Showing papers in "Biological Journal of The Linnean Society in 1988"

Insect diversity: facts, fiction and speculation*

Abstract: Biologists are still trying to grasp the global dimensions of the phylum Arthropoda and its major class the Insecta, in spite of the fact that over a million species of arthropods have been described. The canopy of rain forest trees is believed by many to hold the key to the immense diversity of insects. In recent years the use of knock-down insecticides to sample insects from rain forest canopy has revealed information on the canopy’s arthropod inhabitants and community structure. The sampling techniques involved are outlined and data reviewed on taxonomic and guild structure, species abundance, body size and biomass ofinsects, and the faunal similarity of trees. Calculations by Erwin (1982), based on knock-down insecticide studies ofthe beetle fauna ofone species ofcentral American tree, suggest there may be 30 million species of tropical forest arthropods. Reanalysis of these calculations, using additional data, produces a range of possible estimates from about 10 to 80 million. The unknown range of plant host-specificities of tropical insects is the main weakness of this method of calculation. Assessment of the faunal importance of the canopy in relation to that of other rain forest biotopes requires comparative quantitative studies. The preliminary results of one such simple study suggest that over 42 million arthropods may be found in a hectare of Seram rain forest (at the time of study), and that 70% occur in the soil and leaf litter and 14% in the canopy. They also suggest that Collembola and Acarina are the dominant groups in this hectare, and that there are as many ants as all the other insects (excluding Collembola),

Variation in mutualisms : the spatio temporal mosaic of a pollinator assemblage

Abstract: Variation in time (annual and seasonal) and space (between- and within-population) is examined for the pollinator assemblage of Lavandula latifolia (Labiatae), an insect-pollinated, summerflowering, evergreen shrub of Mediterranean woodlands in southeastern Spain. Lauandula latiflia is pollinated there by nearly 85 species of comparatively long-tongued Hymenoptera, Diptera and Lepidoptera. The diversity, composition and abundance of pollinators varied markedly between years. Lepidopteran taxa prevailed numerically in one year, hymenopterans in four years, and both groups had similar abundance in the remaining year. The vast majority of pollinators exhibited significant annual differences in average abundance. Only 35.7% of taxa were recorded in all of the six study years. The pollinator assemblage had marked seasonal dynamics. Important changes in species richness, abundance and composition took place over the flowering season of L. latiflia (nearly 3 months). Only 21.7y0 of insect taxa occurred throughout the flowering season. Lavandula latiflia populations at different distances from water courses differed broadly in the abundance and composition of pollinators. Hymenopterans dominated numerically at the water-distant sites, whereas lepidopterans prevailed in populations growing in the vicinity of streams. Only 40.7% of taxa were recorded at all of the four study populations. Within the same plant population, horizontal distances of the order of 25 m resulted in significant changes in pollinator composition. Variations at the four scales considered combine to produce, from the viewpoint of L. latifolia, a spatio-temporal mosaic of pollinators. As these differ broadly in frequency of pollen transfer, size of pollen loads deposited, and between-flower flight distance patterns, spatio-temporal variation will predictably result in inconsistent or contradictory selective pressures on the plant, thus hindering specialization in relation to particular pollinators.

Food selection by two South-east Asian colobine monkeys (Presbytis rubicunda and Presbytis melalophos) in relation to plant chemistry.

Abstract: The diets of the banded leaf monkey (Presbytis melalophos) at Kuala Lompat in the Krau Game Reserve of West Malaysia and the red leaf monkey (Presbytis rubicunda) in Sepilok Virgin Jungle Reserve, Sabah, East Malaysia have been examined in relation to plant chemistry. Both monkeys spent about half their time eating foliage, and about half their time eating fruits and seeds. They both favoured leaves with high digestibility (due to relatively low levels of fibre) and high levels of protein, a combination found predominantly in young leaves and some flowers. The monkeys appeared to favour seeds and fruits with high concentrations of storage carbohydrates or fats, but not those rich in simple sugars. Selection of seeds and fruits was not correlated with protein content. An analysis of the fibre and protein contents of foods showed that, on an annual basis, the two monkeys achieved a comparable intake for both items. However, these diets were obtained in radically different ways. Presbytis melalophos was able to eat foliage from many of the common tree species in its home range, whereas P. rubicunda relied on rare trees and lianas. This difference is attributed to the very high density of Dipterocarpaceae at Sepilok, a tree family that provides little food for colobines. The rarity of P. rubicunda's food plants at Sepilok is considered to be the main reason for the greater home range size and lower population density in comparison to P. melalophos. Finally, the biochemical profiles of the young leaf diet of these two monkeys were compared with previously published information on two African and one south Indian colobines. In many respects the intake of supposed critical components, protein and fibre, showed marked similarities between different animals, considering contrasts in their habitats.

Testes size, ejaculate quality and sperm competition in birds

Abstract: The relationship between testes size, ejaculate quality (volume, sperm concentration, number of sperm per ejaculate) and sperm competition in birds was analysed using data collected in artificial insemination studies. I hypothesized that ejaculate quality, because of natural selection, should be superior in species with intense sperm competition compared with other species. In regression analyses, testes weight increased with body weight, with an exponent less than one, and ejaculate volume increased with testes weight with an exponent not significantly different from one, whereas sperm number per ejaculate increased with testes weight with an exponent larger than one. Species with relatively large testes also produced ejaculates with a high sperm concentration. Monogamous species with a relatively low intensity of sperm competition copulate rarely, but deliver ejaculates with a relatively large number of sperm. Monogamous species with a high intensity of sperm competition copulate frequently, but produce ejaculates with a relatively small number of sperm. Males of polygynous species, which also experience intense sperm competition, copulate rarely with specific females, but produce many ejaculates per male each with a relatively small number of sperm.

Evolutionary patterns of host utilization by ichneumonoid parasitoids (Hymenoptera: Ichneumonidae and Braconidae)*

Abstract: The four major biological strategies of ichneumonoid parasitoids, koinobiont and idiobiont, ecto-and endoparasitism, are discussed and the evolutionary radiations of the two families Ichneumonidae and Braconidae compared in an attempt to relate differences in patterns of host utilization to differences in evolutionary history. The most primitive members of both families are idiobiont ectoparasitoids of hosts concealed in plant tissue. Idiobiont ectoparasitic braconids are all still primarily associated with such hosts, but idiobiont ectoparasitic ichneumonids have radiated to attack hosts in other situations, such as in aculeate nests or in cocoons. A shift in emphasis between the behavioural steps, host habitat location and host location, is envisaged as being important in such evolutionary change. Idiobiont endoparasitism is postulated as having arisen amongst ectoparasitoids attacking cocooned hosts, as an adaptation that allows them to exploit pupae and puparia in relatively exposed positions; it is a fairly common strategy in the Ichneumonidae, but virtually unknown in the Braconidae. Koinobiosis is perceived as having evolved in association with hosts which feed in a relatively weakly concealed position, but pupate in a more secluded and safe location. The strategy is advantageous as it allows a parasitoid to oviposit on an easily discoverable host, but to use the host's pupation concealment to complete its own development. The evolution of koinobiosis has allowed parasitoids to exploit hosts that feed in exposed positions, and to attack hosts at a younger and numerically more common stage in the host's life cycle. Koinobiont ectoparasitism is envisaged, in some braconid and ichneumonid groups, to occupy an evolutionary transitional position between idiobiosis and endoparasitic koinobiosis; only in the Ichneumonidae have large radiations of koinobiont ectoparasitoids occurred. Endoparasitic koinobiosis is hypothesized as having arisen in the Braconidae in association with lepidopterous/coleopterous hosts, whilst in the major lineage of endoparasitic koinobiont ichneumonids, this habit is hypothesized as having arisen in association with symphytan hosts. The great majority of braconids are koinobiont endoparasitoids, but only about 50% of the Ichneumonidae have this habit. Very few koinobiont braconids develop as endoparasitoids of hymenopterous hosts, although many endoparasitic ichneumonids attack Hymenoptera. However, lineages of the Braconidae have radiated to exploit adult insects and exopterygote nymphs; ichneumonids do not utilize such hosts.

Hostplants and classification: a review of nymphalid butterflies

Abstract: In reviewing the hostplant associations of nymphalid butterflies, particular emphasis is placed on the intractable problem of nymphalid classification. Although offering few certain conclusions, if used in conjunction with more formal morphological characters, the data presented should contribute toward a resolution of the inter-relationships of the many widely recognized groupings within the Nymphalidae, several of which seem to be broadly characterized by typical host families. As a direct result of this analysis, the presumed association between larval hostplants and unpalatability is re-appraised.

Patterns of variation in tail ornament size in birds

Abstract: In recent years several different kinds of sexual selection models have been developed, and tail ornaments in birds have frequently been used as an example of a sexually selected character where the models might apply. However, very little is known about intra- and interpopulation variation in ornament size. We have studied the elongated tail ornaments in four species of whydahs Vidua, the forktailed flycatcher Tyrannus savana and the Asian paradise flycatcher Terpsiphone paradisi. Ornaments were relatively longer in males with the longest tarsi (‘heterogony’ with positive allometry). Also, tail lengths were remarkably variable within each geographical area, the coefficient of variation (average = 11%) being three times as high as for body size characters. Models, with female preference of ornaments bearing no relation to male viability, usually generate lines of neutral equilibria. Thus, they predict extraordinary variation in ornaments between populations. However, elongated tail ornaments did not show higher geographical variation than the body size characters, suggesting that there is no line of equilibria for these ornaments.

Frugivorous and animalivorous bats (Microchiroptera): dental and cranial adaptations

Abstract: The most derived fruit-eating bats have small canines, wide palates and molars with a distinctive labial rim. Paracone and metacone have moved from a dilambdodont position in the middle of the tooth to the labial side of the tooth where they form the labial cutting edge. Along with the well-developed and close fi tting labial cutting edges of the premolars and canines, this cutting edge skirts nearly the entire perimeter of the palate. The labial rim of the lower teeth fi t inside the labial rim of the upper teeth like two cookie cutters nesting one inside the other. Frugivores have a greater allocation of tooth area at the anterior end of the toothrow, while animalivorous species have more at the posterior end of the toothrow. The area occupied by canines of predators of struggling prey is greater than that for bats that eat non-struggling prey like fruit. In addition, frugivores have wider palates than long while many carnivores have longer palates than wide. Omnivores appear to have a more equal allocation of space to more kinds of teeth, particularly the incisors and non-molariform premolars, on the toothrow than do frugivores or animalivores. The mechanical nature of different food items is discussed and the suggestion made that describing foods in terms of their texture may be more important in tooth design than whether they are fruit or insect or vertebrate.

A comparison of the floristics and leaf chemistry of the tree flora in two Malaysian rain forests and the influence of leaf chemistry on populations of colobine monkeys in the Old World

Abstract: A comparative floristic survey of lowland tropical rain forest at Kuala Lompat, Krau Game Reserve, West Malaysia and at Sepilok Virgin Jungle Reserve, Sabah, East Malaysia, revealed significant differences in the composition of canopy species, notably in the relative paucity of Dipterocarpaceae and abundance of Leguminosae at Kuala Lompat. A further comparison with data from other sites in Malesia indicated that Kuala Lompat was rather atypical, representing an extreme example of the relatively legume rich and dipterocarp poor forests of West Malaysia. Conversely, the predominance of dipterocarps and lack of legumes at Sepilok was similar to other sites in Borneo. Young and mature leaves were collected from a representative sample of the canopy flora of the two study sites; the levels of nitrogen, fibre, total phenolics and condensed tannins, and degree of digestibility by cellulase and pepsin, were measured. Levels of fibre, nitrogen and phenolics differed significantly between the two floras; foliage from Kuala Lompat being, on the whole, richer in nitrogen but lower in phenolics and fibre. As a consequence of the lower levels of fibre, and less certainly of phenolics, the Kuala Lompat foliage was comparatively more digestible. Possible reasons for the differences in foliar biochemistry between the two sites are discussed. Differences in rainfall and in soil quality are regarded as potential evolutionary causes. A further comparison of the chemical profiles with data for other forests in India and Africa indicate that Sepilok is a site in which trees invest heavily in quantitative defences such as fibre and phenolics whereas trees at Kuala Lompat place a lower emphasis on the production of these. Finally, attention is drawn to the use of foliar chemical profiles in predicting the carrying capacity for colobine monkeys at each of the five Old World forests where data are available. It was found that the ratio of nitrogen to fibre correlated well with the biomass of colobines at the five sites.

Ecological and evolutionary implications of diet in monitor lizards

Abstract: A survey of 35 species indicates that monitor lizards (Varanus) typically hunt over large areas, search in particular microhabitats, and feed frequently on a wide variety of prey, many of which are relatively small. There is ontogenetic, seasonal, and geographic variation in diet. With some exceptions, invertebrates are the predominant prey, but rare predation on vertebrates is often energetically significant. A few monitors specialize on prey types that occur as occasional items in the diet of species with more generalized diets; these include crabs, snails, orthopterans, lizards, and large mammals. For most species, prey specialization occurs via habitat selection and a variety of prey types and sizes are eaten, as expected for widely searching predators. Comparisons with other anguimorphans suggest that derived features of Varanus are associated with high body temperature and activity levels; specialized chemoreception; and rapid, skillful capture of hidden and/or potentially hard to catch prey. Occasional ingestion of moderately large prey is primitive for Varanoidca (Helodermatidae +Varanidae), accentuating a trend that is perhaps primitive for anguimorphan lizards. Reduction of very large prey prior to ingestion is a derived attribute within Varanus, seen infrequently in several larger species and commonly in V. komodoensis. This study illustrates the synthesis of comparative natural history in a phylogenetic context, a method that addresses the history of organismal change.

Behavioural variation in natural populations. V. Morphological correlates of locomotion in the garter snake (Thamnophis radix)

Abstract: A series of morphologieal and locomotor performance variables was measured in a population of newborn garter snakes to determine whether performance capacity has a significant morphological basis in these animals. Morphological traits measured were body length and mass, number of body and tail vertebrae and numbers of vertebral abnormalities. Locomotor performances included burst and mid-distance speed and distance and time crawled before anti-predator displays were assumed. All performance variables were repeatable in daily replicate trials (P < 0.001). Individual burst speed, mid-distance speed, and distance crawled were significantly correlated pairwise (P < 0.01). Most morphological and performance variables had a significant mass dependence (static allometry), although the effects were rather weak (r2 < 0.1, except for body length): larger animals performed better and had fewer abnormalities. There were significant associations between some morphological traits and locomotor performance. Morphological factors accounted for 19% of the variation in mid-distance speed and 14% of the variation in antipredator behavior by multiple regression analysis. Canonical correlation of all performance and morphological variables simultaneously accounted for 24% of the observed variation in performance. Numbers of body and tail vertebrae (assayed by scale counts) had an interactive effect on speed of locomotion.

Evolutionary rates: effects of stress upon recombination

Abstract: There is increasing evidence that ecological variables involving stress are important in determining evolutionary rates. This paper incorporates recombination into this scenario. In Drosophila melanogaster, recombination increases at developmental temperatures above and below normal culture temperatures, giving a U-shaped curve which is most pronounced in centromeric regions; however, at near lethal temperature extremes there is some evidence for a fall in recombination. More limited data from other organisms are generally consistent with this conclusion. Nutritional stress in the form of starvation increases recombination in D. melanogaster, and behavioural stress has been found to increase recombination in male mice. In natural populations recombination is under complex genetic control analogous to other quantitative traits. In D. melanogaster in a novel environment, there is evidence that additive genetic variability for recombination is higher than in a standard laboratory environment. During selection in populations exposed to extreme stress increased recombination may occur; this implies that in marginal (stressful) habitats, variability generated by recombination may increase. In D. melanogaster, structural heterozygosity due to inversions in one part of the genome tends to increase recombination in the remainder of the genome in a qualitatively similar manner to, and cumulative with, direct environmental effects especially temperature. Substantial recombination should be inducible under combinations of karyotypes and environments deviating from existing circumstances, especially if the suggestion that effects are often synergistic due to a dependence upon available energy levels can be confirmed.

Myriapod metamerism and arthropod segmentation

Abstract: Outstanding progress in understanding segmentation of tracheate arthropods (Atelocerata), i.e. Chilopoda, Diplopoda, Pauropoda, Symphyla and Insecta, has been gained through experimental studies carried out on a single, very derivative organism, i.e. Drosophila. We stress the need for a broader comparative approach. We have studied the segmental structure of the trunk in geophilomorph centipedes, where we can identify morphogenetic units of two, four, eight or 16 segments. Accordingly, we sketch an improved model for arthropod segmentation, with the following initial steps: (a) biochemical marking of a very few repetitive units (eosegments); (b) iterative duplications of this first periodicity, until the embryo acquires an array of biochemical markings matching the whole number of segments of the future larva or juvenile specimen; (c) transpatterning, stabilization and interpretation of this ‘segmental’ arrangement; (d) possible repatterning, to give a final repetitive pattern we define as metasegmental. Finally, we express some doubt about the homology between annelid and arthropod segmentation.

Evolution of life histories and yields in experimental populations of Daphnia magna

Abstract: An experiment on life-history evolution is described in which replicated populations of the Cladoceran Daphnia magna, made up of a standard mixture of clones, were subjected to two contrasting culling regimes, involving removal of small or large individuals. After approximately 150 days of culling, analysis of the life histories showed that genetic differences had emerged between culling regimes. Clones selected by culling small sized individuals grew rapidly through small size classes, whereas those selected by culling large sizes grew slowly through small size classes, with the result that the age at which they became vulnerable to harvesting was delayed. In addition, there was some redistribution of reproduction towards size classes that were not culled. This evolution is consistent with a major decline observed in the yield from populations in which large individuals were culled. We argue that changes of this general kind are likely to take place in exploited populations and that serious consideration should be given to evolutionary aspects of the management of such populations.

Genetic parallelism of protein polymorphism in nature: ecological test of the neutral theory of molecular evolution

Abstract: We have conducted an ecological test of protein polymorphism in 13 unrelated genera of plants, invertebrates and vertebrates, involving 21 species, 142 populations and 5474 individuals. Each was tested, on average, for 27 enzymatic gene loci. These species varied in population size and structure, life histories and biogeographical origins, but they largely share a geographically short (260 km) and ecologically stressful gradient of increasing aridity in Israel, both eastward and (mainly) southward. We found genetic parallelism across most taxa, and most loci. Observed average heterozygosity, H, and gene diversity, He, were positively and overall significantly correlated with rainfall variation. This result corroborates the environmental theory of genetic diversity, primarily the niche-width variation hypothesis in both space and time. Our results are inconsistent with the neutral theory of molecular evolution and suggest that natural selection appears to be an important differentiating evolutionary force at the protein level.

The evolution of vertebrate flight

Abstract: Flight–defined as the ability to produce useful aerodynamic forces by flapping the wings–is one of the most striking adaptations in vertebrates. Its origin has been surrounded by considerable controversy, due in part to terminological inconsistencies, in part to phylogenetic uncertainty over the sister groups and relationships of birds, bats and pterosaurs, and in part to disagreement over the interpretation of the available fossil evidence and over the relative importance of morphological, mechanical and ecological specializations. Study of the correlation between functional morphology and mechanics in contemporary birds and bats, and in particular of the aerodynamics of flapping wings, clarifies the mechanical changes needed in the course of the evolution of flight. This strongly favours a gliding origin of tetrapod flight, and on mechanical and ecological grounds the alternative cursorial and fluttering hypotheses (neither of which is at present well-defined) may be discounted. The argument is particularly strong in bats, but weaker in birds owing to apparent inconsistencies with the fossil evidence. However, study of the fossils of the Jurassic theropod dinosaur Archaeopteryx, the sister-group of the stem-group proto-birds, supports this view. Its morphology indicates adaptation for flapping flight at the moderately high speeds which would be associated with gliding, but not for the slow speeds which would be required for incipient flight in a running cursor, where the wingbeat is aerodynamically and kinematically considerably more complex. Slow flight in birds and bats is a more derived condition, and vertebrate flapping flight apparently evolved through a gliding stage.

Sexual and asexual Taraxacum species

Abstract: The genus Taraxacum is a widely dispersed, ecologically variable taxon of some 2000 sexual and apomictic (agarnospermous) species. Data from numerous studies are used to examine the influences sexuality and apomixis have had on its evolution, geographical distribution and ecological diversification. A new explanation is given of the geographical distribution of sexual and apomictic forms, and the role of polyploidy in buffering apomicts against the effects of an accumulation of deleterious mutations is examined.

Microgeographical variation in shell size of the land snail Chondrina clienta

Abstract: Microgeographical variation in shell morphology of the rock-dwelling land snail Chondrina clienta, collected from 30 sites within an area of 0.5 km2 on the island of Oland, Sweden, was examined in relation to its own population density and that of a potential competitor (the land snail Balea perversa) and to environmental factors. Dispersal of marked individuals averaged 83 cm per year within a stone pile and 291 cm in an area of exposed bedrock. Local population density of active C. clienta ranged from 5 to 794 individuals per m2. Shell characters were highly intercorrelated, both within and between populations. Principal component analysis revealed that most of the interpopulational variation could be expressed by the single character of shell height, which ranged from 5.54 to 6.94 mm. In all populations, snails of a given size had the same whorl number. Shell size was not influenced by habitat type (exposed rock surface, stone pile or stone wall) or proportion of calcareous stones within habitat. It was, however, negatively correlated with local population density, indicating intraspecific competition, and positively correlated with the degree of plant cover within the habitat. Analysis of variance revealed additionally a density effect of B. perversa on shell size in C. clienta, probably as a result of interspecific competition. Breeding experiments using C. clienta from different sites and carried out under unifrom conditions caused most of the phenotypic variation to disappear, demonstrating the high phenotypic plasticity of the species.

Ant‐mimicry in some Brazilian salticid and clubionid spiders (Araneae: Salticidae, Glubionidae)*

Abstract: This paper describes the morphological and behavioural adaptations responsible for ant-like appearance in eight species (genera zunigu, Synemosyna, Sphecotypus, and Myrmecium) of salticid and clubionid spiders studied in Amazonian and SE Brazil. All ant-mimicking spiders have body and legs thin, and the shiny integument typical of their models. Light horizontal hair bands and constrictions of the cephalothorax and abdomen simulate, respectively, the head-thorax joint and segmented gaster of ants. The petiole and postpetiole of the ants are usually mimicked by a lengthened pedicel, together with a narrowing of the posterior cephalothorax and/or anterior abdomen. The prominent pedipalps of the spiders often simulate ant mandibles, but they may also be strikingly similar to an ant’s head. All ant-mimicking spiders walked in a zig-zag ant-like pattern, and frequently raised and moved about the first pair of legs as ‘antennae’. The mimics were found in the same microhabitats (foliage or ground) as their models, and displayed strong avoidance reactions toward the latter both in the field and in captivity. The inoffensive characteristics of the mimics and the noxious traits of their models (strong mandibles, potent sting, hard integument, venomous secretions) strongly suggest that the spiders are Batesian ant-mimics. The detailed structural and behavioural adaptations enhancing ant-mimicry provide strong circumstantial evidence that the selective agents involved must have good visual acuity, and are probably small insectivorous vertebrates (e.g. birds, lizards and toads) or arthropods (e.g. wasps and spiders) which avoid ants.

A model for the evolution and control of generative apomixis

Abstract: A model is presented for the evolution and control of generative apomixis—a collective term for apomixis in animals and diplosporous apomixis in flowering plants. Its development takes into account data obtained from studies of apomictic-like processes in sexual organisms and in non-apomictic parthenogens, as well as data obtained from studies of generative apomicts. This approach provides insights into the evolution and control of generative apomixis that cannot be obtained from studies of generative apomicts alone. It is argued that the control of the avoidance of meiotic reduction during egg production in generative apomicts resides at a single locus, the identity of which can vary between lineages. This variation accounts for the observed variation between taxa in the pattern of avoidance of meiotic reduction. The affected locus contains a wild-type allele that codes for meiotic reduction and excess copies of a mutant allele that codes for its avoidance. The dominance relationship between these is determined by their ratio and by the environment. Environmental differences between female generative cells and somatic cells are such that the phenotypic expression of the mutant allele is favoured in the former, while that of the wild-type allele is favoured in the latter. This is important, for the locus is also involved in the control of mitosis which would be disrupted by the expression of the mutant allele in somatic cells. The requirement to maintain a viable pattern of growth and development explains why the wild-type allele is retained by generative apomicts, and this in turn explains why the ability to produce meiotically reduced eggs is retained by facultative forms and why it appears to be suppressed in, rather than absent from, obligate forms. The requirement for excess copies of the mutant allele in generative cells explains why generative apomicts are typically polyploid, as this condition provides a simple and effective means of generating the correct balance of mutant and wild-type alleles. Environmental effects can also lead to the dominance relationship between wild-type and mutant alleles varying between generative cells. In plants, this can lead to the apomixis gene being expressed, and thus to meiotic reduction being avoided, in only some ovules. Meiotically reduced, as well as meiotically unreduced, eggs are produced when this occurs. If compatible and viable pollen is available the meiotically reduced eggs may be fertilized, resulting in these organisms reproducing as facultative apomicts. It is argued that the control and evolution of parthenogenesis in generative apomicts varies between taxa. In some, the parthenogenetic initiation of embryos may result from the acquisition of a parthenogenesis gene or genes; but there is no reason to believe that this is either a general or a common requirement. Indeed, in some it may be an ancestral trait, these apomicts differing from their sexual ancestors in the ability to mature, rather than in the ability to initiate, embryos from unfertilized eggs; or it may result from physiological or developmental changes induced, for example, by polyploidization, hybridization, or the avoidance of meiotic reduction. In some plants it may be induced by pollination (without fertilization) or by the activity of a developing endosperm. Although it is argued that most generatively apomictic lineages may have acquired this form of reproduction relatively easily, by the acquisition of a mutation at a single locus, it is argued that newly initiated lineages may often be reproductively inefficient. These will begin to accumulate mutations that improve the efficiency of apomictic reproduction. Thus several loci may be involved in the control of generative apomixis in established lineages, even though only a single locus was involved in its initiation in these lineages. Care must be taken to distinguish between these initiator and modifier genes when considering the evolution of generative apomixis. Finally, it is argued that although generatively apomictic lineages have easily acquired this form of reproduction, its evolution in some taxa may be so difficult, requiring the acquisition of mutations simultaneously at two or more loci, that these may never acquire it. Thus, evidence obtained from taxa that have successfully made the transition from sexual reproduction to generative apomixis that its evolution was straightforward should not be used as evidence that its evolution will always be relatively easily achieved. Its uneven taxonomic distribution indicates that it is much more easily evolved by some taxonomic groups than by others.

Homologies in the colour patterns of the genus Heliconius (Lepidoptera: Nymphalidae)

Abstract: The colour patterns of Heliconius butterflies are composed from a relatively simple set of pattern elements whose homologues are recognizable throughout the genus. Although Heliconius colour patterns look quite different from those of most nymphalids, these pattern elements are seen to derive from the generalized nymphalid groundplan. The differences arise primarily from the loss or positional shift of certain pattern elements, a high degree of fusion between individual pattern elements, and, in the forewing, asymmetries of the pattern elements relative to the wing-cell midline. The scheme of homologies we present is consistent with what is currently known about the comparative morphology and developmental physiology of colour pattern formation in Lepidoptera, and provides a framework for the interpretation of developmental, evolutionary and genetic studies in Heliconius.

Colony genetic structure and queen mating frequency in fire ants of the subgenus Solenopsis (Hymenoptera: Formicidae)

Abstract: Colony genetic structure was studied in natural populations of three fire ant taxa, Solenopsis richteri Forel, S. geminata (Fabr.), and hybrid S. invicta/richteri, using allozyme markers. All colonies studied exhibited arrays of female genotypes predicted under a model of monogyny (single functional queen) and monoandry (single insemination of queens). Males produced in the colonies appear to originate exclusively from the foundress queen, rather than from any virgin females present in the colonies. Thus these social insect colonies represent simple, albeit enormous, family groups. Single insemination and foundress parentage of males appear to be conserved reproductive traits in the subgenus Solenopsis, whereas another major determinant of colony genetic structure, the number of functional queens, is evolutionarily labile in this group.

The giant tadpole of Pseudis paradoxa

Abstract: Pseudis paradoxa has an extremely large tadpole with a long, deep tail- These features are also found in overwintering tadpoles of temperate species where low temperatures delay development and prolong prolactin exposure. Pseudis paradoxa does not occur in localities with marked seasonal temperature fluctuations. Low temperature cannot be implicated in the development of the tadpole. However, the parallels in shape between Pseudis tadpoles and those of temperate overwintering species suggest that Pseudis tadpoles may have a prolonged exposure to prolactin, higher levels of prolactin during development or an increased sensitivity to prolactin.

Variation in hostplant relations and courtship signals of weed‐associated populations of the brown planthopper, Nilaparvata lugens (Stål), from Australia and Asia: a test of the recognition species concept

Abstract: Populations of the morphological species, Nilaparvata lugens (Stal), were found to breed and feed on the grass, Leersia hexandra Schwartz, at six sites in Queensland, Australia. They differ from sympatric rice-feeding populations in characters of pulse repetition frequencies of male and female acoustic courtship signals. The two host-derived populations hybridize freely in the laboratory, but in mate choice experiments show very significant preferences for homogametic matings. No indication of field hybridization has been found, so that the two morphologically inseparable populations represent sympatric biological species in Australia. Populations from L. hexandra are also reported from four localities in Sri Lanka and one in Orissa, India. These resemble previously studied populations from the Philippines. They differ significantly in courtship call characters, both from sympatric rice-associated populations and from allopatric Leersia-associated populations from Australia. The geographical variation reported for acoustic signals is not consistent with Paterson's recognition concept of species, but may be interpreted in terms of theories of allopatric speciation involving sexual selection for mate recognition signals.

On the gynodioecious polymorphism in Saxifraga granulata L. (Saxifragaceae).

Abstract: Sexual and vegetative fitness components in hermaphrodite and female plants of the self-compatible, perennial herb Saxifraga granulata are compared using material derived from a gynodioecious population in northern England. Females produced only 57% as many seeds as hermaphrodites, but their ovule offspring were 1.28 times as fit as those of hermaphrodites, and females were more vegetatively vigorous. The advantages to females in ovule offspring quality and in vegetative reproduction counteract their disadvantages in pollen and seed production and therefore probably play a role in the maintenance of the gynodioecious polymorphism. Pollination ecology, resource reallocation and inbreeding depression all appear to contribute to the observed sex differences in fitness.

Mimicry and mutualism in carnivorous pitcher plants (Sarraceniaceae, Nepenthaceae, Cephalotaceae, Bromeliaceae)

Abstract: The carnivorous pitcher plants of the Sarraceniaceae, Nepenthaceae and Cephalotaceae seem to have developed attraction and rewarding systems which resemble those of many flowers. Contrary to a previous view, Batcsian mimicry does not appear to apply to pitcher traps because: (a) there is no evidence of insects visiting traps by mistake, (b) no model is known for any one of the traps, (c) the duration of the trap activity and the structure of the plant community are not compatible with a deceptive mimetic status, (d) nectar which is a real reward is provided to visiting insects by the pitcher plants. It is suggested that these plants perform mutual rather than deceptive relations with the insect communities in their habitats. Insects benefit from nectar which is provided by pitchers and which serves for their nutrition in habitats where sources of floral nectar might be either uncommon or absent. At the same time the insects pay the plants in a small portion of their community which is ‘sacrificed’ as prey and consumed by the plants, which grow in nutrient deficient soils.

Microgeographic variation in shell characters of Littorina saxatilis Olivi—a question mainly of size?

Abstract: The question of whether there are shape differences between populations of Littorina saxatilis living in different environments is examined by multivariate analyses of 13 morphological characters. Principal component analysis reveals that morphologic differences between populations from habitats with contrasting degrees of wave exposure are mainly due to a general size factor, including shell thickness. Utilizing the group structure among the snails, canonical variate analysis discloses that the main character excluding size that influences subpopulation differentiation is pointedness.

Temporal patterns of reproduction and outcrossing in weedy populations of Echium plantagineum.

Abstract: Single and multilocus estimates of outcrossing rates were made in three populations of Echium plantagineum. Despite spatial separation, variations in population size (though not density) and reproductive output, no statistically significant difference was detected in outcrossing rates between the populations. Similarly, only slight differences in outcrossing rates were detected within populations when estimates were based on seed collected from flowers open at different times in the flowering season. The earliest flowers tended to have a lower estimated rate of outcrossing. In all cases, multilocus outcrossing rates were high, ranging from 0.81 to 1.05. In all three populations flower production extended over a period of more than 2 months but the majority of seed was produced by flowers that opened during the first third of the (lowering season. This was largely caused by a high rate of flower production during the early part of the season and not changes in the number of seeds set per flower. The average number of seeds produced per flower varied both between individuals within populations, and between the different populations, but neither of these differences were significant. Controlled pollination of flowers with self or outcross pollen, applied either singly or together, failed to detect any differences in the likelihood that either type of pollen would give rise to fertile seeds.

A functional analysis of the shift from gill to lung breathing during the evolution of land crabs crustacea decapoda

Abstract: The evolution of air-breathing in land crabs is associated with a progressive shift in the primary site of respiratory gas exchange from the diffusion-limited gills used for water-breathing, via a simple ‘cutaneous’ lung surface to the perfusion-limited, invaginated lung described in the mountain crab, Pseudothelphusa garmani. The reduced diffusion limitation over the lungs facilitates oxygen transfer from air to the tissues at lower ventilation rates but is associated with accumulation of carbon dioxide. A potential respiratory acidosis is buffered by the respiratory pigment haemocyanin and by elevation of haemolymph bicarbonate levels. These changes parallel those described in vertebrates but air-breathing crustaceans maintain relatively low carbon dioxide levels in the haemolymph, either by retaining an aquatic route for its elimination over the reduced gills or by blowing it off across the lung. Maintenance of low carbon dioxide levels may be associated with a limited capacity to buffer against an acidosis due to low levels of circulating haemocyanin (i.e. crustaceans lack red blood cells). This may ultimately limit their survival in air as an acidosis will reduce oxygen transport due to a marked Bohr effect on haemocyanin. The primary role of an invaginated lung may be to reduce rates of water loss in air.