Sex in an Evolutionary Perspective: Just Another Reaction Norm

It is common to refer to all sorts of clear-cut differences between the sexes as something that is biologically almost inevitable. Although this does not reflect the status of evolutionary theory on sex determination and sexual dimorphism, it is probably a common view among evolutionary biologists as well, because of the impact of sexual selection theory. To get away from thinking about biological sex and traits associated with a particular sex as something static, it should be recognized that in an evolutionary perspective sex can be viewed as a reaction norm, with sex attributes being phenotypically plastic. Sex determination itself is fundamentally plastic, even when it is termed “genetic”. The phenotypic expression of traits that are statistically associated with a particular sex always has a plastic component. This plasticity allows for much more variation in the expression of traits according to sex and more overlap between the sexes than is typically acknowledged. Here we review the variation and frequency of evolutionary changes in sex, sex determination and sex roles and conclude that sex in an evolutionary time-frame is extremely variable. We draw on recent findings in sex determination mechanisms, empirical findings of morphology and behaviour as well as genetic and developmental models to explore the concept of sex as a reaction norm. From this point of view, sexual differences are not expected to generally fall into neat, discrete, pre-determined classes. It is important to acknowledge this variability in order to increase objectivity in evolutionary research.     

Direct use values and passive use values: implications for conservation of large carnivores

We made a quantitative analysis of the responses of urban and rural residents in Sweden to arguments supporting and opposing conservation of large carnivores. The most important arguments in favour of conservation were: “I want them [the large carnivores] to exist in Sweden, even if I will never see any of them”, “Sweden should share the responsibility of conserving the large carnivores” and “We owe it [conservation of large carnivores] to future generations”. We found only small differences between rural and urban residents. For arguments opposing conservation, the difference between rural and urban areas was slightly greater. The most important arguments opposing conservation of large carnivores were: “They may have serious negative impact on livestock farming”, “They may have serious negative impact on reindeer husbandry” and “May inflict suffering on injured livestock”. We conclude that there seems to be less support for direct use values such as hunting, ecotourism or just experiencing large carnivores, this may imply that the minimum viable population size can be used as a long-term management goal for large carnivore populations, possibly with an exception for bears. We also conclude that a separate conservation or management plan is needed for each species, since the conflicts with human interests vary greatly between the different carnivore species.     

Ecological explanations for successful invasion of exotic plants

The intentional introduction of exotic species can increase the level of local biodiversity, enrich people’s material lives, and bring significant social and economic benefits that are also the symbols of human progress. However, along with the frequent intercourse among countries and regions, the frequency of uncontrolled crossregional migration of species is increased and there is a lack of scientific management strategy for the intentional introduction of exotic species. Exotic species invasion, which is behind habitat fragmentation, has become the second largest threatening factor to the maintenance of the global-scale level of biological diversity. Exotic species invasion can destroy the structure of an ecosystem, disturb the economic life of a society, and do harm to human health. In this paper, the authors review some of the ecological explanations for issues such as “what causes or mechanisms have led to the successful invasion of exotic species”, including the “ideal weeds characteristics”, “biodiversity resistance hypothesis”, “enemies release hypothesis”, “evolution of increased competitive ability hypothesis”, “niche opportunity hypothesis”, and “novel weapon hypothesis”. The authors also analyze and evaluate the background and theoretical basis of the hypotheses, providing explanations for some phenomena, as well as the deficiencies of these explanations.     

Evolutionary biology and feminism

Evolutionary biology and feminism share a variety of philosophical and practical concerns. I have tried to describe how a perspective from both evolutionary biology and feminism can accelerate the achievement of goals for both feminists and evolutionary biologists. In an early section of this paper I discuss the importance of variation to the disciplines of evolutionary biology and feminism. In the section entitled “Control of Female Reproduction” I demonstrate how insight provided by participation in life as woman and also as a feminist suggests testable hypotheses about the evolution of social behavior—hypotheses that are applicable to our investigations of the evolution of social behavior in nonhuman animals. In the section on “Deceit, Self-deception, and Patriarchal Reversals” I have overtly conceded that evolutionary biology, a scientific discipline, also represents a human cultural practice that, like other human cultural practices, may in parts and at times be characterized by deceit and self-deception. In the section on “Femininity” I have indicated how questions cast and answered and hypotheses tested from an evolutionary perspective can serve women and men struggling with sexist oppression. Patricia Adair Gowaty studies the evolution of social behavior, particularly mating systems and sex allocation, primarily in birds. She is most well-known for her long-term studies of eastern bluebirds, which began in 1977 and are on-going. She was an undergraduate at H. Sophie Newcomb College of Tulane University (1963–1967). In the late sixties and early seventies, while employed at the Bronx Zoo (New York Zoological Society), she belonged to a feminist “consciousness-raising” group. She started graduate school in 1974 at the University of Georgia and received her Ph.D. from Clemson University (1980). She had a postdoctoral position at the University of Oklahoma (1982–1983) and a visiting faculty position at Cornell University through the Visiting Professorships for Women NSF program (1983–1984) before returning to her bluebird study sites at Clemson in 1985. She has supported herself and her research efforts throughout her academic career on a series of awards and grants. She is currently (1990–1995) supported by a Research Scientist Development Award from The National Institute of Mental Health.     

Sex differences in optimal incubation temperatures in a scincid lizard species

Most theoretical models for the evolution of temperature-dependent sex determination (TSD) rely upon differential fitness of male and female offspring incubated under different thermal regimes. However, there are few convincing data on this topic. We studied incubation effects in a lizard species (Bassiana duperreyi, Scincidae) with genotypic sex determination, so that we could separate effects due to incubation temperatures from those due to offspring gender. We incubated eggs under two different fluctuating-temperature regimes that simulated hot and cold natural nest-sites. The effects of our incubation treatments on phenotypes of the hatchling lizards (morphology and locomotor performance) differed between the sexes. Females emerging from eggs exposed to the “hot nest” treatment (diel cycling, 23–31°C) were larger, and ran faster, than did their sisters from the “cold nest” treatment (16–24°C). Males showed a smaller and less consistent phenotypic response than females. These incubation-induced responses were relatively stable during the first few weeks of life post-hatching, at least in captive lizards maintained under laboratory conditions. These kinds of sex differences in the phenotypic responses of hatchling reptiles to incubation conditions provide a plausible basis for the evolution of temperature-dependent sex determination in reptiles. Received: 07 May 1998 / Accepted: 16 November 1998     

The advantages and disadvantages of being introduced

Introduced species, those dispersed outside their natural ranges by humans, now cause almost all biological invasions, i.e., entry of organisms into habitats with negative effects on organisms already there. Knowing whether introduction tends to give organisms specific ecological advantages or disadvantages in their new habitats could help understand and control invasions. Even if no specific species traits are associated with introduction, introduced species might out-compete native ones just because the pool of introduced species is very large (“global competition hypothesis”). Especially in the case of intentional introduction, high initial propagule pressure might further increase the chance of establishment, and repeated introductions from different source populations might increase the fitness of introduced species through hybridization. Intentional introduction screens species for usefulness to humans and so might select for rapid growth and reproduction or carry species to suitable habitats, all which could promote invasiveness. However, trade offs between growth and tolerance might make introduced species vulnerable to extreme climatic events and cause some invasions to be transient (“reckless invader hypothesis”). Unintentional introduction may screen for species associated with human-disturbed habitats, and human disturbance of their new habitats may make these species more invasive. Introduction and natural long-distance dispersal both imply that species have neither undergone adaptation in their new habitats nor been adapted to by other species there. These two characteristics are the basis for many well-known hypotheses about invasion, including the “biotic resistance”, “enemy release”, “evolution of increased competitive ability” and “novel weapon” hypotheses, each of which has been shown to help explain some invasions. To the extent that biotic resistance depends upon local adaption by native species, altering selection pressures could reduce resistance and promote invasion (“local adaptation hypothesis”), and restoring natural regimes could reverse this effect.     

Fame,fortune, and fitness at the Academy Awards

People across many societies routinely participate in physical or intellectual competitions in the absence of immediate substantial monetary or other apparent material rewards. But increased fame and social status associated with awards, such as the “Oscars”, need not be necessarily and solely a cultural construct unrelated to natural selection. Rather, prizes might be badges of honor if they are also honest indicators of evolutionary fitness. Analyses of reported reproductive success data, from a survey of well-known female and male actors, followed previously reported patterns of biological fitness in this sample of a human population. In addition, the numbers of Academy Awards received for acting were positively associated with reported numbers of biological children for both genders. The association of increased fitness with more awards received was statistically consistent even when considering that this subset of the population conformed to the Bateman effect in human reproduction: male actors had a more positive correlation than females between cumulative numbers of married partners and overall numbers of children. Honest signals of reproductive quality that are displayed by both sexes are expected to occur in humans and other species with costly biparental care and mutual mate choice.     

Abnormal births and other “ill omens”

We summarize the ethnographic literature illustrating that “abnormal birth” circumstances and “ill omens” operate as cues to terminate parental investment. A review of the medical literature provides evidence to support our assertion that ill omens serve as markers of biological conditions that will threaten the survival of infants. Daly and Wilson (1984) tested the prediction that children of demonstrably poor phenotypic quality will be common victims of infanticide. We take this hypothesis one stage further and argue that some children will be poor vehicles for parental investment yet are not of demonstrably poor quality at birth. We conclude that when people dispose of infants due to “superstitious beliefs” they are pursuing an adaptive strategy in eliminating infants who are poor vehicles for parental investment. Catherine Hill lectures in biological anthropology/human sciences at Durham University’s University College, Stockton. She trained in biological anthropology at University College, London. Her current research interests include human and nonhuman primate socioecology and human resource ecology and development issues. Helen Ball lectures in biological anthropology/human sciences at Durham University’s University College, Stockton. She trained in biological anthropology at the University of Massachusetts at Amherst. Her current research interests include nonhuman primate behavior and socioecology, reproductive biology, and evolutionary issues.     

Modelling mate-finding Allee effects and populations dynamics,with applications in pest control

In this paper, we review how mate-finding Allee effects enter population dynamical models that consider both sexes, highlight possible limitations of the more widely used “one-sex” models, and outline the links between the different model classes. We further explore interactions between the mate-finding Allee effect and other mechanisms relevant to pest-control strategies: release of natural enemies, sterile male release, and culling. Many of these strategies impose an additional component Allee effect on the population, and we discuss which of them might be efficient in the control of pest species that also suffer from the failure to locate mates. We focus primarily on eradication thresholds; our simple models show that most of the strategies yield similar results, and depending on the costs, one strategy or a combination of several can lead to the most efficient control.     

Indicators for plant species richness in pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) forests of Germany

Forestry is obliged to record as well as maintain and/or enhance biological diversity in forests due to national and international agreements. Accordingly, it is necessary to work out methodological approaches for the assessment of biodiversity in forests. In the study presented here, we focus on the total plant species pool (563 vascular plant and bryophyte species) of pine (Pinus sylvestris L.) forests in NE Germany to identify indicators for plant species richness. We distinguished several groups like “herb”, “grass-like”, “woody”, “endangered”, and “exotic species”, for which we detected indicators for low (class #1), intermediate (class #2), and high (class #3) species numbers. From a total of 84 species, which were identified by a three-step procedure, most indicators were found for class #3. Only few indicators have been revealed for intermediate species numbers, i.e. class #2. With help of Ellenberg’s ecological indicator values and information on the main occurrence in Central European vegetation types and plant communities, respectively, we characterized the indicator species ecologically. The ecological site preferences of the indicator species in general reflect the fact that species richness is highest in base-rich, light, and anthropogenically disturbed pine forests. On the contrary, species-poor forests were revealed by indicators, which mainly occur on acidic sites. It is concluded that a considerable set of indicators for species richness can help facilitate biodiversity assessments in forestry and ecosystem restoration practice. Electronic Supplementary Material  The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

Blogging Evolution

A weblog (“blog”) is an publication on the WorldWideWeb in which brief entries are displayed in date order, much like a diary or journal. I describe the general characteristics of blogs, contrasting blogs with other of WWW formats for self-publishing. I describe four categories for blogs about evolutionary biology: “professional,” “amateur,” “apostolic,” and “imaginative.” I also discuss blog networks. I identify paradigms of each category. Throughout, I aim to illuminate blogs about evolutionary biology from the point of view of a user looking for information about the topic. I conclude that blogs are not the best type of source for systematic and authoritative information about evolution, and that they are best used by the information-seeker as a way of identifying what issues are of interest in the community of evolutionists and for generating research leads or fresh insights on one’s own work.

From “Us vs. Them” to “Shared Risk”: Can Animals Help Link Environmental Factors to Human Health?

Linking human health risk to environmental factors can be a challenge for clinicians, public health departments, and environmental health researchers. While it is possible that nonhuman animal species could help identify and mitigate such linkages, the fields of animal and human health remain far apart, and the prevailing human health attitude toward disease events in animals is an “us vs. them” paradigm that considers the degree of threat that animals themselves pose to humans. An alternative would be the development of the concepts of animals as models for environmentally induced disease, as well as potential “sentinels” providing early warning of both noninfectious and infectious hazards in the environment. For such concepts to truly develop, critical knowledge gaps need to be addressed using a “shared risk” paradigm based on the comparative biology of environment–host interactions in different species.     

Human neuropsychology and the concept of culture

American anthropology is distinguished by a four-fields approach in which biological, cultural, archaeological, and linguistic dimensions of behavior are examined in evolutionary and cross-cultural perspective. Nevertheless, assumptions of mind-body dualism pervade scholarly thinking in anthropology and have prevented the development of a truly integrated science of human experience. This dualism is most exemplified by the lack of consideration of the role of the brain in both “physical” and “mental” processes, including phenomena labeled as cultural. In this paper, I review neural mechanisms of learning, communication, and emotion, and discuss the implications of these findings for culture theory. Lee Xenakis Blonder is an assistant professor in the Department of Behavioral Science and the Sanders-Brown Center on Aging, University of Kentucky Medical School, Lexington. She is currently examining the effects of stroke in different regions of the brain on language, nonverbal communication, and emotional processing in an attempt to better understand human brain and behavior relations. Recent publications include “Neuropsychological Functioning in Hemiparkinsonism” (with R. E. Gur, R. C. Gur, A. J. Saykin, and H. I. Hurtig),Brain and Cognition 9:244–257 (1989).     

Apomorphy-based definition also pinpoints a node, and PhyloCode names prevent effective communication

Acceptable methods of defining taxon (or clade) names in the draft PhyloCode, or so-called phylogenetic nomenclature, are “node based,” “stem based,” and “apomorphy based.” All of them define a clade name by pinpointing a node; whereas node-based and stem-based definitions require two or more taxon “specifiers” to define names, an apomorphy-based definition requires two specifiers of different types; namely, a single-taxon specifier and a character specifier. The taxon specifier in an apomorphy-based definition is completely different from the “type” in the Linnaean system. Taxon (or clade) names in the PhyloCode are characterized in two entirely different manners: One is a name that does not change, either in its orthography or in the contents of the taxon referred to by it (or its meaning) over time; the other is a name that is just like a pure mark and thus has no meaning. Communication through such PhyloCode names is very ineffective or impossible.     

Genomic screening in dioecious “yerba mate” tree (<Emphasis Type="Italic">Ilex paraguariensis</Emphasis> A. St. Hill., Aquifoliaceae) through representational difference analysis

The “yerba mate” tree, Ilex paraguariensis, is a functionally dioecious crop species with economic relevance in several South American countries. We report a genomic screening accomplished through representational difference analysis (RDA) in male and female I. paraguariensis trees. The aim of the present paper was to investigate the occurrence of sex-related genomic differences in order to develop an early gender detection molecular method that could help reducing energy inputs during the “yerba mate” processing and that could be suitable for breeding programs. An intra-experiment redundancy was detected via SSCP analysis and sequence characterization. Taking together both reciprocal RDA assays, fragments isolated can be discriminated into three main categories. The first category of fragments shows spurious affinities with available deposited sequences and could be considered as specific to I. paraguariensis. The second category comprises sequences identified as organellar or ribosomal plant DNA. Sequences grouped in the third category involve clones akin to conserved domains of retrotransposons (RNaseH, integrases and/or chromodomains) from at least two distinct lineages of Ty3/Gypsy retrotransposons and one from Ty1/Copia retroelements, which in addition are associated to sex determination regions of the Solanaceae, Caricaceae and Salicaceae. A contig sequence was assembled that codes for an integrase core domain and a chromodomain. The phylogenetic analysis of the so-called IPRE (for I. paraguariensis retroelement) integrase domain indicates that it belongs to the Del lineage of the Chromoviridae. This is the first report of mobile elements isolated and detected from the “yerba mate” tree. Although RDA derived fragments, so far tested, have been retrieved from both sexes with similar sequences, association to sex related regions cannot be completely discarded. Implications of present results are further discussed.     

Recognition of sex in the acoustic communication of the grasshopper Chorthippus biguttulus (Orthoptera, Acrididae)

Many gomphocerine grasshoppers communicate acoustically: a male's calling song is answered by a female which is approached phonotactically by the male. Signals and recognition mechanisms were investigated in Chorthippus biguttulus with regard to the cues which allow sex discrimination. (1) The stridulatory files on the hindfemur of both sexes are homologous in that they are derived from the same row of bristles, but convergent with respect to the “pegs”. In males the pegs are derived from the bristles, and in females from the wall of the bristle's cup. (2) Male and female songs are generated by similar, probably homologous motor programs, but differ in the duration, intensity, “gappyness” of syllables, risetime of pulses, and the frequency spectra. The hindleg co-ordination during stridulation and the resulting temporal song patterns are less variable in males than in females. (3) For both sexes, recognition of a mate's signal depends on species-specific syllable structure. For males it is essential that the female syllables consist of distinct short pulses, whereas females reject “gappy” syllables. Males strongly prefer “ramped” pulses, females respond to syllables irrespective of steeply or slowly rising ramps. Males react only to the low-frequency component, whereas females prefer spectra containing both, low and high frequency components. Accepted: 20 November 1996     

Inventory, differentiation, and proportional diversity: a consistent terminology for quantifying species diversity

Almost half a century after Whittaker (Ecol Monogr 30:279–338, 1960) proposed his influential diversity concept, it is time for a critical reappraisal. Although the terms alpha, beta and gamma diversity introduced by Whittaker have become general textbook knowledge, the concept suffers from several drawbacks. First, alpha and gamma diversity share the same characteristics and are differentiated only by the scale at which they are applied. However, as scale is relative––depending on the organism(s) or ecosystems investigated––this is not a meaningful ecological criterion. Alpha and gamma diversity can instead be grouped together under the term “inventory diversity.” Out of the three levels proposed by Whittaker, beta diversity is the one which receives the most contradictory comments regarding its usefulness (“key concept” vs. “abstruse concept”). Obviously beta diversity means different things to different people. Apart from the large variety of methods used to investigate it, the main reason for this may be different underlying data characteristics. A literature review reveals that the multitude of measures used to assess beta diversity can be sorted into two conceptually different groups. The first group directly takes species distinction into account and compares the similarity of sites (similarity indices, slope of the distance decay relationship, length of the ordination axis, and sum of squares of a species matrix). The second group relates species richness (or other summary diversity measures) of two (or more) different scales to each other (additive and multiplicative partitioning). Due to that important distinction, we suggest that beta diversity should be split into two levels, “differentiation diversity” (first group) and “proportional diversity” (second group). Thus, we propose to use the terms “inventory diversity” for within-sample diversity, “differentiation diversity” for compositional similarity between samples, and “proportional diversity” for the comparison of inventory diversity across spatial and temporal scales. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

An ethological analysis of types of agonistic interaction in a captive group of Java-monkeys (Macaca fascicularis)

The primate literature provides many indications not only that the nature of dyadic interactions is to a large extent determined by the relations of the interacting animals with others and between these others, but also of the existence of polyadic interactions in which more than two individuals are simultaneously involved. The objectives of the present study are to obtain a quantitative categorization of the agonistic interaction types of captive Java-monkeys and an analysis of their dynamics. After having described the agonistic behaviour patterns of Java-monkeys we shall discuss the categorization of agonistic interaction types (depending on the number of involvees: “dyads”, “triads” and “polyads”), the way in which these types can be further differentiated on the basis of the nature and the direction of the behaviours shown (e.g., different types of alliances), and the existence of so-called “sub-directed” behaviours (i.e., non-agonistic behaviours which are shown towards a dominant third animal more or less simultaneously with aggressive behaviour directed towards an opponent). The analysis indicates that agonistic behaviour is different both in its form and its regulation in interactions of different complexity. This research was supported in part by a government grant (i.e.: Beleidsruimte project: 16-21-06, “Brain and Behaviour”) to the first author. The investigation was supported by a grant from the Beleidsruimtemiddelen Hersenen en Gedrag to the first author.     

Grandmothers,hunters and human life history

This paper critiques the competing “Grandmother Hypothesis” and “Embodied Capital Theory” as evolutionary explanations of the peculiarities of human life history traits. Instead, I argue that the correct explanation for human life history probably involves elements of both hypotheses: long male developmental periods and lives probably evolved due to group selection for male hunting via increased female fertility, and female long lives due to the differential contribution women’s complex foraging skills made to their children and grandchildren’s nutritional status within groups provisioned by male hunting.     

Psychological Altruism Vs. Biological Altruism: Narrowing The Gap With The Baldwin Effect

This paper defends the position that the supposed gap between biological altruism and psychological altruism is not nearly as wide as some scholars (e.g., Elliott Sober) insist. Crucial to this defense is the use of James Mark Baldwin's concepts of “organic selection”and “social heredity” to assist in revealing that the gap between biological and psychological altruism is more of a small lacuna. Specifically, this paper argues that ontogenetic behavioral adjustments, which are crucial to individual survival and reproduction, are also crucial to species survival. In particular, it is argued that human psychological altruism is produced and maintained by various sorts of mimicry and self-reflection in the aid of both individual and species survival. The upshot of this analysis is that it is possible to offer an account of psychological altruism that is closelytethered to biological altruism without reducing entirely the former to thelatter.