Frequency-dependent food selection by arthropods: a review

We review published studies in which arthropods were presented with a choice of food types at a range of relative densities. Twenty-four of 28 studies showed frequency-independent preference; 11 studies also showed frequency-dependent preference. Both pro- and anti-apostatic selection were found. In the light of these studies we discuss possible behavioural mechanisms which produce such selection; the potential benefits to the predators and the consequences of such selection for diversity both within and between species.     

Red-Tailed Guenons (Cercopithecus ascanius) and Strychnos mitis: Evidence for Plant Benefits Beyond Seed Dispersal

I report data collected on red-tailed guenon (Cercopithecus ascanius schmidti) fruit processing behaviors between June 1993 and April 1994. Red-tailed guenons consumed the fruit of Strychnos mitis in 542 of 2,930 fruit-eating events (FEEs). The monkeys spat out cleaned seeds of Strychnos mitis in a majority of these records (477/542; 88%); seeds were occasionally swallowed whole, but only when pulp was unripe (69/542; 12%). In 83% of the FEEs on Strychnos, the red-tailed guenons spat out seeds within 10 m of the removal site; they typically stayed in the same tree while processing fruit, and in 56% of the FEEs, they moved <1 m before spitting seeds. I monitored spat seeds to evaluate the impact of monkey fruit processing on seed fate. Results indicate that 83% of seeds spat out by the red-tailed guenons germinated, while only 12% of unprocessed seeds survived to germination (p < 0.01). Of the processed seeds that germinated, 60% survived to germination and seedling establishment, while only 5% of unprocessed seeds survived to seedling establishment (p < 0.01). Unprocessed seeds were also more likely to be attacked by seed predators (p < 0.01) and fungus (p < 0.01). Although there is generally high mortality in seeds/seedlings, mature trees of Strychnos mitis are found in groves of adults, under which dense populations of seedlings and saplings can occur. These data suggest that Strychnos mitis does not conform to expectations of the Janzen-Connell model of seed escape from parent trees. Instead, I suggest that by removing pulp, a process that results in a reduction of fungal pathogen attack, red-tailed guenons positively effect the seed survivorship of Strychnos mitis. Although this effect has been observed in pulp-cleaning ant species, it is a hitherto undescribed effect of primates on their fruit resources.     

A surplus of seeds: High rates of post-dispersal seed predation in a flooded grassland in monsoonal Australia

Abstract Oryza meridionalis is an annual emergent wetland grass which produces between 26 and 260 kg seeds ha^-1 annually. Seed shed occurs at the end of the wet season, when the plains are usually still partially flooded. The juvenile recruitment of key native vertebrate species, such as the Magpie Goose (Anseranus semipalmata) and the Dusky Plains Rat (Rattus colletti), coincides with seed shed. This study investigated predation of O. meridionalis seeds at two sites on the South Alligator River floodplain in monsoonal Australia. The effects of inundation and the presence of a background density of seeds on seed removal were investigated by stratified sampling with respect to position down the topographic slope, to include the ‘dry’ floodplain margin, ‘damp’Oryza zone, and ‘flooded’Oryza zone. The effect of seed lot size on the proportion of seed removed was also investigated, and exclosures were used to identify the principal predator group. The proportion of seeds removed was not affected by the presence/absence of a background of seeds nor the number of seeds placed in experimental ‘lots’. The majority of seeds (75%) was consumed by vertebrate predators (most likely the abundant Dusky Plains Rat). Inundation afforded some protection from predation. Despite high losses of seeds exposed to predators, O. meridionalis is an abundant and widespread species on these floodplains, possibly because of the protection from predation afforded by inundation to those seeds which are shed into the water column. It is likely that there is a complex interaction between topography, rainfall and predator and prey relationships, which ultimately determines the importance of seed removal for the maintenance of populations of O. meridionalis. These high losses of seeds to predation have implications for wetland rehabilitation where seed broadcasting is proposed.     

Perceptual Processes and the Maintenance of Polymorphism Through Frequency-dependent Predation

One of the key challenges of both ecology and evolutionary biology is to understand the mechanisms that maintain diversity. Negative frequency-dependent selection is a powerful mechanism for maintaining variation in the population as well as species diversity in the community. There are a number of studies showing that this type of selection, where individuals of a rare type (i.e. a rare morph or a rare species) experience higher survival than those of more common type(s). However, it is still not clear how frequency-dependent selection operates. Search image formation has been invoked as a possible, proximate explanation. Although the conceptual link between search image and frequency-dependent predation is often assumed in ecological and evolutionary studies, a review of the literature reveals a paucity of evidence demonstrating the occurrence of both in a natural predator-prey system. Advances in the field of psychology strongly support the existence of search image, yet these findings are not fully recognized in the realm of ecology and evolutionary biology, in part, we feel because of confusion and inconsistencies in terminology. Here we try to simplify the language, clarify the advances in the study of frequency-dependent predation and search image, and suggest avenues for future research. We feel that the investigations of both proximate (perceptual mechanisms) and ultimate (pattern of predation) processes are necessary to fully understand the importance of individual behavioural processes for mediating evolutionary and ecological diversity.Co-ordinating editor: O. Leimar     

Selection by wild birds on artificial dimorphic prey on varied backgrounds

Our aim was to test the effects of prey frequency and background composition on selection by free-ranging birds. We did three series of experiments with populations of grey and orange pastry prey scattered among coloured stones that made the prey either conspicuous or inconspicuous. Series 1 tested whether the predicted equilibrium frequency of the two prey types was influenced by the frequency of matching grey and orange stones. Birds at a single site were given a random sequence of combinations of prey frequency and stone frequency. Selection was dependent on background and the effect of prey frequency also varied with background. In series 2, we explored the frequency-independent effect of background: birds at five sites were given equal numbers of the two prey in three frequencies of matching stones and two of non-matching. There was a higher risk of predation for prey that matched rarer stones. In series 3 we attempted to measure, at a single site, the actual equilibrium prey frequencies in three different backgrounds: two extreme stone frequencies and one intermediate. Each experiment started with a population of equal numbers of grey and orange prey. After half the prey had been eaten we calculated the frequencies of the survivors and presented a new population of the original size but with the new prey frequencies; each experiment ran for 25 such 'generations'. The results suggested that at equilibrium the commoner 'morph' was the one that resembled the commoner colour of stone. Overall, our findings support the idea that visual selection can result in morph frequencies becoming related to the proportions of their matching background components and that this equilibrium will 'track' temporal or spatial changes in the background.     

Missing and forbidden links in mutualistic networks

Ecological networks are complexes of interacting species, but not all potential links among species are realized. Unobserved links are either missing or forbidden. Missing links exist, but require more sampling or alternative ways of detection to be verified. Forbidden links remain unobservable, irrespective of sampling effort. They are caused by linkage constraints. We studied one Arctic pollination network and two Mediterranean seed-dispersal networks. In the first, for example, we recorded flower-visit links for one full season, arranged data in an interaction matrix and got a connectance C of 15 per cent. Interaction accumulation curves documented our sampling of interactions through observation of visits to be robust. Then, we included data on pollen from the body surface of flower visitors as an additional link ‘currency’. This resulted in 98 new links, missing from the visitation data. Thus, the combined visit–pollen matrix got an increased C of 20 per cent. For the three networks, C ranged from 20 to 52 per cent, and thus the percentage of unobserved links (100 − C) was 48 to 80 per cent; these were assumed forbidden because of linkage constraints and not missing because of under-sampling. Phenological uncoupling (i.e. non-overlapping phenophases between interacting mutualists) is one kind of constraint, and it explained 22 to 28 per cent of all possible, but unobserved links. Increasing phenophase overlap between species increased link probability, but extensive overlaps were required to achieve a high probability. Other kinds of constraint, such as size mismatch and accessibility limitations, are briefly addressed.     

A seed predator drives the evolution of a seed dispersal mutualism

Although antagonists are hypothesized to impede the evolution of mutualisms, they may simultaneously exert selection favouring the evolution of alternative mutualistic interactions. We found that increases in limber pine (Pinus flexilis) seed defences arising from selection exerted by a pre-dispersal seed predator (red squirrel Tamiasciurus hudsonicus) reduced the efficacy of limber pine's primary seed disperser (Clark's nutcracker Nucifraga columbiana) while enhancing seed dispersal by ground-foraging scatter-hoarding rodents (Peromyscus). Thus, there is a shift from relying on primary seed dispersal by birds in areas without red squirrels, to an increasing reliance on secondary seed dispersal by scatter-hoarding rodents in areas with red squirrels. Seed predators can therefore drive the evolution of seed defences, which in turn favour alternative seed dispersal mutualisms that lead to major changes in the mode of seed dispersal. Given that adaptive evolution in response to antagonists frequently impedes one kind of mutualistic interaction, the evolution of alternative mutualistic interactions may be a common by-product.     

帽儿山国家森林公园30年前后植物区系比较研究

该研究采用样方结合样线调查的方法,并综合文献资料,于1983年和2013年2次对帽儿山国家森林公园植物种类进行实地调查,并进行了区系特征的对比分析,为评估帽儿山国家森林公园植物保护状况提供依据。结果表明:(1)2013年研究区的植物科数量和组成与1983年相同,但2013年比1983年少20属60种,不同等级科组成差异较小,而不同等级属差异较大。(2)优势科组成差异较小,但2013年兰科(Orchidacea)由优势科变成了非优势科,而藜科(Chenopodiaceae)和松科(Pinaceae)由非优势科变成优势科;优势属组成差异较大,2013年比1983年少了8个优势属。(3)2次调查表征科组成变化较大,但表征属组成差异较小,2013年比1983年少了2个表征属。(4)1983年和2013年科、属、种的分布型均以温带性分布为主;科的分布型完全相同,属的分布型2013年比1983年少了地中海区-西亚至中亚分布型;种的分布型,2013年比1983年多了亚洲-北美-北极分布。研究认为,30年间帽儿山国家森林公园的植物区系构成基本稳定,说明对这一区域的植物保护总体有效,但属和种的数量明显减少,30年间物种多样性显著降低,尤其是某些重要的科,如兰科植物的减少,也说明植物保护现状依然不容乐观。     

Massive polymorphism and natural selection in Donacilla cornea (Poli, 1791) (Bivalvia: Mesodesmatidae)

We describe a massive polymorphism for shell colour and pattern in Donacilla cornea , a shallow-burrowing bivalve which inhabits the mid-littoral of sandy and stony beaches in southern and western Europe. Of the four sites studied, the one where the substrate was a mixture of small stones and sand contained the highest densities of Donacilla. At this site the densities of stones and animals and the frequencies of shell and stone colour phenotypes are both highly correlated. Although the four populations sampled comprise a similar range of phenotypes, the frequencies of most morphs and population diversity indices differ significantly between sites. At two sites the similar range of colour and pattern variation between shells and substrate suggests selection for crypsis and masquerade. On the other hand, at two other sites, one or two common phenotypes (and a number of rare ones at all sites) fail to mimic any aspect of the background. We therefore suggest that if frequency-dependent selection by predators is responsible for the maintenance of polymorphism in Donacilla , then the mechanisms involved include crypsis and masquerade in some cases and selective advantages from being different in others.