SPECIES RECOGNITION AND SEXUAL SELECTION AS A UNITARY PROBLEM IN ANIMAL COMMUNICATION

We investigated patterns of mating call preference and mating call recognition by examining phonotaxis of female túngara frogs, Physalaemus pustulosus, in response to conspecific and heterospecific calls. There are four results: females always prefer conspecific calls; most heterospecific calls do not elicit phonotaxis; some heterospecific calls do elicit phonotaxis and thus are effective mate recognition signals; and females prefer conspecific calls to which a component of a heterospecific call has been added to a normal conspecific call. We use these data to illustrate how concepts of species recognition and sexual selection can be understood in a unitary framework by comparing the distribution of signal traits to female preference functions.     

SEX DIFFERENCES IN MATE RECOGNITION AND CONSPECIFIC PREFERENCE IN SPECIES WITH MUTUAL MATE CHOICE

Sexual isolation is often assumed to arise because choosy females recognize and reject heterospecific males as mates. Yet in taxa in which both males and females are choosy, males might also recognize and reject heterospecific females. Here, we asked about the relative contribution of the sexes to the strong sexual isolation found in limnetic–benthic species pairs of threespine sticklebacks, which show mutual mate choice. We asked whether males and females of the two species recognize conspecifics and also prefer to mate with them. We found evidence for mate recognition by both sexes but only females prefer conspecifics. The nature of male courtship depended on which species of female they were courting, indicating that males recognized conspecific females and differentiated them from heterospecifics. However, males courted both species of females with equal vigor and changed courtship in a manner that would increase the chance of mating with heterospecifics. Females both recognized conspecifics and strongly preferred them. They responded very little to heterospecific male courtship and almost never mated with them. Therefore, males are likely to undermine sexual isolation, but females uphold it. Despite mutual mate choice and mate recognition in both sexes, females are primarily responsible for sexual isolation in these taxa.     

INTRASEXUAL COMPETITION ALONE FAVORS A SEXUALLY DIMORPHIC ORNAMENT IN THE RUBYSPOT DAMSELFLY HETAERINA AMERICANA

I studied the sex-limited red spots on the wings of male rubyspot damselflies (Hetaerina americana) in relation to territoriality and fitness in the wild. Both observational and experimental (wing spot manipulation) studies indicated that wing spots were selected through competition among males for mating territories, not through female choice or direct competition for females. Males with naturally or artificially large wing spots were more successful at holding territories and consequently mated at higher rates than males with relatively small wing spots. In contrast, sexual selection on male body size appeared to operate among nonterritorial males at the clasping stage of the mating sequence, perhaps because larger males were better at clasping females forcibly. Of four models proposed to explain the evolution of ornaments through territory competition, only the agonistic handicap model makes predictions consistent with the results of this study.     

VARIATION IN MATE CHOICE AND MATING PREFERENCES: A REVIEW OF CAUSES AND CONSEQUENCES

The aim of this review is to consider variation in mating p among females. We define mating p as the sensory and behavioural properties that influence the propensity of individuals to mate with certain phenotypes. Two properties of mating p can be distinguished: (i) ‘preference functions’–the order with which an individual ranks prospective mates and (2)‘choosiness’ -the effort an individual is prepared to invest in mate assessment. Patterns of mate choices can be altered by changing the costs of choosiness without altering the preference function. We discuss why it is important to study variation in female mating behaviour and identify five main areas of interest: Variation in mating p and costs of choosiness could (i) influence the rate and direction of evolution by sexual selection, (2) provide information about the evolutionary history of female p, (3) help explain inter-specific differences in the evolution of secondary sexual characteristics, (4) provide information about the level of benefits gained from mate choice, (5) provide information about the underlying mechanisms of mate choice. Variation in mate choice could be due to variability in preference functions, degree of choosiness, or both, and may arise due to genetic differences, developmental trajectories or proximate environmental factors. We review the evidence for genetic variation from genetic studies of heritability and also from data on the repeatability of mate-choice decisions (which can provide information about the upper limits to heritability). There can be problems in interpreting patterns of mate choice in terms of variation in mating p and we illustrate two main points. First, some factors can lead to mate choice patterns that mimic heritable variation in p and secondly other factors may obscure heritable p. These factors are divided into three overlapping classes, environmental, social and the effect of the female phenotype. The environmental factors discussed include predation risk and the costs of sampling; the social factors discussed include the effect of male–male interactions as well as female competition. We review the literature which presents data on how females sample males and discuss the number of cues females use. We conclude that sexual-selection studies have paid far less attention to variation among females than to variation among males, and that there is still much to learn about how females choose males and why different females make different choices. We suggest a number of possible lines for future research.     

THE GEOGRAPHY OF MITOCHONDRIAL DNA VARIATION,POPULATION STRUCTURE,HYBRIDIZATION, AND SPECIES LIMITS IN THE FOX SPARROW (PASSERELLA ILIACA)

Geographic variation in mitochondrial DNA (mtDNA) restriction sites was studied in the fox sparrow (Passerella iliaca). Seventy-eight haplotypes were found. Haplotypes fall into four phylogeographic groups that correspond to groups defined by plumage characters. The geographic distribution of these four groups does not appear congruent with mtDNA patterns in other vertebrates. Within each group, there is little geographic variation in mtDNA restriction sites, although there is geographic variation in plumage coloration and body size. The evolution of mtDNA diversity in fox sparrows seems best explained by vicariant events rather than isolation by distance. The mtDNA evidence suggests that Passerella megarhyncha and Passerella schistacea, two nonsister taxa that occur in western North America, have independently undergone bottlenecks. Hybridization is limited between all pairs of taxa except P. megarhyncha and P. schistacea, where mtDNA evidence suggests a narrow contact zone along the interface of the Great Basin and Sierra Nevada/Cascades. Morphometric characters intergrade over a broader area, suggesting that different processes are responsible for the two gradients. The occurrence of limited backcrossing among taxa suggests that cytoplasmic-nuclear incompatibility is lacking. The number of biological species would range from one to four, depending on the degree of hybridization tolerated. The mtDNA and plumage characters suggest four phylogenetic species: P. iliaca, P. megarhyncha, P. unalaschcensis, and P. schistacea.     

SPECIES ISOLATION,GENITAL MECHANICS,AND THE EVOLUTION OF SPECIES-SPECIFIC GENITALIA IN THREE SPECIES OF MACRODACTYLUS BEETLES (COLEOPTERA,SCARABEIDAE, MELOLONTHINAE)

The question asked was why male genitalic structures have diverged in three syntopic species of Macrodactylus beetles. Four hypotheses were evaluated: 1. The ways in which male genitalia mesh with internal female structures indicate that selection for species isolation via mechanical exclusion (“lock and key”) is unlikely to explain the genitalic differences. 2. The specific mate recognition hypothesis also clearly fails to explain genitalic differences due to the implausibility of postulated environmental effects on genitalia, and lack of postulated coevolution of male and female morphologies. 3. Selection for species isolation via differences in genitalic stimulation (sensory lock and key) is unlikely due to relatively infrequent cross-specific pair formation and intromission in the field, and “excessive” numbers of species-specific genitalic structures and male courtship behavior patterns which nevertheless occasionally fail. It also fails to explain the frequent failure of intraspecific copulations to result in sperm transfer. This hypothesis cannot, however, be rejected as confidently as the previous hypotheses. 4. Conditions under which sexual selection by cryptic female choice could take place are common. Females frequently exercise their ability to prevent sperm transfer by conspecific males even after intromission has occurred, and females generally mate repeatedly, probably with different males. Males behave as if cryptic female choice is occurring, courting assiduously while their genitalia are within the female. Sexual selection by female choice could thus contribute to the divergence in genitalic structures.     

CULTURAL INHERITANCE OF SONG AND ITS ROLE IN THE EVOLUTION OF DARWIN'S FINCHES

Songs of Darwin's finches were studied on the Galápagos Island of Daphne Major from 1976 to 1995. A single, structurally simple, and unvarying song is sung throughout life by each male of the two common species, Geospiza fortis (medium ground finch) and G. scandens (cactus finch). Songs of the two species differ strongly in quantitative features, and individual variation among males is much broader in G. fortis than in G. scandens. Although there are exceptions, songs of sons strongly resemble the songs of their fathers. They also resemble the songs of their paternal grandfathers, but not their maternal grandfathers, indicating that they are culturally inherited and not genetically inherited. Female G. fortis display a tendency to avoid mating with males that sing the same type of song as their father. They also avoid mating with males that sing heterospecific song, with very rare exceptions. Thus song, an evolving, culturally inherited trait, is an important factor in species recognition and mate choice. It constrains the mating of females to conspecifics, even when there is no genetic penalty to interbreeding, and thus may play a crucial role in species formation by promoting genetic isolation on secondary contact. The barrier is leaky in that occasional errors in song transmission result in misimprinting, which leads to a low incidence of hybridization and introgression. Introgression slows the rate of postzygotic isolation, but can produce individuals in novel genetic and morphological space that can provide the starting point of a new evolutionary trajectory.     

中国红■属研究及新种记述(蜻蜓目,■科)

系统记述了红■属Pyrrhosoma Charpentier的研究历史,明确了本属的属征,描述了中国的种类,并确定一个采自中国云南省的新种Pyrrhosoma latiloba sp.nov.,模式标本保存于南开大学生命科学学院昆虫学研究所昆虫标本室。     

SIMULATING RANGE EXPANSION: MALE SPECIES RECOGNITION AND LOSS OF PREMATING ISOLATION IN DAMSELFLIES

Prolonged periods of allopatry might result in loss of the ability to discriminate against other formerly sympatric species, and can lead to heterospecific matings and hybridization upon secondary contact. Loss of premating isolation during prolonged allopatry can operate in the opposite direction of reinforcement, but has until now been little explored. We investigated how premating isolation between two closely related damselfly species, Calopteryx splendens and C. virgo , might be affected by the expected future northward range expansion of C. splendens into the allopatric zone of C. virgo in northern Scandinavia. We simulated the expected secondary contact by presenting C. splendens females to C. virgo males in the northern allopatric populations in Finland. Premating isolation toward C. splendens in northern allopatric populations was compared to sympatric populations in southern Finland and southern Sweden. Male courtship responses of C. virgo toward conspecific females showed limited geographic variation, however, courtship attempts toward heterospecific C. splendens females increased significantly from sympatry to allopatry. Our results suggest that allopatric C. virgo males have partly lost their ability to discriminate against heterospecific females. Reduced premating isolation in allopatry might lead to increased heterospecific matings between taxa that are currently expanding and shifting their ranges in response to climate change.     

PHYLOGENY, REPRODUCTIVE ISOLATION AND KIN RECOGNITION IN THE SOCIAL AMOEBA DICTYOSTELIUM PURPUREUM

Little is known about the population structure of social microorganisms, yet such studies are particularly interesting for the ways that genetic variation impacts their social evolution. Dictyostelium , a eukaryotic microbe widely used as a developmental model, has a social fruiting stage in which some formerly independent individuals die to help others. To assess genetic variation within the social amoeba Dictyostelium purpureum , we sequenced ∼4000 base pairs of ribosomal DNA (rDNA) from 37 isolates collected in Texas, Virginia, and Japan. Our analysis showed extensive genetic variation between populations and clear evidence of phylogenetic structure. We identified three major phylogenetic groups that were more different than other accepted species pairs. Tests using pairs of clones showed that both sexual macrocyst and asexual fruiting body formation were influenced by genetic divergence. Macrocysts were less likely to form between pairs of clones from different groups than from the same group. There was also a correlation between the genetic divergence of a pair of clones and their degree of mixing within fruiting bodies. These observations suggest that cryptic species might occur within D. purpureum and, more importantly, reveal how genetic variation impacts social interactions.     

中国云南奈春蜓属一新种(蜻蜓目,春蜓科)

记述奈春蜓属1新种,二齿奈春蜓Nychogomphus bidentatus sp.nov.,模式标本存放在云南大理学院生命科学与化学学院。     

FIVE GENERATIONS OF ENFORCED SELFING AND OUTCROSSING IN MIMULUS GUTTATUS: INBREEDING DEPRESSION VARIATION AT THE POPULATION AND FAMILY LEVEL

The focus of this study was to examine the consequences of five sequential generations of enforced selfing and outcrossing in two annual populations of the mixed-mating Mimulus guttatus. Our primary goal was to determine whether purging of deleterious recessive alleles occurs uniformly between populations and among families, and thus gain insights into the mode of gene action (dominance, overdominance, and/or epistasis) governing the expression of inbreeding depression at both the population and family levels across the life cycle.     

中国黑额蜓属一新种记述(蜻蜓目,蜓科)(英文)

记述黑额蜓属1新种,刘氏黑额蜓Planaeschna liui sp.nov..新种区别于黑额蜓属其它种的主要特征如下:1)上唇中基部具1对黄绿色斑,上颚基部具1黄绿色小斑;2)腹部第9节背面端半部黄褐色,侧面端下部土黄色,腹部第10节背面具1黄褐色大斑,侧面下部土黄色;3)上肛附器在中部扭曲,下肛附器长度超过上肛附器一半,其基端宽度与腹部第10节高度相等;4)阳茎末节近方形,前端稍平截,基部两侧具1对稍向前弯曲的大耳状突.正模♂,福建武夷山,2008-07-16,邱志鹏采.模式标本存放在福建农林大学作物病虫生物防治研究所.     

MEASURING SELECTION ON A POPULATION OF DAMSELFLIES WITH A MANIPULATED PHENOTYPE

The estimation of the relationship between phenotype and fitness in natural populations is constrained by the distribution of phenotypes available for selection to act on. Because selection is blind to the underlying genotype, a more variable phenotypic distribution created by using environmental effects can be used to enhance the power of a selection study. I measured selection on a population of adult damselflies (Enallagma boreale) whose phenotype had been modified by raising the larvae under various levels of food availability and density. Selection on body size (combination of skeletal and mass at emergence) and date of emergence was estimated in two consecutive episodes. The first episode was survival from emergence to sexual maturity and the second was reproductive success after attaining sexual maturity. Female survival to sexual maturity was lower, and therefore opportunity for selection greater, than males in both years. Opportunity for selection due to reproductive success was greater for males. The total opportunity for selection was greater for males one year and for females the other. Survival to sexual maturity was related to mass gain between emergence and sexual maturity. Females gained more mass and survived less well than males in both years but there was no linear relationship between size at emergence and survival for females in either year. However, females in the tails of the phenotype distribution were less likely to survive than those near the mean. In contrast, small males consistently gained more mass than large males and survived less well in one year. There was significant selection on timing of emergence in both years, but the direction of selection changed due to differences in weather; early emerging females were more successful one year and late emerging males and females the other. The number of clutches laid by females was independent of body size. Because the resources used to produce eggs are acquired after emergence and this was independent of size at emergence, female fitness did not increase with size. Small males may have had lower survival to sexual maturity but they had higher mating success than large males. Resources acquired prior to sexual maturity are essential for reproductive success and may in some species alter their success in inter- and intrasexual competition. Therefore, ignoring the mortality associated with resource acquisition will give an incomplete and potentially misleading picture of selection on the phenotype.     

HYBRIDIZATION AND CHLOROPLAST DNA VARIATION IN A PINUS SPECIES COMPLEX FROM ASIA

Heterologous hybridization of chloroplast DNA (cpDNA) involving 30 endonucleaseprobe combinations was used to analyze cpDNA variation in multiple individuals and populations of Pinus tabulaeformis (Carr.), Pinus yunnanensis (Franchèt) and Pinus massoniana (Lamb.). Restriction fragment patterns detected by several combinations distinguished among the three species. The obtained cpDNA markers were subsequently used to examine cpDNA variation of Pinus densata (Masters), a putative tertiary hybrid between P. tabulaeformis and P. yunnanensis. The analysis demonstrated that P. densata populations harbor three different haplotypes. Two of these haplotypes are characteristic of P. tabulaeformis and P. yunnanensis. However, the third haplotype found in P. densata appears to be absent in other extant Asian Pinus species. It is suggested that the observed cpDNA composition of P. densata populations is a result of past hybridization involving P. tabulaeformis, P. yunnanensis, and a third unknown or extinct taxon. Chloroplast DNA polymorphism in P. densata was much greater than that for nuclear allozyme markers in this and the other Pinus species. Population differentiation was also substantial in P. densata and exceeded that for allozyme markers. In contrast, no cpDNA polymorphism was detected in populations of P. tabulaeformis, P. yunnanensis, and P. massoniana. The study suggests that interspecific gene exchange may lead to the creation of stable cpDNA polymorphism in conifer hybrids.     

肉苁蓉(Cistanche deserticola)种内的变异式样

用比较形态解剖学的方法对肉苁蓉的营养器官和繁殖器官进行比较,研究了名贵中药肉苁蓉营养器官和繁殖器官的变异式样。发现肉苁蓉种内具有不同形态特点的类型,造成中药肉苁蓉中在其药理、药效等方面也存在着差异。     

MUTATION,SELECTION, AND THE MAINTENANCE OF LIFE-HISTORY VARIATION IN A NATURAL POPULATION

In an effort to provide insight into the role of mutation in the maintenance of genetic variance for life-history traits, we accumulated spontaneous mutations in 10 sets of clonal replicates of Daphnia pulex for approximately 30 generations and compared the variance generated by mutation with the standing level of variation in the wild population. Mutations for quantitative traits appear to arise at a fairly high rate in this species, on the order of at least 0.6 per character per generation, but have relatively small heterozygous effects, changing the phenotype by less than 2.5% of the mean. The mean persistence time of a new mutation affecting life-history/body-size traits is approximately 40 generations in the natural population, which requires an average selection coefficient against new mutations of approximately 3% in the heterozygous state. These data are consistent with the idea that the vast majority of standing genetic variance for life-history characters may be largely a consequence of the recurrent introduction of transient cohorts of mutations that are at least conditionally deleterious and raise issues about the meaning of conventional measures of standing levels of variation for fitness-related traits.     

REPRODUCTIVE ISOLATION AND DISTINCT POPULATION STRUCTURES IN TWO TYPES OF THE FRESHWATER SHRIMP PALAEMON PAUCIDENS

Twenty local populations of the Japanese freshwater shrimp Palaemon paucidens were electrophoretically and morphologically surveyed. Based on the diagnostic distributions of some alleles at Gpi, Mpi, Mdh-1, and Mdh-2, these populations were largely classified into two types (A and B). The A type occurred in lakes, ponds, and rivers, while the B type was observed only in rivers. Average Nei's genetic distance (D) between the two types fell into the subspecies range . The coefficient of gene differentiation, G_ST, varied considerably between the two types. In 12 populations of the A type, with a G_ST value of 0.281, nine pond and lake populations showed a higher G_ST (0.246) than the three river populations (0.151). On the other hand, G_ST was 0.036 for the eight local populations of the B type. The lower rostrum tooth number had a mode of two in type A and three in type B. Type-A populations largely varied in the upper rostrum tooth number and egg size but type B did not. Under laboratory conditions, mating frequently occurred within each type, but not between types. Furthermore, no embryonic development was observed in the few cases of intertype mating. These results indicate that the A and B types had experienced cladogenic separation with pre- or postmating isolation, whereafter the A type, under geographic isolation, underwent genetic and phenotypic differentiation, while the B type, under extensive gene flow, did not undergo differentiation.     

紊蒿属一新种和对该属分类及演化的讨论

对新种多头紊蒿(Elachanthemum polycephalum Z.Y.Zhu et C.Z.Liang)的形态特征进行了描述,并从形态和生态特征、地理分布及其区系起源等方面探讨了紊蒿属(Elachanthemum Y.Ling et Y.R.Ling)的分类地位。结果表明：紊蒿属与百花蒿属(Stilpnolepis Krasch.)在形态上虽有相近之处,但花、果实和花粉等形态差异明显,因而不能并入百花蒿属,而应独立成属。该属起源于第三纪亚洲北部的古蒿类群(Pro-Artemisia L.),是蒿类中较为原始的属,为古地中海东部残遗的旱生成分,也是现代亚洲中部荒漠(戈壁荒漠)的持有属;多头紊蒿新种的发现,使紊蒿属从单种属变成寡种(双种)属,表明荒漠植物区系的物种分化仍在进行。