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Arhynchobdellida (Annelida: Oligochaeta: Hirudinida): phylogenetic relationships and evolution.

A remarkable diversity of life history strategies, geographic distributions, and morphological characters provide a rich substrate for investigating the evolutionary relationships of arhynchobdellid leeches. The phylogenetic relationships, using parsimony analysis, of the order Arhynchobdellida were investigated using nuclear 18S and 28S rDNA, mitochondrial 12S rDNA, and cytochrome c oxidase subunit I sequence data, as well as 24 morphological characters. Thirty-nine arhynchobdellid species were selected to represent the seven currently recognized families. Sixteen rhynchobdellid leeches from the families Glossiphoniidae and Piscicolidae were included as outgroup taxa. Analysis of all available data resolved a single most-parsimonious tree. The cladogram conflicted with most of the traditional classification schemes of the Arhynchobdellida. Monophyly of the Erpobdelliformes and Hirudiniformes was supported, whereas the families Haemadipsidae, Haemopidae, and Hirudinidae, as well as the genera Hirudo or Aliolimnatis, were found not to be monophyletic. The results provide insight on the phylogenetic positions for the taxonomically problematic families Americobdellidae and Cylicobdellidae, the genera Semiscolex, Patagoniobdella, and Mesobdella, as well as genera traditionally classified under Hirudinidae. The evolution of dietary and habitat preferences is examined.

Animals↗

Is morphology still relevant?

The utility of morphological data in modern systematics has recently been challenged because strong selection pressures are thought to create widespread patterns of convergent evolution at this level. This concern has led to suggestions that morphological data should be excluded either from all analyses or at least from analyses where there is conflict with molecular data. These concerns, however, are generally unwarranted and excluding data is not a defensible strategy for dealing with problems that do exist. We emphasize the importance of empirical responses, such as collecting additional and diverse data and exploring taxa and data set interactions, rather than the implementation of a priori assumptions, to overcoming many of the concerns associated with combining morphological and molecular data. Numerous factors may create biases in both molecules and morphology. While these biases are prevalent enough to cause widespread incongruence, they highlight the importance of combining, rather than separating, data. Morphological data also offer distinct advantages over molecular data, such as the inclusion of fossil taxa, cost-effectiveness and presence of biases different from those in molecular data.

Animals↗

Molecular phylogenetic analysis of Onchocerca lupi and its Wolbachia endosymbiont.

The morphology of Onchocerca lupi, responsible for canine ocular onchocercosis, is unique within the genus. Earlier analyses of the 5S ribosomal RNA gene spacer region sequence of the parasite and the 16S ribosomal RNA gene sequence of its Wolbachia endosymbiotic bacteria (Rickettsiales) supported the morphological and biological arguments that O. lupi is a distinct species. However, the exact phylogenetic position of O. lupi and its endosymbiont could not be unambiguously determined. Herein we report analyses based on the mitochondrial cytochrome oxidase I (COI) gene of the filarial species and the Wolbachia surface protein (wsp) and the bacterial cell-cycle ftsZ genes of their wolbachiae. Our results indicate that O. lupi separated from other Onchocerca spp. early in evolution. This is in line with the previous morphological analysis demonstrating that O. lupi is an atypical Onchocerca species showing both primitive and evolved characters. The phylogenetic trees generated for the COI sequences of filariae and the wsp and ftsZ sequences of their wolbachiae were congruent with each other, which supports the hypothesis that nematodes and their Wolbachia endobacteria share a long co-evolutionary history.

Animals↗

A biogeographic pattern in sparrow bill morphology: parallel adaptation to tidal marshes.

The study of ecological convergence, the evolution of similar traits on multiple occasions in response to similar conditions, is a powerful method for developing and testing adaptive hypotheses. However, despite the great attention paid to geographic variation and the foraging ecology of birds, surprisingly few cases of convergent or parallel feeding adaptations have been adequately documented. In this study, we document a biogeographic pattern of parallel bill morphology across 10 sparrow taxa endemic to tidal marshes. All North American tidal marsh sparrows display parallel differentiation from close relatives in other habitats, suggesting that selection on bill morphology is strong. Relative to their body mass, tidal marsh sparrows have longer, thinner bills than their non-tidal marsh counterparts, which is likely an adaptation for consuming more invertebrates and fewer seeds, as well as for probing in sediment crevices to capture prey. Published data on tidal marsh food resources and diet of the relevant taxa support this hypothesis. This morphological differentiation is most pronounced between sister taxa with the greatest estimated divergence times, but is found even in taxa that show little or no structure in molecular genetic markers. We, therefore, speculate that tidal marsh ecosystems are likely settings for ecological speciation.

Adaptation, Biological↗

Thirty-five years of Tropical biology: a quantitative history.

Citation indices are inappropriate measures of scientific output and impact. For that reason, nonparametric statistics were preferred to analyze 35 years of publication on Tropical biology in the Revista de Biología Tropical. The most frequent subjects are animal taxonomy, human biology-including medicine- ecology and animal behavior. Botany papers are less frequent and mainly deal with morphology and taxonomy. Applied studies are not predominant. In that period, only one case of unethical experimentation with humans was discovered in a paper of Mexican origin. The proportion of foreign institutions publishing in the journal has increased from 23% (1953-1963) to 50% in the last decade; similarly, the number of studies done in the Neotropics is on the rise. English and Spanish are equally frequent, although English is the basic language of ethology and evolution and Spanish predominates in papers on animal morphology and parasitology. Most Costa Rican authors publish in Spanish, in contrast with many of their Latin American colleagues. In recent years, there is a tendency to publish shorter papers written by more than one author. The Revista de Biología Tropical, covered by 18 data bases, accounts for most exchanges and sales in the Universidad de Costa Rica and has worldwide distribution.

Biology↗

The Fox and the thyroid: the amphioxus perspective.

The evolutionary origins of several vertebrate organs are still controversial. The thyroid is classically thought to derive directly from the endostyle (a pharyngeal organ found in urochordates, cephalochordates and lampreys). Several molecular and biochemical lines of evidence agree with this scenario. However, a recent paper,1 describing the expression of a FoxE ortholog in amphioxus, suggests that some molecular pathways might actually have been recruited from an adjacent region of the pharynx. Although additional data from urochordates and lamprey are needed to confirm this hypothesis; these results propose an interesting new scenario for thyroid evolution that involved the reorganisation of genetical and morphological features in the pharyngeal endoderm in order to give rise to a entirely new organ. They also give an indication that the ancestral role of the FoxE gene family was probably limited to the differentiation of part of the pharynx.

Animals↗

Phenotypic evolution and hidden speciation in Candidula unifasciata ssp. (Helicellinae, Gastropoda) inferred by 16S variation and quantitative shell traits.

In an effort to link quantitative morphometric information with molecular data on the population level, we have analysed 19 populations of the conchologically variable land snail Candidula unifasciata from across the species range for variation in quantitative shell traits and at the mitochondrial 16S ribosomal (r)DNA locus. In genetic analysis, including 21 additional populations, we observed two fundamental haplotype clades with an average pairwise sequence divergence of 0.209 +/- 0.009 between clades compared to 0.017 +/- 0.012 within clades, suggesting the presence of two different evolutionary lineages. Integrating additional shell material from the Senckenberg Malacological Collection, a highly significant discriminant analysis on the morphological shell traits with fundamental haplotype clades as grouping variable suggested that the less frequent haplotype corresponds to the described subspecies C. u. rugosiuscula, which we propose to regard as a distinct species. Both taxa were highly subdivided genetically (FST = 0.648 and 0.777 P < 0.001). This was contrasted by the partition of morphological variance, where only 29.6% and 21.9% of the variance were distributed among populations, respectively. In C. unifasciata, no significant association between population pairwise FST estimates and corresponding morphological fixation indices could be detected, indicating independent evolution of the two character sets. Partial least square analysis of environmental factors against shell trait variables in C. u. unifasciata revealed significant correlations between environmental factors and certain quantitative shell traits, whose potential adaptational values are discussed.

Animals↗

Cycads: evolutionary innovations and the role of plant-derived neurotoxins.

Cycads are an important relic from the past and represent the oldest living seed plants. Cycads have been instrumental in our understanding the evolution of angiosperms and gymnosperms because they have recognizable morphological characteristics intermediate between less-recently evolved plants such as ferns and more-derived (advanced) plants including the angiosperms. Cycads also produce several compounds that are carcinogenic and neurotoxic. Because of their unique placement in terrestrial plant evolution, molecular studies should help to define the origins of structures that led to the rise of seed plants and the role of neurotoxic compounds that are found in cycads.

Biological Evolution↗

Sexual selection and genital evolution.

Genitalia are conspicuously variable, even in closely related taxa that are otherwise morphologically very similar. Explaining genital diversity is a longstanding problem that is attracting renewed interest from evolutionary biologists. New studies provide ever more compelling evidence that sexual selection is important in driving genital divergence. Importantly, several studies now link variation in genital morphology directly to male fertilization success, and modern comparative techniques have confirmed predicted associations between genital complexity and mating patterns across species. There is also evidence that male and female genitalia can coevolve antagonistically. Determining mechanisms of genital evolution is an important challenge if we are to resolve current debate concerning the relative significance of mate choice benefits and sexual conflict in sexual selection.

Journal Article↗

Complex and Dynamic Gene-by-Age and Gene-by-Environment Interactions Underlie Functional Morphological Variation in Adaptive Divergence in Arctic Charr (Salvelinus alpinus).

The evolution of adaptive phenotypic divergence requires heritable genetic variation. However, it is underappreciated that trait heritability is molded by developmental processes interacting with the environment. We hypothesized that the genetic architecture of divergent functional traits was dependent on age and foraging environment. Thus, we induced plasticity in full-sib families of Arctic charr (Salvelinus alpinus) morphs from two Icelandic lakes by mimicking prey variation in the wild. We characterized variation in body shape and size at two ages and investigated their genetic architecture with quantitative trait locus (QTL) analysis. Age had a greater effect on body shape than diet in most families, suggesting that development strongly influences phenotypic variation available for selection. Consistent with our hypothesis, multiple QTL were detected for all traits and their location depended on age and diet. Many of the genome-wide QTL were located within a subset of duplicated chromosomal regions suggesting that ancestral whole genome duplication events have played a role in the genetic control of functional morphological variation in the species. Moreover, the detection of two body shape QTL after controlling for the effects of age provides additional evidence for genetic variation in the plastic response of morphological traits to environmental variation. Thus, functional morphological traits involved in phenotypic divergence are molded by complex genetic interactions with development and environment.

Animals↗

The development of crustacean limbs and the evolution of arthropods.

Arthropods exhibit great diversity in the position, number, morphology, and function of their limbs. The evolutionary relations among limb types and among the arthropod groups that bear them (insects, crustaceans, myriapods, and chelicerates) are controversial. Here, the use of molecular probes, including an antibody to proteins encoded by arthropod and vertebrate Distal-less (Dll and Dlx) genes, provided evidence that common genetic mechanisms underlie the development of all arthropod limbs and their branches and that all arthropods derive from a common ancestor. However, differences between crustacean and insect body plans were found to correlate with differences in the deployment of particular homeotic genes and in the ways that these genes regulate limb development.

Amino Acid Sequence↗

The significance of the evolution of the cerebrospinal fluid system.

A study of the comparative morphology of the cerebrospinal fluid (CSF) system has been made in amphioxus, lamprey, dogfish, goldfish, lungfish, frog, salamander, turtle, pigeon, and mouse. Using mainly intracardiac fixation and a careful histological technique, serial sections have been obtained of the brain in situ surrounded by its various membranes and the skull. The ventricular system, the roof of the hindbrain, the meninges and subarachnoid space, the ependyma with its various derivatives, including the choroid plexuses and paraphysis, and the relationship between the various CSF compartments and the cerebrovascular system have all been compared in these animals. The hypothesis has been derived that the CSF system is primarily developed to maintain the chemical environment necessary to the function of the cells of the central nervous system, including the neuroendocrine pathways.

Animals↗

Canalization, developmental stability, and morphological integration in primate limbs.

Canalization and developmental stability refer to the tendency of developmental processes to follow particular trajectories, despite external or internal perturbation. Canalization is the tendency for development of a specific genotype to follow the same trajectory under different conditions (different environments or different genetic backgrounds), while developmental stability is the tendency for the development of a specific genotype to follow the same trajectory under the same conditions. Morphological integration refers to the tendency for structures to show correlated variation because they develop in response to shared developmental processes or function in concert with other structures. All three phenomena are emergent properties of developmental systems that can affect the interaction of development and evolution. In this paper, we review the topics of canalization, developmental stability, and morphological integration and their relevance to primate and human evolution. We then test three developmentally motivated hypotheses about the patterning of variability components in the mammalian limb. We find that environmental variances and fluctuating asymmetries (FA) increase distally along the limb in adult macaques but not in fetal mice. We infer that the greater variability of more distal segments in macaques is due to postnatal mechanical effects. We also find that heritability and FA are significantly correlated when different limb measurements are compared in fetal mice. This supports the idea that the mechanisms underlying canalization and developmental stability are related. Finally, we report that the covariation structure of fore- and hindlimb skeletal elements shows evidence for morphological integration between serially homologous structures between the limbs. This is evidence for the existence of developmental modules that link structures between the limbs. Such modules would produce covariation that would need to be overcome by selection for divergence in hind- and forelimb morphology.

Anatomy, Comparative↗

The evolution of spinal instrumentation for the management of occipital cervical and cervicothoracic junctional injuries.

STUDY DESIGN/SETTING: Independent computer-based literature review of articles pertaining to instrumentation and fusion of junctional injuries of the cervical spine. OBJECTIVE: To review and discuss the evolution of instrumentation techniques and systems used in the treatment of cervical spine junctional injuries. SUMMARY OF BACKGROUND DATA: Instrumentation of junctional injuries of the cervical spine has been limited historically by failure to achieve rigid internal fixation in multiple planes. The evolution of these techniques has required increased insight into the morphology and unique biomechanics of the structures to be instrumented. METHODS: Computer-based literature search of Ovid and PubMed databases. RESULTS: Extensive literature search yielded insights into the evolution of systems initially based on onlay bone graft combined with wiring techniques. Such techniques have come to include systems incorporating rigid, longitudinal struts that accommodate multiplanar screws placed in the lateral masses, pedicles, transarticular regions, and occipital bone. CONCLUSIONS: Despite a rapid evolution of techniques and instrumentation technologies, it remains incumbent on the physician to provide the patient with a surgical procedure that balances the likelihood of a favorable outcome with the risk inherent in the implementation of the procedure.

Cervical Vertebrae↗

Tempo and mode of evolutionary radiation in iguanian lizards.

Identification of general properties of evolutionary radiations has been hindered by the lack of a general statistical and phylogenetic approach applicable across diverse taxa. We present a comparative analytical framework for examining phylogenetic patterns of diversification and morphological disparity with data from four iguanian-lizard taxa that exhibit substantially different patterns of evolution. Taxa whose diversification occurred disproportionately early in their evolutionary history partition more of their morphological disparity among, rather than within, subclades. This inverse relationship between timing of diversification and morphological disparity within subclades may be a general feature that transcends the historically contingent properties of different evolutionary radiations.

Adaptation, Biological↗

Elevated rates of protein secretion, evolution, and disease among tissue-specific genes.

Variation in gene expression has been held responsible for the functional and morphological specialization of tissues. The tissue specificity of genes is known to correlate positively with gene evolution rates. We show here, using large data sets, that when a gene is expressed highly in a small number of tissues, its protein is more likely to be secreted and more likely to be mutated in genetic diseases with Mendelian inheritance. We find that secreted proteins are evolving at faster rates than nonsecreted proteins, and that their evolutionary rates are highly correlated with tissue specificity. However, the impact of secretion on evolutionary rates is countered by tissue-specific constraints that have been held constant over the past 75 million years. We find that disease genes are underrepresented among intracellular and slowly evolving housekeeping genes. These findings illuminate major selective pressures that have shaped the gene repertoires expressed in different mammalian tissues.

Animals↗

Evolution of the angiosperms: calibrating the family tree.

Growing evidence of morphological diversity in angiosperm flowers, seeds and pollen from the mid Cretaceous and the presence of derived lineages from increasingly older geological deposits both imply that the timing of early angiosperm cladogenesis is older than fossil-based estimates have indicated. An alternative to fossils for calibrating the phylogeny comes from divergence in DNA sequence data. Here, angiosperm divergence times are estimated using non-parametric rate smoothing and a three-gene dataset covering ca. 75% of all angiosperm families recognized in recent classifications. The results provide an initial hypothesis of angiosperm diversification times. Using an internal calibration point, an independent evaluation of angiosperm and eudicot origins is performed. The origin of the crown group of extant angiosperms is indicated to be Early to Middle Jurassic (179-158 Myr), and the origin of eudicots is resolved as Late Jurassic to mid Cretaceous (147-131 Myr). Both estimates, despite a conservative calibration point, are older than current fossil-based estimates.

Calibration↗

A molecular timescale for vertebrate evolution.

A timescale is necessary for estimating rates of molecular and morphological change in organisms and for interpreting patterns of macroevolution and biogeography. Traditionally, these times have been obtained from the fossil record, where the earliest representatives of two lineages establish a minimum time of divergence of these lineages. The clock-like accumulation of sequence differences in some genes provides an alternative method by which the mean divergence time can be estimated. Estimates from single genes may have large statistical errors, but multiple genes can be studied to obtain a more reliable estimate of divergence time. However, until recently, the number of genes available for estimation of divergence time has been limited. Here we present divergence-time estimates for mammalian orders and major lineages of vertebrates, from an analysis of 658 nuclear genes. The molecular times agree with most early (Palaeozoic) and late (Cenozoic) fossil-based times, but indicate major gaps in the Mesozoic fossil record. At least five lineages of placental mammals arose more than 100 million years ago, and most of the modern orders seem to have diversified before the Cretaceous/Tertiary extinction of the dinosaurs.

Animals↗