Seed predation

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A strawberry aggregate accessory fruit damaged by a mouse eating the seeds (achenes). Aardbei muizenschade.jpg
A strawberry aggregate accessory fruit damaged by a mouse eating the seeds (achenes).

Seed predation, often referred to as granivory, is a type of plant-animal interaction in which granivores (seed predators) feed on the seeds of plants as a main or exclusive food source, [1] in many cases leaving the seeds damaged and not viable. Granivores are found across many families of vertebrates (especially mammals and birds) as well as invertebrates (mainly insects); [2] thus, seed predation occurs in virtually all terrestrial ecosystems. Seed predation is commonly divided into two distinctive temporal categories, pre-dispersal and post-dispersal predation, which affect the fitness of the parental plant and the dispersed offspring (the seed), respectively. Mitigating pre- and post-dispersal predation may involve different strategies. To counter seed predation, plants have evolved both physical defenses (e.g. shape and toughness of the seed coat) and chemical defenses (secondary compounds such as tannins and alkaloids). However, as plants have evolved seed defenses, seed predators have adapted to plant defenses (e.g., ability to detoxify chemical compounds). Thus, many interesting examples of coevolution arise from this dynamic relationship.

Contents

Seeds and their defenses

Mouse eating seeds Mouse eating seeds.jpg
Mouse eating seeds

Plant seeds are important sources of nutrition for animals across most ecosystems. Seeds contain food storage organs (e.g., endosperm) that provide nutrients to the developing plant embryo (cotyledon). This makes seeds an attractive food source for animals because they are a highly concentrated and localized nutrient source in relation to other plant parts.

Seeds of many plants have evolved a variety of defenses to deter predation. Seeds are often contained inside protective structures or fruit pulp that encapsulate seeds until they are ripe. Other physical defenses include spines, hairs, fibrous seed coats and hard endosperm. Seeds, especially in arid areas, may have a mucilaginous seed coat that can glue soil to seed hiding it from granivores. [3]

Some seeds have evolved strong anti-herbivore chemical compounds. In contrast to physical defenses, chemical seed defenses deter consumption using chemicals that are toxic or distasteful to granivores or that inhibit the digestibility of the seed. These chemicals include toxic non-protein amino acids, cyanogenic glycosides, protease and amylase inhibitors, and phytohemaglutinins. [1] Plants may face trade-offs between allocation toward defenses and the size and number of seeds produced.

Plants may reduce the severity of seed predation by making seeds spatially or temporally scarce to granivores. Seed dispersal away from the parent plant is hypothesized to reduce the severity of seed predation. [4] [5] Seed masting is an example of how plant populations are able to temporally regulate the severity of seed predation. Masting refers to a concerted abundance of seed production followed by a period of paucity. This strategy has the potential to regulate the size of the population of seed predators.

Seed predation vs. seed dispersal

Adaptations to defend seeds against predation can impact seeds' ability to germinate and disperse. Thus anti-predator adaptations often occur in a suite of adaptations for a particular seed life history. For example, chili plants selectively deter mammal seed predators and fungi using capsaicin, which does not deter bird seed dispersers [6] [7] because bird taste receptors do not bind with capsaicin. Chili seeds in turn have higher survival if they pass through a bird's stomach than if they fall to the ground. [8]

Pre- and post-dispersal

Seed predation can occur both before and after seed dispersal. [9]

Pre-dispersal

Pre-dispersal seed predation takes place when seeds are removed from the parent plant before dispersal, and it has been most often reported in invertebrates, birds, and in granivorous rodents that clip fruits directly from trees and herbaceous plants. Post-dispersal seed predation arises once seeds have been released from the parent plant. Birds, rodents, and ants are known to be among the most pervasive postdispersal seed predators. Furthermore, postdispersal seed predation can take place at two contrasting stages: predation on the "seed rain" and predation on the "seed bank". Whereas predation on the seed rain occurs when animals prey on released seeds usually flush with the ground surface, predation on the seed bank takes place after seeds have been incorporated deeply into the soil. [1] Nevertheless, there are important vertebrate pre-dispersal predators, especially birds and small mammals.

Post-dispersal

Post-dispersal seed predation is extremely common in virtually all ecosystems. Given the heterogeneity in both resource type (seeds from different species), quality (seeds of different ages and/or different status of integrity or decomposition) and location (seeds are scattered and hidden in the environment), most post-dispersal predators have generalist habits. [1] These predators belong to a diverse array of animals, such as ants, beetles, crabs, fish, rodents and birds. The assemblage of post-dispersal seed predators varies considerably among ecosystems. [1] A dispersed seed is the first independent life stage of a plant, thus post-dispersal seed predation is the first potential mortality event and one of the first biotic interactions in a plant's life cycle.

Differences

Both pre- and post-dispersal seed predation are common. Pre-dispersal predators differ from post-dispersal predators in most often being specialists, adapted to clustered resources (on the plant). They use specific cues like plant chemistry (volatile compounds), color, and size to locate seeds, and their short life cycles often match the production of seeds by the host plant. Insect groups containing many pre-dispersal seed predators are Coleoptera, Hemiptera, Hymenoptera and Lepidoptera. [1]

Effects on plant demography

The complex relationship between seed predation and plant demography is an important topic of plant-animal interactive studies. Plant population structure and size over time is closely associated with the effectiveness at which seed predators locate, consume, and disperse seeds. In many cases this relationship depends on the type of seed predator (specialist vs. generalist) or the particular habitat in which the interaction is taking place. The role of seed predation on plant demography may be either detrimental or in particular cases actually beneficial to plant populations[ citation needed ].

The Janzen-Connell model concerns how seed density and survival respond to distance from the parent tree and differential rates of seed predation. Seed density is hypothesized to decrease as distance from the parent tree increases. Where seeds are most abundant under the parent tree, seed predation is predicted to be at its highest. As distance from the parent tree increases, seed abundance and thus seed predation are predicted to decrease as seed survival increases. [4] [5]

The degree to which seed predation influences plant populations may vary by whether a plant species is safe site limited or seed limited. If a population is safe site limited it is likely that seed predation will have little impact to the success of the population. In safe site limited populations increased seed abundance does not translate into increased seedling recruitment. However, if a population is seed limited, seed predation has a better chance of negatively affecting the plant population by decreasing seedling recruitment. Maron and Simms [10] found both safe site limited and seed limited populations depending on the habitat in which the seed predation was taking place. In dune habitats seed predators (deer mice) were limiting seedling recruitment in the population, thus negatively affecting the population. However, in grassland habitat the seed predator had little effect on the plant population because it was safe site limited.

In many cases seed predators support plant populations by dispersing seeds away from the parent plant, in effect supporting gene flow between populations. Other seed predators collect seeds and then store or cache them for later consumption. [11] In the case that the seed predator is unable to locate the buried or hidden seed there is a chance that it will later germinate and grow, supporting the species dispersal. Generalist (vertebrate) seed predators may also aid the plant in other indirect ways, for instance by inducing top-down control on host-specific seed predators (termed "intra-guild predation"), and as such negating Janzen-Connell type effects and so benefiting the plant in competition with other plant species. [12]

See also

Related Research Articles

<span class="mw-page-title-main">Food</span> Substances consumed for nutrition

Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin and contains essential nutrients such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is ingested by an organism and assimilated by the organism's cells to provide energy, maintain life, or stimulate growth. Different species of animals have different feeding behaviours that satisfy the needs of their metabolisms and have evolved to fill a specific ecological niche within specific geographical contexts.

<span class="mw-page-title-main">Herbivore</span> Organism that eats mostly or exclusively plant material

A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthparts adapted to rasping or grinding. Horses and other herbivores have wide flat teeth that are adapted to grinding grass, tree bark, and other tough plant material.

<span class="mw-page-title-main">Predation</span> Biological interaction where a predator kills and eats a prey organism

Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation and parasitoidism. It is distinct from scavenging on dead prey, though many predators also scavenge; it overlaps with herbivory, as seed predators and destructive frugivores are predators.

<span class="mw-page-title-main">Biological interaction</span> Effect that organisms have on other organisms

In ecology, a biological interaction is the effect that a pair of organisms living together in a community have on each other. They can be either of the same species, or of different species. These effects may be short-term, or long-term, both often strongly influence the adaptation and evolution of the species involved. Biological interactions range from mutualism, beneficial to both partners, to competition, harmful to both partners. Interactions can be direct when physical contact is established or indirect, through intermediaries such as shared resources, territories, ecological services, metabolic waste, toxins or growth inhibitors. This type of relationship can be shown by net effect based on individual effects on both organisms arising out of relationship.

<span class="mw-page-title-main">Frugivore</span> Organism that eats mostly fruit

A frugivore is an animal that thrives mostly on raw fruits or succulent fruit-like produce of plants such as roots, shoots, nuts and seeds. Approximately 20% of mammalian herbivores eat fruit. Frugivores are highly dependent on the abundance and nutritional composition of fruits. Frugivores can benefit or hinder fruit-producing plants by either dispersing or destroying their seeds through digestion. When both the fruit-producing plant and the frugivore benefit by fruit-eating behavior the interaction is a form of mutualism.

<span class="mw-page-title-main">Biological dispersal</span> Movement of individuals from their birth site to a breeding site

Biological dispersal refers to both the movement of individuals from their birth site to their breeding site, as well as the movement from one breeding site to another . Dispersal is also used to describe the movement of propagules such as seeds and spores. Technically, dispersal is defined as any movement that has the potential to lead to gene flow. The act of dispersal involves three phases: departure, transfer, settlement and there are different fitness costs and benefits associated with each of these phases. Through simply moving from one habitat patch to another, the dispersal of an individual has consequences not only for individual fitness, but also for population dynamics, population genetics, and species distribution. Understanding dispersal and the consequences both for evolutionary strategies at a species level, and for processes at an ecosystem level, requires understanding on the type of dispersal, the dispersal range of a given species, and the dispersal mechanisms involved. Biological dispersal can be correlated to population density. The range of variations of a species' location determines expansion range.

<span class="mw-page-title-main">Anti-predator adaptation</span> Defensive feature of prey for selective advantage

Anti-predator adaptations are mechanisms developed through evolution that assist prey organisms in their constant struggle against predators. Throughout the animal kingdom, adaptations have evolved for every stage of this struggle, namely by avoiding detection, warding off attack, fighting back, or escaping when caught.

<span class="mw-page-title-main">Seed dispersal</span> Movement or transport of seeds away from the parent plant

In spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors, such as the wind, and living (biotic) vectors such as birds. Seeds can be dispersed away from the parent plant individually or collectively, as well as dispersed in both space and time. The patterns of seed dispersal are determined in large part by the dispersal mechanism and this has important implications for the demographic and genetic structure of plant populations, as well as migration patterns and species interactions. There are five main modes of seed dispersal: gravity, wind, ballistic, water, and by animals. Some plants are serotinous and only disperse their seeds in response to an environmental stimulus. These modes are typically inferred based on adaptations, such as wings or fleshy fruit. However, this simplified view may ignore complexity in dispersal. Plants can disperse via modes without possessing the typical associated adaptations and plant traits may be multifunctional.

<span class="mw-page-title-main">Harvester ant</span> Common name for several different ants

Harvester ant, also known as harvesting ant, is a common name for any of the species or genera of ants that collect seeds, or mushrooms as in the case of Euprenolepis procera, which are stored in the nest in communal chambers called granaries. They are also referred to as agricultural ants. Seed harvesting by some desert ants is an adaptation to the lack of typical ant resources such as prey or honeydew from hemipterans. Harvester ants increase seed dispersal and protection, and provide nutrients that increase seedling survival of the desert plants. In addition, ants provide soil aeration through the creation of galleries and chambers, mix deep and upper layers of soil, and incorporate organic refuse into the soil.

<span class="mw-page-title-main">Myrmecochory</span> Seed dispersal by ants

Myrmecochory ( ; from Ancient Greek: μύρμηξ, romanized: mýrmēks and χορεία khoreíā is seed dispersal by ants, an ecologically significant ant–plant interaction with worldwide distribution. Most myrmecochorous plants produce seeds with elaiosomes, a term encompassing various external appendages or "food bodies" rich in lipids, amino acids, or other nutrients that are attractive to ants. The seed with its attached elaiosome is collectively known as a diaspore. Seed dispersal by ants is typically accomplished when foraging workers carry diaspores back to the ant colony, after which the elaiosome is removed or fed directly to ant larvae. Once the elaiosome is consumed, the seed is usually discarded in underground middens or ejected from the nest. Although diaspores are seldom distributed far from the parent plant, myrmecochores also benefit from this predominantly mutualistic interaction through dispersal to favourable locations for germination, as well as escape from seed predation.

Marine larval ecology is the study of the factors influencing dispersing larvae, which many marine invertebrates and fishes have. Marine animals with a larva typically release many larvae into the water column, where the larvae develop before metamorphosing into adults.

<span class="mw-page-title-main">Community (ecology)</span> Associated populations of species in a given area

In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, biological community, ecological community, or life assemblage. The term community has a variety of uses. In its simplest form it refers to groups of organisms in a specific place or time, for example, "the fish community of Lake Ontario before industrialization".

<span class="mw-page-title-main">Defaunation</span> Loss or extinctions of animals in the forests

Defaunation is the global, local, or functional extinction of animal populations or species from ecological communities. The growth of the human population, combined with advances in harvesting technologies, has led to more intense and efficient exploitation of the environment. This has resulted in the depletion of large vertebrates from ecological communities, creating what has been termed "empty forest". Defaunation differs from extinction; it includes both the disappearance of species and declines in abundance. Defaunation effects were first implied at the Symposium of Plant-Animal Interactions at the University of Campinas, Brazil in 1988 in the context of Neotropical forests. Since then, the term has gained broader usage in conservation biology as a global phenomenon.

<span class="mw-page-title-main">Dispersal vector</span> Transporters of biological dispersal units

A dispersal vector is an agent of biological dispersal that moves a dispersal unit, or organism, away from its birth population to another location or population in which the individual will reproduce. These dispersal units can range from pollen to seeds to fungi to entire organisms.

The Janzen–Connell hypothesis is a well-known hypothesis for the maintenance of high species biodiversity in the tropics. It was published independently in the early 1970s by Daniel Janzen, who focused on tropical trees, and Joseph Connell who discussed trees and marine invertebrates. According to their hypothesis, host-specific herbivores, pathogens, or other natural enemies make the areas near a parent tree inhospitable for the survival of seeds or seedlings. These natural enemies are referred to as 'distance-responsive predators' if they kill seeds or seedlings near the parent tree, or 'density-dependent predators' if they kill seeds or seedlings where they are most abundant. Such predators can prevent any one species from dominating the landscape, because if that species is too common, there will be few safe places for its seedlings to survive. Both Janzen and Connell originally proposed that for natural enemies to increase local diversity, they must be host-specific and relatively immobile, such that they disproportionately reduce the density of the more locally common tree species. This prevents any one species from becoming dominant and excluding other species through competition, allowing more species to coexist in small areas. This can be classified as a stabilizing mechanism.

Consumer–resource interactions are the core motif of ecological food chains or food webs, and are an umbrella term for a variety of more specialized types of biological species interactions including prey-predator, host-parasite, plant-herbivore and victim-exploiter systems. These kinds of interactions have been studied and modeled by population ecologists for nearly a century. Species at the bottom of the food chain, such as algae and other autotrophs, consume non-biological resources, such as minerals and nutrients of various kinds, and they derive their energy from light (photons) or chemical sources. Species higher up in the food chain survive by consuming other species and can be classified by what they eat and how they obtain or find their food.

<span class="mw-page-title-main">Evolutionary anachronism</span> Attributes of living species that arose due to coevolution with other now-extinct species

Evolutionary anachronism, also known as "ecological anachronism", is a term initially referring to attributes of native plant species that seemed best explained as having been favorably selected in the past due to their coevolution with plant-eating megafauna that are now extinct. Diminished effectiveness and distance of seed dispersal by fruit-eating mammals inhabiting the same ecosystems today suggest maladaptation. Maladaptation of such fruiting plants will intensify as ongoing climate change shifts the physical and ecological conditions within their current geographic range.

Flowering synchrony is the amount of overlap between flowering periods of plants in their mating season compared to what would be expected to occur randomly under given environmental conditions. A population which is flowering synchronously has more plants flowering at the same time than would be expected to occur randomly. A population which is flowering asynchronously has fewer plants flowering at the same time than would be expected randomly. Flowering synchrony can describe synchrony of flowering periods within a year, across years, and across species in a community. There are fitness benefits and disadvantages to synchronized flowering, and it is a widespread phenomenon across pollination syndromes.

Diplochory, also known as “secondary dispersal”, “indirect dispersal” or "two-phase dispersal", is a seed dispersal mechanism in which a plant's seed is moved sequentially by more than one dispersal mechanism or vector. The significance of the multiple dispersal steps on the plant fitness and population dynamics depends on the type of dispersers involved. In many cases, secondary seed dispersal by invertebrates or rodents moves seeds over a relatively short distance and a large proportion of the seeds may be lost to seed predation within this step. Longer dispersal distances and potentially larger ecological consequences follow from sequential endochory by two different animals, i.e. diploendozoochory: a primary disperser that initially consumes the seed, and a secondary, carnivorous animal that kills and eats the primary consumer along with the seeds in the prey's digestive tract, and then transports the seed further in its own digestive tract.

<span class="mw-page-title-main">Evolution of seed size</span>

The first seeded plants emerged in the late Devonian 370 million years ago. Selection pressures shaping seed size stem from physical and biological sources including drought, predation, seedling-seedling competition, optimal dormancy depth, and dispersal.

References

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Further reading