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

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Publications (27)


FIGURE 1. T. thaii, habitus, specimen collected at the type locality.
FIGURE 18. Eyespan allometry in T. thaii. Males = closed circles, females = open circles. Least-squares regression lines are given for heuristic purposes.
FIGURE 19. Bayesian phylogeny of the Teleopsis genus created using 16S+w+wg+COII. Bayes Posterior Probabilities and Maximum Likelihood Bootstrap Support values are given above and below each branch respectively. Unsupported branches are unlabeled, Posterior Probabilities values below 0.70 and Bootstrap Support values below 50% are represented by "-". The bar represents the number of nucleotide substitutions per site.
FIGURES 2-5. T. thaii sp. n. male genitalia. 1: detailed ventral view with surstyli, gonopods, cerci, 2: lateral view with aedeagal apodeme, hypandrium and epandrium, 3: detailed ventral view with base of hypandrium and parameres, 4: ventral view. Legend: ae: aedeagal apodeme, c: circus, e:epandrium, h: hypandrium, hb: hypandrial bridge, p: paramere, s: surstylus.
FIGURES 6-9. T. whitei male genitalia. 5: detailed ventral view with surstyli, gonopods, cerci, 6: lateral view with aedeagal apodeme, hypandrium and epandrium, 7: detailed ventral view with base of hypandrium and parameres, 8: ventral view.

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A morphological and molecular description of a new Teleopsis species (Diptera: Diopsidae) from Thailand
  • Article
  • Full-text available

October 2007

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376 Reads

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11 Citations

Zootaxa

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Samuel Cotton

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A new species of Teleopsis (Diptera, Diopsidae) from Chiang Mai, Thailand is described and illustrated. Teleopsis thaii Földvári & Carr is shown to be a member of a species group, termed the dalmanni species group, along with three previ-ously described species. Presented here are a morphological description of T. thaii and an allometric comparison of the species with other members of the Teleopsis genus. We also present multi-gene phylogenetic analyses to highlight the possible position of T. thaii within the dalmanni species group.

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

October 2007

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35 Reads

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15 Citations

Zootaxa

A new species of Teleopsis (Diptera, Diopsidae) from Chiang Mai, Thailand is described and illustrated. Teleopsis thaii Földvári & Carr is shown to be a member of a species group, termed the dalmanni species group, along with three previously described species. Presented here are a morphological description of T. thaii and an allometric comparison of the species with other members of the Teleopsis genus. We also present multi-gene phylogenetic analyses to highlight the possible position of T. thaii within the dalmanni species group.


Control of introduced species using Trojan sex chromosomes

October 2007

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108 Reads

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80 Citations

Trends in Ecology & Evolution

To control introduced exotic species that have predominantly genetic, but environmentally reversible, sex determination (e.g. many species of fish), Gutierrez and Teem recently modeled the use of carriers of Trojan Y chromosomes--individuals who are phenotypically sex reversed from their genotype. Repeated introduction of YY females into wild populations should produce extreme male-biased sex ratios and eventual elimination of XX females, thus leading to population extinction. Analogous dynamics are expected in systems in which sex determination is influenced by one or a few major genes on autosomes.



Evolutionary genetics: Sexually selected mutation rates

May 2007

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56 Reads

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3 Citations

Heredity

A long-standing problem in evolutionary biology is whether females derive genetic benefits from mating with well-ornamented males. Male sexual ornaments are believed to be informative signals of male genetic quality as they exhibit condition-dependent expression, with condition being influenced by many loci in a male's genome. But persistent selection on ornaments and condition should cause genetic variation in male fitness to be depleted rapidly, thereby eliminating any advantage to female mate preference – a problem known as the 'lek paradox'. Recently, Petrie and Roberts (2006) proposed that female mate choice indirectly selects for elevated rates of mutation, and claim this increases the amount of available genetic variation faster than sexual selection can erode it. Their solution provides an intriguing resolution to the lek paradox, but can it really account for the maintenance of genetic variation in fitness associated with sexual ornaments?


Table 1 . Evidence for condition-dependent mate preference.
Table 1 . Continued
Figure 2. Examples of species in which condition-dependent mate preferences may operate. Condition-dependent mate preferences may play an important role in sexual selection and the evolution of male ornamental traits in many species, including (A) crickets (with permission from Alex McWilliam), (B) stalk-eyed flies (with permission from Sam Cotton) and (C) topi (with permission from [66]). 
Table 2 . The influence of social dominance and social constraints on mate preference.
Table 2 . Continued
Sexual Selection and Condition-Dependent Mate Preferences

October 2006

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593 Reads

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468 Citations

Current Biology

The last decade has witnessed considerable theoretical and empirical investigation of how male sexual ornaments evolve. This strong male-biased perspective has resulted in the relative neglect of variation in female mate preferences and its consequences for ornament evolution. As sexual selection is a co-evolutionary process between males and females, ignoring variation in females overlooks a key aspect of this process. Here, we review the empirical evidence that female mate preferences, like male ornaments, are condition dependent. We show accumulating support for the hypothesis that high quality females show the strongest mate preference. Nonetheless, this is still an infant field, and we highlight areas in need of more research, both theoretical and empirical. We also examine some of the wider implications of condition-dependent mating decisions and their effect on the strength of sexual selection.


FIGURES 9 & 10: D. comoroensis (9) and D. meigenii (10) wings. The larger apical band is clearly visible in the D. meigenii wing, spanning between the M and R2+3 veins. FIGURES 11 & 12: D. comoroensis (11) and D. meigenii (12) first coxae.
FIGURES 13-16: D. comoroensis male genitalia. 13: ventral view, 14: detailed ventral view with surstyli, gonopods, cerci (no hairs on cerci drawn), 15: detailed lateral view with gonopod and paramere, 16: lateral view with aedeagal apodeme, hypandrium and epandrium. Scale bar 0.4 mm for Fig 13, 16, 0.2 mm for Fig 14 and 0.1 mm for Fig 15.
FIGURES 17-20: D. meigenii male genitalia. 9: ventral view, 10: detailed ventral view with surstyli, gonopods, cerci (no hairs on cerci drawn), 11: detailed lateral view with gonopod and paramere, 12: lateral view with aedeagal apodeme, hypandrium and epandrium. Scale bar 0.4 mm for Fig 17, 20, 0.2 mm for Fig 18 and 0.1 mm for Fig 19.
A Description Of A New Species Of Diasemopsis (Diptera, Diopsidae) From The Comoro Islands With Morphological, Molecular And Allometric Data

May 2006

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195 Reads

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19 Citations

Zootaxa

A new species of Diasemopsis (Diptera, Diopsidae) from Comoro Islands is described and illustrated for the first time with allometric datasets. Diasemopsis comoroensis Carr & Földvári is shown to be genetically close, but morphologically distinct from the widespread Afro-tropical species D. meigenii (Westwood); notably a significant divergence in the degree of sexual dimorphism within eye span has occurred between the two species. A revised molecular phylogeny of the genus Diasemopsis is presented based on the partial sequences of four genes.


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

May 2006

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7 Reads

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6 Citations

Zootaxa

A new specie s of Diasemopsis (Diptera, Diopsidae ) from Comoro Islands is described and illustrated for the first time wi th a llometric datasets . Diasemopsis comoroensis Carr & Földvár i is shown t o be genetically close, but morphologically distinct from th e widesprea d Afro-tropical species D. meigenii (Westwood); notably a significant dive rgence in the degree of sexual dimorphism within eye spa n has occurre d between the two species. A revised molecula r phylogeny of th e genus Diasemopsis is presente d based on the partial sequenc es of four genes.


Variation in preference for a male ornament is positively associated with female eyespan in the stalk-eyed fly Diasemopsis meigenii

February 2006

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175 Reads

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62 Citations

Samuel Cotton

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David W Rogers

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Jennifer Small

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[...]

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Kevin Fowler

There is currently much interest in mate preferences for sexual ornaments. However, few studies have focused on individual variation in mate preference despite its importance for the rate and direction of sexual selection. Females of the sexually dimorphic stalk-eyed fly, Diasemopsis meigenii, exhibit an unambiguous rejection response towards unattractive males bearing small ornaments. We investigated individual mate preferences using repeated sequential sampling of female rejection or acceptance responses to a wide range of male ornament phenotypes. We found significant variation in the strength of individual preference. In addition, preference was positively associated with female eyespan, a condition-dependent trait putatively linked to visual acuity.


Genital disc morphology and Engrailed (EN) expression in D. melanogaster and representative species of three Diopsid genera: Teleopsis, Diasemopsis and Sphyracephala. All discs are bilaterally symmetrical. A. EN expression in male third instar genital disc (ventral view) of D. melanogaster. Note EN expression in two posterior lateral domains (arrowhead indicates the posterior domain on the left side of the disc) derived from the posterior compartments of the tenth abdominal segment. The larger anterior domains (arrow indicates the anterior domain on the left side of the disc) are derived from the posterior compartment of the ninth segment and give rise to parts of the male genitalia in the adult. B. EN expression in female third instar genital disc (ventral view) of D. melanogaster. As in the male disc, EN is expressed in two posterior lateral domains (arrowhead indicates the posterior domain on the left side of the disc) derived from the posterior compartments of the tenth abdominal segment. The larger anterior domains (arrow indicates the anterior domain on the left side of the disc) are derived from the posterior compartment of the eighth segment and give rise to parts of the female genitalia in the adult. C, E & G. Male third instar genital discs of T. dalmanni (C), D. meigenii (E) and S. beccarii (G). D, F & H. Female third instar genital discs of T. dalmanni (D), D. meigenii (F) and S. europaea (H). With the exception of the female discs in Sphyracephala, morphology and EN expression appear similar to that of the male and female discs in Drosophila. On each side of the disc EN is expressed in two domains, one anterior (arrows) and one more posterior and lateral (arrowheads). Anterior is uppermost in all panels.
Distance-based neighbour-joining tree of the Diopsid species clade, constructed using sequences of white and wingless. The tree has been rooted with the sequence from the non-hypercephalic species, T. entabenensis. Representative species of each genera that were analysed in this study and featured in Figure 1 C-H are asterisked. Bootstrap values are positioned on the nodes and are percentages taken from 1,000 replicates. The degree of sexual dimorphism in eyespan is indicated for each species.
Assigning sex to pre-adult stalk-eyed flies using genital disc morphology and X chromosome zygosity

February 2006

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164 Reads

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9 Citations

BMC Developmental Biology

In stalk-eyed flies (Diopsidae) the eyes and antennae are laterally displaced at the ends of elongated eyestalks. Eyespan and the degree of sexual dimorphism in eyespan vary considerably between species and several sexually dimorphic species show sexual selection through female mate preference for males with exaggerated eyespan. The genes on which selection acts to regulate eyespan remain to be identified. This could be achieved by comparing gene expression during eyestalk development in males and females if the sex of pre-adult flies could be reliably assigned. Here we describe two techniques, one morphological and one microsatellite-based, that identify the sex of stalk-eyed fly larvae and pupae. We showed that genital discs of the stalk-eyed fly Teleopsis dalmanni have two highly distinct morphologies, compact ("C") and lobed ("L"). Segment composition (revealed by Engrailed expression) was consistent with C morphology being typical of males and L morphology of females. We confirmed the proposed association between disc morphology and sex by evaluating the combined heterozygosity of four X-linked microsatellite markers. We demonstrated that individuals with C genital discs had hemizygous (male) genotypes while those with L discs were heterozygous (female) genotypes. Similar dimorphism in genital disc morphology was observed in eight other species spanning three representative Diopsid genera. In every case the segment composition supported C morphology being male and L morphology female. We assigned larval sex by C or L morphology and compared cell division frequencies in male and female eye-antennal discs in two species (T. dalmanni and Diasemopsis meigenii) sexually dimorphic for eyespan. The number of mitotic (anti-H3-labelled) cells did not differ between the sexes in either species. We have made novel use of two complementary techniques for identifying the sex of pre-adult stalk-eyed flies. These procedures will facilitate studies of the evolution of sexually dimorphic development in a variety of other species. Morphology and En expression in male and female genital discs are highly conserved within each genus of Diopsidae. Finally, sexual dimorphism for eyespan in two Diopsid species is unlikely to be due to an increased rate of cell division at the third larval instar in males.


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