The role of KNOX genes in the evolution of morphological novelty in Streptocarpus

The genus Streptocarpus comprises species with diverse body plans. Caulescent species produce leaves from a conventional shoot apical meristem (SAM), whereas acaulescent species lack a conventional SAM and produce only a single leaf (the unifoliate form) or clusters of leaves from the base of more mature leaves (the rosulate form). These distinct morphologies reflect fundamental differences in the role of the SAM and the process of leaf specification. A subfamily of KNOTTED-like homeobox (KNOX) genes are known to be important in regulating meristem function and leaf development in model species with conventional morphologies. To test the involvement of KNOX genes in Streptocarpus evolution, two parologous KNOX genes (SSTM1 and SSTM2) were isolated from species with different growth forms. Their phylogenetic analysis suggested a gene duplication before the subgeneric split of Streptocarpus and resolved species relationships, supporting multiple evolutionary origins of the rosulate and unifoliate morphologies. In S. saxorum, a caulescent species with a conventional SAM, KNOX proteins were expressed in the SAM and transiently downregulated in incipient leaf primordia. The ability of acaulescent species to initiate leaves from existing leaves was found to correlate with SSTM1 expression and KNOX protein accumulation in leaves and to reflect genetic differences at two loci. Neither locus corresponded to SSTM1, suggesting that cis-acting differences in SSTM1 regulation were not responsible for evolution of the rosulate and unifoliate forms. However, the involvement of KNOX proteins in leaf formation in rosulate species suggests that they have played an indirect role in the development of morphological diversity in Streptocarpus.     

Sexual isolation evolves faster than hybrid inviability in a diverse and sexually dimorphic genus of fish (Percidae: Etheostoma)

Abstract .Theory predicts that sexual (or behavioral) isolation will be the first form of reproductive isolation to evolve in lineages characterized by sexual selection. Here I directly compare the rate of evolution of sexual isolation with that of hybrid inviability in a diverse and sexually dimorphic genus of freshwater fish. The magnitude of both sexual isolation and hybrid inviability were quantified for multiple pairs of allopatric species. Rates of evolution were inferred by comparing genetic distances of these species pairs with the magnitude of each form of reproductive isolation: the slope of the regression of genetic distance on the magnitude of reproductive isolation represents the rate of evolution. Of the two forms of isolation, the magnitude of sexual isolation exhibited the steeper slope of regression, indicating that sexual isolation will tend to evolve to completion earlier than hybrid inviability, strictly as a by-product of evolution in geographically isolated populations. Additional evidence from the literature is used to qualitatively compare rates of evolution of sexual isolation with that of other forms of reproductive isolation. Preliminary comparisons support the prediction that sexual isolation will evolve more rapidly than other forms. Because Etheostoma is characterized by striking sexual dimorphism, these results are consistent with the hypothesis that sexual selection for exaggerated mate-recognition characters causes the relatively rapid evolution of sexual isolation.     

Understanding proboscidean evolution: a formidable task

A new approach to proboscidean evolution depicts taxa in three major radiations. This approach highlights general proboscidean evolutionary trends and origins more than the specific relationships among them. Data from more than 55 million years of evolution help to interpret how the integration of primitive and derived characters was essential to proboscidean success. Only two, or perhaps three, species remain of approximately 164 that lived in the past. Extinct forms were extremely cosmopolitan, occupying a variety of habitats, from deserts to mountain tops, on all continents except Australia and Antarctica. Challenges for future investigators include a better understanding of structure and function of infrasonic call production and perception, brain features, and reproductive biology in extinct proboscideans based on inferences from living forms.     

Patterns of reproductive‐mode evolution in Old World tree frogs (Anura,Rhacophoridae)

The Old World tree frogs (Anura: Rhacophoridae), with 387 species, display a remarkable diversity of reproductive modes – aquatic breeding, terrestrial gel nesting, terrestrial foam nesting and terrestrial direct development. The evolution of these modes has until now remained poorly studied in the context of recent phylogenies for the clade. Here, we use newly obtained DNA sequences from three nuclear and two mitochondrial gene fragments, together with previously published sequence data, to generate a well‐resolved phylogeny from which we determine major patterns of reproductive‐mode evolution. We show that basal rhacophorids have fully aquatic eggs and larvae. Bayesian ancestral‐state reconstructions suggest that terrestrial gel‐encapsulated eggs, with early stages of larval development completed within the egg outside of water, are an intermediate stage in the evolution of terrestrial direct development and foam nesting. The ancestral forms of almost all currently recognized genera (except the fully aquatic basal forms) have a high likelihood of being terrestrial gel nesters. Direct development and foam nesting each appear to have evolved at least twice within Rhacophoridae, suggesting that reproductive modes are labile and may arise multiple times independently. Evolution from a fully aquatic reproductive mode to more terrestrial modes (direct development and foam nesting) occurs through intermediate gel nesting ancestral forms. This suggests that gel nesting is not only a possible transitional state for the evolution of terrestriality, but also that it is a versatile reproductive mode that may give rise to other terrestrial reproductive modes. Evolution of foam nesting may have enabled rhacophorids to lay a larger number of eggs in more open and drier habitats, where protection from desiccation is important. Terrestrial direct development allows frogs to lay eggs independent of bodies of water, in a diversity of humid habitats, and may represent a key innovation that facilitated the evolution of nearly half of all known rhacophorid species.     

Phylogenetic relationships in the iguanid lizard genus Liolaemus: multiple origins of viviparous reproduction and evidence for recurring Andean vicariance and dispersal

Phylogenetic relationships within the iguanid lizard genus Liolaemus are investigated using 1710 aligned base positions (785 phylogenetically informative) of mitochondrial DNA sequences, representing coding regions for eight tRNAs, ND2, and portions of ND1 and COL Sixty new sequences ranging in length from 1736 to 1754 bases are compared with four previously reported sequences. Liolaemus species form two well-supported monophyletic groups of subgeneric status, Liolaemus and Eulaemus. These subgenera appear to have separated at least 12.6 million years ago based on the amount of molecular evolutionary divergence between them. Hypotheses that species occurring in the Andes, west of the Andes, and east of the Andes, each comprise distinct monophyletic groups are independently rejected statistically. The shortest estimate of phylogeny suggests mat Liolaemus originated either in the Andes or the eastern lowlands. Numerous evolutionary shifts have occurred between the Andes, and the eastern and western lowlands, suggesting recurring vicariance and dispersal. Species occurring at high elevations or high latitudes usually have viviparous reproduction. Depending on whether parity mode is considered reversible in Liolaemus , the most parsimonious reconstruction supports at least six independent origins of viviparity or at least three gains followed by three losses of viviparity among die 60 Liolaemus lineages examined.     

The Caribbean slipper spurge Euphorbia tithymaloides: the first example of a ring species in plants

A ring species arises when a parental population expands around an area of unsuitable habitat in such a way that when the two fronts meet they behave as distinct species while still being connected through a series of intergrading populations. Ring species offer great possibilities for studying the forces causing species divergence (e.g. the nature of pre-zygotic or post-zygotic reproductive isolation) or helping to maintain species integrity (e.g. reinforcement). Yet, ring species are extremely rare, and have only been documented convincingly in animals. Here, we present phylogenetic analyses of two nuclear gene regions from the Caribbean slipper spurge (Euphorbia tithymaloides) species complex that provide evidence that this group forms a ring species. These data show that the species complex originated in the area where Mexico and Guatemala meet, and expanded around the Caribbean basin along two distinct fronts: one eastward through the Yucatan Peninsula and into the Greater Antilles (GA); one southeastward through northern South America and then northward to the Lesser Antilles and eastern GA. The two terminal forms co-occur in the Virgin Islands and appear to be morphologically and ecologically distinct. Thus, our results suggest that Euphorbia tithymaloides is the first compelling example of a ring species in plants.     

A phenetic study of Crepis tectorum in Fennoscandia and Estonia

Morphological variation in Crepis tectorum in Fennoscandia and Estonia was assessed in a multivariate study of herbarium material. The variation pattern revealed was more complex than that detected in earlier studies of the species in this area. There was a gradual transition between the common weed form and a similar form occurring on rocky outcrops throughout the area, particularly along the coasts and on steep hillsides on the mainland. The two endemic alvar forms on the Baltic islands of Oland and Gotland were distinct from each other but overlapped slightly with other groups. An additional outcrop form from the coast of South Finland graded into the Oland alvar form while the alvar form from Gotland was more similar to the widespread forms on the mainland of Fennoscandia and in Estonia. An outgroup comparison suggested that the widespread forms on the mainland have retained a more "ancient" morphology than the geographically restricted Baltic forms. Morphological and distributional data are used to test and extend earlier hypotheses regarding the evolution of C. tectorum in these areas. An eastern route seems likely for the first immigrations of the species into Fennoscandia. I suggest that the Baltic forms and at least some populations of the weed form were derived from non-weedy outcrop populations on the mainland, the first group when the Baltic lowland areas emerged from the sea, and the second group when man created suitable conditions for weed colonizations throughout Fennoscandia. Taxonomic implications of the results are also discussed.     

Molecular population genetics of accessory gland protein genes and testis-expressed genes in Drosophila mojavensis and D. arizonae

Molecular population genetic investigation of Drosophila male reproductive genes has focused primarily on melanogaster subgroup accessory gland protein genes (Acp's). Consistent with observations from male reproductive genes of numerous taxa, Acp's evolve more rapidly than nonreproductive genes. However, within the Drosophila genus, large data sets from additional types of male reproductive genes and from different species groups are lacking. Here we report findings from a molecular population genetics analysis of male reproductive genes of the repleta group species, Drosophila arizonae and D. mojavensis. We find that Acp's have dramatically higher average pairwise Ka/Ks (0.93) than testis-enriched genes (0.19) and previously reported melanogaster subgroup Acp's (0.42). Overall, 10 of 19 Acp's have Ka/Ks > 1 either in nonpolarized analyses or in at least one lineage of polarized analyses. Of the nine Acp's for which outgroup data were available, average Ka/Ks was considerably higher in D. mojavensis (2.08) than in D. arizonae (0.87). Contrasts of polymorphism and divergence suggest that adaptive protein evolution at Acp's is more common in D. mojavensis than in D. arizonae.     

A review of the evolution of viviparity in lizards: structure,function and physiology of the placenta

The aim of this review is to collate data relevant to understanding the evolution of viviparity in general, and complex placentae in particular. The wide range of reproductive modes exhibited by lizards provides a solid model system for investigating the evolution of viviparity. Within the lizards are oviparous species, viviparous species that have a very simple placenta and little nutrient uptake from the mother during pregnancy (lecithotrophic viviparity), through a range of species that have intermediate placental complexities and placental nutrient provision, to species that lay microlecithal eggs and most nutrients are provided across the placenta during development (obligate placentotrophy). In its commonest form, lecithotrophic viviparity, some uptake of water, inorganic ions and oxygen occurs from the mother to the embryo during pregnancy. In contrast, the evolution of complex placentae is rare, but has evolved at least five times. Where there is still predominantly a reliance on egg yolk, the omphaloplacenta seems to be paramount in the provision of nutrition to the embryo via histotrophy, whereas the chorioallantoic placenta is more likely involved in gas exchange. Reliance on provision of substantial organic nutrient is correlated with the regional specialisation of the chorioallantoic placenta to form a placentome for nutrient uptake, particularly lipids, and the further development of the gas exchange capabilities of the other parts of the chorioallantois.     

Habitat use and phenology of the large insular endemic grasshopper Acrostira euphorbiae (Orthoptera: Pamphagidae)

The habitat use and the phenology of the large grasshopper Acrostira euphorbiae García and Oromí endemic to La Palma (Canary Islands) are studied. This grasshopper is entirely dependent on the Canarian endemic shrub Euphorbia lamarckii both for food and to avoid predation. Adults stay on subapical branches during the day, probably to reduce the risk of predation, and climb up to the apex at night to feed. While females seem to ensure the genetic diversity of offspring by waiting for visits by different males, the latter have to move to guarantee their reproductive success. Monophagy in this species may be related to the year-round presence of tender shoots, and to the predator-repellent toxic latex found in the host plant. Unlike related species from continental areas, adults and nymphs of A. euphorbiae are present almost all year round, probably in adaptation to the particular climate of the islands. Nymphs are more abundant in winter, when Euphorbia leaves are most available. However, adults are more abundant than nymphs in spring, summer and autumn. Males develop more quickly than females, an apparent reproductive strategy based on achieving sexual maturity to coincide with females undergoing imaginal moulting. Matings start immediately after adult females emerge. Densities oscillated between 73 and 193 individuals ha(-1), which can be considered a low value compared with other continental pamphagid species.     

Evolutionary aspects of gonadotropin-releasing hormone and its receptor

Summary 1. Gonadotropin-releasing hormone (GnRH) was originally isolated as a hypothalamic peptide hormone that regulates the reproductive system by stimulating the release of gonadotropins from the anterior pituitary. However, during evolution the peptide was subject to gene duplication and structural changes, and multiple molecular forms have evolved.2. Eight variants of GnRH are known, and at least two different forms are expressed in species from all vertebrate classes: chicken GnRH II and a second, unique, GnRH isoform.3. The peptide has been recruited during evolution for diverse regulatory functions: as a neurotransmitter in the central and sympathetic nervous systems, as a paracrine regulator in the gonads and placenta, and as an autocrine regulator in tumor cells.4. Evidence suggests that in most species the early-evolved and highly conserved chicken GnRH II has a neurotransmitter function, while the second form, which varies across classes, has a physiologic role in regulating gonadotropin release.5. We review here evolutionary aspects of the family of GnRH peptides and their receptors.     

Testing the adaptive nature of radiation: growth form and life history divergence in the African grass genus Ehrharta (Poaceae: Ehrhartoideae)

In most documented examples of adaptive radiation, the processes underlying divergence in form and function are poorly explored and remain speculative. Here, data from a comparative seedling growth experiment are used to explore growth form divergence in Ehrharta, a group of grasses that radiated in seasonally arid environments of the Cape region of South Africa. Seedlings of eight Ehrharta species of variable growth form were grown in liquid culture under conditions of high resource availabilty for 56 d, during which time changes in dry mass, allocation, and leaf parameters were measured. The results of this experiment reveal the existence of distinct seedling growth patterns that are associated with differences in adult plant form and seasonal drought survival strategy. Specifically, species that utilize a reseeding strategy have higher seedling growth rates and flower earlier than species that persist by vegetative means. A correlation between species' growth rates and their native substrates suggests that edaphic heterogeneity has been central in directing the evolution of alternative persistence strategies and growth forms. Parsimony reconstruction identifies slow growth and an association with nutrient-deficient sandstone-derived soils as ancestral in Ehrharta, with fast growth evolving after a transition to richer shale- and granite-derived soils. The emergence of annual species in two fast-growth lineages suggests that the latter is a key step in the evolution of an ephemeral strategy. An association between plant function and habitat identifies the radiation of Ehrharta as adaptive.     

Where are all the moms? External fertilization predicts the rise of male parental care in bony fishes

Parental care shows remarkable variation across the animal kingdom, but while maternal and biparental care are common in terrestrial organisms, male‐only care dominates in aquatic species that provide care. Using the most complete phylogenetic tree of bony fishes to date, we test whether the opportunity for external fertilization in aquatic environments can explain the more frequent evolution of male care in this group. We show that paternal care has evolved at least 30 times independently in fish and is found exclusively in externally fertilizing species. Male care is positively associated with pair spawning, suggesting that confidence in paternity is an important determinant of the evolution of care. Crucially, while female care is constrained by other forms of reproductive investment, male care occurs more frequently when females invest heavily in gamete production. Our results suggest that moving control of fertilization outside of the female reproductive tract raises male confidence in parentage and increases the potential for paternal care, highlighting that in an aquatic environment in which fertilization is external, paternal care is an effective reproductive strategy.     

中国蹄盖蕨属的研究I─属下分类

蹄盖蕨属是蕨类植物中分类最困难的大属之一,中国、喜马拉雅和日本为其分布中心。隶属于本属的种实际上是由蹄盖蕨科或广义蹄盖蕨属中其他属被划分出去以后剩余下来的种类集合而成的“大杂烩”,因此,它们被放在一起并非是一个单系起源的自然类群;属下不同类群分化的程度相差也很大,有的类群进化活跃,近缘种很多,有的类群则很孤立,所以属下单位所包括的种数相差悬殊;孢子周壁表面褶皱的有无在本属的属下分类中占有重要的位置。已发表的中国蹄盖蕨属植物名称约有３００多个,它们初步被处理为１１７种和一些变种和杂种,本文建议把这些种分为１４个组和１５个系。     

A comparative phytochemical study of the diterpenes of some species of the genera Euphorbia and Elaeophorbia (Euphorhiaceae)

Nearly 60 species from the genera Euphorbia L. and Elaeophorbia Stapf were investigated for diterpenes of the classes tiglianes, ingenanes and ortho-esters. The diterpenes were isolated as their acetates by a micro-technique from latex or fresh herb material collected from several countries around the world. Authentication of the diterpenes was by chromatographic means using spectroscopic methods as back-up data. These compounds were absent from only 9% of the species examined. The most commonly occurring diterpene was ingenol, followed closely by 12-Deoxy phorbol. The ingenane derivative 5-Deoxy-ingenol was always detected as a minor companion to ingenol from species of the section Tithymalus. Species of the section Euphorbium contained mainly the diterpene ingenol although both tigliane and ortho-ester diterpenes were also present in some species. Species from the sections Anisophylhtm and Poinsettia did not contain diterpenes of the type in question, whilst species from the genus Elaeophorbia yielded ingenol. These results provide additional chemical evidence concerning recent suggestions on the subgeneric and generic status of Anisophyllum, Poinsettia, Tithymalus and Elaeophorbia.     

中国蹄盖蕨属的研究Ⅰ-属下分类

蹄盖蕨属是蕨类植物中分类最困难的大属之一,中国、喜马拉雅和日本为其分布中心。隶属于本属的种实际上是由蹄盖蕨科或广义蹄盖蕨属中其他属被划分出去以后剩余下来的种类集合而成的"大杂烩",因此,它们被放在一起并非是一个单系起源的自然类群;属下不同类群分化的程度相差也很大,有的类群进化活跃,近缘种很多,有的类群则很孤立,所以属下单位所包括的种数相差悬殊;孢子周壁表面褶皱的有无在本属的属下分类中占有重要的位置。已发表的中国蹄盖蕨属植物名称约有300多个,它们初步被处理为117种和一些变种和杂种,本文建议把这些种分为14个组和15个系。     

Reproductive output of invasive versus native plants

Aim Propagule size and output are critical for the ability of a plant species to colonize new environments. If invasive species have a greater reproductive output than native species (via more and/or larger seeds), then they will have a greater dispersal and establishment ability. Previous comparisons within plant genera, families or environments have conflicted over the differences in reproductive traits between native and invasive species. We went beyond a genus‐, family‐ or habitat‐specific approach and analysed data for plant reproductive traits from the global literature, to investigate whether: (1) seed mass and production differ between the original and introduced ranges of invasive species; (2) seed mass and production differ between invasives and natives; and (3) invasives produce more seeds per unit seed mass than natives. Location Global. Methods We combined an existing data set of native plant reproductive data with a new data compilation for invasive species. We used t‐tests to compare original and introduced range populations, two‐way ANOVAs to compare natives and invasives, and an ANCOVA to examine the relationship between seed mass and production for natives and invasives. The ANCOVA was performed again incorporating phylogenetically independent contrasts to overcome any phylogenetic bias in the data sets. Results Neither seed mass nor seed production of invasive species differed between their introduced and original ranges. We found no significant difference in seed mass between invasives and natives after growth form had been accounted for. Seed production was greater for invasive species overall and within herb and woody growth forms. For a given seed mass, invasive species produced 6.7‐fold (all species), 6.9‐fold (herbs only) and 26.1‐fold (woody species only) more seeds per individual per year than native species. The phylogenetic ANCOVA verified that this trend did not appear to be influenced by phylogenetic bias within either data set. Main conclusions This study provides the first global examination of both seed mass and production traits in native and invasive species. Invasive species express a strategy of greater seed production both overall and per unit seed mass compared with natives. The consequent increased likelihood of establishment from long‐distance seed dispersal may significantly contribute to the invasiveness of many exotic species.     

The evolution of plant reproductive systems: how often are transitions irreversible?

Flowering plants are characterized by striking variation in reproductive systems, and the evolutionary lability of their sexual traits is often considered a major driver of lineage diversification. But, evolutionary transitions in reproductive form and function are never entirely unconstrained and many changes exhibit strong directionality. Here, I consider why this occurs by examining transitions in pollination, mating and sexual systems, some of which have been considered irreversible. Among pollination systems, shifts from bee to hummingbird pollination are rarely reversible, whereas transitions from animal to wind pollination are occasionally reversed. Specialized pollination systems can become destabilized through a loss of pollinator service resulting in a return to generalized pollination, or more commonly a reliance on self-pollination. Homomorphic and heteromorphic self-incompatibility systems have multiple origins but breakdown to self-compatibility occurs much more frequently with little evidence for subsequent gains, at least over short time-spans. Similarly, numerous examples of the shift from outcrossing to predominant self-fertilization are known, but cases of reversal are very limited supporting the view that autogamy usually represents an evolutionary dead-end. The evolution of dioecy from hermaphroditism has also been considered irreversible, although recent evidence indicates that the occurrence of sex inconstancy and hybridization can lead to the origin of derived sexual systems from dioecy. The directionality of many transitions clearly refutes the notion of unconstrained reproductive flexibility, but novel adaptive solutions generally do not retrace earlier patterns of trait evolution.     

A mid-cretaceous origin of sociality in xylocopine bees with only two origins of true worker castes indicates severe barriers to eusociality

The origin of sterile worker castes, resulting in eusociality, represents one of the major evolutionary transitions in the history of life. Understanding how eusociality has evolved is therefore an important issue for understanding life on earth. Here we show that in the large bee subfamily Xylocopinae, a simple form of sociality was present in the ancestral lineage and there have been at least four reversions to purely solitary nesting. The ancestral form of sociality did not involve morphological worker castes and maximum colony sizes were very small. True worker castes, entailing a life-time commitment to non-reproductive roles, have evolved only twice, and only one of these resulted in discrete queen-worker morphologies. Our results indicate extremely high barriers to the evolution of eusociality. Its origins are likely to have required very unusual life-history and ecological circumstances, rather than the amount of time that selection can operate on more simple forms of sociality.