Review Article
Adaptive explanation in socio-ecology: lessons from the Equidae
- WAYNE L. LINKLATER
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- Published online by Cambridge University Press:
- 01 February 2000, pp. 1-20
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Socio-ecological explanations for intra- and interspecific variation in the social and spatial organization of animals predominate in the scientific literature. The socio-ecological model, developed first for the Bovidae and Cervidae, is commonly applied more widely to other groups including the Equidae. Intraspecific comparisons are particularly valuable because they allow the role of environment and demography on social and spatial organization to be understood while controlling for phylogeny or morphology which confound interspecific comparisons. Feral horse (Equus caballus Linnaeus 1758) populations with different demography inhabit a range of environments throughout the world. I use 56 reports to obtain 23 measures or characteristics of the behaviour and the social and spatial organization of 19 feral horse populations in which the environment, demography, management, research effort and sample size are also described. Comparison shows that different populations had remarkably similar social and spatial organization and that group sizes and composition, and home range sizes varied as much within as between populations. I assess the few exceptions to uniformity and conclude that they are due to the attributes of the studies themselves, particularly to poor definition of terms and inadequate empiricism, rather than to the environment or demography per se. Interspecific comparisons show that equid species adhere to their different social and spatial organizations despite similarities in their environments and even when species are sympatric. Furthermore, equid male territoriality has been ill-defined in previous studies, observations presented as evidence of territoriality are also found in non-territorial equids, and populations of supposedly territorial species demonstrate female defence polygyny. Thus, territoriality may not be a useful categorization in the Equidae. Moreover, although equid socio-ecologists have relied on the socio-ecological model derived from the extremely diverse Bovidae and Cervidae for explanations of variation in equine society, the homomorphic, but large and polygynous, and monogeneric Equidae do not support previous socio-ecological explanations for relationships between body size, mating system and sexual dimorphism in ungulates. Consequently, in spite of the efforts of numerous authors during the past two decades, functional explanations of apparent differences in feral horse and equid social and spatial organization and behaviour based on assumptions of their current utility in the environmental or demographic context remain unconvincing. Nevertheless, differences in social cohesion between species that are insensitive to intra- and interspecific variation in habitat and predation pressure warrant explanation. Thus, I propose alternative avenues of inquiry including testing for species-specific differences in inter-individual aggression and investigating the role of phylogenetic constraints in equine society. The Equidae are evidence of the relative importance of phylogeny and biological structure, and unimportance of the present-day environment, in animal behaviour and social and spatial organization.
Why do females mate multiply? A review of the genetic benefits
- MICHAEL D. JENNIONS, MARION PETRIE
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- Published online by Cambridge University Press:
- 01 February 2000, pp. 21-64
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The aim of this review is to consider the potential benefits that females may gain from mating more than once in a single reproductive cycle. The relationship between non-genetic and genetic benefits is briefly explored. We suggest that multiple mating for purely non-genetic benefits is unlikely as it invariably leads to the possibility of genetic benefits as well. We begin by briefly reviewing the main models for genetic benefits to mate choice, and the supporting evidence that choice can increase offspring performance and the sexual attractiveness of sons. We then explain how multiple mating can elevate offspring fitness by increasing the number of potential sires that compete, when this occurs in conjunction with mechanisms of paternity biasing that function in copula or post-copulation. We begin by identifying cases where females use pre-copulatory cues to identify mates prior to remating. In the simplest case, females remate because they identify a superior mate and ‘trade up’ genetically. The main evidence for this process comes from extra-pair copulation in birds. Second, we note other cases where pre-copulatory cues may be less reliable and females mate with several males to promote post-copulatory mechanisms that bias paternity. Although a distinction is drawn between sperm competition and cryptic female choice, we point out that the genetic benefits to polyandry in terms of producing more viable or sexually attractive offspring do not depend on the exact mechanism that leads to biased paternity. Post-copulatory mechanisms of paternity biasing may: (1) reduce genetic incompatibility between male and female genetic contributions to offspring; (2) increase offspring viability if there is a positive correlation between traits favoured post-copulation and those that improve performance under natural selection; (3) increase the ability of sons to gain paternity when they mate with polyandrous females. A third possibility is that genetic diversity among offspring is directly favoured. This can be due to bet-hedging (due to mate assessment errors or temporal fluctuations in the environment), beneficial interactions between less related siblings or the opportunity to preferentially fertilise eggs with sperm of a specific genotype drawn from a range of stored sperm depending on prevailing environmental conditions. We use case studies from the social insects to provide some concrete examples of the role of genetic diversity among progeny in elevating fitness. We conclude that post-copulatory mechanisms provide a more reliable way of selecting a genetically compatible mate than pre-copulatory mate choice. Some of the best evidence for cryptic female choice by sperm selection is due to selection of more compatible sperm. Two future areas of research seem likely to be profitable. First, more experimental evidence is needed demonstrating that multiple mating increases offspring fitness via genetic gains. Second, the role of multiple mating in promoting assortative fertilization and increasing reproductive isolation between populations may help us to understand sympatric speciation.
Plant invasions – the role of mutualisms
- DAVID M. RICHARDSON, NICKY ALLSOPP, CARLA M. D'ANTONIO, SUZANNE J. MILTON, MARCEL REJMÁNEK
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- Published online by Cambridge University Press:
- 01 February 2000, pp. 65-93
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Many introduced plant species rely on mutualisms in their new habitats to overcome barriers to establishment and to become naturalized and, in some cases, invasive. Mutualisms involving animal-mediated pollination and seed dispersal, and symbioses between plant roots and microbiota often facilitate invasions. The spread of many alien plants, particularly woody ones, depends on pollinator mutualisms. Most alien plants are well served by generalist pollinators (insects and birds), and pollinator limitation does not appear to be a major barrier for the spread of introduced plants (special conditions relating to Ficus and orchids are described). Seeds of many of the most notorious plant invaders are dispersed by animals, mainly birds and mammals. Our review supports the view that tightly coevolved, plant-vertebrate seed dispersal systems are extremely rare. Vertebrate-dispersed plants are generally not limited reproductively by the lack of dispersers. Most mycorrhizal plants form associations with arbuscular mycorrhizal fungi which, because of their low specificity, do not seem to play a major role in facilitating or hindering plant invasions (except possibly on remote islands such as the Galapagos which are poor in arbuscular mycorrhizal fungi). The lack of symbionts has, however, been a major barrier for many ectomycorrhizal plants, notably for Pinus spp. in parts of the southern hemisphere. The roles of nitrogen-fixing associations between legumes and rhizobia and between actinorhizal plants and Frankia spp. in promoting or hindering invasions have been virtually ignored in the invasions literature. Symbionts required to induce nitrogen fixation in many plants are extremely widespread, but intentional introductions of symbionts have altered the invasibility of many, if not most, systems. Some of the world's worst invasive alien species only invaded after the introduction of symbionts. Mutualisms in the new environment sometimes re-unite the same species that form partnerships in the native range of the plant. Very often, however, different species are involved, emphasizing the diffuse nature of many (most) mutualisms. Mutualisms in new habitats usually duplicate functions or strategies that exist in the natural range of the plant. Occasionally, mutualisms forge totally novel combinations, with profound implications for the behaviour of the introduced plant in the new environment (examples are seed dispersal mutualisms involving wind-dispersed pines and cockatoos in Australia; and mycorrhizal associations involving plant roots and fungi). Many ecosystems are becoming more susceptible to invasion by introduced plants because: (a) they contain an increasing array of potential mutualistic partners (e.g. generalist frugivores and pollinators, mycorrhizal fungi with wide host ranges, rhizobia strains with infectivity across genera); and (b) conditions conducive for the establishment of various alien/alien synergisms are becoming more abundant. Incorporating perspectives on mutualisms in screening protocols will improve (but not perfect) our ability to predict whether a given plant species could invade a particular habitat.
Toxic action/toxicity
- D. E. HATHWAY
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- Published online by Cambridge University Press:
- 01 February 2000, pp. 95-127
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Some six or so physiological systems, essential to normal mammalian life, are involved in poisoning; an intoxication that causes severe injury to any one of them could be life threatening. Reversible chemical reactions showing Scatchard-type binding are exemplified by CO, CN− and cyclodiene neurotoxin insecticide intoxications, and by antigen–antibody complex formation. Haemoglobin (Hb) molecular biology accounts for the allosteric co-operativity and other characteristics of CO poisoning, CN− acts as a powerful cytochrome oxidase inhibitor, and antigen binding in a deep antibody cleft between two domains equipped with epitopes for antigen-binding groups explains hapten-specific immune reactions. Covalent chemical reactions with second-order (SN2) kinetics characterize Hg and Cd poisonings, the reactions of organophosphates and phosphonates with acetylcholinesterase and neurotoxic esterase and the reaction sequence whereby Paraquat accepts electrons and generates superoxide under aerobic conditions. Indirect carcinogens require cytochrome P450 activation to form DNA adducts in target-organ DNA and cause cancer, but a battery of detoxifying enzymes clustered with the P450 system must be overcome. Thus, S-metabolism competes ineffectively with target DNA for reactive vinyl chloride (VC) metabolites, epoxide hydrolase is important to the metabolism and carcinogenicity of aflatoxins and polycyclic aromatic hydrocarbons (benzo[a]pyrene, etc.), and the non-toxic 2-naphthylhydroxylamine N-glucuronide acts as a transport form in 2-naphthylamine bladder cancer. VC liver-cancer pathogenesis is explicable in terms of the presence of the glutathione S-transferase detoxifying system in hepatocytes and its absence from the fibroblastic elements, and of the VC concentrations reaching the liver by different administrative routes. In VC carcinogenicity, chemical reactions give imidazo-cyclization products with nucleoside residues of target DNA, and in benzene leukaemia, Z,Z-muconaldehyde forms cyclic products containing a pyrrole residue linked to purine. Increased HbCO concentrations reduce the O2-carrying capacity of the blood, and the changed shape of the O2-Hb dissociation curve parallels disturbance in O2 unloading. CN− acts on electron transport and paralyses respiration. In telodrin poisoning, preconvulsive glutamine formation abstracts tricarboxylic acid intermediates incommensurately with normal cerebral respiration. Antigen–antibody complexing depletes the antibody titre, available against infection. At high doses of Cd, Cd-thionein filtered through the kidneys is reabsorbed and tubular lesions produced. Some organophosphate insecticides promote irreversible acetylcholinesterase phosphorylation and blockade nerve function, and others react with neurotoxic esterase to cause delayed neuropathy. The evidence for Paraquat pulmonary poisoning suggests a radical mechanism involving three interrelated cyclic reaction stages. The action of N- and O6 (O substituent in 6-position of the purine) demethylases explains deletion mechanisms for DNA-alkyl adducts. DNA-directed synthesis in the presence of ultimate carcinogens provides for an estimation of misincorporations, which implicate the same transversions as those found by direct mutagenicity testing. Chemical carcinogens recognize tissue-sensitive cells and modify their heritable genetic complement. Oncoproteins encoded by activated oncogenes signal the transformation of normal cells into cancer cells. The importance of the H-ras oncogene and p53 tumour-suppressor gene is stressed. Antidotal action is analysed; for example, parenteral glutamine administration to telodrin-intoxicated rats restores the depleted cerebral glutamate level and prevents seizures. Glutamate acts as anticonvulsant in petit mal epilepsy. In general, therefore, the reaction of the toxicant-related substance with the relevant target-tissue macromolecule accounts for the biochemical/biological events at a cellular level and also the symptoms in the living mammal. This mechanism is analogous to mechanisms for diseases such as arthritis and Parkinsonism.
Flight adaptations in Palaeozoic Palaeoptera (Insecta)
- ROBIN J. WOOTTON, JARMILA KUKALOVÁ-PECK
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- Published online by Cambridge University Press:
- 01 February 2000, pp. 129-167
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The use of available morphological characters in the interpretation of the flight of insects known only as fossils is reviewed, and the principles are then applied to elucidating the flight performance and techniques of Palaeozoic palaeopterous insects. Wing-loadings and pterothorax mass/total mass ratios are estimated and aspect ratios and shape-descriptors are derived for a selection of species, and the functional significance of wing characters discussed. Carboniferous and Permian ephemeropteroids (‘mayflies’) show major differences from modern forms in morphology and presumed flight ability, whereas Palaeozoic odonatoids (‘dragonflies’) show early adaptation to aerial predation on a wide size-range of prey, closely paralleling modern dragonflies and damselflies in shape and wing design but lacking some performance-related structural refinements. The extensive adaptive radiation in form and flight technique in the haustellate orders Palaeodictyoptera, Megasecoptera, Diaphanopterodea and Permothemistida is examined and discussed in the context of Palaeozoic ecology.