On the origin of feathers

The appearance of feathers defines the appearance of birds. A number of changes defined, preceded or accompanied the event. The changes were hierarchical in nature and included revolutions in genomic organization (i.e., HOX and the feather keratin genes), protein sequence and shape, the large scale organization of proteins into filaments, and in the geometry of the cells and their roles in the follicle. Changes at each of these levels differ or produced different products than found in its analog in reptiles. They are essentially unique to birds and produced an evolutionary novelty. I used analysis of extant structure and information on development to reconstruct key events in the evolution of feathers. The ancestral reptilian epidermal structure, while probably a scale or tubercles, is still unidentified. The structural genes of feather proteins (φ-keratin) are tandem repeats probably assembled from pre-existing exons. They are unlike the alpha-keratin of vertebrate soft epidermis. Amino-acid composition, shape, and behavior of feather keratins are unique among vertebrates. The 3-dimensional organization of the follicle and the developmental processes are also unique. Although we lack a complete understanding of the appearance and early role of feathers, they are clearly the results of novel events.     

Origins,bottlenecks, and present‐day diversity: Patterns of morphospace occupation in marine bivalves

It has long been known that species should not be distributed randomly in morphospace (a multidimensional trait space), even under simple models of evolution. However, recent studies suggest that position in morphospace can affect aspects of evolution such as the durations of clades and the species richness of their constituent taxa. Here we investigate the dynamics of morphospace occupancy in living and fossil marine bivalves using shell size and aspect ratio, two functionally important traits. Multiple lines of evidence indicate that the center of a family's morphospace today represents a location where taxonomic diversity is maximized, apparently owing to lower extinction rates. Within individual bivalve families, species with narrow geographic ranges are distributed throughout the morphospace but widespread species, which are generally expected to be extinction resistant, tend to be concentrated near the center. The morphospace centers of most species‐rich families today (defined as the median value for all species in the family) tend to be close to the positions of the family founders, further suggesting an association between position in morphospace and net diversification rates. However, trajectories of individual subclades (genera) are inconsistent with the center of morphospace being an evolutionary attractor.     

The European Far West: Miocene mammal isolation,diversity and turnover in the Iberian Peninsula

Aim To analyse the diversity dynamics of Miocene mammalian faunas in the Iberian Peninsula in order to determine whether the patterns are related to the dispersal of taxa from other areas into this region. Location Mainly the Iberian Peninsula, but two close geographical areas (Central Europe and the Eastern Mediterranean) are also considered in some of our calculations. Methods Genus‐level faunal lists for a total of 299 localities from the Iberian Peninsula, covering 10 successive biochronological units [Mammal Neogene (MN) zones] that span from the latest Early Miocene to the early Pliocene (about 17–4 Ma), were compiled. The dataset was expanded with a further 331 localities in Central Europe and the Eastern Mediterranean for the same time span. Next, a taxonomically standardized database was used to create composite faunal lists of micro‐ and macromammalian genera present during each MN zone. Separate genera‐by‐MN‐zone matrices for both micro‐ and macromammals were built for each region. Mean standing diversity as well as origination and extinction rates were calculated for the Iberian Peninsula, and their correlation with preservation rates is discussed. Simpson’s coefficient of faunal similarity with Central Europe and the Eastern Mediterranean was calculated in order to evaluate whether diversity patterns were related to changes in the affinity of the Iberian mammalian faunas with those of other regions. Results Diversity changes in the Iberian macromammalian faunas coincide with periods of increased faunal similarity with other regions, suggesting a relationship to the expansions and contractions of the geographical ranges of the constituent taxa. This pattern is not recognized for micromammals; that is, their diversity trends are not related to changes in geographical ranges. Main conclusions Climatic shifts result in expansions or contractions in the geographical ranges of macromammals, owing to changes in the distribution of their preferred habitats. The lower dispersal ability of micromammals results in a higher extinction risk when habitat fragmentation confines their populations to relatively small environmental patches. Hence, they are more severely affected by climatic changes. Our results thus emphasize the role of climatic forcing in mammalian biogeography and diversity.     

Harmonia axyridis invasions: Deducing evolutionary causes and consequences

I consider evolutionary approaches to deducing factors that have made the ladybird beetle Harmonia axyridis such a successful invader, and the contribution that studies of this species in its native range can make. Work aiming to demonstrate which (pre)adaptations have made the species so successful often fails to compare these putative characters with those of other ladybirds. This has led to a tendency for “argument by design”‐type claims on characters widely shared by non‐invasive coccinellids. There is good evidence from genetic studies that evolutionary change occurred in invasive populations, contributing to their success. There is some evidence for subsequent evolutionary change after the establishment of invasive H. axyridis, primarily in the native organisms with which the ladybird interacts. I show here that there appears to have been little adaptation in H. axyridis, over about 20 generations, to the alkaloids of one North American native intraguild prey, the ladybird Coleomegilla maculata. Studies of H. axyridis in its native range are important, as they provide a snapshot of the ancestral ladybird, unobscured by subsequent evolutionary change related to its invasiveness. They provide baseline data about phenomena such as interactions with natural enemies and intraguild predation, and they also can provide pointers as to how H. axyridis might further adapt in the regions it has colonized. Harmonia axyridis represents an ideal opportunity for greater international co‐operation between scientists studying this species in its native range in Asia and scientists studying it in Europe, America and Africa, where it is an invasive exotic.     

不同水分条件下春小麦种群中个体大小不整齐性及遗传学分析

种群内个体大小不整齐性是种群数量结构的主要指标。本文研究了不同水分条件下,３个品种春小麦种群个体大小不整齐性的建立及变化规律。对春小麦种群不整齐性的遗传学分析表明：遗传结构与随机环境修饰对种群数量结构形成的相对重要性,因水分条件不同而异。种群不整齐性在自然选择中的作用可用下列简单模型表示：ＧＳ０＝ＳＨ×ｈ２ＳＨＧＳ０：自然选择强度;ＳＨ：大小不整齐性;ｈ２ＳＨ：不整齐性的遗传力。     

Macroecology and the hierarchical expansion of evolutionary theory

The constraint envelope describing the relationship between geographical range size and body size has usually been explained by a minimum viable population size model, furnishing a strong argument for species selection if geographical range size turns out to be ‘heritable’. Recent papers have questioned this assumption of nonzero geographical range heritability at a phylogenetic level, meaning that the logic that constraint envelopes provide support for higher‐level selection fails. However, I believe that analysis of constraint envelopes can still furnish insights for the hierarchical expansion of evolutionary theory because the fitness furnished by variation in body size, which is frequently measured as a highly ‘heritable’ trait at the species level, can be partitioned into anagenetic and cladogenetic components. The constraint envelope furnishes an explicit mechanism for large‐body biased extinction rates influencing the distribution of body size. More importantly, it is possible to envisage a scenario in which anagenetic trends driving an increase in body size in higher latitudes within species (Bergmann's rule) are counteracted by available habitat area or continental edges constraining overall species distribution in these higher latitudes, increasing the probability of extinction. Under this combined model, faunas at higher latitudes and under habitat constraints may reach equilibrium points between these opposing hierarchical adaptive forces at smaller body size than faunas with less intense higher‐level constraints and will tend to be more right‐skewed.     

SEX‐SPECIFIC SELECTION AND INTRASPECIFIC VARIATION IN SEXUAL SIZE DIMORPHISM

Sexual size dimorphism (SSD) is thought to evolve due to sex differences in selection on body size, but it is largely unknown whether intraspecific variation in SSD reflects differences in sex‐specific selection among populations. We addressed this question by comparing viability selection between two island populations of the brown anole lizard (Anolis sagrei) that differ in the magnitude of male‐biased SSD. On both islands, females experienced stabilizing selection favoring intermediate size whereas males experienced directional selection favoring larger size. Thus, sex‐specific selection matched the overall pattern of male‐biased SSD, but population differences in the magnitude of SSD were not associated with local differences in selection. Rather, population differences in SSD appear to result from underlying differences in the environmental potential for a rapid growth, coupled with sex‐specific phenotypic plasticity. Males grew more slowly on the island with low SSD whereas growth of females did not differ between islands. Both sexes had substantially lower mass per unit length on the island with low SSD, suggesting that they were in a relatively poorer energetic condition. We propose that this energetic constraint disproportionately impacts growth of males due to their greater absolute energy requirements, thus driving intraspecific variation in SSD.     

浙江和江西二叠/三叠系界线层上下的辐鳍鱼类化石与鱼类的绝灭、复苏和辐射

记述了浙江长兴县煤山剖面、江西修水县四都乡东岭剖面和信丰县铁石口镇铁石口剖面二叠/三叠系界线层上下的辐鳍鱼类6个类别的微体化石,包含2新属2新种,它们是:赵氏浙江鱼(Zhejiangichthys zhaoi gen.et sp.nov.)和小齿葆青鱼(Baoqingichthys microdontus gen.et sp.nov.)。这是包括全球二叠/三叠系界线层型剖面和点位(GSSP)在内的全球二叠/三叠系界线层上下辐鳍鱼类微体化石序列的首次报道。在总结煤山剖面已记述的鱼类大化石和微体化石资料的基础上,分析了长兴煤山剖面二叠纪末鱼类的集群绝灭。绝灭发生得很晚,持续的时间很短,规模也很大,绝灭率高达93%。讨论了华南二叠/三叠系之交大绝灭后鱼类的复苏和辐射,认为鱼类和牙形类一样都是大绝灭后在三叠纪最早复苏的类别,最早复苏的鱼类为裂齿鱼类;鱼类从绝灭期到辐射期仅用了1.3 Ma到4 Ma,从地质时间考虑,大绝灭后鱼类的复苏和辐射是相当快的。华南早三叠世以裂齿鱼类的张氏鱼(Zhangina)和软骨鱼类的弓鲛(Hybodus)为代表的组合替代了晚二叠世以古鳕类的中华扁体鱼(sinoplatysomys)和软骨鱼类的中华尖齿鲨(Sinacrodus)为代表的组合。华南晚二叠世海相地层产出的辐鳍鱼类和软骨鱼类的一些土著属与产于特提斯区三叠纪的一些属非常相近,表明华南下扬子区很可能是后来繁盛于特提斯区的三叠纪鱼类的发源地。     

EPISTASIS,PLEIOTROPY, AND THE MUTATION LOAD IN SEXUAL AND ASEXUAL POPULATIONS

Mutation may impose a substantial load on populations, which varies according to the reproductive mode of organisms. Over the past years, various authors used adaptive landscape models to predict the long‐term effect of mutation on mean fitness; however, many of these studies assumed very weak mutation rates, so that at most one mutation segregates in the population. In this article, we derive several simple approximations (confirmed by simulations) for the mutation load at high mutation rate (U), using a general model that allows us to play with the number of selected traits (n), the degree of pleiotropy of mutations, and the shape of the fitness function (which affects the average sign and magnitude of epistasis among mutations). When mutations have strong fitness effects, the equilibrium fitness of sexuals and asexuals is close to ; under weaker mutational effects, sexuals reach a different regime where is a simple function of U and of a parameter describing the shape of the fitness function. Contrarily to weak mutation results showing that is an increasing function of population size and a decreasing function of n, these parameters may have opposite effects in sexual populations at high mutation rate.     

TEMPO AND MODE OF MULTICELLULAR ADAPTATION IN EXPERIMENTALLY EVOLVED SACCHAROMYCES CEREVISIAE

Multicellular complexity is a central topic in biology, but the evolutionary processes underlying its origin are difficult to study and remain poorly understood. Here we use experimental evolution to investigate the tempo and mode of multicellular adaptation during a de novo evolutionary transition to multicellularity. Multicelled “snowflake” yeast evolved from a unicellular ancestor after 7 days of selection for faster settling through liquid media. Over the next 220 days, snowflake yeast evolved to settle 44% more quickly. Throughout the experiment the clusters evolved faster settling by three distinct modes. The number of cells per cluster increased from a mean of 42 cells after 7 days of selection to 114 cells after 227 days. Between days 28 and 65, larger clusters evolved via a twofold increase in the mass of individual cells. By day 227, snowflake yeast evolved to form more hydrodynamic clusters that settle more quickly for their size than ancestral strains. The timing and nature of adaptation in our experiment suggests that costs associated with large cluster size favor novel multicellular adaptations, increasing organismal complexity.     

EVOLUTION AND EXTINCTION IN A CHANGING ENVIRONMENT: A QUANTITATIVE-GENETIC ANALYSIS

Because of the ubiquity of genetic variation for quantitative traits, virtually all populations have some capacity to respond evolutionarily to selective challenges. However, natural selection imposes demographic costs on a population, and if these costs are sufficiently large, the likelihood of extinction will be high. We consider how the mean time to extinction depends on selective pressures (rate and stochasticity of environmental change, and strength of selection), population parameters (carrying capacity, and reproductive capacity), and genetics (rate of polygenic mutation). We assume that in a randomly mating, finite population subject to density-dependent population growth, individual fitness is determined by a single quantitative-genetic character under Gaussian stabilizing selection with the optimum phenotype exhibiting directional change, or random fluctuations, or both. The quantitative trait is determined by a finite number of freely recombining, mutationally equivalent, additive loci. The dynamics of evolution and extinction are investigated, assuming that the population is initially under mutation-selection-drift balance. Under this model, in a directionally changing environment, the mean phenotype lags behind the optimum, but on the average evolves parallel to it. The magnitude of the lag determines the vulnerability to extinction. In finite populations, stochastic variation in the genetic variance can be quite pronounced, and bottlenecks in the genetic variance temporarily can impair the population's adaptive capacity enough to cause extinction when it would otherwise be unlikely in an effectively infinite population. We find that maximum sustainable rates of evolution or, equivalently, critical rates of environmental change, may be considerably less than 10% of a phenotypic standard deviation per generation.     

PATTERNS IN TREE BALANCE AMONG CLADISTIC,PHENETIC, AND RANDOMLY GENERATED PHYLOGENETIC TREES

I examine patterns in tree balance for a sample of 208 cladograms and phenograms from the recent literature. I provide an expression for expected imbalance under a simple, uniform-rate random speciation model, and I estimate variances by simulation for the same model. Imbalance decreases with tree size (number of included taxa) in both theoretical and literature trees. In contrast to previous suggestions, I find cladistic trees to be no more imbalanced than phenetic trees when confounding variables are appropriately controlled. The degree of imbalance found in literature trees is inconsistent with the uniform-rate speciation model; this is most likely a result of variability in speciation and extinction rates among real lineages. The existence of such variation is a necessary (but not sufficient) condition for the operation of the macroevolutionary processes of species sorting and species selection.     

A STUDY ON MINDARUS KOCH IN CHINA WITH DESCRIPTIONS OF NEW SPECIES AND NEW SUBSPECIES (HOMOPTERA: MIN-DARIDAE)*

Abstract This paper deals with the genus Mindam Koch from China. The phylogeny of the genus is discussed. M. keteleerifoliae is regarded as the most advanced species, M. obliquus and M. piceasuc-tus are more advanced than M. abietinus, M. japonicus and M. victoria. The origin centre of the genus may be in Europe and the differentiation centre of the genus may be in Hengduan Mountains Region of China. Geological distribution of the genus shows island-shaped pattern. Five species from China was described in this paper, including one new species, M. piceasuctus Zhang and Qiao, one new subspecies, M. abietinus triprimesensori Zhang and Qiao and one newly recorded species, M. abietinus abietinus Koch 1856. A key to the Chinese species of Mindarus is given. The specimens of five species from China are deposited in the Institute of Zoology, Academia Sinica.     

ADAPTIVE PHENOTYPIC PLASTICITY AND ITS DIFFERENCE BETWEEN UNIVOLTINE AND MULTIVOLTINE POPULATIONS IN A BRUCHID BEETLE,KYTORHINUS SHARPIANUS

The multivoltine bruchid Kytorhinus sharpianus shows seasonal phenotypic plasticity in adult longevity, the preoviposition period, and the number of eggs laid without feeding between the diapausing and nondiapausing generations. This study compared the norms of reaction in three life-history traits between the univoltine Aomori and multivoltine Mitsuma populations. The directions of response in the norms of reaction were similar in both populations, although their response curves differed between populations. This result indicated a potential for variation in seasonal phenotypic plasticity in the univoltine population. However, the variation in the norms of reaction was small in both populations, suggesting strong selection pressure on the plasticity in the multivoltine population. These results also suggest that the univoltine Aomori population may have originated from a multivoltine population.     

中国纩蚜属研究及新种新亚种记述(同翅目:纩蚜科)(英文)

通过对中国纩蚜属的研究,作者提出纩蚜属的现代地理分布呈岛状分布的格局;并推测该属的起源中心可能在欧洲,现代分化中心可能在中国的横断山脉地区。从动物地理的角度,探讨了该属内的系统发育关系。同时,发现一新种,即云杉纩蚜Mindaurspiceasuctusspnov.。正模为无翅孤雌蚜,副模为5只有翅孤雌蚜和8只无翅孤雌蚜,分布在云南省昆明市,辽宁省沈阳市和熊岳县。一新亚种,即冷杉纩蚜三圈亚种M.abietinustriprimesensorissp.nov.,正模为有翅孤雌蚜,副模为3只有翅孤雌蚜,分布在云南省丽江市玉龙山。一新纪录种,即冷杉纩蚜指名亚种M.abietinusabietinusKoch1856。并且给出种的检索表,模式标本存放在中国科学院动物研究所标本馆。     

BOOM AND BUST: ANCIENT AND RECENT DIVERSIFICATION IN BICHIRS (POLYPTERIDAE: ACTINOPTERYGII), A RELICTUAL LINEAGE OF RAY‐FINNED FISHES

Understanding the history that underlies patterns of species richness across the Tree of Life requires an investigation of the mechanisms that not only generate young species‐rich clades, but also those that maintain species‐poor lineages over long stretches of evolutionary time. However, diversification dynamics that underlie ancient species‐poor lineages are often hidden due to a lack of fossil evidence. Using information from the fossil record and time calibrated molecular phylogenies, we investigate the history of lineage diversification in Polypteridae, which is the sister lineage of all other ray‐finned fishes (Actinopterygii). Despite originating at least 390 million years (Myr) ago, molecular timetrees support a Neogene origin for the living polypterid species. Our analyses demonstrate polypterids are exceptionally species depauperate with a stem lineage duration that exceeds 380 million years (Ma) and is significantly longer than the stem lineage durations observed in other ray‐finned fish lineages. Analyses of the fossil record show an early Late Cretaceous (100.5–83.6 Ma) peak in polypterid genus richness, followed by 60 Ma of low richness. The Neogene species radiation and evidence for high‐diversity intervals in the geological past suggest a “boom and bust” pattern of diversification that contrasts with common perceptions of relative evolutionary stasis in so‐called “living fossils.”     

MODERN AND HISTORICAL EVIDENCE FOR NATURAL HYBRIDIZATION BETWEEN SYMPATRIC SPECIES IN MANDARINA (PULMONATA: CAMAENIDAE)

Geographical and temporal variation in gene exchange between two endemic land snail species, Mandarina aureola and Mandarina ponderosa, was studied on Hahajima Island of the Bonin Islands. Allozyme variation in modern samples, and variation in the color and shell morphology of modern and fossil samples, suggest a complex geographical and historical pattern of hybridization. These two species occur in sympatry, and their shell morphologies and protein genotypes are markedly divergent. However, many specimens of M. aureola, collected from the middle region of the island, exhibit intermediate shell morphologies and possess marker alleles of M. ponderosa. Fossil samples of the two species strongly suggest that these intermediates were hybrids with M. ponderosa that were produced since the end of the Pleistocene. Each of these species, in addition, is subdivided into two genetically and morphologically divergent parapatric races. Interspecific hybridization appears to have produced genetical and morphological admixture among these four distinctive groups of populations. The past distribution and geographic variation of M. ponderosa can be traced in the distribution of M. ponderosa-derived genotypes in current populations of M. aureola. Temporal changes of the color pattern in the fossil populations of hybrids suggest that the traits introduced from M. ponderosa to M. aureola have been affected by natural selection and could replace traits of living species when advantageous. Moreover, these introgressed genes appeared to provide novel properties that enabled M. aureola to advance into a new environment. Relatively independent change in shell color and morphology further suggests mosaic evolution following the hybridization events. Connectively, these data reveal how hybridization events may be an important source of evolutionary novelties and make it clear that the phenomenon of reticulate evolution cannot be ignored.     

Quantifying the effects of migration and mutation on adaptation and demography in spatially heterogeneous environments

How do mutation and gene flow influence population persistence, niche expansion and local adaptation in spatially heterogeneous environments? In this article, we analyse a demographic and evolutionary model of adaptation to an environment containing two habitats in equal frequencies, and we bridge the gap between different theoretical frameworks. Qualitatively, our model yields four qualitative types of outcomes: (i) global extinction of the population, (ii) adaptation to one habitat only, but also adaptation to both habitats with, (iii) specialized phenotypes or (iv) with generalized phenotypes, and we determine the conditions under which each equilibrium is reached. We derive new analytical approximations for the local densities and the distributions of traits in each habitat under a migration–selection–mutation balance, compute the equilibrium values of the means, variances and asymmetries of the local distributions of phenotypes, and contrast the effects of migration and mutation on the evolutionary outcome. We then check our analytical results by solving our model numerically, and also assess their robustness in the presence of demographic stochasticity. Although increased migration results in a decrease in local adaptation, mutation in our model does not influence the values of the local mean traits. Yet, both migration and mutation can have dramatic effects on population size and even lead to metapopulation extinction when selection is strong. Niche expansion, the ability for the population to adapt to both habitats, can also be prevented by small migration rates and a reduced evolutionary potential characterized by rare mutation events of small effects; however, niche expansion is otherwise the most likely outcome. Although our results are derived under the assumption of clonal reproduction, we finally show and discuss the links between our model and previous quantitative genetics models.