Ecological opportunity and sexual selection together predict adaptive radiation

A fundamental challenge to our understanding of biodiversity is to explain why some groups of species undergo adaptive radiations, diversifying extensively into many and varied species, whereas others do not. Both extrinsic environmental factors (for example, resource availability, climate) and intrinsic lineage-specific traits (for example, behavioural or morphological traits, genetic architecture) influence diversification, but few studies have addressed how such factors interact. Radiations of cichlid fishes in the African Great Lakes provide some of the most dramatic cases of species diversification. However, most cichlid lineages in African lakes have not undergone adaptive radiations. Here we compile data on cichlid colonization and diversification in 46 African lakes, along with lake environmental features and information about the traits of colonizing cichlid lineages, to investigate why adaptive radiation does and does not occur. We find that extrinsic environmental factors related to ecological opportunity and intrinsic lineage-specific traits related to sexual selection both strongly influence whether cichlids radiate. Cichlids are more likely to radiate in deep lakes, in regions with more incident solar radiation and in lakes where there has been more time for diversification. Weak or negative associations between diversification and lake surface area indicate that cichlid speciation is not constrained by area, in contrast to diversification in many terrestrial taxa. Among the suite of intrinsic traits that we investigate, sexual dichromatism, a surrogate for the intensity of sexual selection, is consistently positively associated with diversification. Thus, for cichlids, it is the coincidence between ecological opportunity and sexual selection that best predicts whether adaptive radiation will occur. These findings suggest that adaptive radiation is predictable, but only when species traits and environmental factors are jointly considered.     

The effects of competition and predation on diversification in a model adaptive radiation

Much of life's diversity is thought to have arisen through successive rounds of adaptive radiation-the rapid diversification of a lineage into a range of ecologically and phenotypically distinct species. Both resource competition and predation have been suggested as mechanisms driving this process, although the former is better studied than the latter. Here we show experimentally how predation by a protist, Tetrahymena thermophila, affects diversification in a model adaptive radiation of the bacterial prey, Pseudomonas fluorescens. We estimate the frequency-dependent fitness functions of competing niche-specialist prey in the presence and absence of predation, and use these to test hypotheses about the extent (measured as the number of new genotypes) and rate of diversification. Competition and predation independently generated diversifying selection that we show is capable of driving prey diversification to similar extents but at different rates, diversification being markedly delayed in the presence of predators. The cause of this delay stems from weaker diversifying selection due to the reduction in prey density caused by predation. Our results suggest that predation may play an under-appreciated role in driving adaptive radiations.     

Sexual dimorphism and adaptive radiation in Anolis lizards

Sexual dimorphism is widespread and substantial throughout the animal world. It is surprising, then, that such a pervasive source of biological diversity has not been integrated into studies of adaptive radiation, despite extensive and growing attention to both phenomena. Rather, most studies of adaptive radiation either group individuals without regard to sex or focus solely on one sex. Here we show that sexual differences contribute substantially to the ecomorphological diversity produced by the adaptive radiations of West Indian Anolis lizards: within anole species, males and females occupy mostly non-overlapping parts of morphological space; the overall extent of sexual variation is large relative to interspecific variation; and the degree of variation depends on ecological type. Thus, when sexual dimorphism in ecologically relevant traits is substantial, ignoring its contribution may significantly underestimate the adaptive component of evolutionary radiation. Conversely, if sexual dimorphism and interspecific divergence are alternative means of ecological diversification, then the degree of sexual dimorphism may be negatively related to the extent of adaptive radiation.     

Evolution of ecological differences in the Old World leaf warblers.

Sympatric species that belong to the same ecological guild usually differ in their behaviour and morphology, and these differences are often interpreted as adaptations to having to make use of different resources. Evidence supporting this interpretation comes from association between ecology and morphology among species, in which an a priori functional relationship is reasonable. But one problem with such comparisons is that members of a guild may be closely related, so the more closely related species can share a greater similarity in their morphology and ecology simply as a result of the lingering legacy of a common ancestor. In principle, the importance of historical legacy can be evaluated from phylogenetic relationships and times since divergence for all species, but this is rarely possible because these data are not available. Here we use a phylogeny for eight sympatric species of warbler in the genus Phylloscopus, based on their mitochondrial DNA sequences, to remove the effects of historical legacy. Without these effects, we find strong support for adaptive interpretations of among-species variation in habitat selection, prey-size choice and feeding method. Ecological variation along any of these three niche axes is associated with predictable morphological variation. We also find evidence for historical legacy in that more closely related species are often more similar behaviourally and morphologically. This paradoxical result can be reconciled because the most closely related species tend to differ along only one niche axis, habitat choice. In contrast, the evolution of prey-size choice and feeding method occurred rapidly and early in the diversification of this group. Once a new ecological zone was occupied, subsequent morphological change along these niche axes was limited, accounting for the similarity of closely related species.     

Adaptive radiation of multituberculate mammals before the extinction of dinosaurs

The Cretaceous-Paleogene mass extinction approximately 66 million years ago is conventionally thought to have been a turning point in mammalian evolution. Prior to that event and for the first two-thirds of their evolutionary history, mammals were mostly confined to roles as generalized, small-bodied, nocturnal insectivores, presumably under selection pressures from dinosaurs. Release from these pressures, by extinction of non-avian dinosaurs at the Cretaceous-Paleogene boundary, triggered ecological diversification of mammals. Although recent individual fossil discoveries have shown that some mammalian lineages diversified ecologically during the Mesozoic era, comprehensive ecological analyses of mammalian groups crossing the Cretaceous-Paleogene boundary are lacking. Such analyses are needed because diversification analyses of living taxa allow only indirect inferences of past ecosystems. Here we show that in arguably the most evolutionarily successful clade of Mesozoic mammals, the Multituberculata, an adaptive radiation began at least 20 million years before the extinction of non-avian dinosaurs and continued across the Cretaceous-Paleogene boundary. Disparity in dental complexity, which relates to the range of diets, rose sharply in step with generic richness and disparity in body size. Moreover, maximum dental complexity and body size demonstrate an adaptive shift towards increased herbivory. This dietary expansion tracked the ecological rise of angiosperms and suggests that the resources that were available to multituberculates were relatively unaffected by the Cretaceous-Paleogene mass extinction. Taken together, our results indicate that mammals were able to take advantage of new ecological opportunities in the Mesozoic and that at least some of these opportunities persisted through the Cretaceous-Paleogene mass extinction. Similar broad-scale ecomorphological inventories of other radiations may help to constrain the possible causes of mass extinctions.     

Intraspecific competition favours niche width expansion in Drosophila melanogaster

Ecologists have proposed that when interspecific competition is reduced, competition within a species becomes a potent evolutionary force leading to rapid diversification. This view reflects the observation that populations invading species-poor communities frequently evolve broader niches. Niche expansion can be associated with an increase in phenotypic variance (known as character release), with the evolution of polymorphisms, or with divergence into many species using distinct resources (adaptive radiation). The relationship between intraspecific competition and diversification is known from theory, and has been used as the foundation for some models of speciation. However, there has been little empirical proof that niches evolve in response to intraspecific competition. To test this hypothesis, I introduced cadmium-intolerant Drosophila melanogaster populations to environments containing both cadmium-free and cadmium-laced resources. Here I show that populations experiencing high competition adapted to cadmium more rapidly than low competition populations. This provides experimental confirmation that competition in a population can drive niche expansion onto new resources for which competition is less severe.     

Loss and recovery of wings in stick insects

The evolution of wings was the central adaptation allowing insects to escape predators, exploit scattered resources, and disperse into new niches, resulting in radiations into vast numbers of species. Despite the presumed evolutionary advantages associated with full-sized wings (macroptery), nearly all pterygote (winged) orders have many partially winged (brachypterous) or wingless (apterous) lineages, and some entire orders are secondarily wingless (for example, fleas, lice, grylloblattids and mantophasmatids), with about 5% of extant pterygote species being flightless. Thousands of independent transitions from a winged form to winglessness have occurred during the course of insect evolution; however, an evolutionary reversal from a flightless to a volant form has never been demonstrated clearly for any pterygote lineage. Such a reversal is considered highly unlikely because complex interactions between nerves, muscles, sclerites and wing foils are required to accommodate flight. Here we show that stick insects (order Phasmatodea) diversified as wingless insects and that wings were derived secondarily, perhaps on many occasions. These results suggest that wing developmental pathways are conserved in wingless phasmids, and that 're-evolution' of wings has had an unrecognized role in insect diversification.     

Productivity-biodiversity relationships depend on the history of community assembly

Identification of the causes of productivity-species diversity relationships remains a central topic of ecological research. Different relations have been attributed to the influence of disturbance, consumers, niche specialization and spatial scale. One unexplored cause is the history of community assembly, the partly stochastic sequential arrival of species from a regional pool of potential community members. The sequence of species arrival can greatly affect community structure. If assembly sequence interacts with productivity to influence diversity, different sequences can contribute to variation in productivity-diversity relationships. Here we report a test of this hypothesis by assembling aquatic microbial communities at five productivity levels using four assembly sequences. About 30 generations after assembly, productivity-diversity relationships took various forms, including a positive, a hump-shaped, a U-shaped and a non-significant pattern, depending on assembly sequence. This variation resulted from idiosyncratic joint effects of assembly sequence, productivity and species identity on species abundances. We suggest that the history of community assembly should be added to the growing list of factors that influence productivity-biodiversity patterns.     

The role of parasites in sympatric and allopatric host diversification

Exploiters (parasites and predators) are thought to play a significant role in diversification, and ultimately speciation, of their hosts or prey. Exploiters may drive sympatric (within-population) diversification if there are a variety of exploiter-resistance strategies or fitness costs associated with exploiter resistance. Exploiters may also drive allopatric (between-population) diversification by creating different selection pressures and increasing the rate of random divergence. We examined the effect of a virulent viral parasite (phage) on the diversification of the bacterium Pseudomonas fluorescens in spatially structured microcosms. Here we show that in the absence of phages, bacteria rapidly diversified into spatial niche specialists with similar patterns of diversity across replicate populations. In the presence of phages, sympatric diversity was greatly reduced, as a result of phage-imposed reductions in host density decreasing competition for resources. In contrast, allopatric diversity was greatly increased as a result of phage-imposed selection for resistance, which caused populations to follow divergent evolutionary trajectories. These results show that exploiters can drive diversification between populations, but may inhibit diversification within populations by opposing diversifying selection that arises from resource competition.     

山地荒漠草原不同植物群落物种-多度关系分析

在群落样方调查的基础上,计算群落物种重要值,确定群落类型.对群落物种重要值采用分割线段模型(broken stick model)、对数级数分布(Log series distr ibution)、对数正态分布(Lognormal distribution)、几何级数分布(Geometric series distribution)等4个种-多度模型进行拟合.结果表明石质山坡(长芒草Stipa bungeana-冷蒿Artemisia frigida)群落对数级数拟合结果更好,土质山坡群落(短花针茅Stipa brevifora-长芒草Stipa bungeana)对数正态分布拟合更好,说明了2个群落结构是对2个样地环境特点与放牧干扰相互作用的响应.沙化草地(油蒿Artemisiaordosica-刺叶柄棘豆Oxytropis aciphlla)群落中油蒿处于极优势地位,占据整个群落的大多数生态空间.拟合检验表明该山地草地植被物种分布格局较符合随机生态位假说和生态位占领假说,严重的放牧干扰不仅降低了优势物种在群落中的优势程度,使次优势物种及其他物种也没能较好的发育,说明外部非生物环境对群落胁迫程度较大.     

Sophisticated particle-feeding in a large Early Cambrian crustacean

Most Cambrian arthropods employed simple feeding mechanisms requiring only low degrees of appendage differentiation. In contrast, post-Cambrian crustaceans exhibit a wide diversity of feeding specializations and possess a vast ecological repertoire. Crustaceans are evident in the Cambrian fossil record, but have hitherto been known exclusively from small individuals with limited appendage differentiation. Here we describe a sophisticated feeding apparatus from an Early Cambrian arthropod that had a body length of several centimetres. Details of the mouthparts resolve this taxon as a probable crown-group (pan)crustacean, while its feeding style, which allowed it to generate and handle fine food particles, significantly expands the known ecological capabilities of Cambrian arthropods. This Early Cambrian record predates the major expansions of large-bodied, particle-handling crustaceans by at least one hundred million years, emphasizing the importance of ecological context in driving adaptive radiations.     

Speciation along environmental gradients

Traditional discussions of speciation are based on geographical patterns of species ranges. In allopatric speciation, long-term geographical isolation generates reproductively isolated and spatially segregated descendant species. In the absence of geographical barriers, diversification is hindered by gene flow. Yet a growing body of phylogenetic and experimental data suggests that closely related species often occur in sympatry or have adjacent ranges in regions over which environmental changes are gradual and do not prevent gene flow. Theory has identified a variety of evolutionary processes that can result in speciation under sympatric conditions, with some recent advances concentrating on the phenomenon of evolutionary branching. Here we establish a link between geographical patterns and ecological processes of speciation by studying evolutionary branching in spatially structured populations. We show that along an environmental gradient, evolutionary branching can occur much more easily than in non-spatial models. This facilitation is most pronounced for gradients of intermediate slope. Moreover, spatial evolutionary branching readily generates patterns of spatial segregation and abutment between the emerging species. Our results highlight the importance of local processes of adaptive divergence for geographical patterns of speciation, and caution against pitfalls of inferring past speciation processes from present biogeographical patterns.     

土壤养分梯度下黄河三角洲湿地植物的生态位

利用Levins生态位宽度公式和Pianka生态位重叠公式,在土壤有机质、土壤总磷、土壤总氮3个养分梯度上分别对黄河三角洲湿地19个植物种的生态位宽度值和生态位重叠值进行了分析.结果表明:各植物种在3个养分梯度上的生态位宽度值差异不大,优势种柽柳(Tamarix chinensis)、盐地碱蓬(Suaeda salsa)、芦苇(Phragmites australis)的生态位宽度值均较大;植物种之间的生态位重叠一般均较小,说明植物种在土壤养分梯度上产生了生态位分化.     

人工林生态系统生物多样性与生产力的关系

生物多样性与生产力之间的关系是生态学界近来争论的热点之一。笔者简要介绍了现有关于生物多样性影响生态系统功能有关抽样效应、生态位互补、保险效应、冗余机制、基因-环境互作、边际效应及中性理论等作用机制,以及现有的关于生产力和多样性之间关系的主要观点。在此基础上,阐述了人工林生物多样性的研究现状和人工林生物多样性与生态功能的关系,重点讨论了造林树种选择、造林地清理、造林密度、间伐、林农复合经营、轮伐期长短等经营措施对人工林生物多样性的影响。人工林生物多样性的高低,很大程度上取决于林分结构的构建和调控,任何左右林分结构的经营措施都有可能对人工林生物多样性产生影响。因此,从生物多样性维护及生态功能发挥的角度看,决定应该营造什么样的人工林以及该采取怎样的经营措施是人工林可持续经营和长期立地生产力维持的关键。     

Biodiversity improves water quality through niche partitioning

Excessive nutrient loading of water bodies is a leading cause of water pollution worldwide, and controlling nutrient levels in watersheds is a primary objective of most environmental policy. Over the past two decades, much research has shown that ecosystems with more species are more efficient at removing nutrients from soil and water than are ecosystems with fewer species. This has led some to suggest that conservation of biodiversity might be a useful tool for managing nutrient uptake and storage, but this suggestion has been controversial, in part because the specific biological mechanisms by which species diversity influences nutrient uptake have not been identified. Here I use a model system of stream biofilms to show that niche partitioning among species of algae can increase the uptake and storage of nitrate, a nutrient pollutant of global concern. I manipulated the number of species of algae growing in the biofilms of 150 stream mesocosms that had been set up to mimic the variety of flow habitats and disturbance regimes that are typical of natural streams. Nitrogen uptake rates, as measured by using (15)N-labelled nitrate, increased linearly with species richness and were driven by niche differences among species. As different forms of algae came to dominate each unique habitat in a stream, the more diverse communities achieved a higher biomass and greater (15)N uptake. When these niche opportunities were experimentally removed by making all of the habitats in a stream uniform, diversity did not influence nitrogen uptake, and biofilms collapsed to a single dominant species. These results provide direct evidence that communities with more species take greater advantage of the niche opportunities in an environment, and this allows diverse systems to capture a greater proportion of biologically available resources such as nitrogen. One implication is that biodiversity may help to buffer natural ecosystems against the ecological impacts of nutrient pollution.     

Immigration history controls diversification in experimental adaptive radiation

Diversity in biological communities is a historical product of immigration, diversification and extinction, but the combined effect of these processes is poorly understood. Here we show that the order and timing of immigration controls the extent of diversification. When an ancestral bacterial genotype was introduced into a spatially structured habitat, it rapidly diversified into multiple niche-specialist types. However, diversification was suppressed when a niche-specialist type was introduced before, or shortly after, introduction of the ancestral genotype. In contrast, little suppression occurred when the same niche specialist was introduced relatively late. The negative impact of early arriving immigrants was attributable to the historically sensitive outcome of interactions involving neutral competition and indirect facilitation. Ultimately, the entire boom-and-bust dynamics of adaptive radiation were altered. These results demonstrate that immigration and diversification are tightly linked processes, with small differences in immigration history greatly affecting the evolutionary emergence of diversity.     

Ecosystem size determines food-chain length in lakes

Food-chain length is an important characteristic of ecological communities: it influences community structure, ecosystem functions and contaminant concentrations in top predators. Since Elton first noted that food-chain length was variable among natural systems, ecologists have considered many explanatory hypotheses, but few are supported by empirical evidence. Here we test three hypotheses that predict food-chain length to be determined by productivity alone (productivity hypothesis), ecosystem size alone (ecosystem-size hypothesis) or a combination of productivity and ecosystem size (productive-space hypothesis). The productivity and productive-space hypotheses propose that food-chain length should increase with increasing resource availability; however, the productivity hypothesis does not include ecosystem size as a determinant of resource availability. The ecosystem-size hypothesis is based on the relationship between ecosystem size and species diversity, habitat availability and habitat heterogeneity. We find that food-chain length increases with ecosystem size, but that the length of the food chain is not related to productivity. Our results support the hypothesis that ecosystem size, and not resource availability, determines food-chain length in these natural ecosystems.     

Host shift to an invasive plant triggers rapid animal hybrid speciation

Speciation in animals is almost always envisioned as the split of an existing lineage into an ancestral and a derived species. An alternative speciation route is homoploid hybrid speciation in which two ancestral taxa give rise to a third, derived, species by hybridization without a change in chromosome number. Although theoretically possible it has been regarded as rare and hence of little importance in animals. On the basis of molecular and chromosomal evidence, hybridization is the best explanation for the origin of a handful of extant diploid bisexual animal taxa. Here we report the first case in which hybridization between two host-specific animals (tephritid fruitflies) is clearly associated with the shift to a new resource. Such a hybrid host shift presents an ecologically robust scenario for animal hybrid speciation because it offers a potential mechanism for reproductive isolation through differential adaptation to a new ecological niche. The necessary conditions for this mechanism of speciation are common in parasitic animals, which represent much of animal diversity. The frequency of homoploid hybrid speciation in animals may therefore be higher than previously assumed.     

唐家河自然保护区斑羚春冬季对生境的选择

1998年3－4月和1999年1－2月，作者在四川省青川县唐家河自然保护区，应用主成分分析和判别分析的方法，对斑羚（Naemorhedusgoral）春冬季对生境的利用进行了研究。研究结果表明：影响斑羚春季生境选择的主要生态因子为人干扰度、植被型、坡位、郁闭度、水源乔木距离、乔木大小、灌木距离、坡度、灌木大小、坡度、食物丰富度、乔木密度；斑羚春冬季生境选择的分离主要表现为植被型、郁闭度、水源、坡位、坡度、乔木大小、乔木距离、灌木密度等生态因子的分离。