James R. Lovvorn

UNDERSTANDING AND PREDICTING ECOLOGICAL DYNAMICS: ARE MAJOR SURPRISES INEVITABLE

Ecological surprises, substantial and unanticipated changes in the abundance of one or more species that result from previously unsuspected processes, are a common outcome of both experiments and observations in community and population ecology. Here, we give examples of such surprises along with the results of a survey of well-established field ecologists, most of whom have encountered one or more surprises over the course of their careers. Truly surprising results are common enough to require their consideration in any reasonable effort to characterize nature and manage natural resources. We classify surprises as dynamic-, pattern-, or intervention-based, and we speculate on the common processes that cause ecological systems to so often surprise us. A long-standing and still growing concern in the ecological literature is how best to make predictions of future population and community dynamics. Although most work on this subject involves statistical aspects of data analysis and modeling, the frequency and nature of ecological surprises imply that uncertainty cannot be easily tamed through improved analytical procedures, and that prudent management of both exploited and conserved communities will require precautionary and adaptive management approaches.

Diet and body condition of spectacled eiders wintering in pack ice of the Bering Sea

Spectacled eiders (Somateria fischeri) winter among leads in the Bering Sea pack ice, where they dive 40–70 m for benthic prey. During the first icebreaker cruises into that area, esophagi of collected eiders contained only clams, mostly Nuculana radiata, with no trace of the once-dominant Macoma calcarea. Alternative prey used elsewhere (snails, amphipods, other bivalves) were available but not eaten. Eiders ate mainly N. radiata 18–24 mm long, although M. calcarea of this length contained 62% more energy. Percent body lipid of eiders averaged 12±3% (SD) for 26 adult males and 14±3% for 12 adult females. Mean body mass (±SE) of these males in late March (1,688±21 g) was higher than reported for 53 males after arriving at breeding areas in late May (1,494±14 g). Body mass of these females (1,550±35 g) was lower (but not significantly) than reported for 11 females upon arrival at breeding sites (1,623±46 g). In 1999, the last spectacled eiders left the wintering area on 21 April, 4–8 weeks before their typical arrival at breeding sites. Their location is unknown in the interim, when habitats used appear critical to acquiring reserves for reproduction.

Stroke patterns and regulation of swim speed and energy cost in free-ranging Brünnich's guillemots

SUMMARY Loggers were attached to free-ranging Brünnichs guillemots Uria lomvia during dives, to measure swim speeds, body angles, stroke rates, stroke and glide durations, and acceleration patterns within strokes, and the data were used to model the mechanical costs of propelling the body fuselage (head and trunk excluding wings). During vertical dives to 102–135 m, guillemots regulated their speed during descent and much of ascent to about 1.6±0.2 m s–1. Stroke rate declined very gradually with depth, with little or no gliding between strokes. Entire strokes from 2 m to 20 m depth had similar forward thrust on upstroke vs downstroke, whereas at deeper depths and during horizontal swimming there was much greater thrust on the downstroke. Despite this distinct transition, these differences had small effect (<6%) on our estimates of mechanical cost to propel the body fuselage, which did not include drag of the wings. Work stroke–1 was quite high as speed increased dramatically in the first 5 m of descent against high buoyancy. Thereafter, speed and associated drag increased gradually as buoyancy slowly declined, so that mechanical work stroke–1 during the rest of descent stayed relatively constant. Similar work stroke–1 was maintained during non-pursuit swimming at the bottom, and during powered ascent to the depth of neutral buoyancy (about 71 m). Even with adjustments in respiratory air volume of ±60%, modeled work against buoyancy was important mainly in the top 15 m of descent, after which almost all work was against drag. Drag was in fact underestimated, as our values did not include enhancement of drag by altered flow around active swimmers. With increasing buoyancy during ascent above 71 m, stroke rate, glide periods, stroke acceleration patterns, body angle and work stroke–1 were far more variable than during descent; however, mean speed remained fairly constant until buoyancy increased rapidly near the surface. For dives to depths >20 m, drag is by far the main component of mechanical work for these diving birds, and speed may be regulated to keep work against drag within a relatively narrow range.

Food Dispersion and Foraging Energetics: a Mechanistic Synthesis for Field Studies of Avian Benthivores

Much effort has focused on modeling and measuring the energy costs of free existence and the foraging strategies of animals. However, few studies have quantitatively linked these approaches to the patch structure of foods in the field. We developed an individual-based model that relates field measurements of the dispersion of benthic foods to search costs and foraging profitability of diving ducks. On Lake Mattamuskeet, North Carolina, Canvasback ducks (Aythya valisineria) eat only the belowground winter buds of the submerged plant Vallisneria americana. We measured and modeled the patch structure of winter buds at the level of potential foraging loci, defined as contiguous circles 1 m in diameter. In the field and in the model, Canvasbacks make repeated vertical dives in such loci, foraging in the sediments by touch, before surface-swimming to another locus. We quantified first-order patchiness by fitting a negative binomial distribution to core samples taken at 50-m intervals along transects, to yield the frequencies of loci with different bud densities. Second-order patchiness was measured by taking cores at 1-m increments radiating from each sampling point, and regressing bud density at each sampling point on densities at these increments. No significant correlations were found, indicating that Canvasbacks could not predict food densities based on densities in nearby foraging loci. For the model, we generated food grids from the negative binomial distributions of core samples. Energy costs of diving were calculated by applying aerobic efficiencies (mechanical power output + aerobic power input) to biomechanical models. Unlike respirometry alone, this method accounts for effects on dive costs of varying water depth and dive duration. We used measurements of Canvasback intake rates at different bud densities to calculate profitability (energy intake minus expenditure) for each dive. Multivariate uncertainty analyses (Latin hypercube) indicated that profitability for Canvasbacks foraging on Vallisneria buds is determined mainly by food-item size and locomotor costs of descent. Bud metabolizable energy, water temperature, bud dispersion, and search and handling time coefficients of the functional response for intake rate have relatively minor influence. Individual-parameter perturbations indicated that to maintain the same foraging benefits, the total area of Vallisneria habitat would have to increase by 1.4-fold if dry mass per bud decreased from 0.10 to 0.03 g, and by 2.1-fold if water depth increased from 0.5 to 2 m. Our method allows study of interactions between patch structure and foraging energetics without detailed spatial mapping of foods, which is not feasible at appropriate scales for highly mobile benthivores. The model yields estimates of energy balance, contaminant intake, and amount and quality of foraging habitat required to sustain diving duck populations under varying environmental conditions. More accurate prediction of giving-up times and giving-up food densities will require better understanding of the time scale over which ducks balance their energy budgets.

Swim speeds and stroke patterns in wing-propelled divers: a comparison among alcids and a penguin

SUMMARY In diving birds, the volume and resulting buoyancy of air spaces changes with dive depth, and hydrodynamic drag varies with swim speed. These factors are important in the dive patterns and locomotion of alcids that use their wings both for aerial flight and underwater swimming and of penguins that use their wings only for swimming. Using small data-loggers on free-ranging birds diving to 20–30 m depth, we measured depth at 1 Hz and surge and heave accelerations at 32–64 Hz of four species of alcids (0.6–1.0 kg mass) and the smallest penguin species (1.2 kg). Low- and high-frequency components of the fluctuation of acceleration yielded estimates of body angles and stroke frequencies, respectively. Swim speed was estimated from body angle and rate of depth change. Brünnichs (Uria lomvia) and common (Uria aalge) guillemots descended almost vertically, whereas descent of razorbills (Alca torda), rhinoceros auklets (Cerorhinca monocerata) and little penguins (Eudyptula minor) was more oblique. For all species, swim speed during descent was within a relatively narrow range. Above depths of 20–30 m, where they were all positively buoyant, all species ascended without wing stroking. During descent, little penguins made forward accelerations on both the upstroke and downstroke regardless of dive depth. By contrast, descending alcids produced forward accelerations on both upstroke and downstroke at depths of <10 m but mainly on the downstroke at greater depths; this change seemed to correspond to the decrease of buoyancy with increasing depth. The magnitude of surge (forward) acceleration during downstrokes was smaller, and that during upstrokes greater, in little penguins than in alcids. This pattern presumably reflected the proportionally greater mass of upstroke muscles in penguins compared with alcids and may allow little penguins to swim at less variable instantaneous speeds.

Nutrient reserves, probability of cold spells and the question of reserve regulation in wintering canvasbacks

Interpreting body mass and composition of wintering birds is often confounded by the inability to discriminate endogenous regulation of reserves from effects of proximate weather and food conditions. Endogenous regulation is thought to act through genetically controlled set-points that change throughout the year, due to evolutionary adaptation to long-term probabilities of needing reserves at different times. For canvasbacks (Aythya valisineria) wintering in upper Chesapeake Bay, coastal North Carolina, and Louisiana, I calculated over many years the probability of cold spells when canvasbacks likely depend on reserves. I then analysed whether the timing of such cold spells is predictable enough to form the basis for monthly schedules of endogenous reserve regulation in free-ranging canvasbacks

RELATIVE FORAGING VALUE TO LESSER SCAUP DUCKS OF NATIVE AND EXOTIC CLAMS FROM SAN FRANCISCO BAY

Invasions of exotic invertebrates have greatly altered many aquatic communities, but impacts on the foraging energetics of predators seldom have been assessed. In San Francisco Bay, California (USA), a major community change occurred with introduction of the Asian clam (Potamocorbula amurensis) in 1986. This species now greatly outnumbers the previous clam prey of a variety of sharks, rays, sturgeon, flatfish, and crabs, as well as several diving duck species for which the bay is the most important wintering area on the U.S. Pacific Coast. P. amurensis also accumulates much higher levels of some contaminants than the formerly dominant prey. Because alteration of the food base or contaminated foods on wintering areas may be factors in the population decline of scaup ducks, effects of this exotic invasion are important to assess. For Lesser Scaup (Aythya affinis), we studied effects of differences in nutrient content, digestibility, crushing resistance of shells, areal density, size, and depth in the sediments on the relative foraging value of exotic P. amurensis vs. the formerly dominant native clam Macoma balthica. P. amurensis, including shells, had higher nitrogen and energy content per clam of the same length class, and higher digestibility of energy, than M. balthica. Gut retention time did not differ between clam species, so their relative profitability for scaup was determined mainly by the intake rate of digestible nutrients during short, costly dives. For scaup foraging in an aquarium 1.8 m deep, intake rates (number of prey per second) of food items buried in sand-filled trays increased with increasing prey density up to at least 4000 prey/m2. For items buried 3 cm deep, intake rates did not differ for prey <6 mm long vs. prey 6–12 mm long; however, intake rates were much lower when prey were deeper in the sediments (6 cm vs. 3 cm). In the field, a much higher percentage of P. amurensis were in the length range most commonly eaten by Lesser Scaup (<12 mm), and unlike M. balthica, almost all P. amurensis were in the top 5 cm of sediments where scaup intake rates are highest. In tensometer measurements, shells of P. amurensis were much harder to crush than shells of M. balthica, which might partly offset the apparent energetic advantages of P. amurensis. In many respects, the exotic P. amurensis appears to be a more valuable food than the native M. balthica for Lesser Scaup. However, because P. amurensis accumulates much higher levels of some contaminants, this exotic invasion increases the risk of toxicity to scaup and a range of other benthic predators.

Biomechanical conflicts between adaptations for diving and aerial flight in estuarine birds

Diving and aerial flight place conflicting physiological constraints on diving birds depending on their typical dive depths. The amount by which air volumes in the respiratory system and plumage are reduced by hydrostatic pressure decreases rapidly with depth. Thus, birds diving shallowly, and ascending passively by means of positive buoyancy, content with greater work against buoyancy as well as more unstable buoyancy as they move vertically in the water column. The buoyancy of air far exceeds that of tissues or blood, whose buoyancy does not change appreciably with depth. Accordingly, experiments on ducks suggest that birds adapt to shallow diving by increasing blood volume and thus blood oxygen stores while decreasing respiratory volume. During dives, increased inertia from greater mass of blood and associated muscle lowers the costs of foraging at the bottom by resisting the upward buoyant force, but raises the costs of descent because of higher inertial work in accelerating the body with each stroke. Thus, average dive depth (compression of buoyant air spaces), stroke kinematics (inertial effects), and the relative time spent descending versus bottom foraging will determine the appropriate balance between buoyancy and inertia for diving.Greater blood volume also increases wing loading, so elements of dive costs must be balanced against flight costs in optimizing allocation of oxygen stores to blood versus the respiratory system. For example, biomechanical models for ducks suggest that increasing blood volume while decreasing respiratory volume lowers dive costs only for dives to depths <5 m or for dives with extended time at constant depth. If flight costs are also considered, these anti-buoyancy mechanisms reduce daily energy expenditure only if average dive depth is <2 m. High wing-loading in many foot-propelled divers is probably not an adaptation to diving but rather a result of modifications in wing size and shape for high flight-speed. These wing modifications appear possible because competing demands on wing morphology (maneuverability, takeoff ability) are relaxed in open aquatic environments.

Distributional responses of canvasback ducks to weather and habitat change

(1) Loss of the plant-tuber foods of canvasback ducks (Aythya valisineria Wilson) wintering in upper Chesapeake Bay, U.S.A., has resulted in the ducks eating mostly clams. To examine the consequences of this habitat change, I investigated how seasonal availability of tubers and clams affects their use by canvasbacks in the mid-Atlantic region of the U.S.A., and how regional canvasback distributions are influenced by food availability and weather. (2) Estimates of ice cover on Chesapeake Bay from 1956 to 1987 suggest that tubers were often inaccessible for parts of January and February, regardless of their abundance. In North Carolina, where ice is seldom an important factor, sequential use of tubers and then clams by canvasbacks appears to result from initially high foraging efficiency for tubers, which declines as tubers are depleted. (3) Above evidence suggests that historically clams were a principal food of canvasbacks throughout this region during many winters; and that tubers were most consistently important during periods of fattening in late autumn to early winter and perhaps again in early spring. (4) Despite dramatic distributional responses to weather later in winter, a decline in numbers of canvasbacks in Maryland in early January has occurred since the early 1970s, which is independent of weather trends, and appears to have resulted from loss of tuberproducing plants there. (5) The decrease in numbers of canvasbacks in Maryland in early January corresponds to increases in Virginia and North Carolina, but this shift is not related to greater availability of plant foods in the latter two areas. (6) The southward shift of canvasbacks suggests that, with loss of tubers from Maryland, the food base there of clams alone is sometimes inadequate. Effective clam abundance may be greater in the southern areas because of higher clam densities, lower thermoregulation costs for ducks, or both. (7) Clam populations fluctuate widely among years and different areas. Thus, effects of the loss of plant tubers on canvasback populations probably depend on the frequency and geographical extent of shortages of alternative clam foods.: In order to investigate the effects of different irrigation and fertilization on soil microbial properties of summer maize field, we used ZN99 with high nitrogen use efficiency as the test material. The experiment adopted the split plot design which included two irrigation levels (526 mm and 263 mm) as the main plots, three fertilizer types (U, M, UM) and two fertilizer levels (N 100 kg . hm-2 and 200 kg . hm-2) as the subplots. The results showed that the irrigation level affected the regulating effects of fertilizer on soil microbial biomass (carbon and nitrogen) and microbial di- versity. The organic fertilizer application must be under the sufficient irrigation level to increase the soil MBC (14.3%-33.6%), MBN (1.8-2.3 times) and abundance significantly. A moderate rate of irrigation, higher rates of organic fertilizer application or organic manure combined with inorganic fertilizer could increase the nitrogen-fixation species and quantity of Firmicutes, γ-Proteobacteria and α-Proteobacteria in the soil. Under the same N level, there was no significant difference of grain yield between organic manure and inorganic fertilizer treatments. Considering sustainable production, proper organic manure application with moderate irrigation could increase the quantity of the soil microbial biomass and microbial diversity, and improve the capacity of soil to supply water and nutrients.

Algal vs. macrophyte inputs to food webs of inland saline wetlands

Invertebrate food webs in wetlands were traditionally thought to be fueled mainly by decaying macrophytes, but recently it has been recognized that microalgae may be more important. In particular, the paradigm that shredders of vascular plant litter dominate food web processes may not apply to many wetlands where shredders are rare and microalgae more abundant. This issue is complicated by potential consumption of flocs of dissolved organic matter (DOM) released from living plants, and of exopolymer secretions (EPS) from both autotrophic and heterotrophic microbes. In Wyoming, we used gut contents and stable isotopes to investigate organic matter sources for the dominant invertebrates in oligosaline (0.5–5 g/L total dissolved solids) and mesosaline (5–18 g/L) wetlands. We examined the trophic importance of microalgae vs. macrophytes in wetlands with and without emergent vegetation (Scirpus acutus), with different growth forms and species of submersed plants (Chara spp. vs. Potamogeton pectinatus), with do...