Fecundity selection

Fecundity selection, also known as fertility selection, is the fitness advantage resulting from the preference of traits that increase 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]

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 a 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)

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 or 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

• Fecundity
• R/K selection theory

References

1. Pincheira-Donoso, D. and Hunt, J. Fecundity selection theory: concepts and evidence. Biological Reviews 92, 341-356 (2017).
2. Darwin, C. (1874). Descent of man, and selection in relation to sex (Second ed.). London: Murray.
3. Clegg, M. T.; Allard, R. W. (1973). "Viability versus Fecundity Selection in the Slender Wild Oat, Avena barbata L.". Science. 181 (4100): 667–668. doi:10.1126/science.181.4100.667. PMID 17736981.
4. Olsson, Mats; Shine, Richard; Wapstra, Erik; Ujvari, Beata; Madsen, Thomas (July 2002). "Sexual Dimorphism In Lizard Body Shape: The Roles Of Sexual Selection And Fecundity Selection" (PDF). Evolution. 56 (7): 1538–1542. doi:10.1111/j.0014-3820.2002.tb01464.x. PMID 12206252.
5. Serrano-Meneses, Martín-Alejandro; Székely, Tamás (June 2006). "Sexual size dimorphism in seabirds: sexual selection, fecundity selection and differential niche-utilisation". Oikos. 113 (3): 385–394. doi:10.1111/j.0030-1299.2006.14246.x.
6. Vekemans, X.; Schierup, M.H.; Christiansen, F.B. (1998), "Mate Availability and Fecundity Selection in Multi-Allelic Self- Incompatibility Systems in Plants", Evolution, 52 (1): 19–29, doi:10.2307/2410916, JSTOR 2410916, PMID 28568138CS1 maint: uses authors parameter (link)
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8. Ghalambor, CK., and Martin, TE. Fecundity-Survival Trade-Offs and Parental Risk-Taking in Birds. Science 292 (5516), 494-497 (2001).
9. Pinchiera-Donoso, D., Tregenza, T. Fecundity selection and the evolution of reproductive output and sex-specific body size in the Liolaemus lizard adaptive radiation. Evolutionary Biology 38: 197-207 (2011).
10. Reeve, JP. and Fairbairn, DJ. Change in sexual size dimoprhism as a correlated response to selection on fecundity. Heredity 83, 697-706 (1999).
11. Moreau, RE. Clutch-size: a comparative study, with special reference to African birds. Ibis 86, 286-347 (1944)
12. Lack, D. The natural regulation of animal numbers. Clarendon Press, Oxford (1954)
13. Ashmole, NP. The regulation of numbers of tropical oceanic birds. Ibis 103b, 458-473 (1963)
14. Slagsvold, T. Clutch size variation in passerine birds: the nest predation hypothesis. Oecologia 54, 159-169 (1982).
15. Slagsvold, T. Clutch size variation in birds in relation to nest predation: on the cost of reproduction. Journal of Animal Ecology53, 945-953 (1984).
16. Skutch, AF. Do tropical birds rear as many young as they can nourish?. Ibis 91, 430-455 (1949).
17. Griebeler, EM. et. al. Evolution of avian clutch size along latitudinal gradients: do seasonality, nest predation or breeding season length matter? Journal of Evolutionary Biology 23, 888-901 (2010).
18. "Introduction to Natural and Sexual Selection" Archived 2010-06-24 at the Wayback Machine. bio.research.ucsc.edu. Retrieved 2018-03-29.
19. "Natural selection". evolution.berkeley.edu. Retrieved 2018-03-29.
20. Trivers, RL. Parent-Offspring Conflict. Integrative and Comparative Biology 14(1), 249-264 (1974).