Endophytic fungi alter foraging and dispersal by desert seed-harvesting ants

Endophytic fungi are thought to interact mutualistically with host plants by producing alkaloid metabolites that deter herbivory. Since such fungi are transmitted via seed in some grasses, the presence of endophytes may also protect plants from seed predators. We conducted seed choice experiments for two dominant seed harvesting ants, Pogonomyrmex rugosus in the Sonoran desert and Pogonomyrmex occidentalis at a higher elevation, riparian zone in Arizona, USA. Non-infected fescue (Festuca arundinacea) seeds and seeds infected with the endophytic fungus, Acremonium coenophialum, were presented to ant colonies in three different populations. Infected seeds were harvested less frequently than non-infected seed for the two populations of Pogonomyrmex rugosus but not for the population of Pogonomyrmex occidentalis. We also a conducted seed dispersal experiment for one population of Pogonomyrmex rugosus. Of the seeds that were harvested, most of the colonies discarded more infected seeds into refuse piles than expected by chance. Seeds discarded into refuse piles have greater germination success than surrounding areas. The most important interaction of endophytes and grasses may be deterrence of seed predation and enhancing the probability of germinating in favorable sites, since these processes directly increase plant fitness.     

Seeds of the annual grasses Schizachyrium spp. as a food resource for tropical granivorous birds

Seed dynamics of the annual tropical grasses Schizachyrium fragile (R. Br.) A. Camus, S. pachyarthron C. Gardner and S. pseudeulalia (Hosok.) S.T. Blake were studied with the aim of documenting fluxes in granivore food resources. In S. fragile, seed production began in the early dry season, and seed output was primarily influenced by seedling survival. Following seed-fall, there were 651 S. fragile seeds/m² (393 kernels/m²⁾ and 1014 S. pachyarthron seeds/m² (593 kernels/m²) across the study area, with a combined kernel biomass of 14.1 × 10³ g/ha. Seed densities remained stable through the dry season, then declined rapidly after wet season rain. Initial wet season rains of up to 25 mm, 40 mm and 50 mm resulted in seed-bank germination of < 5%, 57%, and 93%, respectively. Some seeds were buried by rain and resurfaced or germinated later, but about 30% was lost. Widespread and abrupt depletion of the seed-bank is likely to occur at the start of about 8% of wet seasons on central Cape York Peninsula, leaving little seed, either for subsequent germination or as food for granivores. Burning early in the dry season, when most seeds were still on the plants, reduced seed densities by 85%. The proportion of seeds with sound kernels was reduced in areas burnt by dry season fires, both directly after the fires and, subsequently, as a result of preferential granivore activity. Areas burnt in the dry season were thus depleted of seed earlier than were unburnt areas. Food availability for granivores should therefore be optimized by fire regimes that include a range of burning histories, including fires in both early dry and early wet seasons as well as keeping other areas unburnt.     

Agricultural habitat-type and the breeding performance of granivorous farmland birds in Britain

Conservation concern about granivorous birds has led to the implication of changing agricultural practices as causes of widespread population decline. We investigate relationships between breeding performance and the agricultural environment for ten granivorous farmland bird species (Stock Dove Columba oenas, Skylark Alauda arvensis, Tree Sparrow Passer montanus, Chaffinch Fringilla coelebs, Greenfinch Carduelis chloris, Linnet C. cannabina, Bullfinch Pyrrhula pyrrhula, Reed Bunting Emberiza schoeniclus, Yellowhammer E. citrinella and Corn Bunting Miliaria calandra). We analyse long-term, extensive data from the British Trust for Ornithology's Nest Record Scheme on breeding performance per breeding attempt with respect to farmland type (arable, grazing or mixed) and time (pre- and post-1975–76). The influence of habitat is investigated at two different scales: within the nesting territory and at the landscape level. Relationships between farmland type and (temporal changes in) breeding performance tended to be species-specific, but a few patterns were each common to some species. Improvements in breeding performance occurred across all three farmland types for four declining species. Grazing farmland seems to have deteriorated as breeding habitat for Linnet and arable/mixed farmland for Reed Bunting. Mixed farming at the territory scale supported better breeding performance for four species, three of which (Bullfinch, Yellowhammer and Corn Bunting) have declined concurrently with mixed farming. Pastoral landscapes supported better breeding performance for up to seven species, six of which have undergone large declines. Arable landscapes supported better breeding only for the stable or increasing Chaffinch and Greenfinch. Different relationships between farming regime and breeding performance were found at the two scales considered.     

Temporal patterns of seed use and availability in a guild of desert ants

ABSTRACT.

• 1 Temporal patterns of seed use were studied from late winter to autumn in three species of seed-harvesting ants in the Sonoran Desert. Measures of effective foraging activity, dietary niche breadth and dietary niche overlaps were obtained each month and were tested for correlation with estimates of the available seed resource.
• 2 Seeds were the only numerically important type of food in the diets of all species.
• 3 The ants partitioned the resource according to both seed species and seed size, although there was considerable overlap.
• 4 Pheidole xerophila had the smallest forager body size and is a specialist on small seeds because it harvested them in greater proportion than their rank in the soils and expanded its diet to larger seeds only when the abundance of small seeds declined.
• 5 When the abundance of the small seeds of Bouteloua barbata decreased, the middle-sized ant, Veromessor pergandei, showed a decrease in foraging activity, increase in niche breadth, and a decrease in overlap with P.xerophila.
• 6 Seed size preferences of V.pergandei did not vary seasonally, except that during the month of highest seed abundance, V.pergandei showed no size preference.
• 7 Pogonomyrmex rugosus was the largest ant; it preferred larger seeds and was inactive when small seeds were most abundant. Seasonal foraging activity and niche parameters were random in relation to seed abundance.
• 8 We suggest that nocturnal foraging by P.rugosus during the summer months was a response to interference with diurnal foraging by either predation frorn horned lizards or competition from V.pergandei.
• 9 Seasonal abundance of small seeds explains most of the seasonal foraging patterns of P.xerophila and V.pergandei. The summertime abundance of larger seeds during years of adequate precipitation may account for the seasonal activity patterns of P.rugosus.

     

Influence of temporal fluctuations in seed abundance on the foraging behaviour of harvester ants (Pogonomyrmex spp.) in the central Monte desert,Argentina

The simultaneous study of the temporal dynamics of foraging behaviour, diet and seed abundance is essential to assess the way in which resources affect the behaviour and ecology of harvester ants. Here, we evaluate how fluctuations in grass seed abundance during three consecutive growing seasons influenced the foraging behaviour and diet of the harvester ants Pogonomyrmex rastratus, P. mendozanus and P. inermis in the central Monte desert, Argentina. Seed abundance of the most consumed grasses varied greatly through ant activity season, and ants altered their foraging behaviour in response to those changes. Foragers spent more time travelling and searching for food, and their foraging trips took longer during the low seed availability season. Foraging distance was very similar among species and, contrary to our expectations, did not vary between seasons. Foraging success of P. rastratus and P. inermis increased during the high availability season. This matched the seasonal pattern of foraging activity, suggesting that colonies may detect seed abundance and regulate their foraging effort with the rate of forager success. Although grass seeds were the main item in the diet of the three species, P. mendozanus, and to a lesser extent P. rastratus, turned more generalist when grass seeds were scarce. In contrast, P. inermis showed a very narrow diet breadth, only harvesting grass seeds in both seasons. Our results indicate the relevance of seed availability on foraging behaviour of harvester ants, which should be taken into account when predicting and evaluating the effect of ants on seed resources as well as numerical responses of harvester ant populations to the temporal and spatial variations in grass seed abundance.     

The influence of food supply on foraging behaviour in a desert spider

We tested the alternative hypotheses that foraging effort will increase (energy maximizer model) or decrease (due to increased costs or risks) when food supply increased, using a Namib desert burrowing spider, Seothyra henscheli (Eresidae), which feeds mainly on ants. The web of S. henscheli has a simple geometrical configuration, comprising a horizontal mat on the sand surface, with a variable number of lobes lined with sticky silk. The sticky silk is renewed daily after being covered by wind-blown sand. In a field experiment, we supplemented the spiders' natural prey with one ant on each day that spiders had active webs and determined the response to an increase in prey. We compared the foraging activity and web geometry of prey-supplemented spiders to non-supplemented controls. We compared the same parameters in fooddeprived and supplemented spiders in captivity. The results support the costs of foraging hypothesis. Supplemented spiders reduced their foraging activity and web dimensions. They moulted at least once and grew rapidly, more than doubling their mass in 6 weeks. By contrast, food-deprived spiders increased foraging effort by enlarging the diameter of the capture web. We suggest that digestive constraints prevented supplemented spiders from fully utilizing the available prey. By reducing foraging activities on the surface, spiders in a prey-rich habitat can reduce the risk of predation. However, early maturation resulting from a higher growth rate provides no advantage to S. henscheli owing to the fact that the timing of mating and dispersal are fixed by climatic factors (wind and temperature). Instead, large female body size will increase fitness by increasing the investiment in young during the period of extended maternal care.     

Agricultural land-use and the spatial distribution of granivorous lowland farmland birds

Current agricultural practices are believed to have contributed to the declines of many farmland bird species, especially seed‐eaters, throughout Europe. We investigated associations between the spatial distribution of fourteen granivorous farmland bird species and agricultural land‐use in Britain, using breeding bird atlas data and national agricultural statistics. Analyses were spatially referenced by 10x10 km square and variation due to broad‐scale geography and spatial auto‐correlation was controlled for. Generalized linear modelling analyses were used to select models describing variation in distribution explained by the available land‐use variables. The results show that relationships between distribution and agriculture tend to be species‐specific, but that some general effects can be identified. Features of intensive arable farming including large areas of sugar beet, wheat and oilseed rape tended to be associated with low frequencies of occurrence for 9–11 species, while large areas of younger (re‐seeded) grassland and high sheep stocking densities were associated with low frequencies in pastoral farmland for up to 12 species. One key feature of lower intensity farming, the presence of larger areas of fallow land, was positively related to frequency index for up to 11 species. The proportion of barley sown in spring and agricultural diversity were each associated with a range of complex relationships with frequency index across species, probably reflecting combinations of positive influences and artefacts of scale and geography. A variable describing the heterogeneity of farmland (the extent to which it is a mix of arable and pastoral land‐use) was negatively related to frequency index for eight species, but other results suggested that farming which is mixed at a smaller spatial scale is widely beneficial. The results reveal relationships between agriculture and the occurrence of granivorous farmland bird species which suggest both hypotheses for the causes of population change and directions for management action. However, data on several key features of agricultural practice (such as pesticide use) were unavailable, so their effects could not be tested, and the effects of the variables included could not be separated from those of other factors which are subject to complex geographical variation. Experimental comparisons of the effects of land‐use at the farm scale are needed to investigate such confounded influences on farmland bird occurrence.     

Density‐mediated indirect interactions alter host foraging behaviour of parasitoids without altering foraging efficiency

1. Foraging decisions of parasitoids are influenced by host density via density‐mediated indirect interactions. However, in the parasitoid's environment, non‐suitable herbivores are also present. These non‐hosts also occur in different densities, which can affect a parasitoid's foraging behaviour. 2. The influence of non‐host densities can be expressed during the first phase of the foraging process, when parasitoids use plant volatiles to locate plants infested by their host. They may also play a role during the second phase, when parasitoids use infochemicals from the host and plant to locate, recognise and accept the host. 3. By using laboratory and field setups, it was studied whether the density of non‐host herbivores influences these two phases of the foraging behaviour of the parasitoid Cotesia glomerata as well as the parasitoid's efficiency to find its host, Pieris brassicae caterpillars. 4. The findings show that a high non‐host density, regardless of the species used, negatively affected parasitoid preference for host‐infested plants, but that the behaviour on the plant and the total host‐finding efficiency of the parasitoids were not influenced by non‐host density. 5. These results are discussed in the context of density‐mediated indirect interactions.     

Grass seed production in the central Monte desert during successive wet and dry years

In desert regions, water availability triggers primary production, which determines seed production, the composition and size of soil seed reserves and the abundance and behaviour of seed-eating animals. In the central Monte desert, large precipitation events (≥10 mm) account for a high proportion of growing season’s rainfall. Our first objective here was to assess whether and how timing and amount of seed production of C₃ and C₄ perennial grasses are linked to spring and summer precipitation pulses and to estimate the seasonal and year-to-year variability in seed production. Our second aim was to calculate grass seed production and compare it with seed requirements by granivorous animals to infer whether the animals can exert top-down effects on plant populations. Seed production of C₃ and C₄ species was triggered by significant spring and summer rainfall, respectively. Such distinct response may be associated with the effect of precipitation during flower development and seed set in both functional groups. In all species, seed production varied among years. Rainfall pulses in the summer triggered and positively affected the magnitude of seed production in most C₄ grasses. However, all perennial grasses were able to produce high amount of seeds even during a year subjected to extreme drought, suggesting that perenniality would allow these species to make large reproductive investment despite harsh environmental conditions. The comparative assessment of seed production and consumer demands suggests that it is unlikely that granivory exerts a top−down control on grasses in the Monte desert.     

Optimal foraging and beyond: how starlings cope with changes in food availability

Foraging adaptations include behavioral and physiological responses, but most optimal foraging models deal exclusively with behavioral decision variables, taking other dimensions as constraints. To overcome this limitation, we measured behavioral and physiological responses of European starlings Sturnus vulgaris to changes in food availability in a laboratory environment. The birds lived in a closed economy with a choice of two foraging modes (flying and walking) and were observed under two treatments (hard and easy) that differed in the work required to obtain food. Comparing the hard with the easy treatment, we found the following differences. In the hard treatment, daily amount of work was higher, but daily intake was lower. Even though work was greater, total daily expenditure was smaller, partly because overnight metabolism was lower. Body mass was lower, but daily oscillation in body mass did not differ. Feces' caloric density was lower, indicating greater food utilization. Energy expenditure rate expressed as multiples of basal metabolic rate (BMR) increased during the working period from 3.5 x BMR (easy) to 5.2 x BMR (hard), but over the 24-h period, it was close to 2.4 x BMR in both treatments. We also found that rate of expenditure during flight was very high in both treatments (52.3 W in easy and 45.5 W in hard), as expected for short (as opposed to cruising) flights. The relative preferences between walking and flying were incompatible with maximizing the ratio of energy gains per unit of expenditure (efficiency) but compatible with maximizing net gain per unit of time during the foraging cycle (net rate). Neither currency explained the results when nonforaging time was included. Time was not a direct constraint: the birds rested more than 90% of the time in both treatments. Understanding this complex picture requires reasoning with ecological, physiological, and cognitive arguments. We defend the role of optimality as an appropriate tool to guide this integrative perspective.     

Influence of environment structure and food availability on the foraging behaviour of the laboratory rat

According to the optimal foraging theory, an animal is expected to enter into a given activity depending on associated costs and benefits. In line with this assumption, numerous studies have suggested that energetic reward is balanced by predation risk in foraging decisions. Therefore, the use of information about indirect cues of predation risk such as physical structure (e.g. cover, escape substrate) can give individuals a selective advantage. We studied foraging behaviour in the laboratory rat in an experimental maze; it allowed us to vary two environmental parameters: food availability and physical structure. In a first experiment, rats were offered a choice between two areas only differing in cover density. In a second experiment, the two areas only differed in food density. In a third experiment, we crossed both parameters. Our results showed that high “cover” patch was preferentially exploited (experiment 1) and that rats foraged more in the high food density patch (experiment 2). The last experiment showed that rats partially trade-off between cover density and food availability, even if the safest area was still preferred. Therefore, we suggest that foraging decisions depend primarily on safety needs, rather than food availability, at least when animals are not severely food-deprived.     

Fruits foraging patterns and seed dispersal effect of frugivorous birds on Hippophae rhamnoides sinensis

Behaviors of 18 species of birds eating fruits of Hippophae rhamnoides spp.sinensis were observed from September 2003 to March 2004.Their foraging patterns were found to be very different and Can be divided into five classes:(1)direct swallowing the fruits on crown of the shrubs and sometimes regurgitating seeds soon after;(2)carrying the fruits to their perching sites and swallowing;(3)pecking the fruits from the shrubs to the ground,eating pulp and seeds but leaving pericarp;(4)pecking through the pericarp,eating pulp and leaving pericarp and seeds;(5)pecking through the pericarp on the top of fruits,and only eating seeds.These foraging patterns have different effects on seed dispersal of H.rhamnoides spp.sinensis.The germination experiment of three groups of seeds(seeds from feces,dry fruits and extracted seeds from dry fruits)was carried out.Although ingestion processes of birds had some adverse effects on the seed germination of H.rhamnoides spp.sinensis,the seeds from feces still have a relatively higher germination ratio.H.rhamnoides spp.sinensis provides food to a variety of frugivorous birds.and the birds disperse its seeds.Thus,a mutually beneficial relationship between the bird and the seed is formed.     

鸟类取食中国沙棘果实的方式及其对种子的传播作用

2003年9月～2004年3月,定期观察取食中国沙棘果实的鸟类及其取食方式。共记录取食中国沙棘果实的鸟类18种,其取食果实的方式主要有：1)直接在树冠上吞食果实,有时候在吞食后将种子呕出;2)将果实从树上衔走后,在栖息处吞食或啄食;3)将果实啄落至地面,然后取食果肉和种子,留下果皮;4)啄破果皮,吸食果肉,留下果皮及种子;5)从顶端将果皮啄破后,仅取食里面的种子。不同的取食方式决定了它们对沙棘种子的传播作用不同。收集鸟粪便中的种子与完整的干果实及人工剥离果肉和内果皮的种子做萌发对比实验,结果表明,鸟类消化道过程对种子的萌发有一定影响,但萌发率仍较高。沙棘为多种鸟类提供食物,而鸟类则为沙棘传播种子,它们之间形成一种互利关系。     

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 of inducing 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-term apparent 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 density and 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.     

Litter decomposition in a sandy Monte desert of western Argentina: Influences of vegetation patches and summer rainfall

Abstract: We tested the hypothesis that shrub canopies interact with monthly rain pulses to control litter decomposition in a sandy Monte desert, in Argentina. We assessed (i) the potential for litter decomposition of soils beneath the canopies of two dominant shrub species (Larrea divaricata and Bulnesia retama, Zygophyllaceae R. Br.) and from bare‐ground microsites or ‘openings’; (ii) litter decomposition at different spatial patches over the summer rainy season; and (iii) the interaction between vegetation patches and monthly rain pulses on short‐term litter decomposition, or decomposition pulses. In a greenhouse experiment, we found buried litter decomposition to be higher in soils from under the canopies of a dominant shrub species compared with soils from openings and sterilized controls. This, and higher nutrient concentration under shrub soils, suggest undercanopy soils may support a microbial community capable of decomposing litter at higher rates than soils in bare openings. However, ?eld trials showed that shrub patches did not affect leaf litter decomposition over the rainy season, at least for short periods. We found an interaction between shrub patches and incubation time at the end of the ?eld experiment, with higher litter decomposition rates under B. retama canopies. In a monthly ?eld experiment, we found monthly rain pulses signi?cantly explained decomposition pulses, irrespective of patch type. Our ?ndings support the hypothesis that shrub soils have a greater potential for litter decomposition, but this is not directly translated to the ?eld possibly due to interactions with abiotic factors. Rain pulses create conditions for decomposition pulses to occur at shorter time scales, whereas rainfall may interact with a dominant shrub undercanopy to control litter mass loss over longer time scales.     

Visual perception and social foraging in birds

Birds gather information about their environment mainly through vision by scanning their surroundings. Many prevalent models of social foraging assume that foraging and scanning are mutually exclusive. Although this assumption is valid for birds with narrow visual fields, these models have also been applied to species with wide fields. In fact, available models do not make precise predictions for birds with large visual fields, in which the head-up, head-down dichotomy is not accurate and, moreover, do not consider the effects of detection distance and limited attention. Studies of how different types of visual information are acquired as a function of body posture and of how information flows within flocks offer new insights into the costs and benefits of living in groups.     

Prey selection and foraging performance of breeding Great Tits Parus major in relation to food availability

We studied the nestling diet and the foraging performance of Great Tits in relation to prey abundance in the field. Numerous experimental studies present data on foraging decisions in captive Great Tits. Little is, however, known about prey selection in the field in relation to the food available and the consequences this has for the food delivery rate to nestlings. Since the foraging performance of the parents is one of the main determinants of fledging weight and juvenile survival, foraging behaviour is an important part of Great Tit reproduction. During the early breeding season up to 75% of the prey biomass delivered to the nestlings were spiders, which is in contrast with other studies. Only when caterpillars reached a size of 10–12 mg (approximately the average size of the spiders caught at that time) did the Great Tits change their preferences and 80–90% of the delivered prey masses were caterpillars, as reported by other authors. This 'switching' between prey occurred within a few days. It was not related to the changes in abundance but to size of caterpillars. The rate at which caterpillars were delivered to the nestlings (in mg/nestling/h) was strongly correlated with the caterpillar biomass available (in mg/m of branches) and nestling growth rate was significantly influenced by the mass of available caterpillars. The results provide evidence why perfect timing of breeding is so important for the Great Tit, and contribute to the understanding of the causal link between food supply, growth and breeding success.     

Uncoupling the effects of seed predation and seed dispersal by granivorous ants on plant population dynamics

Secondary seed dispersal is an important plant-animal interaction, which is central to understanding plant population and community dynamics. Very little information is still available on the effects of dispersal on plant demography and, particularly, for ant-seed dispersal interactions. As many other interactions, seed dispersal by animals involves costs (seed predation) and benefits (seed dispersal), the balance of which determines the outcome of the interaction. Separate quantification of each of them is essential in order to understand the effects of this interaction. To address this issue, we have successfully separated and analyzed the costs and benefits of seed dispersal by seed-harvesting ants on the plant population dynamics of three shrub species with different traits. To that aim a stochastic, spatially-explicit individually-based simulation model has been implemented based on actual data sets. The results from our simulation model agree with theoretical models of plant response dependent on seed dispersal, for one plant species, and ant-mediated seed predation, for another one. In these cases, model predictions were close to the observed values at field. Nonetheless, these ecological processes did not affect in anyway a third species, for which the model predictions were far from the observed values. This indicates that the balance between costs and benefits associated to secondary seed dispersal is clearly related to specific traits. This study is one of the first works that analyze tradeoffs of secondary seed dispersal on plant population dynamics, by disentangling the effects of related costs and benefits. We suggest analyzing the effects of interactions on population dynamics as opposed to merely analyzing the partners and their interaction strength.