Orientation behaviour in the Grayling butterfly: thermoregulation or crypsis?

ABSTRACT.

• 1 Body orientation and wing tilting in Hipparchia semele (L.) have been explained previously both by the need for shadow-minimization (crypsis hypothesis) and by the need to regulate body temperature (thermoregulation hypothesis).
• 2 This study tests the thermoregulation hypothesis by quantifying butterfly orientation in relation to sun position, using vector analysis, and correlating the data with ambient temperature and time of day.
• 3 During sunshine, Graylings tend to present a maximal wing area to the sun's rays when the ambient temperature is relatively low, and a minimal area when it is high ('sunning’and‘heat-avoiding’positions respectively).
• 4 On cool sunny days, butterflies adopt sunning positions throughout the day, whereas on warm sunny days they adopt sunning positions during the outlying periods and heat-avoiding positions in the middle period of the day. Correlations between wing exposure and both ambient temperature and time of day are statistically significant for warm sunny days.
• 5 During overcast weather, butterfly orientation is very variable and possibly random.
• 6 Analysis of sunny day data supports the thermoregulation hypothesis. This is discussed in relation to other known types of thermoregulatory behaviour observed in this species: stilting, shade-seeking, ground-hugging and shivering.
• 7 Conversely, the results do not support the crypsis hypothesis.

     

Infectious disease in the Pleistocene: Old friends or old foes?

The impact of endemic and epidemic disease on humans has traditionally been seen as a comparatively recent historical phenomenon associated with the Neolithisation of human groups, an increase in population size led by sedentarism, and increasing contact with domesticated animals as well as species occupying opportunistic symbiotic and ectosymbiotic relationships with humans. The orthodox approach is that Neolithisation created the conditions for increasing population size able to support a reservoir of infectious disease sufficient to act as selective pressure. This orthodoxy is the result of an overly simplistic reliance on skeletal data assuming that no skeletal lesions equated to a healthy individual, underpinned by the assumption that hunter-gatherer groups were inherently healthy while agricultural groups acted as infectious disease reservoirs. The work of van Blerkom, Am. J. Phys. Anthropol., vol. suppl 37 (2003), Wolfe et al., Nature, vol. 447 (2007) and Houldcroft and Underdown, Am. J. Phys. Anthropol., vol. 160, (2016) has changed this landscape by arguing that humans and pathogens have long been fellow travelers. The package of infectious diseases experienced by our ancient ancestors may not be as dissimilar to modern infectious diseases as was once believed. The importance of DNA, from ancient and modern sources, to the study of the antiquity of infectious disease, and its role as a selective pressure cannot be overstated. Here we consider evidence of ancient epidemic and endemic infectious diseases with inferences from modern and ancient human and hominin DNA, and from circulating and extinct pathogen genomes. We argue that the pandemics of the past are a vital tool to unlock the weapons needed to fight pandemics of the future.     

Gall-induction in insects: evolutionary dead-end or speciation driver?

Background

The tree of life is significantly asymmetrical - a result of differential speciation and extinction - but general causes of such asymmetry are unclear. Differences in niche partitioning are thought to be one possible general explanation. Ecological specialization might lead to increases in diversification rate or, alternatively, specialization might limit the evolutionary potential of specialist lineages and increase their extinction risk. Here we compare the diversification rates of gall-inducing and non-galling insect lineages. Compared with other insect herbivores feeding on the same host plant, gall-inducing insects feed on plant tissue that is more nutritious and less defended, and they do so in a favorable microhabitat that may also provide some protection from natural enemies. We use sister-taxon comparisons to test whether gall-inducing lineages are more host-specific than non-galling lineages, and more or less diverse than non-gallers. We evaluate the significance of diversity bipartitions under Equal Rates Markov models, and use maximum likelihood model-fitting to test for shifts in diversification rates.

Results

We find that, although gall-inducing insect groups are more host-specific than their non-galling relatives, there is no general significant increase in diversification rate in gallers. However, gallers are found at both extremes - two gall-inducing lineages are exceptionally diverse (Euurina sawflies on Salicaceae and Apiomorpha scale insects on Eucalytpus), and one gall-inducing lineage is exceptionally species-poor (Maskellia armored scales on Eucalyptus).

Conclusions

The effect of ecological specialization on diversification rates is complex in the case of gall-inducing insects, but host range may be an important factor. When a gall-inducing lineage has a host range approximate to that of its non-galling sister, the gallers are more diverse. When the non-galler clade has a much wider host range than the galler, the non-galler is also much more diverse. There are also lineage-specific effects, with gallers on the same host group exhibiting very different diversities. No single general model explains the observed pattern.     

How and when did Old World ratsnakes disperse into the New World?

To examine Holarctic snake dispersal, we inferred a phylogenetic tree from four mtDNA genes and one scnDNA gene for most species of the Old World (OW) and New World (NW) colubrid group known as ratsnakes. Ancestral area distributions are estimated for various clades using divergence-vicariance analysis and maximum likelihood on trees produced using Bayesian inference. Dates of divergence for the same clades are estimated using penalized likelihood with statistically crosschecked calibration references obtained from the Miocene fossil record. With ancestral areas and associated dates estimated, various hypotheses concerning the age and environment associated with the origin of ratsnakes and the dispersal of NW taxa from OW ancestors were tested. Results suggest that the ratsnakes originated in tropical Asia in the late Eocene and subsequently dispersed to the Western and Eastern Palearctic by the early Oligocene. These analyses also suggest that the monophyletic NW ratsnakes (the Lampropeltini) diverged from OW ratsnakes and dispersed through Beringia in the late Oligocene/early Miocene when this land bridge was mostly composed of deciduous and coniferous forests.     

Homo erectus: Ancestor or evolutionary side branch?

Controversies in paleoanthropology wax and wane, but substantial interest is currently focused on Homo erectus. This species has traditionally been regarded as a member in good standing of the human family, where it is placed as an evolutionary intermediate between earlier Homo habilis and later Homo sapiens. Recently, however, some workers have questioned whether the species exists at all. If its populations have been transformed slowly toward the modern condition, and if continuity with living people can be demonstrated in many geographic regions, then any separation of Homo erectus from Homo sapiens must be largely arbitrary. In that case, only one species should be recognized and this slowly changing lineage would have to be called Homo sapiens. Other paleontologists adopt a different view, arguing that Homo erectus is not only anatomically distinctive but also restricted in its geographic distribution. They claim that the fossils from Java and China are so specialized in appearance that they cannot lie in the mainstream of human evolution. Homo erectus, strictly defined as limited to the Far East, probably went extinct without issue. If so, more modern populations must have evolved from another source, perhaps one outside of Asia altogether.     

Heterogeneity in shore flies – the case of Glenanthe Haliday (Diptera: Ephydridae) in the Old World

Old World species of the genus Glenanthe are treated comprehensively with an emphasis on the unusually diverse morphological heterogeneity discovered in structures of the male terminalia. As perspective for this treatment, the tribe Lipochaetini, in which Glenanthe is placed, is characterized and discussed, and an annotated key to the four included genera is provided to facilitate their identification. Glenanthe is demonstrated to be a monophyletic lineage within Lipochaetini. Seven Old World species of Glenanthe are now known, including two new Afrotropical species as follows (type locality in parenthesis): G. namibia n. sp. (Namibia. Mukwe: Divuju: Okavango River [18°04’04”S, 21°28’51”E] and G. danielssoni n. sp. (Republic of South Africa. Cape Province: De Hoop Nature Reserve [34°27’S, 28°25’E]. The identity of G. fuscinervis Becker, revised status (raised from synonymy with G. ripicola (Haliday)), is clarified, and found to be conspecific with Hydrina ochracea Oldenberg, which is recognized as a junior synonym. Papp’s synonymy of Glenanthe fasciventris Becker with Hydrellia (Glenanthe) ripicola Haliday is confirmed, and herein we relegate G. iranica Canzoneri & Rampini also as junior synonym of G. ripicola. The remarkable diverse structures of the male terminalia of all included species are described and illustrated and distribution maps are also provided.     

Animal regeneration: ancestral character or evolutionary novelty?

An old question about regeneration is whether it is an ancestral character which is a general property of living matter, or whether it represents a set of specific adaptations to the different circumstances faced by different types of animal. In this review, some recent results on regeneration are assessed to see if they can throw any new light on this question. Evidence in favour of an ancestral character comes from the role of Wnt and bone morphogenetic protein signalling in controlling the pattern of whole‐body regeneration in acoels, which are a basal group of bilaterian animals. On the other hand, there is some evidence for adaptive acquisition or maintenance of the regeneration of appendages based on the occurrence of severe non‐lethal predation, the existence of some novel genes in regenerating organisms, and differences at the molecular level between apparently similar forms of regeneration. It is tentatively concluded that whole‐body regeneration is an ancestral character although has been lost from most animal lineages. Appendage regeneration is more likely to represent a derived character resulting from many specific adaptations.     

Metabolic flexibility: the key to long-term evolutionary success in Bryozoa?

Oxygen consumption (MO2) and activity were evaluated in Antarctic Bryozoa. Three species representing two different morphologies, flat sheet, laminar forms, Isoseculiflustra tenuis and Kymella polaris, and the bush form Camptoplites bicornis were used. In Bryozoa, activity is measured as the proportion of colony zooids with their lophophores extended. In I. tenuis and K. polaris, residual analysis showed that the percentage of zooids with extended lophophores was not correlated with colony MO2. Lophophore extension is, therefore, a poor measure of activity, and other costs (e.g. growth, reproduction, storage) probably form the major metabolic costs. MO2 per unit of ash-free dry mass (AFDM) in the laminar forms was low compared with other Antarctic marine invertebrates, but not lower than brachiopods and echinoderms. However, the lowest rate here, 16.8 microg O2 g AFDM(-1) h(-1) for a K. polaris colony, is (to our knowledge) the lowest for any animal so far reported. MO2 per unit of AFDM for C. bicornis, however, is among the highest reported for sessile or slow moving Antarctic marine ectotherms, with values similar to those for bivalve and gastropod molluscs. The highest rate, 527 microg O2 g AFDM(-1) h(-1) for one colony is (to our knowledge) the highest reported for polar animals of this type. Extreme diversity in metabolic strategy may explain the bryozoan long evolutionary record and great success in shallow marine environments worldwide.     

Determining cancer risk: the evolutionary multistage model or total stem cell divisions?

A recent hypothesis proposed that the total number of stem cell divisions in a tissue (TSCD model) determine its intrinsic cancer risk; however, a different model—the multistage model—has long been used to understand how cancer originates. Identifying the correct model has important implications for interpreting the frequency of cancers. Using worldwide cancer incidence data, we applied three tests to the TSCD model and an evolutionary multistage model of carcinogenesis (EMMC), a model in which cancer suppression is recognized as an evolving trait, with natural selection acting to suppress cancers causing a significant mean loss of Darwinian fitness. Each test supported the EMMC but contradicted the TSCD model. This outcome undermines results based on the TSCD model quantifying the relative importance of ‘bad luck'' (the random accumulation of somatic mutations) versus environmental and genetic factors in determining cancer incidence. Our testing supported the EMMC prediction that cancers of large rapidly dividing tissues predominate late in life. Another important prediction is that an indicator of recent oncogenic environmental change is an unusually high mean fitness loss due to cancer, rather than a high lifetime incidence. The evolutionary model also predicts that large and/or long-lived animals have evolved mechanisms of cancer suppression that may be of value in preventing or controlling human cancers.     

Complex multicellularity in fungi: evolutionary convergence,single origin,or both?

Complex multicellularity represents the most advanced level of biological organization and it has evolved only a few times: in metazoans, green plants, brown and red algae and fungi. Compared to other lineages, the evolution of multicellularity in fungi follows different principles; both simple and complex multicellularity evolved via unique mechanisms not found in other lineages. Herein we review ecological, palaeontological, developmental and genomic aspects of complex multicellularity in fungi and discuss general principles of the evolution of complex multicellularity in light of its fungal manifestations. Fungi represent the only lineage in which complex multicellularity shows signatures of convergent evolution: it appears 8–11 times in distinct fungal lineages, which show a patchy phylogenetic distribution yet share some of the genetic mechanisms underlying complex multicellular development. To explain the patchy distribution of complex multicellularity across the fungal phylogeny we identify four key observations: the large number of apparently independent complex multicellular clades; the lack of documented phenotypic homology between these clades; the conservation of gene circuits regulating the onset of complex multicellular development; and the existence of clades in which the evolution of complex multicellularity is coupled with limited gene family diversification. We discuss how these patterns and known genetic aspects of fungal development can be reconciled with the genetic theory of convergent evolution to explain the pervasive occurrence of complex multicellularity across the fungal tree of life.     

The predatory behaviour of the thylacine: Tasmanian tiger or marsupial wolf?

The extinct thylacine (Thylacinus cynocephalus) and the extant grey wolf (Canis lupus) are textbook examples of convergence between marsupials and placentals. Craniodental studies confirm the thylacine's carnivorous diet, but little attention has been paid to its postcranial skeleton, which would confirm or refute rare eyewitness reports of a more ambushing predatory mode than the pack-hunting pursuit mode of wolves and other large canids. Here we show that thylacines had the elbow morphology typical of an ambush predator, and propose that the 'Tasmanian tiger' vernacular name might be more apt than the 'marsupial wolf'. The 'niche overlap hypothesis' with dingoes (Canis lupus dingo) as a main cause of thylacine extinction in mainland Australia is discussed in the light of this new information.     

Animal behaviour: emotion in invertebrates?

Bees exposed to vigorous shaking designed to simulate a dangerous event judge ambiguous stimuli as predicting a negative outcome - a 'pessimistic' cognitive bias that is characteristic of anxious or depressed humans and other vertebrates in putative negative emotional states.     

Inefficient nitrogen resorption in genets of the actinorhizal nitrogen fixing shrubComptonia peregrina: physiological ineptitude or evolutionary tradeoff?

Nutrient resorption was measured in an actinorhizal nitrogen-fixing shrub,Comptonia peregrina, for five years in the understory of a deciduous oak forest in Rhode Island, USA. Mean resorption of nitrogen was extremely inefficient (11%) compared to most deciduous species (50%+), yet resorption of phosphorus was efficient (53%) and comparable to other species. Of the seven additional nutrients studied, only copper (6%) and zinc (10%) were resorbed from senescing leaves. Resorption of nitrogen (5%–20%) and phosphorus (40%–71%) varied significantly among years. Copper was resorbed from leaves in three years and accreted into leaves in two years. Five-year resorption means differed among individual genets by as much as a factor of 2.5 for nitrogen, and 1.3 for phosphorus. Resorption of nitrogen, copper, and zinc were highly correlated, yet resorption of phosphorus remained autonomous from other nutrients. The ecophysiological tradeoffs inComptonia which have resulted in the cooccurence of actinorhizal nitrogen fixation, inefficient nitrogen resorption, and efficient phosphorus resorption suggest that plant nutrient status does have an impact on resorption efficiency and that the evolution of nutrient conservation strategies is nutrient-specific.