Associations among cranial traits

Multivariate analysis of associations among seven cranial characteristics originally presented by Hertzog ('68) suggests that the appearance of the circumparietal accessory ossicles, parietal foramina, and form of the fronto-temporal suture are largely independent of each other both within and across geographical races.     

Foregut morphology and ontogeny of the spider crab Maja brachydactyla (Brachyura,Majoidea, Majidae)

We describe the morphology of the foregut of the spider crab Maja brachydactyla Balss, 1922, from first larval stage to adult, with detailed stage‐specific documentation using light and scanning electron microscopy. A total of 40 ossicles have been identified in the foregut of adults of M. brachydactyla using Alizarin‐Red staining. The morphological pattern of the ossicles and gastric mill is very similar to other Majoidea species with only a few variations. The foregut of the zoeae stages appeared as a small and simple cavity, with a cardio‐pyloric valve that separates the stomach into cardiac and pyloric regions. The pyloric filter is present from the first zoea, in contrast to the brachyuran species which have an extended larval development. Calcified structures have been identified in the cardio‐pyloric valve and pyloric region of the zoeal stages. The most significant changes in foregut morphology take place after the metamorphosis from ZII to megalopa, including the occurrence of the gastric mill. In the megalopa stage, the foregut ossicles are recognizable by their organization and general morphology, but are different from the adult phase in shape and number. Moreover, the gastric teeth show important differences: the cusps of the lateral teeth are sharp (no molariform); the dorsal tooth have a small, dentate cusp (not a well‐developed quadrangular cusp); and the accessory teeth are composed of one sharp peak (instead of four sharp peaks). The gastric mill ontogeny from megalopa to adult reveals intermediate morphologies during the earlier juvenile stages. The relationship between gastric mill structures with food preferences and their contribution to the brachyuran phylogeny are briefly discussed. J. Morphol. 276:1109–1122, 2015. © 2015 Wiley Periodicals, Inc.     

喜马拉雅旱獭的洞穴特征及其生态意义

穴居性动物的洞穴对其生存有着重要的影响。为了解喜马拉雅旱獭（Marmota himalayana）利用洞穴特征及其生态意义,于2017年5月至10月以及2018年4月至5月在四川省阿坝州若尔盖县班佑乡多玛村对30对喜马拉雅旱獭繁殖对所利用的洞穴进行了定量研究。按照当年被频繁利用的洞穴（夜宿洞与繁殖洞）与临时利用的洞穴1∶2的比例取样,共测量了90个洞穴的7个物理特征参数,比较了不同类型洞穴的物理特征差异。结果表明：（1）当地喜马拉雅旱獭利用洞穴外径的长轴和短轴分别为59.3 ± 21.9 (16~100) 和45.8 ± 20.5 (18~100) cm,内径内横径和内竖径分别为24.4 ± 4.0 (16~39) 和19.7 ± 3.5 (15~39) cm;（2）洞道首段长为127.3 ± 43.3 (60~240) cm;（3）洞道首段倾斜角度为45.7° ± 9.5° (20°~72°);（4）洞口朝向角度为195.8° ± 96.2° (3°~356°)。除了夜宿洞与繁殖洞临时洞的内竖径有显著性差异,其它不同洞穴类型之间的物理特征基本一致。喜马拉雅旱獭利用洞穴物理特征参数表明其洞穴具有躲避天敌、抵御不良天气、良好的排水性能以及维持洞穴小气候稳定的功能。大部分洞穴之间的物理特征基本一致结合行为观察,说明喜马拉雅旱獭存在年际间的换洞行为。     

Arm joint articulations in the ophiuran brittlestars (Echinodermata: Ophiuroidea): a morphometric analysis of ontogenetic, serial, and interspecific variation

The ophiuroid arm contains a series of vertebral ossicles that form an articulated internal skeleton. Ontogenetic, serial, and interspecific variation in these skeletal elements are investigated using morphometric data from 35 species of brittle-stars (Order Ophiurae). Multiple ossicles were sampled from each individual and several individuals were sampled from each species to reconstruct serial and ontogenetic changes in vertebral morphology. Within species, ontogenetic and serial allometries are not statistically different. These data support 'Jackson's law of localized stages' (Jackson, 1899; Clark, 1914), which proposes that serial variation along the arm reflects ontogenetic stages of ossicle growth.
A multivariate analysis of interspecific variation shows two major vertebral forms: ossicles with a proximal depression and distal keel, and ossicles lacking these features. Variation within these groups is largely continuous, but individual species show distinct shape differences and unique allometric patterns of serial variation. These results suggest that vertebral ossicle variation among species can be described by: 1) variation in initial shape; and 2) variation in the allometric trajectory along the proximal-distal axis.
In all species, the most proximal ossicles within the disk show a non-keeled morphology. In species with keeled arm ossicles, however, there is an abrupt transition within the disk between non-keeled and keeled vertebral forms. A single ossicle, having features of both vertebral types, occurs at this site. The taxonomic distribution of the two vertebral forms and the anatomical transition between forms is discussed with reference to current classification systems and recent phylogenetic schemes for the Ophiuroidea.     

Functional morphology of synarthrial articulations in the crinoid stem

Synarthrial fulcral ridges are found in crinoid columnals from the mid-Ordovician to the present and in all four subclasses. Similar articulations did not become common in the cirri until the Mesozoic. Synarthrial stem articulations fall into two broad groups. Type I articulations have a fulcral ridge in the centre of the articular facet. In elliptical ossicles this ridge corresponds to either the long (IA) or short (IB) axis of the facet. Although both are functionally similar, Type IA ossicles are more common. Type II articulations have an excentric fulcral ridge, parallel to either the long (IIA) or short (IIB) axis of the articular facet. Type IIA articulations are found in crinoid stems capable of coiling. Type II articulations are particularly common in the cirri of articulates and are well adapted for attachment to hard and soft substrates. Columnals with Type I articulations often have divergent fulcra, giving the stem flexibility in all directions, but this feature is not seen in cirri or in coiled stems, where it would impair normal functions. Only the cirri of isocrinids and comatulids are muscular, so the movement of columns with fulcra must be passive.     

Acoustic communication and the evolution of hearing in fishes

Fishes have evolved a diversity of sound-generating organs and acoustic signals of various temporal and spectral content. Additionally, representatives of many teleost families such as otophysines, anabantoids, mormyrids and holocentrids possess accessory structures that enhance hearing abilities by acoustically coupling air-filled cavities to the inner ear. Contrary to the accessory hearing structures such as Weberian ossicles in otophysines and suprabranchial chambers in anabantoids, sonic organs do not occur in all members of these taxa. Comparison of audiograms among nine representatives of seven otophysan families from four orders revealed major differences in auditory sensitivity, especially at higher frequencies (> 1 kHz) where thresholds differed by up to 50 dB. These differences showed no apparent correspondence to the ability to produce sounds (vocal versus non-vocal species) or to the spectral content of species-specific sounds. In anabantoids, the lowest auditory thresholds were found in the blue gourami Trichogaster trichopterus, a species not thought to be vocal. Dominant frequencies of sounds corresponded with optimal hearing bandwidth in two out of three vocalizing species. Based on these results, it is concluded that the selective pressures involved in the evolution of accessory hearing structures and in the design of vocal signals were other than those serving to optimize acoustic communication.     

Expression of glycoproteins in the vomeronasal organ reveals a novel spatiotemporal pattern of sensory neurone maturation

The main olfactory and the accessory olfactory systems are both anatomically and functionally distinct chemosensory systems. The primary sensory neurones of the accessory olfactory system are sequestered in the vomeronasal organ (VNO), where they express pheromone receptors, which are unrelated to the odorant receptors expressed in the principal nasal cavity. We have identified a 240 kDa glycoprotein (VNO(240)) that is selectively expressed by sensory neurones in the VNO but not in the main olfactory neuroepithelium of mouse. VNO(240) is first expressed at embryonic day 20.5 by a small subpopulation of sensory neurones residing within the central region of the crescent-shaped VNO. Although VNO(240) was detected in neuronal perikarya at this age, it was not observed in the axons in the accessory olfactory bulb until postnatal day 3.5. This delayed appearance in the accessory olfactory bulb suggests that VNO(240) is involved in the functional maturation of VNO neurones rather than in axon growth and targeting to the bulb. During the first 2 postnatal weeks, the population of neurones expressing VNO(240) spread peripherally, and by adulthood all primary sensory neurones in the VNO appeared to be expressing this molecule. Similar patterns of expression were also observed for NOC-1, a previously characterized glycoform of the neural cell adhesion molecule NCAM. To date, differential expression of VNO-specific molecules has only been reported along the rostrocaudal axis or at different apical-basal levels in the neuroepithelium. This is the first demonstration of a centroperipheral wave of expression of molecules in the VNO. These results indicate that mechanisms controlling the molecular differentiation of VNO neurones must involve spatial cues organised, not only about orthogonal axes, but also about a centroperipheral axis. Moreover, expression about this centroperipheral axis also involves a temporal component because the subpopulation of neurones expressing VNO(240) and NOC-1 increases during postnatal maturation.     

TENDRILS OF PASSIFLORA FOETIDA: HISTOGENESIS AND MORPHOLOGY

Passiflora foetida bears an unbranched tendril, one or two laterally situated flowers, and one accessory vegetative bud in the axil of each leaf. The vegetative shoot apex has a single-layered tunica and an inner corpus. The degree of stratification in the peripheral meristem, the discreteness of the central meristem, and its centric and acentric position in the shoot apex are important plastochronic features. The procambium of the lateral leaf trace is close to the site of stipule initiation. The main axillary bud differentiates at the second node below the shoot apex. Adaxial to the bud 1–3 layers of cells form a shell-zone delimiting the bud meristem from the surrounding cells. A group of cells of the bud meristem adjacent to the axis later differentiates as an accessory bud. A second accessory bud also develops from the main bud opposite the previous one. A bud complex then consists of two laterally placed accessory bud primordia and a centrally-situated tendril bud primordium. The two accessory bud primordia differentiate into floral branches. During this development the initiation of a third vegetative accessory bud occurs on the axis just above the insertion of the tendril. This accessory bud develops into a vegetative branch and does not arise from the tissue of the tendril and adjacent two floral buds. The trace of the tendril bud consists of two procambial strands. There is a single strand for the floral branch trace. The tendril primordium grows by marked meristematic activity of its apical region and general intercalary growth.     

Extreme tadpoles: the morphology of the fossorial megophryid larva, Leptobrachella mjobergi

The bizarre larvae of Leptobrachella mjobergi are fossorial and live in the gravel beds of small streams. These tadpoles are vermiform in body shape. Here we present details on their skeleton and musculature, particularly of the head. The entire cranium and its associated musculature are reconstructed in three dimensions from serial histological sections. The hyobranchial apparatus is highly reduced. The head of the L. mjobergi larva is more mobile than in other anuran species. This mobility can largely be ascribed to the exclusion of the notochord from the cranial base and an articulation of the foramen magnum floor with the atlas of the tadpole. The articulation is unique among anuran species, but design parallels can be drawn to salamanders and the articulation between atlas and axis in mammals. In L. mjobergi, the atlas forms an anterior dens that articulates with the basal plate in an accessory, third occipital articular face. The muscle arrangements deviate from the patterns found in other tadpoles: For instance, epaxial and ventral trunk muscles reach far forward onto the skull. The post-cranial skeleton of L. mjobergi is considerably longer than that of other anurans: it comprises a total of 35 vertebrae, including more than 20 post-sacral perichordal centra. Despite a number of features in cranial and axial morphology of L. mjobergi, which appear to be adaptations to its fossorial mode of life, the species clearly shares other features with its megophryid and pelobatid relatives.     

The skeleton of the eye: II. Overlap of the scleral ossicles of the domestic fowl

The scleral ossicles, a ring of overlapping membrane bones, lie just outside the corneal margin in the eyes of domestic fowl. Eighty percent of the bony rings contain 14 bones; less than 1% have 13 bones; 19% have 15 bones; about 1% have 16 bones. Each bone is foreshadowed during development by a transient papilliform thickening in the overlying conjunctival epithelium. These conjunctival papillae appear on the eighth day of incubation and disappear on the twelfth day, when the corresponding preosseous membranes begin to ossify. Observations, and experiments involving the removal of specific papillae early or late in their maturation (in order to delete, or to reduce the size of, individual bones), demonstrated the following constraints on the morphogenesis of the scleral ossicular ring. (1) The number of ossicles is a function, not only of the number of papillae, but also of the distance between adjacent papillae; when two papillae lie close together, a single ossicle may arise beneath the pair. (2) There are three regions in the ring: nasal, dorsal (in both of which the bones overlap in one direction) and temporal (in which the bones overlap in the opposite direction). (3) The determinants of the direction of overlap between adjacent bones are extrinsic to the ossicles themselves and are distributed throughout each region, rather than confined to the discrete locations within each region where overlap normally occurs. (4) The three places in the ring where these regions meet are characterized by the loss (in 13-membered rings) or the addition (in 15- and 16-membered rings) of papillae and of their corresponding bones, and by transitions in the direction of overlap between the bones.     

The middle and inner ears of the Palaeogene golden mole Namachloris: A comparison with extant species

Many living species of golden moles (Chrysochloridae) have greatly enlarged middle ear ossicles, believed to be used in the detection of ground vibrations through inertial bone conduction. Other unusual features of chrysochlorids include internally coupled middle ear cavities and the loss of the tensor tympani muscle. Our understanding of the evolutionary history of these characteristics has been limited by the paucity of fossil evidence. In this article, we describe for the first time the exquisitely preserved middle and inner ears of Namachloris arenatans from the Palaeogene of Namibia, visualised using computed tomography, as well as ossicles attributed to this species. We compare the auditory region of this fossil golden mole, which evidently did not possess a hypertrophied malleus, to those of three extant species with similarly sized ear ossicles, Amblysomus hottentotus, Calcochloris obtusirostris, and Huetia leucorhinus. The auditory region of Namachloris shares many common features with the living species, including a pneumatized, trabeculated basicranium and lateral skull wall, arteries and nerves of the middle ear contained in bony tubes, a highly coiled cochlea, a secondary crus commune, and no identifiable canaliculus cochleae for the perilymphatic duct. However, Namachloris differs from extant golden moles in the apparent absence of a basicranial intercommunication between the right and left ears, the possession of a tensor tympani muscle and aspects of ossicular morphology. One Namachloris skull showed what may be pneumatization of some of the dorsal cranial bones, extending right around the brain. Although the ossicles are small in absolute terms, one of the Huetia leucorhinus specimens had a more prominent malleus head than the other. This potentially represents a previously unrecognised subspecific difference.     

The foregut-ossicle system of Dromia wilsoni, Dromia personata and Lauridromia intermedia (Decapoda, Brachyura, Dromiidae), studied with a new staining method

The ossicles of the cardiac and pyloric foregut of three species of decapods: Dromia wilsoni [Fulton and Grant, 1902], Dromia personata [Linnè, 1758] and Lauridromia intermedia [Lauri, 1906] are described and illustrated in detail. Five new ossicles are recognized based on a specific staining-method with Alizarin-Red. The ossicle terminology of the brachyuran foregut is revised now to include 37 ossicles. All described ossicles are documented by drawings and photographs. The two new cardiac ossicles, the prepterocardiac and the postpterocardiac ossicles are narrow ossicles associated with the pterocardiac ossicles at the antero-dorsal margin of the cardiac foregut. The pyloric foregut includes the newly described lateral mesopyloric- and the posterior uropyloric ossicles. These pyloric ossicles are clearly recognizable only in the stained condition. Based on our data we provide new additional evidence for brachyuran monophyly and the paraphyletic status of the podotrematan crabs.     

Skeletal morphology of the mud star,Ctenodiscus crispatus (Echinodermata: Asteroidea)

The morphology of the following eight major ossicle types is described and illustrated for the goniopectinid asteroid Ctenodiscus crispatus: terminal plates, superomarginal and inferomarginal ossicles, adambulacral and ambulacral ossicles, odontophores, oral intermediate plates, and superambulacral ossicles. Development, variation, and relationships with soft body-parts and with other ossicles are embphasized. Each ossicle type is distinguished by numerous structures related to its function and to articulation with adjoining skeletal elements. Because major structures (such as pustules, alveoli, and articulation surfaces) distinguishing ossicle types develop early during ontogeny, immature ossicles are readily identifiable. However, changes in form and orientation of these structures occur during ossicle growth. Ontogenetic changes are influenced by development of associated skeletal and soft parts. Ambulacral and adambulacral ossicles near the peristome are highly modified but retain the basic characteristics of structure and orientation which define these major types of skeletal elements.     

Structure,molting, and mineralization of the dorsal ossicle complex in the gastric mill of the blue crab,Callinectes sapidus

This study examined the mesocardiac and urocardiac ossicles in the gastric mill of the blue crab to describe its structure, mineralization, and dynamics throughout the molt cycle, and to assess its possible utility in age determination. Morphologically, the mineralized ossicles are similar to the calcified dorsal carapace having a lamellate structure comprised of sheets of chitin/protein fibrils. Staining with acridine orange showed the same arrangement of an epicuticle, exocuticle, and endocuticle. In much of the mesocardiac and urocardiac ossicles, the endocuticle is very reduced, with the exocuticle predominating; the reverse of the dimensions of the exoskeleton. The lamellate structure of the ossicles was confirmed with scanning electron microscopy; however, elemental mapping by energy‐dispersive analysis of X‐rays revealed that the ossicles are mineralized with calcium phosphate, in contrast to the calcium carbonate biomineral of the exoskeleton. The medial tooth of the urocardiac ossicle is not calcified, but the epicuticle is highly elaborated and impregnated with silica. Histological examination of the ossicles demonstrated that they are molted during ecdysis, so despite the appearance of bands in the mesocardiac ossicle, it is difficult to hypothesize how the bands could represent a record of chronological age. J. Morphol. 276:1358–1367, 2015. © 2015 Wiley Periodicals, Inc.     

The eye of the magellanic penguin (Spheniscus magellanicus): structure of the anterior segment

We undertook a light and scanning electron microscopic study of the eye in the Magellanic penguin (Spheniscus magellanicus). The anatomical peculiarities of the eyeball shape in Sphenisciformes have been previously described by others; here, we show that they are accompanied by several modifications in the organization of the anterior segment of the eye. The main change was found in the portion of opaque sclera extending from the cornea to the anterior border of the scleral ossicles, which was much broader than in other avian eyes. This scleral region was made of a very dense fibrous tissue and was as difficult to cut as the ossicles. The corneo-scleral boundary was also different from that of other birds, since the aqueous humor channel and the pectinate ligament were located 1.0-1.5 mm posterior to the cornea. The osseous ring was formed by 13 bones, including three pairs of over- and underplates. There was a single ciliary muscle, with meridionally oriented striated fibers. They were inserted on a circumference along the boundary between the fibrous sclera and the ossicles, far away from the wall of the aqueous humor channel. On their posterior end, the muscle fibers formed a tendinous structure attached to the inner surface of the sclera and to the outer surface of the ciliary body. Only short zonular fibrils were observed. These anatomical features are probably relevant for the adaptation of penguin eyes to vision on land and in the aquatic environment.     

The fine structure and development of calcified skeletal elements in the body wall of holothurian echinoderms

The calcareous ossicles and iron-rich calcified granules in the integument of sea cucumbers (Echinodermata: Holothuroida) have been examined by light and electron microscopy. Most ossicles are perforated, platelike structures that measure 100–500 μm long and consist of magnesium-rich calcite. The formation of ossicles occurs within multinucleated syncytia of sclerocytes that are situated in the dermal layer of the body wall. Sclerocytes possess fairly large numbers of mitochondria and are characteristically enveloped by an external lamina. Each ossicle arises as a minute rodlike spicule that branches and develops into a fenestrated skeletal element. When viewed by SEM, fully developed ossicles appear non-crystalline. Following decoration with synthetically grown calcite seeds, however, each ossicle reveals an ordered array of crystallites that seem to be aligned perpendicular to the ossicle's original plane of growth. Examinations of sectioned ossicles indicate that the mineral phase in each skeletal element lacks a well developed matrix of EDTA-insoluble organic substances. Ossicles first arise in the ciliated, lecithotrophic larvae of Eupentacta quinquesimita at about 1 week postfertilization. Two-week-old specimens adopt an epibenthic existence and subsequently become enveloped by imbricated ossicles that are secreted by sclerocytes resembling those found in adults. In molpadiid holothurians, the adult body wall contains numerous reddish-brown granules that are chiefly composed of iron-rich subunits. The dermal granules differ from calcitic ossicles in developing extracellularly in association with finely filamentous material.     

Developmental and morphological variation in the teleost craniofacial skeleton reveals an unusual mode of ossification

We investigated the morphology and development of the scleral ossicles within the eyes of three species from three basal teleost orders, namely, the alewife (Alosa pseudoharengus; Clupeiformes), the surface morph of the Mexican tetra (Astyanax mexicanus; Characiformes) and zebrafish (Danio rerio; Cypriniformes). Two morphologies, circular and elongated, and one variation, fused elements, were identified. Zebrafish have small circular ossicles, whereas the alewife and the Mexican tetra have elongated ossicles. Surprisingly in the Mexican tetra these elements fuse at one end forming a continuous element with an antero-ventral opening; this may be typical for the Order Characiformes. Regardless of morphology, the ossicles develop via unilateral perichondral ossification of the scleral cartilage from two centers opposite one another in the eye. This unilateral type of ossification, in which only the perichondrium furthest from the retina contributes to the ossicles, has not previously been reported in any vertebrate. Because either the perichondrium and/or an extension of the perichondrium can transform into the scleral ossicle, we refer to the transitional tissue as periskeletal. Although the functional significance of the different shaped ossicles is unclear, skeletal muscle attaches directly to these bones, implying voluntary control. The morphological and developmental variation of teleost scleral ossicles makes them an ideal system for determining the genetic basis underlying phenotypic variation as well as for studying mechanisms underlying osteogenic and chondrogenic processes in teleosts. These data support our previous finding that scleral ossicles in teleosts may not be homologous to those in other vertebrates, such as reptiles.     

Linear growth in spines from Acanthaster planci (L.) involving growth lines and periodic pigment bands

We report the first observations of a linear growth pattern in aboral spine ossicles of adult Acanthaster planci (L.). This is unlike the spine development of other echinoderms. Growth in aboral spine ossicles of A. planci is essentially by addition of stereom at the base and the spine's growth history is preserved along its length. There are numerous growth lines perpendicular to the long axis of the ossicle. These are clearly evident in longitudinal spine sections and apparently caused by frequent growth episodes. There are periodic pigment bands which are parallel to the growth lines and evident on the surface of the ossicle. Basal growth of the spine ossicle and the nature of the growth lines were confirmed by tetracycline staining. Size/frequency analyses of a population of A. planci from Davies Reef (GBR) found spine ossicle growth, but not body diameter growth, over the six month period between sampling dates. The additional pigment banding in spine ossicles of 4 individuals recaptured after 6 months suggests that pigment bands are laid down seasonally. If pigment band cyclicity is validated, it offers a simple method for ageing adults of A. planci in field populations.     

A probable homologue of the clavicle in the holostean fish Amia calva

The two pairs of appendages associated with the throat region in Amia are described.
The posterior serrated appcndage of Amia calva is considered a homologue of the clavicle. This theory is based on topographical similarity (a bony element situated in front of the ventral part of the cleithrum and overlapping the anterior margin of the adjoining part of the cleithrum), structural similarity (cellular bone with ridges bearing denticles arranged in distinct patterns), ontogenetic similarity (early ontogenctic ossification with other dements of the dermal shoulder girdle prior to the appearance of scales), and phylogenetic cvidcnce (presence of similar elements in the caturid Furo ).
Although the anterior serrated appendage of A. calva exhibits structural and topographical similarities to both scales and clavicles, the ontogenetic evidence favours an homology with scales.
The ossicles located dorsolateral and lateral to the anteroventral part of the cleithrum in Lepisosteus oculatus share structural and topographical similarities to both scales and clavicles. However, ontogenetic evidence indicates that these ossicles are scales rather than fragmented clavicles.     

A morphometric and comparative study of the malleus.

This study has attempted to research, in detail, the dimensions and form of the malleus, and to indicate possible differences with regard to race, sex and side of origin (left or right). The ossicles were obtained from 75 adult cadavers and the dimensions were determined with the aid of a reflection microscope. Clear statistically significant differences were found between the Negroid and Caucasoid races, as well as between the right and left ossicles. No meaningful differences were determined between male and female ossicles.