Eco-evolution from deep time to contemporary dynamics: The role of timescales and rate modulators

Eco-evolutionary dynamics, or eco-evolution for short, are often thought to involve rapid demography (ecology) and equally rapid heritable phenotypic changes (evolution) leading to novel, emergent system behaviours. We argue that this focus on contemporary dynamics is too narrow: Eco-evolution should be extended, first, beyond pure demography to include all environmental dimensions and, second, to include slow eco-evolution which unfolds over thousands or millions of years. This extension allows us to conceptualise biological systems as occupying a two-dimensional time space along axes that capture the speed of ecology and evolution. Using Hutchinson's analogy: Time is the ‘theatre’ in which ecology and evolution are two interacting ‘players’. Eco-evolutionary systems are therefore dynamic: We identify modulators of ecological and evolutionary rates, like temperature or sensitivity to mutation, which can change the speed of ecology and evolution, and hence impact eco-evolution. Environmental change may synchronise the speed of ecology and evolution via these rate modulators, increasing the occurrence of eco-evolution and emergent system behaviours. This represents substantial challenges for prediction, especially in the context of global change. Our perspective attempts to integrate ecology and evolution across disciplines, from gene-regulatory networks to geomorphology and across timescales, from today to deep time.     

Darwin's ‘mystery of mysteries’: the role of sexual selection in plant speciation

Sexual selection is considered one of the key processes that contribute to the emergence of new species. While the connection between sexual selection and speciation has been supported by comparative studies, the mechanisms that mediate this connection remain unresolved, especially in plants. Similarly, it is not clear how speciation processes within plant populations translate into large-scale speciation dynamics. Here, we review the mechanisms through which sexual selection, pollination, and mate choice unfold and interact, and how they may ultimately produce reproductive isolation in plants. We also overview reproductive strategies that might influence sexual selection in plants and illustrate how functional traits might connect speciation at the population level (population differentiation, evolution of reproductive barriers; i.e. microevolution) with evolution above the species level (macroevolution). We also identify outstanding questions in the field, and suitable data and tools for their resolution. Altogether, this effort motivates further research focused on plants, which might potentially broaden our general understanding of speciation by sexual selection, a major concept in evolutionary biology.     

甜菊叶愈伤组织诱导过程中叶绿体的超微结构变化

观察了甜菊(Stevia rebaudiana Bertoni)叶外植体愈伤组织诱导过程中叶绿体的超微结构变化。结果表明,当叶外植体转移到培养基上培养后,叶绿体的片层结构逐渐退化。在叶绿体发生退化的过程中伴有叶绿体出芽和原质体的形成。推测新产生的原质体来自叶绿体产生的芽状体。而叶绿体本身最后完全解体消失。叶绿体超微结构的这种变化与高度液泡化的叶肉细胞脱分化至分生状态是平行的。随着培养的进行,分生状态的细胞发生液泡化变为薄壁细胞时,在愈伤组织表层的细胞中,质体重新形成片层结构,而内部细胞的质体则充满淀粉粒。     

Biogeography of the octocorallian coelenterate fauna of southern Africa

The geographical and bathymetric distribution of southern African octocorals is analysed. Of the over 200 estimated species of regional octocorals, 81 confirmed and adequately described species are studied using a radial sector method. Two primary faunal components are recognized–endemic (53.3% of the fauna by numbers of species) and Indo-Pacific (39.4%). An Atlantic component contributes only minimally (about 1.7%), while the remaining fauna is made up of cosmopolites (2.8%) and scattered species (2.8%). A subantarctic component is not evident for the present-day, although evidence for previous contact is presented. A sister-group analysis using genera as a guide to sister species, shows the biogeographic affinities for the present-day fauna as a whole to be 45% Indo-Pacific, 31% cosmopolitan, 10% endemic, 10% Atlantic and 4% southern oceans (subantarctic). Applying the same method to only those genera with endemic species shows the affinities of the present-day endemic fauna to be 27.5% Indo-Pacific, 27.5% endemic, 24% cosmopolitan, 14% Atlantic and 7% subantarctic. Clearly defined boundaries for west, south, and east coast faunas (as recognized by previous authors in describing various intertidal faunas) are found not be present with regard to the octocoral fauna (largely due to its overwhelmingly subtidal nature). Instead two primary zoogeographic provinces are recognized–the Cape Endemic Province (extending from Liideritz to Inhaca Island) and the south-western fringe of the Indo-Pacific Province from East London north-eastwards. An overlap zone between these two is recognized between East London and Inhaca Island, with the region in the vicinity of Richards Bay having an essentially evenly mixed fauna (roughly 50% Cape Endemic Province and 50% Indo-Pacific). Of the 84 octocoral genera recorded for the region, seven (or 8.3%) are endemic, and of these, five are monotypic while two are ditypic. The fauna is shown to be predominantly sublittoral (about 95% by numbers of species), the shallow sublittoral (< 100 m in depth) being the region with highest species richness. Pennatulaceans are eurybathic (intertidal to 4756 m) and clearly show a high proportion of cosmopolites (20% of presently identified species). Soft corals are stenobathic and restricted to the intertidal, continental shelf and uppermost portion of the continental slope (<500m), while gorgonians are intermediate in depth distributon (intertidal to 1200 m). No cosmopolitan alcyonaceans are presently recorded. The centre of the Cape Endemic Province is the Agulhas Bank–an extensive region of shallow continential shelf (< 200 m in depth) between Cape Town and East London. Two regions of octocoral radiation for southern Africa are postulated–the Agulhas Bank and the western Indian Ocean.     

DIVERGENT AMBULATORY AND GROOMING BEHAVIOR IN SERIALLY BOTTLENECKED LINES OF THE HOUSEFLY

The extent of genome-wide restructuring predicted in bottleneck models of speciation is addressed in assays of non-reproductive behavior in lines of the housefly. After five serial founder-flush cycles of one of three sizes (1, 4, or 16 pairs), each bottleneck line showed significant differentiation from the outbred control in ambulatory levels and grooming sequences in videotaped records of precopulatory activity. Only one line (4-pair) showed overall lethargy which was associated to inbreeding depression in egg-to-adult viability, thus exemplifying a case of probable extinction due to bottlenecks. The two most hyperactive lines (1- and 16-pair) showed very similar directions of differentiation from the control in locomotor activity and grooming behavior, as well as in mating behavior evaluated from a separate study. This high congruence suggested that directional selection toward the phenotypic optima of the ancestor operated on the bottleneck populations and that a 10-fold difference in theoretical inbreeding coefficients did not affect the magnitude of response. The remaining two bottleneck lines showed some independence from these general trajectories, their divergence along minor axes of ancestral intercorrelation structure possibly being more important to the formation of new species. Significant perturbations of the thresholds for execution of grooming and locomotor movements suggested increased evolutionary potential for ritualization (i.e., sexual selection for adoption of non-reproductive behavior into courtship repertoire) due to bottlenecks.     

GENETIC DIFFERENTIATION OF POLISH POPULATIONS OF SOREX ARANEUS L. III. INTERCHROMOSOMAL RECOMBINATION IN A HYBRID ZONE

Two parapatric chromosomal races of the common shrew (Sorex araneus) in Poland differ in their complement of metacentric arm combinations: hk, io, gr, nm (race IV), and hi, ko, gm, np (race II). In hybrids, these eight race-diagnostic metacentrics form two randomly segregating complexes. The first complex (C₁) occurs in the form of a ring configuration ok/kh/hi/io, or a chain o/ok/kh/hi/i (when there is Robertsonian polymorphism of the element io). The second complex (C₂) always takes the form of a six-element chain configuration r/rg/gm/mn/np/p. The C₂ complex may be shortened to five or even four elements, when acrocentrics g, m and n are present. In the contact zone we found shrews of pure races (race II or IV), as well as hybrids with C₁ or C₂ complexes, and recombinants hi, ko, gr, nm. Complex heterozygotes are likely to suffer reduced fertility due to malsegregation at meiosis. However, the C₁ hybrids with ring configurations occur with a high frequency throughout the contact zone. This suggest that their fitness is only slightly lowered relative to pure race individuals, in contrast to the hybrids with C₁ or C₂ chain configurations, which presumably have a more heavily reduced fertility. On the other hand, at the center of the zone there is a high proportion of recombinants, which, being chromosomal homozygotes, should display normal meiotic segregation. Furthermore, the high frequencies of recombinants within the contact zone should facilitate gene flow between the races. The occurrence of recombinants plays a similar role as the appearance of the maximum frequencies of acrocentric homozygotes described in several contact zones of S. araneus.     

DEVELOPMENTAL MUTANTS OF VOLVOX: DOES MUTATION RECREATE THE PATTERNS OF PHYLOGENETIC DIVERSITY?

The nature of the variation which is created by mutation can show how the direction of evolution is constrained by internal biases arising from development and pre-existing design. We have attempted to quantify these biases by measuring eight life history characters in developmental mutants of Volvox carteri. Most of the mutants in our sample were inferior to the wild type, but deviated by less than tenfold from the wild-type mean. Characters differed in mutability, suggesting different levels of canalisation. Most correlations between life history characters among strains were positive, but there was a significant negative correlation between the size and the number of reproductive cells, suggesting an upper limit to the total quantity of germ produced by individuals. The most extreme phenotypes in our sample were very vigorous, showing that not all mutations of large effect are unconditionally deleterious. We investigated the effect of developmental constraints on the course of evolution by comparing the variance and covariance patterns among mutant strains with those among species in the family Volvocaceae. A close correspondence between patterns at these two levels would suggest that pre-existing design has a strong influence on evolution, while little or no correspondence shows the action of selection. The variance generated by mutation was equal to that generated by speciation in the family Volvocaceae, the genus Volvox, or the section Merillosphaera, depending on the character considered. We found that mutation changes the volume of somatic tissue independently of the quantity of germ tissue, so that the interspecific correlation between soma and germ can be attributed to selection. The negative correlation between size and number of germ cells among mutants of V. carteri is also seen among the larger members of the family (Volvox spp.), but not among the smaller members, suggesting a powerful design constraint that may be responsible for the absence of larger forms in the entire group.     

Reconstructing the Pleistocene geography of the Aphelocoma jays (Corvidae)

Understanding historical distributions of species and evolving lineages has been a topic of considerable interest, yet methods used to date have not provided detailed, quantitative distributional hypotheses. Here, we present a technique based on models of species’ ecological niches and Pleistocene climate reconstructions that provides such hypotheses, providing the example of reconstructions for the Aphelocoma jays. We demonstrate in general a greater degree than expected of stability in jay species’ distributional areas back through at least the most recent glaciation event, and that existing patterns of genetic differentiation may date to before the Late Pleistocene glaciations. More generally, the method offers the potential for reconstructing historical distributions of species or lineages, and providing a detailed geographic framework for addressing many biogeographic and systematic questions.