Evolution of a dental character in the beaver Castor fiber L. (Mammalia: Rodentia)

Variation in morphology of beaver cheek teeth has been examined in respect of ontogeny, geographical variation and geological time. When the effects of ontogeny are taken into account, certain patterns of dental variation are found to be restricted in space and time. An isolated loop of enamel ubiquitously present in the lower third molars of European beavers from the present interglacial seems to be a "new" acquisition which is lacking in Recent Siberian and North American beavers, as well as in Western European specimens from earlier interglacials.
Whilst the genetic basis of such morphological details is presumed rather than established, they may be helpful in tracing past population movements and are also of potential stratigraphic importance. Changes in range associated with Pleistocene climatic oscillations have led to the "sudden" appearance of this and other castorid dental characters in the Western European fossil record. The morphological discontinuity in the fossil record should be taken as evidence of climatic change rather than evidence for the operation of a certain pattern of evolution.     

Ontogenic study of the brachiopod Dicoelosia by geometric morphometrics and morphing techniques

The distinctive brachiopod Dicoelosia King 1850 is characterized by a strongly bilobed outline. To date, studies have concentrated on its functional morphology, taxonomy and evolution; little attention has been paid to its ontogeny. Here, we map population variation by principal component analysis for over 80 specimens distributed across five species of Dicoelosia. Using geometric morphometrics with landmarks for some 40 specimens, the ontogenic trends in D. sp. nov. are compared with those of Dicoelosia biloba. In addition, the ontogenic pathway in D. sp. nov. is investigated by morphing with control points, a new technique introduced here to palaeontology. Combining the results above, the ontogeny of the key character of the genus, emargination, is modelled. Within single populations, taxa may develop from broad weakly emarginate forms into those that are elongate and deeply emarginate. As the identification of the genus and its species depends on external morphological characters, the definition of ontogenetic trends in each species is essential for taxonomic discrimination. Substantial population variation exists in many of its species; however, the morphing technique provides a method of simulation, predicting the full range of ontogenetic variation in given populations.     

Features of inheritance of several morphological characters in the hybrids between Siberian sturgeon <Emphasis Type="Italic">Acipenser baerii</Emphasis> and beluga <Emphasis Type="Italic">A. huso</Emphasis> (Acipenseridae) and in gynogenetic individuals of Siberian sturgeon

Comparative analysis of several morphological characters is conducted in experimentally obtained hybrids between Siberian sturgeon Acipenser baerii and beluga A. huso, in parental species, and in gynogenetic progeny of Siberian sturgeon. The gynogenetic individuals are similar to Siberian sturgeon by all characters used for differentiation between Siberian sturgeon and beluga that supports gynogenetic origin of these individuals. Based on the data obtained on the hybrid specimens, the features of inheritance of the studied characters in sturgeons and their variation in postlarval ontogeny are discussed.     

Variation in heteroblastic succession among populations of Crepis tectorum

Eleven populations of the monocarpic species, Crepis tectorurn (Asteraceae) in South Sweden differed in the extent to which different leaf characters changed during plant ontogeny. Multivariate analyses on sequences of leaf samples collected from greenhouse-grown plants of different ages revealed a group structure of populations that was different from that revealed by analysing variation in leaves from adult plants. Ontogenetic data suggested that populations within a weed and an alvar ecotype constituted more natural groups than combinations of populations from both eco-types that had similar leaves at adult stages, supporting the hypothesis that these ecotypes may have different migration histories in this area. Variation among the populations in leaf characteristics was partly due to differences that were present at all ontogenetic stages and partly due to variation in the rate of development of some of the leaf characters during plant ontogeny. Plants in some populations appeared to have more 'juvenile' leaves at adult stages than plants in other populations, at least in some characters.     

Phylogenetic interpretation of ontogenetic change: sorting out the actual and artefactual in an empirical case study of centrarchid fishes

Hypothesized relationships between ontogenetic and phylogenetic change in morphological characters were empirically tested in centrarchid fishes by comparing observed patterns of character development with patterns of character evolution as inferred from a representative phylogenetic hypothesis. This phylogeny was based on 56–61 morphological characters that were polarized by outgroup comparison. Through these comparisons, evolutionary changes in character ontogeny were categorized in one of eight classes (terminal addition, terminal deletion, terminal substitution, non-terminal addition, non-terminal deletion, non-terminal substitution, ontogenetic reversal and substitution). The relative frequencies of each of these classes provided an empirical basis from which assumptions underlying hypothesized relationships between ontogeny and phylogeny were tested. In order to test hypothesized relationships between ontogeny and phylogeny that involve assumptions about the relative frequencies of terminal change (e.g. the use of ontogeny as a homology criterion), two additional phylogenies were generated in which terminal addition and terminal deletion were maximized and minimized for all characters. Character state change interpreted from these phylogenies thus represents the maxima and minima of the frequency range of terminal addition and terminal deletion for the 8.7 × 10³⁶ trees possible for centrarchids. It was found for these data that terminal change accounts for c. 75% of the character state change. This suggests either that early ontogeny is conserved in evolution or that interpretation and classification of evolutionary changes in ontogeny is biased in part by the way that characters are recognized, delimited and coded. It was found that ontogenetic interpretation is influenced by two levels of homology decision: an initial decision involving delimitation of the character (the ontogenetic sequence), and the subsequent recognition of homologous components of developmental sequences. Recognition of phylogenetic homology among individual components of developmental sequences is necessary for interpretation of evolutionary changes in ontogeny as either terminal or non-terminal. If development is the primary criterion applied in recognizing individual homologies among parts of ontogenetic sequences, the only possible interpretation of phylogenetic differences is that of terminal change. If homologies of the components cannot be ascertained, recognition of the homology of the developmental sequence as a whole will result in the interpretation of evolutionary differences as substitutions. Particularly when the objective of a study is to discover how ontogeny has evolved, criteria in addition to ontogeny must be used to recognize homology. Interpretation is also dependent upon delimitation within an ontogenetic sequence. This is in part a function of the way that an investigator ‘sees’ and codes characters. Binary and multistate characters influence interpretation differently and predictably. The use of ontogeny for determining phylogenetic polarity as previously proposed rests on the assumptions that ancestral ontogenies are conserved and that character evolution occurs predominantly through terminal addition. It was found for these data that terminal addition may comprise a maximum of 51.9% of the total character state change. It is concluded that the ontogenetic criterion is not a reliable indicator of phylogenetic polarity. Process and pattern data are collected simultaneously by those engaged in comparative morphological studies of development. The set of alternative explanatory processes is limited in the process of observing development. These form necessary starting points for the research of developmental biologists. Separating ‘empirical’ results from interpretational influences requires awareness of potential biases in the course of character selection, coding and interpretation. Consideration of the interpretational problems involved in identifying and classifying phylogenetic changes in ontogeny leads to a re-evaluation of the purpose, usefulness and information conveyed by the current classification system. It is recommended that alternative classification schemes be pursued.     

Morphological variation in Gyrinus sericeolimbatus Rég. in New Guinea and description of a new Gyrinus species (Coleoptera: Gyrinidae)

Abstract. Samples of Gyrinus sericeolimbatus from twenty-six localities in New Guinea vary greatly in metric body characters as well as in elytral reticulation patterns. The variation is smaller within than between populations. Although the characters of adjacent populations are usually similar, certain proximate populations exhibit significant differences, probably resulting from some kind of dispersal barrier. Similarities between populations at comparable latitudes suggest that some of the morphological variation is altitude-related. In addition, evidence is found for habitat-related morphological differentiation. Changes in phenotypes in response to possible environmental factors in ontogeny and constraints imposed by various selection pressures are discussed.
A new species, G.brincki sp.n., from western New Guinea is described.     

ONTOGENETIC VARIATION IN PATTERNS OF DEVELOPMENTAL AND FUNCTIONAL INTEGRATION IN SKULLS OF SIGMODON FULVIVENTER

Patterns of variation and covariation within populations can influence how characters respond to natural selection and random genetic drift and so constrain the ability of natural selection to modify the phenotype. We examined several potential developmental and functional explanations of character covariation throughout ontogeny using known-age samples of the cotton rat (Sigmodon fulviventer) to identify the causes of covariation and to assess the variability of patterns of covariation throughout postnatal growth. Competing developmental and functional models were fit to samples of orofacial and neurocranial measures by confirmatory factor analysis and evaluated for their ability to reconstruct observed variance-covariance matrices. Samples of successive ages were simultaneously fit to a common model to test the hypothesis that the patterns of developmental and functional integration were invariant between ages. Orofacial characters derived from the same branchial-arch primordium covary early in ontogeny. Subsequently, there is a repatterning of integration that may reflect a transition from developmental to functional sources of integration. Neurocranial characters exhibit even more variation in patterns of covariation: initially, characters appear to comprise a single integrated unit; before puberty, they appear to respond to localized bone growth; after puberty, they form separate calvarial and basicranial components. This ontogenetic variation in patterns of covariation suggests that developmental constraints are transient and flexible and that the consequences of selection may depend upon the age at which it acts.     

Variable segment number in centipedes: population genetics meets evolutionary developmental biology

The case studies of population genetics focus on intraspecific variation, but most cases--at least where the variation is polymorphic--deal with characters that are not directly linked to organismic structure or ontogeny. Conversely, the case studies of evolutionary developmental biology focus directly on structure/ontogeny, but usually involve only interspecific comparisons. To integrate these complementary approaches, it is desirable to have a model system that permits study of intraspecific variation in development, using a character whose genetic basis either is already known or can be elucidated. Segment number in geophilomorph centipedes is proposed as a possible model system of this kind. Segment number is variable in natural populations of geophilomorphs, while in the other centipede orders it is fixed, either completely (scutigeromorphs, lithobiomorphs), or at least within species (scolopendromorphs). Statistical analysis of data on the extent of variation in different geophilomorph species suggests that segment number may be of selective importance, rather than the variation being merely an inevitable consequence of the difficulty of achieving a high degree of repeatability when there is a large number of segments.     

Morphological evolution in Ceratophryinae frogs (Anura,Neobatrachia): the effects of heterochronic changes during larval development and metamorphosis

Heterochrony produces morphological change with effects in shape, size, and/or timing of developmental events of a trait related to an ancestral ontogeny. This paper analyzes heterochrony during the ontogeny of Ceratophryinae (Ceratophrys, Chacophrys, and Lepidobatrachus), a monophyletic group of South American frogs with larval development, and uses different approaches to explore their morphological evolution: (1) inferences of ancestral ontogenies and heterochronic variation from a cladistic analysis based on 102 morphological larval and adult characters recorded in ten anuran taxa; (2) comparisons of size, morphological variation, and timing (age) of developmental events based on a study of ontogenetic series of ceratophryines, Telmatobius atacamensis, and Pseudis platensis. We found Chacophrys as the basal taxon. Ceratophrys and Lepidobatrachus share most derived larval features resulting from heterochrony. Ceratophryines share high rates of larval development, but differ in rates of postmetamorphic growth. The ontogeny of Lepidobatrachus exhibits peramorphic traits produced by the early onset of metamorphic transformations that are integrated in an unusual larval morphology. This study represents an integrative examination of shape, size, and age variation, and discusses evolutionary patterns of metamorphosis. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 752–780.     

Morphological variation in some lower palaeozoic dalmanellid brachiopods

Three predictions arising from the niche-variation model concerning morphological variation are tested. These are: increasing diversity accompanied by reduced species variability, variation directly proportional to species abundance and eurytopic species more variable than stenotopic forms. Using Ordovician and Silurian dalmanellid brachiopods, measurements were taken at different intervals through ontogeny of 5 brachial characters on 11 species ofBancroftina, Dalmanella andOnniella (all Ordovician andIsorthis (Silurian). Morphological variability decreases through ontogeny but is not related to species abundance as predicted. Indeed, evidence suggests the opposite. Less abundant species are generally more variable, this is again the opposite of the predictions. There is no apparent relationship between morphological variability, eurytopy and stenotopy. These results considered with other data suggest that other Lower Palaeozoic animals, particularly suspension feeders, have similar niche requirements and these may be fairly broad.     

Quantitative handling of characters useful in snake systematics with particular reference to intraspecific variation in the Ringed Snake Natrix natrix (L.)

This paper is based on the numerous characters used in a biometric study of the intraspecific variation of Natrix natrix (L.). These characters are drawn from the internal morphology, scalation, colour pattern, body proportions, dentition and dermal sense organs. The criteria involved in selecting these characters for taxonomic work are considered.
The alternative methods for quantifying the above features are discussed and so are the theoretical problems involved in coding the continuous characters.
The value of the characters in studying the geographic variation, ontogenetic variation and sexual dimorphism in N. natrix is indicated.
The use of a wider range of characters than just the scalation and colour pattern is urged when diagnosing the lower taxonomic levels of ophidians. The neglected internal morphology may well be able to supply characters to this end.     

Craniofacial ontogeny in centrosaurine dinosaurs (Ornithischia: Geratopsidae): taxonomic and behavioral implications

Centrosaurine ceratopsians are characterized by well developed nasal horncores or bosses, relatively abbreviated supraorbital horncores or bosses, and adorned parietosquamosal frills. Recent study of several paucispecific (low diversity) bonebed assemblages in Alberta and Montana has contributed greatly to our understanding of ontogenetic and taxonomic variation in the skulls of centrosaurines. Relative age determination of centrosaurines is now possible through examination of ontogenetic change in several characters, including the surface bone morphology of specific skeletal elements. The within-group taxonomy of centrosaurines is based almost entirely on characters of the skull roof, relating particularly to horns and frills. Juvenile and sub-adult centrosaurines are characterized by relatively simple, unadorned skulls compared to their adult counterparts. As in numerous living taxa, the cranial ornaments of centrosaurines developed late in ontogeny, as individuals approached or attained adult size. An important implication arising directly from this study is that juvenile and sub-adult centrosaurines are difficult to distinguish taxonomically at the specific level. Two monospecific genera represented only by immature materials, Brachyceratops montanensis and Monoclonius crassus , cannot be defended and should be considered nomina dubia . The late ontogenetic development and diverse taxonomic variation of horn and frill morphologies support the contention diat these structures are best interpreted as reproductive characters employed in mate competition.     

Hundred species of the genus Peronopsis Hawle et Corda, 1847

The data on the genus Peronopsis, including 116 species and subspecies, that is, at least half of Middle Cambrian agnostians, are completely revised. The revision is based on the order of the formation of pygidial characters in ontogeny. A convenient taxonomic system of abundant species of this genus is proposed. It is shown that the sequence of the formation of characters in ontogeny generally corresponds to the chronological sequence of the appearance of species with appropriate morphology. The study of the genus Peronopsis resulted in a revision of the species composition of some other genera of the family Peronopsidae, clarification of the periods of existence of the majority of genera of this family. New diagnoses of genera and subgenera of the family Peronopsidae are provided.     

La Ferrassie 6 and the development of Neandertal pubic morphology

Functional interpretations of the mediolateral elongation and superoinferior flattening of the superior pubic ramus of Neandertals require knowledge of its ontogeny. Metric comparisons between the La Ferrassie 6 Neandertal infant, aged 3-5 years, and a sample of modern infants reveal that the acetabulosymphyseal length of La Ferrassie 6, relative to femoral length and iliac breadth, falls at the limits of the range of variation of the modern infants, while the relative height of its superior pubic ramus is indistinguishable from that of the modern sample. It appears that acetabulosymphyseal elongation of the Neandertal pubis is a feature which is expressed quite early in ontogeny, well before puberty. Superoinferior flattening and ventral margin thinning of the superior ramus probably appear later in Neandertal ontogeny.     

Ontogenetic variation in the mandibular ramus of great apes and humans

Considerable variation exists in mandibular ramus form among primates, particularly great apes and humans. Recent analyses of adult ramal morphology have suggested that features on the ramus, especially the coronoid process and sigmoid notch, can be treated as phylogenetic characters that can be used to reconstruct relationships among great ape and fossil hominin taxa. Others have contended that ramal morphology is more influenced by function than phylogeny. In addition, it remains unclear how ontogeny of the ramus contributes to adult variation in great apes and humans. Specifically, it is unclear whether differences among adults appear early and are maintained throughout ontogeny, or if these differences appear, or are enhanced, during later development. To address these questions, the present study examined a broad ontogenetic sample of great apes and humans using two‐dimensional geometric morphometric analysis. Variation within and among species was summarized using principal component and thin plate spline analyses, and Procrustes distances and discriminant function analyses were used to statistically compare species and age classes. Results suggest that morphological differences among species in ramal morphology appear early in ontogeny and persist into adulthood. Morphological differences among adults are particularly pronounced in the height and angulation of the coronoid process, the depth and anteroposterior length of the sigmoid notch, and the inclination of the ramus. In all taxa, the ascending ramus of the youngest specimens is more posteriorly inclined in relation to the occlusal plane, shifting to become more upright in adults. These results suggest that, although there are likely functional influences over the form of the coronoid process and ramus, the morphology of this region can be profitably used to differentiate among great apes, modern humans, and fossil hominid taxa. J. Morphol. 275:661–677, 2014. © 2013 Wiley Periodicals, Inc.     

Patterns of intraspecific variation through ontogeny: a case study of the Cretaceous nautilid Eutrephoceras dekayi and modern Nautilus pompilius

The magnitude and ontogenetic patterns of intraspecific variation can provide important insights into the evolution and development of organisms. Understanding the intraspecific variation of organisms is also a key to correctly pursuing studies in major fields of palaeontology. However, intraspecific variation has been largely overlooked in ectocochleate cephalopods, particularly nautilids. Furthermore, little is known regarding the evolutionary pattern. Here, we present morphological data for the Cretaceous nautilid Eutrephoceras dekayi (Morton) and the modern nautilid Nautilus pompilius Linnaeus through ontogeny. The data are used to describe conch morphology and to elucidate the evolutionary patterns of intraspecific variation. We discovered a similar overall pattern of growth trajectories and the presence of morphological changes at hatching and maturity in both taxa. We also found that intraspecific variation is higher in earlier ontogeny than in later ontogeny in both taxa. The high variation in earlier ontogeny may imply increased flexibility in changing the timing of developmental events, which probably played an important role in nautilid evolution. We assume that the decrease in variation in later ontogeny reflects developmental constraints. Lastly, we compared the similarity/dissimilarity of ontogenetic patterns of variation between taxa. Results reveal that the similarity/dissimilarity of the ontogenetic pattern differs between E. dekayi and N. pompilius. We conclude that this shift in the ontogenetic pattern of variation may be rooted in changes in the developmental programme of nautilids through time. We propose that studying ontogenetic patterns of intraspecific variation can provide new insights into the evolution and development of organisms.     

Tests for the presence of gametoclonal variation in barley and wheat doubled haploids produced using the Hordeum bulbosum system

Summary To investigate whether the Hordeum bulbosum system of doubled haploid production generates gametoclonal variation, populations of second generation doubled haploid lines were developed from first generation doubled haploid lines of two barley varieties and three wheat genotypes. In barley, no variation between doubled haploids from doubled haploids was detected for a range of quantitative characters, suggesting the absence of any gametoclonal effects. However, the original selfed-seed stocks were shown to contain cryptic allelic variation for some of the characters investigated. In wheat, gametoclonal variation was detected for ear emergence time, plant height and yield, and its components for two out of the three genotypes investigated. The type and range of variation was similar to that reported from studies of somaclonal variation from immature embryos and gametoclonal variation from anther culture. Generally, the effects appeared to reduce the yield performance of individual lines. The difference in response between the two species and the consequences for the use of the doubled haploid system in breeding programmes are discussed.