Constraints on the morphological evolution of marsupial shoulder girdles

Throughout their evolutionary histories, marsupial mammals have been taxonomically and morphologically less diverse than their sister taxa the placentals. Because of this, it has been proposed that the evolution of marsupials has been constrained by the functional requirements of their mode of reproduction. Marsupials give birth after short gestation times to immature neonates that immediately crawl, under the power of their precociously developed shoulder girdles, to the teat where they attach and complete their early development. Using a novel approach incorporating adult and embryological morphological data, this study is the first to both: (1) statistically support adult patterns of morphological divergence consistent with the constraint hypothesis, and (2) identify ontogenetic patterns of morphological change that demonstrate that the constraint was responsible, at least in part, for their formation. As predicted by the marsupial constraint, the shoulder girdles of adult marsupials are less diverse than those of adult placentals, and adult marsupial scapulae are less morphologically diverse than adult marsupial pelves. Furthermore, marsupials that complete an extensive crawl to the teat are restricted to a common pattern of ontogenetic scapular shape change, strongly supporting the hypothesis that the morphological development of the marsupial scapula has been limited evolutionarily by its obligate role in the crawl to the teat. Because this study establishes that ontogenetic and evolutionary morphological change is correlated within mammalian scapulae, it is probable that the marsupial constraint also restricted the morphological divergence of the scapula over evolutionary time by limiting ontogenetic change in the scapula. These findings, coupled with the importance of the shoulder girdle in mammalian locomotor specialization, support the conclusion that the low morphological diversity of marsupial forms over evolutionary time could be directly due to the constraint on marsupial morphological evolution caused by the functional requirements of the crawl to the teat.     

Growth rules based on the modularity of the Canarian Aeoniwm (Crassulaceae) and their phylogenetic value

Growth forms of 22 species of Aeonium (Crassulaceae) were quantified. Since all species are simple in their modular construction, models were developed to predict module length, branching mode and flowering probability using linear and logistic regression. When combined, the parameters of these models are species specific. A discriminant analysis generates a statistically significant separation of species at the level of phylogenetic sections. The results therefore demonstrate the phylogenetic value of growth rules in plants. This dynamic approach strongly contrasts with the traditional static view on forms in systematics and morphology. It also leaves scope for predicting the evolutionary pathways of morphological change which have caused the great diversity of growth forms in the genus Aeonium.     

Rapid evolution of an ancient lake species flock: Freshwater limpets (Gastropoda: Ancylidae) in the Balkan Lake Ohrid

Ancient lakes have long been recognized as evolutionary theatres and hot spots of endemism; the evolution of their morphologically often highly diverse species flocks has received much attention. However, as each ancient lake has its own geological and evolutionary history, modes of speciation may differ from system to system. Ancient lakes can act as evolutionary reservoirs that assure the survival of relict species, but at the same time extant species may evolve through intralacustrine speciation. Other aspects of interest are the actual rates of immigration, diversification or extinction as well as the temporal framework of morphological change. Many of these questions have been addressed in the African (e.g. Lake Tanganyika) and Asian (e.g. Lake Baikal) ancient lakes. For an European ancient lakes (e.g. Lakes Ohrid and Prespa), such studies are largely missing. In the present paper, extraordinarily shaped endemic freshwater limpets of the genus Ancylus from the Balkan Lake Ohrid are used in a phylogeographic and phylogenetic context to test whether they represent an ancient lake species flock, to study the mode of speciation, and to assess the timing of morphological change. Based on DNA data from two mitochondrial genes (COI, LSU rDNA), it has been found that the Lake Ohrid Ancylus species form an endemic monophyletic group. In addition, the lake's feeder springs are inhabited by another, undescribed Ancylus species. All other studied waterbodies within the watershed do not support their own Ancylus lineages but are inhabited by a widespread Mediterranean taxon. The split between the species endemic to the lake and its sister taxon is dated to 1.4±0.6 million years ago. The study presents the first genetic confirmation for the existence of a species flock in a European ancient lake. Contrary to the prevailing opinion it shows that, concerning Ancylus, Lake Ohrid represents a site of intralacustrine speciation rather than an evolutionary reservoir. Moreover, it provides the first evidence for rapid morphological change in an European ancient lake species flock. See also Electronic Supplement at: http://www.senckenberg.de/odes/06-12.htm.     

Evidence of constrained phenotypic evolution in a cryptic species complex of agamid lizards

Lineages that exhibit little morphological change over time provide a unique opportunity to explore whether nonadaptive or adaptive processes explain the conservation of morphology over evolutionary time scales. We provide the most comprehensive evaluation to date of the evolutionary processes leading to morphological similarity among species in a cryptic species complex, incorporating two agamid lizard species (Diporiphora magna and D. bilineata). Phylogenetic analysis of mitochondrial (ND2) and nuclear (RAG-1) gene regions revealed the existence of eight deeply divergent clades. Analysis of morphological data confirmed the presence of cryptic species among these clades. Alternative evolutionary hypotheses for the morphological similarity of species were tested using a combination of phylogenetic, morphological, and ecological data. Likelihood model testing of morphological data suggested a history of constrained phenotypic evolution where lineages have a tendency to return to their medial state, whereas ecological data showed support for both Brownian motion and constrained evolution. Thus, there was an overriding signature of constrained evolution influencing morphological divergence between clades. Our study illustrates the utility of using a combination of phylogenetic, morphological, and ecological data to investigate evolutionary mechanisms maintaining cryptic species.     

Role of development in the evolution of the scapula of the giant sthenurine kangaroos (Macropodidae: Sthenurinae)

Extinct giant sthenurine kangaroos possessed scapulae morphologically distinct from those of all other extant and extinct adult macropodids, but qualitatively resembling those of newborn macropodids. The similarity between adult sthenurine and neonatal macropodid scapulae suggests that a developmental process, such as heterochrony, might have been behind the evolution of the unique sthenurine scapular morphology. By incorporating adult and ontogenetic data, this study examines the evolution and development of the sthenurine scapula. This study quantitatively upholds the previous qualitative morphological observations of macropodid scapulae and finds that ontogenetic and evolutionary morphological changes are correlated in macropodids. The similarity of scapula morphology in sthenurines and newborn macropodids, the correlation between ontogenetic and evolutionary morphological change, and information from other sources (i.e., sthenurine evolutionary history) suggests that pedomorphic shifts in morphology, most likely due to neotenic processes, occurred within the development of the scapula of giant sthenurines.     

Phylogenetic analysis of sexual dimorphism and eye-span allometry in stalk-eyed flies (Diopsidae)

Eye stalks and their scaling relationship with body size are important features in the mating system of many diopsid species, and sexual selection is a critical force influencing the evolution of this exaggerated morphology. Interspecific variation in eye span suggests there has been significant evolutionary change in this trait, but a robust phylogenetic hypothesis is required to determine its rate and direction of change. In this study, the pattern of morphological evolution of eye span is assessed in a phylogenetic framework with respect to its function in the sexual system of these flies. Specifically, we examine within the family Diopsidae the pattern of increase and decrease in sexual dimorphism, the morphological coevolution of eye span between males and females, and the evolutionary flexibility of eye-span allometry. Based on several different methods for reconstructing morphological change, results suggest a general pattern of evolutionary flexibility, particularly for eye-span allometry. Sexual dimorphism in eye span has evolved independently at least four times in the family and this trait also has undergone several reductions within the genus Diasemopsis. Despite most species being dimorphic, there is a strong phylogenetic correlation between males and females for mean eye span. The coevolution between the sexes for eye-span allometry, however, is significantly weaker. Overall, eye-span allometry exhibits significantly more change on the phylogeny than the other morphological traits. The evolutionary pattern in eye-span allometry is caused primarily by changes in eye-span variance. Therefore, this pattern is consistent with recent models that predict a strong relationship between sexual selection and the variance of ornamental traits and highlights the significance of eye-span allometry in intersexual and intrasexual signaling.     

Inferring speciation modes in a clade of Iberian chafers from rates of morphological evolution in different character systems

Background  

Studies of speciation mode based on phylogenies usually test the predicted effect on diversification patterns or on geographical distribution of closely related species. Here we outline an approach to infer the prevalent speciation mode in Iberian Hymenoplia chafers through the comparison of the evolutionary rates of morphological character systems likely to be related to sexual or ecological selection. Assuming that mitochondrial evolution is neutral and not related to measured phenotypic differences among the species, we contrast hypothetic outcomes of three speciation modes: 1) geographic isolation with subsequent random morphological divergence, resulting in overall change proportional to the mtDNA rate; 2) sexual selection on size and shape of the male intromittent organs, resulting in an evolutionary rate decoupled to that of the mtDNA; and 3) ecological segregation, reflected in character systems presumably related to ecological or biological adaptations, with rates decoupled from that of the mtDNA.     

An evaluation of a possible phylogenetic relationship between the Euglenophyta and Kinetoplastida

Winthout close phylogenetic ties to any of the other algae, the Euglenophyta are a taxonomic enigma. The argument is made here that the euglenophytes have extensive morphological homology with the zooflagellte trypanosomatids, bodonids, andIsonema. A phylogenetic sequence is postulated linking these group; the available data suggest that the euglenophytes had their origin in these zooflagellates and that the introduction of the chloroplast and a phototrophic mode of nutrition was a relatively late step in this evolutionary process.     

Extra-oral digestion and the problem of parasitism in Parasitengona mites (Acariformes)

An analysis of the extra-oral digestion in Parasitengona being a highly specialized group of Acariform mites is carried out from the viewpoint of functional morphology and ecology. The significance of the extra-oral digestion in the life strategy of these mites and their larval parasitism is also evaluated. The morphological pre-adaptations of this mode of feeding as well as its probable evolutionary consequences are demonstrated by an example of trombiculid mites (Trombiculidae). It is shown, in particular, that parasitism in general may be considered as a particular life scenario implying that the feeding preferences of the organism are evolutionary formed in a close association with other organisms, which provide a parasite with a feeding substrate mainly already prepared for utilization. Based on this assumption, all Parasitengona, including both larval and post-larval instars, irrespectively of the size of their potential victim, preferably should not be considered as parasites, but as micro-predators.     

Phenotypic plasticity and morphological integration in a marine modular invertebrate

Background  

Colonial invertebrates such as corals exhibit nested levels of modularity, imposing a challenge to the depiction of their morphological evolution. Comparisons among diverse Caribbean gorgonian corals suggest decoupling of evolution at the polyp vs. branch/internode levels. Thus, evolutionary change in polyp form or size (the colonial module sensu stricto) does not imply a change in colony form (constructed of modular branches and other emergent features). This study examined the patterns of morphological integration at the intraspecific level. Pseudopterogorgia bipinnata (Verrill) (Octocorallia: Gorgoniidae) is a Caribbean shallow water gorgonian that can colonize most reef habitats (shallow/exposed vs. deep/protected; 1–45 m) and shows great morphological variation.     

Transposable elements and an epigenetic basis for punctuated equilibria

Evolution is frequently concentrated in bursts of rapid morphological change and speciation followed by long‐term stasis. We propose that this pattern of punctuated equilibria results from an evolutionary tug‐of‐war between host genomes and transposable elements (TEs) mediated through the epigenome. According to this hypothesis, epigenetic regulatory mechanisms (RNA interference, DNA methylation and histone modifications) maintain stasis by suppressing TE mobilization. However, physiological stress, induced by climate change or invasion of new habitats, disrupts epigenetic regulation and unleashes TEs. With their capacity to drive non‐adaptive host evolution, mobilized TEs can restructure the genome and displace populations from adaptive peaks, thus providing an escape from stasis and generating genetic innovations required for rapid diversification. This “epi‐transposon hypothesis” can not only explain macroevolutionary tempo and mode, but may also resolve other long‐standing controversies, such as Wright's shifting balance theory, Mayr's peripheral isolates model, and McClintock's view of genome restructuring as an adaptive response to challenge.     

The role of post-natal ontogeny in the evolution of phenotypic diversity in Podarcis lizards

Understanding the role of the developmental pathways in shaping phenotypic diversity allows appreciating in full the processes influencing and constraining morphological change. Podarcis lizards demonstrate extraordinary morphological variability that likely originated in short evolutionary time. Using geometric morphometrics and a broad suite of statistical tests, we explored the role of developmental mechanisms such as growth rate change, ontogenetic divergence/convergence/parallelism as well as morphological expression of heterochronic processes in mediating the formation of their phenotypic diversity during the post-natal ontogeny. We identified hypermorphosis - the prolongation of growth along the same trajectory - as the process responsible for both intersexual and interspecific morphological differentiation. Albeit the common allometric pattern observed in both sexes of any species constrains and canalizes their cephalic scales variation in a fixed portion of the phenotypic space, the extended growth experienced by males and some species allows them to achieve peramorphic morphologies. Conversely, the intrasexual phenotypic diversity is accounted for by non-allometric processes that drive the extensive morphological dispersion throughout their ontogenetic trajectories. This study suggests a model of how simple heterochronic perturbations can produce phenotypic variation, and thus potential for further evolutionary change, even within a strictly constrained developmental pathway.     

SPECIES DISCRIMINATION AND EVOLUTIONARY MODE OF BUCHIA (BIVALVIA: BUCHIIDAE) FROM UPPER JURASSIC–LOWER CRETACEOUS STRATA OF GRASSY ISLAND, BRITISH COLUMBIA, CANADA

Abstract: Buchiid bivalves are geographically widespread in Upper Jurassic and Lower Cretaceous strata of the Northern Hemisphere. They are often abundant and their short stratigraphic ranges make them ideal biostratigraphic index fossils; these characteristics also render them useful for study of evolutionary patterns. We used multivariate methods to determine if we could discriminate between species of Buchia and examine how morphological characters change through time within the genus. Using ten morphological characters to describe shell shape and size, we tested for taxonomic differences and morphologic change in populations of buchiids collected from a single stratigraphic section on Grassy Island, located along the west coast of Vancouver Island, British Columbia. Morphometric analysis utilized traditional morphological metrics and techniques, including linear and angular measurements as well as Fourier (outline shape) analyses. Phenetic discrimination revealed considerable overlap among the recognized species in the morphospace, as well as a fairly low discriminatory power between species when compared as a group using a step‐wise canonical variate analysis. Step‐wise discriminant analyses between species pairs gave rise to much higher classification rates, suggesting that different characters are important for distinguishing between different species pairs. Our results also indicate that single individuals and small sample sizes of Buchia specimens are insufficient for biostratigraphic discrimination (unless other rarely preserved features such as the hinge and bysuss ear are available) and that a number of previously described species variants may not be taxonomically valid. A biolog using the multivariate axis that best discriminates between species (CV1) and a random walk‐based test using a Hurst estimate analysis indicate a gradualistic evolutionary mode for the Buchia species of Grassy Island. Shell shape and size of buchiids do not appear to be closely tied to lithofacies changes over the c. 10 myr time interval, suggesting that ecophenotypic variation (as it relates to substrate changes) probably had minimal influence on morphology.     

Deconstructing morphology

Scholtz, G. 2010. Deconstructing morphology. —Acta Zoologica (Stockholm) 91 : 44–63 Morphology as the science of form is, in particular, related to the overwhelming diversity of animal forms. Due to its long pre‐Darwinian tradition, organismic morphology is partly burdened by ahistorical typological views. On the other hand, the study of organismic form has always implied concepts of transformation, which helped to pave the way for evolutionary theories. This contradictory history and the fact that we need words to describe organismic form lead in many cases to morphological concepts implying a mixture of structural, functional, developmental, ecological, typological, and evolutionary aspects in current morphological approaches. Because these mixed views lead to contradictory and misleading interpretations of animal form, I stress the need to deconstruct morphological concepts at all levels. I propose a morphology that analyses transformation of animal forms strictly at the structural level in combination with genealogical thinking. Function and other biological aspects of form should be considered in an independent second analytical step. A comparative pattern approach, including developmental patterns, of animal structure in an evolutionary framework allows for the analysis of morphological change, in particular, phylogenetic reconstructions, homology assessment, and the recognition of evolutionary independent morphological units.     

Developmental quantitative genetic models of evolutionary change

Discussions about evolutionary change in developmental processes or morphological structures are predicated on specific quantitative genetic models whose parameters predict whether evolutionary change can occur, its relative rate and direction, and if correlated change will occur in other related and unrelated structures. The appropriate genetic model should reflect the relevant genetical and developmental biology of the organisms, yet be simple enough in its parameters so that deductions can be made and hypotheses tested. As a consequence, the choice of the most appropriate genetic model for polygenically controlled traits is a complex tissue and the eventual choice of model is often a compromise between completeness of the model and computational expediency. Herein, we discuss several developmental quantitative genetic models for the evolution of development and morphology. The models range from the classical direct effects model to complex epigenetic models. Further, we demonstrate the algebraic equivalency of the Cowley and Atchley epigenetic model and Wagner's developmental mapping model. Finally, we propose a new multivariate model for continuous growth trajectories. The relative efficacy of these various models for understanding evolutionary change in developmental and morphological traits is discussed. © 1994 Wiley-Liss, Inc.     

浅谈病理学实验教学中多种教学手段的运用体会

病理学是一门直观性很强的形态学学科,在医学教育和临床医疗中都发挥着重要作用。传统的病理实验教学模式大多为验证性实验教学,教学形式单调,不利于学生对各种知识点的理解及纵向和横向知识的融合。笔者所在的病理教研室尝试综合利用多媒体辅助教学、数字切片和显微数码互动系统改变传统的病理学实验教学模式,旨在提高病理实验教学的效果和质量。     

BACKGROUND MATCHING AND COLOR-CHANGE PLASTICITY IN COLONIZING FRESHWATER SCULPIN POPULATIONS FOLLOWING RAPID DEGLACIATION

Anthropogenic-induced change is forcing organisms to shift their distributions and colonize novel habitats at an increasing rate, which leads to complex interactions among evolutionary processes. Coastrange sculpin ( Cottus aleuticus ) have colonized recently deglaciated streams of Glacier Bay in Alaska within the last 220 years. We examined divergence among populations in background matching coloration and tested the hypothesis that observed variation is due to morphological color plasticity. To examine how color-change plasticity has interacted with other evolutionary processes, we also determined the influence of colonization on neutral genetic diversity. We observed clinal variation in substrate-matching fish color along the chronological continuum of streams. Microsatellites provided little evidence of genetic subdivision among sculpin populations. Fish color was significantly correlated to substrate color, but was not correlated to neutral population genetic structure. Furthermore, a laboratory experiment revealed that morphological color plasticity could explain much, but not all, of the observed fish color divergence. Our study demonstrates that sculpin in Glacier Bay have colonized and adapted to recently deglaciated habitat and suggests that color change plasticity has aided in this process. This research, therefore, highlights the important role phenotypic plasticity may play in the adaptation of species to rapid climate change.     

The evolution of Strepsiptera (Hexapoda)

An evolutionary scenario for the enigmatic group Strepsiptera is provided, based on the results of a comprehensive cladistic analysis of characters of all life stages. A recently described fossil--+Protoxenos janzeni--the most archaic strepsipteran, sheds new light on the early evolution of the group and reduces the "morphological gap" between Strepsiptera and other insects. It weakens both current hypotheses--Coleoptera+Strepsiptera and Diptera+Strepsiptera (="Halteria"). The splitting into +Protoxenos (Protoxenidae) and the remaining Strepsiptera was linked with a distinct size reduction and many morphological changes. Unlike males of extant strepsipteran species, +Protoxenos was still able to process food. Mengeidae (+Mengea), with two small species, is the sister group of extant Strepsiptera. A unique characteristic of extant males (Strepsiptera s. str.) is the mouthfield sclerite. It is part of an air uptake apparatus which belongs to an extremely modified air-filled "balloon gut". Besides this, male strepsipterans possess specialised antennae and compound eyes, a strongly developed flight apparatus, large testes, and a sperm pump, whereas other organ systems are strongly reduced (e.g., fat body, malpighian tubules). Males are designed to find females within a few hours and to copulate. A dramatic change is linked with the split into Mengenillidae and Stylopidia. The change to pterygote hosts and the permanent endoparasitism of the females are evolutionary novelties acquired by the latter clade, and linked with far-reaching morphological transformations, e.g. the presence of unique brood organs. Hairy tarsal adhesive devices are present in males and guarantee efficient attachment to the host during copulation. A well-founded clade within Stylopidia is Stylopiformia, which are characterised by a unique fissure-shaped birth opening. The evolutionary development towards the most specialised and successful forms (parasites of aculeate Hymenoptera [e.g., Xenidae+Stylopidae], ca. 46% of the species) is a stepwise process. The presented evolutionary scenario comprises a complex network of functionally correlated morphological changes in primary larvae, secondary larvae, females and males.     

Influence of Evolutionary Allometry on Rates of Morphological Evolution and Disparity in strictly Subterranean Moles (Talpinae,Talpidae, Lipotyphla,Mammalia)

The adaptation to a particular function could directly influence the morphological evolution of an anatomical structure as well as its rates. The humeral morphology of moles (subfamily Talpinae) is highly modified in response to intense burrowing and fully fossorial lifestyle. However, little is known of the evolutionary pathways that marked its diversification in the two highly fossorial moles tribes Talpini and Scalopini. We used two-dimensional landmark-based geometric morphometrics and comparative methods to understand which factors influenced the rates and patterns of the morphological evolution of the humerus in 53 extant and extinct species of the Talpini (22 extant plus 12 extinct) and Scalopini (six extant plus 13 extinct) tribes, for a total of 623 humeri. We first built a synthetic phylogeny of extinct and extant taxa of the subfamily Talpinae based on all the available information from known phylogenies, molecular data, and age ranges of fossil records. We tested for evolutionary allometry by means of multivariate regression of shape on size variables. Evolutionary allometric trajectories exhibited convergence of humeral shape between the two tribes, even when controlling for phylogeny, though a significant differences in the evolutionary rates was found between the two tribes. Talpini, unlike Scalopini, seem to have reached a robust fossorial morphology early during their evolution, and their shape disparity did not change, if it did not decrease, through time. Furthermore, the basal Geotrypus spp. clearly set apart from the other highly fossorial moles, exhibiting a significant acceleration of evolutionary shifts toward higher degree of fossorial adaptation. Our observations support the hypothesis that the evolution of allometry may reflect a biological demand (in this case functional) that constrains the rates of evolution of anatomical structures.     

Evolutionary dynamics of a sexual ornament in the house sparrow (Passer domesticus): the role of indirect selection within and between sexes

The relative contribution of sexual and natural selection to evolution of sexual ornaments has rarely been quantified under natural conditions. In this study we used a long-term dataset of house sparrows in which parents and offspring were matched genetically to estimate the within- and across-sex genetic basis for variation and covariation among morphological traits. By applying two-sex multivariate "animal models" to estimate genetic parameters, we estimated evolutionary changes in a male sexual ornament, badge size, from the contribution of direct and indirect selection on correlated traits within males and females, after accounting for overlapping generations and age-structure. Indirect natural selection on genetically correlated traits in males and females was the major force causing evolutionary change in the male ornament. Thus, natural selection on female morphology may cause indirect evolutionary changes in male ornaments. We observed however no directional phenotypic change in the ornament size of one-year-old males during the study period. On the other hand, changes were recorded in other morphological characters of both sexes. Our analyses of evolutionary dynamics in sexual characters require application of appropriate two-sex models to account for how selection on correlated traits in both sexes affects the evolutionary outcome of sexual selection.