Harder,better, faster,stronger: Weapon size is more sexually dimorphic than weapon biomechanical components in two freshwater anomuran species

Sexual selection influences the evolution of morphological traits that increase the likelihood of monopolizing scarce resources. When such traits are used during contests, they are termed weapons. Given that resources are typically linked to monopolizing mating partners, theory expects only males to bear weapons. In some species, however, females also bear weapons, although typically smaller than male weapons. Understanding why females bear smaller weapons can thus help us understand the selective pressures behind weapon evolution. However, most of our knowledge comes from studies on weapon size, while the biomechanics of weapons, such as the size of the muscles, efficiency, and shape are seldom studied. Our goal was to test if the theoretical expectations for weapon size sexual dimorphism also occur for weapon biomechanics using two aeglid crab species. Males of both species had larger claws which were also stronger than female claws. Male claws were also more efficient than females' claws (although we used only one species in this analysis). For weapon shape, though, only one species differed in the mean claw shape. Regarding scaling differences, in both species, male claws had higher size scaling than females, while only one species had a higher shape scaling. However, male weapons did not have higher scaling regarding strength and efficiency than females. Thus, males apparently allocate more resources in weapons than females, but once allocated, muscle and efficiency follow a similar developmental pathway in both sexes. Taken together, our results show that sexual dimorphism in weapons involves more than differences in size. Shape differences are especially intriguing because we cannot fully understand its causes. Yet, we highlight that such subtle differences can only be detected by measuring and analysing weapon shape and biomechanical components. Only then we might better understand how weapons are forged.     

BACTERIAL AND CHEMICAL WARFARE-The Current Status

For fourteen years public attention has been focused so sharply on atomic weapons as to lose sight of other, less spectacular but equally significant advances in the art of warfare. In the shadows cast by brilliant research in nuclear physics are hidden startling advances in the field of chemical and biological weapons. These weapons, as now developed, are not only capable of producing mass casualties quite comparable with those of atomic bombs, but they also possess certain advantages which may make them the weapons of choice for an unscrupulous enemy. If war should come, it is the medical profession which will have the sole responsibility for protecting the citizens of California against these weapons, and we can therefore delay no longer in acquainting ourselves with their potentialities and characteristics. In this task, we are working under two serious handicaps. The first is that our classical medical training affords little appreciation of the real danger, and the second is the cloak of secrecy surrounding the entire subject.     

BACTERIAL AND CHEMICAL WARFARE—The Current Status

For fourteen years public attention has been focused so sharply on atomic weapons as to lose sight of other, less spectacular but equally significant advances in the art of warfare.In the shadows cast by brilliant research in nuclear physics are hidden startling advances in the field of chemical and biological weapons. These weapons, as now developed, are not only capable of producing mass casualties quite comparable with those of atomic bombs, but they also possess certain advantages which may make them the weapons of choice for an unscrupulous enemy.If war should come, it is the medical profession which will have the sole responsibility for protecting the citizens of California against these weapons, and we can therefore delay no longer in acquainting ourselves with their potentialities and characteristics.In this task, we are working under two serious handicaps. The first is that our classical medical training affords little appreciation of the real danger, and the second is the cloak of secrecy surrounding the entire subject.     

Is diversification in male reproductive traits driven by evolutionary trade-offs between weapons and nuptial gifts?

Many male animals have evolved exaggerated traits that they use in combat with rival males to gain access to females and secure their reproductive success. But some male animals invest in nuptial gifts that gains them access to females. Both these reproductive strategies are costly in that resources are needed to produce the weapon or nuptial gift. In closely related species where both weapons and nuptial gifts are present, little is known about the potential evolutionary trade-off faced by males that have these traits. In this study, we use dobsonflies (order Megaloptera, family Corydalidae, subfamily Corydalinae) to examine the presence and absence of enlarged male weapons versus nuptial gifts within and among species. Many dobsonfly species are sexually dimorphic, and males possess extremely enlarged mandibles that they use in battles, whereas in other species, males produce large nuptial gifts that increase female fecundity. In our study, we show that male accessory gland size strongly correlates with nuptial gift size and that when male weapons are large, nuptial gifts are small and vice versa. We mapped weapons and nuptial gifts onto a phylogeny we constructed of 57 species of dobsonflies. Our among-species comparison shows that large nuptial gift production evolved in many species of dobsonfly but is absent from those with exaggerated weapons. This pattern supports the potential explanation that the trade-off in resource allocation between weapons and nuptial gifts is important in driving the diversity of male mating strategies seen in the dobsonflies, whereas reduced male–male competition in the species producing large spermatophores could be an alternative explanation on their loss of male weapons. Our results shed new light on the evolutionary interplay of multiple sexually selected traits in animals.     

Experimental manipulation reveals a trade‐off between weapons and testes

Theory predicts a trade‐off between sexually selected weapons used to secure mates and post‐copulatory traits used to maximize fertilization success. However, individuals that have a greater capacity to acquire resources from the environment may invest more in both pre‐ and post‐copulatory traits, and trade‐offs may not be readily apparent. Here, we manipulate the phenotype of developing individuals to examine allocation trade‐offs between weapons and testes in Mictis profana (Hemiptera: Coreidae), a species where the hind legs are sexually selected weapons used in contests over access to females. We experimentally prevented males from developing weapons by inducing them to autotomize their hind legs before the final moult to adulthood. We compared trait expression in this group to males where autotomy was induced in the mid‐legs, which are presumably not under sexual selection to the same extent. We found males without weapons invested proportionally more in testes mass than those with their mid‐legs removed. Males that developed to adulthood without weapons did not differ from the mid‐leg removal group in other traits potentially under precopulatory sexual selection, other post‐copulatory traits or naturally selected traits. In addition, a sample of adult males from the same population in the wild revealed a positive correlation between investment in testes and weapons. Our study presents a critical contribution to a growing body of literature suggesting the allocation of resources to pre‐ and post‐copulatory sexual traits is influenced by a resource allocation trade‐off and that this trade‐off may only be revealed with experimental manipulation.     

Preface

Abstract

Major naval weapons and weapons systems are examined as they relate to naval strategy and the law of the sea. Although there are some legal limitations on the development and use of nuclear and other mass destruction weapons on the seabed, and in nuclear free zones such as Latin America, and on naval weaponry in Antarctica, the use of naval weapons systems is largely unrestricted by international law and the law of the sea. Weapons systems from mines to large submarines and carriers are related to areas, to categories of states, and to the main sub‐fields of international relations. The conclusion to be drawn is that a third Hague Conference on the laws of war should be called and arms control and disarmament issues in naval weaponry should be addressed urgently. Detailed data on naval weapons, missile systems, and maritime zones are presented in three appendixes. Glossary of Abbreviations for Naval Weaponry immediately precedes the appendices.     

Scientific progress and new biological weapons

The biological weapons are different from conventional weapons, because living germs hold an extraordinary and predictable potential for multiplication, propagation and genetic variation during their dissemination in a susceptible population. Only natural pathogens (1rst generation weapons) have been used in the past (smallpox virus, plague, anthrax, toxins...). However, new threats are emerging, due to the rapid progress of scientific knowledge and its exponential worldwide diffusion. It is possible to synthesize microorganisms from in silico sequences widely diffused on Internet (poliovirus, influenza...), thus resulting in the accessibility of very dangerous virus confined today in high-security laboratories (virus Ebola...). It is possible also to "improve" pathogens by genetic manipulations, becoming more resistant or virulent (2nd generation weapons). Finally, one can now create de novo new pathogens by molecular breeding (DNA shuffling), potentially highly dangerous for naive populations (3rd generation weapons). Making biological weapons does not require too much technological resources and appears accessible to terrorists, due to low cost and easy use. Although the destructive consequences are difficult to predict, the psychological and social damages should be considerable, because of the highly emotional burden in the population associated to the transgression by man of a taboo of life.     

Ownership-dependent mating tactics of minor males of the beetle <Emphasis Type="Italic">Librodor japonicus</Emphasis> (Nitidulidae) with intra-sexual dimorphism of mandibles

Intra-sexual dimorphism is found in the weapons of many male beetles. Different behavioral tactics to access females between major and minor males, which adopt fighting and alternative tactics, respectively, are thought to maintain the male dimorphism. In these species major males have enlarged weapons that they use in fights with rival males. Minor males also have small weapons in some of these species, and it is unclear why these males possess weapons. We examined the hypothesis that minor males might adopt a fighting tactic when their status was relatively high in comparison with that of other males (e.g., ownership of a territory). We observed the behavioral tactics of major and minor males of the beetle Librodor japonicus, whose males have a dimorphism of their mandibles. Major males fought for resources, whereas minor males adopted two status-dependent tactics, fighting and sneaking, to access females, depending on their ownership of a sap site. We suggest that ownership status-dependent mating tactics in minor males may maintain the intra-sexual dimorphism in this beetle.     

The Effects of Extra-Somatic Weapons on the Evolution of Human Cooperation towards Non-Kin

Human cooperation and altruism towards non-kin is a major evolutionary puzzle, as is ‘strong reciprocity’ where no present or future rewards accrue to the co-operator/altruist. Here, we test the hypothesis that the development of extra-somatic weapons could have influenced the evolution of human cooperative behaviour, thus providing a new explanation for these two puzzles. Widespread weapons use could have made disputes within hominin groups far more lethal and also equalized power between individuals. In such a cultural niche non-cooperators might well have become involved in such lethal disputes at a higher frequency than cooperators, thereby increasing the relative fitness of genes associated with cooperative behaviour. We employ two versions of the evolutionary Iterated Prisoner''s Dilemma (IPD) model – one where weapons use is simulated and one where it is not. We then measured the performance of 25 IPD strategies to evaluate the effects of weapons use on them. We found that cooperative strategies performed significantly better, and non-cooperative strategies significantly worse, under simulated weapons use. Importantly, the performance of an ‘Always Cooperate’ IPD strategy, equivalent to that of ‘strong reciprocity’, improved significantly more than that of all other cooperative strategies. We conclude that the development of extra-somatic weapons throws new light on the evolution of human altruistic and cooperative behaviour, and particularly ‘strong reciprocity’. The notion that distinctively human altruism and cooperation could have been an adaptive trait in a past environment that is no longer evident in the modern world provides a novel addition to theory that seeks to account for this major evolutionary puzzle.     

Selection on defensive traits in a sterile caste – caste evolution: a mechanism to overcome life‐history trade‐offs?

SUMMARY During development and evolution individuals generally face a trade-off between the development of weapons and gonads. In termites, characterized by reproductive division of labor, a caste evolved—the soldiers—which is completely sterile and which might be released from developmental trade-offs between weapons and testes. These soldiers are exclusively dedicated to defense. First, we investigated whether defensive traits are under selection in sterile termite soldiers using allometric analyses. In soldiers of the genus Cryptotermes phragmotic traits such as a sculptured and foreshortened head evolve rapidly but were also lost twice. Second, we compared the scaling relationships of these weapons with those in solitary insects facing a trade-off between weapons and gonads. Defensive traits consistently had lower slopes than nondefensive traits which supports the existence of stabilizing selection on soldier phragmotic traits in order to plug galleries. Moreover, soldier head widths were colony specific and correlated with the minimum gallery diameter of a colony. This can proximately be explained by soldiers developing from different instars. The scaling relationships of these termite soldiers contrast strikingly with those of weapons of solitary insects, which are generally exaggerated (i.e., overscaling) male traits. These differences may provide important insights into trait evolution. Trade-offs constraining the development of individuals may have been uncoupled in termites by evolving different castes, each specialized for one function. When individuals in social insect are "released" from developmental constraints through the evolution of castes, this certainly contributed to the ecological and evolutionary success of social insects.     

A proposal for the classification of biological weapons sensu lato

Due to historical and legislation reasons, the category of bioweapons is rather poorly defined. Authors often disagree on involving or excluding agents like hormones, psychochemicals, certain plants and animals (such as weeds or pests) or synthetic organisms. Applying a wide definition apparently threatens by eroding the regime of international legislation, while narrow definitions abandon several important issues. Therefore, I propose a category of ‘biological weapons sensu lato’ (BWsl) that is defined here as any tool of human aggression whose acting principle is based on disciplines of biology including particularly microbiology, epidemiology, medical biology, physiology, psychology, pharmacology and ecology, but excluding those based on inorganic agents. Synthetically produced equivalents (not necessarily exact copies) and mock weapons are also included. This definition does not involve any claim to subject all these weapons to international legislation but serves a purely scholarly purpose. BWsl may be properly categorized on the base of the magnitude of the human population potentially targeted (4 levels: individuals, towns, countries, global) and the biological nature of the weapons’ intended effects (4 levels: agricultural-ecological agents, and non-pathogenic, pathogenic, or lethal agents against humans).     

Social selection and the evolution of a female weapon in queens of the ant Messor pergandei (Hymenoptera: Formicidae)

The evolution of weaponry occurs less frequently in females than in males and is most often important for protecting ecological resources or offspring rather than winning mates. The purpose of female weapons is often confounded by the presence of similar weapons in males, so cases where only females need weapons provide important tests of our understanding of how and why weapons evolve. In some populations of the ant, Messor pergandei (Mayr), newly mated queens initiate new nests in social groups that subsequently break down when queens engage in battles for control. The incipient social environment differs geographically, so that lethal fighting occurs in some populations but not others. Consistent with the hypothesis that queens in populations where lethal fighting occurs should show selection for weaponry (broad heads and strong mandibles), we found that heads of queens from sites where lethal fighting occurs were broader than those at sites with non‐fighting queens and a site with solitary queens. Evolution of weaponry is specific to queens, because regression results from workers often did not follow this pattern. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 1011–1020.     

Weapon performance drives weapon evolution

Many sexually selected traits function as weapons, and these weapons can be incredibly diverse. However, the factors underlying weapon diversity among species remain poorly understood, and a fundamental hypothesis to explain this diversity remains untested. Although weapons can serve multiple functions, an undeniably important function is their role in fights. Thus, a crucial hypothesis is that weapon diversification is driven by the evolution of weapon modifications that provide an advantage in combat (e.g. causing more damage). Here, we test this fighting-advantage hypothesis using data from 17 species of coreid bugs. We utilize the fact that male–male combat in coreids often results in detectable damage, allowing us to link different weapon morphologies to different levels of damage among species. We find that certain weapon morphologies inflict much more damage than others, strongly supporting the fighting-advantage hypothesis. Moreover, very different weapon morphologies can inflict similarly severe amounts of damage, leading to a weapon performance landscape with multiple performance peaks. This multi-peak pattern could potentially drive different lineages towards divergent weapon forms, further increasing weapon diversity among species. Overall, our results may help explain how sexually selected weapons have evolved into the diversity of forms seen today.     

Bioterrorism: responding to an emerging threat

Only a few years ago bioterrorism was considered a remote concern but few today are complacent about the possibility of biological agents being intentionally used to cause widespread panic, disruption, disease and death. By its very nature, the biological weapons threat - with its close links to naturally occurring infectious agents and disease - requires a different paradigm than that for conventional terrorism, military strikes or attacks caused by other weapons of mass destruction. This evolving threat presents the medical, public health and scientific communities (importantly including biotechnology) with a set of difficult and pressing challenges. This article provides a brief overview of the threat from biological weapons, the nature of a bioterrorist attack and some of the issues that need to be addressed if we are to make meaningful progress to prevent or contain this disturbing and potentially catastrophic danger.     

Maximum shields: the assembly and function of the bacterial spore coat

Spores produced by bacilli and clostridia are surrounded by a multilayered protein shell called the coat. As the armor-like appearance of the coat suggests, this structure, along with others within the spore, confers the remarkable resistance properties that make Bacillus anthracis spores such potent biological weapons. Here, I review recent studies of coat assembly in the model organism Bacillus subtilis, and explore the implications of these findings for coat assembly in B. anthracis and for defense against biological weapons.     

Anti-predator behaviour depends on male weapon size

Tonic immobility and escape are adaptive anti-predator tactics used by many animals. Escape requires movement, whereas tonic immobility does not. If anti-predator tactics relate to weapon size, males with larger weapons may adopt tonic immobility, whereas males with smaller weapons may adopt escape. However, no study has investigated the relationship between weapon size and anti-predator tactics. In this study, we investigated the relationship between male weapon size and tonic immobility in the beetle Gnathocerus cornutus. The results showed that tonic immobility was more frequent in males with larger weapons. Although most studies of tonic immobility in beetles have focused on the duration, rather than the frequency, tonic immobility duration was not affected by weapon size in G. cornutus. Therefore, this study is the first, to our knowledge, to suggest that the male weapon trait affects anti-predator tactics.     

Hazards,threats and risks: lessons from the past to a defensive attitude for the future

Since ever infectious diseases have been a major hazard for the armed forces in operations. Nowadays our nations are facing the threat of terrorism, including bioterrorism. This threat is much more related to the potential disorganization of the society than to the lethal effects of the agents. Biological weapons are considered more like terror weapons than like mass destruction weapons, hence the importance of preparing specific defence measures. The know-how acquired from the struggle against natural infectious diseases is a useful help to face the biological weapon threats and risks. Likewise, the defence attitude is based on three pillars: anticipating, managing, and restoring. This military as well as civilian defence attitude applies to six important functions: (1) alert, (2) detection, diagnosis and identification, (3) medical countermeasures (drugs, vaccines and sera), (4) medical care in hospital, (5) training and information, (6) research and development of dedicated technologies.     

The Evolution of Positive Allometry of Weaponry in Horned Lizards (Phrynosoma)

The allometry of weapons and other conspicuous structures has long fascinated biologists. Recently, a debate has arisen about the roles of sexual and natural selection in driving the allometry of structures, with some authors suggesting that positive allometry is exclusively the result of sexual selection. Although some studies, often focusing on weapons, support this hypothesis, others have shown that many naturally selected structures also exhibit positive allometry. We study the allometry of the cranial horns in 14 species of horned lizards (Phrynosoma). These horns are purely defensive weapons and so are under natural, as opposed to sexual, selection. In almost all cases, the length of these horns is positively allometric through ontogeny (intraspecifically) and through evolution (interspecifically). Our findings demonstrate that positive allometry can be the product of natural selection, suggesting that the function of structures may dictate allometry and not the type of selection. For example, it is possible that weapons tend to be positively allometric. Our findings also suggest that longer horns may arise through the prolongation of horn growth, and that the horns that are most effective at defense from predators attacking a lizard from above are integrated with one another. Finally, we find that species with unusual horn morphologies have diverged from other species in their horn allometries, indicating that the evolution of morphological diversity can be mediated through the evolution of allometry.     

Mating success and potential male-worker conflict in a male-dimorphic ant

Background  

Males of many species adjust their reproductive tactics with regard to their condition and status. For example, large males may develop weapons and fight for access to females, whereas small or undernourished males do not express costly weapons or ornaments and sneak copulations. Different condition-dependent reproductive tactics may be associated with unequal average fitness, but the tactic chosen by a given male under given circumstances is thought to result in the highest possible fitness return.