Samuel Cotton’s research while affiliated with University College London and other places

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Publications (27)


The distribution of second male sperm precedence (P2) among 22 dam-sire pair families in the stalk-eyed fly, Teleopsis dalmanni. Open bars depict the frequency of families with P2 significantly different from 0.5 (using 2-tailed binomial tests); black bars denote those with P2 not significantly different from 0.5. The grey bar denotes the sperm precedence of a single family where the observed P2 of zero was not significantly different from P2 = 0.5. However, the brood size of this family was small (n = 5), so this result should be treated with caution.
Highly variable sperm precedence in the stalk-eyed fly, Teleopsis dalmanni
  • Article
  • Full-text available

February 2006

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157 Reads

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23 Citations

BMC Evolutionary Biology

Laura S Corley

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Samuel Cotton

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Ellen McConnell

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[...]

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When females mate with different males, competition for fertilizations occurs after insemination. Such sperm competition is usually summarized at the level of the population or species by the parameter, P2, defined as the proportion of offspring sired by the second male in double mating trials. However, considerable variation in P2 may occur within populations, and such variation limits the utility of population-wide or species P2 estimates as descriptors of sperm usage. To fully understand the causes and consequences of sperm competition requires estimates of not only mean P2, but also intra-specific variation in P2. Here we investigate within-population quantitative variation in P2 using a controlled mating experiment and microsatellite profiling of progeny in the multiply mating stalk-eyed fly, Teleopsis dalmanni. We genotyped 381 offspring from 22 dam-sire pair families at four microsatellite loci. The mean population-wide P2 value of 0.40 was not significantly different from that expected under random sperm mixing (i.e. P2 = 0.5). However, patterns of paternity were highly variable between individual families; almost half of families displayed extreme second male biases resulting in zero or complete paternity, whereas only about one third of families had P2 values of 0.5, the remainder had significant, but moderate, paternity skew. Our data suggest that all modes of ejaculate competition, from extreme sperm precedence to complete sperm mixing, occur in T. dalmanni. Thus the population mean P2 value does not reflect the high underlying variance in familial P2. We discuss some of the potential causes and consequences of post-copulatory sexual selection in this important model species.

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Do insect sexual ornaments demonstrate heightened condition dependence?

September 2005

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19 Reads

These proceedings contain 18 papers that discuss topics on speciation and adaptation; life history, evolution, phenotypic plasticity and genetics; sexual selection and reproductive biology; insect-plant interactions; insect-natural enemy interactions; and social insects. A series of empirical case studies in evolutionary ecology using insects as model systems are also presented.



Figure 1. Singing for success. Males with the most complex songs leave the most descendents in the song sparrow Melospiza melodia. (Photo courtesy of Doug Backlund.) 
Sexual Selection: The Importance of Long-Term Fitness Measures

Current Biology

New results from a 20-year study of free-living song sparrows confirm that attractive males contribute more offspring than less attractive males. They also reveal that the offspring of preferred males produce more descendents themselves. Females prefer males with a large song repertoire, which further work shows is a condition-dependent indicator of male quality.


Figure 5: Comparisons between eyespan (ES) and wing length of Cyrtodiopsis dalmanni in their response to food treatment after controlling for body size. Least squares means estimates were standardized to unity in the 0.12 g treatment group to ease comparisons between different sized traits. Least squares means from other treatments are expressed as proportions of the standardized 0.12 g groups. Error bars are omitted for clarity. Asterisks indicate significance of within-trait, between-adjacent treatment comparisons after sequential Bonferroni correction: **P < 0.01, *P < 0.05.
Heightened condition dependence is not a general feature of male eyespan in stalk-eyed flies (Diptera: Diopsidae)

December 2004

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81 Reads

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43 Citations

Journal of Evolutionary Biology

Stalk-eyed flies are exemplars of sexual selection leading to the evolution of exaggerated male ornaments (eyespan). In Sphyracephala beccarri, there is no evidence for female mate choice for exaggerated male eyespan and only minor sex differences in eyespan. We used S. beccarri to test whether heightened condition dependence only evolves when male eyespan becomes sexually exaggerated. Male eyespan showed heightened condition dependence under food stress compared with a control trait (wing length). However, female eyespan displayed a similar pattern and there was no sex difference in the degree of increased eyespan sensitivity. The finding that eyespan is a sensitive indicator of food stress, even in an unexaggerated state, suggests that this may have acted as a pre-adaptation to the role of eyespan in sexual signalling in other Diopsid species. These results are consistent with handicap theory and Fisher's view of how sexual selection is initiated.


TABLE 1 . Absolute (mm) trait sizes (mean SD) of flies grown in each environment. Sample sizes are given in parentheses. 
TABLE 3 . Z-values for between-trait comparisons of allometric dis- persion (AD). For AD-values refer to Figure 6. 
Condition dependence of sexual ornament size and variation in the stalk-eyed fly Cyrtodiopsis dalmanni (Diptera : Diopsidae)

June 2004

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465 Reads

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212 Citations

Evolution

We used the stalk-eyed fly Cyrtodiopsis dalmanni to examine predictions made by condition-dependent handicap models of sexual selection. Condition was experimentally varied by manipulation of larval food availability. Cyrtodiopsis dalmanni is a highly dimorphic species exhibiting strong sexual selection, and the male sexual ornament (exaggerated eyespan) showed strong condition-dependent expression relative to the homologous trait in females and nonsexual traits. Male eyespan also showed a great increase in standardized variance under stress, unlike nonsexual traits. The inflated variance of the male ornament was primarily attributable to condition-dependent (but body-size-independent) increase in variance. Thus, evaluation of male eyespan allows females to gain additional information about male condition over and above that given by body size. These findings accord well with condition-dependent handicap models of sexual selection.


Do sexual ornaments demonstrate heightened condition-dependent expression as predicted by the handicap hypothesis?

May 2004

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225 Reads

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588 Citations

The handicap hypothesis of sexual selection predicts that sexual ornaments have evolved heightened condition-dependent expression. The prediction has only recently been subject to experimental investigation. Many of the experiments are of limited value as they: (i) fail to compare condition dependence in sexual ornaments with suitable non-sexual trait controls; (ii) do not adequately account for body size variation; and (iii) typically consider no stress and extreme stress manipulations rather than a range of stresses similar to those experienced in nature. There is also a dearth of experimental studies investigating the genetic basis of condition dependence. Despite the common claim that sexual ornaments are condition-dependent, the unexpected conclusion from our literature review is that there is little support from well-designed experiments.


Citations (25)


... Nearly all species of Diopsidae (Bilberg, 1820) are well-known for their exaggerated eye stalks (Shillito 1971). There are approximately 160-8000 species and 10-15 genera containing stalk-eyed flies in the family Diopsidae (Shillito 1971;Steyskal 1972;Carr et al. 2006;Ovtshinnikova and Galinskaya 2016;Roskov et al. 2019). Although both males and females in Diopsinae have eyes that are laterally displaced from the central head, the level of sexual dimorphism varies between and within species (Burkhardt and de la Motte 1985; Wilkinson and Dodson 1997;Meier and Hilger 2000). ...

Reference:

Description of the karyotype of Sphyracephala detrahens (Diptera, Diopsidae)
A description of a new species of Diasemopsis (Diptera, Diopsidae) from the Comoro Islands with morphological, molecular and allometric data

Zootaxa

... The geographically close habitat ranges of Bactrocera and the potential donor genus Teleopsis led to the proposal that the horizontal transfer may have occurred in New Guinea. The species composition of Teleopsis is under debate [29][30][31], however here it is used within its broadest sense to include the putatively nested or synonymous genera Cyrtodiopsis and Megalobops. ...

A morphological and molecular description of a new Teleopsis species (Diptera: Diopsidae) from Thailand
  • Citing Article
  • October 2007

Zootaxa

... It has long been recognized that the genetic relationship between male sexual display traits and components of nonsexual fitness such as condition is an important empirical pursuit in understanding the evolution of female mating preferences (Cotton et al., 2004;Hunt et al., 2004;Tomkins et al., 2004). Our results suggest that if we are to understand the evolution of male mating preferences, analyses of male preferences for female sexual display traits will need to be complemented with a detailed understanding of their genetic basis. ...

Do insect sexual ornaments demonstrate heightened condition dependence?
  • Citing Article
  • September 2005

... Although less studied than female mate choice, there is evidence that male mate choice is common across taxa [15][16][17][18][19] and is often based on female traits that are correlated with fecundity [17][18][19][20][21]. Despite reproductive senescence causing female fecundity to decline with age, there is mixed evidence of how male mate choice is affected by female age. ...

Male mate preference for female eyespan and fecundity in the stalk-eyed fly, Teleopsis dalmanni

Behavioral Ecology

... Females have evolved responses or preferences to prevent them from mating with SGE carriers in some cases. In stalk-eyed flies, females preferentially mate with males with wider eye spans, whereas SGE-carrying males are generally associated with smaller eye spans (Cotton et al., 2014;Wilkinson et al., 1988). The presence of sperm-killing meiotic drive also correlates with the rate of polyandry in populations, where meiotic drive is more common in areas with low polyandry (Pinzone & Dyer, 2013;Price et al., 2014). ...

Male eyespan size is associated with meiotic drive in wild stalk-eyed flies (Teleopsis dalmanni)

Heredity

... Chaetodiopsis Séguy (Diopsinae). An endemic genus of two described species: C. comoroensis (Carr & Földvári, 2006), described from Comoros (Carr et al. 2006); and C. meigenii, the commonest sub-Saharan diopsid, widely distributed in the continental Afrotropics and even extending into Oman in the Southern Arabian Peninsula . The genus is one of the four genera comprising the Diasemopsis genus-group. ...

A Description Of A New Species Of Diasemopsis (Diptera, Diopsidae) From The Comoro Islands With Morphological, Molecular And Allometric Data

Zootaxa

... Common ancestry makes species means nonindependent of each other (Felsenstein, 1985); thus, we also performed the same sets of analyses described above with phylogeny taken into account. Because there is no published phylogeny that includes all of the taxa we studied, we created a composite phylogeny (Fig. 1) from Baker & Wilkinson (2001), Wright et al. (2004), Swallow et al. (2005), and Földvári et al. (2007, and set branch lengths equal to 1 because the divergence times among the different species are unclear. We then used the PDAP:PDTREE module (Garland, Midford & Ives, 1999; Midford, Garland & Maddison, 2005) in MESQUITE, version 2.72 (Maddison & Maddison, 2009) to calculate standardized phylogenetically independent contrasts (Felsenstein, 1985). ...

A morphological and molecular description of a new Teleopsis species (Diptera: Diopsidae) from Thailand

Zootaxa

... Biological ornaments are conspicuous traits considered to function as condition-dependent signals of mate quality during mate selection (Hill, 2014;Winters, 2018), though they may serve a dual role as armaments (i.e., weapons or status badges; Berglund et al., 1996). Examples of visual ornamentation can be found across many taxa, such as the colourful iridescent tail feathers of peacocks (Pavo cristatus; Dakin and Montgomerie, 2013) and the long eye stalks of stalk-eyed flies (Teleopsis dalmanni; Cotton et al., 2010). As condition-dependent signals, ornaments are reported to have higher quality under more favourable environmental conditions (e.g., head ornaments of unparasitised vs parasitised male red jungle fowl, Gallus gallus, progenitor of the domestic fowl; Zuk et al., 1990). ...

Eyespan reflects reproductive quality in wild stalk-eyed flies

Evolutionary Ecology

... This may also explain why the relationship between the vibrational interaction and frontal leg length found for losers have similar but non-significant trends as the ones found for winners. In animal contests, it is common for the larger rival to be more aggressive than the smaller one (e.g. Brown et al. 2006;Small et al. 2009;Graham et al. 2020) including spider contests (Whitehouse 1997;Taylor et al. 2001;Elias et al. 2008). In T. clavipes, in particular, males escalate the interaction to physical contact more frequently when the contest involves two individuals of greater size than two individuals of smaller size (Constant et al. 2011). ...

Small J, Cotton S, Fowler K, Pomiankowski A.. Male eyespan and resource ownership affect contest outcome in the stalk-eyed fly, Teleopsis dalmanni. Anim Behav 78: 1213-1220

Animal Behaviour

... The direct benefits and costs of polyandry have been documented for several species of insects, even when only using two males as the polyandry treatment Wedell 1998, Worthington andKelly 2016). Nonetheless, in other cases, the direct benefits of polyandry are not detected or are minimal (Jennions et al. 2007, Harley et al. 2010, as in this experiment. Regarding the possible indirect benefits of polyandry beyond egg viability, like inbreeding avoidance, increased genetic compatibility, and increased offspring fitness (Ivy and Sakaluk 2005) it was not possible to detect them with our design, and requires further scrutiny (Jennions et al. 2007, Slatyer et al. 2012. ...

No Detectable Fertility Benefit from a Single Additional Mating in Wild Stalk-Eyed Flies