MITOCHONDRIAL-DNA DIVERSITY AND THE ORIGIN OF THE MENIDIA CLARKHUBBSI COMPLEX OF UNISEXUAL FISHES (ATHERINIDAE)

Restriction-endonuclease analysis of mitochondrial DNA (mtDNA) of the unisexual M. clarkhubbsi complex and close sexual relatives indicated that the unisexual complex arose through multiple, nonreciprocal hybridizations involving females of M. peninsulae. High-resolution analyses using restriction endonucleases that cleave at 4-bp sites revealed mtDNA sequence diversity that was low among unisexuals but high among individuals of M. peninsulae. The identification of M. peninsulae as a parent of the unisexuals conflicts with some details of previous allozyme comparisons. One possibility is that the unisexuals were derived from hybridization involving M. beryllina and a recently extinct form of M. peninsulae. In contrast to the unisexuals, contemporary hybrids of M. peninsulae and M. beryllina are formed by reciprocal matings. The origins of extant unisexual lineages from nonreciprocal hybridizations, together with their low mtDNA diversity relative to the maternal ancestor, implies strong constraints on origins of unisexuality via hybridization. Data on reproduction by contemporary F₁ hybrids reveal one form of genetic/developmental constraint: M. peninsulae and M. beryllina may now have diverged beyond the point where the hybrid origin of new unisexual lineages is possible.     

SEX RATIO SELECTION AND THE EVOLUTION OF ENVIRONMENTAL SEX DETERMINATION IN LABORATORY POPULATIONS OF MENIDIA MENIDIA

What happens when a population with environmental sex determination (ESD) experiences a change to an extreme environment that causes a highly unbalanced sex ratio? Theory predicts that frequency-dependent selection would increase the proportion of the minority sex and decrease the level of ESD in subsequent generations. We empirically modeled this process by maintaining five laboratory populations of a fish with temperature-dependent sex determination (the Atlantic silverside, Menidia menidia) in extreme constant temperature environments that caused highly skewed sex ratios to occur initially. Increases in the minority sex consistently occurred from one generation to the next across all five populations, first establishing and then maintaining a balanced sex ratio until termination of the experiment at 8 to 10 generations. The extent to which the level of ESD changed as balanced sex ratios evolved, however, was not consistent. Two populations that experienced high temperatures each generation displayed a loss of ESD, and in one of these ESD was virtually eliminated. This suggests that temperature-insensitive, sex-determining genes were being selected. In populations maintained in low temperature environments, however, the level of ESD did not decline. Instead, the response of sex ratio to temperature was adjusted upward or downward, perhaps by selection of sex-determining genes sensitive to higher (or lower) temperatures. The two different outcomes at low versus high temperatures occurred independent of the geographic origin of the founding population. Our results demonstrate that ESD is capable of evolving in response to selection.     

VARIATION IN ENVIRONMENTAL AND GENOTYPIC SEX-DETERMINING MECHANISMS ACROSS A LATITUDINAL GRADIENT IN THE FISH,MENIDIA MENIDIA

Models of environmental sex determination (ESD) usually assume that genetic influences on sex are polygenic, but the validity of this (or any other) form of genotype-environment interaction is virtually unknown. In the Atlantic silverside, Menidia menidia, sex is determined by an interaction between temperature and genotype and the response of sex ratio to temperature differs among populations from different latitudes. We examined the genetic basis of this pattern by measuring among family variation in the proportion of females, F/(F + M), within and among high (21°C) and low (15°C) temperatures for two populations: one from Nova Scotia (NS) where the level of ESD is low, and another from South Carolina (SC) where the level of ESD is high. In NS fish, temperature had a significant influence on sex ratio in only 1 of 23 families. The distribution of the fraction of females within temperatures for families from NS was highly heterogeneous and tended to fall into distinct classes (0.0, 0.25, 0.5, 1.0) like that expected from Mendelian segregation of a major sex factor(s). In contrast, temperature had a highly significant influence on sex ratio in all SC families examined (N = 24). Family sex ratios within temperatures were highly heterogeneous and, at least at 15°C, did not conform to simple Mendelian ratios. At 21°C, the proportion of females in most SC families was near zero and so the underlying sex tendencies of different families could not be discerned. Based on a previous study, mid-latitude fish appear to have an intermediate form of sex determination: simple Mendelian sex-ratio patterns exist and there is a moderate thermal influence on sex ratio in most but not all families. We suggest that sex determination in M. menidia is controlled by an interaction between major genetic factors, polygenic factors, and temperature and that the relative importance of each component differs with latitude. High latitude populations appear to have evolved a major sex-determining factor(s) that overrides the effect of temperature, and this factor(s) is lacking in low latitude populations.     

EVOLUTION OF INTRINSIC GROWTH AND ENERGY ACQUISITION RATES. I. TRADE-OFFS WITH SWIMMING PERFORMANCE IN MENIDIA MENIDIA

Abstract Latitudinal populations of the Atlantic silverside, Menidia menidia , show substantial genetic variation in rates of energy acquistion and allocation. Reared in common environments, silversides from northern latitudes consume more food, grow faster and more efficiently, store more energy, and produce greater quantities of eggs than their southern conspecifics. The persistence of seemingly inferior southern genotypes in the face of ostensibly superior northern genotypes suggest that there are hidden evolutionary trade-offs associated with these elevated acquisition and allocation rates. We tested the hypothesis that rapid growth and high levels of food consumption trade-off against locomotory performance in M. menidia . We compared both aerobic (prolonged and endurance) and anaerobic (burst) swimming capacities between intrinsically fast-growing fish from the north (Nova Scotia, NS) and intrinsically slow-growing fish from the south (South Carolina, SC) and between growth-manipulated phenotypes within each population. We also compared swimming speeds and endurance between fasted and recently fed fish within populations. Maximum prolonged and burst swimming speeds of NS fish were significantly lower than those of SC fish, and swimming speeds of fast-growing phenotypes were lower than those of slow-growing phenotypes within populations. Fed fish had lower burst speeds and less endurance than fasted fish from the same population. Thus, high rates of growth and the consumption of large meals clearly diminish swimming performance, which likely increases vulnerability to predation and decreases survival and relative fitness. The submaximal growth rate of southern M. menidia appears to be adaptive, resulting from balancing selection on rates of somatic growth.     

EVOLUTION OF INTRINSIC GROWTH AND ENERGY ACQUISITION RATES. II. TRADE-OFFS WITH VULNERABILITY TO PREDATION IN MENIDIA MENIDIA

Abstract The Atlantic silverside ( Menidia menidia ) exhibits countergradient latitudinal variation in somatic growth rate along the East Coast of North America. Larvae and juveniles from high-latitude populations display higher intrinsic rates of energy consumption and growth than genotypes from low-latitude populations. The existence of submaximal growth in some environments suggests that trade-offs must counter the oft-cited theoretical benefits of energy and growth maximization (e.g., "bigger is better,""faster is better") in the immature life stages. We hypothesized that energy and growth maximization trades off against investment in defense from predators. We conducted laboratory selection experiments to compare vulnerability to predation of silversides from: (1) fast-growing northern (Nova Scotia, NS) versus slow-growing southern (South Carolina, SC) source populations; (2) phenotypically manipulated fast-growing versus moderately-growing NS fish; and (3) recently fed versus unfed NS and SC fish. Tests involved fish drawn from common-garden environments and were conducted by subjecting mixed-treatment schools of size-matched silversides to natural, common piscine predators. NS silversides suffered significantly higher predation mortality than SC silversides. Parallel results were found in phenotypic manipulation of growth: NS silversides reared on a fast-growth trajectory (∼1.0 mm/day) were significantly more vulnerable to predation than those growing at a moderate rate (∼0.5 mm/day). Food consumption also affected vulnerability to predators: Silversides with large meals in their stomachs suffered significantly higher predation mortality than unfed silversides. Differences in predation vulnerability were likely due to swimming performance, not attractiveness to predators. Our findings demonstrate that maximization of energy intake and growth rate engenders fitness costs in the form of increased vulnerability to predation.     

鱼类线粒体DNA的遗传与进化

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EVOLUTION AND COEXISTENCE OF POLLINATION ECOTYPES IN AN AFRICAN GLADIOLUS (IRIDACEAE)

Pollinator‐mediated selection has been suggested as a key driver of speciation in plants. We examined the potential role of hawkmoth pollinators in driving allopatric divergence and maintaining sympatric coexistence of morphotypes in the African iris Gladiolus longicollis. Floral tube length in this species varies from 35 mm to 130 mm across its geographic range and reflects the prevailing tongue lengths of local hawkmoth assemblages. The distribution of floral tube lengths is bimodal with two relatively discrete categories—long (about 90 mm) or short (about 50 mm)—that match the bimodal distribution of hawkmoth tongue lengths in eastern South Africa. At a contact site between these two floral morphs, we found few individuals of intermediate length, suggesting limited gene flow between morphs despite their interfertility. A difference in flowering phenology appears to be the main isolating barrier between morphs at this site. Long‐ and short‐tubed morphs differed markedly in the chemical composition of their floral fragrance, a trait that could be used as a cue for morph‐specific foraging by hawkmoths. Positive directional selection on tube length was found to occur in both morphs.     

THE EVOLUTION OF INSECT-MEDIATED SELF-POLLINATION FROM AN OUTCROSSING SYSTEM IN CALYPTRIDIUM (PORTULACACEAE)

Calyptridium monospermum (Portulacaceae) comprises two biological races visited by different groups of insects. One race, pollinated by a bumblebee, Bombus vosnesenskii (Apidae), is outcrossing, while the other race is pollinated casually by a variety of other insects whose visits result in insect-mediated self-pollination and geitonogamy within the inflorescence. Insect-mediated selfing is also the mode of pollination in the closely related species, C. umbellatum. While all plants of both species are self-compatible, insect visitation is necessary for good seed set. Selfing populations of C. monospermum exhibit many floral adaptations for insect-mediated self-pollination which parallel those of the selfing C. umbellatum, while retaining vegetative characteristics typical of C. monospermum. These floral traits involve flower color and odor, style and stamen orientation, size of stigmatic surfaces, number of pollen grains produced, density of open flowers on inflorescences, and presence or absence of protogyny. Artifical pollination experiments show that these floral adaptations are effective in increasing the probability of insect-mediated self-pollination. Insect-mediated selfing in C. umbellatum and in some populations of C. monospermum is encouraged and probably made necessary by the inconstant pollinating behavior of the insects which visit these plants, as shown by the large percentage of non-Calyptridium pollen recovered from the bodies of insects captured while visiting these plants.     

亚洲土著类群硅藻鼠科的演化

硅藻鼠科是仅分布于亚洲的啮齿类类群,头骨具豪猪型咬肌结构,下颌则为松鼠型。该类群代表了一个从梳趾鼠类分化出的区域性支系,虽与豪猪次目相近,但并不是它的成员。最早的硅藻鼠化石记录发现于Baluchistan渐新世陆相地层中,稍晚的记录发现于南业晚渐新世。印度次大陆连续的地层为该科的演化历史提供了主要的化石依据。早中新世晚期至中中新世,硅藻科化石也发现了泰国、中国以及日本。其后的硅藻鼠种类少且鲜有化石记录。晚中新世早期在巴基斯坦有一个种,中国南部晚中新世也可能有一个种。尽管在最早的化石地点发现的标本较多,但其后的时段内除泰国李盆地外化石并不丰富,这指示了其特殊的生活习性或其生活区域化石保存的偏差。已知的化石记录并没有指示出该类群的多样性,也可能是缺少石化作用的结果。硅藻鼠类在水系外围区域延续生存与最近在老挝中部多岩石地带发现的现生Laonastes也是一个硅藻鼠的假设相一致。     

A CYTOLOGICAL CONSPECTUS OF THE GENUS CALYCADENIA (ASTERACEAE): AN EXAMPLE OF CONTRASTING MODES OF EVOLUTION

The chromosome numbers of 12 taxa of Calycadenia are documented by determinations in 77 widespread populations of the genus. Results of biosystematic studies indicate that hybrids among C. multiglandulosa, C. ciliosa, C. pauciflora, C. hispida, C. spicata, and C. oppositifolia characteristically have multiple associations of chromosomes producing complex meiotic configurations, whereas hybrids among C. villosa, C. mollis, C. truncata, and C. tenella generally yield few or no bivalents and a predominance of univalents during meiosis. Hybrids between C. multiglandulosa and C. hispida exhibit strict bivalent pairing and high pollen stainability. It is demonstrated that C. pauciflora and C. ciliosa comprise several sibling taxa that are morphologically cryptic, yet highly differentiated cytologically. Conversely, it is concluded that C. multiglandulosa and C. hispida comprise at least five morphologically distinct taxa that so far as known are highly interfertile (as judged by pollen stainability of hybrids) and cytologically uniform. Reproductive biology, chromosome mutation rates, habitats, flowering time, and genetic recombination mechanisms of the C. pauciflora-C. ciliosa and C. multiglandulosa-C. hispida complexes are compared in an attempt to account for the different modes of evolution observed in these closely related groups of Calycadenia. Lacking other plausible explanations, it is suggested that differential methods for limiting recombination of critical adaptive gene complexes might account for the divergent evolutionary patterns encountered in Calycadenia.     

EVOLUTIONARY INTERACTIONS BETWEEN SEXUAL AND ALL-FEMALE TAXA OF CYPRINOTUS (OSTRACODA: CYPRIDIDAE)

Freshwater ostracodes show both an exceptionally high incidence of transitions to unisexuality and, in some cases, an extraordinary level of clonal diversity. There is no understanding of the agents promoting these transitions to thelytoky, although it has been suggested that their frequency may set the stage for sexual taxa to infuse clonal diversity into unisexuals. This study examines the nature and origins of clonal diversity in the unisexual ostracode Cyprinotus incongruens. A combination of allozyme and cytogenetic studies revealed the presence of two diploid clones of this species at three temperate sites and ten clones at one arctic site including three diploids, five triploids, and two tetraploids. The low heterozygosity (0%–20%) of its diploid clones suggests that parthenogenesis has arisen spontaneously in C. incongruens rather than through hybridization, as in vertebrate asexuals. Polyploid clones appear to owe their origin to genome additions from sexual taxa, although subsequent mutational divergence has played a role in further enhancing diversity. Two triploid clones have apparently originated from the incorporation of a haploid genome from the sexually reproducing C. glaucus, as evidenced by their high heterozygosity and possession of alleles otherwise found only in that species. Other polyploid clones have likely arisen as a result of interbreeding between bisexual and unisexual C. incongruens. These results suggest that both the incidence of spontaneous transitions to clonality and the frequency of interbreeding with relatives may be the key processes that govern clonal diversity in unisexual ostracodes.