Fecundity selection

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Fecundity selection, also known as fertility selection, is the fitness advantage resulting from selection on traits that increases the number of offspring (i.e. fecundity). [1] Charles Darwin formulated the theory of fecundity selection between 1871 and 1874 to explain the widespread evolution of female-biased sexual size dimorphism (SSD), where females were larger than males. [2]

Contents

Along with the theories of natural selection and sexual selection, fecundity selection is a fundamental component of the modern theory of Darwinian selection. Fecundity selection is distinct [3] in that large female size relates to the ability to accommodate more offspring, and a higher capacity for energy storage to be invested in reproduction. Darwin's theory of fecundity selection predicts the following: [1]

  1. Fecundity depends on variation in female size, which is associated with fitness.
  2. Strong fecundity selection favors large female size, which creates asymmetrical female-biased sexual size dimorphism.

Although sexual selection and fecundity selection are distinct, it still may be difficult to interpret whether sexual dimorphism in nature is due to fecundity selection, or to sexual selection. [4] [5] Examples of fecundity selection in nature include self-incompatibility flowering plants, where pollen of some potential mates are not effective in forming seed, [6] as well as bird, lizard, fly, and butterfly and moth species that are spread across an ecological gradient. [7] [8] [9] [10]

Moreau-Lack's rule

Moreau (1944) suggested that in more seasonal environments or higher latitudes, fecundity depends on high mortality. [11] Lack (1954) suggested differential food availability and management across latitudes play a role in offspring and parental fitness. [12] Lack also highlighted that more opportunities for parents to collect food due to an increase in day-length towards the poles is an advantage. This means that moderately higher altitudes provide more successful conditions to produce more offspring. However, extreme day-lengths (i.e. at the poles) may work against parental survival as repetitive food searching would exhaust the parent.

Together, the Moreau-Lack rule hypothesizes that fecundity increases with increasing latitude. [1] Evidence supporting and doubting this claim has led to the consolidation of other predictions, which may better explain Moreau-Lack's rule.

Seasonality and Ashmole's hypothesis

Ashmole (1963) suggested (bird) fecundity depends on seasonality patterns. [13] Food differences in availability between seasons are greater towards higher latitudes, so birds are predicted to experience low survival during the winter due to limited resources. This decline in population may be advantageous for survivors, since there is more food available by the next breeding season. This leads to an enhancement of energy when invested in fitness as a result of higher fecundity. [1] [13] Therefore, Ashmole's hypothesis is dependent upon resource availability as a factor fecundity. [1]

Differences in nest predation

Areas with severe nest predation tend to be those of large clutches/litters, especially in the tropics, [1] as they are more noticeable to predators (frequent parental care, noisier offspring [14] [15] [16] ). This predation pressure may lead to the selection for multiple nests of smaller size, with shorter development time.

A criticism of this hypothesis is that it indirectly assumes that these nest-predators are visually-oriented, however, they may be chemically oriented, too, with heightened olfactory senses.

Length of breeding season (LBS) hypothesis

Populations at higher latitudes experience an increasing seasonality and shorter warm seasons. As a result, these populations have more chances of having multiple reproductive episodes. [1] Intense fecundity selection depends on the length of breeding season (LBS). Factors that may delay LBS or the start of breeding season, are snow cover or delayed food growth, which, in turn, minimizes the chance for these populations to reproduce.

Long breeding seasons towards the tropics favor smaller clutches since females are able to balance energy reserved for reproduction, and the risk of predation. [1] [16] Fecundity selection acts by favoring early reproduction and higher clutch size in species that reproduce frequently. The opposite trend is seen in populations that reproduce less frequently, where delayed reproduction is favored.

The 'bet-hedging strategy' hypothesis

The total fecundity per year depends on the length of breeding season (LBS), which also determines the number of breeding episodes. In addition, the total fecundity also depends on nest predation, as it describes differential survival over a variety of populations. [1] [17] When food is limited, and the breeding season is long, and nest predation is intense, selection tends to favor a 'bet-hedging' strategy, where the risk of predation is spread over many smaller clutches. This means that the success of the number of offspring depends on whether they are large in size or not. The strategy suggests that fewer, but larger, clutches in higher latitudes are a result of food seasonality, nest predation, and LBS.

In nature

The findings below are based on individual research studies.

Southern and Northern Hemisphere birds

It has been assumed that parents of fewer offspring, with a high probability of adult survival, should permit less risk to themselves. Even though this compromises their young, the overall fitness of their offspring is reduced, which is a strategy to invest in producing more offspring in the future. It was found that within and between regions, there is a negative correlation between clutch size and adult survival. Southern-Hemisphere parents were inclined to reduce mortality risk to themselves, even at a cost to their offspring, whereas Northern parents experienced greater risk to themselves to reduce risk to their offspring. [8]

(Edward B. Poulton, 1890). Differences in wing size, wing shape, wing color pattern, the size and shape of antennae and of body hairs, as well as abdominal characteristics in butterfly. Females (right) are overall much larger than males (left) Mimicry in South African Butterflies - chromolithographic frontispiece of The Colours of Animals by Edward Bagnall Poulton, 1890.jpg
(Edward B. Poulton, 1890). Differences in wing size, wing shape, wing color pattern, the size and shape of antennae and of body hairs, as well as abdominal characteristics in butterfly. Females (right) are overall much larger than males (left)

Liolaemus lizard

Liolaemus species span from the Atacama Desert to austral rain forests and Patagonia, and across a wide range of altitudes. Due to radiation, life history strategies have diversified within this genus. [9] In turn, it was found that increased fecundity does not lead to female-biased SSD, which is also not effected by latitude-elevation.

Drosophila melanogaster

In lines of D. melanogaster selected for increased fecundity (i.e. more eggs laid over an 18-hour period), females experienced an increase in thorax and abdomen width than males. [10] In general, SSD increased with selection for increased fecundity. These results support the hypothesis that in response to fecundity selection, SSD can evolve rapidly. [10]

Lepidoptera butterfly and moth species

Female-biased SSD in many Lepidopteran species are initiated during their developmental period. Since females of this species, as in many other species, reserve their larval resources for reproduction, fecundity depends on larger (female) size. In this way, larger females can enhance fecundity as well as their survival by having multiple partners. [7]

Other types of selection

Natural selection is defined as the differential survival and/or reproduction of organisms as a function of their physical attributes, where their 'fitness' is the ability to adapt to the environment and produce more (fertile) offspring. [18] The trait(s) that contribute to survival or reproduction of offspring has a higher chance of being expressed in the population. [19]

Sexual selection acts to refine secondary sexual (i.e. non-genital) phenotypes, such as the morphological differences between males and females (sexual dimorphism), or even differences between species of the same sex. [18] As a refinement to Darwin's theory of selection, Trivers (1974) observed that: [18] [20]

  1. Females are the limiting sex and invest more in offspring than males
  2. Because males tend to be in excess, males tend to develop ornaments for attracting mates (female choice), as well competing with other males.

See also

Related Research Articles

<span class="mw-page-title-main">Sexual selection</span> Mode of natural selection involving the choosing of and competition for mates

Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with, and compete with members of the same sex for access to members of the opposite sex. These two forms of selection mean that some individuals have greater reproductive success than others within a population, for example because they are more attractive or prefer more attractive partners to produce offspring. Successful males benefit from frequent mating and monopolizing access to one or more fertile females. Females can maximise the return on the energy they invest in reproduction by selecting and mating with the best males.

<span class="mw-page-title-main">Sexual dimorphism</span> Condition where males and females exhibit different characteristics

Sexual dimorphism is the condition where sexes of the same species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most animals and some plants. Differences may include secondary sex characteristics, size, weight, color, markings, or behavioral or cognitive traits. Male–male reproductive competition has evolved a diverse array of sexually dimorphic traits. Aggressive utility traits such as "battle" teeth and blunt heads reinforced as battering rams are used as weapons in aggressive interactions between rivals. Passive displays such as ornamental feathering or song-calling have also evolved mainly through sexual selection. These differences may be subtle or exaggerated and may be subjected to sexual selection and natural selection. The opposite of dimorphism is monomorphism, when both biological sexes are phenotypically indistinguishable from each other.

Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to produce offspring, measured by the number of gametes (eggs), seed set, or asexual propagules.

<span class="mw-page-title-main">Viviparous lizard</span> Species of lizard

The viviparous lizard, or common lizard,, is a Eurasian lizard. It lives farther north than any other species of non-marine reptile, and is named for the fact that it is viviparous, meaning it gives birth to live young. Both "Zootoca" and "vivipara" mean "live birth," in Greek and Latin respectively. It was called Lacerta vivipara until the genus Lacerta was split into nine genera in 2007 by Arnold, Arribas & Carranza.

<span class="mw-page-title-main">Reproductive success</span> Passing of genes on to the next generation in a way that they too can pass on those genes

Reproductive success is an individual's production of offspring per breeding event or lifetime. This is not limited by the number of offspring produced by one individual, but also the reproductive success of these offspring themselves.

<span class="mw-page-title-main">Parental investment</span> Parental expenditure (e.g. time, energy, resources) that benefits offspring

Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure that benefits offspring. Parental investment may be performed by both males and females, females alone or males alone. Care can be provided at any stage of the offspring's life, from pre-natal to post-natal.

<span class="mw-page-title-main">Siberian jay</span> Species of bird

The Siberian jay is a small jay with a widespread distribution within the coniferous forests in North Eurasia. It has grey-brown plumage with a darker brown crown and a paler throat. It is rusty-red in a panel near the wing-bend, on the undertail coverts and on the sides of the tail. The sexes are similar. Although its habitat is being fragmented, it is a common bird with a very wide range so the International Union for Conservation of Nature has assessed its conservation status as being of "least concern".

<span class="mw-page-title-main">Common side-blotched lizard</span> Species of lizard

The common side-blotched lizard is a species of side-blotched lizard in the family Phrynosomatidae. The species is native to dry regions of the western United States and northern Mexico. It is notable for having a unique form of polymorphism wherein each of the three different male morphs utilizes a different strategy in acquiring mates. The three morphs compete against each other following a pattern of rock paper scissors, where one morph has advantages over another but is outcompeted by the third.

<span class="mw-page-title-main">Clutch (eggs)</span> Grouping of eggs in a nest

A clutch of eggs is the group of eggs produced by birds, amphibians, or reptiles, often at a single time, particularly those laid in a nest.

Life history theory is an analytical framework designed to study the diversity of life history strategies used by different organisms throughout the world, as well as the causes and results of the variation in their life cycles. It is a theory of biological evolution that seeks to explain aspects of organisms' anatomy and behavior by reference to the way that their life histories—including their reproductive development and behaviors, post-reproductive behaviors, and lifespan —have been shaped by natural selection. A life history strategy is the "age- and stage-specific patterns" and timing of events that make up an organism's life, such as birth, weaning, maturation, death, etc. These events, notably juvenile development, age of sexual maturity, first reproduction, number of offspring and level of parental investment, senescence and death, depend on the physical and ecological environment of the organism.

Monogamous pairing in animals refers to the natural history of mating systems in which species pair bond to raise offspring. This is associated, usually implicitly, with sexual monogamy.

Cooperative breeding is a social system characterized by alloparental care: offspring receive care not only from their parents, but also from additional group members, often called helpers. Cooperative breeding encompasses a wide variety of group structures, from a breeding pair with helpers that are offspring from a previous season, to groups with multiple breeding males and females (polygynandry) and helpers that are the adult offspring of some but not all of the breeders in the group, to groups in which helpers sometimes achieve co-breeding status by producing their own offspring as part of the group's brood. Cooperative breeding occurs across taxonomic groups including birds, mammals, fish, and insects.

<span class="mw-page-title-main">Parental care</span>

Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal kingdom. There is great variation in different animal groups in terms of how parents care for offspring, and the amount of resources invested by parents. For example, there may be considerable variation in the amount of care invested by each sex, where females may invest more in some species, males invest more in others, or investment may be shared equally. Numerous hypotheses have been proposed to describe this variation and patterns in parental care that exist between the sexes, as well as among species.

In behavioral ecology, adaptive behavior is any behavior that contributes directly or indirectly to an individual's reproductive success, and is thus subject to the forces of natural selection. Examples include favoring kin in altruistic behaviors, sexual selection of the most fit mate, and defending a territory or harem from rivals.

Polygyny is a mating system in which one male lives and mates with multiple females but each female only mates with a single male. Systems where several females mate with several males are defined either as promiscuity or polygynandry. Lek mating is frequently regarded as a form of polygyny, because one male mates with many females, but lek-based mating systems differ in that the male has no attachment to the females with whom he mates, and that mating females lack attachment to one another.

<span class="mw-page-title-main">Avian clutch size</span>

Clutch size refers to the number of eggs laid in a single brood by a nesting pair of birds. The numbers laid by a particular species in a given location are usually well defined by evolutionary trade-offs with many factors involved, including resource availability and energetic constraints. Several patterns of variation have been noted and the relationship between latitude and clutch size has been a topic of interest in avian reproduction and evolution. David Lack and R.E. Moreau were among the first to investigate the effect of latitude on the number of eggs per nest. Since Lack's first paper in the mid-1940s there has been extensive research on the pattern of increasing clutch size with increasing latitude. The proximate and ultimate causes for this pattern have been a subject of intense debate involving the development of ideas on group, individual, and gene-centric views of selection.

<span class="mw-page-title-main">Sexual selection in amphibians</span> Choice of and competition for mates

Sexual selection in amphibians involves sexual selection processes in amphibians, including frogs, salamanders and newts. Prolonged breeders, the majority of frog species, have breeding seasons at regular intervals where male-male competition occurs with males arriving at the waters edge first in large number and producing a wide range of vocalizations, with variations in depth of calls the speed of calls and other complex behaviours to attract mates. The fittest males will have the deepest croaks and the best territories, with females making their mate choices at least partly based on the males depth of croaking. This has led to sexual dimorphism, with females being larger than males in 90% of species, males in 10% and males fighting for groups of females.

Polyandry in fishes is a mating system where females mate with multiple males within one mating season. This type of mating exists in a variety of animal species. Polyandry has been found in both oviparous and viviparous bony fishes and sharks. General examples of polyandry occur in fish species, such as green swordtails and Trinidadian guppies. Specific types of polyandry have also been classified, such as classical polyandry in pipefish cooperative polyandry in cichlids and convenience polyandry in sharks.

<span class="mw-page-title-main">Polyandry in nature</span>

In behavioral ecology, polyandry is a class of mating system where one female mates with several males in a breeding season. Polyandry is often compared to the polygyny system based on the cost and benefits incurred by members of each sex. Polygyny is where one male mates with several females in a breeding season . A common example of polyandrous mating can be found in the field cricket of the invertebrate order Orthoptera. Polyandrous behavior is also prominent in many other insect species, including the red flour beetle and the species of spider Stegodyphus lineatus. Polyandry also occurs in some primates such as marmosets, mammal groups, the marsupial genus' Antechinus and bandicoots, around 1% of all bird species, such as jacanas and dunnocks, insects such as honeybees, and fish such as pipefish.

<span class="mw-page-title-main">Parental care in birds</span>

Parental care refers to the level of investment provided by the mother and the father to ensure development and survival of their offspring. In most birds, parents invest profoundly in their offspring as a mutual effort, making a majority of them socially monogamous for the duration of the breeding season. This happens regardless of whether there is a paternal uncertainty.

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