Epidermal differentiation in the developing scales of embryos of the Australian scincid lizard Lampropholis guichenoti

Formation of the first epidermal layers in the embryonic scales of the lizard Lampropholis guichenoti was studied by optical and electron microscopy. Morphogenesis of embryonic scales is similar to the general process in lizards, with well‐developed overlapping scales being differentiated before hatching. The narrow outer peridermis is torn and partially lost during scale morphogenesis. A second layer, probably homologous to the inner peridermis of other lizard species, but specialized to produce lipid‐like material, develops beneath the outer peridermis. Two or three lipogenic layers of this type develop in the forming outer surface of scales near to the hinge region. These layers form a structure here termed “sebaceous‐like secretory cells.” These cells secrete lipid‐like material into the interscale space so that the whole epidermis is eventually coated with it. This lipid‐like material may help to reduce friction and to reduce accumulation of dirt between adjacent extremely overlapping scales. At the end of their differentiation, the modified inner periderm turns into extremely thin cornified cells. The layer beneath the inner peridermis is granulated due to the accumulation of keratohyalin‐like granules, and forms a shedding complex with the oberhautchen, which develops beneath. Typically tilted spinulae of the oberhautchen are formed by the aggregation of tonofilaments into characteristically pointed cytoplasmic outgrowths. Initially, there is little accumulation of β‐keratin packets in these cells. During differentiation, the oberhautchen layer merges with cells of the β‐keratin layer produced underneath, so that a typical syncytial β‐keratin layer is eventually formed before hatching. Between one‐fourth distal and the scale tip, the dermis under epidermal cells is scarce or absent so that the mature scale tip is made of a solid rod of β‐keratinized cells. At the time of hatching, differentiation of a mesos layer is well advanced, and the epidermal histology of scales corresponds to Stage 5 of an adult shedding cycle. The present study confirms that the embryonic sequence of epidermal stratification observed in other species is basically maintained in L. guichenoti. J. Morphol. 241:139–152, 1999. © 1999 Wiley‐Liss, Inc.     

Epidermal differentiation during ontogeny and after hatching in the snake Liasis fuscus (Pythonidae,Serpentes, Reptilia), with emphasis on the formation of the shedding complex

Differentiation and localization of keratin in the epidermis during embryonic development and up to 3 months posthatching in the Australian water python, Liasis fuscus, was studied by ultrastructural and immunocytochemical methods. Scales arise from dome-like folds in the skin that produce tightly imbricating scales. The dermis of these scales is completely differentiated before any epidermal differentiation begins, with a loose dermis made of mesenchymal cells beneath the differentiating outer scale surface. At this stage (33) the embryo is still unpigmented and two layers of suprabasal cells contain abundant glycogen. At Stage 34 (beginning of pigmentation) the first layers of cells beneath the bilayered periderm (presumptive clear and oberhautchen layers) have not yet formed a shedding complex, within which prehatching shedding takes place. At Stage 35 the shedding complex, consisting of the clear and oberhautchen layers, is discernible. The clear layer contains a fine fibrous network that faces the underlying oberhautchen, where the spinulae initially contain a core of fibrous material and small beta-keratin packets. Differentiation continues at Stage 36 when the beta-layer forms and beta-keratin packets are deposited both on the fibrous core of the oberhautchen and within beta-cells. Mesos cells are produced from the germinal layer but remain undifferentiated. At Stage 37, before hatching, the beta-layer is compact, the mesos layer contains mesos granules, and cells of the alpha-layer are present but are not yet keratinized. They are still only partially differentiated a few hours after hatching, when a new shedding complex is forming underneath. Using antibodies against chick scale beta-keratin resolved at high magnification with immunofluorescent or immunogold conjugates, we offer the first molecular confirmation that in snakes only the oberhautchen component of the shedding complex and the underlying beta cells contain beta-keratin. Initially, there is little immunoreactivity in the small beta-packets of the oberhautchen, but it increases after fusion with the underlying cells to produce the syncytial beta layer. The beta-keratin packets coalesce with the tonofilaments, including those attached to desmosomes, which rapidly disappear in both oberhautchen and beta-cells as differentiation progresses. The labeling is low to absent in forming mesos-cells beneath the beta-layer. This study further supports the hypothesis that the shedding complex in lepidosaurian reptiles evolved after there was a segregation between alpha-keratogenic cells from beta-keratogenic cells during epidermal renewal.     

Scanning electron microscopy of changes in epidermal structure occurring during the shedding cycle in squamate reptiles

Previous studies of squamate epidermal structure have focused on either histology and ultrastructure or oberhautchen surface texture as revealed by scanning electron microscopy (SEM). Using SEM data drawn from a variety of lizard taxa (primarily iguanids, but also agamids, chamaeleonids, and scincids), as well as amphisbaenians and colubrid snakes, we relate the surfaces encountered in gross dissection of squamate skin to histologically identifiable layers, and characterize their surface structure. Only the oberhautchen bears the repeating pattern of ornamentation noted by previous authors. Because the clear layer is a perfect template of the oberhautchen surface, it is the only layer with which the oberhautchen might be confused. However, the clear layer can be identified by its tendency to curl and crack during preparation. All other surfaces encountered were relatively featureless, except for impressions left by dermal “papillae” associated with mechanoreceptors. Using a method for examining preserved specimens to determine the stage in the shedding cycle, we assess two sources of variation in epidermal surface structure: stage in the shedding cycle and wear. Examination of immature renewal-phase epidermis suggests that the oberhautchen does not mature synchronously across a single scale or across body regions. Comparing inner- and outer-generation oberhautchen in sheddingphase epidermis, we conclude that changes in surface appearance caused by natural wear fall into two categories: discrete scratches and accumulation of debris. We see no evidence of overall “buffing” on a microscopic level, though surface structure may be obscured by scratches and gouges. Many squamate taxa show a gradient from low relief surface structure on elevated regions such as keels to high relief patterns at scale edges. This gradient is not due to wear; its significance is unknown.     

Ultrastructure of the embryonic snake skin and putative role of histidine in the differentiation of the shedding complex.

The morphogenesis and ultrastructure of the epidermis of snake embryos were studied at progressive stages of development through hatching to determine the time and modality of differentiation of the shedding complex. Scales form as symmetric epidermal bumps that become slanted and eventually very overlapped. During the asymmetrization of the bumps, the basal cells of the forming outer surface of the scale become columnar, as in an epidermal placode, and accumulate glycogen. Small dermal condensations are sometimes seen and probably represent primordia of the axial dense dermis of the growing tip of scales. Deep, dense, and superficial loose dermal regions are formed when the epidermis is bilayered (periderm and basal epidermis) and undifferentiated. Glycogen and lipids decrease from basal cells to differentiating suprabasal cells. On the outer scale surface, beneath the peridermis, a layer containing dense granules and sparse 25-30-nm thick coarse filaments is formed. The underlying clear layer does not contain keratohyalin-like granules but has a rich cytoskeleton of intermediate filaments. Small denticles are formed and they interdigitate with the oberhautchen spinulae formed underneath. On the inner scale surface the clear layer contains dense granules, coarse filaments, and does not form denticles with the aspinulated oberhautchen. On the inner side surface the oberhautchen only forms occasional spinulae. The sloughing of the periderm and embryonic epidermis takes place in ovo 5-6 days before hatching. There follow beta-, mesos-, and alpha-layers, not yet mature before hatching. No resting period is present but a new generation is immediately produced so that at 6-10 h posthatching an inner generation and a new shedding complex are forming beneath the outer generation. The first shedding complex differentiates 10-11 days before hatching. In hatchlings 6-10 h old, tritiated histidine is taken up in the epidermis 4 h after injection and is found mainly in the shedding complex, especially in the apposed membranes of the clear layer and oberhautchen cells. This indicates that a histidine-rich protein is produced in preparation for shedding, as previously seen in lizard epidermis. The second shedding (first posthatching) takes place at 7-9 days posthatching. It is suggested that the shedding complex in lepidosaurian reptiles has evolved after the production of a histidine-rich protein and of a beta-keratin layer beneath the former alpha-layer.     

Observations on the histochemistry and ultrastructure of the epidermis of the tuatara,Sphenodon punctatus (Sphenodontida,Lepidosauria, Reptilia): a contribution to an understanding of the lepidosaurian epidermal generation and the evolutionary origin of the squamate shedding complex

Histochemical and TEM analysis of the epidermis of Sphenodon punctatus confirms previous histological studies showing that skin-shedding in this relic species involves the periodic production and loss of epidermal generations, as has been well documented in the related Squamata. The generations are basically similar to those that have been described in the latter, and their formation involves a cyclic alternation between beta- and alpha-keratogenesis. The six differences from the previously described squamate condition revealed by this study include: 1) the absence of a well-defined shedding complex; 2) the persistence of plasma membranes throughout the mature beta-layer, including the oberhautchen; 3) the concomitant presence of lipogenic lamellar bodies and PAS-positive mucous granules in most presumptive alpha-keratinizing cells; 4) the presence of the secreted contents of these organelles in the intercellular domains of the three derived tissues, the homologues of the squamate mesos, alpha-, and lacunar cells; 5) the paucity of lamellated lipid deposits in such domains; 6) the presence of keratohyalin-like granules (KHLG) in the presumptive lacunar, clear, and oberhautchen cells. In toto, the absence of many of the precisely definable, different pathways of cytogenesis discernible during squamate epidermal generation production might be interpreted as primitive for lepidosaurs. However, when the evolutionary significance of each of the six differences listed is evaluated separately, it becomes clear that the epidermis of S. punctatus possesses primitive amniote, shared and derived lepidosaurian, and some unique characters. This evaluation further elucidates the concept of a lepidosaurian epidermal generation as a derived manifestation of the sauropsid synapomorphy of vertical alternation of keratin synthesis and shows that further study of keratinocyte differentiation in the tuatara may contribute to our understanding of the origin and evolution of beta-keratinization in sauropsid amniotes.     

Histidine uptake in the epidermis of lizards and snakes in relation to the formation of the shedding complex

Mammalian epidermis utilizes histidine-rich proteins (filaggrins) to aggregate keratin filaments and form the stratum corneum. Little is known about the involvement of histidine-rich proteins during reptilian keratinization. The formation of the shedding complex in the epidermis of snakes and lizards, made of the clear and the oberhautchen layers, determines the cyclical epidermal sloughing. Differently from snakes, keratohyalin-like granules are present in the clear layer of lizards. The uptake of tritiated histidine into the epidermis of two lizards and one snake has been studied by autoradiography in sections at progressive post-injection periods. At 40 min and 1 hr post-injection keratohyalin-like granules were not or poorly labeled. At 3-22 hr post-injection most of the labeling was present over suprabasal cells destined to form the shedding complex, in keratohyalin-like granules of the clear layer, and in the forming a-layer but was low in the forming b-layer, and in superficial keratinized layers. The analysis of the shedding complex in the pad lamellae (a specialized scale used for climbing) of a gecko showed that the setae and the cytoplasm of clear cells among them are main sites of histidine uptake at 4 hr post-injection. In the snake most of the labeling at 4 hr post-injection was localized in the shedding complex along the boundary between the clear and oberhautchen layers. The present study suggests that, in the epidermis of lepidosaurian reptiles, the synthesis of a histidine-rich protein is involved in the formation of the shedding layer and, as in mammals, in a-keratinization.     

Ontogeny and homology of the skeletal elements that form the sucking disc of remoras (Teleostei,Echeneoidei, Echeneidae)

The sucking disc of the sharksuckers of the family Echeneidae is one of the most remarkable and most highly modified skeletal structures among vertebrates. We studied the development of the sucking disc based on a series of larval, juvenile, and adult echeneids ranging from 9.3 mm to 175 mm standard length. We revisited the question of the homology of the different skeletal parts that form the disc using an ontogenetic approach. We compared the initial stages of development of the disc with early developmental stages of the spinous dorsal fin in a representative of the morphologically basal percomorph Morone. We demonstrate that the “interneural rays” of echeneids are homologous with the proximal‐middle radials of Morone and other teleosts and that the “intercalary bones” of sharksuckers are homologous with the distal radials of Morone and other teleosts. The “intercalary bones” or distal radials develop a pair of large wing‐like lateral extensions in echeneids, not present in this form in any other teleost. Finally the “pectinated lamellae” are homologous with the fin spines of Morone and other acanthomorphs. The main part of each pectinated lamella is formed by bilateral extensions of the base of the fin spine just above its proximal tip, each of which develops a row of spinous projections, or spinules, along its posterior margin. The number of rows and the number of spinules increase with size, and they become autogenous from the body of the lamellae. We also provide a historical review of previous studies on the homology of the echeneid sucking disc and demonstrate that the most recent hypotheses, published in 2002, 2005 and 2006, are erroneous. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

Cartilage in the cloaca: Phallodeal spicules in Caecilians (Amphibia: Gymnophiona)

The presence of spicules (also termed spines, teeth, or denticles) on the intromittent organ, a character unique to male members of some species in the African caecilian family Scolecomorphidae, has long been known (Noble, '31; Taylor, '68; Wake,'72; Nussbaum,'85). However, their organization and structure has not been examined. Series of males of three species of Scolecomorphus were examined in order to determine the gross and histological organization of the spicules. Spicule morphology changes with ontogeny within species and differs grossly among species. The cellular organization of the spicules is remarkably similar among species, however. The spicules are composed of chondroid cartilage. The large discoid cells are arranged in “stacks” to form the projections, which emerge from an extensive base plate of the same cartilage. The spicules have a connective tissue sheath, which is not continuous with the cloacal epithelium. The sheath of the spicules may or may not be mineralized; the cartilage itself does not appear to be calcified. Mineralization apparently is not correlated with age, reproductive status, or season. Cartilage in the cloaca of these few species poses a series of questions about the interactions of developmental, structural, functional, and phylogenetic properties in the evolution of the reproductive biology of caecilians. J. Morphol. 237:177–186, 1998. © 1998 Wiley-Liss, Inc.     

Ontogeny of the complex sperm in the macrostomid flatworm Macrostomum lignano (Macrostomorpha,Rhabditophora)

Spermiogenesis in Macrostomum lignano (Macrostomorpha, Rhabditophora) is described using light‐ and electron microscopy of the successive stages in sperm development. Ovoid spermatids develop to highly complex, elongated sperm possessing an undulating distal (anterior) process (or “feeler”), bristles, and a proximal (posterior) brush. In particular, we present a detailed account of the morphology and ontogeny of the bristles, describing for the first time the formation of a highly specialized bristle complex consisting of several parts. This complex is ultimately reduced when sperm are mature. The implications of the development of this bristle complex on both sperm maturation and the evolution and function of the bristles are discussed. The assumed homology between bristles and flagellae questioned. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Prezygapophyseal articular facet shape in the catarrhine thoracolumbar vertebral column

Two contrasting patterns of lumbar vertebral morphology generally characterize anthropoids. “Long‐backed” monkeys are distinguished from “short‐backed” apes [Benton: The baboon in medical research, Vol. 2 (1967:201)] with respect to several vertebral features thought to afford greater spinal flexibility in the former and spinal rigidity in the latter. Yet, discussions of spinal mobility are lacking important functional insight that can be gained by analysis of the zygapophyses, the spine's synovial joints responsible for allowing and resisting intervertebral movements. Here, prezygapophyseal articular facet (PAF) shape in the thoracolumbar spine of Papio, Hylobates, Pongo, Gorilla, and Pan is evaluated in the context of the “long‐backed” versus “short‐backed” model. A three‐dimensional geometric morphometric approach is used to examine how PAF shape changes along the thoracolumbar vertebral column of each taxon and how PAF shape varies across taxa at corresponding vertebral levels. The thoracolumbar transition in PAF shape differs between Papio and the hominoids, between Hylobates and the great apes, and to a lesser extent, among great apes. At the level of the first lumbar vertebra, the PAF shape of Papio is distinguished from that of hominoids. At the level of the second lumbar vertebra, there is variation to some extent among all taxa. These findings suggest that morphological and functional distinctions in primate vertebral anatomy may be more complex than suggested by a “long‐backed” versus “short‐backed” dichotomy. Am J Phys Anthropol 142:600–612, 2010. © 2010 Wiley‐Liss, Inc.     

Three‐dimensional reconstruction of black tiger prawn (Penaeus monodon) spermatozoa using serial block‐face scanning electron microscopy

Serial Block‐Face Scanning Electron Microscopy (SBF‐SEM) was used in this study to examine the ultrastructural morphology of Penaeus monodon spermatozoa. SBF‐SEM provided a large dataset of sequential electron‐microscopic‐level images that facilitated comprehensive ultrastructural observations and three‐dimensional reconstructions of the sperm cell. Reconstruction divulged a nuclear region of the spermatophoral spermatozoon filled with decondensed chromatin but with two apparent levels of packaging density. In addition, the nuclear region contained, not only numerous filamentous chromatin elements with dense microregions, but also large centrally gathered granular masses. Analysis of the sperm cytoplasm revealed the presence of degenerated mitochondria and membrane‐less dense granules. A large electron‐lucent vesicle and “arch‐like” structures were apparent in the subacrosomal area, and an acrosomal core was found in the acrosomal vesicle. The spermatozoal spike arose from the inner membrane of the acrosomal vesicle, which was slightly bulbous in the middle region of the acrosomal vesicle, but then extended distally into a broad dense plate and to a sharp point proximally. This study has demonstrated that SBF‐SEM is a powerful technique for the 3D ultrastructural reconstruction of prawn spermatozoa, that will no doubt be informative for further studies of sperm assessment, reproductive pathology and the spermiocladistics of penaeid prawns, and other decapod crustaceans. J. Morphol. 277:565–574, 2016. © 2016 Wiley Periodicals, Inc.     

Functional morphology of amplexus (clasping) in spinicaudatan clam shrimps (Crustacea,Branchiopoda) and its evolution in bivalved branchiopods: A video‐based analysis

Male clam shrimps (Crustacea: Branchiopoda: Laevicaudata, Spinicaudata, and Cyclestherida) have their first one or two trunk limb pairs modified as “claspers,” which are used to hold the female during mating and mate guarding. Clasper morphology has traditionally been important for clam shrimp taxonomy and classification, but little is known about how the males actually use the claspers during amplexus (clasping). Homologies of the various clasper parts (“movable finger,” “large palp,” “palm,” “gripping area,” and “small palp”) have long been discussed between the three clam shrimp taxa, and studies have shown that only some structures are homologous while others are convergent (“partial homology”). We studied the clasper functionality in four spinicaudatan species using video recordings and scanning electron microscopy, and compared our results with other clam shrimp groups. General mating behavior and carapace morphology was also studied. Generally, spinicaudatan and laevicaudatan claspers function similarly despite some parts being nonhomologous. We mapped clasper morphology and functionality aspects on a branchiopod phylogeny. We suggest that the claspers of the three groups were adapted from an original, simpler clasper, each for a “stronger” grip on the female's carapace margin: 1) Spinicaudata have two clasper pairs bearing an elongated apical club/gripping area with one setal type; 2); Cyclestherida have one clasper pair with clusters of molariform setae on the gripping area and at the movable finger apex; and 3) Laevicaudata have one clasper pair, but have incorporated an additional limb portion into the clasper palm and bear a diverse set of setae. J. Morphol. 278:523–546, 2017. © 2017 Wiley Periodicals, Inc.     

Erratum: Darwin,race, and the AAPA 1997 Charles R. Darwin lifetime achievement award,acceptance address. Am. J. Phys. Anthropol. 104:553–555.

ERRATUM: Philip V. Tobias (1997) Darwin, Race, and the AAPA 1997 Charles R. Darwin lifetime achievement award, acceptance address. Am. J. Phys. Anthropol. 104:553–555. Page 553, column 2, paragraph 3, line 2: “analyses” should be “analysts.” Page 554, column 1, line 4: Darwin's “great black bug of the Pampas” is Triatoma infestans. Page 554, column 1, paragraph 3, line 2: “century” should read “centenary.” The AJPA apologizes for these and other errors that crept into the printed version of Philip V. Tobias's Acceptance Address. © 1998 Wiley-Liss, Inc.     

Corallite wall and septal microstructure in scleractinian reef corals: Comparison of molecular clades within the family Faviidae

Recent molecular phylogenies conflict with traditional scleractinian classification at ranks ranging from suborder to genus, challenging morphologists to discover new characters that better agree with molecular data. Such characters are essential for including fossils in analyses and tracing evolutionary patterns through geologic time. We examine the skeletal morphology of 36 species belonging to the traditional families Faviidae, Merulinidae, Pectiniidae, and Trachyphylliidae (3 Atlantic, 14 Indo‐Pacific, 2 cosmopolitan genera) at the macromorphological, micromorphological, and microstructural levels. Molecular analyses indicate that the families are not monophyletic groups, but consist of six family‐level clades, four of which are examined [clade XV = Diploastrea heliopora; clade XVI = Montastraea cavernosa; clade XVII (“Pacific faviids”) = Pacific faviids (part) + merulinids (part) + pectiniids (part) + M. annularis complex; clade XXI (“Atlantic faviids”) = Atlantic faviids (part) + Atlantic mussids]. Comparisons among molecular clades indicate that micromorphological and microstructural characters (singly and in combination) are clade diagnostic, but with two exceptions, macromorphologic characters are not. The septal teeth of “Atlantic faviids” are paddle‐shaped (strong secondary calcification axes) or blocky, whereas the septal teeth of “Pacific faviids” are spine‐shaped or multidirectional. Corallite walls in “Atlantic faviids” are usually septothecal, with occasional trabeculothecal elements; whereas corallite walls in “Pacific faviids” are usually trabeculothecal or parathecal or they contain abortive septa. Exceptions include subclades of “Pacific faviids” consisting of a) Caulastraea and Oulophyllia (strong secondary axes) and b) Cyphastrea (septothecal walls). Diploastrea has a diagnostic synapticulothecal wall and thick triangular teeth; Montastraea cavernosa is also distinct, possessing both “Pacific faviid” (abortive septa) and “Atlantic faviid” (paddle‐shaped teeth) attributes. The development of secondary axes is similar in traditional Atlantic faviids and mussids, supporting molecular results placing them in the same clade. Subclades of “Pacific faviids” reveal differences in wall structure and the arrangement and distinctiveness of centers of rapid accretion. J. Morphol. 272:66–88, 2011. © 2010 Wiley‐Liss, Inc.     

Presence of a glycine-cysteine-rich beta-protein in the oberhautchen layer of snake epidermis marks the formation of the shedding layer

The complex differentiation of snake epidermis largely depends on the variation in the production of glycine-cysteine-rich versus glycine-rich beta-proteins (beta-keratins) that are deposited on a framework of alpha-keratins. The knowledge of the amino acid sequences of beta-proteins in the snake Pantherophis guttatus has allowed the localization of a glycine-cysteine-rich beta-protein in the spinulated oberhautchen layer of the differentiating shedding complex before molting takes place. This protein decreases in the beta-layer and disappears in mesos and alpha-layers. Conversely, while the mRNA for a glycine-rich beta-protein is highly expressed in differentiating beta-cells, the immunolocalization for this protein is low in these cells. This discrepancy between expression and localization suggests that the epitope in glycine-rich beta-proteins is cleaved or modified by posttranslational processes that take place during the differentiation and maturation of the beta-layer. The present study suggests that among the numerous beta-proteins coded in the snake genome to produce epidermal layers with different textures, the glycine-cysteine-rich beta-protein marks the shedding complex formed between alpha- and beta-layers that allows for molting while its disappearance between the beta- and alpha-layers (mesos region for scale growth) is connected to the formation of the alpha-layers.     

Immunogold labeling shows that glycine‐cysteine‐rich beta‐proteins are deposited in the Oberhäutchen layer of snake epidermis in preparation to shedding

Shedding in snakes is cyclical and derives from the differentiation of an intraepidermal shedding complex made of two different layers, termed clear and Oberhäutchen that determine the separation between the outer from the inner epidermal generation that produces a molt. The present comparative immunocytochemical study on the epidermis and molts of different species of snakes shows that a glycine‐cysteine‐rich corneous beta‐protein in a snake is prevalently accumulated in cells of the Oberhäutchen layer and decreases in those of the beta‐layer. The protein is variably distributed in the mature beta‐layer of species representing some snake families when the beta‐layer merges with the Oberhäutchen but disappears in alpha‐layers. Therefore, this protein represents an early marker of the transition between the outer and the inner epidermal generations in the epidermis of snakes in general. It is hypothesized that specific gene activation for glycine‐cysteine‐rich corneous beta‐proteins occurs during the passage from the clear layer of the outer epidermal generation to the Oberhäutchen layer of the replacing inner epidermal generation. It is suggested that in the epidermis of most species glycine‐cysteine‐rich corneous beta‐proteins form part of the dense corneous material that rapidly accumulates in the differentiating Oberhäutchen cells but decreases in the following beta‐layer of the inner epidermal generation destined to be separated from the previous outer generation in the process of shedding. The regulation of the synthesis of these and other proteins is, therefore, crucial in timing the different stages of the shedding cycle in lepidosaurian reptiles. J. Morphol. 276:144–151, 2015. © 2014 Wiley Periodicals, Inc.     

Antennal morphology of the endogean carabid genus typhlocharis (Coleoptera: Carabidae: Anillini): Description of sensilla and taxonomic implications

The antennal morphology and chaetotaxy were studied in 52 species of the endogean carabid genus Typhlocharis, using scanning electron microscopy and light microscopy. The antennae are composed of 11 antennomeres (scape, pedicel, and nine flagellomeres). We found considerable variation between species in the third antennomere, with short‐stem and long‐stem forms, and flagellomere morphology, distinguishing two morphs: rounded (subovoid, subspheric and subquadrate, morph 1) and reniform shapes (morph 2). Antennal sensilla are grouped in six types of sensilla trichodea, three types of sensilla basiconica, one type of sensilla coeloconica, and one type of sensilla campaniformia. The distribution of sensilla along the antennomeres is described. The “rings” of trichoid sensilla in the antennomere body are affected by its shape and there is interspecific variation in the pattern of sensilla coeloconica in antennomere 11°, a novelty for the genus. The types of sensilla found in Typhlocharis are compared to those described in other Carabidae and the potential functionality and taxonomic interest of those variable antennal features are discussed. A correlation between the flagellomere morphology and the presence/absence of a stridulatory organ is suggested. The study also allowed comparing the observation of antennal features by SEM and light microscopy. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Linking cranial morphology to prey capture kinematics in three cleaner wrasses: Labroides dimidiatus,Larabicus quadrilineatus,and Thalassoma lutescens

Cleaner fishes are well known for removing and consuming ectoparasites off other taxa. Observers have noted that cleaners continuously “pick” ectoparasites from the bodies of their respective client organisms, but little is known about the kinematics of cleaning. While a recent study described the jaw morphology of cleaners as having small jaw‐closing muscles and weak bite forces, it is unknown how these traits translate into jaw movements during feeding to capture and remove ectoparasites embedded in their clients. Here, we describe cranial morphology and kinematic patterns of feeding for three species of cleaner wrasses. Through high‐speed videography of cleaner fishes feeding in two experimental treatments, we document prey capture kinematic profiles for Labroides dimidiatus, Larabicus quadrilineatus, and Thalassoma lutescens. Our results indicate that cleaning in labrids may be associated with the ability to perform low‐displacement, fast jaw movements that allow for rapid and multiple gape cycles on individually targeted items. Finally, while the feeding kinematics of cleaners show notable similarities to those of “picker” cyprinodontiforms, we find key differences in the timing of events. In fact, cleaners generally seem to be able to capture prey twice as fast as cyprinodontiforms. We thus suggest that the kinematic patterns exhibited by cleaners are indicative of picking behavior, but that “pickers” may be more kinematically diverse than previously thought. J. Morphol. 276:1377–1391, 2015. © 2015 Wiley Periodicals, Inc.     

Formation of large micro‐ornamentations in developing scales of agamine lizards

The embryonic scales of two Australian agamine lizards (Hypsilurus spinipes and Physignatus lesueuerii) derive from the undulation of the epidermis to form dome‐shaped scale anlage that later become asymmetric and produce keratinized layers. Glycogen is contained in basal and suprabasal cells of the forming outer scale surface that are destined to differentiate into β‐keratin cells. The outer peridermis is very flat, but the second epidermal layer, provisionally identified as an inner peridermis, is composed of large cells that accumulate vesicular bodies and a network of coarse filaments. The sequence of epidermal layers produced beneath the inner peridermis in these agamine lizards corresponds to that of previously studied lizards, but the first subperidermal layer has characteristics of both clear (keratohyalin‐like granules) and oberhautchen (dark β‐keratin packets) cells. This layer is here identified as an oberhautchen since it fuses with the underlying β‐keratinizing cells forming large spinulae as the entire tissue becomes syncytial so that the units appear to increase in size. These spinulae very likely represent sections of honeycomb‐shaped micro‐ornamentations. A mesos layer appears underneath the β‐layer before hatching. J. Morphol. 240:251–266, 1999. © 1999 Wiley‐Liss, Inc.     

Pelvic and hindlimb myology of the basal archosaur Poposaurus gracilis (archosauria: Poposauroidea)

The discovery of a largely complete and well preserved specimen of Poposaurus gracilis has provided the opportunity to generate the first phylogenetically based reconstruction of pelvic and hindlimb musculature of an extinct nondinosaurian archosaur. As in dinosaurs, multiple lineages of basal archosaurs convergently evolved parasagittally erect limbs. However, in contrast to the laterally projecting acetabulum, or “buttress erect” hip morphology of ornithodirans, basal archosaurs evolved a very different, ventrally projecting acetabulum, or “pillar erect” hip. Reconstruction of the pelvic and hindlimb musculotendinous system in a bipedal suchian archosaur clarifies how the anatomical transformations associated with the evolution of bipedalism in basal archosaurs differed from that of bipedal dinosaurs and birds. This reconstruction is based on the direct examination of the osteology and myology of phylogenetically relevant extant taxa in conjunction with osteological correlates from the skeleton of P. gracilis. This data set includes a series of inferences (presence/absence of a structure, number of components, and origin/insertion sites) regarding 26 individual muscles or muscle groups, three pelvic ligaments, and two connective tissue structures in the pelvis, hindlimb, and pes of P. gracilis. These data provide a foundation for subsequent examination of variation in myological orientation and function based on pelvic and hindlimb morphology, across the basal archosaur lineage leading to extant crocodilians. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc.