THE ORIGIN OF AN ORGAN: PHYLOGENETIC ANALYSIS OF EVOLUTIONARY INNOVATION IN THE DIGESTIVE TRACT OF FLIES (INSECTA: DIPTERA)

The cardia, a prominent digestive tract organ consisting of several specialized cell types, occurs throughout the “higher” or muscoid flies, division Schizophora of order Diptera. Phylogenetic analysis of cellular organization in 65 insect species from 36 families indicates that this organ originated within the order Diptera from ancestrally undifferentiated tissues. “Lower” flies, suborder “Nematocera,” display little or no epithelial cell specialization at the corresponding site. Scorpionflies of the outgroup order Mecoptera are similarly unspecialized. Intermediate levels of cellular specialization occur in Tabanomorpha, Asilomorpha and Aschiza, dipteran taxa that diverge between “Nematocera” and Schizophora. The distribution of epithelial characteristics suggests that the cardia evolved through a sequence of simple tissue transformations, combining changes in epithelial configuration with local differentiation of cell structure and function. The evolution of locally specialized cell types implies the emergence of structural genes and regulatory mechanisms through the modification of an ancestral genome that had not supported such extensive differentiation. Comparison of localized gene expression in Drosophila melanogaster with that in other fly species having greater or lesser degrees of cell specialization may provide a practical model system for studying specific patterns of mutation associated with such evolutionary innovation.     

THE ORIGIN OF TETRAPODS-PAST AND PRESENT HYPOTHESES

HistoricalreviewoneshouldexpectthatanewtheorychangesorimprovestheunderstandingofPhylogeneticquestions.ThatdoesnotseemtobetrueoftheoriginoftetrapodsasRosenetal.(l98l)havealreadyshowninthecaseoftheapPearanceofDarwin's'ontheoriginofsPeciesinl859.Incontrast,thehistoryofthedevelopmentofhypothesesontheoriginoftetrapodsdemonstratesthatdiscoveryofnewextantorfos-silforms(Tab.l)shapesourunderstandingoftherelationshipoftetrapodstofishes.Thefrstextantlungfishwasdiscoveredinl836inSouthAmerica(Fitzinge…     

OPTIMUM BROOD SIZE: TESTS OF ALTERNATIVE HYPOTHESES

The most productive litter size (five) was not as common as expected in a free-living population of white-footed mice. I evaluated four competing hypotheses that can explain this pattern. Reproductive costs and annual variation in recruitment appear to be insufficient explanations for the empirical distribution of litter size. Optimal investment of reproductive resources that vary among parents is supported by some tests, but not by all. The abundance of litters less than the apparent optimum is at least partially explained by asymmetric survival in large litters (the cliff-edge hypothesis). Hypotheses that explain the empirical distribution of brood size are not mutually exclusive. Several mechanisms can act alone, or interact, to create an average brood size less than that which appears to produce the greatest number of descendants.     

THE ORIGIN OF MORPHOLOGICAL CHARACTERS AND THE BIOLOGICAL BASIS OF HOMOLOGY

A homolog is a part of the phenotype that is homologous to equivalent parts in other species. A biological homology concept is expected to explain three properties of homologs: 1) the conservation of those features that are used to define a homolog, 2) the individualization of the homolog with regard to the rest of the body, and 3) the uniqueness of homologs, i.e., their specificity for monophyletic groups. The main obstacle to describing a mechanistic basis for homology is the variability of the developmental pathways of undoubtedly homologous characters. However, not all aspects of the developmental pathway are of equal importance. The only organizational features of the developmental system that matter are those that have been historically acquired and cause developmental constraints on the further evolutionary modification of the characters. Two main factors contribute to historically acquired developmental constraints: generative rules of pattern formation and ontogenetic networks. In particular, hierarchical and cyclical inductive networks have the required properties to explain homology. How common such networks are is an open empirical question. The development and variation of pectoral fin hooks in blenniid fishes is presented as a model for the study of a simple ontogenetic network.     

EVOLUTIONARY CHANGE IN THE MORPHOLOGICAL COMPLEXITY OF THE MAMMALIAN VERTEBRAL COLUMN

The notion that morphological complexity increases in evolution is widely accepted in biology and paleontology. Several possible explanations have been offered for this trend, among them the suggestion that it has an active forcing mechanism, such as natural selection or the second law of thermodynamics. No such mechanism has yet been empirically demonstrated, but testing is possible: if a forcing mechanism has operated, the expectation is that complexity would have increased in evolutionary lineages more frequently than it decreased. However, a quantitative analysis of changes in the complexity of the vertebral column in a random sample of mammalian lineages reveals a nearly equal number of increases and decreases. This finding raises the possibility that no forcing mechanism exists, or at least that it may not be as powerful or pervasive as has been assumed. The finding also highlights the need for more empirical tests.     

THE FUNCTION OF CALL ALTERNATION IN ANURAN AMPHIBIANS: A TEST OF THREE HYPOTHESES

Males of many species of anurans alternate calls with those of their neighbors in a chorus. This pattern of calling reduces signal interference and may: 1) facilitate intermale spacing because males can better gauge the intensity of neighbors' calls if these calls do not overlap with their own; 2) help preserve species-specific temporal information in calls required to attract females; and/or 3) make it easier for females to localize males in the chorus. I tested these hypotheses with three species that exhibit call alternation, Hyla crucifer, H. versicolor, and H. microcephala. Males of all three species gave more aggressive calls to high-intensity synthetic stimuli that alternated with their calls than to those that overlapped their calls. These results support the first hypothesis. Results of four-speaker female choice experiments using alternating and overlapping calls indicate that preservation of signal integrity also is important in H. versicolor and H. microcephala, species that have fine-scale temporal information in their calls. However, the third hypothesis was not supported; females failed to discriminate among alternating and overlapping calls if the problem of signal disruption was eliminated or irrelevant.     

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.     

A HYPOTHESIS ON THE EVOLUTIONARY ORIGIN OF THE SQUAMOSAL BONE IN THE THERIAN MAMMALS

Theatheriansandthetherianshavebothinheritedthesquamosa1fromthecyno-dontidancestry,theprimitivepropertyofthesquamosa1intheatherianlineagehavebeenmaintainedfromthebeginning(theLateTriassic,Morganucodontids)totheend(Cretaceous,Triconodontids).Inthetherianlineage,thelowertherianshavethesamesquamosalastheatherians,however,thehighertherianshaveapeculiarsquamosal,so,thechangeofthepropertyofthesquamosaImaytakeplacelntheevolutionaryphasefromthelowertherianmammals(Symmetrodonta,Eupan-totheria,Multitu…     

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.     

中国寄蝇科狭颊寄蝇属研究(双翅目:寄蝇科)

综合研究中国狭颊寄蝇属73种的鉴别方法、分布、寄主种类;研究并发现一些种类的变异特点,♀、♂异型等现象;文中附有73种的检索表及特征图,15个新种的描述;订正了5种为新异名.重建了狭颊寄蝇属的分类系统.     

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.