EVOLUTION OF PELVIC REDUCTION IN THREESPINE STICKLEBACK FISH: A TEST OF COMPETING HYPOTHESES

Reimchen hypothesized that pelvic reduction in threespine stickleback is favored by an absence of piscivorous fishes and the resulting increase in predation by insects, but Giles hypothesized that the predation regime is unimportant and that a low dissolved calcium concentration favors evolution of pelvic reduction. Substantial pelvic reduction in threespine stickleback sampled from 179 lakes around Cook Inlet, Alaska is strongly associated both with an absence of predatory fishes and a low calcium concentration. However, the association of pelvic reduction with low calcium concentration appears to be contingent on the absence of predatory fishes. These results emphasize the importance of interactions between seemingly unrelated environmental variables for selection of a single trait. However, these results also conflict with some observations elsewhere and do not rule out the possibility that other environmental factors are important for selection for pelvic reduction in threespine stickleback.     

A CLADISTIC TEST OF THE TAXON CYCLE AND TAXON PULSE HYPOTHESES

Abstract— A species' habitat preference is intcrpretablc both as a response to present-day conditions and as a result of evolutionary response to historical conditions. The taxon cycle and taxon pulse have been proposed as hypotheses that allow prediction of patterns of habitat specialization within a lineage. Both are based on common assumptions: (1) habitat specialization is largely irreversible in a lineage, (2) ecological specializations arise in a center of origin, and (3) dispersal events leading to current distributions can be ascertained. Eight taxa of Antillean, Mexican, and Central American Carabidae, for which cladistic hypotheses of relationship have been proposed, are used to test the generality of the taxon cycle and pulse. The patterns of habitat utilization predicted by the taxon cycle and taxon pulse hypotheses are tested by comparing cladistic transformations of habitat preference to randomly generated patterns of data generated under a null hypothesis. Evolutionary changes in habitat are interpreted using Camin-Sokal coding, which assumes irreversible habitat shifts and a predetermined ecological ground state (assumptions of the taxon cycle and taxon pulse). An observed pattern is considered to demonstrate the taxon cycle or pulse when it results in an explanation of the habitat shifts that is more parsimonious than 95 % of the randomly generated patterns. Of the eight carabid groups, only one exhibits a statistically significant pattern supporting the taxon cycle and pulse. The failure of the taxon cycle and pulse as generally predictive hypotheses may be due to historical changes in climate that permit episodes of range expansion for species previously restricted to small ranges, and habitat shifts and specialization that do not progress in a linear transformation series. Habitat shifts are also analyzed using Farris optimization, resulting in the most parsimonious transformation series of habitats, subject to a predefined ordering of habitats, while allowing reversals. Significance of an observed pattern of habitat shifts under Farris optimization implies habitat constancy relative to cladogenesis, and step-wise changes in habitat preference. Two of the eight groups exhibit significant patterns of habitat utilization under Farris optimization, indicating that vicariance of areas of like habitat has been the predominant factor generating species diversity in these groups. The other groups do not exhibit habitat constancy, suggesting rapid changes in habitat preference relative to speciation.     

LICHEN HERBIVORE PREFERENCE: A TEST OF TWO HYPOTHESES

Field data from a saxicolous lichen community in Shenandoah National Park, Virginia, suggested that the slug Pallifera varia grazed more frequently on certain lichen species than would be expected if it fed randomly. Two hypotheses might explain this grazing pattern. The preference hypothesis, that slugs select lichens of highest quality, was tested by measuring the concentration of essential elements in lichens eliciting high and low preference by slugs. Lichens with the highest element contents were assumed to be of the highest quality to herbivores. The avoidance hypothesis, that preference by slugs actually results from the rejection of unpalatable species, was tested by offering slugs choices of baited filter-paper disks impregnated with secondary products of the lichen species in question. Results suggested that preferred lichens had significantly lower concentrations of N, P, and Ca, and that avoided lichens produced secondary products that effectively inhibited Pallifera grazing activity. These results suggested that the avoidance hypothesis was the better explanation for nonrandom Pallifera grazing patterns. Furthermore, they suggested that lichens with the highest concentrations of essential elements are most likely to produce defense compounds, an observation that supports predictions to explain patterns of chemical defense in vascular plants.     

EVOLUTIONARY ORIGIN OF THE CYCLORRHAPHA (DIPTERA): TEST OF ALTERNATIVE MORPHOLOGICAL HYPOTHESES

Abstract— The higher flies, infraorder Cyclorrhapha [=Muscomorpha (McAlpine, 1989)], have undergone enormous radiation since the Cretaceous (∼100 Myr). Rapid morphological evolution in cyclorrhaphans has made their phylogenetic placement with respect to more primitive clades a long-standing problem in dipteran systematics. Of the two most plausible hypotheses, one treats the Cyclorrhapha as sister group to the orthorrhaphous superfamily Empidoidea [=Empidiformia (Hennig, 1948), Orthogenya (Brauer, 1883)], while the other places them within the empidoids. The debate over cyclorrhaphan origin has heretofore focused on homology interpretations for a few character systems, particularly the male genitalia. We provide the first attempt to assemble and quantify all of the available morphological evidence. By cladistic analysis of these data under alternative codings of genitalic features reflecting opposing homology theories, and then excluding these features altogether, we sought to judge which genitalic theory is better supported by the evidence as a whole, and how much the debate matters to resolving cyclorrhaphan origins. Using the analog of a factorial design, we also measured the effect of alternative transformation series in several other controversial characters, of outgroup choice and of successive weighting. Under all manipulations, including both genitalic codings, the Cyclorrhapha originate within the Empidoidea, near the family Atelestidae. However, trees in which the Empidoidea are constrained to be monophyletic are only 1-6 steps longer (out of ∼150), a fit not significantly worse under a permutation test for monophyly. Adult morphological data may not suffice to settle either the placement of Cyclorrhapha or the debate over genitalic homology. Moreover, the issue of genitalic homology does not appear critical to that of cyclorrhaphan origin.     

THE REGULATION OF ALTERNATION OF GENERATION IN FLOWERING PLANTS

The developmental changes involved in the alternation of generation represent the major gene-switching events in the life history of plants. While a large number of genes are common to both sporophyte and gametophyte, many thousand sequences are specifically expressed in each generation; indeed, certain key constituents (e.g. tubulin) are encoded by different genes in each generation, indicating that sporophyte and gametophyte are responding to different evolutionary pressures. Evidence is accumulating that major gene-switching events in plants, such as flowering, are regulated by complex control systems which ensures that development occurs only in the correct groups of cells at the appropriate time. A similar, or more sophisticated system might thus be expected to regulate alternation of generation. It is not possible to manipulate alternation of generation in a similar fashion to flowering, but study of apparent aberrations of development occurring in nature and in vitro suggests that alternation only occurs in cells which have become competent to receive particular developmental stimuli. Further, in certain cases, competent cells may be switched either into sporophytic or gametophytic developmental pathways depending upon the nature of the stimulus. Acquisition of competence seems to involve isolation of cells from the symplast, some cytoplasmic dedifferentiation, and perhaps cell cycle arrest or transition. The stimuli in vivo appear metabolic in nature, although embryogenesis may be activated by specific classes of glycoproteins. Interestingly, examination of agamospermic systems suggests that fertilization of the egg per se is not the signal which activates sporophytic development. Once competent cells have received the stimulus they start to develop, with no delay in a ‘determined’ state. Sporophytic and gametophytic development in vivo and in vitro both start with an asymmetric division, except for the female gametophyte which may arise via a range of developmental pathways, depending on the species.     

INFLORESCENCE SIZE: TEST OF THE MALE FUNCTION HYPOTHESIS

One explanation for low fruit sets in plants with hermaphroditic flowers is that total flower production by a plant is controlled primarily by selection through male function. This male function hypothesis presupposes that success in pollen donation increases more strongly with flower number than does seed set. I tested this prediction by measuring male and female components of reproductive success as functions of flower number in natural populations of the self-incompatible, perfect flowered plant, Ipomopsis aggregata. Fruit set in this hummingbird-pollinated plant averaged 4.9 to 40.3% across the 4 years of study. Both the total amount of pollen donated and the total amount received, as estimated by movement of fluorescent powdered dyes, increased linearly with number of flowers on a plant. Total seed production, however, increased disproportionately quickly because plants with larger floral displays were more likely to set at least one fruit. An estimate of the functional femaleness of a plant, based on pollen donation and seed production, increased with flower number. These results do not support the male function hypothesis.     

GEOGRAPHIC VARIATION OF SEXUAL DIMORPHISM IN SIZE OF SAVANNAH SPARROWS (PASSERCULUS SANDWICHENSIS): A TEST OF HYPOTHESES

The Savannah Sparrow (Passerculus sandwichensis) is a widespread and common North American bird that shows both geographic variation and sexual dimorphism in size. I used information from 24 measurements on 1,791 individuals from 51 populations to test two hypotheses (sexual-selection and niche-partitioning) about the evolution of sexual dimorphism. Throughout their range male Savannah Sparrows are larger, on average, than females. This doubtless reflects Darwinian sexual selection, for territorial fights usually involve males, many of whom fail to obtain mates. In some parts of their range, Savannah Sparrows are commonly polygynous, whereas in others they are characteristically monogamous. Among species of American sparrows (subfamily Emberizinae) sexual size dimorphism is generally greater in polygynous species than in monogamous ones. However, I did not find a similar trend among populations of Savannah Sparrows. The amount of dimorphism in all populations of Savannah Sparrows is equivalent in magnitude to that of other species of sparrows that are commonly or regularly polygynous, and it is greater than that of other sparrow species that are characteristically monogamous. The amount of sexual dimorphism, either in overall size or in bill size, does not correlate with species diversity and does not differ between island and mainland populations. These results do not support the niche-variation hypothesis. Size dimorphism is relatively great in populations of Savannah Sparrows that are resident in southwestern salt marshes, and these birds are the only sparrow-like birds that generally breed in these marshes. Dimorphism is, in general, relatively great in marsh-dwelling species in the family Emberizidae. These species are commonly, but not always, polygynous; the mating systems of the salt-marsh Savannah Sparrows are not known. There are no significant differences in the extent of dimorphism among populations of salt-marsh sparrows, and there are few among the non-salt-marsh ones, probably reflecting conservatism in the evolution of size dimorphism.     

DO CONSTANT ENVIRONMENTS PROMOTE COMPLEXITY OF FORM?: THE DISTRIBUTION OF BRYOZOAN POLYMORPHISM AS A TEST OF HYPOTHESES

The distribution of cheilostome bryozoans on the Caribbean reefs of Panama was surveyed to test the hypothesis that physically constant environments favor increased morphologic complexity, expressed as the number of zooid types within a colony. The proportion of species within defined grades of complexity did not vary significantly with locality, depth, or substratum. Some differences were found in grade-specific ecological success, measured by colony abundance and spatial cover, but these were not consistently related to habitat type. There was no inverse correlation between morphologic complexity and range of distribution: morphologically specialized cheilostomes were not more stenotopic than generalized forms. Patterns of distribution and total space occupation indicate a sensitivity to local habitat conditions, but relative success of species was not correlated with level of polymorphism. In a bryozoan fauna from Florida, the frequency of polymorphic species was weakly associated with constancy of habitat. In estuaries, polymorphic cheilostomes are almost absent at salinities below 18‰, but this pattern is strongly confounded taxonomically. All species tolerant of low salinities are encrusting anascans; within this group, polymorphism does not decrease significantly with declining salinity. Bryozoan faunas from different biogeographic zones may vary in frequency of avicularian polymorphism, but not along a simple latitudinal cline. These large-scale comparisons may be strongly biased historically and taxonomically. The distribution of cheilostome polymorphism on a local and geographic scale provides no evidence for a causal relationship between habitat constancy and morphologic specialization at the zooidal level. This is in striking contrast to the strong habitat dependence of colony form, which suggests that selective processes may operate differently at the zooidal and colonial levels.     

A TEST OF SEVERAL HYPOTHESES FOR THE DETERMINATION OF SEED NUMBER IN AMELANCHIER ARBOREA,USING SIMULATED PROBABILITY DISTRIBUTIONS TO EVALUATE DATA

The number of seeds per fruit is variable within Amelanchier arborea trees. Because A. arborea flowers are five-carpellate and each carpel contains two ovules, we were able to use the pattern of seed maturation within fruits to test whether the failure of some ovules to develop into seeds is determined by mechanisms operating at the level of carpels, such as stigma-clogging, or by mechanisms operating at the level of ovules, such as ovule infertility. The presence of one-, two-, and zero-seeded carpels demonstrated that the number of ovules developing into seeds was not due entirely to carpel-level phenomena. In order to test the hypothesis of carpel independent seed development, without the assumption that all ovules have the same probability of developing into seeds, it was necessary to use simulation, since no conventional statistical models were appropriate. Analysis of this simulation allowed us to reject carpel independent processes as the only determinant of seed number. A mixed model of seed development, in which some carpels fail entirely and ovules in the remaining carpels develop equiprobably, was shown to be consistent with the data.     

DIFFERING SELECTION ON PLANT PHYSIOLOGICAL TRAITS IN RESPONSE TO ENVIRONMENTAL WATER AVAILABILITY: A TEST OF ADAPTIVE HYPOTHESES

I used phenotypic selection analysis to test the prediction from functional and comparative studies of plants that smaller leaves and more efficient water use are adaptive in drier environments. I measured selection gradients on leaf size and instantaneous water-use efficiency (a measure of carbon gain per unit water loss) in experimental populations of Cakile edentula var. lacustris placed into wet and dry environments in the field. Linear and nonlinear selection differed significantly between the two environments as predicted. Water-use efficiency was selected to be higher, and leaf area was selected toward a small intermediate optimum, in the dry environment. There was also significant positive correlational selection on water-use efficiency and leaf size, suggesting that the optimum leaf size in the dry environment is greater for plants with higher water-use efficiency. In contrast, neither leaf size nor water-use efficiency were selected in the wet environment, though larger leaves resulted in greater vegetative biomass. Path analysis of the linear selection gradients found that water-use efficiency affected plant fitness primarily because it increased vegetative biomass, as suggested by the hypotheses about the function of physiological traits. These results were not only consistent with the functional hypotheses but also with the observed genetic differentiation in water-use efficiency and leaf size between wet and dry site populations.     

ALTERNATION OF GENERATIONS IN FERNS: MECHANISMS AND SIGNIFICANCE

Arguments against the compiling of generalized life cycles summarizing alternation of generations in ferns are presented, and some common misconceptions about breeding systems addressed. What little is known or can be deduced about time frames, mechanisms and significance of alternation events in the lives of two species: bracken fern ( Pteridium ) and Killarney fern ( Trichomanes speciosum ) is presented. Evidence is provided that gametophytes may play a more important role in survival of both these species than previously suspected, and the need for more long-term studies and experiments/measurements of ferns in natural conditions/populations is stressed.     

A DEVELOPMENTAL ANALYSIS OF AN EVOLUTIONARY TREND: DIGITAL REDUCTION IN AMPHIBIANS

In this study, we integrate information from phylogeny, comparative ontogeny, and experimental embryology in an attempt to elucidate the mechanisms controlling evolutionary trends towards digital reduction and loss observed in amphibians. Frogs and salamanders that have lost phalanges and even whole toes have done so in a very ordered manner, i.e., certain skeletal elements are lost prior to others. This pattern of morphological diversity is described and trends elucidated. It is concluded that the process is characterized by striking intraordinal convergences coupled with substantial differences between the trends observed in frogs as compared to urodeles. We argue that this pattern is essentially a reflection of the differences in the ontogenies of the two orders. Similarly, the convergences within urodeles and within anurans can be explained as the result of regulation of developmental parameters in a resilient developmental program. We further explore this hypothesis by experimentally perturbing the number of cells in the embryonic limb primordium to show that reduction in the number of mesenchymal cells secondarily affects the developmental process of pattern formation causing a rearrangement of the skeletal morphology of the foot. The same experimental manipulation has different effects in frogs as compared to salamanders. However, in both cases, the experimentally generated morphologies tend to parallel the phenotypes and trends observed in nature. Our conclusion is that most of the patterns of diversity in the digital morphology of amphibians can be explained as a reflection of developmental properties. In general, we present a methodology that attempts to empirically address the issue of identifying developmental constraint in morphological evolution.     

中国化石两栖类和有鳞类研究进展:过去15年回顾

过去15年中,我国的化石两柄类和有鳞类研究取得了加速进展。共报道了16种两栖类和22种有鳞类,其中15种两栖类和11种有鳞类为新种,包括我国最古老的四足动物宁夏晚泥盆世的潘氏中国螈,湖北中三叠世的大型全椎两栖类宽头远安鲵,东北晚侏罗世/早白垩世的多种滑体两栖类(如无尾类三燕丽蟾和北票中蟾,有尾类奇异热河螈和东方塘螈),华北占近纪的蜥蜴类(如淅川短齿蜥和垣曲响蜥),以及内蒙古晚白垩世的多种蜥蜴类等。部分旧属种被修订,多与热河生物群的种类有关,如细小矢部龙过去依据幼年个体被认为是一种小型细弱的蜥蜴,新材料显示它实为大型强壮的种类。一批新的两栖类和有鳞类重要化石点被发现,如辽宁北票市四合屯、陆家屯和葫芦岛市水口子,河北丰宁县炮仗沟以及内蒙古宁城县道虎沟和乌拉特后旗巴彦满达呼。我国晚侏罗世/早白垩世两栖类和有鳞类对研究滑体两栖类和有鳞类相关类群的起源和早期演化提供了重要信息,但系统发育研究工作尚处于初级阶段。