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1.  Basal superoxide as a sex-specific immune constraint 
Biology Letters  2011;7(6):906-908.
There is increasing evidence that reactive oxygen species (ROS), a group of unstable and highly reactive chemical molecules, play a key role in regulating and maintaining life-history trade-offs. Upregulation of ROS in association with immune activation is costly because it may result in an imbalance between pro- and antioxidants and, hence, oxidative damage. Previous research aimed at quantifying this cost has mostly focused on changes in the pro-/antioxidant balance subsequent to an immune response. Here, we test the hypothesis that systemic ROS may constrain immune activation. We show that systemic, pre-challenge superoxide (SO) levels are negatively related to the strength of the subsequent immune response towards the mitogen phytohaemagglutinin in male, but not female painted dragon lizards (Ctenophorus pictus). We therefore suggest that systemic SO constrains immune activation in painted dragon males. We speculate that this may be due to sex-specific selection pressures on immune investment.
doi:10.1098/rsbl.2011.0350
PMCID: PMC3210656  PMID: 21632618
reactive oxygen species; immunity; Ctenophorus pictus; phytohaemagglutinin
2.  Proximate determinants of telomere length in sand lizards (Lacerta agilis) 
Biology Letters  2010;6(5):651-653.
Telomeres are repeat sequences of non-coding DNA that cap the ends of chromosomes and contribute to their stability and the genomic integrity of cells. In evolutionary ecology, the main research target regarding these genomic structures has been their role in ageing and as a potential index of age. However, research on humans shows that a number of traits contribute to among-individual differences in telomere length, in particular traits enhancing cell division and genetic erosion, such as levels of free radicals and stress. In lizards, tail loss owing to predation attempts results in a stress-induced shift to a more cryptic lifestyle. In sand lizard (Lacerta agilis) males, telomere length was compromised by tail regrowth in a body size-related manner, so that small males, which already exhibit more cryptic mating tactics, were less affected than larger males. Tail regrowth just fell short of having a significant relationship with telomere length in females, and so did age in males. In females, there was a significant positive relationship between age and telomere length. We conclude that the proximate effect of compromised antipredation and its associated stress seems to have a more pronounced effect in males than in females and that age-associated telomere dynamics differ between the sexes.
doi:10.1098/rsbl.2010.0126
PMCID: PMC2936144  PMID: 20356883
autotomy; telomere length; predation stress
4.  Consistent male–male paternity differences across female genotypes 
Biology Letters  2009;5(2):232-234.
In a recent paper, we demonstrated that male–female genetic relatedness determines male probability of paternity in experimental sperm competition in the Peron's tree frog (Litoria peronii), with a more closely related male outcompeting his rival. Here, we test the hypothesis that a male–male difference in siring success with one female significantly predicts the corresponding difference in siring success with another female. With male sperm concentration held constant, and the proportion of viable sperm controlled statistically, the male–male difference in siring success with one female strongly predicted the corresponding difference in siring success with another female, and alone explained more than 62 per cent of the variance in male–male siring differences. This study demonstrates that male siring success is primarily dictated by among-male differences in innate siring success with less influence of male–female relatedness.
doi:10.1098/rsbl.2008.0796
PMCID: PMC2665838  PMID: 19324659
genetic compatibility; good genes; paternity; relatedness; sperm traits; amphibian
5.  Free radicals run in lizard families 
Biology Letters  2008;4(2):186-188.
In the ageing individual, the production of reactive oxygen species (ROS) accelerates with cell senescence. Depending on the heritability of the underlying processes that determine net ROS levels, this may influence ageing per se and its evolutionary direction and rate of change. In order to understand the inheritance and evolution of net ROS levels in free-ranging lizards, we used flow cytometry together with ROS-sensitive fluorogenic probes to measure ROS in lizard blood cells. We measured basal levels of (i) non-specific ROS (superoxide, singlet oxygen, H2O2 and peroxynitrite), (ii) superoxide specifically and (iii) superoxide after CCCP treatment, which elevated ROS production in the mitochondria. The cumulative level of non-specific ROS was higher in adults than juveniles and superoxide level showed high heritability and variability among families. We suggest that the evolution of ROS dynamics may be ROS species specific and perhaps depend on the relative degree of uni- or biparental inheritance of ROS main regulatory pathways.
doi:10.1098/rsbl.2007.0611
PMCID: PMC2429935  PMID: 18211861
reactive oxygen species; mitochondrial inheritance; heritability
6.  Sons are made from old stores: sperm storage effects on sex ratio in a lizard 
Biology Letters  2007;3(5):491-493.
Sperm storage is a widespread phenomenon across taxa and mating systems but its consequences for central fitness parameters, such as sex ratios, has rarely been investigated. In Australian painted dragon lizards (Ctenophorus pictus), we describe elsewhere that male reproductive success via sperm competition is largely an effect of sperm storage. That is, sperm being stored in the female reproductive tract out-compete more recently inseminated sperm in subsequent ovarian cycles. Here we look at the consequences of such sperm storage for sex allocation in the same species, which has genetic sex determination. We show that stored sperm have a 23% higher probability of producing sons than daughters. Thus, shifts in sex ratio, for example over the reproductive season, can partly be explained by different survival of son-producing sperm or some unidentified female mechanism taking effect during prolonged storage.
doi:10.1098/rsbl.2007.0196
PMCID: PMC2391176  PMID: 17650477
sperm storage; sex ratios; lizard
7.  Consistent sex ratio bias of individual female dragon lizards 
Biology Letters  2006;2(4):569-572.
Sex ratio evolution relies on genetic variation in either the phenotypic traits that influence sex ratios or sex-determining mechanisms. However, consistent variation among females in offspring sex ratio is rarely investigated. Here, we show that female painted dragons (Ctenophorus pictus) have highly repeatable sex ratios among clutches within years. A consistent effect of female identity could represent stable phenotypic differences among females or genetic variation in sex-determining mechanisms. Sex ratios were not correlated with female size, body condition or coloration. Furthermore, sex ratios were not influenced by incubation temperature. However, the variation among females resulted in female-biased mean population sex ratios at hatching both within and among years.
doi:10.1098/rsbl.2006.0526
PMCID: PMC1834013  PMID: 17148290
sex ratio; sex allocation; TSD; Ctenophorus pictus
8.  Differential sex allocation in sand lizards: bright males induce daughter production in a species with heteromorphic sex chromosomes 
Biology Letters  2005;1(3):378-380.
In sand lizards (Lacerta agilis), males with more and brighter nuptial coloration also have more DNA fragments visualized in restriction fragment length polymorphism analysis of their major histocompatibility complex class I loci (and, hence, are probably more heterozygous at these loci). Such males produce more viable offspring, with a particularly strong viability effect on daughters. This suggests that females should adjust both their reproductive investment and offspring sex ratio in relation to male coloration (i.e. differential allocation). Our results show that experimental manipulation of partner coloration in the wild results in significantly higher maternal effort and a 10% higher proportion of daughters than sons. This supports the hypothesis that females increase their maternal energetic expenditure and adjust their offspring sex ratio in response to high-quality partners. However, it also suggests that this has probably evolved through natural selection for increased offspring viability (primarily through production of daughters), rather than through increased mate attraction (e.g. sexy sons).
doi:10.1098/rsbl.2005.0327
PMCID: PMC1617163  PMID: 17148211
sex allocation; maternal allocation; male attractiveness; major histocompatibility complex; heteromorphic sex chromosomes
9.  Costly parasite resistance: a genotype-dependent handicap in sand lizards? 
Biology Letters  2005;1(3):375-377.
Male sand lizards (Lacerta agilis) with a specific restriction fragment length polymorphism fragment in their major histocompatibility complex (MHC) genotype (‘O-males’) are more resistant to ectoparasites (a tick, Ixodes ricinus) than are males that lack this fragment (‘NO-males’). However, emerging evidence suggests that such adaptive immune responses are costly, here manifested by reduced body condition and a compromised defence against secondary infections by haemoprotid parasites that use the ticks as vectors. Subsequent to tick encounter, O-males suffer from a higher leucocyte–erythrocyte ratio, and higher haemoprotid parasitaemia, in particular in relation to vector encounter rate. Furthermore, O-males (i.e. successful tick defenders) with more haemoprotid parasites remaining in their blood stream were in better body condition, whereas this did not apply in NO-males, demonstrating that the adaptive immunoreaction can—in the short term—be energetically even more costly than being moderately parasitized. In agreement with Zahavian handicap theory, O-males had a (marginally) higher reproductive success than males that lacked this fragment.
doi:10.1098/rsbl.2005.0339
PMCID: PMC1617138  PMID: 17148210
MHC; costly parasite resistance; handicap; sand lizard

Results 1-9 (9)