Morphology and biomechanical implications of isolated discocystinid plates (Edrioasteroidea, Echinodermata) from the Carboniferous of North America

Detailed examination of isolated thecal plates belonging to three discocystinid edrioasteroids, Spiraclavus nacoensis Sumrall, Hypsiclavus kinsleyi Sumrall, and Giganticlavus bennisoni Sumrall and Bowsher, reveals striking similarity in morphology among these species. Stereom observed in the ambulacral floor plates indicates that ligamentous connective tissue opened the ambulacral cover plates and muscle tissue closed them. The ambulacral floor plates are interpreted as rigid supports for the oral surface with the interambulacral areas acting as flexible integuments of plates. The aboral surface is interpreted as flexible and highly contractile. All discocystinid thecal openings are consistent in morphology with adaptations for thecal pressurization. Extension and contraction of the theca was accomplished by pumping water in and out of an inflatable sac associated with the periproctial opening. The pedunculate zone is interpreted as passively expanding and contracting by relaxing of mutable collagenous tissue and stiffening when the theca was in the desired position. All of these features illustrate that discocystinid edrioasteroids have highly–evolved morphology and function.     

Ambulacral growth allometry in edrioasteroids: functional surface‐volume change in ontogeny and phylogeny

McKinney, M. L. & Sumrall, C. D. 2010: Ambulacral growth allometry in edrioasteroids: functional surface‐volume change in ontogeny and phylogeny. Lethaia, Vol. 44, pp. 102–108. Most organisms do not maintain geometric similarity as they grow, in large part because of surface‐volume interactions. Because respiratory and food‐acquiring organs are dependent on surface area, which increases more slowly than volume, organisms have evolved many strategies to increase the efficiency of, and/or the functional surface areas of these organs. Here, we report some preliminary results comparing area of the feeding apparatus (ambulacra) versus the volume of the theca for a suite of isorophid edrioasteroid species at various ontogenetic sizes. Regression of log (ambulacral area) on log (thecal volume) indicates a strongly constrained allometric pattern whereby the ontogenies of all measured species fall on or near the same line (r² = 0.962, n = 55), with a slope (k) = 0.654 (± 0.018). This slope and associated 0.05 standard error (0.618–0.690) is within the bounds of that expected from increasing surface area to keep pace with the metabolic demands of increasing thecal size. This allometric value is also comparable to the size scaling of metabolism and respiratory and feeding surfaces in many living and a few fossil invertebrates (k ～ 0.5–0.9). The edrioasteroid species analysed show a very wide variety of changes in ambulacral length, width and sinuosity, documenting many different patterns for achieving the same goal of increasing surface area to keep pace with volume increase. There is no evidence of increasing feeding efficiency. □Edrioasteroid, allometry, ambulacrum, surface‐volume, echinoderm.     

Reinterpretation of thecal plate homology and phylogeny in the Class Crinoidea

Since the Class Crinoidea was erected in 1821 there has been a consistent failure to resolve the phylogeny of this major group on even the coarsest scale. Reinterpretation of crinoid thecal plate homology, using the orientation of the stem rather than the position of the arms as a reference point for the theca, indicates that two-circlet (monocyclic) crinoids may be derived from three-circlet (dicyclic) taxa by the loss of any one of the three plate circlets in the theca rather than just the lowest circlet as has been assumed previously. Cladistic analysis utilizing this new homology, which is supported by evidence from ontogeny and from the position of other plates in the theca, suggests that Aethocrinus is sister group to all other crinoids and that the Cladida are primitive sister group to both the Disparida and Camerata. The Disparida, Hybocrinida and Perittocrinidae together represent a monophyletic clade. The Camerata also are monophyletic, but the orders Monobathrida and Diplobathrida can no longer be considered to represent natural taxa, with two-circlet camerates probably having evolved more than once. This reinterpretation of thecal plate homology sheds new light on the relationships between the major crinoid groups and the pattern of early crinoid evolution. □ Echinodermata, Crinoidea, homology, phylogeny.     

ISOLATED GRAPTOLITES FROM THE LITUIGRAPTUS CONVOLUTUS BIOZONE (SILURIAN, LLANDOVERY) OF DALARNA, SWEDEN

Abstract:  Isolated material of 13 graptolite species from the Aeronian (middle Llandovery) Lituigraptus convolutus Biozone is described. A considerable amount of late astogenetic peridermal thickening is revealed in Normalograptus scalaris and Rivagraptus bellulus . As a result, in the former, thecal morphology is modified from climacograptid to pseudoglyptograptid; in both species, the virgella becomes robust. In Metaclimacograptus minimus and Me. sp., it is shown that the dorsal metathecal wall forms the genicular hood, whereas in N. nikolayevi , the infragenicular wall of the succeeding theca forms the distal thecal apertural margin. Pribylograptus argutus exhibits typical pribylograptid thecae along the length of the available rhabdosome fragments. Characters differentiating Campograptus lobiferus from C. harpago include the greater dorso-ventral width and more rapid increase in dorso-ventral width of the former and the greater recurving of distal thecae and presence of thecal spines/processes on all thecae of the latter. Lituigraptus convolutus has rastritiform thecae proximally; thecal apertures throughout the rhabdosome are crescentic and laterally expanded.     

Devonian features in Ludlovian graptolites — no guide to ancestry

A diverse group of Ludlovian Monograptus species is characterized by the retarded growth of thecal hoods, either proximally [ M. colonus (Barrande), M. chimaera (Barrande) and M. haupti Kühne] or throughout the rhabdosome [M. micropoma (Jaekel)]. In all these species the hoods are secondary structures, i.e. later growth additions to the thecae. By contrast, in the Devonian Monograptus species the thecal hoods are primary structures, i.e. the outward extension of the dorsal thecal wall. and are formed by undelayed growth that typically is completed before growth of the succeeding theca begins. It is among Ludlovian ( M. uncinatus Tullberg) and Pridolian ( M. simih Přibyl) forms with homologous hoods that the roots of the majority of the Devonian Monograptus species might be found. The Ludlovian M. micropoma and M. haupti should not be affiliated to the group of Monograptus vomerinus (Nicholson), which is highly characteristic of, and restricted to, the late Llandovery and Wenlock graptolite faunas.     

贵州寒武纪始海百合类萼部形态定量评价及其意义

始海百合类是贵州寒武纪杷榔动物群和凯里生物群代表性化石类群。产出杷榔动物群和凯里生物群的杷榔组和凯里组主要由灰绿色、灰色的泥、页岩组成,始海百合类化石保存在2个组中上部的泥、页岩中,化石保存好,丰度高,是陆棚斜坡较深水环境生态类群的典型代表。本文基于来自杷榔组的36块Guizhoueocrinus yui,29块Protogloboeocrinus yini标本,和凯里组的20块Sinoeocrinus lui,20块Globoeocrinus globulus标本,采用几何形态测量方法和传统形态测量方法,对每块标本萼部选定了4个界标点(landmark),进行数理统计和几何形态的定量分析。发现寒武纪始海百合类的萼部尺寸上差异较小,在形态上,杷榔组的Protogloboeocrinus yini和Guizhoueocrinus yui萼形相似,均偏细长形,而凯里组的Sinoeocrinus lui和Globoeocrinus globulus萼形差异较大,口部区域宽度均出现增大,萼茎连接处宽度则出现变大和变小两种趋势。结果说明:同一生态环境下寒武纪早期的Guizhoueocrinus yui,和中期的Sinoeocrinus lui的萼在演化上具连续性,细长的萼与长茎能够让始海百合个体更有效捕食上部水域的食物,而寒武纪早期的Protogloboeocrinus yini和中期的Globoeocrinus globulus的萼在形态上具有演化上的连续性。球状的萼配合粗短的茎,可以增强始海百合个体的稳定性,以适应更强的水动力环境,表明出现了生态位的扩张。     

The skeletal structure of Desmophyllum cristagalli: the use of deep-water corals in sclerochronology

Skeletal banding has been found in the deep-water scleractinian coral Desmophyllum cristagalli , an important animal in studies of climate change. This banding pattern sheds light on skeletogenesis and suggests methods by which the record of climate change contained within the coral skeletons may be interpreted. A central wall built of trabeculae forms the interior of the septa and rings the theca. Lamellae form a sheath over the trabecular frame, showing continuity from thecal edge to septum. Skeletal bands are added by the tissue layer, which overlaps and seals the internal coral and upper portion of the outer theca. Truncated inner bands on the outer theca indicate a pattern of skeletal deposition and dissolution dependent on the presence or absence of the live tissue layer. A long-term record will be difficult to collect from D. cristagalli since lamellae are less than 10 μm thick and band position is unpredictable. Density banding in shallow-water coral skeletons has long been recognized as a valuable paleo-oceanographic tool, and deep-water corals are now being used to reconstruct deep-ocean environments. Pattern of skeletal growth must be carefully considered if deep-water corals are to be used as proxy climate recorders.     

Smooth muscle cells in the walls of ovarian follicles in the Japanese quail

Summary The walls of pre-ovulatory follicles of the Japanese quail were examined at the ultrastructural level for the presence of cells displaying the typical morphological features of smooth muscle cells. These characteristics were found in the cells of the chordae, the tunica albuginea, and the theca externa. Small, elongated cells, containing microfilaments, were observed in the theca of prelampbrush follicles localized in the ovarian cortex. These thecal cells were considered as the putative precursors of the thecal smooth muscle cells of the pre-ovulatory follicle. The difference between the smooth muscle cells of the pre-ovulatory follicle and those in the wall of the most recent post-ovulatory follicle is the contracted state of the latter, which is most evident in the cells of the theca externa. It can be concluded that the cells of the theca externa are smooth muscle cells which are mainly contracted during the ovulatory process. A comparison was made with other vertebrate species.     

DIVERSITY AND NICHE EVOLUTION ALONG ARIDITY GRADIENTS IN NORTH AMERICAN LIZARDS (PHRYNOSOMATIDAE)

Deserts occupy approximately 12% of the Earth's land surface, and are thought to have species poor but highly specialized biotas. However, few studies have examined the evolutionary origins of desert biotas and of diversity patterns along aridity gradients. Further, it is unclear if species occurring in more extreme conditions on a given niche axis (i.e., precipitation) are more specialized for those conditions (i.e., have narrower niche breadths). We address these questions here using a time‐calibrated phylogeny and climatic data for 117 species of phrynosomatid lizards. Phrynosomatids are the most species‐rich family of lizards in North America, and are found from deserts to rainforests. Surprisingly, we find that phrynosomatids have higher richness in more arid environments. This pattern occurs seemingly because they have been present in more arid habitats longer (～55 million years), and lineages in mesic environments are recently derived from more arid‐dwelling ancestors. We find little support for the hypothesis that species in more extreme environments are more specialized. Instead, many desert‐dwelling species are broadly distributed, and species in the most mesic environments have the broadest niche breadths. In summary, phrynosomatids offer a counterexample to the idea that arid regions are inhabited by a small number of recent and highly specialized lineages.     

Functional morphology of the theca interna of the vesicular follicle in human ovary]

A study of histological serial sections of human ovaries and histochemical reactions established that several cell types differentiate in the theca interna during the formation and development of tertiary follicles (2nd growth period). This leads, in turn, to triple-layering of the theca interna in follicles of the 2nd resting period. The inner thecal layer consists of fusiform cells from the basement membrane, which are apposed to the membrana granulosa and show a strong positive reaction to alkaline phosphatases. A tropic function for the follicular epithelium must be assigned to this layer. The middle layer consists of epitheloid thecal cells, which show a strong positive reaction to 3beta-ol-steroid hydrogenase. These represent the estrogen glands of the follicle. The outer layer of the theca interna, whose transition to the theca externa is indistinct, also has fusiform cells, which are available as reserve material for the differentiation of further epitheloid thecal cells to pre-ovulatory follicles in later periods of development.     

Mast cell histamine and ovarian follicular growth in the lizard Anolis carolinensis.

The hypothesis is proposed that histamine released from mast cells in the theca of ovarian follicles increases thecal hyperemia and vessel permeability, and thus plays a role in follicular growth in Anolis carolinensis. Mast cells are present in the stroma and theca, and the number of thecal mast cells increases as follicles grow. The levels of histamine in follicular walls varies with follicular size. Histamine causes vasodilatation of thecal vessels. Antihistamine blocks the effects of histamine and, when given alone, stimulates vasoconstriction. Antihistamine also blocks estradiol-induced growth of large follicles. These findings are consistent with the above hypothesis.     

Ultrastructural changes in the theca interna during follicular atresia in sheep

Sixteen antral follicles, 1.8--4.2 mm in diameter, at various stages of atresia, were studied by electron microscopy. Deletion of theca interna cells by condensation followed by fragmentation (apoptosis), with subsequent engulfment of the fragments by remaining thecal cells, was present at all stages, but was most marked during secondary and tertiary atresia. In primary and secondary atresia, the relative numbers of thecal cells whose cytoplasm was rich in tubular endoplasmic reticulum were higher than in non-atretic follicles of comparable size. During tertiary atresia the number of cell layers in the theca interna was reduced, and cells rich in tubular endoplasmic reticulum became proportionately less numerous. Degenerating cellular material was present within the lumina of thecal capillaries at all stages of atresia. Such material was rarely seen in primary atresia, and increased in incidence progressively in later stages. It was concluded that during atresia a large number of theca interna cells are deleted by apoptosis, and many thecal capillaries become blocked with cellular debris.     

Hemichordate skeletal growth: shared patterns in Rhabdopleura and graptoloids

Rhabdopleura shows several features of skeleton growth that are also seen in graptoloids. The similarities between the growth patterns, in terms of the plan towards which the zooids aimed and of their response to environmental disturbance, are profound. Both demonstrate a high degree of genetic control, not only on the gross morphology of the tube or theca but also on the pattern of increments by which this must be achieved. The execution of this 'blueprint' is facilitated by spatial awareness in the zooids of both groups. The main variable left to the graptoloid zooid was the number of increments used in building a theca. Variations in the number of increments probably reflect differences in the productivity of the environment and hence the amount of spare energy in the colony budget. An important new observation is that mortality is common amongst zooids in both Rhabdopleura and graptoloids, with new animals taking over tube or thecal building from where the previous zooid left off. This is identifiable in the increment patterns of tubes and thecae. Several generations of zooids can inhabit a rhabdopleuran tube and can be demonstrated to have inhabited a graptolite theca. This means that innate senescence was not a major cause of death for graptolite colonies. It also means that all thecae might have been continuously occupied and that the colony could have survived significantly bleak environmental conditions by large-scale zooid mortality followed by regeneration. □ Graptolite, ecology, hemichordate, pterobranch, RHABDOPLEURA, growth.     

A STUDY OF CELL WALL AND FLAGELLA FORMATION DURING CELL DIVISION IN THE SCALY GREEN ALGA,SCHERFFELIA DUBIA (CHLOROPHYTA)1

The thecate green flagellate Scherffelia dubia (Perty) Pascher divides within the parental cell wall into two progeny cells. It sheds all four flagella before cell division, and the maturing progeny cells regenerate new walls and flagella. By synchronizing cell division, we observed mitosis, cytokinesis, cell maturation, flagella extension, and cell wall formation via differential interference contrast microscopy of live cells and serial thin‐section EM. Synthesis of thecal and flagellar scales is spatially and temporally strictly separated. Flagellar scales are collected in a pool during late interphase. Before prophase, Golgi stacks divide, flagella are shed, the parental theca separates from the plasma membrane, and flagellar scales are deposited on the plasma membrane near the flagellar bases. At prophase, Golgi bodies start to synthesize thecal scales, continuing into interphase after cytokinesis. During cytokinesis, vesicles containing thecal scales coalesce near the cell posterior, forming a cleavage furrow that is initially oriented slightly diagonal to the longitudinal cell axis but later becomes transverse. After the progeny nuclei have moved into opposite directions, resulting in a “head to tail” orientation of the progeny cells, theca biogenesis is completed and flagellar scale synthesis resumes. Progeny cells emerge through a hole near the posterior end of the parental theca with four flagella of about 8 μm long. The precise timing of flagellar and thecal scale synthesis appears to be an evolutionary adaptation in a scaly green flagellate for the thecal condition, necessary for the evolution of the phycoplast and thus multicellularity in the Chlorophyta.     

Ultrastructure of the theca interna of ovarian follicles in sheep

Summary The theca interna of non-atretic ovarian follicles from 2.0 mm in diameter up to the stage shortly following ovulation was studied by light and electron microscopy.In follicles <3.0mm in diameter, the theca interna consisted of about 8–12 layers of flattened cells, together with many capillaries and small bundles of collagen. Two main forms of cellular differentiation were seen. These were towards either fibroblast-like cells or presumed steroidogenic cells whose cytoplasm contained large amounts of predominantly smooth tubular endoplasmic reticulum, to which some ribosomes were attached. The majority of cells were of relatively undifferentiated or intermediate structure.In larger follicles up to the early stages of oestrus the theca interna cells became larger and less flattened, and cells rich in tubular endoplasmic reticulum became proportionately more numerous. By 18 h after the onset of oestrus the theca interna was oedematous, and many cells possessed pseudopodia. Many cells also contained numerous lipid droplets, but there were no signs of thecal cell degeneration or death. Shortly after ovulation the basal lamina of the membrana granulosa was incomplete, and it became more difficult to distinguish between theca and granulosa layers. Structural heterogeneity, with two major cell types and cells of intermediate structure, was present at all stages.It was concluded that: (1) the theca interna of 2.0–2.9 mm follicles contained many cells whose structure was compatible with a steroidogenic capacity; (2) changes in the differentiated thecal cells up to the early stages of oestrus were quantitative rather than qualitative, and suggestive of an increased steroidogenic capacity; (3) the accumulation of lipid in many cells of the theca interna by 18 h after the onset of oestrus probably reflected a reduction in steroidogenic activity; and (4) there was no evidence of any structural specialization to facilitate the transport of steroids from the theca interna to the membrana granulosa.     

Paired pleural disks in Dicaulograptus cumdiscus n. sp.

A diplograptid graptolite with strongly introverted thecae and unusual thecal processes was isolated from limestones of the Ordovician Pratt Ferry Formation of Alabama and is described as a new species Dicaulograptus cumdiscus. The thecal processes are referred to as pleural disks. They occur in pairs above every thecal aperture and are inferred to have functioned as suspension processes receiving uplift from small-scale turbulent water motion. Such a function is compatible with a passively buoyant mode of life and is difficult to reconcile with an automobile mode of life.     

Interrelationship of growth differentiation factor 9 and inhibin in early folliculogenesis and ovarian tumorigenesis in mice

To investigate the interrelationship of inhibin alpha and growth differentiation factor 9 (GDF9) during early folliculogenesis, we generated mice lacking both inhibin alpha and GDF9. Our findings on these Inha Gdf9 double-mutant mice are as follows: 1). females develop ovarian tumors and a cachexia-like wasting syndrome, resembling mice lacking inhibin alpha alone. This indicates that the granulosa cells are competent to proliferate despite the lack of GDF9; 2). follicular development progresses to multiple-layer follicle stages before tumorigenesis. This demonstrates that the up-regulation of inhibin alpha in the Gdf9 knockout ovary directly prevents the proliferation of the granulosa cells at the primary follicle stage, an effect that is released in the absence of inhibin alpha; 3). a morphological theca forms around the preantral follicles with no detectable selective theca markers [i.e. 17alpha-hydroxylase (Cyp17), LH receptor (Lhr), and Kit]. These results indicate that the theca recruitment can occur independently of GDF9, but the differentiation of thecal cells is blocked; and 4). inhibin/activin subunits betaA, betaB, and Kit ligand (Kitl) mRNA are highly up-regulated, suggesting that the increased activins and KITL play functional roles in early folliculogenesis. Thus, GDF9 appears to function indirectly to regulate early granulosa cell proliferation and theca recruitment in vivo.     

Phylogenetic relationships within the turacos (Musophagidae)

Cytochrome b sequences were used to investigate both the systematic position of the Musophagidae and the species relationships within the family. Phylogenetic analyses (neighbour-joining, maximum likelihood and maximum parsimony) supported the hypothesis that the Musophagidae are more closely related to the cuckoos, parrots and the Hoatzin than the gamebirds. Within the family, three major clades are evident that correspond to the three subfamilies generally recognized. The 'grey turacos' and the turacin-bearing turacos form two separate monophyletic groups. Corythaixoides leucogaster appears to be basal within the subfamily Criniferinae (the 'grey turacos'). This analysis yields a hypothesis for the controversy surrounding the position of Gallirex porphyreolophus and Tauraco johnstoni, which appear to be closely related and basal to the other turacin-bearing turacos. The Musophaga species are placed within the Tauraco clade and appear to be specialized forms of typical turacos. The superspecies persa forms a monophyletic group, within which T. hartlaubi falls. This is an association that has not been suggested previously.     

Oral Region Homologies in Paleozoic Crinoids and Other Plesiomorphic Pentaradial Echinoderms

The phylogenetic relationships between major groups of plesiomorphic pentaradial echinoderms, the Paleozoic crinoids, blastozoans, and edrioasteroids, are poorly understood because of a lack of widely recognized homologies. Here, we present newly recognized oral region homologies, based on the Universal Elemental Homology model for skeletal plates, in a wide range of fossil taxa. The oral region of echinoderms is mainly composed of the axial, or ambulacral, skeleton, which apparently evolved more slowly than the extraxial skeleton that forms the majority of the body. Recent phylogenetic hypotheses have focused on characters of the extraxial skeleton, which may have evolved too rapidly to preserve obvious homologies across all these groups. The axial skeleton conserved homologous suites of characters shared between various edrioasteroids and specific blastozoans, and between other blastozoans and crinoids. Although individual plates can be inferred as homologous, no directly overlapping suites of characters are shared between edrioasteroids and crinoids. Six different systems of mouth (peristome) plate organization (Peristomial Border Systems) are defined. These include four different systems based on the arrangement of the interradially-positioned oral plates and their peristomial cover plates, where PBS A1 occurs only in plesiomorphic edrioasteroids, PBS A2 occurs in plesiomorphic edrioasteroids and blastozoans, and PBS A3 and PBS A4 occur in blastozoans and crinoids. The other two systems have radially-positioned uniserial oral frame plates in construction of the mouth frame. PBS B1 has both orals and uniserial oral frame plates and occurs in edrioasterid and possibly edrioblastoid edrioasteroids, whereas PBS B2 has exclusively uniserial oral frame plates and is found in isorophid edrioasteroids and imbricate and gogiid blastozoans. These different types of mouth frame construction offer potential synapomorphies to aid in parsimony-based phylogenetics for exploring branching order among stem groups on the echinoderm tree of life.     

Morphology and phylogenetic interpretation of a new Cambrian edrioasteroid (Echinodermata) from Spain

Abstract: The edrioasteroid, Aragocystites belli gen. nov. sp. nov. from the middle Cambrian Murero Formation of Spain, is described based on a small number of very well‐preserved specimens. Important anatomical characteristics include star‐shaped and pseudoclavate theca, rare or absent epispires, well‐developed interradially positioned oral plates and several unorganized cover plates associated with each widely exposed flooring plate. A phylogenetic analysis including several Cambro–Ordovician species shows it is more derived than Stromatocystites and Totiglobus but is a sister group to a clade comprising Cambraster and Edriodiscus. Ontogenetic observations based on juveniles of 5 mm in diameter suggest that this species changed thecal shape markedly during growth. A. belli gen. nov. sp. nov. probably lived in quiet environments where it attached directly to the sea floor on stabilized substrates.