INTERSPECIFIC AND INTRASPECIFIC MORPHOLOGICAL VARIATION IN MANATEES (SIRENIA: TRICHECHUS)

The three living species of Trichechus are clearly defined and well exemplify the degree of variability and taxonomic value of morphological characters in a well-understood mammalian genus. Statistical analysis of the largest sample of manatee skulls yet studied has allowed us to identify small suites of characters that effectively distinguish these species. The two subspecies of T. manatus proposed by Hatt (1934) can likewise be distinguished, and their use as taxonomic categories seems justified. This suggests that the cool winters of the northern Gulf Coast, on the one hand, and the deep water and strong currents of the Straits of Florida, on the other, are effective barriers to gene flow between Florida and Antillean manatees. Alleged taxonomic distinctions within T. senegalensis , however, have no demonstrated basis. No significant sexual dimorphism was detected in skulls of any of the species. Many interspecific differences can be correlated with feeding ecology, but others remain unexplained. Features of tooth crown morphology are among the most constant characters examined, but some osteological characters are equally good. T. manatas and T. senegalensis (which are phenetically the most similar) also seem to share a more recent common ancestor than either does with T. inunguis. However, the three species probably separated from each other at nearly the same time. T. inunguis has since become the most derived species, while T. senegalensis has changed the least.     

Phylogeography, phylogeny and hybridization in trichechid sirenians: implications for manatee conservation

Abstract The three living species of manatees, West Indian (Trichechus manatus), Amazonian (Trichechus inunguis) and West African (Trichechus senegalensis), are distributed across the shallow tropical and subtropical waters of America and the western coast of Africa. We have sequenced the mitochondrial DNA control region in 330 Trichechus to compare their phylogeographic patterns. In T. manatus we observed a marked population structure with the identification of three haplotype clusters showing a distinct spatial distribution. A geographic barrier represented by the continuity of the Lesser Antilles to Trinidad Island, near the mouth of the Orinoco River in Venezuela, appears to have restricted the gene flow historically in T. manatus. However, for T. inunguis we observed a single expanding population cluster, with a high diversity of very closely related haplotypes. A marked geographic population structure is likely present in T. senegalensis with at least two distinct clusters. Phylogenetic analyses with the mtDNA cytochrome b gene suggest a clade of the marine Trichechus species, with T. inunguis as the most basal trichechid. This is in agreement with previous morphological analyses. Mitochondrial DNA, autosomal microsatellites and cytogenetic analyses revealed the presence of hybrids between the T. manatus and T. inunguis species at the mouth of the Amazon River in Brazil, extending to the Guyanas and probably as far as the mouth of the Orinoco River. Future conservation strategies should consider the distinct population structure of manatee species, as well as the historical barriers to gene flow and the likely occurrence of interspecific hybridization.     

LONG-TERM TRENDS IN ABUNDANCE AND DISTRIBUTION OF MANATEES (TRICHECHUS MANATUS) IN THE NORTHERN BANANA RIVER, BREVARD COUNTY, FLORIDA

Four aerial survey projects were conducted between 1977 and 1986 to determine the abundance, density and distribution of West Indian manatees (Trichechus manatus), in the northern Banana River, Kennedy Space Center, Florida. Manatee density and distribution within selected portions of the 78.5 km² study area were determined. Peak numbers of manatees occurred in spring of each year. The maximum counts increased from 56 in 1978 to 297 in 1986. Manatee abundance was lowest in the winter of each year. Mean density per flight increased from 0.52 manatees/km2 in 1977–78 to 2.73/km² in 1984–86. This increase may reflect increases in the east coast population or shifts in the population distribution. Distributional changes were observed in the study area through time, with a lower percentage of manatees occurring in industrial areas and a correspondingly higher percentage of manatees in nonindustrial areas by 1985.     

Palaeoecological and morphofunctional interpretation of bone mass increase: an example in Late Cretaceous shallow marine squamates

Bone mass increase (BMI; i.e. osteosclerosis with possible additional pachyostosis) is characteristically displayed by many Late Cretaceous squamates that adapted to shallow marine environments—plesiopelvic mosasauroids, stem‐ophidians and pachyophiids. A combined morphological and microanatomical analysis of vertebrae and, to a lesser extent, ribs of these fossil squamates provides new data about the distribution and variability of this osseous specialization in these taxa. Classical thin sections and third generation synchrotron microtomography and laminography were used for the microanatomical analysis. Following the explanation of the likely involvement of this specialization in the control of buoyancy, body trim and Carrier's constraint, new palaeoecological inferences and new hypotheses about the locomotor abilities and life environment of these organisms are produced. The taxa displaying BMI are considered to have undertaken long dives, hovering slowly and maintaining a horizontal trim, in shallow and protected water environments. Conversely, marine stem‐ophidians deprived of this specialization are regarded as slow surface swimmers able to live in more open marine environments. This study highlights the importance of microanatomical data for palaeoecological studies. It also discusses the significance of the use of this specialization as a character in phylogenetic studies.     

QUANTITATIVE GENETICS OF SHAPE IN CRICKET WINGS: DEVELOPMENTAL INTEGRATION IN A FUNCTIONAL STRUCTURE

The role of developmental and genetic integration for evolution is contentious. One hypothesis states that integration acts as a constraint on evolution, whereas an alternative is that developmental and genetic systems evolve to match the functional modularity of organisms. This study examined a morphological structure, the cricket wing, where developmental and functional modules are discordant, making it possible to distinguish the two alternatives. Wing shape was characterized with geometric morphometrics, quantitative genetic information was extracted using a full‐sibling breeding design, and patterns of developmental integration were inferred from fluctuating asymmetry of wing shape. The patterns of genetic, phenotypic, and developmental integration were clearly similar, but not identical. Heritabilities for different shape variables varied widely, but no shape variables were devoid of genetic variation. Simulated selection for specific shape changes produced predicted responses with marked deflections due to the genetic covariance structure. Three hypotheses of modularity according to the wing structures involved in sound production were inconsistent with the genetic, phenotypic, or developmental covariance structure. Instead, there appears to be strong integration throughout the wing. The hypothesis that genetic and developmental integration evolve to match functional modularity can therefore be rejected for this example.     

COMPARATIVE METHODS FOR THE ANALYSIS OF CONTINUOUS VARIABLES: GEOMETRIC INTERPRETATIONS

Abstract This study is concerned with statistical methods used for the analysis of comparative data (in which observations are not expected to be independent because they are sampled across phylogenetically related species). The phylogenetically independent contrasts (PIC), phylogenetic generalized least‐squares (PGLS), and phylogenetic autocorrelation (PA) methods are compared. Although the independent contrasts are not orthogonal, they are independent if the data conform to the Brownian motion model of evolution on which they are based. It is shown that uncentered correlations and regressions through the origin using the PIC method are identical to those obtained using PGLS with an intercept included in the model. The PIC method is a special case of PGLS. Corrected standard errors are given for estimates of the ancestral states based on the PGLS approach. The treatment of trees with hard polytomies is discussed and is shown to be an algorithmic rather than a statistical problem. Some of the relationships among the methods are shown graphically using the multivariate space in which variables are represented as vectors with respect to OTUs used as coordinate axes. The maximum‐likelihood estimate of the autoregressive parameter, ρ, has not been computed correctly in previous studies (an appendix with MATLAB code provides a corrected algorithm). The importance of the eigenvalues and eigenvectors of the connection matrix, W, for the distribution of ρ is discussed. The PA method is shown to have several problems that limit its usefulness in comparative studies. Although the PA method is a generalized least‐squares procedure, it cannot be made equivalent to the PGLS method using a phylogenetic model.     

QUANTITATIVE GENETICS OF RESPONSE TO COMPETITORS IN NEMOPHILA MENZIESII: A GREENHOUSE STUDY

To determine the potential for adaptation to a local biotic environment, we examined the magnitude and nature of genetic variation in response to neighboring plants within a natural population of the native California annual, Nemophila menziesii. A total of 22 plants from a natural population were crossed in three reciprocal factorials. The progeny were grown in a greenhouse in nine treatments that varied in conspecific density and in the density of a naturally co-occurring grass species, Bromus diandrus. Increasing the density of each species significantly reduced individual survival, fruit number, and dry weight. Among survivors, we found small to moderate heritability of dry weight within treatments. Additive genetic correlations (r_A) of dry weight between competitive regimes were generally large and positive. In no case were they significantly different from 1, as expected under the null hypothesis that the relative performance of the genotypes under consideration is the same in all environments. On the basis of these results, we cannot conclude that the structure of genetic covariation within this population would promote genetic differentiation in response to locally varying conditions of density of these two species. Aspects of the experiment that may have compromised our ability to detect r_A differing from 1 are discussed.     

晋西北黄土区人工林土壤水分动态的定量研究

本文根据１９８８－１９９０年在定位观测资料,对晋西北黄土丘陵区河北杨林、刺槐林和柠条藻木林的土壤水分动态规律进行了定量分析。依次就水分分布分层、水分剖面特征、水分季节动态、水分循环模式与水分循环水分等问题进行了探讨。本文采用的新的分析途径和方法将有助于对黄土区不同植被类型土壤水分生态研究的进一步深化。     

中国-中新世湖泊的古生态:根据山旺盆地植硅体的初步解释

山东山旺中新世山旺组湖相沉积中富含保存精美的古生物化石。对于湖相沉积中的昆虫、植物和哺乳动物化石,前人已经做了大量的研究工作。其中植物大化石和孢粉的研究结果表明,中新世山旺组在沉积期间,山旺湖周围的植被处于湿润的暖温带到亚热带的古生态环境下。哺乳动物蝙蝠和貘化石的研究结果也支持这个结论。然而,迄今为止,还没有关于草本植物的种类组成及含量的研究报道。本文首次对山旺组样品中的植硅体进行了研究,为山旺中新世植被的研究提供了新的证据。对从6块山旺组样品中分析出的植硅体进行研究的结果表明,山旺中新世植硅体组合含有丰富的保存极好的草本植物植硅体,代表森林成分的双子叶植物植硅体,以及偶尔出现的棕榈科植硅体。草本植物的植硅体主要来自于C_3/C_4禾本科的PACCAD类群和早熟禾亚科的植物,少数可能来自于在林下封闭生活环境中生长的草本植物。山旺中新世植硅体组合显示,在湖边森林生活环境下,伴生大量的适合沼生到中生环境的草本植物,在较干旱的地区生长着早熟禾(和PACCAD)的草本植物。本文的研究结果进一步支持山旺中新世气候比现在更加湿润而且变动较小的观点。山旺中新世植硅体组合与土耳其和北美大平原中新世湖相沉积中的植硅体组合不同,后者的植硅体组合主要是由多种C_3早熟禾亚科草本和不同种类的PACCAD类群草本植物组成。今后的研究工作将进一步揭示这种差异是否反映了草本植物群落在大尺度的生物地理上的差异,或者是由于地区性的局部气候不同所导致。     

DATA AND DATA INTERPRETATION IN THE STUDY OF LIMB EVOLUTION: A REPLY TO GALIS ET AL. ON THE REEVOLUTION OF DIGITS IN THE LIZARD GENUS BACHIA

Galis and collaborators (2010) claim that our recent paper ( Kohlsdorf and Wagner 2006 ), presenting statistical evidence for the reevolution of digits in the genus Bachia, may be flawed. Their reanalysis of the data does not support the possibility of a reevolution of digits and the authors also argue that such a reevolution would be implausible on functional and developmental grounds. In response, we reanalyzed our data with additional outgroup species. Our results differ from the one published in 2006, but this incongruence is not statistically significant. In contrast, the hypothesis presented by Galis et al. is significantly worse. An analysis of digit number evolution, using novel techniques to test for irreversibility of character loss ( Goldberg and Igic 2008 ), confirmed our original conclusion that there is strong evidence for reevolution of digits in Bachia. We also point out that this result is not in conflict with the hypothesis by Galis and Metz (2001) that mutations affecting the initial digit patterning are associated with strong negative pleiotropic effects and thus unlikely to be fixed in evolution. An important avenue of future research will be to directly test whether reevolved digits develop from conserved digit condensations retained after digit loss.     

正常小鼠骨髓血红蛋白含量的测定

小鼠骨髓血红蛋白含量的变化可以间接地反映骨髓微循环系统形态和功能的状况。按Burger and Knyszynski(1969)方法操作繁复,限制了它的推广应用及正常值的问世。最近,我们建立的简易测定方法,为成批标本的测定和正常值的确定创造了条件。 正常小鼠骨髓血红蛋白含量测定的目的:1)在较大量标本的测定中进一步验证该方法的可靠性;2)确定青、成年小鼠骨髓血红蛋白的正常值范围;3)分析其可能的影响因素,以便更好地控制实验条件和判断骨髓微循环障碍的程度。     

SKIN DENSITY AND ITS INFLUENCE ON BUOYANCY IN THE MANATEE (TRICHECHUS MANATUS LATIROSTRIS), HARBOR PORPOISE (PHOCOENA PHOCOENA), AND BOTTLENOSE DOLPHIN (TURSIOPS TRUNCATUS)

This study investigated how skin contributes to buoyancy control in the Florida manatee ( Trichechus manatus latirostris ), harbor porpoise ( Phocoena phocoena ), and bottlenose dolphin ( Tursiops truncatus ). Manatees are shallow divers and control their position in the water column hydrostatically. The two cetaceans are relatively deep divers that control their buoyancy hydrodynamically. Although the cetacean skin had been hypothesized to lower total body density ( e. g. , Dearolf et al. 2000, Nowacek et al. 2001), its buoyant force had not been calculated. The density of manatee skin, and its contribution to buoyancy, was unknown. Skin densities of 27 manatees, five harbor porpoises, and five bottlenose dolphins were measured volumetrically. Skin mass and density were used to calculate buoyant force. Harbor porpoise (952 kg/m³) and bottlenose dolphin (969 kg/m³) skins were less dense than seawater, and added 9 and 25 N of positive buoyant force, respectively, to total body buoyancy. By contrast, manatee skin (1,121 kg/m³) contributed 56 N of negative buoyant force, which equaled 70% of the negative buoyant force of their dense, pachyosteosclerotic ribs. Calculation of buoyant forces of the skeleton, skin and lungs demonstrates that the manatee is positively buoyant at the surface and negatively buoyant at depths of less than 10 m.     

大鼠中枢神经系统衰老的形态学研究Ⅳ.大脑皮质锥体细胞树突棘的数量变化

用6、12与31个月的雄性Wistar大鼠的大脑Krieg 2、3区皮质,对其V层大锥体细胞的五段50μm长度内的树突棘做形态学定量研究。在Golgi法的切片中共计数了三个年龄组的151个细胞的725段树突的棘密度。结果表明,老年大鼠比成年和青年大鼠的棘密度普遍下降。其中以基树突与侧树突棘度下降最显著(减少24％左右),顶树突只中段有明显减少。老年大鼠锥体细胞还常出现胞体、树突及其分支的明显形态改变。     

THE QUANTITATIVE GENETICS OF SEXUAL DIMORPHISM IN SILENE LATIFOLIA (CARYOPHYLLACEAE). II. RESPONSE TO SEX-SPECIFIC SELECTION

A well-established theoretical relationship exists between genetic correlations between the sexes and the dynamics of response to sex-specific selection. The present study investigates the response to sex-specific selection for two sexually dimorphic traits that have been documented to be genetically variable, calyx diameter and flower number, in Silene latifolia. Following the establishment of a base generation with a known genetic background, selection lines were established and two generations of sex-specific selection were imposed. Calyx diameter responded directly to sex-specific selection, and the positive genetic correlation between the sexes was reflected in correlated responses in the sex that was not the basis for selection within a particular line. Flower number showed a more erratic response to sex-specific selection in that selection in some lines was initially in the wrong direction, that is, selection for a decrease in flower number resulted in an increase. These erratic responses were attributable to genotype-environment interaction as reflected in significant heteroscedasticity in variance among families. Correlated responses to selection in the sex that was not the immediate basis for selection indicated the possible existence of a negative genetic correlation between the sexes for this trait. These results test for the first time the impact of genetic correlations between the sexes on the evolutionary dynamics of sexually dimorphic traits in a plant species.     

FISHER'S GEOMETRIC MODEL OF ADAPTATION MEETS THE FUNCTIONAL SYNTHESIS: DATA ON PAIRWISE EPISTASIS FOR FITNESS YIELDS INSIGHTS INTO THE SHAPE AND SIZE OF PHENOTYPE SPACE

The functional synthesis uses experimental methods from molecular biology, biochemistry and structural biology to decompose evolutionarily important mutations into their more proximal mechanistic determinants. However these methods are technically challenging and expensive. Noting strong formal parallels between R.A. Fisher's geometric model of adaptation and a recent model for the phenotypic basis of protein evolution, we sought to use the former to make inferences into the latter using data on pairwise fitness epistasis between mutations. We present an analytic framework for classifying pairs of mutations with respect to similarity of underlying mechanism on this basis, and also show that these data can yield an estimate of the number of mutationally labile phenotypes underlying fitness effects. We use computer simulations to explore the robustness of our approach to violations of analytic assumptions and analyze several recently published datasets. This work provides a theoretical complement to the functional synthesis as well as a novel test of Fisher's geometric model.     

THE EVOLUTION OF THRESHOLD TRAITS: A QUANTITATIVE GENETIC ANALYSIS OF THE PHYSIOLOGICAL AND LIFE-HISTORY CORRELATES OF WING DIMORPHISM IN THE SAND CRICKET

Many traits are phenotypically discrete but polygenically determined. Such traits can be understood using the threshold model of quantitative genetics that posits a continuously distributed underlying trait, called the liability, and a threshold of response, individuals above the threshold displaying one morph and individuals below the threshold displaying the alternate morph. For many threshold traits the liability probably consists of a hormone or a suite of hormones. Previous experiments have implicated juvenile hormone esterase (JHE), a degratory enzyme of juvenile hormone, as a physiological determinant of wing dimorphism in the crickets Gryllus rubens and G. firmus. The present study uses a half-sib experiment to measure the heritability of JHE in the last nymphal stadium of G. firmus and its genetic correlation with fecundity, a trait that is itself genetically correlated with wing morph. The phenotypic and genetic parameters are consistent with the hypothesis that JHE is a significant component of the liability. Comparison of sire and dam estimates suggest that nonadditive effects may be important. Two models have been proposed to account for the fitness differences between morphs: the dichotomy model, which assumes that each morph can be characterized by a particular suite of traits, and the continuous model, which assumes that the associated fitness traits are correlated with the liability rather than the morphs themselves. The latter model predicts that the fitness differences will not be constant but change with the morph frequencies. Variation in fecundity and flight muscle histolysis are shown to be more consistent with the continuous model. Data from the present experiment on JHE are inconclusive, but results from a previous selection experiment also suggest that variation in JHE is consistent only with the continuous model.     

EVOLUTIONARY ENDOCRINOLOGY OF JUVENILE HORMONE ESTERASE: FUNCTIONAL RELATIONSHIP WITH WING POLYMORPHISM IN THE CRICKET,GRYLLUS FIRMUS

The existence, nature, and physiological consequences of genetic variation for juvenile hormone esterase (JHE) activity was studied in the wing-polymorphic cricket, Gryllus firmus. Hemolymph (blood) JHE activity was sixfold lower in nascent short-winged (SW) females, relative to nascent long-winged (LW) females during the last juvenile stadium (stage). Morph-associated genetic variation for JHE activity had two causes, variation in loci: (1) regulating whole-organism enzyme activity; and (2) controlling the degree to which JHE is secreted into the blood Reduced JHE activity in nascent SW-selected individuals was associated with reduced in vivo juvenile hormone catabolism. This suggests that variation in JHE activity during juvenile development may have important physiological consequences with respect to the regulation of blood levels of juvenile hormone and consequent specification of wing morph. This is the first definitive demonstration of genetic variation for hormonal metabolism in any insect and a genetic association between hormone metabolism and the subsequent expression of morphological variation (wing morph). However, we have not yet firmly established whether these associations represent causal relationships In contrast to the clear association between JHE activity and wing morph development, we observed no evidence indicating that variation in JHE activity plays any direct or indirect role in causing the dramatic differences in ovarian growth between adult wing morphs. Variation in JHE activity also does not appear to be important in coordinating the development of wing morph with the subsequent expression of reproductive differences between adult morphs. Finally genetic variation for the developmental profiles of JHE activity during juvenile and adult stages are remarkably similar in three Gryllus species. This suggests that genetic correlations between JHE activities during different periods of development, which underlie these activity profiles, have been conserved since the divergence of the three Gryllus species.     

FUNCTIONAL INCOMPATIBILITY BETWEEN THE FERTILIZATION SYSTEMS OF TWO ALLOPATRIC POPULATIONS OF CALLOSOBRUCHUS MACULATUS (COLEOPTERA: BRUCHIDAE)

Abstract Recent studies indicate that postcopulatory sexual selection may represent an important component of the speciation process by initiating reproductive isolation via the evolutionary divergence of fertilization systems. Using two geographically isolated populations of the polyandrous beetle Callosobruchus maculatus, we investigated divergence in fertilization systems by determining the extent of postcopulatory functional incompatibility. Through reciprocal, cross‐population matings we were able to separately estimate the effects of male and female population origin and their interaction on the extent of last‐male sperm precedence, female receptivity to further copulation and female oviposition. Our results indicate partial incompatibility between the fertilization systems of the two populations at all three functional levels. Males derived from the same population as females outcompete rival, allopatric males with respect to sperm preemption, sperm protection, and ability to stimulate female oviposition. This pattern is reciprocated in both populations indicating that postcopulatory, prezygotic events represent important mechanisms by which between‐population gene flow is reduced. We suggest the partial gametic isolation observed is a by‐product of the coevolution of male and female fertilization systems by a process of cryptic female choice. Our results are consistent with a mechanism akin to conventional mate choice models although they do not allow us to reject antagonistic sexual coevolution as the mechanism of cryptic female choice.