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The origin and early evolution of snakes has long been studied, but little research has focused on soft-tissue organs such as the brain. I report data from dissections and 3D reconstructions of the endocasts of diverse species, including the Cretaceous stem snake Dinilysia patagonica in order to provide a comparative evolutionary framework for the snake brain. Snakes are a special case among reptiles because the braincase almost entirely encloses the whole brain, so endocasts provide realistic representations of brain size and shape. Diversity of brain gross anatomy among snakes is remarkable, encompassing two major cerebrotypes occurring in surface-dwelling and burrowing species. The repeated acquisition of the burrowing cerebrotype in different and phylogenetically distant snake clades suggests that brain gross anatomy is surprisingly evolutionary labile in snakes. Brain gross anatomy and other features such as body size and the absence of any unequivocal osteological feature related to burrowing is interpreted as evidence that D. patagonica was surface-dwelling, and that at least some of the early history of snakes occurred above ground.
Snakes have distinct body plans that can be traced to the origin of the clade. It remains unresolved whether ancestral snakes were adapted to terrestrial environments as burrowers, or to marine environments as swimmers. Recently, new approaches have been used to infer fossorial and aquatic specialists in the early evolution of snakes, using virtual CT models of the ear of fossils. This chapter reviews variation in the osseous part of the ear of major snake lineages. Vestibules are relatively large in fossorial species and small in aquatic snakes. Using quantitative analyses of bony labyrinth geometry, it has been suggested that putative stem snakes, such as Dinilysia patagonica, were fossorial. Improvements to testing correlations between bony labyrinth morphology and ecology can be made in the refinement of quantitative approaches to capturing and analysing shape variations, as well as better classifications of ecology. Using inner and middle ear morphology to improve the accuracy and precision of inferences of the ecology of the ancestral snake will depend also upon robust, well-resolved phylogenies for extinct and extant taxa, and denser taxonomic and ecomorphological sampling.
This chapter assesses morphological characters proposed to support the Pythonomorph Hypothesis— a purported close relationship between snakes and mosasaurians. With an emphasis on early diverging (non-mosasaurid) mosasaurians and mosaurids, new morphological data (including from high-resolution CT) for well-preserved dolichosaurid and mosasaurid fossils are presented. Details of the skull and mandibles are interpreted as supporting the monophyly of Mosasauria as the proximal outgroup of Varanoidea, to the exclusion of snakes. However, mosasaurians do deviate from the typical varanoid condition in aspects of their infraorbital foramen, ventral part of the lacrimal and its relationship with the prefrontal, anterior ramus of the ectopterygoid and its contact with the maxilla and jugal, lack of plicidentine, and (at least in early diverging mosasaurians) anguinoidean tooth replacement. We consider most characters previously reported as supporting the Pythonomorph Hypothesis to be problematic, because of incomplete fossil preparation, artefacts of taphonomy, limited comparisons, misinterpretations of anatomy, incomplete taxon sampling, or inadequate character formulation and/or scoring.
Mosasaurian phylogenetics has been one of the most controversial topics in squamate systematics, with various studies and authors arguing in favor of a varanoid affinity (the Varanoid Hypothesis), a snake affinity (the Pythonomorph and Ophidiomorph Hypotheses) or only distant affinities to these lineages (the Stem-scleroglossan Hypothesis). We review the classification history of mosasaurians over the past two centuries, focusing on non-mosasaurid mosasaurians (dolichosaurs and aigialosaurs). A reappraisal is provided based on a new phylogenetic analysis. Our results clearly support the Varanoid Hypothesis. The Pythonomorph and Ophidiomorph Hypotheses are reviewed, and characters traditionally inferred to support these hypotheses are discussed and reinterpreted. Taxonomic sampling and fossil completeness likely play a major role—our (hopefully improved) phylogenetic hypothesis being based on denser taxon sampling and more complete character scoring resulting from additional studies, including the application of modern imaging techniques. Based on current data, our interpretation is that a particularly close phylogenetic relationship between mosasaurians and snakes can be rejected.
Pachyophiidae fossils are among the most complete known for snakes, and include the earliest snakes with fully developed hindlimbs. Pachyophiids have been historically seen as suitable morphological intermediates between lizards and extant snakes, supporting the hypothesis that snakes originated in a marine setting from a macrophagous common ancestor with mosasaurian lizards. Pachyophiids have been subject to conflicting interpretations of their anatomy, fuelling renewed debate on snake origins and early diversification. We revisit pachyophiid cranial anatomy, providing additional evidence from new preparations, high resolution CT scans, and Synchrotron images. We address challenges posed by fossil (in)completeness to the study and interpretation of these specimens, and reassess phylogenetic affinities. We critically reassess morphological evidence supporting the Marine Hypothesis, concluding that (i) snakes are not especially closely related to mosasaurians, and (ii) pachyophiids are relatively deeply nested within the snake crown, so that they are of greater importance for understanding early crown-snake evolutionary history than they are for understanding snake origins.
Exposure to statistical patterns of language use affects language production and comprehension. In this longitudinal study of English language learner (ELL) university students, we examined the interplay between language experience and language statistics as a window into the formation and stability of morphological representations in memory. We hypothesized that within-participant change in sensitivity to distributional properties of complex words on written production would reflect changes in morphological knowledge. At two timepoints, separated by 8 months of language exposure, a sample of ELLs (n = 196) completed a written suffix completion task. The largest gains in production accuracy were observed for derived words ending in less productive suffixes. In addition, across both timepoints we found a consistent effect of derivational family entropy, such that derived words belonging to morphological families with equally dominant members were less accurately produced. Both effects indicate that ELLs exploit distributional cues to morphological structure and shed light on two aspects of morphological knowledge in ELLs. First, knowledge of suffixes becomes more entrenched in memory, independently of knowledge of the full forms of derived words. Second, ELLs draw upon interlexical connections between morphological family members during written word production.
Cruzia tentaculata is a helminth parasite of marsupials and has a wide geographic distribution from Mexico to Argentina. The aim of this study was to analyse the genetic population structure of this nematode along the Atlantic Forest biome. Cruzia tentaculata specimens were recovered from Didelphis aurita, Didelphis albiventris and Philander quica in 9 localities. Morphological and morphometric data were investigated for phenotypic diversity among localities and hosts using multivariate discriminant analysis of principal components. Phylogenetic relationships of C. tentaculata were determined using maximum likelihood and Bayesian inference. The population structure was analysed by fixation indices, molecular variance analysis, Tajima's D and Fu's Fs neutrality tests, Mantel tests and Bayesian clustering analysis. A higher significant morphometric difference for males was observed between localities. In the haplogroup networks, 2 groups were recovered, separating locations from the north and from the south/southeast. The morphometric variation in C. tentaculata between different localities was compatible with this north and southeast/south pattern, suggesting adaptation to different ecological conditions. Population genetic analyses suggested a pattern of evolutionary processes driven by Pleistocene glacial refugia in the northeast and southeast of the Atlantic Forest based on the distribution of genetic diversity.
Phonology/phonetics and morphology interact in such a way as to render difficult any clear-cut dividing line between these subfields. Romanists have long observed that phonetics play a crucial role in language change. From this point of view there is nothing exceptional in the fact that phonetics/phonology may provide the system with the very substance of morphological oppositions. The number and the extension of the morphological processes amenable to phonetic principles in the Romance domain are so wide that only a few typical phenomena are treated in this chapter. Furthermore, it is not always evident how one can establish the extent to which a given morphological alternation is phonologically driven, whether we are dealing with a purely phonological phenomenon or whether we should recognize some lexical conditioning in the choice of the allomorphs. The examples discussed (allomorphy in the definite article, subject clitics and affixes, possessives, and the nominal, verbal, and adjectival stems) show that the phonetic impulse for a morphological alternation may no longer be transparent ; in other cases, the trigger of a given pattern is no longer available or can only be identified following a process of diachronic reconstruction.
This chapter elaborates on two case studies in structural variation to illustrate how the comparison of closely related grammatical systems fuels research questions on general theoretical issues. Our first case study regards subject clitics in central Romance dialects. Subject clitics have been studied extensively over recent decades, but they still raise several questions concerning the nature of null subject languages. Analogously, there is a huge literature on the selection of perfective auxiliaries – the second case study in our chapter – and, as in the case of subject clitics, lesser-known non-standard dialects display a kaleidoscope of auxiliation options whose rationalization poses fascinating analytical challenges and yields insights into basic issues of linguistic theory. The core question raised by our case studies concerns the modelling of linguistic diversity: do the above phenomena result from a finite set of discrete parameters or emerge from random language-specific options? We argue that the otherwise ‘hyperastronomical’ number of possible grammars is aptly constrained by syntactic factors, although inflexional morphology – which syntax cannot control entirely – may have a role in the realization of specific auxiliary or subject clitic forms in each dialect and for each person.
This chapter uses data from a range of Romance languages to illustrate the different definitions of the notion of suppletion in the linguistic literature, and to offer a typology of suppletion (notable the difference between ‘incursive’ and phonologically induced suppletion). Suppletion may be most usefully viewed simply as an extreme contrast between unity of meaning, on the one hand, and disunity of the forms expressing that meaning, on the other. The typology and distribution of Romance suppletions is described, for example, from the numeral system, from the system of marking comparatives in adjectives, from the inflexional morphology of personal pronouns, from the inflexional morphology or verbs, nouns, and adjectives. While the Romance languages provide cross-linguistically typical illustrations of suppletion in its different manifestations, the Romance data are particularly thought-provoking with regard to, among other things, (i) the particular role of synonymy between lexemes in determining the emergence of incursive suppletion in diachrony; (ii) the role of existing abstract patterns of alternation in providing ‘templates’ for the paradigmatic distribution of suppletive alternants; and (iii) the role of phonological resemblance as a determinant of incursive suppletion.
This paper synthesizes evidence for the origin and spread of the Indo-European languages from three disciplines – genomic research, archaeology, and, especially, linguistics – to reassess the validity of the Anatolian and Steppe Hypotheses. Research on ancient DNA reveals a massive migration off the steppe c. 2500 BCE, providing exceptionally strong support for the Steppe hypothesis. However, intriguing questions remain, such as why ancient Greek and Indo-Iranian populations had a smaller proportion of steppe ancestry, and Anatolian apparently had none at all. Lexical and archaeological evidence for wheels and looms provides essential clues about the early separation of Anatolian from the Indo-European community and the late entrance of Greek into the Aegean area. Evidence from the morphologies of the Indo-European languages supports these findings: the morphological patterns of the Anatolian languages show clear archaism, implying earlier separation, while the morphologies of Indo-Iranian and Greek display an array of similarities pointing to relatively late areal contact. Both the lexical and the morphological evidence, then, alongside the genomic and archaeological record, suggests that the Steppe hypothesis offers a preferable solution. Ultimately, these conclusions demonstrate the need for more dynamic models of change, including considerations of contact, stratification, and cross-disciplinary approaches.
Debate around inflectional morphology in language acquisition has contrasted various rule- versus analogy-based approaches. This paper tests the rule-based Tolerance Principle (TP) against a new type of pattern in the acquisition of the possessive suffix -im in Northern East Cree. When possessed, each noun type either requires or disallows the suffix, which has a complex distribution throughout the lexicon. Using naturalistic video data from one adult and two children – Ani (2;01–4;03) and Daisy (3;08–5;10) – this paper presents two studies. Study 1 applies the TP to the input to extrapolate two possible sets of nested rules for -im and make predictions for child speech. Study 2 tests these predictions and finds that each child’s production of possessives over time is largely consistent with the predictions of the TP. This paper finds the TP can account for the acquisition of the possessive suffix and discusses implications for language science and Cree language communities.
The tongue is a fundamental organ in feeding, vocalization, and grooming. It is characterized by evolutionary adaptations reflected by diet, habitat, and function. Rodents are a very diverse mammalian order and the tongue's morphology varies in size, form, and presence of papillae. This work aimed to describe the morphological and ultrastructural aspects of the tongue of Spix's yellow-toothed cavy (Galea spixii, Wagler, 1831). Tongues of Spix's yellow-toothed cavies were analyzed with light microscopy, scanning, and transmission electron microscopy. The results showed that the tongue was divided into apex, body, and root. There were different types of papillae, such as vallate, foliate, laterally placed fungiform, fungiform, filiform, and robust filiform. The epithelium was organized into layers, including keratinized, granulous, spinous, and basal, below were lamina propria, and musculature, which evolved mucous and serous gland clusters. The tongue of Spix's yellow-toothed cavy was structurally and ultrastructurally similar to other rodents and had papillae with similar morphologies to other Caviidae species. However, the presence of robust filiform papillary lines and laterally placed fungiform papillae showed the main differences from other species. This was the first description of the tongue of Spix's yellow-toothed cavy.
Are the dimensions of morphological diversity dependent on the cognitive pathways for processing, storage, and learning of word structure, and if so, how? Conversely, are languages that differ in their morphological structure processed and learned in different ways? This volume examines the relationship between linguistic cognition and the morphological diversity found in the world’s languages. As the idea that domain-general cognitive processes and morphological typology are inextricably linked has moved into the mainstream of linguistics, the field has diversified conceptually and methodologically. This introduction to the volume offers an overview of conceptual issues that underpin the volume’s papers and some of the methodological trends they reflect. It thus serves as a roadmap for the papers that follow.
Morphological structures interact dynamically with lexical processing and storage, with the parameters of morphological typology being partly dependent on cognitive pathways for processing, storage and generalization of word structure, and vice versa. Bringing together a team of well-known scholars, this book examines the relationship between linguistic cognition and the morphological diversity found in the world's languages. It includes research from across linguistic and cognitive science sub-disciplines that looks at the nature of typological diversity and its relationship to cognition, touching on concepts such as complexity, interconnectedness within systems, and emergent organization. Chapters employ experimental, computational, corpus-based and theoretical methods to examine specific morphological phenomena, and an overview chapter provides a synthesis of major research trends, contextualizing work from different methodological and philosophical perspectives. Offering a novel perspective on how cognition contributes to our understanding of word structure, it is essential reading for psycholinguists, theoreticians, typologists, computational modelers and cognitive scientists.
The morphology of quaggas, their taxonomy, habitats, and behaviors are described. Photographs, illustrations, and taxidermy specimens are employed to examine representations and descriptions critically, for example, did quaggas have horse-like tails, and were Daniell and Harris correct in their illustrations of body striping? My analysis of taxidermy specimens shows sexual dimorphism: stallions were slightly smaller than mares, which is unlike the situation in other subspecies of plains zebras. Quaggas, named Equus quagga by both Boddaert and Gmelin, lived in a variety of habitats in the Cape Colony and Orange Free State (areas that are now part of the Western Cape, Eastern Cape, Northern Cape, and the Free State provinces) including: the nama Karoo, succulent Karoo, fynbos, Albany thicket, savanna, and grassland biomes. A quagga breeding group consisting of a stallion, several mares, and their foals often joined other breeding groups to form a herd. Quaggas often grazed with wildebeests (gnus) and ostriches, and all three species probably benefitted from this association; these animals often migrated together in large numbers seeking grazing and water.
It is often assumed that pre-schoolers learn a second language (L2) with ease, even for structures that are absent in their L1, such as Mandarin-speaking pre-schoolers learning L2 English grammatical inflections (e.g., ducks, horses). However, while the results from Study 1 showed that such learners can imitate plural words (age = 3;5, N = 20), Studies 2 and 3 showed that they cannot yet generate or comprehend plural morphology (Study 2: age = 4;8, N = 20; Study 3: age = 4;1, N = 20), raising questions about when this is achieved. These findings have important implications for school readiness, as well as for identifying those at risk of developmental language disorders.
This article aims to examine to what extent English and Jordanian Arabic (JA) have the same classification of N + N compounds based on their degree of compositionality. It also attempts to propose a universally applicable classification of compositionality in N + N compounds. I suggest a modified version of the degree of compositionality based on previous classifications by Fernando (1996), Dirven and Verspoor (1998), and Kavka (2009). The new classification is based on the semantic contribution of the head and the non-head to the meaning of the whole compound. After I have applied the new scale to the JA data, I argue that English and JA have compounds that exhibit the four degrees of compositionality; namely completely compositional, semi-compositional, semi non-compositional and completely non-compositional. The article concludes with some recommendations for future research.
An important goal for humanoid robots is to achieve fast, flexible and stable walking. In previous research, the structure and walking algorithms evolved separately, resulting in a slow evolution speed and lack of an initial design basis. This paper proposes comprehensively considering body morphology and walking patterns, exploring the relationship between them and their influence on the motion ability. The method parameterizes the body morphology and walking patterns. Then a response surface model is established to describe the complex relationship between these parameters and finally obtain the optimized parameters, which provides a reference for the structural design and gait generation.
It is true that most teachers have limited knowledge of how words work in English. Linguistics hasn’t been a feature of their own schooling or their teacher education, and you can’t teach what you don’t know. The good news is that it isn’t hard to build the knowledge – in fact, it’s fun. In this chapter we look more closely at the linguistic threads that contribute to the rich tapestry of each word: etymology, orthography, phonology and morphology.