Effects of climate change on phenology, frost damage, and floral abundance of montane wildflowers

The timing of life history traits is central to lifetime fitness and nowhere is this more evident or well studied as in the phenology of flowering in governing plant reproductive success. Recent changes in the timing of environmental events attributable to climate change, such as the date of snowmelt at high altitudes, which initiates the growing season, have had important repercussions for some common perennial herbaceous wildflower species. The phenology of flowering at the Rocky Mountain Biological Laboratory (Colorado, USA) is strongly influenced by date of snowmelt, which makes this site ideal for examining phenological responses to climate change. Flower buds of Delphinium barbeyi, Erigeron speciosus, and Helianthella quinquenervis are sensitive to frost, and the earlier beginning of the growing season in recent years has exposed them to more frequent mid-June frost kills. From 1992 to 1998, on average 36.1% of Helianthella buds were frosted, but for 1999-2006 the mean is 73.9%; in only one year since 1998 have plants escaped all frost damage. For all three of these perennial species, there is a significant relationship between the date of snowmelt and the abundance of flowering that summer. Greater snowpack results in later snowmelt, later beginning of the growing season, and less frost mortality of buds. Microhabitat differences in snow accumulation, snowmelt patterns, and cold air drainage during frost events can be significant; an elevation difference of only 12 m between two plots resulted in a temperature difference of almost 2 degrees C in 2006 and a difference of 37% in frost damage to buds. The loss of flowers and therefore seeds can reduce recruitment in these plant populations, and affect pollinators, herbivores, and seed predators that previously relied on them. Other plant species in this environment are similarly susceptible to frost damage so the negative effects for recruitment and for consumers dependent on flowers and seeds could be widespread. These findings point out the paradox of increased frost damage in the face of global warming, provide important insights into the adaptive significance of phenology, and have general implications for flowering plants throughout the region and anywhere climate change is having similar impacts.     

Scale-dependent responses of plant biodiversity to nitrogen enrichment

Experimental studies demonstrating that nitrogen (N) enrichment reduces plant diversity within individual plots have led to the conclusion that anthropogenic N enrichment is a threat to global biodiversity. These conclusions overlook the influence of spatial scale, however, as N enrichment may alter beta diversity (i.e., how similar plots are in their species composition), which would likely alter the degree to which N-induced changes in diversity within localities translate to changes in diversity at larger scales that are relevant to policy and management. Currently, it is unclear how N enrichment affects biodiversity at scales larger than a small plot. We synthesized data from 18 N-enrichment experiments across North America to examine the effects of N enrichment on plant species diversity at three spatial scales: small (within plots), intermediate (among plots), and large (within and among plots). We found that N enrichment reduced plant diversity within plots by an average of 25% (ranging from a reduction of 61% to an increase of 5%) and frequently enhanced beta diversity. The extent to which N enrichment altered beta diversity, however, varied substantially among sites (from a 22% increase to an 18% reduction) and was contingent on site productivity. Specifically, N enrichment enhanced beta diversity at low-productivity sites but reduced beta diversity at high-productivity sites. N-induced changes in beta diversity generally reduced the extent of species loss at larger scales to an average of 22% (ranging from a reduction of 54% to an increase of 18%). Our results demonstrate that N enrichment often reduces biodiversity at both local and regional scales, but that a focus on the effects of N enrichment on biodiversity at small spatial scales may often overestimate (and sometimes underestimate) declines in regional biodiversity by failing to recognize the effects of N on beta diversity.     

Intraspecific variation in a predator affects community structure and cascading trophic interactions

Intraspecific phenotypic variation in ecologically important traits is widespread and important for evolutionary processes, but its effects on community and ecosystem processes are poorly understood. We use life history differences among populations of alewives, Alosa pseudoharengus, to test the effects of intraspecific phenotypic variation in a predator on pelagic zooplankton community structure and the strength of cascading trophic interactions. We focus on the effects of differences in (1) the duration of residence in fresh water (either seasonal or year-round) and (2) differences in foraging morphology, both of which may strongly influence interactions between alewives and their prey. We measured zooplankton community structure, algal biomass, and spring total phosphorus in lakes that contained landlocked, anadromous, or no alewives. Both the duration of residence and the intraspecific variation in foraging morphology strongly influenced zooplankton community structure. Lakes with landlocked alewives had small-bodied zooplankton year-round, and lakes with no alewives had large-bodied zooplankton year-round. In contrast, zooplankton communities in lakes with anadromous alewives cycled between large-bodied zooplankton in the winter and spring and small-bodied zooplankton in the summer. In summer, differences in feeding morphology of alewives caused zooplankton biomass to be lower and body size to be smaller in lakes with anadromous alewives than in lakes with landlocked alewives. Furthermore, intraspecific variation altered the strength of the trophic cascade caused by alewives. Our results demonstrate that intraspecific phenotypic variation of predators can regulate community structure and ecosystem processes by modifying the form and strength of complex trophic interactions.     

Terrestrial habitat selection and strong density-dependent mortality in recently metamorphosed amphibians

To predict the effects of terrestrial habitat change on amphibian populations, we need to know how amphibians respond to habitat heterogeneity, and whether habitat choice remains consistent throughout the life-history cycle. We conducted four experiments to evaluate how the spatial distribution of juvenile wood frogs, Rana sylvatica (including both overall abundance and localized density), was influenced by habitat choice and habitat structure, and how this relationship changed with spatial scale and behavioral phase. The four experiments included (1) habitat manipulation on replicated 10-ha landscapes surrounding breeding pools; (2) short-term experiments with individual frogs emigrating through a manipulated landscape of 1 m wide hexagonal patches; and habitat manipulations in (3) small (4-m2); and (4) large (100-m2) enclosures with multiple individuals to compare behavior both during and following emigration. The spatial distribution of juvenile wood frogs following emigration resulted from differences in the scale at which juvenile amphibians responded to habitat heterogeneity during active vs. settled behavioral phases. During emigration, juvenile wood frogs responded to coarse-scale variation in habitat (selection between 2.2-ha forest treatments) but not to fine-scale variation. After settling, however, animals showed habitat selection at much smaller scales (2-4 m2). This resulted in high densities of animals in small patches of suitable habitat where they experienced rapid mortality. No evidence of density-dependent habitat selection was seen, with juveniles typically choosing to remain at extremely high densities in high-quality habitat, rather than occupying low-quality habitat. These experiments demonstrate how prediction of the terrestrial distribution of juvenile amphibians requires understanding of the complex behavioral responses to habitat heterogeneity. Understanding these patterns is important, given that human alterations to amphibian habitats may generate extremely high densities of animals, resulting in high density-dependent mortality.     

Cloning of urea transporters from the kidneys of two batoid elasmobranchs: evidence for a common elasmobranch urea transporter isoform

One of the two phloretin-sensitive, facilitated urea transporters identified from the kidneys of the myliobatiform, euryhaline elasmobranch, Dasyatis sabina, a 379 amino acid protein ([D. sabina]strUT-2), was very similar to the 380 amino acid isoform (shUT) present in the kidney of the squaliform, dogfish shark, Squalus acanthias (a species that can be considered marginally euryhaline since it utilizes upper estuarine, as well as ocean habitats). To test the proposal that this isoform is a conserved urea transporter (UT) expressed in the kidneys of diverse elasmobranchs, UTs were cloned from the kidneys of a rajiform elasmobranch, the stenohaline skate, Leucoraja ocellata and another dasyatid stingray, the marginally euryhaline, Dasyatis say. Utilizing 5′/3′ RACE, a 2,060 nt cDNA that encoded a phloretin-sensitive, 378 amino acid skate urea transporter ([L. ocellata]skUT-2) and a 1,683 nt cDNA that encoded a stingray 379 amino acid UT ([D. say]strUT-2) were obtained. These deduced UTs have a very high sequence identity with the known elasmobranch Uts. [L. ocellata]skUT-2 was 86% identical to [D. sabina]strUT-2 and 84% identical to [S. acanthias]shUT. [D. say]strUT-2 was 97% identical to the [D. sabina]strUT-2. These findings support the hypothesis that a conserved UT isoform is present in the kidneys of marine elasmobranchs, and is an important pathway for facilitated urea transport in the kidneys of marine elasmobranchs. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. H.A. Gefroh and E.E. Cwengros contributed equally to this study.     

Spatial variation of otolith elemental signatures among juvenile gray snapper (<Emphasis Type="Italic">Lutjanus griseus</Emphasis>) inhabiting southern Florida waters

Juvenile gray snapper, Lutjanus griseus, are believed to use bays and estuaries in southern Florida as nurseries before moving out to the adjoining reef tract as adults. Using high-resolution sector field-inductively coupled plasma-mass spectrometry (SF-ICP-MS), the elemental chemistry of the otoliths of juveniles from five nursery regions was resolved by establishing elemental “signatures” for each region. In this study we simultaneously analyzed 32 elements including a suite of rare earth elements. A stepwise variable selection procedure retained a subset of eight elements that contributed substantially to separating otolith samples, including two rare earth elements; this is one of the first studies in which rare earth elements in otoliths have contributed to separation of fish stocks. The classification success rate in assigning fishes to the correct region of origin was 82%. Resolution of sites less than 10 km apart suggested high site fidelity in juvenile gray snapper and little mixing of water masses between sites. The juvenile nursery signatures will be used to determine the relative contribution of different nurseries to the adult population on an adjoining reef tract.     

Direct versus indirect methods of quantifying herbivore grazing impact on a coral reef

Two methods were used to assess the grazing impact of roving herbivorous fishes across a coral reef depth gradient within Pioneer Bay, Orpheus Island, Great Barrier Reef. The first technique employed was a method traditionally used to quantify herbivory on coral reefs via the (indirect) inference of herbivore impact from biomass estimates and reported feeding rates. The second method (one of a range of direct approaches) used remote underwater video cameras to film the daily feeding activity of roving herbivores in the absence of divers. Both techniques recorded similar patterns and relative levels of herbivore biomass across five reef zones at the study site. Indirect estimates of the grazing impact across the reef depth gradient of the three predominant species of herbivore broadly coincided with levels quantified directly by remote underwater video, indicating that, to a large extent, presence does correspond to function. However, the video data suggested that, for individual species in particular reef zones, the absolute level of impact may be less than that inferred from presence. In the case of the parrotfish Scarus rivulatus, the video recordings suggested that, at the reef crest, an average of 52% (±18 SE) of each m² area of reef would be grazed each month, compared with an area of 109% (±41 SE) suggested by inferring grazing activity from presence alone. Potential biases associated with remote video recorders may explain some of the discrepancy between values. Overall, the results suggest that, for some fish groups, the indirect method of inferring function from presence can provide a good indication of relative levels of herbivore impact across a coral reef. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Size–frequency dynamics of NE Pacific abyssal ophiuroids (Echinodermata: Ophiuroidea)

The 17-year time-series study at Station M in the NE Pacific has provided one of the longest datasets on deep-sea ophiuroids to date. Station M is an abyssal site characterized by low topographical relief and seasonal and interannual variation in surface-derived food inputs. From 1989 to 2005, over 31,000 ophiuroid specimens were collected. Size–frequency distributions of the four dominant species, Ophiura bathybia, Amphilepis patens, Amphiura carchara and Ophiacantha cosmica, were examined for recruitment and the role of surface-derived food supplies on body size distributions. Juveniles were collected in sediment traps and used to investigate settlement patterns and seasonality. Trawl samples showed no indication of seasonal changes in recruitment to larger size classes; however, there was evidence of seasonal settling of juveniles. Interannual differences in median disk diameters and size distributions of trawl-collected adults are greater than those at the seasonal scale. Three of the four species, O. bathybia, A. patens and O. cosmica, had co-varying monthly median disk diameters, suggesting they may have a similar factor(s) controlling their growth and abundance. Interannual differences in monthly size distributions were generally greater than those between seasons. Cross-correlations between the particulate organic carbon (POC) flux (food supply) and size distribution indices for O. bathybia, A. patens and O. cosmica all were significant indicating that increases in food supply were followed by increases in the proportion of smaller size classes after approximately 17–22 months. These findings suggest that food inputs are indeed an important factor influencing deep-sea ophiuroid populations on interannual time scales, more generally supporting the long-hypothesized connection between food availability and population size structure in the deep sea. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Does the risk of encountering lions influence African herbivore behaviour at waterholes?

A central question in the study of predator–prey relationships is to what extent prey behaviour is determined by avoidance of predators. Here, we test whether the long-term risk of encountering lions and the presence of lions in the vicinity influence the behaviour of large African herbivores at waterholes through avoidance of high-risk areas, increases in group size, changes in temporal niche or changes in the time spent in waterhole areas. In Hwange National Park, Zimbabwe, we monitored waterholes to study the behaviour of nine herbivore species under different risks of encountering lions. We radio-collared 26 lions in the study area which provided the opportunity to monitor whether lions were present during observation sessions and to map longer-term seasonal landscapes of risk of encountering lions. Our results show that the preferred prey species for lions (buffalo, kudu and giraffe) avoided risky waterholes. Group size increased as encounter risk increased for only two species (wildebeest and zebra), but this effect was not strong. Interestingly, buffalo avoided the hours of the day which are dangerous when the long-term and short-term risks of encountering lions were high, and all species showed avoidance of waterhole use at night times when lions were in the vicinity. This illustrates well how prey can make temporal adjustments to avoid dangerous periods coinciding with predator hunting. Additionally, many herbivores spent more time accessing water to drink when the long- and short-term risks of encountering lions were high, and they showed longer potential drinking time when the long-term risk of encountering lions was high, suggesting higher levels of vigilance. This study illustrates the diversity of behavioural adjustments to the risk of encountering a predator and how prey respond differently to temporal variations in this risk.     

Probabilistic movement model with emigration simulates movements of deer in Nebraska, 1990–2006

Movements of deer can affect population dynamics, spatial redistribution, and transmission and spread of diseases. Our goal was to model the movement of deer in Nebraska in an attempt to predict the potential for spread of chronic wasting disease (CWD) into eastern Nebraska. We collared and radio-tracked >600 white-tailed deer (Odocoileus virginianus) and mule deer (Odocoileus hemionus) in Nebraska during 1990–2006. We observed large displacements (>10 km) for both species and sexes of deer, including migrations up to 100 km and dispersals up to 50 km. Average distance traveled between successive daily locations was 166 m for male and 173 for female deer in eastern Nebraska, and 427 m for male and 459 for female deer in western Nebraska. Average daily displacement from initial capture point was 10 m for male and 14 m for female deer in eastern Nebraska, and 27 m for male and 28 m for female deer in western Nebraska. We used these data on naturally occurring movements to create and test 6 individual-based models of movement for white-tailed deer and mule deer in Nebraska, including models that incorporated sampling from empirical distributions of movement lengths and turn angles (DIST), correlated random walks (CRW), home point fidelity (FOCUS), shifting home point (SHIFT), probabilistic movement acceptance (MOVE), and probabilistic movement with emigration (MOVEwEMI). We created models in sequence in an attempt to account for the shortcomings of the previous model(s). We used the Kolmogrov–Smirnov goodness-of-fit test to verify improvement of simulated annual displacement distributions to empirical displacement distributions. The best-fit model (D = 0.07 and 0.08 for eastern and western Nebraska, respectively) included a probabilistic movement chance with emigration (MOVEwEMI) and resulted in an optimal daily movement length of 350 m (maximum daily movement length of 2800 m for emigrators) for eastern Nebraska and 370 m (maximum of 2960 m) for western Nebraska. The proportion of deer that moved as emigrators was 0.10 and 0.13 for eastern and western Nebraska, respectively. We propose that the observed spread of CWD may be driven by large movements of a small proportion of deer that help to establish a low prevalence of the disease in areas east of the current endemic area. Our movement models will be used in a larger individual-based simulation of movement, survival, and transmission of CWD to help determine future surveillance and management actions.