Showing papers in "Journal of Morphology in 2007"
TL;DR: A new and robust view of jaw muscle homology is provided and the first synthesized nomenclature of sauropsid musculature is introduced using multiple lines of evidence, indicating multiple topological criteria are necessary for interpretations of soft‐tissue homology and warrant further investigation into character congruence and developmental connectivity.
Abstract: The homologies of jaw muscles among archosaurs and other sauropsids have been unclear, con- founding interpretation of adductor chamber morphology and evolution. Relevant topological patterns of muscles, nerves, and blood vessels were compared across a large sample of extant archosaurs (birds and crocodylians) and outgroups (e.g., lepidosaurs and turtles) to test the utility of positional criteria, such as the relative position of the trigeminal divisions, as predictors of jaw muscle homol- ogy. Anatomical structures were visualized using dissec- tion, sectioning, computed tomography (CT), and vascular injection. Data gathered provide a new and robust view of jaw muscle homology and introduce the first synthesized nomenclature of sauropsid musculature using multiple lines of evidence. Despite the great divergences in ce- phalic morphology among birds, crocodylians, and out- groups, several key sensory nerves (e.g., n. anguli oris, n. supraorbitalis, n. caudalis) and arteries proved useful for muscle identification, and vice versa. Extant crocodylians exhibit an apomorphic neuromuscular pattern counter to the trigeminal topological paradigm: the maxillary nerve runs medial, rather than lateral to M. pseudotemporalis superficialis. Alternative hypotheses of homology necessi- tate less parsimonious interpretations of changes in to- pology. Sensory branches to the rictus, external acoustic meatus, supraorbital region, and other cephalic regions suggest conservative dermatomes among reptiles. Differ- ent avian clades exhibit shifts in some muscle positions, but maintain the plesiomorphic, diapsid soft-tissue topo- logical pattern. Positional data suggest M. intramandibu- laris is merely the distal portion of M. pseudotemporalis separated by an intramuscular fibrocartilaginous sesa- moid. These adductor chamber patterns indicate multiple topological criteria are necessary for interpretations of soft-tissue homology and warrant further investigation into character congruence and developmental connectiv-
204 citations
TL;DR: The morphological differences observed are probably not caused by exploitative competition between the species, but rather appear attributable to the modification of the relative influence of sexual and natural selection on both sexes.
Abstract: Podarcis bocagei and P. carbonelli are two closely related lacertid species, very similar morphologi- cally and ecologically. We investigated sexual dimor- phism patterns presented by both species in allopatry and in sympatry. Sexual size and shape dimorphism pat- terns were analyzed using both multivariate and geo- metric morphometric techniques. Multivariate morpho- metrics revealed a marked sexual dimorphism in both species—males being larger with more robust habitus and females presenting a longer trunk. General patterns of sexual size dimorphism are not modified in sympatry, although there is evidence for some morphological change in male head size. The application of geometric morphometrics offered a more detailed image of head shape and revealed that males present a more developed tympanic area than do females, while females have a more rounded head. Differences in the degree of sexual shape dimorphism were detected in sympatry, but no consistent patterns were observed. From the results of the study, and based on previous knowledge on the pop- ulations studied, we conclude that the morphological dif- ferences observed are probably not caused by exploita- tive competition between the species, but rather appear attributable to the modification of the relative influence of sexual and natural selection on both sexes. J. Mor- phol. 268:152-165, 2007 2007 Wiley-Liss, Inc.
135 citations
TL;DR: Analysis of arm morphology from the standpoint of biomechanics suggests that the transverse musculature is responsible for elongation of the arms, the longitudinal musculatures are responsible for shortening, and the oblique muscle layers and associated connective tissues create torsion.
Abstract: The morphology of the musculature and connective tissues of the arms of Octopus bimaculoides was analyzed with light microscopy. We also studied O. briareus and O. digueti, which possess relatively more elongate and less elongate arms, respectively. The morphology of the arms was found to be remarkably uniform among species. The arms consist of a densely packed three-dimensional arrangement of muscle fibers and connective tissue fibers surrounding a central axial nerve cord. Three primary muscle fiber orientations were observed: 1) transverse muscle fibers oriented in planes perpendicular to the long axis of the arm; 2) longitudinal muscle fibers oriented parallel to the long axis; and 3) oblique muscle fibers arranged in helixes around the arm. The proportion of the arm cross section occupied by each of these muscle fiber groups (relative to the total cross sectional area of the musculature) remains constant along the length of the arm, even though the arm tapers from base to tip. A thin circular muscle layer wraps the arm musculature on the aboral side only. Much of this musculature has its origin and insertion on several robust connective tissue sheets including a layer surrounding the axial nerve cord and crossed-fiber connective tissue sheets located on the oral and the aboral sides of the arm. An additional thin layer of connective tissue wraps the arm musculature laterally and also serves as a site of origin and insertion of some of the muscle fibers. The fibers of the oral and aboral crossed-fiber connective tissue sheets are arranged oblique to the long axis of the arm with the same fiber angle as the oblique muscle layers that originate and insert on the sheets. The oblique muscle layers and the crossed-fiber connective tissue sheets thus form composite right- and left-handed helical fiber arrays. Analysis of arm morphology from the standpoint of biomechanics suggests that the transverse musculature is responsible for elongation of the arms, the longitudinal musculature is responsible for shortening, and the oblique muscle layers and associated connective tissues create torsion. Arm bending may involve unilateral contraction of longitudinal muscle bundles in combination with resistance to arm diameter increase due to contraction of the transverse musculature or passive stiffness of the arm tissues. The arms may also be bent by a combination of decrease in diameter due to contraction of the transverse musculature and maintenance of constant length on one side of the arm by unilateral activity of longitudinal muscle bundles. An increase in flexural stiffness of the arm may be achieved by cocontraction of the transverse and longitudinal muscle. Torsional stiffness may be increased by simultaneous contraction of both the right- and left-handed oblique muscle layers.
127 citations
TL;DR: It is suggested that active control of the swimbladder may not occur until after the formation of the two chambers and subsequent development and maturation of vasculature, musculature and innervation of these structures at about 28–30 dph.
Abstract: Many teleosts including zebrafish, Danio rerio, actively regulate buoyancy with a gas-filled swimbladder, the volume of which is controlled by autonomic reflexes acting on vascular, muscular, and secretory effectors. In this study, we investigated the morphological development of the zebrafish swimbladder together with its effectors and innervation. The swimbladder first formed as a single chamber, which inflated at 1-3 days posthatching (dph), 3.5-4 mm body length. Lateral nerves were already present as demonstrated by the antibody zn-12, and blood vessels had formed in parallel on the cranial aspect to supply blood to anastomotic capillary loops as demonstrated by Tie-2 antibody staining. Neuropeptide Y-(NPY-) like immunoreactive (LIR) fibers appeared early in the single-chambered stage, and vasoactive intestinal polypeptide (VIP)-LIR fibers and cell bodies developed by 10 dph (5 mm). By 18 dph (6 mm), the anterior chamber formed by evagination from the cranial end of the original chamber; both chambers then enlarged with the ductus communicans forming a constriction between them. The parallel blood vessels developed into an arteriovenous rete on the cranial aspect of the posterior chamber and this region was innervated by zn-12-reactive fibers. Tyrosine hydroxylase- (TH-), NPY-, and VIP-LIR fibers also innervated this area and the lateral posterior chamber. Innervation of the early anterior chamber was also demonstrated by VIP-LIR fibers. By 25-30 dph (8-9 mm), a band of smooth muscle formed in the lateral wall of the posterior chamber. Although gas in the swimbladder increased buoyancy of young larvae just after first inflation, our results suggest that active control of the swimbladder may not occur until after the formation of the two chambers and subsequent development and maturation of vasculature, musculature and innervation of these structures at about 28-30 dph.
121 citations
TL;DR: It is suggested that otolith variation between populations results predominantly from geographic separation and may be suitable for assessing taxonomy and diversity in other species‐rich groups like the atheriniforms and many perciforms because these groups display otolith Bauplans that are similar to those seen in killifishes.
Abstract: Systematic assignment of fossil otoliths is virtually always based on studies of otolith morphology and subsequent comparisons with otoliths from collections and/or literature. Although this usually represents a practical method, comparisons and subsequent evaluation may be biased by subjective criteria used in the individual descriptions. Quantitative morphometric studies focusing on variations in the otolith morphology of extant fishes have been conducted in fisheries research, mostly based on Fourier shape analysis and related methods. However, with regard to fossil otoliths, these approaches are generally not suitable, mainly due to preservation-related problems. Here we present a new approach for quantifying otolith variation between species and populations of killifish (cyprinodontiforms) in the genera Aphanius Nardo and †Prolebias Sauvage that can be used with both extant and fossil otoliths. Our new approach includes the definition of 10 variables from linear and angle measurements of an otolith and statistical analyses. Best results were obtained by presorting the otoliths into three groups based on sulcus shape (straight, bent, S-shaped). In this case, canonical discriminant analysis (CDA) with jackknifed cross-validation yielded an overall species classification success of 86–96%. The three groups based on sulcus shape separate according to zoogeographic patterns (i.e., Mediterranean Aphanius, Arabian Aphanius, European †Prolebias) and probably reflect phylogenetic lineages. Application of CDA to compare otolith variation between populations resulted in an overall classification success (jackknifed) of 33–83%. High levels of variation were observed for Aphanius dispar and †Prolebias malzi, but not for A. fasciatus and †P. weileri. We suggest that otolith variation between populations results predominantly from geographic separation. Combination of qualitative characters (sulcus morphology) with quantitative approaches (otolith morphometry) presents a new approach for obtaining a better understanding of the taxonomy, diversity, and zoogeography of both fossil and extant killifishes. Moreover, the method may also be suitable for assessing taxonomy and diversity in other species-rich groups like the atheriniforms and many perciforms because these groups display otolith Bauplans that are similar to those seen in killifishes. J. Morphol., 2007. © 2007 Wiley-Liss, Inc.
103 citations
TL;DR: The occurrence of fully‐differentiated neoselachian enameloids microstructure in Chlamydoselachus anguineus, a basal Squalean with teeth that are functionally “cladodont,” is evidence that triple‐layered enameloid microst structure was a preadaption to the cutting and gouging function of many neose lachian teeth, and thus may have played an integral role in the Mesozoic radiation of the neose
Abstract: A systematic SEM survey of tooth micro- structure in (primarily) fossil taxa spanning chon- drichthyan phylogeny demonstrates the presence of a su- perficial cap of single crystallite enameloid (SCE) on the teeth of several basal elasmobranchs, as well as on the tooth plates of Helodus (a basal holocephalan). This sug- gests that the epithelial-mesenchymal interactions required for the development of enameloid during odonto- genesis are plesiomorphic in chondrichthyans, and most likely in toothed gnathostomes, and provides phylogenetic support for the homology of chondrichthyan and actino- pterygian enameloid. Along the neoselachian stem, we see a crownward progression, possibly modulated by hetero- chrony, from a monolayer of SCE lacking microstructural differentiation to the complex triple-layered tooth ename- loid fabric of neoselachians. Finally, the occurrence of fully-differentiated neoselachian enameloid microstruc- ture (including compression-resistant tangle fibered enameloid and bending-resistant parallel fibered ename- loid) in Chlamydoselachus anguineus, a basal Squalean with teeth that are functionally ''cladodont,'' is evidence that triple-layered enameloid microstructure was a prea- daption to the cutting and gouging function of many neo- selachian teeth, and thus may have played an integral role in the Mesozoic radiation of the neoselachian crown group. J. Morphol. 268:33-49, 2007. ! 2006 Wiley-Liss, Inc.
92 citations
TL;DR: The rainbow trout, Oncorhynchus mykiss, is a salmoniform fish that spawns once per year, andipe females that had ovulated naturally, and those induced to ovulate using salmon gonadotropin‐releasing hormone were studied to determine whether follicles were forming at the time of spawning and to describe the process of folliculogenesis.
Abstract: The rainbow trout, Oncorhynchus mykiss (Walbaum, 1792), is a salmoniform fish that spawns once per year. Ripe females that had ovulated naturally, and those induced to ovulate using salmon gonadotropin-releasing hormone, were studied to determine whether follicles were forming at the time of spawning and to describe the process of folliculogenesis. After ovulation, the ovaries of postspawned rainbow trout were examined histologically, using the periodic acid-Schiff procedure, to stain basement membranes that subtend the germinal epithelium and to interpret and define the activity of the germinal epithelium. After spawning, the ovary contained a few ripe oocytes that did not ovulate, numerous primary growth oocytes including oocytes with cortical alveoli, and postovulatory follicles. The germinal epithelium was active in postspawned rainbow trout, as determined by the presence of numerous cell nests, composed of oogonia, mitotic oogonia, early diplotene oocytes, and prefollicle cells. Cell nests were separated from the stroma by a basement membrane continuous with that subtending the germinal epithelium. Furthermore, follicles containing primary growth oocytes were connected to the germinal epithelium; the basement membrane surrounding the follicle joined that of the germinal epithelium. After ovulation, the basement membrane of the postovulatory follicle was continuous with that of the germinal epithelium. We observed consistent separation of the follicle, composed of an oocyte and surrounding follicle cells, from the ovarian stroma by a basement membrane. The follicle is derived from the germinal epithelium. As with the germinal epithelium, follicle cells derived from it never contact those of the connective tissue stroma. As with epithelia, they are always separated from connective tissue by a basement membrane. J. Morphol., 2007. © 2007 Wiley-Liss, Inc.
80 citations
TL;DR: It appears reasonable to reconstruct the neck utilization and the feeding strategies of dinosaurs with long necks by deriving the flexibility of the neck from preserved vertebrae.
Abstract: The gross morphology and the flexibility along the neck of the ostrich (Struthio camelus) were examined using fresh tissue as well as neck skeletons. The results of the morphologic studies were compared with results from observations of living ostriches. The investigation was focused on differences in the morphology and the function between different sections of the neck. Additionally, the function of major dorsal neck ligaments was examined, including measurements of force-strain-relations. Comparative studies of giraffes (Giraffa camelopardalis) and camels (Camelus bactrianus) were conducted to find relations between the flexibility along the neck and the general feeding strategy. The examinations revealed that the neck of the ostrich can be divided into four sections with different functions. The first is the atlas-axis-complex which is responsible for torsion. The adjacent cranial section of the neck is flexible in dorsoventral and lateral directions but this part of the neck is usually kept straight at rest and during feeding. Dorsoventral flexibility is highest in the middle section of the neck, whereas the base of the neck is primarily used for lateral excursions of the neck. For giraffes and camels, the posture and utilization of the neck are also reflected in the flexibility of the neck. For all three species, it is possible to reconstruct the pattern of flexibility of the neck by using the neck skeletons alone. Therefore, it appears reasonable to reconstruct the neck utilization and the feeding strategies of dinosaurs with long necks by deriving the flexibility of the neck from preserved vertebrae. For Diplodocus carnegii the neck posture and the feeding strategy were reconstructed. Two neck regions, one around the 9th neck vertebra and the second at the base of the neck, indicate that Diplodocus, like the ostrich, adopted different neck postures. The neck was probably kept very low during feeding. During interruptions of the feeding, e.g., in an alert, the head could have been lifted in an economic way by raising the cranial section of the neck. During standing and locomotion the head was probably located well above the shoulders.
78 citations
TL;DR: Together the teeth, dental ligament, and jaws form an integrated system that may be beneficial to the feeding ecology of C. plagiosum, allowing for a diet that includes prey of varying hardness and elusiveness.
Abstract: The teeth of white-spotted bamboo sharks (Chiloscyllium plagiosum) are used to clutch soft-bodied prey and crush hard prey; however, the dual function is not evident from tooth morphology alone. Teeth exhibit characteristics that are in agreement with a clutching-type tooth morphology that is well suited for grasping and holding soft-bodied prey, but not for crushing hard prey. The dual role of this single tooth morphology is facilitated by features of the dental ligament and jaw joint. Tooth attachment is flexible and elastic, allowing movement in both sagittal and frontal planes. During prey capture spike-like tooth cusps pierce the flesh of soft prey, thereby preventing escape. When processing prey harder than the teeth can pierce the teeth passively depress, rotating inward towards the oral cavity such that the broader labial faces of the teeth are nearly parallel to the surface of the jaws and form a crushing surface. Movement into the depressed position increases the tooth surface area contacting prey and decreases the total stress applied to the tooth, thereby decreasing the risk of structural failure. This action is aided by a jaw joint that is ventrally offset from the occlusal planes of the jaws. The offset joint position allows many teeth to contact prey simultaneously and orients force vectors at contact points between the jaws and prey in a manner that shears or rolls prey between the jaws during a bite, thus, aiding in processing while reducing forward slip of hard prey from the mouth. Together the teeth, dental ligament, and jaws form an integrated system that may be beneficial to the feeding ecology of C. plagiosum, allowing for a diet that includes prey of varying hardness and elusiveness.
77 citations
TL;DR: Digital cranial endocasts from CT scans of a growth series of skulls of Monodelphis domestica, the gray short‐tailed opossum, are extracted to provide an anatomical description to document developmental changes in endocranial morphology and to examine ontogenetic and individual variation with respect to phylogenetic characters of endocrinial cavities that are known to be variable between different mammalian taxa.
Abstract: Intraspecific variation (eg, ontogenetic, individual, sexual dimorphic) is rarely examined among cranial endocasts (infillings of the braincase cavity) because of the difficulty in obtaining multiple specimens of a species, particularly fossil taxa We extracted digital cranial endocasts from CT scans of a growth series of skulls of Monodelphis domestica, the gray short-tailed opossum, as a preliminary assessment of the amount of intraspecific variation in mammalian endocranial morphology The goals of this study were 1) to provide an anatomical description to document developmental changes in endocranial morphology of M domestica and 2) to examine ontogenetic and individual variation with respect to phylogenetic characters of endocranial cavities that are known to be variable between different mammalian taxa In this study, "ontogenetic variation" refers to variation between specimens of different ages whereas "individual variation" (ie, polymorphism) is restricted to variation between specimens of comparable age Aside from size, changes in shape account for the greatest amount of morphological variation between the endocasts of different ages Endocast length, width, and volume increase with age for the growth series Relative olfactory bulb cast size increases with age in the growth series, but the relative size of the parafloccular casts shows a slight negative allometric trend through ontogeny More than one-third of the phylogenetic characters of the endocranial cavity we examined showed some sort of variation (ontogenetic, individual, or both) This suggests that although endocasts are potentially informative for systematics, both ontogenetic and individual variation affect how endocranial characters are scored for phylogenetic analysis Further studies such as this are necessary to determine the taxonomic extent of significant intraspecific variation of these endocranial characters
71 citations
TL;DR: The present findings may serve as a basis for revising the currently used avian nomenclature so that it will reflect homologies of muscles with their non‐avian counterparts.
Abstract: Homologies of muscles of the m. longissimus and m. iliocostalis groups in the dorsal and cervical regions, as well as those of the subvertebral muscles and mm. intercostales externi that continue from the dorsal into the cervical regions, in extant Diapsida are proposed based on detailed dissections and published accounts of lepidosaurs, crocodylians, and birds. The morphology of tendons and innervation patterns suggest that the avian "m. iliocostalis" in the dorsal region include the homologs of both m. longissimus and m. iliocostalis in non-avian diapsids. The conserved nature of the morphology of tendons in palaeognath birds also revealed that the avian mm. intertransversarii in the cervical region consist of muscles of the both m. longissimus and m. iliocostalis groups despite having been treated as a single series of muscles, and thus are not homologous with muscles of the same name in Lepidosauria or Crocodylia. The avian mm. inclusi that lie medial to mm. intertransversarii are homologous with mm. intercostales externi in Lepidosauria and mm. intercostales externi and m. scalenus combined in Crocodylia. Innervation patterns suggest that a muscle ("m. iliocostalis capitis") connecting the atlas rib and occiput in Crocodylia includes contributions from the subvertebral layer and m. cucullaris complex, and possibly m. iliocostalis as well. The present findings may serve as a basis for revising the currently used avian nomenclature so that it will reflect homologies of muscles with their non-avian counterparts.
TL;DR: Antler regeneration illustrates that extensive appendage regeneration in a postnatal mammal can be achieved by a developmental process that differs in several fundamental aspects from limb regeneration in urodeles.
Abstract: The periodic replacement of antlers is an exceptional regenerative process in mammals, which in general are unable to regenerate complete body appendages. Antler regeneration has traditionally been viewed as an epimorphic process closely resembling limb regeneration in urodele amphibians, and the terminology of the latter process has also been applied to antler regeneration. More recent studies, however, showed that, unlike urodele limb regeneration, antler regeneration does not involve cell dedifferentiation and the formation of a blastema from these dedifferentiated cells. Rather, these studies suggest that antler regeneration is a stem-cell-based process that depends on the periodic activation of, presumably neural-crest-derived, periosteal stem cells of the distal pedicle. The evidence for this hypothesis is reviewed and as a result, a new concept of antler regeneration as a process of stem-cell-based epimorphic regeneration is proposed that does not involve cell dedifferentiation or transdifferentiation. Antler regeneration illustrates that extensive appendage regeneration in a postnatal mammal can be achieved by a developmental process that differs in several fundamental aspects from limb regeneration in urodeles.
TL;DR: The results therefore do not provide any clues as to the origin of the chordate neural tube relative to the dorsal‐ventral orientation of the enteropneusts.
Abstract: Ripe specimens of Ptychodera flava were collected at Paiko Peninsula, Oahu, Hawaii, USA, and the development from egg to tornaria larva was followed in the laboratory. To complete the series, large tornaria larvae were collected from the plankton off the nearby Ala Moana Beach, and followed through metamorphosis to a juvenile stage with four pairs of gill slits. Ciliary band development was examined by scanning electron microscopy, and the development of the serotonergic nervous system was followed by means of immunostaining. The development of the apical tuft and neotroch (circumoral/perioral ciliary band) and their subsequent degeneration accorded fully with previous descriptions. A perianal ciliary ring of separate cilia develops just after hatching. This later develops a midventral extension, the neurotroch, extending to the neotroch posterior to the mouth. The cilia of this ring apparently beat diaplectically, with the effective stroke in the clockwise direction when seen from behind. An additional ring of cilia develops several days later anterior to the perianal ring. This opisthotroch (called telotroch by previous authors) consists at first of separate cilia, but later they became organized as large compound cilia. The apical tuft disappears after about a week, the neotroch degenerates at the transition to the Agassiz stage, and the opisthotroch degenerates just after metamorphosis. The serotonergic nervous system of the fully grown tornaria consists of an apical ganglion with many perikarya, a paired lateral group of perikarya on the postoral ciliary band, and scattered perikarya along the opisthotroch. Serotonergic processes are found along the ciliary bands except for the ventral and perianal ciliary bands and are scattered along the epidermis. At the Spengel stage and at metamorphosis (Agassiz stage), the processes along the ciliary bands are concentrated in the three ciliated food grooves so as to form three separate nerves, and are retained on the proboscis at least until 2–3 gill slit stage. No serotonergic processes were found to extend from the proboscis to the collar region, and no serotonergic neurons were observed in the collar cord or in the ventral nerve cord. Our results therefore do not provide any clues as to the origin of the chordate neural tube relative to the dorsal-ventral orientation of the enteropneusts. J. Morphol., 2007. © 2007 Wiley-Liss, Inc.
TL;DR: This evolutionary shift was accompanied by a loss of digit mobility and flexibility, and pedal posture was modified to better align the pes with the main direction of the ground reaction force, thus becoming well suited to support high loads.
Abstract: The evolution of ornithopod dinosaurs pro- vides a well-documented example of the transition from digitigrady to subunguligrady. During this transition, the ornithopod pes was drastically altered from the plesiomor- phic dinosaurian morphology (four digits, claw-shaped unguals, strongly concavo-convex joints, phalanges longer than wide, excavated collateral ligament fossae, presence of sagittal ridge, and prominent processes for the attach- ment of tendons) to a more derived condition (tridactyly, modification of the unguals into hooves, phalanges wider and thinner than long, lack of collateral ligament fossae, loss of sagittal ridge and tendon attachment processes, relatively flattened articular surfaces). These changes are particularly noteworthy given the overall conservatism in pedal morphology seen across Dinosauria. But what are the functional consequences of these specific morphologi- cal transitions? To study them, we examine aw ide range of pedal morphologies in four non-avian dinosaurs and two birds. Our analyses of the external morphology, two- dimensional models (using Finite Element Analysis), and internal bone structure demonstrate that this evolution- ary shift was accompanied by a loss of digit mobility and flexibility. In addition, pedal posture was modified to bet- ter align the pes with the main direction of the ground reaction force, thus becoming well suited to support high loads. These conclusions can be applied to other, parallel evolutionary changes (in both dinosaurs and mammals) that involved similar transitions to a subunguligrade pos-
TL;DR: The Emu showed a marked reduction in the number of muscles in the wing, even compared with other ratites, and relaxed selection on vestigial organs should allow more variation to persist in the population, and corresponds to what is observed here.
Abstract: Emus have reduced their wing skeleton to only a single functional digit, but the myological changes associated with this reduction have never been properly described. Moreover, the intraspecific variability associated with these changes has not previously been examined, dissections having been restricted in the past to only one or two individuals. In this paper, the myology and osteology of the Emu wing is described for a sample of five female birds. The Emu showed a marked reduction in the number of muscles in the wing, even compared with other ratites. Many wing muscles showed diversity in structure, origin and insertion sites, number of heads, as well as presence–absence variation. This variability dramatically exceeds that found in flying birds. Evolutionary theory predicts that relaxed selection on vestigial organs should allow more variation to persist in the population, and corresponds to what is observed here. A large amount of fluctuating asymmetry was also detected, indicating reduced canalization of the wing during development. J. Morphol. © 2007 Wiley-Liss, Inc.
TL;DR: A detailed description of the morphology and adult ontogeny of the skeleton of A. crepitans is provided, and novel morphologies, interesting postmetamorphic developmental patterns, and intraspecific skeletal variation are discussed.
Abstract: Acris crepitans is a small, semiaquatic member of the treefrog family Hylidae. Much recent attention has been paid to this species because of reports of population declines and malformations, yet few works have considered the skeletal anatomy of this common North American frog. Herein, we provide a detailed description of the morphology and adult ontogeny of the skeleton of A. crepitans, and discuss novel morphologies, interesting postmetamorphic developmental patterns, and intraspecific skeletal variation. The reduced amount of adult ossification, as well as several novel morphologies present in this species, are consistent with patterns of miniaturization seen in other anurans. For example, the skull is poorly ossified, but most of the cranial cartilages are heavily mineralized, the nasal bones are fused to endochondral ossification of the tectum nasi, the palatines are reduced, and the prootics and exoccipitals are not fused to one another (although the prootics are well-developed and ornamented). In addition, several specimens exhibit abnormalities, which might indicate that: (1) the population was under an acute malformation outbreak, (2) a high incidence of small skeletal malformations is normal in this species, (3) the population is under stress because of habitat fragmentation, (4) there is environmental deterioration in the region where the specimens were collected, and/or (5) the species is now showing signs of decline in southern Missouri. Regardless of the cause, it is clear that further examination of skeletal variability in A. crepitans, including ossification patterns and the frequency of abnormalities, is warranted.
TL;DR: This contribution presents the first comprehensive investigation of the gross morphology of the scorpion vascular system, based on a survey of species representing all major lineages of the order, using classical and modern non‐destructive techniques in combination with three‐dimensional reconstruction.
Abstract: Although scorpions are one of the better known groups of Arthropoda, detailed knowledge of their anatomy remains superficial. This contribution presents the first comprehensive investigation of the gross morphology of the scorpion vascular system, based on a survey of species representing all major lineages of the order, using classical and modern non-destructive techniques in combination with three-dimensional reconstruction. The investigation reveals that the hemolymph vascular system (HVS) of Scorpiones comprises a central pumping heart which extends the entire length of the mesosoma and is enclosed in a pericardium. Several arteries branch off the heart to supply different organs and body regions. Two different anterior aorta major branching patterns are identified among the species investigated. Arteries that branch off the anterior aorta system supply the appendages (chelicerae, pedipalps, and walking legs) and the central nerve mass with a complex arterial network. This study of the HVS of scorpions provides further evidence that the vascular systems of euarthropods can be highly complex. Use of the term "open circulatory system" within arthropods is re-emphasized, as it refers to the general organization of the body cavity (i.e. mixocoely) rather than to the complexity of the circulatory system.
TL;DR: Using a developmental series of the direct‐developing Philautus silus (Ranidae: Rhacophorinae) from Sri Lanka, features of cranial morphology that are part of a suite of adaptations that facilitate feeding in free‐living tadpoles, but have been changed or lost in other direct‐ developinging lineages are examined.
Abstract: Direct development has evolved in rhaco- phorine frogs independently from other anuran lineages, thereby offering an opportunity to assess features associ- ated with this derived life history. Using a developmen- tal series of the direct-developing Philautus silus (Ranidae: Rhacophorinae) from Sri Lanka, we examine features of cranial morphology that are part of a suite of adaptations that facilitate feeding in free-living tadpoles, but have been changed or lost in other direct-developing lineages. Larval-specific upper jaw cartilages, which are absent from many non-rhacophorine direct-developing species (such as Eleutherodactylus coqui), develop in embryos of P. silus. Similarly, lower jaw cartilages ini- tially assume a larval morphology, which is subsequently remodeled into the adult jaw configuration before hatch- ing. However, the cartilaginous jaw suspension and hyobranchial skeleton never assume a typical larval morphology. The palatoquadrate, which suspends the lower jaw, lacks the posterior connections to the brain- case found in many metamorphosing species. Unlike in metamorphosing species, bone formation in P. silus begins before hatching. However, the sequence of bone formation resembles that of metamorphosing anurans more than that of other direct developers. In particular, P. silus does not exhibit precocious ossification of the lower jaw, which is characteristic of some frogs and cae- cilians that lack a free-living tadpole. These data reveal some similarities between Philautus and other direct- developing anurans. However, the departure of Phi- lautus embryos from the generalized tadpole skeletal morphology is less pronounced than that observed in other direct-developing taxa. J. Morphol. 268:715-725, 2007. 2007 Wiley-Liss, Inc.
TL;DR: A comparative study identifies a number of distinct morphologies, some of which appear highly correlated with ecological niche, for example, inhabiting swift‐moving waters appears to be a key factor in the encapsulation of the anterior gas bladder in some cobitids, balitorids, and gobionin cyprinids.
Abstract: Cypriniformes (which includes the minnows, carps, loaches, algae-eaters, stone loaches, and suckers) is a morphologically diverse and incredibly speciose order of teleosts. It has been suggested that a number of evolutionary innovations, key to improved hearing and feeding, have played an important role in cypriniform fishes' success. One such innovation, the Weberian apparatus, is a novel assemblage of vertebral elements and modified ribs that relay and amplify sound pressure changes from the gas bladder to the inner ear. The Weberian apparatus unites Cypriniformes with other major orders into an extremely species-rich group of fishes, the Otophysi. Together, otophysan fishes comprise one of the largest groups of fishes in the world, as well as the majority of freshwater fishes. Here we present a detailed comparison of the Weberian apparatus in a number of cypriniform families using cleared and stained specimens. We present data regarding inter- and intrafamilial morphological variation within Cypriniformes. With few, but evolutionarily important, exceptions we find that diagnostic features of the Weberian apparatus characterize each family. Interspecific variation within each of the families Balitoridae, Gyrinocheilidae, and Catostomidae is only slight, whereas variation among subfamilies within Cyprinidae and Cobitidae is far more significant. This comparative study identifies a number of distinct morphologies, some of which appear highly correlated with ecological niche. For example, inhabiting swift-moving waters appears to be a key factor in the encapsulation of the anterior gas bladder in some cobitids, balitorids, and gobionin cyprinids. J. Morphol., 2007. © 2007 Wiley-Liss, Inc.
TL;DR: An updated discussion on the homologies and evolution of the osteichthyan pectoral muscles is provided, based on the authors' own analyses and on a survey of the literature, both old and recent.
Abstract: The Osteichthyes, including bony fishes and tetrapods, is a highly speciose group of vertebrates, comprising more than 42,000 living species. The anatomy of osteichthyans has been the subject of numerous comparative studies, but most of these studies concern osteological structures; much less attention has been paid to muscles. The most detailed comparative analyses of osteichthyan pectoral muscles that were actually based on a direct observation of representatives of various major actinopterygian and sarcopterygian groups were provided several decades ago by authors such as Howell and Romer. Despite the quality of their work, these authors did not have access to much information that is now available. In the present work, an updated discussion on the homologies and evolution of the osteichthyan pectoral muscles is provided, based on the authors' own analyses and on a survey of the literature, both old and recent. It is stressed that much caution should be taken when the results obtained in molecular and developmental studies concerning the pectoral muscles of model actinopterygians such as the teleostean zebrafish are discussed and compared with the results obtained in studies concerning model sarcopterygians from clades such as the Amphibia and/or the Amniota. This is because, as shown here, as a result of the different evolutionary routes followed within the actinopterygian and the sarcopterygian clades none of the individual muscles found, for example, in derived actinopterygians such as teleosts is found in derived sarcopterygians such as tetrapods. It is hoped that the information provided in the present work may help in paving the way for future analyses of the pectoral muscles in taxa from different osteichthyan groups and for a proper comparison between these muscles in those taxa.
TL;DR: This work contrasts wing loading, aspect ratio, and planform shape among five pelecaniform seabirds and correlate morphological diversity with known differences in flight strategies and found that the three boobies were more similar than either the tropicbird or the frigatebird.
Abstract: The selective pressures associated with flight are significant factors in shaping the morphology of volant forms. Tropical seabirds are of particular interest because of their long foraging bouts, which can last hundreds of kilometers in search of unpredictable (spatially and temporally) resources. Here, we contrast wing loading (WL), aspect ratio (AR), and planform shape among five pelecaniform seabirds and correlate morphological diversity with known differences in flight strategies. Overall, WL and AR scaled with body mass. The Great Frigratebird had lower WL than that predicted, whereas the Red-tailed Tropicbird had higher WL than that predicted. The tropicbird also exhibited a lower AR than that predicted. Visualization of planform shape was accomplished by using Thin-plate spline relative warp analysis (TPS/RWA), and three major regions of variations were discovered: wing base, mid-wing, and distal wing/wing tip. As expected, the three boobies were more similar than either the tropicbird or the frigatebird. The tropicbird had a broader distal wing and more rounded wing tip, associated with its greater use of flapping flight. The frigatebird showed the greatest deviation in the distal wing and wing tip associated with the high maneuverability required for aerial pursuit and kleptoparasitism. By using TPS/RWA, important differences were detected in planform shape that would have otherwise gone unnoticed when using only WL and AR. These differences correlated strongly with parameters such as maneuverability, flapping, and soaring flight.
TL;DR: Significant differences in bone‐density levels between normal and myostatin‐deficient mice, coupled with the multivariate differences in patterns of plasticity between the corpus and symphysis, underscore the need for a comprehensive analysis of the plasticity of masticatory tissues vis‐à‐vis altered mechanical loads.
Abstract: Compared with the normal or wild-type condition, knockout mice lacking myostatin (Mstn), a negative regulator of skeletal muscle growth, develop significant increases in relative masticatory muscle mass as well as the ability to generate higher maximal muscle forces. Wild-type and myostatin-deficient mice were compared to assess the postweaning influence of elevated masticatory loads because of increased jaw-adductor muscle and bite forces on the biomineralization of mandibular cortical bone and dental tissues. Microcomputed tomography (microCT) was used to quantify bone density at a series of equidistant external and internal sites in coronal sections for two symphysis and two corpus locations. Discriminant function analyses and nonparametric ANOVAs were used to characterize variation in biomineralization within and between loading cohorts. Multivariate analyses indicated that 95% of the myostatin-deficient mice and 95% of the normal mice could be distinguished based on biomineralization values at both symphysis and corpus sections. At the corpus, ANOVAs suggest that between-group differences are due to the tendency for cortical bone mineralization to be higher in myostatin-deficient mice, coupled with higher levels of dental biomineralization in normal mice. At the symphysis, ANOVAs indicate that between-group differences are related to significantly elevated bone-density levels along the articular surface and external cortical bone in the knockout mice. Both patterns, especially those for the symphysis, appear because of the postweaning effects of increased masticatory stresses in the knockout mice versus normal mice. The greater number of symphyseal differences suggest that bone along this jaw joint may be characterized by elevated plasticity. Significant differences in bone-density levels between normal and myostatin-deficient mice, coupled with the multivariate differences in patterns of plasticity between the corpus and symphysis, underscore the need for a comprehensive analysis of the plasticity of masticatory tissues vis-a-vis altered mechanical loads. J. Morphol., 2007. © 2007 Wiley-Liss, Inc.
TL;DR: The flow of nutrients through the digestive tract of Gryllus bimaculatus is regulated by the proventriculus, which effectively triturates the partially digested food coming from the crop and shoves the mushy nutrient mass into the space between the paired caeca.
Abstract: The flow of nutrients through the digestive tract of Gryllus bimaculatus is regulated by the proventriculus, which effectively triturates the partially digested food coming from the crop and shoves the mushy nutrient mass into the space between the paired caeca. The many folds at the base of the caeca form a sieve, and only fine food particles (4-10 microm) and fluids in the mush are filtered under pressure (produced by proventricular peristalsis) into the caeca. Combined with the release of enzymes in the caeca and the influx of water, the caeca are rapidly inflated on day 1 after the terminal molt. The remaining, mostly undigested food is shoved into a tube formed by the peritrophic membrane, which is first formed at the anterior end of the ventriculus. A mucous membrane (peritrophic gel) covers the caecal epithelium, and seems to merge with the true peritrophic membrane at the beginning of the ventriculus. The Type I peritrophic membrane is dragged posteriorly through the entire ventriculus and ileum by the posterior movement of the food bolus, which is shoved posteriorly at a rate of 6 mm/h by proventricular pressure. The growth rate of the peritrophic membrane is about 3 mm/h. Peristalsis does not occur in the midgut or ileum; the muscles in these regions function solely to counteract the internal pressure produced by the proventriculus. The exo- and endoperitrophic space in newly molted animals is open and fluids can flow in both directions. The endoperitrophic space becomes filled on day 1, and leads to a great reduction of the exoperitrophic space. In the ileal pouch (exoperitrophic space) the peritrophic membrane separates the mass of bacteria from the waste bolus within the endoperitrophic space. Feathery bristles arising from the cuticular covering of the finger-like invaginations of the ileal wall hold most of the bacterial mass in place. The crop weight decreases from day 1 to day 3 as the weight of caeca, ventriculus, and ileum increases. After day 3, food uptake and the weight of the entire gut system decrease in female crickets, partly in response to space restrictions in the abdomen caused by rapid ovarial growth.
TL;DR: The development of the osteocranium of the suckermouth armored catfish Ancistrus cf.
Abstract: The development of the osteocranium of the suckermouth armored catfish Ancistrus cf. triradiatus is described based on specimens ranging from prehatching stages to juvenile stages where the osteocranium is more or less fully formed. The first bony elements that arise are the opercle, jaws, and lateralmost branchiostegal rays, as well as the basioccipital and parasphenoid in the skull floor. The supracleithrum and the membranous and peri- chondral pterotic components form one large, double-lay- ered skull bone during ontogeny, without clear evidence of the involvement of a supratemporal. The Baudelot's liga- ment ossifies from two sides, i.e., from the basioccipital medially and the supracleithrum laterally. The lower jaw consists of a dentary, mentomeckelian, and angulo-articu- lar, which all soon fuse. The parurohyal, formed by the fusion of a ventral sesamoid bone and a dorsal cartilage element associated with the first basibranchial, is pierced by a vein, unlike in some other siluriforms. The interhyal cartilage disappears during ontogeny; medially of it, a small sesamoid bone appears in a ligament. The largest, canal-bearing cheek plate is not homologous to the intero- percle. The results of the present research, with emphasis on bone formations and homologies, are compared with studies on related catfishes. J. Morphol. 268:254-274, 2007. 2007 Wiley-Liss, Inc.
TL;DR: The main difference in homoscleromorph metamorphosis compared with Demospongiae is the persistence of the flagellated epithelium throughout this process and even in the adult since exo‐ and endopinacoderm remain flageLLated.
Abstract: The metamorphosis of the cinctoblastula of Homoscleromorpha is studied in five species belonging to three genera. The different steps of metamorphosis are similar in all species. The metamorphosis occurs by the invagination and involution of either the anterior epithelium or the posterior epithelium of the larva. During metamorphosis, morphogenetic polymorphism was observed, which has an individual character and does not depend on either external or species specific factors. In the rhagon, the development of the aquiferous system occurs only by epithelial morphogenesis and subsequent differentiation of cells. Mesohylar cells derive from flagellated cells after ingression. The formation of pinacoderm and choanoderm occurs by the differentiation of the larval flagellated epithelium. This is possibly due to the conservation of cell junctions in the external surface of the larval flagellated cells and of the basement membrane in their internal surface. The main difference in homoscleromorph metamorphosis compared with Demospongiae is the persistence of the flagellated epithelium throughout this process and even in the adult since exo- and endopinacoderm remain flagellated. The antero-posterior axis of the larva corresponds to the baso-apical axis of the adult in Homoscleromorpha.
TL;DR: Sonic morphology in the fawn cusk‐eel Lepophidium profundorum is investigated to determine morphology potentially associated with high frequency sound production and quantified development and sexual dimorphism of sonic structures.
Abstract: Recent reports of high frequency sound production by cusk-eels cannot be explained adequately by known mechanisms, i.e., a forced response driven by fast sonic muscles on the swimbladder. Time to complete a contraction-relaxation cycle places a ceiling on frequency and is unlikely to explain sounds with dominant frequencies above 1 kHz. We investigated sonic morphology in the fawn cusk-eel Lepophidium profundorum to determine morphology potentially associated with high frequency sound production and quantified development and sexual dimorphism of sonic structures. Unlike other sonic systems in fishes in which muscle relaxation is caused by internal pressure or swimbladder elasticity, this system utilizes antagonistic pairs of muscles: ventral and intermediate muscles pull the winglike process and swimbladder forward and pivot the neural arch (neural rocker) above the first vertebra backward. This action stretches a fenestra in the swimbladder wall and imparts strain energy to epineural ribs, tendons and ligaments connected to the anterior swimbladder. Relatively short antagonistic dorsal and dorsomedial muscles pull on the neural rocker, releasing strain energy, and use a lever advantage to restore the winglike process and swimbladder to their resting position. Sonic components grow isometrically and are typically larger in males although the tiny intermediate muscles are larger in females. Although external morphology is relatively conservative in ophidiids, sonic morphology is extremely variable within the family. J. Morphol., 2007. © 2007 Wiley-Liss, Inc.
TL;DR: A detailed description of the jaw and hyobranchial musculature in larval stages of a caecilian is provided and the cranial muscles described for adult caecilians are compared and homologized.
Abstract: Within the Gymnophiona (caecilians) oviparous species with biphasic life-cycles possess a free living semi-aquatic larval stage that feeds in aquatic habitats. The larvae pass through a metamorphosis to a purely terrestrial adult stage. It is likely that the cranial morphology of caecilian larvae has specializations for aquatic feeding. However, little is known about the cranial morphology, and the cranial musculature is especially neglected in the literature. This study provides a detailed description of the jaw and hyobranchial musculature in larval stages of a caecilian. We studied late embryonic and early larval specimens of Ichthyophis kohtaoensis. Furthermore, we compared and homologized the cranial muscles found in larval I. kohtaoensis with the muscles described for adult caecilians. Most cranial muscles of larval I. kohtaoensis are also present in the adult, except for the m. levator mandibulae externus and the m. subarcualis obliquus II. Our results were compared with the data available for larval frogs and salamanders in order to hypothesize the cranial musculature in the larva of the most recent common ancestor of the Lissamphibia. Larval caecilians, frog tadpoles, and salamander larvae share many characters in their cranial musculature, which, consequently, can be assigned to the lissamphibian ground pattern. However, the m. pterygoideus and the m. levator quadrati are unique to the Gymnophiona.
TL;DR: The terminology critical to understanding cranial morphology in polypterids is reviewed and a new description of the skull of E. calabaricus is presented as observed through classical methods of skeletal preparation, X‐radiographic microfocus computed tomography, and 3D‐digital reconstruction.
Abstract: Polypterid fishes are considered the basalmost group of extant actinopterygians and may be a direct link to understanding the systematics and evolution of the first bony fishes. Several investigations have been conducted on one member genus, Polypterus; however, since the first specimens of its sister taxon Erpetoichthys calabaricus were described, remarkably little work has been done on the species. We review terminology critical to understanding cranial morphology in polypterids and present a new description of the skull of E. calabaricus as observed through classical methods of skeletal preparation, X-radiographic microfocus computed tomography, and 3D-digital reconstruction. Differences among E. calabaricus and at least three species of Polypterus (P. bichir, P. senegalus, and P. endlicheri), besides the gross variation in size, include an overall elongation of the skull roof observable in most elements of E. calabaricus with a shortening of most associated processes. In addition, several elements present in species of Polypterus are absent in E. calabaricus. As a result, Polypterus should not be used as a proxy for the family Polypteridae to the exclusion of E. calabaricus in phylogenetic studies, which examine early actinopterygians. Each should be treated separately, to resolve inter- and intrarelationships of Polypteridae.
TL;DR: The digestive system of several species of sea spiders was studied by electron microscopy and found that in the absence of a hepatopancreas, the midgut serves both digestive and absorptive functions.
Abstract: The digestive system of several species of sea spiders (Pycnogonida, Arthropoda) was studied by electron microscopy. It is composed of the foregut inside a long proboscis, a midgut and a hindgut. Lips near the three jaws at the tip of the proboscis receive several hundred ductules originating from salivary glands. These previously undetected glands open on the lips, a fluted, projecting ridge at the external hinge line of the jaws, i.e., to the outside of the mouth. This disposition suggests affinities to the chelicerate line. The trigonal esophagus within the proboscis contains a complex, setose filter device, operated by dedicated muscles, that serves to reduce ingested food to subcellular dimensions. The midgut has diverticula into the bases of all legs. Its cells differentiate from the basal layer and contain a bewildering array of secretion droplets, lysosomes and phagosomes. In the absence of a hepatopancreas, the midgut serves both digestive and absorptive functions. The cuticle-lined hindgut lies in the highly reduced, peg-like abdomen.
Traditionally, pycnogonids have been claimed to have no excretory organ at all. Such a structure, however, has been located in at least one ammotheid, Nymphopsis spinosissima, in which a simple, but standard, excretory gland has been found in the scape of the chelifore. It consists of an end sac, a straight proximal tubule, a short distal tubule, and a raised nephropore. The end sac is a thin-walled and polygonal chamber, about 150 μm in cross section, suspended in the hemocoel of the appendage, its edges radially tethered to the cuticle at more than half a dozen locations. This wall consists of a filtration basement membrane, 1–4 μm thick, facing the hemocoel, and internally of a continuous carpet of podocytes and their pedicels. The podocytes, measuring maximally 10 by 15 μm, have complex contents, of which a labyrinthine system of connected intracellular channels stands out. These coated cisternae open into a central vacuole that often rivals the nucleus in size. The design of the organ closely approximates that of the primitive crustacean Hutchinsoniella macracantha. J. Morphol., 2007. © 2007 Wiley-Liss, Inc.
TL;DR: In this paper, developmental stages critical to the formation of the chondrocranium of the Red-eared Slider, Trachemys scripta (Testudines: Emydidae), are described and illustrated, with particular attention given to ontogenetic changes that take place in the orbitotemporal region of the skull.
Abstract: Few descriptions of the formation of the chelonian chondrocranium exist. Herein, developmental stages critical to the formation of the chondrocranium of the Red-eared Slider, Trachemys scripta (Testudines: Emydidae), are described and illustrated, with particular attention given to ontogenetic changes that take place in the orbitotemporal region of the skull. Morphological descriptions are based on cleared and double-stained and serially-sectioned embryos. These specimens allowed for a detailed evaluation of the developmental morphology of the trabeculae, interorbital septum, pilae metoptica, taeniae marginalis, acrochordal cartilage, pilae antotica, parachordal cartilages, and crista sellaris. Additionally, the formation of the chondrocranium of T. scripta is compared to those of Chrysemys picta (Emydidae) and Caretta caretta (Chelonidae). Overall, the patterns of formation and remodeling of the chondrocranium are quite similar among these species, with the most conspicuous differences observed in remodeling of the posterior orbital cartilages (specifically, the pila metoptica). Reorganization of these cartilages is discussed briefly in the context of associated extraocular muscles for T. scripta and C. caretta. A prominent intertrabecula is reported in T. scripta, supporting previous observations of this structure in emydid turtles.