversity more strongly than does diversity of foraging niches. Although his case is circumstantial at this point, Martin argues compellingly for considering predation more seriously in studies of community ecology. Indeed, many patterns attributed to competition for food resources may arise, instead, from the partitioning of predator-escape space.     

Nest predation and the evolution of nestling begging calls

Begging by nestling birds can be conspicuous and loud. Such displays are thought to function in signalling nestling condition and securing parental care, but they also may inadvertently attract the attention of predators. We compared the structure of nestling begging calls to the risk of predation among 24 species of birds breeding in a forest community in central Arizona. After controlling for body size and phylogeny, we found that species subject to greater nest predation had calls with higher frequency (pitch) and lower amplitude (loudness) than species subject to lower rates of nest predation. As these acoustic features make it difficult for potential predators to pinpoint the source of a sound, our results suggest that an increased risk of predation has led to the evolution of begging calls that minimize locatability. The relationship between call structure and the risk of predation also supports the hypothesis that attracting predators is a direct cost of begging and that such costs can constrain any evolutionary escalation in the intensity of nestling begging.     

Changes in soil biochemical properties associated with Ligularia virgaurea spreading in grazed alpine meadows

Ligularia virgaurea, a toxic perennial weed, has become a dominant species in the heavily-grazed alpine meadows of the eastern Qinghai?CTibetan Plateau over recent decades. We investigated changes in soil biochemical properties associated with L. virgaurea spreading in grazed alpine meadows at three sites. Soil and root biomass samples were taken at depths of 0?C8?cm, 8?C16?cm and 16?C24?cm from patches where L. virgaurea was dominant and from areas between L. virgaurea patches, with only other native species. Across sites, root biomass, acid-extracted carbohydrate C, organic C and N mineralization, and microbial biomass C and N concentrations in the top 8-cm layer were significantly higher while nitrate N concentration was significantly lower in L. virgaurea patches than in areas between L. virgaurea patches. The increased activities of dehydrogenase, ???Cglucosidase, urease and phosphatase in the top 8-cm layer under L. virgaurea were associated with enhanced soil microbial biomass. Our results indicate a close association between changes in soil biochemical properties in the top 8-cm layer and the spread of L. virgaurea in grazed alpine meadows of the eastern Qinghai?CTibetan Plateau.     

Nest predation patterns in ground-nesting passerines on the Iberian Peninsula

The idea that ground-nesting birds have a high risk of predation is widely accepted This paper analyzes the nest predation patterns of ground-nesting passerines on the Iberian Peninsula The nest predation rates are higher m open land species x= 71%) than m countryside farmland (10%) or forest birds (29%) The latter species have no differences in daily survival between ground and above-ground nests The higher nest predation in ground-nesting open land species was similar m peninsular Spain and North America We conclude that it is incorrect to generalize that all ground nesters have high nest predation rates, and we discuss the possible link of the high nest predation rates of open land birds to the decline observed in their population trends in Spain     

Optimal management of the rare Gladiolus imbricatus in Estonian coastal meadows indicated by its population structure

Questions: What is the best grassland management regime for the threatened plant species Gladiolus imbricatus; is the stage structure of local populations a feasible indicator of the effect of changed management. Location: Coastal meadow system in southwestern Estonia. Methods : The effect of five management regimes was studied in a long‐term (three‐year) field experiment: (1) mowing in late July, (2) grazing by cattle, (3) grazing by sheep, (4) sheep grazing during the first year and mowing during subsequent years, (5) no management (control). Results: The population density increased significantly in response to the mowing treatment and to the mowing after sheep grazing treatment. The proportion of grazed plant individuals was higher in the sheep‐grazed than in the cattle‐grazed treatment. Generative and vegetative adult individuals of G. imbricatus were significantly more damaged by cattle herbivory than juveniles. All management regimes shifted the population structure towards a dynamic state where juvenile stages dominate, while the not managed control retained a regressive population structure. Conclusions: Population stage structure was a useful indicator of different management conditions, even in the case where population density did not differ. As indicated by population stage structure, the best management regime for G. imbricatus was either mowing in late July only, or alternation of grazing and mowing in different years.     

Nest success and cause-specific nest failure of grassland passerines breeding in prairie grazed by livestock

Livestock grazing is a widespread source of habitat modification, and may affect populations of ground-nesting grassland birds by influencing rates of nest failure. Nesting attempts can fail for various reasons, and determining risk of failure from specific causes associated with livestock grazing would enhance development of range management practices in areas managed for threatened grassland bird populations. Domestic livestock may influence nest failure by affecting vegetation structure, numerical or functional responses of predators, or directly by trampling nests. We hypothesized stocking rate may influence nest fate because it affects the amount and distribution of remaining vegetation, and the number of large herbivores to which nests are exposed. In 2007 and 2008, we evaluated nest fates for savannah sparrows and horned larks under 4 stocking rates experimentally applied in 40-ha paddocks in northeastern Oregon, USA. In addition to stocking rate, we evaluated variables such as vegetation structure and predator abundance and activity to help clarify mechanisms responsible for nest failure. We used a discrete competing risks framework to estimate daily probability of nest survival and failure from specific causes. These algorithms, implemented in a stand-alone graphical user interface-driven model, allow incorporation of covariates within an information theoretic approach to model inference. Although stocking rate influenced vegetation structure, the only nest failures related to stocking rate were from trampling. Trampling events were too infrequent to test for treatment effects (only 1 nest of each species), but occurred in the moderate and high stocking treatments. Additional variables were related to variation in nest failure from predation, but we found no support for the hypothesis that these causes of failure were affected by stocking rate. For savannah sparrows, daily probability of nest success (95% CI) = 0.97 (0.96–0.98); predation = 0.018 (0.008–0.028); and trampling = 0.001 (0.000–0.004). For horned larks, daily probability of nest success = 0.96 (0.95–0.98); predation = 0.029 (0.012–0.045); and trampling = 0.003 (0.000–0.007). Our results suggest grasslands managed for livestock may generally be compatible with grassland songbird conservation, at least for the species and stocking rates examined here. The most effective conservation strategies for improving nest success will involve decreasing risk of nest predation. However, we found no evidence that management of stocking rate is an effective method for doing so. © 2012 The Wildlife Society.     

Nest predation and habitat change interact to influence Siberian jay numbers

We studied Siberian jays, breeding in northern Sweden, to examine the potential for interactions between nest predation and reduced vegetation heterogeneity around nest sites to cause a decrease in jay numbers. Parent behaviour and nests are highly cryptic in the species. Our 12-year data showed, however, that nests had a probability of only 0.46 to be successful and produce at least one nestling. Nest predation was intense and a main cause of nest failure. All predators that could be identified were visually oriented hunters, mostly other corvids able to colonize taiga forest only close to human settlements. Consistent with the idea that predators used visual cues, nest predation increased with parental activity, which suggests that predators used parental provisioning trips to locate nests. Furthermore, a reduction in daily nest survival rates with decreasing amount of nesting cover was more pronounced in areas with high corvid activity as predicted when cover mediates the hunting efficiency of visual oriented predators. Declining temperatures interacted with the effects of habitat characteristics to further reduce daily nest survival rates suggesting that parents were not able to increase nest visitation rates to satisfy the higher energy demands of their nestlings without endangering the nest. Our results identify a mechanism through which predation and human-induced reduction in nesting cover on a larger scale may interact to cause a reduction in Siberian jay numbers larger than expected from habitat loss alone.     

Nest predation in Reed Buntings Emberiza schoeniclus: an experimental study

Predation is a major cause of nest failure in many bird species. High levels of nest loss may be a consequence of habitat fragmentation, leading to increased amounts of edge habitat. Yet the evidence for generally high nest predation rates along edges in fragmented landscapes is ambiguous. Using real nests of Reed Buntings Emberiza schoeniclus in which artificial Reed Bunting and real Japanese Quail Coturnix japonica eggs were placed, we experimentally tested for edge effects on nest predation in highly fragmented reed Phragmites sp. habitats in the Swiss lowlands. We also examined seasonal patterns of predation and the impacts of nest visits by observers. We found evidence for an edge effect at the water-sided reed edge, with nests located closer to the water being more likely to be predated than those further away. Predation probability increased from early to late season, suggesting that nest predation may be density dependent. Probability of nest predation was only weakly influenced by whether or not a nest was visited. Our results suggest that the intensive reed management currently applied in Swiss nature reserves may result in unnaturally high levels of nest losses in the Reed Bunting, because reed bands are not wide enough to allow nest placement at a safe distance from reed edges.     

Vegetation changes in semi-natural meadows after abandonment in coastal northern Finland

A research project on traditional cultural landscapes in coastal northern Finland provided the opportunity to compare presently managed meadow sites to a series of corresponding abandoned sites. Twenty-two grassland habitats. either grazed or mown, were studied to analyze differences between species compositions The data were classified with TWINSPAN and ordinated with Detrended Correspondence Analysis and Canonical Correspondence Analysis. The multivariate analysis recognized successfully habitats that are either presently used or differ considerably from the others by their moisture gradient although a few very coininon generalist species were present in almost every seral stage studied. The studied deviant habitats resemble most of the previously described vegetation types in northern Finland. Results suggest that drought can delay the invasion of tall broad-leaved species. The results also support the idea that abandoned meadows of corresponding moisture gradients begin to resemhle each other in later stages of succession. Multivariate methods can be used to determine the succession stages in grassland habitats and thus help to determine the urgency for restoration practices.     

Nest trampling and ground nesting birds: Quantifying temporal and spatial overlap between cattle activity and breeding redshank

Conservation grazing for breeding birds needs to balance the positive effects on vegetation structure and negative effects of nest trampling. In the UK, populations of Common redshank Tringa totanus breeding on saltmarshes declined by >50% between 1985 and 2011. These declines have been linked to changes in grazing management. The highest breeding densities of redshank on saltmarshes are found in lightly grazed areas. Conservation initiatives have encouraged low‐intensity grazing at <1 cattle/ha, but even these levels of grazing can result in high levels of nest trampling. If livestock distribution is not spatially or temporally homogenous but concentrated where and when redshank breed, rates of nest trampling may be much higher than expected based on livestock density alone. By GPS tracking cattle on saltmarshes and monitoring trampling of dummy nests, this study quantified (i) the spatial and temporal distribution of cattle in relation to the distribution of redshank nesting habitats and (ii) trampling rates of dummy nests. The distribution of livestock was highly variable depending on both time in the season and the saltmarsh under study, with cattle using between 3% and 42% of the saltmarsh extent and spending most their time on higher elevation habitat within 500 m of the sea wall, but moving further onto the saltmarsh as the season progressed. Breeding redshank also nest on these higher elevation zones, and this breeding coincides with the early period of grazing. Probability of nest trampling was correlated to livestock density and was up to six times higher in the areas where redshank breed. This overlap in both space and time of the habitat use of cattle and redshank means that the trampling probability of a nest can be much higher than would be expected based on standard measures of cattle density. Synthesis and applications: Because saltmarsh grazing is required to maintain a favorable vegetation structure for redshank breeding, grazing management should aim to keep livestock away from redshank nesting habitat between mid‐April and mid‐July when nests are active, through delaying the onset of grazing or introducing a rotational grazing system.     

Nest predation,food, and female age explain seasonal declines in clutch size

The selection pressures responsible for intra- and interspecific variation in avian clutch size have been debated for over half a century. Seasonal declines in clutch size represent one of the most robust patterns in avian systems, yet despite extensive research on the subject, the mechanisms underlying this pattern remain largely unknown. We tested a combination of experimental and observational predictions to evaluate ten hypotheses, representing both evolutionary and proximate mechanisms proposed to explain seasonal declines in avian clutch size. In line with long held life-history theory, we found strong support for both an evolved and proximate response to food availability for young. We also found evidence consistent with predictions that proximate level experiential nest predation influences seasonal declines in clutch size. Finally, older females appear to invest more in reproduction (initiate nests earlier and lay larger clutches) and choose better territories than younger females. Our results highlight the importance of examining multiple hypotheses in a theoretical context to elucidate the ecological processes underlying commonly observed patterns in life history.     

Refuge use by fish as a function of body length-related metabolic expenditure and predation risks

Theory predicts that staying in a refuge has benefits in terms of predator avoidance and costs in terms of lost feeding opportunities. In this study, we investigated how the relative importance of these costs and benefits changes with increasing body length. This is of particular interest in animals such as fish, which show continuous growth throughout their lives. Our results suggest that larger fish are subject to lower predation risks and are less affected by food deprivation than small fish, with fish decreasing their responses to food-deprivation treatments more strongly with increasing body length than to predation treatments. This may explain our observation that large fish emerged later from a refuge than small ones and spent shorter times outside the refuge. The key role of differential responses to food deprivation was further illustrated by the finding that the relative weight loss of individual fish was strongly correlated with a reduction in hiding time even in the absence of body length differences. The importance of inter-individual differences in metabolic rates for the decision-making behaviour of animals is discussed.     

Nest predation in forest birds: influence of predator type and predator's habitat quality

We used the introduction of a generalist nest predator, the red squirrel Tamiasciurus hudsonicus, and of a large herbivore, the Sitka black-tailed deer Odocoileus hemionus sitkensis, to the islands of Haida Gwaii (Queen Charlotte Islands, British Columbia, Canada) to study how predator assemblage and habitat quality and structure influenced nest predation in forest birds. We compared losses of natural nests to predators on islands with and without squirrels. We selected nine islands with or without squirrel or deer and used 506 artificial nests put on the ground or in shrubs to further analyse variation of nest predation with predator assemblage and habitat quality for the predators. For both natural and artificial nests predation risk was higher in presence of squirrels. But predation risk varied within island categories. In presence of squirrels it was highest in stands with mature conifers where it fluctuated from year to year, in response to fluctuations in squirrel abundance. Vegetation cover around the nest had little effect on nest predation by squirrels. Where squirrels were absent, nest predation concentrated near predictable food sources for corvids, the main native predators, and increased with decreasing vegetation cover, suggesting that removal of the vegetation by deer increased the risk of predation by native avian nest predators that use visual cues. Predation risk in these forests therefore varies in space and time with predator composition and with quality of the habitat from the predators' perspective. This temporal and spatial variation in predation risk should promote trade-offs in the response of birds to nest predation, rather than fine-tuned adaptations to a given predation pattern.     

Nest predation in hole-nesting birds in relation to habitat edge: an experiment

We experimentally tested the hypothesis that nest predation rate in hole-nesting birds, especially the common goldeneye Bucephala clangula , was related to the edge between aquatic and terrestrial habitats We also studied whether nest predation rate was related to habitat patch (lake) size In three study areas nest-boxes were erected in pairs one nest-box was erected at the shoreline and the other one inside the forest at a varying distance (range from 14 to 140 m) from the shoreline One chicken egg was placed in each nest-box (dummy nest) Overall predation rate on dummy nests varied from 34 6% to 52 6% depending on the study area The pine marten Martes manes was the primary nest predator m all study areas Daily predation rates did not differ between natural and dummy nests Predation rate on dummy nests was not related to the distance of the nest-box from the shoreline in any study area Predation rate was not related to lake size in two study areas but in one area predation rate on dummy nests was lower around largest lakes However, the result may be accidental and not because of lake size per se because the proportion of forest was exceptionally low in that part of the study area     

Predicting group size in primates: foraging costs and predation risks

We present a direct test of the long-standing hypothesis thatfood competition limits primate group size. Group size is acritical social variable because it constrains most other aspectsof social organization. We develop a simple population-specificindex of indirect feeding competition based on daily foragingcosts. This index explains nearly two-thirds of between-populationvariation in mean group sizes of mostly fruit-eating (but notof mostly leaf-eating) primates. Group size is also significantlyrelated to body size and terrestriality (or use of open country),which are suspected correlates of predation risk, although feedingcompetition remains an important predictor of group size evenwhen these correlates are controlled. Phylogeny also appearsto be important: the differences between observed mean populationgroup sizes and those predicted using ecological factors aremost positive for the Old World monkeys and most negative forthe lemuroids in our sample. The weak relationship between groupsize and feeding competition found for folivorous species maybe explained either by the energetic constraints of a leafydiet or by limits to group size imposed by infanticide as ahabitual male reproductive strategy.     

Nest predation reduces benefits to early clutch initiation in northern cardinals Cardinalis cardinalis

Life history theory and empirical studies suggest that early breeding confers higher reproductive success, but the extent to which this advantage can be generalized to human‐dominated systems and across species is less well understood. We studied the fitness consequences of clutch initiation for 181 female northern cardinals Cardinalis cardinalis and 1228 nests in forests within urban and rural landscapes of Ohio, USA between 2004–2007. Cardinals that bred earlier made significantly more nesting attempts, but cumulative number of young fledged was similar to that of later‐breeding individuals. The expected number of fledglings produced per successful nest was unrelated to date and remained ~1.8 fledglings across the season, despite the fact that nest survival rates improved dramatically as the season progressed. Because the probability of resighting breeding individuals in subsequent years was unrelated to first clutch initiation date, we have no evidence that clutch initiation affected adult survival. The absence of a clear benefit to early breeding appears to be a consequence of high rates of nest predation early in the breeding season.