Cranial muscles of the anurans leiopelma hochstetteri and ascaphus truei and the homologies of the mandibular adductors in lissamphibia and other gnathostomes

The frogs Ascaphus truei and Leiopelma hochstetteri are members of the most basal lineages of extant anurans. Their cranial muscles have not been previously described in full and are investigated here by dissection. Comparison of these taxa is used to review a controversy regarding the homologies of the jaw adductor muscles in Lissamphibia, to place these homologies in a wider gnathostome context, and to define features that may be useful for cladistic analysis of Anura. A new muscle is defined in Ascaphus and is designated m. levator anguli oris. The differences noted between Ascaphus and Leiopelma are in the penetration of the jaw adductor muscles by the mandibular nerve (V³). In the traditional view of this anatomy, the paths of the trigeminal nerve branches define homologous muscles. This scheme results in major differences among frogs, salamanders, and caecilians. The alternative view is that the topology of origins, insertions, and fiber directions are defining features, and the nerves penetrate the muscle mass in a variable way. The results given here support the latter view. A new model is proposed for Lissamphibia, whereby the adductor posterior (levator articularis) is a separate entity, and the rest of the adductor mass is configured around it as a folded sheet. This hypothesis is examined in other gnathostomes, including coelacanth and lungfish, and a possible sequence for the evolution of the jaw muscles is demonstrated. In this system, the main jaw adductor in teleost fish is not considered homologous with that of tetrapods. This hypothesis is consistent with available data on the domain of expression of the homeobox gene engrailed 2, which has previously not been considered indicative of homology. Terminology is discussed, and “adductor mandibulae” is preferred to “levator mandibulae” to align with usage in other gnathostomes. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.     

Cranial ontogeny of otariid seals

Over 2000 otariid skulls were measured for a morphometric study of cranial ontogeny in fur seals and sea lions. Few interspecific differences in cranial ontogeny were observed in the Otariidae, with only minor differences in rates of growth. Sexual dimorphism was significant in all otariids but was more apparent in the larger species. Female otariids of each species showed monophasic development in all characteristics, whereas males expressed monophasic growth for some characters and biphasic growth for others. Biphasic development in skulls of male otariids occurred well after physical maturity had been reached, usually at a suture index of 27. The rate of development varied between skull characters; components relating to the nervous system completed growth well before the rest of the skull, whereas those related to feeding, respiration and vocalisation developed in synchrony with the overall growth of total skull length. Sutures of the calvaria, or braincase, were the first to show partial closure and those uniting the facial bones were usually the last to fuse. As with the calvaria, the orbits and otic capsules developed quickly, suggesting a need for good hearing and vision early in independent postnatal life. Development of the rostral and palatal regions required significantly longer to complete growth.     

Occurrence and regional distribution of striated muscle fibers in the rat pineal gland

Summary In a total of 96 rat pineals studied 31 were found to contain striated muscle fibers or their precursors. The muscle fibers were most frequently present in the stalk region and more frequently found in the left than in the right hemisphere. Size measurements revealed that the lengths of pineal muscle cell nuclei differ only slightly from those of the sphincter muscle of the iris. However, the yellowish appearance of pineal muscle cell nuclei under darkfield investigation, a phenomenon observed in all muscular tissues of mesenchymal origin and connective tissue cells, may support the hypothesis that pineal musculature is of mesenchymal rather than ectodermal origin.Supported by a grant (Vo 135/4) of the Deutsche Forschungsgemeinschaft within the Schwerpunktprogramm Neuroendokrinologie     

The larval head anatomy of Rhyacophila (Rhyacophilidae) with discussion on mouthpart homology and the groundplan of Trichoptera

The external and internal features of the larval head of Rhyacophila fasciata (Trichoptera: Rhyacophilidae) were described in detail. Anatomical examinations were carried out using a multimethod approach including histology, scanning electron microscopy, confocal laser‐scanning microscopy, microcomputed tomography, and computer‐based three‐dimensional reconstructions. Additionally, the information on the larval head of Limnephilus flavicornis (Limnephilidae) and Hydropsyche angustipennis (Hydropsychidae) available in the literature were reinvestigated. These anatomical data were used to address major questions of homology and terminology, that is, the ventral closure of the head capsule, the sclerites, and appendages of labium and maxilla and their muscles. These topics were discussed by summarizing the main hypotheses present in the literature and a critical inclusion of new findings. Consequently, the inner lobe of the maxilla very likely represents the galea. The distal maxillary sclerite (palpifer) is an anatomical composite formation at least including dististipes and lacinia. Based on these homology hypotheses several potential groundplan features of the larval head of Trichoptera were reconstructed. The head of Rhyacophila shows several presumably plesiomorphic features as for instance the prognath orientation of the mouthparts, the well‐developed hypocranial bridge, the triangular submentum and eyes composed of seven stemmata. Derived features of Rhyacophila are the reduced antennae, the anterior directing of three stemmata and the shift of the tentorio‐stipital muscle to the mentum. J. Morphol. 276:1505–1524, 2015. © 2015 Wiley Periodicals, Inc.     

Classical taxonomy and modern methodology

Classical rotifer taxonomy and recent approaches are reviewed. -taxonomy (morphology) remains the most widely-used technique, and most prone to subjective errors. Computer technology, and genetic or biochemical methods are not widely available in areas of most need — those with developing limnology programs. Strict adherence to the International Code of Zoological Nomenclature, uniform treatment of ecotypic variation and polymorphism, communication with and between systematists, and establishment of a centralized data base are among the points discussed. A standardized nomenclature is proposed.     

Apodemes associated with limbs support serial homology of claws and jaws in onychophora (velvet worms)

Although the onychophoran jaw blades are believed to be derivatives of foot claws, serial homology of these structures has not been demonstrated. To shed light on the evolutionary origin of the onychophoran jaws, we searched for morphological landmarks and compared the internal and external anatomy of jaws and distal leg portions in representatives of the two major onychophoran subgroups, the Peripatidae and Peripatopsidae. Our data revealed hitherto unknown structures associated with the onychophoran limbs, such as a soft diastemal membrane separating the anterior and posterior portions of the inner jaw blade (present only in Peripatidae), apodemes associated with feet, an eversible dorsal sac at the basis of each foot claw, and a specific arrangement of musculature associated with the sclerotised claws, jaws and their apodemes. Specific correspondences in structure and position of apodemes support serial homology of claws and jaws, suggesting that the onychophoran jaw evolved from the distal portion rather than the entire limb in the last common ancestor of Onychophora. J. Morphol. 274:1180–1190, 2013. © 2013 Wiley Periodicals, Inc.     

Cranial variation in the European badger Meles meles (Carnivora,Mustelidae) in Scandinavia

Morphometric variation in 30 craniometric characters of 465 skulls of the European badgers (Meles meles) from across Europe was analysed. Multivariate analyses revealed that the populations from Norway, Sweden, and Finland differ from other European populations in having smaller skulls. The analyses also revealed significant differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ forms. On average, badgers from south‐west Norway were smaller than those of the remaining Fennoscandia. Morphological differences between the ‘south‐western Norwegian’ and ‘main Fennoscandian’ populations of M. meles suggest a possible in situ semisympatric divergence since the beginning of the Holocene warming, or a complex history of two groups involving at least two colonization routes. The small‐sized Scandinavian badgers may be close to the ancestral form that used to be widespread in Denmark and throughout Europe. The animals from south‐west Norway may instead be descendants of ancestors that were the first to penetrate the southern parts of the Scandinavian Peninsula. The ‘main Fennoscandian’ badgers are likely to have been the descendants of the second wave of recolonization of Scandinavia. Specifically, they might have colonized the Scandinavian Peninsula from the east after the last glaciation.     

Cranial and mandibular morphology of Middle Pleistocene cave bears (Ursus deningeri): implications for diet and evolution

ABSTRACT

Deninger’s bears (Ursus deningeri) have been studied less frequently than Ursus spelaeus s.l. Our objective is to present, for the first time, an analysis of the skull shape of U. deningeri.

Bear crania and mandibles were digitised with a Microscribe or CT-scanned and the surface models subsequently landmarked. The landmarks were chosen based on a compromise between functional morphology and sample size.

Results show that U. deningeri and U. spelaeus mandibles display very similar morphologies and allometric trajectories, both to each other and to Ailuropoda melanoleuca. It is inferred that masticatory adaptations to a herbivorous diet were already present in the Middle Pleistocene. U. deningeri displays a cranial morphology that is similar to that of U. spelaeus when comparing all species, but U. deningeri has a relatively narrower and dorsoventrally lower zygomatic arch than U. spelaeus, although the masticatory signal is less strong in the skull.

We observe intraspecific differences between different populations of U. deningeri, which could parallel the genetic diversity found in U. spelaeus. The intraspecific differences found within U. deningeri may be temporal and/or geographical in nature and could be related to the evolution of the Late Pleistocene cave bear, but this hypothesis remains to be tested.     

Cranial allometry and geographic variation in slow lorises (Nycticebus)

A series of 20 craniodental measurements was obtained for two sister taxa: Nycticebus coucang (common slow loris) and N. pygmaeus (pygmy slow loris). Multivariate analysis of variance was performed with adult data to describe patterns of subspecific and specific variation in this genus. The geometric mean of adult cranial dimensions was compared to field data on latitudinal coordinates for available specimens to investigate if size variation in Nycticebus is clinal in nature. Ontogenetic series for larger-bodied N. coucang and smaller-bodied N. pygmaeus were compared to test the hypothesis that species and subspecific variation in skull form results from the differential extension of common patterns of relative growth. A MANOVA provides independent support of Groves's [pp. 44–53 in Proceedings of the Third International Congress on Primatology, Vol. 1 (Basel: S. Karger), 1971] classification of Nycticebus into two species, with four subspecies in the common slow loris and one form of the pygmy slow loris. Within N. coucang, cranial proportions for all four subspecies are ontogenetically scaled, and size differentiation is mainly clinal (Bergmann's Rule). N. c. bengalensis represents the most northerly disposed and the largest form. N. c. javanicus represents the next-largest form and is located in a southerly direction the next-farthest away from the equator. N. c. coucang and N. c. menagensis are both equatorial; however, the latter subspecies is the smallest. A genetic basis for some of the taxonomic variation between N. c. coucang and N. c. menagensis is supported by such nonclinal variation in body size. Variation in the presence/absence of I² is not size-related but rather tracks geographic proximity and isolating factors which predate the most recent inundation of the Sunda Shelf. Although they inhabit a nonequatorial environment, pygmy slow lorises are the smallest of all Nycticebus. As N. pygmaeus is sympatric with N. c. bengalensis, the largest slow loris, it appears that the evolution of its smaller body size represents a case of character displacement. Unlike N. coucang, skull size becomes significantly smaller in more northern N. pygmaeus. This may also reflect character displacement between sympatric sister taxa underlain by a cline-dependent ecological factor which is marked in more northerly latitudes. On the other hand, the negative correlation between body size and latitude in N. pygmaeus could be due to the influence of nonprimate fauna, such as predators, which themselves evince a similar clinal pattern. Analyses of relative growth indicate that skull proportions in the two species of Nycticebus are ontogenetically scaled in two-thirds of the cases. All but one of the seven comparisons (interorbital breadth) which do not indicate ontogenetic scaling represent part of the masticatory complex. This likely reflects a reorganization of N. pygmaeus maxillomandibular proportions linked to smaller size and changes in diet. Am. J. Primatol. 45:225–243, 1998. © 1998 Wiley-Liss, Inc.     

Cranial shape variation and molecular phylogenetic structure of crested newts (Triturus cristatus superspecies: Caudata,Salamandridae) in the Balkans

In the present study, we investigated the degree of congruence between phylogeny, as inferred from mitochondrial (mt)DNA sequences, and cranium shape variation of crested newts (Triturus cristatus superspecies) in the Balkans. These newts belong to four phylogenetic clades defined by mtDNA analysis, and significantly differed in cranial shape. Allometry explained a high percentage of shape variation in crested newts. The clade‐specific allometric slopes significantly diverged for both the ventral cranium and dorsal cranium, indicating that differences in shape between clades could not be a simple consequence of their difference in size. The analysis of hierarchical and spatial variation showed similarity in the patterns of global and spatially localized hierarchical variation of cranial shape. We also found significant congruence between the pattern of cranial shape variation and molecular phylogeny. The differences in morphology of Triturus dobrogicus in comparison to other crested newt clades, including marked differences in cranium shape, is discussed in the context of the evolution and ecology of crested newts. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 348–360.     

Cranial anatomy of Shunosaurus, a basal sauropod dinosaur from the Middle Jurassic of China

Shunosaurus, from the Middle Jurassic of China, is probably the best‐known basal sauropod and is represented by several complete skeletons. It is unique among sauropods in having a small, bony club at the end of its tail. New skull material provides critical information about its anatomy, brain morphology, tooth replacement pattern, feeding habits and phylogenetic relationships. The skull is akinetic and monimostylic. The brain is relatively small, narrow and primitively designed. The tooth replacement pattern exhibits back to front replacement waves in alternating tooth position. The teeth are spatulate, stout and show well‐developed wear facets indicative of coarser plant food. Upper and lower tooth rows interdigitate and shear past each other. Tooth morphology, skull architecture, and neck posture indicate that Shunosaurus was adapted to ground feeding or low browsing. Shunosaurus exhibits the following cranial autapomorphies: emargination of the ventral margin of the jugal/quadratojugal bar behind the tooth row; postorbital contains a lateral pit; vomers do not participate in the formation of the choanae; pterygoid is extremely slender and small with a dorsal fossa; quadrate ramus of the pterygoid is forked; quadratojugal participates in the jaw articulation; tooth morphology is a combination of cylindrical and spatulate form; basipterygoid process is not wrapped by the caudal process of the pterygoid; trochlear nerve has two exits; occlusal level of the maxillary tooth row is convex downward, whereas that of the dentary is concave upward, acting like a pair of garden shears; dentary tooth count is 25 or more; and the replacing teeth invade the labial side of the functional teeth. Cranial characters among the basal sauropods are reviewed. As Shunosaurus is the earliest sauropod for which cranial remains are known, it occupies an important position phylogenetically, showing the modification of skull morphology from the prosauropod condition. Although the skull synapomorphies of Sauropoda are unknown at present, 27 cranial synapomorphies are known for the clade Eusauropoda. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 145?169.     

The fine structure of striated muscle fibres of tunica muscularis of the intestine in some teleosts

Summary The muscular layer of the fish oesophagus is made up exclusively of striated muscles, which form a circular and a longitudinal layer. The intestinal muscular layer of the tench, which is an exception among the vertebrates in this respect, consists of two layers of striated muscles and a layer of smooth ones. The organization of striated muscle fibres is described from the muscular layer of the alimentary canal in four species: the tench, crucian carp, gudgeon and stone-loach. The fibres of the oesophagus differ from each other in diameter, length of sarcomeres, glycogen content and, in the case of one species, organization of SR. On the basis of the organization of sarcomeres all the fibres examined must be counted among the slow-contracting ones (broad Z line). The sarcoplasmic reticulum is organized according to the Z type except for the fibres of the oesophageal muscles in which the SR represents the A-I type. Two types, en grappe and en plaque, of nerve endings have been observed. The presence of relatively large dense core vesicles seems to indicate their adrenergic origin.This work has been supported by the Polish Academy Sciences.     

Cranial anatomy of the stem salamander Kokartus honorarius (Amphibia: Caudata) from the Middle Jurassic of Kyrgyzstan

The cranial anatomy of the stem salamander Kokartus honorarius, from the Middle Jurassic of Kyrgyzstan, is redescribed in detail on the basis of all available material using conventional methods and high‐resolution computed microtomography. In contrast to previous interpretations, the skull is characterized by posteriorly positioned external nostrils, the presence of an internarial fenestra between the premaxillae, the maxillae are not shortened, the overlapping of the frontal by the nasal and the parietal by the frontal, a dentate vomer, a dentate pterygoid with a long, strongly arcuate and anteromedially orientated anterior process, the absence of a fontanelle between frontals and pineal pit on the ventral surface of the frontals, and an inferred unique attachment of the adductor mandibulae internus to the lateral border and ventral ridge of the frontals and parietals. Kokartus is closely related to Karaurus and both these taxa constitute the monophyletic taxon Karauridae, which forms the sister group of crown‐group salamanders (Urodela). Karaurids were neotenic forms and neoteny was probably the ancestral life history strategy of salamanders. The morphological analysis of Kokartus supports the current phylogenetic hypothesis that Permian amphibamids (including branchiosaurids) are the closest relatives of salamanders amongst known non‐lissamphibian temnospondyls. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 816–838.     

Morphological integration and serial homology: A case study of the cranium and anterior vertebrae in salamanders

Serial homology or the repetition of equivalent developmental units and their derivatives is a phenomenon encountered in a variety of organisms, with the vertebrate axial skeleton as one of the most notable examples. Serially homologous structures can be viewed as an appropriate model system for studying morphological integration and modularity, due to the strong impact of development on their covariation. Here, we explored the pattern of morphological integration of the cranium and the first three serially homologous structures (atlas, first, and second trunk vertebrae) in salamandrid salamanders, using micro-CT scanning and three-dimensional geometric morphometrics. We explored the integration between structures at static and evolutionary levels. Effects of allometry on patterns of modularity were also taken into account. At the static level (within species), we analyzed inter-individual variation in shape to detect functional modules and intra-individual variation to detect developmental modules. Significant integration (based on inter-individual variation) among all structures was detected and allometry is shown to be an important integrating factor. The pattern of intra-individual, asymmetric variation indicates statistically significant developmental integration between the cranium and the atlas and between the first two trunk vertebrae. At the evolutionary level (among species), the cranium, atlas, and trunk vertebrae separate as different modules. Our results show that morphological integration at the evolutionary level coincides with morphological and functional differentiation of the axial skeleton, allowing the more or less independent evolutionary changes of the cranial skeleton and the vertebral column, regardless of the relatively strong integration at the static level. The observed patterns of morphological integration differ across levels, indicating different impacts of developmental and phylogenetic constraints and functional demands.