DEVELOPMENTAL DECOUPLING OF ALTERNATIVE PHENOTYPES: INSIGHTS FROM THE TRANSCRIPTOMES OF HORN‐POLYPHENIC BEETLES

Developmental mechanisms play an important role in determining the costs, limits, and evolutionary consequences of phenotypic plasticity. One issue central to these claims is the hypothesis of developmental decoupling, where alternate morphs result from evolutionarily independent developmental pathways. We address this assumption through a microarray study that tests whether differences in gene expression between alternate morphs are as divergent as those between sexes, a classic example of developmental decoupling. We then examine whether genes with morph‐biased expression are less conserved than genes with shared expression between morphs, as predicted if developmental decoupling relaxes pleiotropic constraints on divergence. We focus on the developing horns and brains of two species of horned beetles with impressive sexual‐ and morph‐dimorphism in the expression of horns and fighting behavior. We find that patterns of gene expression were as divergent between morphs as they were between sexes. However, overall patterns of gene expression were also highly correlated across morphs and sexes. Morph‐biased genes were more evolutionarily divergent, suggesting a role of relaxed pleiotropic constraints or relaxed selection. Together these results suggest that alternate morphs are to some extent developmentally decoupled, and that this decoupling has significant evolutionary consequences. However, alternative morphs may not be as developmentally decoupled as sometimes assumed and such hypotheses of development should be revisited and refined.     

Photoperiod modifies thermal reaction norms for growth and development in the red poplar leaf beetle Chrysomela populi (Coleoptera: Chrysomelidae)

Regression lines of development rate on temperature appeared significantly different between long (22 h) and short (12 h) day conditions and intersected each other at 23.8 °С. Thus, the rate of growth and development was higher at temperatures below the intersection point under short-day but above the intersection point it was higher under long day. Ecological relevance of this effect seems as follows: in autumn, as nights become longer and average daily temperature decreases, larvae have to speed up their development because it is only imago that overwinters. Conversely, midsummer offers long days and usually higher temperature, so again it is advantageous to develop as fast as possible in order to have at least one more generation per year. These results are compared with other studies showing interactions between photoperiod and temperature, and some possible general patterns are outlined. The lower thermal threshold for larval development depended on photoperiodic conditions; therefore rate isomorphy must be violated in this species. Development at higher temperatures generally resulted in smaller adults, as is usual with ectotherms according to the “temperature-size rule”, but body weight depended significantly on temperature only under short day. Our estimates of the lower temperature thresholds for growth and development in both cases did conform to the generalization made previously by Walters and Hassall (2006) in spite of another formula used by us. We briefly discuss this phenomenon and argue that relative position of these thresholds can be explained mathematically and per se may lack any biological sense.     

Microhabitat preferences drive phylogeographic disparities in two Australian funnel web spiders

Comparative phylogeography is underpinned by the assumption that sympatrically‐distributed taxa will have experienced similar environmental histories, resulting in broadly congruent spatial structuring of phylogenetic lineages, particularly if they inhabit similar niches. However, divergent local conditions, specifically those related to microhabitat, may produce significantly divergent systematic signatures of demographic histories. In the present study, we compare the phylogenetic and population genetic spatial patterns displayed by two species of niche‐separated (but sympatrically distributed) Australian funnel web spiders (Mygalomorphae: Hexathelidae). We demonstrate that an apparently minor disparity in habitat niche has led to divergent experiences of a common environmental history in the saproxylic Hadronyche cerberea and the ground‐burrowing Atrax sutherlandi. Furthermore, we take a crucial first step in documenting the molecular systematics of a group that has traditionally suffered from a dearth of research interest. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 805–819.     

地面节肢动物营养类群对土地覆被变化和管理扰动的响应

近几十年来,黑河中游荒漠绿洲过渡带大面积天然沙质草地被转变为农田和防风固沙人工林,然而目前尚缺乏土地利用/覆被变化和管理措施对地面节肢动物多样性形成和维持影响的系统研究。以天然沙质草地转变为的人工梭梭灌木林、人工杨树林、人工樟子松林和农田为研究对象,以天然草地为对照,通过对5种样地地面节肢动物群落的调查及非生物环境因子的测定,采用典范对应分析(CCA)等方法,系统探讨了土地覆被和管理强度(天然草地和人工梭梭林实施零管理、杨树和樟子松人工林实施灌溉管理、农田实施灌溉和施肥管理)变化对地面节肢动物营养类群组成、数量和物种丰富度的影响。主要结果是:(1) 在干旱环境下,单纯改变土地覆被并不会导致土壤环境的显著变化,而高强度的管理扰动加速了土壤环境的演变过程。(2)不同土地覆被和管理措施对营养类群组成及其数量和丰富度的调控作用不同,将天然草地转变为杨树和樟子松人工林和农田后,显著提高了捕食性和植食性类群的比例,降低了'腐食性+杂食性’类群的比例;转变为人工梭梭林后显著降低了3种营养类群的数量;转变为人工杨树林后显著降低了'腐食性+杂食性’类群数量;转变为人工樟子松林后显著增加了植食性类群数量而降低了其它类群的数量;转变为农田后显著增加了植食性类群数量而降低了'腐食性+杂食性’类群数量。(3)影响捕食性动物分布的关键环境因子是地面温度、土壤硝态氮和田间持水量,影响植食性动物分布的关键因子是土壤粘粉粒、土壤硝态氮、土壤容重和pH值,影响'腐食性+杂食性’动物分布的关键因子是地面温度和田间持水量。主要结论是:土地覆被变化与管理措施相互作用对地面节肢动物群落组装过程及其多样性形成具有重要调控作用,尽管其相对重要性不同。     

Transnational Sacralizations: When Daoist Monks meet Global Spiritual Tourists

This article examines the production of sacrality in the context of globalization, through the case of encounters between international spiritual tourists and Chinese monks. The sacred mountain of Huashan has historically been localized in the context of Daoist cosmology, Chinese imperial civilizing, socialist nation-building and, now, global capitalism. While the monks experience Huashan as a gateway for embeddedness into Daoist lineage, ritual, and cosmology, the spiritual tourists approach it as a fountain of raw, consumable energy on a path of disembedding and individuation. But encounters between the two groups lead to the mutual interference and interpenetration of both trajectories. Undermining dichotomist concepts of the sacred which define it as either essentially Other or as socially constructed and contested, the sacrality of Huashan serves as both an anchor for multiple centralizing projects and forces, and as a catalyst and node for the formation of interconnecting and expanding horizontal networks.     

Floral polymorphism and the fitness implications of attracting pollinating and florivorous insects

Background and Aims

Floral polymorphism is frequently attributed to pollinator-mediated selection. Multiple studies, however, have revealed the importance of non-pollinating visitors in floral evolution. Using the polymorphic annual daisy Ursinia calenduliflora, this study investigated the importance of different insect visitors, and their effects on fitness, in the maintenance of floral polymorphism.

Methods

The spatial structure of a discrete floral polymorphism was characterized based on the presence/absence of anthocyanin floret spots in U. calenduliflora. A 3-year observational study was then conducted in polymorphic populations to investigate differences in visitation rates of dominant visitors to floral morphs. Experiments were performed to explore the floral preference of male and female Megapalpus capensis (the dominant insect visitor) and their effectiveness as pollinators. Next, floral damage by antagonistic florivores and the reproductive success of the two floral morphs were surveyed in multiple populations and years.

Key Results

Floral polymorphism in U. calenduliflora was structured spatially, as were insect visitation patterns. Megapalpus capensis males were the dominant visitors and exhibited strong preference for the spotted morph in natural and experimental observations. While this may indicate potential fitness benefits for the spotted morph, female fitness did not differ between floral morphs. However, as M. capensis males are very efficient at exporting U. calenduliflora pollen, their preference may likely increase the reproductive fitness of the spotted morph through male fitness components. The spotted morph, however, also suffered significantly greater costs due to ovule predation by florivores than the spotless morph.

Conclusions

The results suggest that pollinators and florivores may potentially exert antagonistic selection that could contribute to the maintenance of floral polymorphism across the range of U. calenduliflora. The relative strength of selection imposed by each agent is potentially determined by insect community composition and abundance at each site, highlighting the importance of community context in the evolution of floral phenotypes.     

Mechanical limits to maximum weapon size in a giant rhinoceros beetle

The horns of giant rhinoceros beetles are a classic example of the elaborate morphologies that can result from sexual selection. Theory predicts that sexual traits will evolve to be increasingly exaggerated until survival costs balance the reproductive benefits of further trait elaboration. In Trypoxylus dichotomus, long horns confer a competitive advantage to males, yet previous studies have found that they do not incur survival costs. It is therefore unlikely that horn size is limited by the theoretical cost–benefit equilibrium. However, males sometimes fight vigorously enough to break their horns, so mechanical limits may set an upper bound on horn size. Here, I tested this mechanical limit hypothesis by measuring safety factors across the full range of horn sizes. Safety factors were calculated as the ratio between the force required to break a horn and the maximum force exerted on a horn during a typical fight. I found that safety factors decrease with increasing horn length, indicating that the risk of breakage is indeed highest for the longest horns. Structural failure of oversized horns may therefore oppose the continued exaggeration of horn length driven by male–male competition and set a mechanical limit on the maximum size of rhinoceros beetle horns.     

Fluid Dynamics of Flapping Insect Wing in Ground Effect

The fluid dynamics of flapping insect wing in ground effect is investigated numerically in this study. To model the insect wing cross-section in forward-flight mode, the laminar flow over a NACA0012 airfoil animated by a combination of harmonic plunge and pitch rotation is considered. To implement the simulation, the proposed immersed boundary-lattice Boltzmann method is employed. By fixing the Reynolds number and the amplitude of motion, we systematically examine the influences of the distance between the foil and the ground and the flapping frequency on the flow behaviors. As compared to the situation out of ground effect, the forces for foil placed in close proximity to the ground show some differences. The mean drag coefficient is increased at low frequency and decreased at high frequency. Meanwhile, the mean lift coefficient is increased at both low and high frequencies and decreased at middle frequency. Moreover, an interesting phenomenon with oblate vortices due to vortex interaction with the ground is observed.     

Composition and distribution of ground-dwelling beetles among oak fragments and surrounding pine plantations in a temperate forest of North China

In this study, we compared ground-dwelling beetle assemblages （Coleoptera） from a range of different oak fragments and surrounding conifer plantations to evaluate effects of forest size and surrounding matrix habitat in a temperate forest of north China. During 2000, beetles were sampled via pitfall traps within two large oak fragments （ca. 2.0-4.0 ha）, two small oak fragments （ca. 0.2-0.4 ha） and two surrounding matrices dom- inated by pine plantations （〉4 ha） in two sites of different aspects. Overall, no significantly negative effects from forest patch size and the surrounding matrix habitat were detected in total species number and abundance of ground-dwelling beetles. However, compared with small oak patches or pine plantations, more species were associated with an affinity for at least one large oak patch of the two aspects. Multivariate regression trees showed that the habitat type better determined the beetle assemblage structure than patch size and aspect, indicating a strong impact of the surrounding matrix. Linear mixed models indicated that species richness and abundance of all ground-dwelling beetles or beetle families showed different responses to the selected environmental variables. Our results suggest that more disturbed sites are significantly poorer in oak forest specialists, which are usually more abundant in large oak fragments and decrease in abundance or disappear in small fragments and surrounding matrix habitats. Thus, it is necessary to preserve a minimum size of forest patch to create conditions characteristic for forest interior, rather than the more difficult task of increasing habitat connectivity.