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Eye stalks and their scaling relationship with body size are important features in the mating system of many diopsid species, and sexual selection is a critical force influencing the evolution of this exaggerated morphology. Interspecific variation in eye span suggests there has been significant evolutionary change in this trait, but a robust phylogenetic hypothesis is required to determine its rate and direction of change. In this study, the pattern of morphological evolution of eye span is assessed in a phylogenetic framework with respect to its function in the sexual system of these flies. Specifically, we examine within the family Diopsidae the pattern of increase and decrease in sexual dimorphism, the morphological coevolution of eye span between males and females, and the evolutionary flexibility of eye-span allometry. Based on several different methods for reconstructing morphological change, results suggest a general pattern of evolutionary flexibility, particularly for eye-span allometry. Sexual dimorphism in eye span has evolved independently at least four times in the family and this trait also has undergone several reductions within the genus Diasemopsis. Despite most species being dimorphic, there is a strong phylogenetic correlation between males and females for mean eye span. The coevolution between the sexes for eye-span allometry, however, is significantly weaker. Overall, eye-span allometry exhibits significantly more change on the phylogeny than the other morphological traits. The evolutionary pattern in eye-span allometry is caused primarily by changes in eye-span variance. Therefore, this pattern is consistent with recent models that predict a strong relationship between sexual selection and the variance of ornamental traits and highlights the significance of eye-span allometry in intersexual and intrasexual signaling.

Abstract.-A phylogenetic hypothesis of relationships among 33 species of stalk-eyed ies was generated from a molecular data set comprising three mitochondrial and three nuclear gene regions. A combined analysis of all the data equally weighted produced a single most-parsimonious cladogram with relatively strong support at the majority of nodes. The phylogenetic utility of different classes of molecular data was also examined. In particular, using a number of different measures of utility in both a combined and separate analysis framework, we focused on the distinction between mitochondrial and nuclear genes and between faster-evolving characters and slower-evolving characters. For the rst comparison, by nearly any measure of utility, the nuclear genes are substantially more informative for resolving diopsid relationships than are the mitochondrial genes. The nuclear genes exhibit less homoplasy, are less incongruent with one another and with the combined data, and contribute more support to the combined analysis topology than do the mitochondrial genes. Results from the second comparison, however, provide little evidence of a clear difference in utility. Despite indications of rapid divergence and saturation, faster-evolving characters in both the nuclear and mitochondrial data sets still provide substantial phylogenetic signal. In general, inclusion of the more rapidly evolving data consistently improves the congruence among partitions. [Diopsidae; incongruence; nuclear genes; partitioned Bremer support; saturation.] Flies in the family Diopsidae are characterized by the elongation of the head into long stalks, with the eyes and antennae laterally displaced at the ends of these stalks. In several species, this elongation is so extreme that the length of their eye stalks exceeds the length of their body. Several dipteran families possess hypercephalic species (Grimaldi and Fenster, 1989;Wilkinson and Dodson, 1997), but the Diopsidae are unique in that both males and females of all the species within the family have some degree of head modi cation. Recently, experiments examining the importance of eye stalks in the mating system of diopsids have provided considerable information about their adaptive signicance (Burkhardt and de la Motte, 1985Motte, , 1988Lorch et al., 1993;Wilkinson, 1993;Burkhardt et al., 1994;Wilkinson and Reillo, 1994;Wilkinson and Dodson, 1997). For the majority of diopsid species, males have markedly larger eye stalks than females, and several studies have demonstrated that this increased eye span functions as an ornament for both male combat (Burkhardt and de la Motte, 1983Motte, , 1987Lorch et al., 1993) and 4 Present address (and address for correspondence):

Aging is often perceived as a degenerative process caused by random accrual of cellular damage over time. In spite of this, age can be accurately estimated by epigenetic clocks based on DNA methylation profiles from almost any tissue of the body. Since such pan-tissue epigenetic clocks have been successfully developed for several different species, it is difficult to ignore the likelihood that a defined and shared mechanism instead, underlies the aging process. To address this, we generated 10,000 methylation arrays, each profiling up to 37,000 cytosines in highly-conserved stretches of DNA, from over 59 tissue-types derived from 128 mammalian species. From these, we identified and characterized specific cytosines, whose methylation levels change with age across mammalian species. Genes associated with these cytosines are greatly enriched in mammalian developmental processes and implicated in age-associated diseases. From the methylation profiles of these age-related cytosines, we successfully constructed three highly accurate universal mammalian clocks for eutherians, and one universal clock for marsupials. The universal clocks for eutherians are similarly accurate for estimating ages (r>0.96) of any mammalian species and tissue with a single mathematical formula. Collectively, these new observations support the notion that aging is indeed evolutionarily conserved and coupled to developmental processes across all mammalian species - a notion that was long-debated without the benefit of this new and compelling evidence.