The foraging behavior of granivorous rodents and short-term apparent competition among seeds

The foraging behavior of a predator species is thought to bethe cause of short-term apparent competition among those preyspecies that share the predator. Short-term apparent competitionis the negative indirect effect that one prey species has onanother prey species via its effects on predator foraging behavior.In theory, the density-dependent foraging behavior of granivorousrodents and their preference for certain seeds are capable ofinducing short-term apparent competition among seed species.In this study, I examined the foraging behavior of two heteromyidrodent species (family Heteromyidae), Merriam's kangaroo rats(Dipodomys merriami) and little pocket mice (Perognathus longimembris).In one experiment I tested the preferences of both rodent speciesfor the seeds of eight plant species. Both rodent species exhibiteddistinct but variable preferences for some seeds and avoidanceof others. However, the differences in preference appearedto have only an occasional effect on the strength of the short-termapparent competition detected in a field experiment. In anotherexperiment, I found that captive individuals of both rodentspecies had approximately equal foraging effort (i.e., timespent foraging) in patches that contained a highly preferredseed type (Oryzopsis hymenoides) regardless of seed densityand the presence of a less preferred seed type (Astragalus cicer)in the patches. The rodents also harvested a large proportionof O. hymenoides seeds regardless of initial seed density;this precluded a negative indirect effect of A. cicer on O.hymenoides. But there was a negative indirect effect of O.hymenoides on A. cicer caused by rodents having a lower foragingeffort in patches that only contained A. cicer seeds than inpatches that contained A. cicer and O. hymenoides seeds. Theindirect interaction between O. hymenoides and A. cicer thusrepresented a case of short-term apparent competition thatwas non-reciprocal. Most importantly, it was caused by theforaging behavior of the rodents.     

Root foraging for patchy resources in eight herbaceous plant species

The root foraging strategy of a plant species can be characterized by measuring foraging scale, precision, and rate. Trade-offs among these traits have been predicted to contribute to coexistence of competitors. We tested for trade-offs among root foraging scale (total root mass and length of structural roots), precision (ln-ratio of root lengths in resource-rich and resource-poor patches), and rate (days required for roots to reach a resource-rich patch, or growth rate of roots within a resource-rich patch) in eight co-occurring species. We found that root foraging scale and precision were positively correlated, as were foraging scale and the rate of reaching patches. High relative growth rate of a species did not contribute to greater scale, precision, or rate of root foraging. Introduced species had greater foraging scale, precision, and rate than native species. The positive correlations between foraging scale and foraging precision and rate may give larger species a disproportionate advantage in competition for patchy soil resources, leading to size asymmetric competition below ground.     

Spatial models of foraging in clonal plant species

Ramets of some clonal plant species alter their internode lengths or their frequency of lateral branching in response to their immediate microenvironment. Such “plant foraging” responses are thought to allow clones to concentrate in favorable portions of their environment. Despite widespread interest among ecologists in plant foraging, few realistic models have been developed to examine conditions under which plant foraging responses are likely to provide clones with ecological benefit. In this paper, we develop spatially explicit, stochastic simulation models to examine consequences of both empirical and hypothetical plant foraging responses. We construct a hierarchical series of models in which we incorporate effects of resource heterogeneity on spacer lengths, angles of growth, and lateral branch production. We also vary the number, size, and arrangement of patches, and the presence or absence of ramet mortality. Simulations based on hypothetical data demonstrated the potential importance of shortening spacer lengths in favorable habitat. In these simulations, ramet crowding increased significantly, implying a potential cost to plant foraging responses whose magnitude is large enough to cause ramets to concentrate in favorable patches. Models calibrated with empirical data suggest that when clonal plants were able to concentrate in favorable habitat, this was usually caused by increased daughter ramet production in the favorable habitat. Variation in clonal growth angles had little impact on the ability of ramets or clones to locate favorable patches, but did increase the ability of clones to remain in favorable patches once found. Alterations in the number and size of patches strongly influenced the effectiveness of the foraging response. The spatial arrangement of patches also was important: clumped distributions of patches decreased the success with which plants located favorable patches, especially at the genet level and when the number of patches was low. Finally, when ramet mortality varied with patch quality, there was an increase in the percentage of ramets located in favorable patches; differential ramet mortality also lessened the impact of other effects, such as the decreased success of clones when patches are clumped. Overall, our models indicate that the effectiveness of plant foraging responses is variable and is likely to depend on a suite of environmental conditions.     

Seed dormancy and germination of five selected NATURA-2000 plant species from Croatia showing different germination strategies

Abstract

NATURA-2000, the ecological network of protected areas in the European Union that has been included in the Croatian legislation, defines Community important plant species with imperative on their conservation ex/in situ. In the Botanical Garden of the Faculty of Science, University of Zagreb, five NATURA-species have been selected as research subjects for germination study to shed light on the topic of their seed ecology and consequently advance their conservation efforts: Degenia velebitica, Scilla litardierei, Klasea lycopifolia, Ligularia sibirica and Genista holopetala. The freshly matured seeds of each species were exposed to cold or warm stratification in duration of four to sixteen weeks, and their germination was investigated through different regimes of incubation parameters, i.e. illumination (light/dark) and temperature (5, 15/6, 23?°C). All species had higher germination values after cold stratification, with the exception of G. holopetala. We concluded that D. velebitica, K. lycopifolia and presumably L. sibirica seeds has non-deep physiological dormancy while S. litardierei has deep complex morphophysiological dormancy and G. holopetala has physical dormancy. The observed patterns in seeds’ behaviour are consistent with the conditions in their natural habitats in Croatia and the knowledge of these patterns is vital for successful conservation strategies in the future.     

Root traits explain different foraging strategies between resprouting life histories

Drought and fire are prevalent disturbances in Mediterranean ecosystems. Plant species able to regrow after severe disturbances (i.e. resprouter life history) have higher allocation to roots and higher water potential during the dry season than coexisting non-resprouting species. However, seedlings of non-resprouters have a higher survival rate after summer drought. We predict that, to counteract their shallow-rooting systems and to maximize seedling survival, non-resprouters have root traits that confer higher efficiency in soil resource acquisition than resprouters. We tested this prediction in seedlings of less than 1.5 months old. We select 13 coexisting woody species (including both resprouters and non-resprouters), grew them in a common garden and measured the following root traits: length, surface, average diameter, root tissue density (RTD), specific root length (SRL), surface:volume ratio (SVR), specific tip density (STD), tip distribution in depth, internal links ratio (ILR), and degree of branching. These root traits were compared between the two resprouting life histories using both standard cross-species and phylogenetic-informed analysis. Non-resprouters showed higher SRL and longer, thinner and more branched laterals, especially in the upper soil layers. The external links (i.e. the most absorptive root region) were also more abundant, longer, thinner and with higher SVR for non-resprouters. The results were supported by the phylogenetic-informed analysis for the root traits most strongly related to soil resource acquisition (SRL, SVR and branching pattern). The seedling root structure of non-resprouters species allows them to more efficiently explore the upper soil layer, whereas seedling roots of resprouters will permit both carbon storage and deep soil penetration.     

Spatial distributions of multiple plant species affect herbivore foraging selectivity

Spatial distribution of food resources is an important factor determining herbivore foraging. Previous studies have demonstrated that clumped distribution of preferred species increases its consumption by herbivores in single‐ or two‐species systems. However, the potential impact of distribution pattern of less preferred species on foraging was ignored. In natural grasslands with high species diversity and complexity, the spatial distribution of preferred species impacts on herbivore foraging may be strongly correlated with the distribution of less preferred species. Our aims were to determine the effect of distribution of both preferred and other plant species on herbivore foraging under conditions close to a native, multi‐species foraging environment, and conceptualize the relationships between spatial distribution of food resources and herbivore consumption. We hypothesized that random distribution of non‐preferred species reduces herbivore consumption of preferred species because the dispersion of less preferred species likely disturbs herbivore foraging. We conducted an experiment using three species with five combinations of clumped and random distribution patterns. Three species Lathyrus quinquenervius, Phragmites australis and Leymus chinensis, were of high, intermediate and low preferences by sheep, respectively. Results showed that distribution of low preferred species, but not that of high preferred one, affected the consumption of preferred species. Sheep obtained higher consumption of high preferred species when low preferred species followed a clumped distribution than a random distribution. Distance between aggregations of high and low preferred species did not affect sheep foraging. It was concluded that the effects of spatial distribution of preferred species on its consumption are dependent on herbivore foraging strategy, and sheep can consume more preferred species when there is a consistent spatial pattern between preferred species and the entire food resource, and that the random dispersion of low preferred species in grassland may reduce herbivore consumption of high preferred species, thus minimizing selective grazing.     

Harvester ant foraging and plant species distribution in annual grassland

Summary The harvester antVeromessor andrei Mayr is a major seed predator on annual grassland growing on serpentine soil at Jasper Ridge, N. California. Ants forage intensively during morning and evening sessions in areas surrounding nests. Activity is at its most intense in early summer, coinciding with peak seed release for most annual plant species. Ants show strong preferences for seeds of non-dominant species, notablyMicroseris douglasii (DC.) Sch.-Bip., but preferences alter over the season in response to seed availability. Seeds of the dominant annual species,Lasthenia californica DC ex Lindley are not foraged until later in the summer when seeds of other species are less abundant.Seedling densities and species compositions on ant nests differ markedly from surrounding areas with species relative abundances being similar to those found on gopher mounds. An exclosure experiment in areas adjacent to nests indicated that ants significantly reduced the densities of species with preferred seeds. Ants may therefore significantly affect plant distribution and abundance within the serpentine grassland.     

Isotopes reveal contrasting water use strategies among coexisting plant species in a Mediterranean ecosystem

Variation in the stable carbon and oxygen isotope composition (δ(13) C, Δ(18) O) of co-occurring plant species may reflect the functional diversity of water use strategies present in natural plant communities. We investigated the patterns of water use among 10 coexisting plant species representing diverse taxonomic groups and life forms in semiarid southeast Spain by measuring their leaf δ(13) C and Δ(18) O, the oxygen isotope ratio of stem water and leaf gas exchange rates. Across species, Δ(18) O was tightly negatively correlated with stomatal conductance (g(s) ), whereas δ(13) C was positively correlated with intrinsic water use efficiency (WUE(i) ). Broad interspecific variation in Δ(18) O, δ(13) C and WUE(i) was largely determined by differences in g(s) , as indicated by a strong positive correlation between leaf δ(13) C and Δ(18) O across species The 10 co-occurring species segregated along a continuous ecophysiological gradient defined by their leaf δ(13) C and Δ(18) O, thus revealing a wide spectrum of stomatal regulation intensity and contrasting water use strategies ranging from 'profligate/opportunistic' (high g(s) , low WUE(i) ) to 'conservative' (low g(s) , high WUE(i) ). Coexisting species maintained their relative isotopic rankings in 2?yr with contrasting rainfall, suggesting the existence of species-specific 'isotopic niches' that reflect ecophysiological niche segregation in dryland plant communities.     

Germination and water dispersal of seeds from a threatened plant species Penthorum chinense

To conserve a threatened plant species (Penthorum chinense Pursh) in Japan, seed germination responses to pretreatment (imbibition and/or chilled), temperature and light, and seed dispersal by water were examined. The seeds collected from abandoned paddy fields in a warm temperate region, central Japan, germinated in light (14 h photoperiod; light 22°C, dark 21°C) after a moist-chilled treatment. After this pretreatment, the seeds germinated well at 10–25°C (optimum temperature 15°C), but did not germinate in darkness even at the optimum temperature. Most of the seeds floated on distilled water, but 20–60% of the seeds that were collected from several populations sank in distilled water, indicating dimorphism in seed dispersal by water. The floating and sunken seeds did not show significant differences in weight and germination rate within a population. The addition of a surface-active agent in distilled water submerged the seeds, indicating that the buoyancy of the seeds is attributable to an oil coating on the seed surface that enhances the interfacial tension on the seeds. Three times the number of seeds sank in river water collected from a rural area than in distilled water. A greater number of seeds also sank in water that had increasing concentrations of linear alkylbenzenesulfonate, which is a major component of synthetic detergents. This suggests that the water dispersal of this species is suppressed by surface-active agents, including detergents, in river water.     

Utilization of phosphate from different sources by six plant species

Summary Six plant species, wheat, paspalum grass (Paspalum plicatulum), maize, molasses grass (Melinis minutiflora), soybean and buckwheat (Fagopyrum esculentum) were compared for their abilities to utilize phosphate from superphosphate, a calcined aluminum phosphate and four rock phosphates.Buckwheat showed an exceptional behaviour in that it could utilize all phosphates. For the other plants, only the calcined aluminum phosphate and one rock phosphate (hyperphosphate) had significant fertilizing values. Their efficiencies, relative to superphosphate, were 0.45 and 0.11 for wheat, 0.73 and 0.43 for paspalum grass, 0.50 and 0.37 for maize, 0.46 and 0.42 for molasses grass, 0.28 and 0.38 for soybean, and 0.72 and 1.08 for buckwheat, respectively.For three P sources, superphosphate, calcined aluminum phosphate and hyperphosphate, a relationship between soil acidity and P uptake was found. Soil pH in its turn was negatively related to the ratio of total equivalents of cations and those of anions absorbed. Consequenly, P uptake was positively related to the ratio of total equivalents of cations to those of anions absorbed. The same effect of plant species on soil pH could also explain the difference in uptake of P from sparingly soluble phosphates. The relative efficiencies of calcined aluminum phosphate and hyperphosphate for the various plant species were closely related to the ratio of total cations and total anions absorbed by these plants.on leave at the Agricultural University during 1977.     

Spatial and reversal learning in congeneric lizards with different foraging strategies

Environmental demands that require intensive search for mates, food and nest sites are correlated with efficient spatial memory in many mammalian and avian species. This convergence of evidence has led to the view that spatial memory, and the neurological structures associated with it, have been selected in niches that require memory for the location of goal objects. Whether such evolutionary demands are also correlated with nonspatial abilities that require flexible use of associations similar to those required for spatial memory has not been well studied. In addition, correlations between niche types and the use of spatial or nonspatial memory have not been investigated in nonmammalian, nonavian taxa. In this study, we investigated the relationship between foraging strategies and performance on two tasks, one spatial and the other nonspatial, in congeneric lizard species: Acanthodactylus boskianus, an active forager that collects clumped sedentary prey, Acanthodactylus scutellatus, a sit-and-wait predator that collects distributed mobile prey. The two species did not differ in their performance of a spatial memory task, but A. boskianus, the active forager, performed better on the reversal of a visual discrimination, a nonspatial task. These findings question the generality of the spatial adaptation model for vertebrates. We present the pliancy hypothesis, which we developed to account for these results. Copyright 1999 The Association for the Study of Animal Behaviour.     

Optimal foraging: the difficulty of exploiting different feeding strategies simultaneously

Summary The foraging efficiency of a visually feeding fish, perch (Perca fluviatilis) was studied on two prey species (Daphnia magna and Chaoborus obscuripus) presented either separately or combined. It is shown that when both prey species are present, the foraging efficiency of the predator is reduced. This is due to the predator's inability to simultaneously cope with prey species with different anti-predatory behaviour. In the mixed-meal experiment the predator captured both prey species in equal proportions in disagreement with optimal foraging models assuming that handling time and encounter rate for a prey species are independent of other prey species. The results are, however, in agreement with optimal foraging models assuming that handling time and encounter rate are influenced by short time learning.     

The behavior of ants of different species during joint foraging

The interaction between ants with different levels of social and territorial organization (Formica polyctena, Myrmica rubra, M. ruginodis, M. schencki, Lasius niger, and Formica fusca) was studied in the Ugra National Park (Kaluga Prov., Russia). The behavior of ants competing for an abundant source of food (carbohydrate feeders) was analyzed. It was found that the possibility of ants coexisting in a multi-species association was ensured, in particular, by different strategies of feeding behavior. A species with effective mobilization may achieve a greater competitive success as compared to more aggressive but less effectively mobilizing ones. Species characterized by non-aggressive behavior and ineffective mobilization may ensure their biological success due to an efficient system of foraging.     

Together but different: co-occurring dune plant species differ in their water- and nitrogen-use strategies

Stress factors may severely constrain the range of plant physiological responses in harsh environments. Convergence of traits is expected in coastal dunes because of environmental filtering imposed by severe abiotic factors. However, the wide range of morphological and phenological traits exhibited by coexisting dune species suggests considerable variation in functional traits. We hypothesized that the constraints imposed by structural traits ought to translate into physiological differences. Five dominant species with different morphological traits, but coexisting in a homogeneous dune area in Northwest Spain, were selected for study. Soil characteristics and leaf functional traits were measured in April, June and November 2008. Integrated water-use efficiency (assessed by C isotope discrimination) and N acquisition and use strategies (estimated by N isotope composition) varied significantly among species and the differences changed over time. Species differences in specific leaf area, relative water content, leaf N and C:N ratio, also varied over time. The species differed in stomatal density but not in soil characteristics, with the exception of pH. Species differences in functional traits related to the use of resources suggest species niche segregation. Species-specific temporal effects on the use of these resources support temporal niche differentiation. Somewhat in contrast to the findings of previous studies on harsh environments, this study revealed a considerable level of functional diversity and complexity, suggesting that dune plant species have evolved species-specific strategies to survive by partitioning growth-limiting resources.     

Nutrient foraging traits in 10 co-occurring plant species of contrasting life forms

1 Responses to spatial heterogeneity of soil nutrients were tested in 10 plant species that differ in life form and successional status, but which co-occur in the South Carolina coastal plain. The morphological responses of the root system were tested by assessing scale (represented by root mass and root length densities), precision (preferential proliferation of roots in nutrient-rich patches compared with less fertile patches) and discrimination (ability to detect and proliferate within the richest patches when patches vary in nutrient concentration). We also investigated sensitivity (growth benefits gained as spatial heterogeneity of nutrients increases, measured as total biomass).
2 Ten individuals of each species were grown in pots under four treatments that had differing nutrient distribution but the same overall nutrient addition. Plants were harvested when roots reached pot edge.
3 We observed high variation between species in scale, precision and sensitivity. No significant discrimination responses were observed, although greatest root mass density occurred at intermediate fertility levels for all species.
4 We rejected the hypothesis that scale and precision are negatively correlated. Indeed, in herbaceous species alone, scale and precision were positively correlated.
5 Sensitivity was not closely related to precision, indicating that proliferation of roots in fertile patches does not always yield growth benefits in heterogeneous soils. Further, some sensitive species had very low precision, suggesting that a positive growth response in heterogeneous environments may be related to plasticity in physiology or root life span, rather than morphology.
6 Plant life form was not correlated with precision or sensitivity. However, scale of response was greater in herbs than in woody plants, possibly because the two life forms develop root systems at different rates.     

Antioxidant composition of selected high alpine plant species from different altitudes

Selected high alpine plant species were collected from different elevations in the Obergurgl/Ötztal subnival and nival regions in Austria to investigate the content of antioxidants in plants growing under the particular in vivo conditions experienced in this area (e.g. chilling stress, short vegetation period and high irradiation). The contents of antioxidants (ascorbic acid, tocopherol and glutathionc) and photosynthetic pigments were measured throughout the day. The contents of most compounds were found to follow a diurnal rhythm, with the maximum occurring at midday and the minimum during the night. It was not clear whether these fluctuations were temperature-dependent or light-dependent. Analyses of the antioxidant spectrum in the same plant species at different altitudes (and thus under different environmental conditions: as altitude increases, for example, day temperature decreases and light intensity increases) revealed that the total amount of antioxidants increases as altitude increases. This enhancement was mainly due to ascorbic acid contents. Each plant species displayed a specific reaction to the increase in stress that accompanies an increase in altitude, resulting in a broad adaptation spectrum for these plants. The present study suggests that the combined effect of lower temperature and higher light intensity induces higher antioxidant contents.     

Phytochemical variation within a single plant species influences foraging behavior of deer

To determine how black‐tailed deer Odocoileus hemionus columbianus respond to phytochemical cues while browsing in heterogeneous phytochemical environments, we offered captive and free‐range deer cloned rooted cuttings and seedlings of western redcedar Thuja plicata selected for varying monoterpene content. Black‐tailed deer were thus allowed to browse among a controlled array of phytochemical cues in a series of experiments designed to evaluate foraging behavior at fine (within plot) and coarse (plot selection) scales. Within‐plot diet selection experiments demonstrated that browse preference for individual western redcedar plants was a function of foliar monoterpene concentration. Individual plant palatability combined with momentary maximization foraging strategy promoted survival of heavily defended plants. Among‐plot foraging experiments demonstrated that coarse‐scale foraging preferences were strongly influenced by distributions of high monoterpene‐containing western redcedar in available plots. Olfaction may play a significant role in both fine and coarse‐scale browse behaviors of deer as they employ a risk‐averse foraging strategy.     

啮齿动物对不同大小和种皮特征种子的取食和搬运

在宁夏六盘山区的辽东栎灌丛,以不同大小的辽东栎种子、野李和华山松种子为材料,研究了种子大小和种皮特征对啮齿动物取食和搬运行为的影响.结果表明:辽东栎小种子和华山松种子的就地取食率均显著高于辽东栎大种子和野李种子;具有厚而坚硬种皮(内果皮)的野李种子被啮齿动物搬运后的取食率和贮藏率均最高.辽东栎大种子被啮齿动物搬运后取食的距离最大(3.10 m),被搬运后贮藏的距离(6.48 m)显著大于其他类型种子.除野李种子外,其他3种类型种子的单个种子取食点都在80％以上,所有种子的单个种子贮藏点都在90％以上,含2个以上种子的取食点和贮藏点较少.较高比例的华山松种子在灌丛基部和洞穴以外的其他环境被取食,其他3种类型种子被啮齿动物搬运后主要集中在灌丛基部和洞穴中取食;种皮(内果皮)厚而坚硬的种子以土壤埋藏方式贮藏的比例偏高.