Genetic diversity, host-specificity and unusual phylogeography of a cryptic, host-associated species complex of gall-inducing scale insects

Abstract.  1. Australia has a unique and speciose gall-inducing scale insect fauna that is primarily associated with Myrtaceae. Much of the diversity is currently undescribed or uncharacterised.
2. This study concerns Apiomorpha munita (Hemiptera), a scale insect that induces characteristic four-horned galls on eucalypts of subgenus Symphyomyrtus and exhibits extraordinary karyotypic diversity (2n = 6 – 2n > 100). The three described subspecies of A. munita are each confined to hosts in different sections of Eucalyptus . Previous chromosomal data, however, cast doubt on the validity of the groupings, as two of the subspecies share multiple, different karyotypes (2n = 6, 20, 22, and 24).
3. Allozyme data were used to examine species delimitation, chromosome evolution, host associations and population structure in A. munita .
4. A cryptic-species radiation was revealed, with at least five taxa each restricted to a discrete set of host eucalypt species. This is consistent with host-associated speciation.
5. Karyotypic variation within A. munita partially fits the five distinct genetic groups, but there are additional chromosomal changes that are not accompanied by detected genetic differentiation.
6. The population structure of taxa within the A. munita species complex suggests that there are high levels of inbreeding, as would be expected for scale insects in which adult females are sessile. Some genotypes, however, are found over great distances (up to 1100 km). This is an unusual population structure because it combines low mobility and local differentiation with occasional long-distance dispersal, probably mediated by wind-dispersal of first-instar nymphs.     

The Evolution of Parasite Recognition Genes in the Innate Immune System: Purifying Selection on <Emphasis Type="Italic">Drosophila melanogaster</Emphasis> Peptidoglycan Recognition Proteins

Genes involved in the recognition of parasites by the acquired immune system are often subject to intense selection pressures. In some cases, selection to recognize a diverse range of parasites has resulted in high levels of polymorphism, while elsewhere the protein sequence has changed rapidly under directional selection. We tested whether parasite recognition genes in the innate immune system show similar patterns of evolution. We sequenced seven peptidoglycan recognition protein genes (PGRPs) from 12 lines of Drosophila melanogaster and one line of D. simulans and used a variety of tests to determine whether the observed mutations were selectively neutral. We were unable to detect either balancing or directional selection. This suggests that the molecular cues used by insects to detect parasites are highly conserved and probably under strong functional constraints which prevent their evolving to evade the host immune response. Therefore, interactions between these genes are unlikely to be the focus of host–parasite coevolution, at least in Drosophila. We also found evidence of gene conversion occurring between two genes, PGRP-SC1A and PGRP-SC1B.     

Correlated evolution of male and female morphologles in water striders

Sexually antagonistic coevolution may be an important force in the evolution of sexual dimorphism. We undertake a comparative study of correlated evolution of male and female morphologies in a clade of 15 water strider species in the genus Gerris (Heteroptera: Gerridae). Earlier studies have shown that superfluous matings impose costs on females, including increased energetic expenditure and predation risk, and females therefore resist males with premating struggles. Males of some species possess grasping structures and females of some species exhibit distinct antigrasping structures, which are used to further the interests of each sex during these premating struggles. We use this understanding, combined with coevolutionary theory, to derive a series of a priori predictions concerning both the types of traits in the two sexes that are expected to coevolve and the coevolutionary dynamics of these traits expected under sexually antagonistic coevolution. We then assess the actual pattern of correlated evolution in this clade with new morphometric methods combined with standard comparative techniques. The results were in agreement with the a priori predictions. The level of armament (different abdominal structures in the two sexes) was closely correlated between the sexes across species. Males are well adapted to grasping females in species in which females are well adapted to thwart harassing males and vice versa. Furthermore, our comparative analyses supports the prediction that correlated evolution of armament in the two sexes should be both rapid and bidirectional.     

Coevolution of color pattern and thermoregulatory behavior in polymorphic pygmy grasshoppers Tetrix undulata

Ectothermic organisms, such as insects and reptiles, rely on external heat sources to control body temperature and possess physiological and behavioral traits that are temperature dependent. It has therefore been hypothesised that differences in body temperature resulting from phenotypic properties, such as color pattern, may translate into selection against thermally inferior phenotypes. We tested for costs and benefits of pale versus dark coloration by comparing the behaviors (i.e., basking duration and bouts) of pygmy grasshopper (Tetrix undulata) individuals exposed to experimental situations imposing a trade-off between temperature regulation and feeding. We used pairs consisting of two full-siblings of the same sex that represented different (genetically coded) color morphs but had shared identical conditions from the time of fertilization. Our results revealed significant differences in behavioral thermoregulation between dark and pale individuals in females, but not in males. Pale females spent more time feeding than dark females, regardless of whether feeding was associated with a risk of either hypothermia or overheating. In contrast, only minor differences in behavior (if any) were evident between individuals that belonged to the same color morph but had been painted black or gray to increase and decrease their heating rates. This suggests that the behavioral differences between individuals belonging to different color morphs are genetically determined, rather than simply reflecting a response to different heating rates. To test for effects of acclimation on behaviors, we used pairs of individuals that had been reared from hatchlings to adults under controlled conditions in either low or high temperature. The thermal regime experienced during rearing had little effect on behaviors during the experiments reported above, but significantly influenced the body temperatures selected in a laboratory thermal gradient. In females (but not in males) preferred body temperature also varied among individuals born to mothers belonging to different color morphs, suggesting that a genetic correlation exists between color pattern and temperature preferences. Collectively, these findings, at least in females, are consistent with the hypothesis of multiple-trait coevolution and suggest that the different color morphs represent alternative evolutionary strategies.     

BUTTERFLIES AND PLANTS: A PHYLOGENETIC STUDY

A database on host plant records from 437 ingroup taxa has been used to test a number of hypotheses on the interaction between butterflies and their host plants using phylogenetic methods (simple character optimization, concentrated changes test, and independent contrasts test). The butterfly phylogeny was assembled from various sources and host plant clades were identified according to Chase et al.'s rbcL-based phylogeny. The ancestral host plant appears to be associated within a highly derived rosid clade, including the family Fabaceae. As fossil data suggest that this clade is older than the butterflies, they must have colonized already diversified plants. Previous studies also suggest that the patterns of association in most insect-plant interactions are more shaped by host shifts, through colonization and specialization, than by cospeciation. Consequently, we have focused explicitly on the mechanisms behind host shifts. Our results confirm, in the light of new phylogenetic evidence, the pattern reported by Ehrlich and Raven that related butterflies feed on related plants. We show that host shifts have generally been more common between closely related plants than between more distantly related plants. This finding, together with the possibility of a higher tendency of recolonizing ancestral hosts, helps to explain the apparent large-scale conservation in the patterns of association between insects and their host plants, patterns which at the same time are more flexible on a more detailed level. Plant growth form was an even more conservative aspect of the interaction between butterflies and their host plants than plant phylogeny. However, this is largely explained by a higher probability of colonizations and host shifts while feeding on trees than on other growth forms.     

非人灵长类肠道微生物研究进展与展望

肠道微生物组被誉为动物的“第二套基因组”,与动物的个体发育、营养获取、生理功能、免疫调节等重要活动密切相关。非人灵长类在生态位、社会结构、地理分布以及进化上与人类相近,开展其肠道微生物研究不仅有助于了解灵长类的生态、保护和进化,而且对深入了解肠道微生物在人类进化中所发挥的作用也具有重要的参考价值。本文总结了影响非人灵长类肠道微生物变化的因素,包括系统发育、觅食、栖息地破碎化、年龄和性别、圈养方式以及社群生活,并探讨了肠道微生物研究在非人灵长类生态、行为、保护以及适应性进化方面的应用。未来,非人灵长类肠道微生物研究将为灵长类生态、进化和人类健康的研究提供新的视角,为灵长类的保护提供新的理论基础和研究方法。     

Evolution of fruit characters and dispersal modes in the tropical family Rubiaceae

We investigated the evolution of fruit characters, animals versus abiotic dispersal modes, life forms and geographical distribution, in the large, mostly tropical, family Rubiaceae. As a basis for our analysis we used a phylogenetic tree derived from chloroplast DNA variation. Fleshy fruits have evolved independently at least 12 times in the family. Most of these originations appear to have occurred during Eocene to Oligocene, i.e. the radiation period for some animal taxa (bird families, mammal orders) comprising most extant dispersers of Rubiaceae fruits. Changes of dispersal modes may be of more recent origin in a few cases, e.g. evolution of drupes in some lineages, and shifts from drupes to nuts. The distribution of fruit characters suggested that in several lineages animal-dispersed fruits, such as berries and drupes have remained largely unaltered since the time of origination. This is in contrast to the occurrence of winged seeds in capsules, and pterophylls, i.e. enlarged calyx lobes promoting wind dispersal of fruits, which apparently have shifted more frequently during evolution, indicating a difference in 'phylogenetic plasticity' between modes of animal and wind dispersal.
Animal dispersal was over-represented among genera dominated by shrubs, whereas abiotic dispersal was most prevalent among herbaceous genera. Drupes were over-represented in groups with transoceanic distributions, and on islands, indicating dispersal over long distances, probably by birds. In contrast, no evidence was found to support the view that animal dispersal in general enhances long distance dispersal. We also analysed geographical patterns on the tribal level but these were too complex to yield any resolved area cladograms due to the occurrence of many widespread taxa and area redundancy.     

Host resistance and pathogen virulence across a plant hybrid zone

Hybridization between locally adapted plant populations has been postulated to have significant evolutionary consequences, and, in particular, may influence host-pathogen interactions with respect to resistance and virulence structure. This study investigated patterns of resistance and virulence in a hybrid zone between ”bog” and ”hill” ecotypes of the native Australian flax, Linum marginale, where the host is subject to attack by the rust pathogen, Melampsora lini. Analysis of the resistance structure of adjoining bog, hill and hybrid populations found that bog plants were generally susceptible to pathogen isolates taken from all these sites, but that hybrids exhibited resistance levels similar to the more resistant hill plants. Similarly, the virulence structure of rust isolates collected from the hybrid population was more similar to that of the hill isolates than the bog. Controlled crosses between bog and hill plants showed that crosses in one direction (bog females×hill males) were much more successful than the other. A multi-year reciprocal transplant study further indicated that bog plants had significantly higher survivorship than hill plants, regardless of site. It is suggested that likelihood of differential gene flow and survivorship for bog and hill plants may at least partially explain the maintenance of a relatively narrow hybrid zone. Received: 5 October 1999 / Accepted: 1 July 1999     

Differential Susceptibility of Two Melolontha Populations to Infections by the Fungus Beauveria brongniartii

The susceptibility of second-instar larvae of the European cockchafer, Melolontha melolontha L., originating from two different geographical locations was investigated with eight isolates of the insect pathogenic fungus Beauveria brongniartii (Sacc.) Petch. 77-94% of larvae from northern Italy (Auer, South Tyrolia) succumbed to mycosis with an average survival time (AST) of 22.9-40.1 days. Infections in larvae from south-western Switzerland (Bramois, Valais) were 28-72% and the ASTs varied between 34.7-56.4 days. Differences in susceptibility between the two host populations may be explained by the historical ages of the two populations and the presence of B. brongniartii resulting in a coevolution of tolerance between the host and pathogen. The Italian population is occasionally infected by B. brongniartii; in comparison, the Swiss population has existed for at least 50 years and regular infections by the pathogen are observed. Coevolution between B. brongniartii and M. melolontha from Switzerland may explain the apparent resistance of the host towards this pathogen in laboratory assays.     

The Robust Statistical Bases of the Coevolution Theory of Genetic Code Origin

A paper (Amirnovin R, J Mol Evol 44:473–476, 1997) seems to undermine the validity of the coevolution theory of genetic code origin by shedding doubt on the connection between the biosynthetic relationships between amino acids and the organization of the genetic code, at a time when the literature on the topic takes this for granted. However, as a few papers cite this paper as evidence against the coevolution theory, and to cast aside all doubt on the subject, we have decided to reanalyze the statistical bases on which this theory is founded. We come to the following conclusions: (1) the methods used in the above referred paper contain certain mistakes, and (2) the statistical foundations on which the coevolution theory is based are extremely robust. We have done this by critically appraising Amirnovin's paper and suggesting an alternative method based on the generation of random codes which, along with the method reported in the literature, allows us to evaluate the significance, in the genetic code, of different sets of amino acid pairs in biosynthetic relationships. In particular, by using this method and after building up a certain set of amino acid pairs reflecting the expectations of the coevolution theory, we show that the presence of this set in the genetic code would be obtained, purely by chance, with a probability of 6 × 10⁻⁵. This observation seems to provide particularly strong support to the coevolution theory. Received: 28 June 1999 / Accepted: 23 October 1999