A Review of Environmental Factors Affecting Elk Winter Diets

Abstract: Decades of research have produced substantial data on elk (Cervus elaphus) diets in winter, when foraging conditions are most likely to affect population dynamics. Using data from 72 studies conducted in western North America between 1938 and 2002, we collated data on elk diets and environmental variables. We used these data to quantify diet selection by elk and to test whether variation in elk diets is associated with habitat type, winter severity, period of winter, human hunting, and study method. Graminoids (grasses and grass-like plants such as sedges) dominated elk diets and consistently occurred at a higher proportion in the diet than in elk foraging habitats, indicating preference. Forbs commonly made up ≤5% of the diet, with no evidence for preference; we conclude that forb use is largely incidental to grazing for graminoids. Browse was consumed in proportion to its availability, implying that the amount of browse in the diet was primarily determined by habitat use rather than selection. Comparing the diets of elk and sympatric ruminants, elk consistently selected graminoids more strongly than sympatric ruminants with the exception of bison (Bison bison), suggesting that elk are not environmentally forced to adopt the graminoid-biased diet that they normally select. The proportion of open meadows and grasslands on winter ranges was strongly and positively associated with graminoid consumption by elk. The proportion of graminoids in the diet was significantly lower in elk experiencing severe winter conditions or predation risk from human hunting. The period of winter (early, middle, and late) had only small effects on elk diets, as did the method by which the diet was determined. Overall, variation in elk diets is well-explained by a consistent tendency to select graminoids if available, modified by winter habitat type, predation risk, and winter severity, which can constrain habitat selection and access to grazing opportunities. To fully understand variation in foraging behavior, biologists should recognize these broad patterns when interpreting resource selection data. Managers should recognize that inconspicuous behavioral responses to environmental stimuli can alter the diet in ways that probably carry nutritional consequences.     

Responses of cetaceans to anthropogenic noise

• 1 Since the last thorough review of the effects of anthropogenic noise on cetaceans in 1995, a substantial number of research reports has been published and our ability to document response(s), or the lack thereof, has improved. While rigorous measurement of responses remains important, there is an increased need to interpret observed actions in the context of population‐level consequences and acceptable exposure levels. There has been little change in the sources of noise, with the notable addition of noise from wind farms and novel acoustic deterrent and harassment devices (ADDs/AHDs). Overall, the noise sources of primary concern are ships, seismic exploration, sonars of all types and some AHDs.
• 2 Responses to noise fall into three main categories: behavioural, acoustic and physiological. We reviewed reports of the first two exhaustively, reviewing all peer‐reviewed literature since 1995 with exceptions only for emerging subjects. Furthermore, we fully review only those studies for which received sound characteristics (amplitude and frequency) are reported, because interpreting what elicits responses or lack of responses is impossible without this exposure information. Behavioural responses include changes in surfacing, diving and heading patterns. Acoustic responses include changes in type or timing of vocalizations relative to the noise source. For physiological responses we address the issues of auditory threshold shifts and ‘stress’, albeit in a more limited capacity; a thorough review of physiological consequences is beyond the scope of this paper.
• 3 Overall, we found significant progress in the documentation of responses of cetaceans to various noise sources. However, we are concerned about the lack of investigation into the potential effects of prevalent noise sources such as commercial sonars, depth finders and fisheries acoustics gear. Furthermore, we were surprised at the number of experiments that failed to report any information about the sound exposure experienced by their experimental subjects. Conducting experiments with cetaceans is challenging and opportunities are limited, so use of the latter should be maximized and include rigorous measurements and or modelling of exposure.

     

Temporal dynamics and spatial variability in the enhancement of canopy leaf area under elevated atmospheric CO2

Increased canopy leaf area (L) may lead to higher forest productivity and alter processes such as species dynamics and ecosystem mass and energy fluxes. Few CO₂ enrichment studies have been conducted in closed canopy forests and none have shown a sustained enhancement of L. We reconstructed 8 years (1996–2003) of L at Duke's Free Air CO₂ Enrichment experiment to determine the effects of elevated atmospheric CO₂ concentration ([CO₂]) on L before and after canopy closure in a pine forest with a hardwood component, focusing on interactions with temporal variation in water availability and spatial variation in nitrogen (N) supply. The dynamics of L were reconstructed using data on leaf litterfall mass and specific leaf area for hardwoods, and needle litterfall mass and specific leaf area combined with needle elongation rates, and fascicle and shoot counts for pines. The dynamics of pine L production and senescence were unaffected by elevated [CO₂], although L senescence for hardwoods was slowed. Elevated [CO₂] enhanced pine L and the total canopy L (combined pine and hardwood species; P<0.050); on average, enhancement following canopy closure was ～16% and 14% respectively. However, variation in pine L and its response to elevated [CO₂] was not random. Each year pine L under ambient and elevated [CO₂] was spatially correlated to the variability in site nitrogen availability (e.g. r²=0.94 and 0.87 in 2001, when L was highest before declining due to droughts and storms), with the [CO₂]‐induced enhancement increasing with N (P=0.061). Incorporating data on N beyond the range of native fertility, achieved through N fertilization, indicated that pine L had reached the site maximum under elevated [CO₂] where native N was highest. Thus closed canopy pine forests may be able to increase leaf area under elevated [CO₂] in moderate fertility sites, but are unable to respond to [CO₂] in both infertile sites (insufficient resources) and sites having high levels of fertility (maximum utilization of resources). The total canopy L, representing the combined L of pine and hardwood species, was constant across the N gradient under both ambient and elevated [CO₂], generating a constant enhancement of canopy L. Thus, in mixed species stands, L of canopy hardwoods which developed on lower fertility sites (～3 g N inputs m^?2 yr^?1) may be sufficiently enhanced under elevated [CO₂] to compensate for the lack of response in pine L, and generate an appreciable response of total canopy L (～14%).     

Molecules,morphology and Mimeoma scarabs: evolutionary and taxonomic implications for a palm‐associated scarab group

Cyclocephaline scarabs, the second largest tribe of rhinoceros beetles, are important pollinators of early‐diverging angiosperm families in the tropics. The evolutionary history of cyclocephaline genera is poorly resolved and several genera are thought to be nonmonophyletic. We assess the monophyly of Mimeoma Casey, a group of Neotropical palm‐feeding scarabs, and its relationship to Cyclocephala with a phylogenetic analysis of 2899 bp of DNA sequence data and 18 morphological characters. All five species of Mimeoma were included in analyses along with species of Cyclocephala Dejean, Dyscinetus Harold and Tomarus Erichson as outgroup taxa. Nearly complete 28S, 12S and CO1 data were collected from 26 of 29 specimens, of which 16 samples were pinned, museum specimens. 28S data strongly support a nonmonophyletic Mimeoma; mitochondrial data (CO1 and 12S) suggest that Mimeoma species are nested within an apical clade of other Cyclocephala species; combined molecular and morphological data identify two strongly supported clades of Mimeoma species but do not support their sister relationship. Combined data show that Mimeoma species are nested within Cyclocephala, thus rendering Cyclocephala paraphyletic. Mimeoma is synonymized within Cyclocephala resulting in the following new combinations: Cyclocephala acuta Arrow n.comb ., Cyclocephala englemani (Ratcliffe) n.comb ., Cyclocephala maculata Burmeister n.comb ., Cyclocephala nigra (Endrödi) n.comb . and Cyclocephala signatoides Höhne n.comb . Our results demonstrate that pinned, museum specimens can be used to obtain DNA sequence data (particularly high‐copy gene regions) for evolutionary studies, and provide the first empirical support that host‐plant associations within cyclocephaline scarab clades are conserved at the plant family‐level.     

Phosphorus recycling in photorespiration maintains high photosynthetic capacity in woody species

Leaf photosynthetic CO₂ responses can provide insight into how major nutrients, such as phosphorus (P), constrain leaf CO₂ assimilation rates (A_net). However, triose‐phosphate limitations are rarely employed in the classic photosynthesis model and it is uncertain as to what extent these limitations occur in field situations. In contrast to predictions from biochemical theory of photosynthesis, we found consistent evidence in the field of lower A_net in high [CO₂] and low [O₂] than at ambient [O₂]. For 10 species of trees and shrubs across a range of soil P availability in Australia, none of them showed a positive response of A_net at saturating [CO₂] (i.e. A_max) to 2 kPa O₂. Three species showed >20% reductions in A_max in low [O₂], a phenomenon potentially explained by orthophosphate (P_i) savings during photorespiration. These species, with largest photosynthetic capacity and P_i > 2 mmol P m^?2, rely the most on additional P_i made available from photorespiration rather than species growing in P‐impoverished soils. The results suggest that rarely used adjustments to a biochemical photosynthesis model are useful for predicting A_max and give insight into the biochemical limitations of photosynthesis rates at a range of leaf P concentrations. Phosphate limitations to photosynthetic capacity are likely more common in the field than previously considered.     

The effect of fire and permafrost interactions on soil carbon accumulation in an upland black spruce ecosystem of interior Alaska: implications for post‐thaw carbon loss

High‐latitude regions store large amounts of organic carbon (OC) in active‐layer soils and permafrost, accounting for nearly half of the global belowground OC pool. In the boreal region, recent warming has promoted changes in the fire regime, which may exacerbate rates of permafrost thaw and alter soil OC dynamics in both organic and mineral soil. We examined how interactions between fire and permafrost govern rates of soil OC accumulation in organic horizons, mineral soil of the active layer, and near‐surface permafrost in a black spruce ecosystem of interior Alaska. To estimate OC accumulation rates, we used chronosequence, radiocarbon, and modeling approaches. We also developed a simple model to track long‐term changes in soil OC stocks over past fire cycles and to evaluate the response of OC stocks to future changes in the fire regime. Our chronosequence and radiocarbon data indicate that OC turnover varies with soil depth, with fastest turnover occurring in shallow organic horizons (～60 years) and slowest turnover in near‐surface permafrost (>3000 years). Modeling analysis indicates that OC accumulation in organic horizons was strongly governed by carbon losses via combustion and burial of charred remains in deep organic horizons. OC accumulation in mineral soil was influenced by active layer depth, which determined the proportion of mineral OC in a thawed or frozen state and thus, determined loss rates via decomposition. Our model results suggest that future changes in fire regime will result in substantial reductions in OC stocks, largely from the deep organic horizon. Additional OC losses will result from fire‐induced thawing of near‐surface permafrost. From these findings, we conclude that the vulnerability of deep OC stocks to future warming is closely linked to the sensitivity of permafrost to wildfire disturbance.