Showing papers in "Journal of Morphology in 2010"

Cranial functional morphology of fossil dogs and adaptation for durophagy in Borophagus and Epicyon (Carnivora, Mammalia).

Abstract: Morphological specialization is a complex interplay of adaptation and constraint, as similarly specialized features often evolve convergently in unrelated species, indicating that there are universally adaptive aspects to these morphologies. The evolutionary history of carnivores offers outstanding examples of convergent specialization. Among larger predators, borophagine canids were highly abundant during the tertiary of North America and are regarded as the ecological vicars of Afro-Eurasian hyenas. Borophaginae is an extinct group of 60+ species, the largest forms evolving robust skulls with prominently domed foreheads, short snouts, and hypertrophied fourth premolars. These specializations have been speculated to enhance bone cracking. To test the extent that the skulls of derived borophagines were adapted for producing large bite forces and withstanding the mechanical stresses associated with bone cracking relative to their nonrobust sister clades, we manipulated muscle forces in models of six canid skulls and analyzed their mechanical response using 3D finite element analysis. Performance measures of bite force production efficiency and deformation minimization showed that skulls of derived borophagines Borophagus secundus and Epicyon haydeni are particularly strong in the frontal region; maximum stresses are lower and more evenly distributed over the skull than in other canids. Frontal strength is potentially coupled with a temporalis-driven bite to minimize cranial stress during biting in the two derived genera, as tensile stress incurred by contracting temporalis muscles is dissipated rostro-ventrally across the forehead and face. Comparison of estimated masticatory muscle cross section areas suggests that the temporalis-masseter ratio is not strongly associated with morphological adaptations for bone cracking in Borophagus and Epicyon; larger body size may explain relatively larger temporalis muscles in the latter. When compared with previous studies, the overall cranial mechanics of the derived borophagines is more similar to bone-cracking hyaenids and percrocutids than to their canid relatives, indicating convergence in both morphological form and functional capability. J. Morphol., 2010. © 2010 Wiley-Liss, Inc.

Lung size and thoracic morphology in shallow‐ and deep‐diving cetaceans

Abstract: Shallow-diving, coastal bottlenose dolphins (Tursiops truncatus) and deep-diving, pelagic pygmy and dwarf sperm whales (Kogia breviceps and K. sima) will experience vastly different ambient pressures at depth, which will influence the volume of air within their lungs and potentially the degree of thoracic collapse they experience. This study tested the hypotheses that lung size will be reduced and/or thoracic mobility will be enhanced in deeper divers. Lung mass (T. truncatus, n = 106; kogiids, n = 18) and lung volume (T. truncatus, n = 5; kogiids, n = 4), relative to total body mass, were compared. One T. truncatus and one K. sima were cross-sectioned to calculate lung, thoracic vasculature, and other organ volumes. Excised thoraxes (T. truncatus, n = 3; kogiids, n = 4) were mechanically manipulated to compare changes in thoracic cavity shape and volume. Kogiid lungs were half the mass and one-fifth the volume of those of similarly sized T. truncatus. The lungs occupied only 15% of the total thoracic cavity volume in K. sima and 37% in T. truncatus. The kogiid and dolphin thoraxes underwent similar changes in shape and volume, although the width of the thoracic inlet was relatively constrained in kogiids. A broader phylogenetic comparison demonstrated that the ratio of lung mass to total body mass in kogiids, physeterids, and ziphiids was similar to that of terrestrial mammals, while delphinids and phocoenids possessed relatively large lungs. Thus, small lung size in deep-diving odontocetes may be a plesiomorphic character. The relatively large lung size of delphinids and phocoenids appears to be a derived condition that may permit the lung to function as a site of respiratory gas exchange throughout a dive in these rapid breathing, short-duration, shallow divers.

Comparative punting kinematics and pelvic fin musculature of benthic batoids.

Abstract: Although the majority of batoid elasmo- branchs, skates and rays, are benthically associated, benthic locomotion has been largely overlooked in this group. Only skates have been previously described to per- form a form of benthic locomotion termed ''punting.'' While keeping the rest of the body motionless, the skate's pelvic fins are planted into the substrate and then retracted caudally, which thrusts the body forward. In this study, we demonstrate that this form of locomotion is not confined to the skates, but is found across a range of phylogenetically and morphologically diverse batoid species. However, only the clearnose skate, Raja eglanteria, and the lesser electric ray, Narcine brasiliensis, performed ''true punting,'' in which only the pelvic fins were engaged. The yellow stingray, Urobatis jamaicensis, and the Atlantic stingray, Dasyatis sabina, performed ''augmented punting,'' in which pectoral fin movement was also used to generate thrust. Despite this supplemental use of pectoral fins, the augmented punters failed to exceed the punting capabilities of the true punters. The urobatid and the true punters all punted approximately half their disc length per punt, whereas the dasyatid punted a significantly shorter dis- tance. The skate punted significantly faster than the other species. Examination of the pelvic fin musculature revealed more specialized muscles in the true punters than in the augmented punters. This concordance of musculature with punting ability provides predictive power regarding the punting kinematics of other elasmobranchs based upon gross muscular examinations. In contrast to previous assumptions, our results suggest that benthic locomotion is widespread among batoids. J. Morphol. 271:1219-1228, 2010. 2010 Wiley-Liss, Inc.

Ontogeny of the collar cord: neurulation in the hemichordate Saccoglossus kowalevskii.

Abstract: The chordate body plan is characterized by a central notochord, a pharynx perforated by gill pores, and a dorsal central nervous system Despite progress in recent years, the evolutionary origin of each of theses characters remains controversial In the case of the nervous system, two contradictory hypotheses exist In the first, the chordate nervous system is derived directly from a diffuse nerve net; whereas, the second proposes that a centralized nervous system is found in hemichordates and, therefore, predates chordate evolution Here, we document the ontogeny of the collar cord of the enteropneust Saccoglossus kowalevskii using transmission electron microscopy and 3D-reconstruction based on completely serially sectioned stages We demonstrate that the collar cord develops from a middorsal neural plate that is closed in a posterior to anterior direction Transversely oriented ependymal cells possessing myofilaments mediate this morphogenetic process and surround the remnants of the neural canal in juveniles A mid-dorsal glandular complex is present in the collar The collar cord in juveniles is clearly separated into a dorsal saddle-like region of somata and a ventral neuropil We characterize two cell types in the somata region, giant neurons and ependymal cells Giant neurons connect via a peculiar cell junction that seems to function in intercellular communication Synaptic junctions containing different vesicle types are present in the neuropil These findings support the hypotheses that the collar cord constitutes a centralized element of the nervous system and that the morphogenetic process in the ontogeny of the collar cord is homologous to neurulation in chordates Moreover, we suggest that these similarities are indicative of a close phylogenetic relationship between enteropneusts and chordates J Morphol, 2010 ©2010 Wiley-Liss, Inc

Exoskeletal chitin scales isometrically with body size in terrestrial insects

Abstract: The skeletal system of animals provides the support for a variety of activities and functions. For animals such as mammals, which have endoskeletons, research has shown that skeletal investment (mass) scales with body mass to the 1.1 power. In this study, we ask how exoskeletal investment in insects scales with body mass. We measured the body mass and mass of exoskeletal chitin of 551 adult terrestrial insects of 245 species, with dry masses ranging from 0.0001 to 2.41 g (0.0002-6.13 g wet mass) to assess the allometry of exo- skeletal investment. Our results showed that exoskeletal chitin mass scales isometrically with dry body mass across the Insecta as Mchitin 5 aM b , where b 5 1.03 6 0.04, indicating that both large and small terrestrial insects allocate a similar fraction of their body mass to chitin. This isometric chitin-scaling relationship was also evident at the taxonomic level of order, for all insect orders except Coleoptera. We additionally found that the relative exoskeletal chitin investment, indexed by the coefficient, a, varies with insect life history and phylog- eny. Exoskeletal chitin mass tends to be proportionally less and to increase at a lower rate with mass in flying than in nonflying insects (Mflying insect chitin 52 0.56 3 M 0:97 dry ;M nonflying insect chitin 52 0.55 3 M

The formation of the nervous system during larval development in Triops cancriformis (Bosc) (crustacea, Branchiopoda): An immunohistochemical survey.

Abstract: We provide data of the development of thenervous system during the first five larval stages of Triops cancriformis. We use immunohistochemical labeling (against acetylated α-tubulin, serotonin, histamine, and FMRFamide), confocal laser scanning microscopy analysis, and 3D-reconstruction. The development of the nervous system corresponds with the general anamorphic development in T. cancriformis. In larval stage I (L I), all brain parts (proto-, deuto-, and tritocerebrum), the circumoral connectives, and the mandibular neuromere are already present. Also, the frontal filaments and the developing nauplius eye are already present. However, until stage L III, the nauplius eye only consists of three cups. Throughout larval development, the protocerebral network differentiates into distinct subdivisions. In the postnaupliar region, additional neuromeres and their commissures emerge in an anteroposterior gradient. The larval nervous system in L V consists of a differentiated protocerebrum including a central body, a nauplius eye comprising four cups, a circumoral nerve ring, mandibular- and postnaupliar neuromeres up to the seventh thoracic segment, each featuring an anterior and a posterior commissure, and two parallel connectives. The presence of a protocerebral bridge is questionable. The distribution of neurotransmitters in L I is restricted to the naupliar nervous system. Over the course of the five stages of development, neurotransmitter distribution also follows an anteroposterior gradient. Each neuromere is equipped with two ganglia innervating the locomotional appendages and possesses a specific neurotransmitter distribution pattern. We suggest a correlation between neurotransmitter expression and locomotion.

The simplicity of males: dwarf males of four species of Osedax (Siboglinidae; Annelida) investigated by confocal laser scanning microscopy.

Abstract: Dwarf males of the bone-eating worms Osedax (Siboglinidae, Annelida) have been proposed to develop from larvae that settle on females rather than on bone. The apparent arrest in somatic development and resemblance of the males to trochophore larvae has been posited as an example of paedomorphosis. Here, we present the first investigation of the entire muscle and nervous system in dwarf males of Osedax frankpressi, O. roseus, O. rubiplumus, and O. ''spiral'' analyzed by multistaining and confocal laser scanning microscopy. Sperm shape and spermiogenesis, the sperm duct and internal and external ciliary patterns were likewise visualized. The males of all four species possess morpho- logical traits typical of newly settled siboglinid larvae: a prostomium, a peristomium with a prototroch, one elon- gate segment and a second shorter segment. Each seg- ment has a ring of eight long-handled hooked chaetae. The longitudinal muscles are distributed as evenly spaced strands forming a grid with the thin outer circu- lar muscles. Oblique protractor and retractor muscles are associated with each of the chaetal sacs. The nerv- ous system comprises a cerebral ganglion, a prototroch nerve ring, paired dorsolateral longitudinal nerves, five ventral longitudinal nerves with paired, posterior gan- glia and a terminal commissure, as well as a net of fine peripheral transverse plexuses surrounding the first seg- ment. Internal ciliation occurs as paired ventrolateral bands along the first segment. The bands appear to lead the free mature sperm to a ciliated duct and seminal vesicle lying just behind the prototroch region. A duct then runs from the seminal vesicle into the dorsal part of the prostomium. The similarity of Osedax males to the larvae of Osedax and other siboglinid annelids as well as similarities shown here to the neuromuscular organization seen in other annelid larvae supports the hypothesis of paedomorphosis in males of Osedax .J. Morphol. 271:127-142, 2010. 2009 Wiley-Liss, Inc.

Morphological correlates of sprint swimming speed in five species of spadefoot toad tadpoles: Comparison of morphometric methods

Abstract: The relationship between vertebrate morphology and swimming performance has long interested biologists. Recent work on predator-induced morphological plasticity of anuran tadpoles has increased this interest. Here, I use data on five species of spadefoot toad tadpoles (Scaphiopodidae) to compare linear and geometric morphometrics. Linear measures explain only 7-26% of the variation in swimming speed, depending on species, whereas geometric morphometrics could explain 24-46% of the same variation. I also compare two methods for examining how similar the morphology-swimming speed relationship is among species. A canonical variate derived from a MANCOVA approach successfully detected species differences in these relationships, whether using linear or geometric methods, but a canonical correlation approach failed in both cases. Overall, tadpoles with smaller bodies, larger tails, and larger tail muscles are faster swimmers but the details of how these shape changes are achieved differed among species. For example, in some species a smaller body was achieved primarily by reducing abdomen size, whereas in others both the head and abdomen are smaller.Faster swimmers also had deeper tails, especially in the posterior half of the tail. This pattern would have been missed in standard linear morphometrics which usually only measures maximum tail depth.

Development of the cypriniform protrusible jaw complex in Danio rerio: constructional insights for evolution.

Abstract: Studies on the evolution of complex biological systems are difficult because the construction of these traits cannot be observed during the course of evolution. Complex traits are defined as consisting of multiple elements, often of differing embryological origins, with multiple linkages integrated to form a single functional unit. An example of a complex system is the cypriniform oral jaw apparatus. Cypriniform fishes possess an upper jaw characterized by premaxillary protrusion during feeding. Cypriniforms effect protrusion via the kinethmoid, a synapomorphy for the order. The kinethmoid is a sesamoid ossification suspended by ligaments attaching to the premaxillae, maxillae, palatines, and neurocranium. Upon mouth opening, the kinethmoid rotates as the premaxillae move anteriorly. Along with bony and ligamentous elements, there are three divisions of the adductor mandibulae that render this system functional. It is unclear how cypriniform jaws evolved because although the evolution of sesamoid elements is common, the incorporation of the kinethmoid into the protrusible jaw results in a function that is atypical for sesamoids. Developmental studies can show how biological systems are assembled within individuals and offer clues about how traits might have been constructed during evolution. We investigated the development of the protrusible upper jaw in zebrafish to generate hypotheses regarding the evolution of this character. Early in development, the adductor mandibulae arises as a single unit. The muscle divides after ossification of the maxillae, on which the A1 division will ultimately insert. A cartilaginous kinethmoid first develops within the intermaxillary ligament; it later ossifies at points of ligamentous attachment. We combine our structural developmental data with published kinematic data at key developmental stages and discuss potential functional advantages in possessing even the earliest stages of a system for protrusion.

Structure and occurrence of cyphonautes larvae (bryozoa, ectoprocta).

Abstract: We have studied larvae of the freshwater ctenostome Hislopia malayensis with scanning electron microscopy (SEM), confocal laser scanning microscopy (CLSM), and LM of serial sections. Some additional observations on larvae of M. membranacea using SEM and CLSM are also reported. The overall configuration of muscles, nerves, and cilia of the two larvae are identical. However, the larva of H. malayensis is much smaller than that of M. membranacea, which may explain most of the differences observed. Although all major nerves and muscle strands are present in H. malayensis, they are generally composed of fewer fibers. The H. malayensis larva lacks the anterior and posterior intervalve cilia. Its pyriform organ is unciliated with only a small central depression. The adhesive epithelium is not invaginated as an adhesive sac and lacks the large muscles interpreted as adhesive sac muscles in the M. membranacea larva. The velum carries two rows of ciliated cells, though the lower “row” consists of only one or two cells. Both rows of ciliated cells are innervated by nerves, which have not been detected in the M. membranacea larva. The ciliated ridge of H. malayensis lacks the frontal cilia. The planktotrophic cyphonautes larvae in a number of ctenostome clades and in the “basal” cheilostome clade Malacostega (and probably in the earliest cheilostomes) support the idea that the cyphonautes larva is the ancestral larval type of the Eurystomata. It may even represent the ancestral larval type of the bryozoans (= ectoprocts). J. Morphol. 271:1094-1109, 2010. © 2010 Wiley-Liss, Inc.

Functional implications of dicynodont cranial suture morphology.

Abstract: Cranial suture morphology of Lystrosaurus and the generalized dicynodont Oudenodon was investigated to determine the strain environment during mastication, which in turn may indicate a difference in cranial function between the two taxa. Finite element (FE) analysis indicated that less strain accumulated in the cranium of Lystrosaurus during orthal bite simulations than in Oudenodon. Despite the overall difference in strain magnitude, moderate to high FE-predicted strain accumulated in similar areas of the cranium of both taxa. The suture morphology in these cranial regions of Lystrosaurus and Oudenodon was investigated further by examination of histological sections and supplemented by observations of serial sections and computed tomography (CT) scans. The predominant type of strain from selected blocks of finite elements that contain sutures was determined, enabling comparison of suture morphology to strain type. Drawing from strain-suture correlations established in extant taxa, the observed patterns of sutural morphology for both dicynodonts were used to deduce cranial function. The moderate to high compressive and tensile strain experienced by the infraorbital bar, zygomatic arch, and postorbital bar of Oudenodon and Lystrosaurus may have been decreased by small adjustive movements at the scarf sutures in those regions. Disparities in cranial suture morphology between the two taxa may reflect differences in cranial function. For instance, the tongue and groove morphology of the postorbital-parietal suture in Oudenodon could have withstood the higher FE-predicted tensile strain in the posterior skull roof. The scarf premaxilla-nasal suture of Lystrosaurus provided an additional region of sutural mobility in the anterior surface of the snout, suggesting that Lystrosaurus may have employed a different biting regime than Oudenodon. The morphology of several sutures sampled in this study correlated with the FE-predicted strain, although other cranial functional hypotheses remain to be tested.

Thermally induced plasticity of body shape in adult zebrafish Danio rerio (Hamilton, 1822)

Abstract: We examined the effect of temperature during the early development on the phenotypic plasticity of Danio rerio. The effect of temperature was examined during two different early developmental periods of 280°d (the product of days × temperature) each, 28-308°d or 280-560°d, by subjecting the experimental populations to three different water temperatures (22°C, 28°C, and 32°C). Before and after the end of the 280°d period of the different thermal exposure, all populations were cultured in standard temperature (28°C). Five to 10 months after exposure to the different thermal regimes, the body shape of the adults was analyzed by geometric morphometrics. In both ontogenetic windows and experimental repetitions, the results showed that developmental temperature and sex significantly affected the body shape of adult zebrafish. Thermally induced shape variation discriminated the fish that developed at 22°C from those developed at 28°C-32°C. In the early developmental period (DP1, 28-308°d postfertilization), dorsal, anal, and caudal fin structures differed between the animals that developed at 22°C and 28°C-32°C. In the later developmental period (DP2, 280-560°d postfertilization), caudal, anal, pectoral, and pelvic fins, as well as the gill cover and lower jaw, were affected when animals developed at different temperatures. These results show that thermal history during a short period of embryonic and larval life affects the body form of adult zebrafish with potentially functional consequences. Based on previous data on the effects of temperature on fish development, we suggest thermally induced muscle and bone remodelling as possible mechanism underlying the observed plasticity.

Comparative cranial osteology of fossorial lizards from the tribe gymnophthalmini (Squamata, Gymnophthalmidae)

Abstract: Squamates (lizards, snakes and amphisbaenians) are represented by a large number of species distributed among a wide variety of habitats. Changes in body plan related to a fossorial habit are a frequent trend within the group and many morphological adaptations to this particular lifestyle evolved convergently in nonrelated species, reflecting adaptations to a similar habitat. The fossorial lifestyle requires an optimal morphological organization for an effective use of the available resources. Skeleton arrangement in fossorial squamates reflects adaptations to the burrowing activity, and different degrees of fossoriality can be inferred through an analysis of skull morphology. Here, we provide a detailed description of the skull morphology of three fossorial gymnophthalmid species: Calyptommatus nicterus, Scriptosaura catimbau, and Nothobachia ablephara.

Comparative anatomy and osteohistology of hyperelongate neural spines in the sphenacodontids Sphenacodon and Dimetrodon (Amniota: Synapsida).

Abstract: Osteohistological investigations of hyperelongate vertebral spinous processes (neural spines) are presented to elucidate previously unknown aspects of dorsal sail form and function in two, closely related genera of "sail-backed" synapsids: Sphenacodon and Dimetrodon. Although recent and classic surveys of bone histology in extinct vertebrates have sampled the genus Dimetrodon, new sectioning of Sphenacodon material allows a comparative analysis of these structures among Sphenacodontidae for the first time. Variability within the histological organization of the neural spine is assessed by examining multiple regions along its length, and implications for soft tissue correlates, growth and mechanics are considered here. Both genera exhibit extensive parallel-fibered and fibrolamellar bone, in addition to lamellar bone. Several features vary along the length of the spine in each species. Muscle scars and extensive Sharpey's fibers are present at the base of the spine; no scars and fewer fibers are manifested ∼55-60 mm above the zygapophyses in mature individuals. The distal cortex of the spine does not exhibit greater vascularity than the proximal region in either genus. However, both genera manifest distinct vascular grooves of variable size along the distal periosteal surface, some of which become incorporated into the distal cortex. The observed histovariability appears to record the transition from the proximal (epaxial muscle embedded) to the distally protruding portion of the spine. These observations and independent pathological evidence support the existence of a short dorsal crest in Sphenacodon and possibly other basal sphenacodontids. Although the thermoregulatory capacity of such a crest remains uncertain, developmental and mechanical features are readily interpretable and are discussed with respect to the origins and early evolution of the dorsal sail in sphenacodontid synapsids.

Morphology and function of the reproductive system of representatives of the genus Uca

Abstract: The morphology of the reproductive organs of three species of fiddler crabs, Uca ecuadoriensis, Uca c.f. forcipata, and Uca tangeri were investigated to subsequently produce a model of their mode of operation. Vulva, vagina, and spermatheca in females, and the first and second gonopods in males were examined by applying histological techniques and electron microscopy. In all three species, vulva and vagina conform to the concave type, and the spermatheca complies with the ventral type. The tissue of the oviduct orifice is enlarged and bulges into the lumen of the spermatheca. Differences between the three species are apparent in the organization of the spermatheca, especially in the distribution and structure of glandular epithelium: In U. ecuadoriensis and U. c.f. forcipata the largest proportion of the spermathecal wall is lined with cuticle and only a small area consists of glandular epithelium, while in U. tangeri almost all of the lining is glandular. Furthermore, the glandular epithelia of the species differ in their histology and ultrastructure: In U. ecuadoriensis it is tubular and multilayered, while in U. c.f. forcipata it is mono-layered. U. tangeri finally has both forms of this tissue. In the males, the terminal segments of the first gonopod exhibit a tight fit to female organs and narrow, tightly sealed sperm channels. These features suggest a tendency towards minimizing loss of fluids, which can be interpreted as an adaptation to mating on land. The tight fit of male gonopod and female opening seem to be protection from interbreeding, which points toward a strong sexual selection. In the terrestrial environment, these originally aquatic organisms experience serious competition for resources; therefore there is pressure on successful reproduction. According to the current results a model of the process of fertilization and egg-laying involving the investigated organs was generated. J. Morphol., 2010. © 2010 Wiley-Liss, Inc.

Placental specializations of the mountain spiny lizard Sceloporus jarrovi.

Abstract: The lizard Sceloporus jarrovi (Phrynosomatidae) is one of the most widely studied viviparous reptiles of North America. Past research has assumed that placentation in this species is relatively simple and functions mainly in gas exchange. Our examination of the late stage placenta via transmission electron microscopy reveals that S. jarrovi has a unique combination of placental characteristics, with unusual specializations for secretion and absorption. In the chorioallantoic placenta, chorionic and uterine tissues are directly apposed through eggshell loss, and their epithelia are greatly attenuated, enhancing gas exchange; this placenta shows evidence of both nutrient transfer and endocrine function. Contrary to past inferences, a yolk sac placenta forms from the avascular omphalopleure and persists through the end of gestation. The uterine epithelium is enlarged and secretory, and the fetal omphalopleure shows branching absorptive channels and other specializations for uptake. Elsewhere, the omphalopleure develops elongated folds that protrude into a coagulum of degenerating shell membrane and other organic material. Uterine tissue in this region shows specializations for absorption. Placental features in S. jarrovi have unexpected functional implications, and challenge assumptions that specializations for nutrient transfer are confined to matrotrophic species. J. Morphol. 271:1153–1175, 2010. © 2010 Wiley-Liss, Inc.

Chirally dimorphic male genitalia in praying mantids (Ciulfina: Liturgusidae)

Abstract: Although male polymorphisms occur widely in nature and have received considerable recent attention from studies of alternative mating strategies, male genital polymorphisms are less well known Here, we describe a dimorphism in the orientation of the male genitalic complex of the praying mantid genus Ciulfina Populations of Ciulfina species vary in the proportion of males with dextral (right-oriented) and sinistral (left-oriented) genitalia, ranging from directional asymmetry (single orientation only) to apparent antisymmetry (equal proportions of both orientations) The proportion of dextral males varied between species (C baldersoni: 46%; C rentzi: 24%; C klassi: 100%; C biseriata: 83%) and between populations We used elliptic Fourier analysis to quantify shape and size variation between the genitalia of dextral and sinistral males and determined that the two forms were mirror images of one another in two species We found that the level of mechanical reproductive isolation between heterospecific populations of opposite genital orientation was no greater than that between heterospecific populations with the same orientation or of mixed orientation Genital orientation therefore did not influence premating isolation between these species, despite complete postmating isolation The geographic proximity of populations to heterospecifics also showed no particular pattern with respect to genital orientation These results suggest that reversible trait asymmetry in Ciulfina is not driven by reproductive isolation, and add to the growing evidence against the species isolation hypothesis for rapid genital evolution

Sperm ultrastructure of the spider crab Maja brachydactyla (Decapoda: Brachyura)

Abstract: This study describes the morphology of the sperm cell of Maja brachydactyla, with emphasis on localizing actin and tubulin. The spermatozoon of M. brachydactyla is similar in appearance and organization to other brachyuran spermatozoa. The spermatozoon is a globular cell composed of a central acrosome, which is surrounded by a thin layer of cytoplasm and a cup-shaped nucleus with four radiating lateral arms. The acrosome is a subspheroidal vesicle composed of three concentric zones surrounded by a capsule. The acrosome is apically covered by an operculum. The perforatorium penetrates the center of the acrosome and has granular material partially composed of actin. The cytoplasm contains one centriole in the subacrosomal region. A cytoplasmic ring encircles the acrosome in the subapical region of the cell and contains the structures-organelles complex (SO-complex), which is composed of a membrane system, mitochondria with few cristae, and microtubules. In the nucleus, slightly condensed chromatin extends along the lateral arms, in which no microtubules have been observed. Chromatin fibers aggregate in certain areas and are often associated with the SO-complex. During the acrosomal reaction, the acrosome could provide support for the penetration of the sperm nucleus, the SO-complex could serve as an anchor point for chromatin, and the lateral arms could play an important role triggering the acrosomal reaction, while slightly decondensed chromatin may be necessary for the deformation of the nucleus. J. Morphol., 2010. © 2009 Wiley-Liss, Inc.

Developmental Expression of Neuromodulators in the Central Complex of the Grasshopper Schistocerca gregaria

Abstract: The central complex is a major integrative region within the insect brain with demonstrated roles in spatial orientation, the regulation of locomotor behavior, and sound production. In the hemimetabolous grasshopper, the central complex comprises the protocerebral bridge, central body (CB), ellipsoid body, noduli, and accessory lobes, and this modular organization develops entirely during embryogenesis. From a biochemical perspective, a range of neuroactive substances has been demonstrated in these modules of the adult central complex, but little is known about their developmental expression. In this study, we use matrix-assisted laser desorption/ionization-imaging mass spectrometry on single brain slices to confirm the presence of several peptide families (tachykinin, allatostatin, periviscerokinin/pyrokinin, FLRFamide, and neuropeptide F) in the adult central complex and then use immunohistochemistry and histology to examine their developmental expression, together with that of the indolamin serotonin, and the endogenous messenger nitric oxide (NO; via its synthesizing enzyme). We find that each neuromodulator is expressed according to a unique, stereotypic, pattern within the various modules making up the central complex. Neuropeptides such as tachykinin (55%) and allatostatin (65%), and the NO-synthesizing enzyme diaphorase (70%), are expressed earlier during embryonic development than the biogenic amine serotonin (80%), whereas periviscerokinin-like peptides and FLRFamide-like peptides begin to be expressed only postembryonically. Within the CB, these neuroactive substances are present in tangential projection neurons before they appear in columnar neurons. There is also no colocalization of serotonin-positive and peptide-positive projections up to the third larval instar during development, consistent with the clear dorsoventral layering of the neuropil we observe. Our results provide the first neurochemical fingerprint of the developing central complex in an hemimetabolous insect.

Uterine and eggshell structure and histochemistry in a lizard with prolonged uterine egg retention (Lacertilia, Scincidae, Saiphos)

Abstract: The eggshell of lizards is a complex structure composed of organic and inorganic molecules secreted by the oviduct, which protects the embryo by providing a barrier to the external environment and also allows the exchange of respiratory gases and water for life support. Calcium deposited on the surface of the eggshell provides an important nutrient source for the embryo. Variation in physical conditions encountered by eggs results in a tradeoff among these functions and influences eggshell structure. Evolution of prolonged uterine egg retention results in a significant change in the incubation environment, notably reduction in efficiency of gas exchange, and selection should favor a concomitant reduction in eggshell thickness. This model is supported by studies that demonstrate an inverse correlation between eggshell thickness and length of uterine egg retention. One mechanism leading to thinning of the eggshell is reduction in size of uterine shell glands. Saiphos equalis is an Australian scincid lizard with an unusual pattern of geographic variation in reproductive mode. All populations retain eggs in the uterus beyond the embryonic stage at oviposition typical for lizards, and some are viviparous. We compared structure and histochemistry of the uterus and eggshell of two populations of S. equalis, prolonged egg retention, and viviparous to test the hypotheses: 1) eggshell thickness is inversely correlated with length of egg retention and 2) eggshell thickness is positively correlated with size of shell glands. We found support for the first hypothesis but also found that eggshells of both populations are surprisingly thick compared with other lizards. Our histochemical data support prior conclusions that uterine shell glands are the source of protein fiber matrix of the eggshell, but we did not find a correlation between size of shell glands and eggshell thickness. Eggshell thickness is likely determined by density of uterine shell glands in this species. J. Morphol., 2010. © 2010 Wiley-Liss, Inc.

Cranial shape varies along an elevation gradient in Gambel's white-footed mouse (Peromyscus maniculatus gambelii) in the Grinnell Resurvey Yosemite transect.

Abstract: Environmental variation over a species's range creates differing pressures to which organisms must adjust in order to survive. Taxa can respond to these pressures at population and individual levels, leading to localized phenotypic differentiation. Assessing the spatial distribution of phenotypic variation can illuminate how dramatically varying environmental factors shape phenotypes and may forecast a taxon's ability to adapt should conditions change. We characterized morphological variation along a transect sampled in the Grinnell Resurvey project to determine whether Gambel's white-footed mouse (Peromyscus maniculatus gambelii), a generalist taxon inhabiting the full elevational range of habitats in Yosemite National Park and surrounding areas, has responded morphologically to variation in its environment. We quantified variation in modern P. m. gambelii cranial shape using 2D generalized Procrustes analysis and Euclidean distance matrix-based geometric morphometrics. We performed multivariate regression of shape coordinates on elevation to test for environmental influences on shape within the principal geographic dimension of change along the transect. We observe a statistically significant correlation with shape on elevation for occlusal and lateral views of the cranium, explaining a small percentage of the overall variation in shape. Modern P. m. gambelii crania show a pattern of flexion in which the angle of the cranial base decreases at higher elevations. Results of EDMA parallel these findings, but highlight additional areas of the cranium that vary with elevation. Collectively, the patterns of variation detected suggest a biological response to the environment that warrants further study. This work lays the foundation for comparison with morphological data from historical specimens, which can address evolutionary scenarios generated from our findings, and for investigation of other taxa included in the resurvey project. J. Morphol. 271:897–909, 2010. © 2010 Wiley-Liss, Inc.

The histological structure of glyptosaurine osteoderms (Squamata: Anguidae), and the problem of osteoderm development in squamates.

Abstract: Glyptosaurinae, a fossil clade of anguid lizards, possess robust osteoderms, with granular ornamentation. In this study, the structural and histological features of these osteoderms were described in order to reconstruct their developmental pattern and further document the degree of homology that could exist between vertebrate integumentary skeletons. Glyptosaurine osteoderms have a diploe architecture and display an unusually complex structure that includes four tissue types: a core of woven-fibered bone intensely remodeled; a peripheral formation of the same tissue containing dense bundles of long Sharpey fibers; a thick basal layer of lamellar bone; and a superficial layer of a non-osseous material that belongs to the category of hypermineralized tissues such as ganoine, or enameloid and enamel tissues. The growth pattern of glyptosaurine osteoderms involved appositional processes due to osteoblast activity. In early growth stages, osseous metaplasia might have also been involved, but this possibility is not substantiated by histological observations. The superficial layer of the osteoderms must have resulted from epidermal contribution, a conclusion that would support previous hypotheses on the role of epidermal-dermal interactions in the formation of squamate osteoderms.

A comparison of the external morphology of the membranous inner ear in elasmobranchs.

Abstract: Studies on the elasmobranch inner ear have focused predominantly on a small group of sharks, particularly, carcharhinids. As a result, subsequent studies in other species have subdivided species into two main groups: those typical and those atypical of carcharhinid sharks. This study proposes a different set of inner-ear morphology groupings to those previously suggested. The inner ears from 17 species of elasmobranchs (representing both sharks and rays) are examined in this study and based on morphometric data some groups include both rays and sharks. Four groups are now proposed based predominantly on the shape and dimensions of the membranous otoconial organs, and characteristics of the semicircular canals. Evident morphological differences between the ear types belonging to the new groups include the membranes of the semicircular canals being bound to the otoconial organs in some species, while only being connected via the canal ducts in others, as well as clear variation present in saccular organ size. Previous studies examining variation in the inner ear have attributed differences to either phylogeny or functional significance. Results from this study suggest that neither phylogeny nor feeding strategy solely accounts for the morphological diversity present in the external morphology of the elasmobranch inner ear. J. Morphol., 2010. © 2010 Wiley-Liss, Inc.

Fine structure of the branchial epithelium in the teleost Oreochromis niloticus.

Abstract: We have studied the gill epithelium of Oreochromis niloticus using transmission electron microscopy with the particular interested relationship between cell morphology and osmotic, immunoregulatory, or other non-regulatory functions of the gill. Pavement cells covered the filament epithelium and lamellae of gills, with filament pavement cells showing distinct features from lamellar pavement cells. The superficial layer of the filament epithelium was formed by osmoregulatory elements, the columnar mitochondria-rich, mucous and support cells, as well as by their precursors. Light mitochondria-rich cells were located next to lamellae. They exhibited an apical crypt with microvilli and horizontal small dense rod-like vesicles, sealed by tight junctions to pavement cells. Dark mitochondria-rich cells had long dense rod-like vesicles and a small apical opening sealed by tight junctions to pavement cells. The deep layer of the filament epithelium was formed by a network of undifferentiated cells, containing neuroepithelial and myoepithelial cells, macrophage and eosinophil-like cells and their precursors, as well as precursors of mucous cells. The lateral-basal surface was coated by myoepithelial cells and a basal lamina. The lamellar blood lacunae was lined by pillar cells and surrounded by a basal lamina and pericytes. The data presented here support the existence of two distinct types of pavement cells, mitochondria-rich cells, and mitochondria-rich cells precursors, a structural role for support cells, a common origin for pavement cells and support cells, a paracrine function for neuroepithelial cells in the superficial layer, and the control of the lamellar capillary base by endocrine and contractile cells. Data further suggest that the filament superficial layer is involved in gill osmoregulation, that may interact, through pale mitochondria-rich cells, with the deep layer and lamellae, whereas the deep layer, through immune and neuroendocrine systems, acts in the regeneration and defense of the tissue. J. Morphol. 2010. © 2010 Wiley-Liss, Inc.

Cephalic morphology of Pythonichthys macrurus (Heterenchelyidae: Anguilliformes): specializations for head-first burrowing.

Abstract: The Heterenchelyidae, a family of Anguil- liformes, are highly specialized fossorial eels. This study was conducted to evaluate the cranial specialization in relation to head-first burrowing behavior in the heteren- chelyid, Pythonichthys macrurus. Thereby, detailed descriptions are provided of the cranial myology and os- teology of P. macrurus and its differences with that of representatives of three families: the Moringuidae (Mor- ingua edwardsi), a head-first burrower; the Anguillidae (Anguilla anguilla), a nonburrowing representative and the Ophichthidae (Pisodonophis boro), a head and tail- first burrower. This comparison may help to get a better understanding of the cranial specialization of head-first burrowers in heterenchelyids and moringuids. We recog- nize as morphological adaptations to burrowing: reduced eye size, a caudoventral orientation of the anteromedial section of the adductor mandibulae muscle complex, the posterior position of the quadrate-mandibular joint, a solid conical skull, large insertion sites of epaxial and hypaxial muscle on the neurocranium, a widened ce- phalic lateral line canals extending into the dermal cav- ities, and a ventral position of the gill opening. J. Mor- phol. 271:1053-1065, 2010. 2010 Wiley-Liss, Inc.

Cranial architecture of tube-snouted gasterosteiformes (Syngnathus rostellatus and Hippocampus capensis).

Abstract: The long snout of pipefishes and seahorses (Syngnathidae, Gasterosteiformes) is formed as an elon- gation of the ethmoid region. This is in contrast to many other teleosts with elongate snouts (e.g., butterflyfishes) in which the snout is formed as an extension of the jaws. Syngnathid fishes perform very fast suction feed- ing, accomplished by powerful neurocranial elevation and hyoid retraction. Clearly, suction through a long and narrow tube and its hydrodynamic implications can be expected to require certain adaptations in the cra- nium, especially in musculoskeletal elements of the feed- ing apparatus. Not much is known about which skeletal elements actually support the snout and what the effect of elongation is on related structures. Here, we give a detailed morphological description of the cartilaginous and bony feeding apparatus in both juvenile and adult Syngnathus rostellatus and Hippocampus capensis. Our results are compared with previous morphological stud- ies of a generalized teleost, Gasterosteus aculeatus .W e found that the ethmoid region is elongated early during development, with the ethmoid plate, the hyosymplectic, and the basihyal cartilage being extended in the chon- drocranium. In the juveniles of both species almost all bones are forming, although only as a very thin layer. The elongation of the vomeral, mesethmoid, quadrate, metapterygoid, symplectic, and preopercular bones is already present. Probably, because of the long and speci- alized parental care which releases advanced develop- mental stages from the brooding pouch, morphology of the feeding apparatus of juveniles is already very simi- lar to that of the adults. We describe morphological fea- tures related to snout elongation that may be considered adaptations for suction feeding; e.g. the peculiar shape of the interhyal bone and its saddle-shaped articulation with the posterior ceratohyal bone might aid in explosive hyoid retraction by reducing the risk of hyoid dislocation. J. Morphol. 271:255-270, 2010. 2009 Wiley-Liss, Inc.

The heart of the South American rattlesnake, Crotalus durissus.

Abstract: Most anatomical and physiological studies of the sauropsid heart have focused on species with extraordinary physiologies, and detailed anatomical descriptions of hearts from sauropsids with more com- mon physiologies are therefore warranted. Here, we present a comprehensive study of the cardiac anatomy of the South American rattlesnake (Crotalus durissus). The cardiovascular physiology of this species has been investigated in a number of studies, whereas only a few cursory studies exist on the cardiac anatomy of viperid snakes. The heart of C. durissus is typically squamate in many regards. Both atria are thin-walled sacs, and the right atrium is the most voluminous. The single ven- tricle contains three major septa; the vertical septum, the muscular ridge (MR), and the bulbuslamelle. These partially divide the ventricle into three chambers; the systemic and left-sided cavum arteriosum (CA), the pul- monary and right-sided cavum pulmonale, and the medial cavum venosum (CV). The MR is the most devel- oped septum, and several additional and minor septa are found within the CA and CV. An extraordinary thin cortical layer encloses the ventricle, and it is irrigated by a remarkably rich arborization of coronary arteries. Previous studies show high degrees of blood flow separa- tion in the Crotalus heart, and this can only be explained by the coordinated actions of the septa and the prominent atrioventricular valves. J. Morphol. 271:1066-1077, 2010. 2010 Wiley-Liss, Inc.

Size dimorphism and avian-perceived sexual dichromatism in a New Zealand endemic bird, the whitehead Mohoua albicilla.

Abstract: Sex differences in behavior, morphology, and physiology are common in animals. In many bird species, differences in the feather colors of the sexes are apparent when judged by human observers and using physical measures of plumage reflectance, cryptic (to human) plumage dichromatism has also been detected in several additional avian lineages. However, it remains to be confirmed in almost all species whether sexual dichromatism is perceivable by individuals of the studied species. This latter step is essential because it allows the evaluation of alternative hypotheses regarding the signaling and communication functions of plumage variation. We applied perceptual modeling of the avian visual system for the first time to an endemic New Zealand bird to provide evidence of subtle but consistent sexual dichromatism in the whitehead, Mohoua albicilla. Molecular sexing techniques were also used in this species to confirm the extent of the sexual size dimorphism in plumage and body mass. Despite the small sample sizes, we now validate previous reports based on human perception that in male whiteheads head and chest feathers are physically brighter than in females. We further suggest that the extent of sexual plumage dichromatism is pronounced and can be perceived by these birds. In contrast, although sexual dimorphism was also detectable in the mass among the DNA-sexed individuals, it was found to be less extensive than previously thought. Sexual size dimorphism and intraspecifically perceivable plumage dichromatism represent reliable traits that differ between female and male whiteheads. These traits, in turn, may contribute to honest communication displays within the complex social recognition systems of communally breeding whitehead and other group-breeding taxa

Shape of articular surface of crocodilian (Archosauria) elbow joints and its relevance to sauropsids.

Abstract: The determination of area and shape of articular surfaces on the limb bones of extinct archosaurs is difficult because of postmortem decomposition of the fibrous tissue and articular cartilages that provide the complex three-dimensional joint surfaces in vivo. This study aims at describing the shape of the articular cartilages in the elbow joints of six crocodilian specimens; comparing its structure with that of four birds, three testudines, and five squamates; and comparing the shapes of the surfaces of the calcified and the articular cartilages in the elbow joints of an Alligator specimen. The shapes of the articular cartilages of crocodilian elbow joint are shown to resemble those of birds. The humerus possesses an olecranon fossa positioned approximately at the midportion of the distal epiphysis and bordering the margin of the extensor side of the articular surface. The ulna possesses a prominent intercotylar process at approximately the middle of its articular surface, and splits the surface into the radial and ulnar cotylae. This divides the articular cartilage into an articular surface on the flexor portion, and the olecranon on the extensor portion. The intercotylar process fits into the olecranon fossa to restrict elbow joint extension. Dinosaurs and pterosaurs, phylogenetically bracketed by Crocodylia and Aves (birds), may have possessed a similar olecranon fossa and intercotylar process on their articular cartilages. Although these shapes are rarely recognizable on the bones, their impressions on the surfaces of the calcified cartilages provide an important indication of the extensor margin of the articular surfaces. This, in turn, helps to determine the maximum angle of extension of the elbow joint in archosaurs. J. Morphol., 2010. © 2010 Wiley-Liss, Inc.

Morphology and function of the hyoid apparatus of fossil xenarthrans (mammalia).

Abstract: The analysis of the hyoid apparatus of fossil xenarthrans provides insight on the form of the tongue and its function in food intake and intraoral processing. The hyoid ap- paratus of xenarthrans is notable for fusion among its ele- ments. The presence of a V-bone, a complex consisting of fused basihyal and thyrohyal bones, is a consistent and probably synapomorphic feature of xenarthrans. Fusion of other ele- ments is variable in fossil xenarthrans. Most fossil sloths retain independent elements, as in living dasypodids and mammals generally. Among nothrotheriids, the elements are slender and their articular surfaces indicate considerable mo- bility, and the relatively long and horizontal orientation of the geniohyoid muscle suggests considerable tongue protru- sion. Among mylodontines, such as Paramylodon and Glossotherium, the elements indicate relatively mobile articu- lations, except between the stylo- and epihyals. The relatively posterior placement of the apparatus and the length and alignment of the geniohyoid muscle indicate considerable capacity for tongue protrusion. Scelidotherium, however, had rigidly articulated stylohyal and epihyal, and the apparatus lies farther anteriorly, which together with the elongated, steeply inclined mandibular symphysis, indicates a relatively shorter geniohyoid muscle and thus more limited capacity for tongue protrusion. A similar situation is indicated for Megatherium, casting doubt on the classical reconstruction of this sloth as having a long prehensile tongue. Among cingu- lates Prozaedyus resembles living dasypodids, indicating con- siderable tongue protrusion important in food acquisition and intake. More extensive fusion of hyoid elements occurs in the cingulates Glyptodon and Proeutatus, in which the stylohyal and epihyal at least, are fused into a single element termed the sigmohyal. The presence of this element supports recent proposals of a sister-group relationship between glyptodonts and eutatines. The rigidity of the apparatus suggests limited tongue protrusion, but the tongue, in glyptodonts at least, was a powerful structure important for intraoral manipulation of food. J. Morphol. 271:1119-1133, 2010. 2010 Wiley-Liss, Inc.