Shallow Traces (Pits) in the Test of the Irregular Echinoid Echinocorys scutata Leske from the Chalk (Upper Cretaceous) of the United Kingdom

Specimens of epifaunal irregular echinoids in the Upper Cretaceous of northern Europe have been reported with patterns of circular, nonpenetrative parabolic pits, Oichnus paraboloides (Bromley), in the apical region. Specimens of Echinocorys scutata Leske from the Chalk at two sites in southeast England were commonly penetrated by this trace, generated by an indeterminate pit-forming organism. Pits commonly surround the apical system and, less commonly, occur within it; they occur preferentially anteriorly. Pits occur within plates, not along margins or sutures. Crosscutting of pits indicates that multiple spatfalls probably occurred. The host echinoid added new test plates adjacent to the apical system; thus, plates bearing O. paraboloides were moved abapically. The reduction in number of pits away from the apex, including those with echinoid tubercles reestablished on the base, indicates that, following death of an infester, the echinoid “reclaimed” and infilled them with calcite. The pit-former was most probably an unmineralized invertebrate that used E. scutata as a domicile which provided good access to food-rich currents for suspension feeding. Although the systematic position of the pit-former is unknown, similar infestations are known from other Upper Cretaceous echinoids and Mississippian crinoids.     

Testing reduced evolutionary rates during the Late Palaeozoic Ice Age using the crinoid fossil record

In 2003, Stanley & Powell reported depressed rates of origination and extinction in marine invertebrates during the Late Palaeozoic Ice Age (LPIA). Using a database of crinoid genera, rates of origination, extinction and genus duration were calculated at the stage level from the Early Devonian to the Late Permian. This 165 m.y. time span includes non‐glacial intervals before and after the LPIA, which spanned the Serpukhovian to Sakmarian, providing background rates for comparison. Data generated on crinoid evolutionary rates during the Middle to Late Palaeozoic were analysed and compared to Stanley & Powell's data to determine whether crinoid evolutionary patterns support their findings or suggest an alternative hypothesis. Rates of origination and extinction in all crinoid clades were reduced during the LPIA compared to the combined background intervals before and after the LPIA. However, crinoid diversity was higher during the LPIA than the surrounding time intervals. The difference in diversity trends between crinoids and other marine invertebrates is due to the advanced cladids clade. Unstable, fluctuating environmental conditions during the LPIA may have created habitats suitable for opportunistic crinoid genera that reduced both the probability of origination and extinction. The increased diversity of the advanced cladids is likely due to their unique adaptation of muscular arm articulations, which allowed them to thrive in marine settings with increased siliciclastic influx brought on by the Alleghenian orogeny. Despite the advanced cladids’ departure from the expected diversity count, the results of analyses performed on the updated crinoid database provide independent confirmation of Stanley & Powell's original hypothesis of depressed evolutionary rates in marine invertebrates during the LPIA.     

Lucky rugose corals on crinoid stems: unusual examples of subepidermal epizoans from the Devonian of Morocco

Berkowski, B & Klug, C. 2011: Lucky rugose corals on crinoid stems: unusual examples of subepidermal epizoans from the Devonian of Morocco. Lethaia, Vol. 45, pp. 24–33. In the fossil record, evidence for true epizoans, i.e. living animals inhabiting other living host‐animals, is rather rare. A host reaction is usually needed to proof the syn vivo‐settling of the epizoan. Herein, we provide a first report of such an epizoan biocoenosis from various strata of the Early Devonian of Hamar Laghdad, the world‐renowned Moroccan mud‐mound locality. In this case, solitary rugose corals settled as larvae on crinoid stems, perhaps at a spot where the epidermis was missing for some reason (injury, disease). Both the crinoid and the coral began to grow around each other. By doing so, the affected crinoid columnals formed a swelling, where ultimately only an opening slightly larger than the coral orifice remained. We discuss both macroecological and small‐scale synecological aspects of this biocoenosis. The coral profited from its elevated home because it reached into more rapid currents providing the polyp with more food than at the densely populated seafloor, which was probably covered by a coral‐meadow around the mounds and hydrothermal vents. □Corals, crinoids, Early Devonian, epizoans, Morocco, Rugosa.     

Host-specific pit-forming epizoans on Silurian crinoids

Brett, Carlton E. 197807 15: Host-specific pit-forming epizoans on Silurian crinoids. Lethaia , Vol. 11. pp. 217–232. Oslo. ISSN 0024–1164.
Circular-parabolic pits occur commonly on the endoskeletal remains of certain Paleozoic crinoids. Detailed study of several hundred specimens, representing about 30 pelmatozoan species from the Upper Silurian Rochester Shale of New York and Ontario, reveals that such pits occur exclusively in seven species of crinoids. Furthermore, there are consistent differences in the morphology and orientation of holes occurring on the different crinoid species. This suggests that distinct epizoan species settled selectively on given hosts. The relationship between the hole-producing epizoans and crinoid hosts is inferred to have been a form of dependent commensalism. Preliminary surveys of other Paleozoic crinoid assemblages reveal similar host-selectivity by pit-producing epizoans. Crinoidepizoan pairs apparently co-evolved through considerable spans of geologic time as related genera and species of different ages, from Silurian to Pennsylvanian, exhibit similar pits.     

Fixed,free, and fixed: The fickle phylogeny of extant Crinoidea (Echinodermata) and their Permian–Triassic origin

Although the status of Crinoidea (sea lilies and featherstars) as sister group to all other living echinoderms is well-established, relationships among crinoids, particularly extant forms, are debated. All living species are currently placed in Articulata, which is generally accepted as the only crinoid group to survive the Permian–Triassic extinction event. Recent classifications have recognized five major extant taxa: Isocrinida, Hyocrinida, Bourgueticrinina, Comatulidina and Cyrtocrinida, plus several smaller groups with uncertain taxonomic status, e.g., Guillecrinus, Proisocrinus and Caledonicrinus. Here we infer the phylogeny of extant Crinoidea using three mitochondrial genes and two nuclear genes from 59 crinoid terminals that span the majority of extant crinoid diversity. Although there is poor support for some of the more basal nodes, and some tree topologies varied with the data used and mode of analysis, we obtain several robust results. Cyrtocrinida, Hyocrinida, Isocrinida are all recovered as clades, but two stalked crinoid groups, Bourgueticrinina and Guillecrinina, nest among the featherstars, lending support to an argument that they are paedomorphic forms. Hence, they are reduced to families within Comatulida. Proisocrinus is clearly shown to be part of Isocrinida, and Caledonicrinus may not be a bourgueticrinid. Among comatulids, tree topologies show little congruence with current taxonomy, indicating that much systematic revision is required. Relaxed molecular clock analyses with eight fossil calibration points recover Articulata with a median date to the most recent common ancestor at 231–252 mya in the Middle to Upper Triassic. These analyses tend to support the hypothesis that the group is a radiation from a small clade that passed through the Permian–Triassic extinction event rather than several lineages that survived. Our tree topologies show various scenarios for the evolution of stalks and cirri in Articulata, so it is clear that further data and taxon sampling are needed to recover a more robust phylogeny of the group.     

Diversity dynamics of post‐Palaeozoic crinoids – in quest of the factors affecting crinoid macroevolution

Following the end‐Permian biotic crisis which led to the near extinction of crinoids, this echinoderm class rebounded rapidly during the Mesozoic, resulting in forms with important morphological and behavioural novelties. However, quantitative patterns of crinoid diversity during the Mesozoic remain largely unexplored. Here, we report results of analyses of the evolutionary dynamics of post‐Palaeozoic crinoid genera spanning a time interval between 250 and 70 Myr. We show that crinoids reached their Mesozoic peak of genus‐level richness during the Late Jurassic. We also document a major reorganization of different ecological crinoid groups in the Mesozoic. More specifically, the diversity of sessile forms generally increased towards the mid‐Mesozoic but decreased significantly starting in the Cretaceous, whereas the number of motile crinoid genera increased linearly during the Mesozoic. The possible role of biotic and abiotic factors in crinoid evolution is discussed.     

The potential effects of sex,posture and living condition on lateralized behaviors in ring-tailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

This research explores the effects of posture, sex, and living condition on hand and side preferences of semi-free-ranging, adult ring-tailed lemurs (Lemur catta) housed at the Duke University Primate Center in Durham, NC. Data were collected on 11 adult individuals (five females and six males) during normal daily activities over a ten-week period from May–July 2001. Variables analyzed in this study include unimanual behaviors (i.e., reach, hold, and limb used to start locomotion) and other potentially lateralized behaviors that do not involve handuse (i.e., whole-body turning and tail position). The data were analyzed to investigate potential individual and population level side biases for each behavior; potential sex biases in side preference for each behavior; and for ‘reach’, potential effects of posture (sitting, tripedal stance, or bipedal stance) on individual hand preferences. Additionally, to investigate potential effects of living condition on lateral biases, the data from this study were compared to data collected on the same individual Lemurs living under more restrictive living conditions during the previous year. Largely, as predicted based on available literature, we found that there was a significant sex difference across all hand-use categories and for whole-body turning, and that posture was a significant factor in the expression of hand preference for reaching. Contrary to previous research, the effect of living condition on lateral preferences was minimal, and no side preferences were found at the population level for any of the behaviors analyzed.     

Survival of crinoid stems following decapitation: evidence from the Ordovician and palaeobiological implications

It was recently discovered that the stems of extant crinoids may survive after detachment of the crown, presumably feeding by the absorption of nutrients through the ectoderm. Previously, only one analogous, albeit morphologically dissimilar, pattern of crownless survival has been recognized from the fossil record. Certain Upper Ordovician (Cincinnatian) crinoid pluricolumnals from Kentucky, Ohio and Indiana, derived from the disparids Cincinnaticrinus spp., have rounded terminations reminiscent of some modern bourgueticrinid overgrowths. Such specimens have hitherto been interpreted as distal terminations of mature individuals that have become detached from their attachment structures and taken to an eleutherozoic existence. However, it is considered more probable that they represent overgrowths of the column following predatory decapitation. If this new interpretation is correct, then post-decapitation survival of crinoid stems is now recognized for most of the history of the crinoids, 'lethal' predation on crinoid crowns occurred at least as early as the Late Ordovician and ancient crinoid populations can no longer be determined merely by counting crowns.     

Unusual tabulate‐crinoid biocoenosis from the Lower Devonian of Morocco

The Early Devonian (Pragian: sulcatus to pireneae conodont zones) crinoid–coral biocoenosis from Hamar Laghdad, Morocco contains fragments of crinoid stalks of various taxa encrusted by spherical and ellipsoidal coralla of the tabulate coral Hamarilopora minima. These corals were encrusting host crinoids syn vivo, and this is evidenced by pluricolumnals exceeding 30 elements overgrown from all sides. Most known to date crinoid–epibiont associations display various types of reaction to the epibiont, such as swellings and deformations. In the case discussed here, no clear interaction is visible; therefore, this association can be classified as paroecia. It can be inferred, however, that due to a change in mechanical properties of the crinoid stalk (losing flexibility), the epizoan influence on the host was negative, while the coral was profiting from the elevated position over the seafloor and nutrient‐bearing water currents. It can be supposed that this interaction was close to parasitism. No strict species‐specific relationship between the epizoan and the host was observed.     

The ichnogenus Tremichnus in the Lower Silurian of western New York

Discovery of Tremichnus cysticus Brett on a crinoid pluricolumnal from the Willowvale Shale of western New York extends the range of this ichnogenus into the Lower Silurian. Pits attributable to Tremichnus apparently represent embedment sites of parasites or commensalistic, filter-feeding organisms of unknown affinities. Embedment of T. cysticus may have been accomplished by localized inhibition of stereom growth within the pits aided by secretion of substances that stimulated accretion of stereom by the host pelmatozoan in areas immediately surrounding the pits.     

Tube foot preservation in the Devonian crinoid Codiacrinus from the Lower Devonian Hunsrück Slate,Germany

Fossilized tube feet are described on Codiacrinus schultzei Follmann from the Lower Devonian Hunsrück Slate of Germany. This is the first definitive proof of tube feet on any fossil crinoid. Three lightly pyritized, flattened tube feet are preserved in a single interray of this cladid crinoid. The tube feet were at least 7 mm long. Their preservation is very similar to the tube feet reported previously from a Hunsrück ophiuroid, except that the Codiacrinus tube feet have small papillae, similar to living crinoids.     

Assessment of hidden diversity of crinoids and their symbionts in the Bay of Nhatrang,Vietnam

Crinoid associates represent an abundant and diverse, but poorly explored, component of the hidden biodiversity of coral-reef ecosystems. We studied data from 5 years of collecting in the Bay of Nhatrang (BN), Vietnam, to assess the diversity of crinoids and their symbionts, to compare it with other areas of the Indo-West Pacific, and to elucidate the extent to which the observed diversity of crinoids and their symbionts corresponds to their true diversity. In total, about 2,287 specimens of symbionts belonging to 70 species were found on 203 specimens of crinoids belonging to 33 species. Among the crinoids, the most numerous species were Himerometra robustipinna (36 specimens) and Cenometra bella (29 specimens), among the symbionts the polychaete Paradyte crinoidicola (c. 850 specimens) and the galatheid crustacean Allogalathea elegans (180 specimens). Species accumulation curves suggest that we have sampled most of the crinoid diversity in the BN, whereas the diversity of their symbionts remained undersampled. Estimated species richness of crinoids was higher than previously observed richness, and varied from 39 (estimated by bootstrap) to 46 (jackknife 2). Estimated species richness of symbionts was higher than observed richness, and varied from 71 (bootstrap) to 93 (jackknife 2). We suggest a slight increase in the number of crinoid species to result from more detailed studies of nocturnal species, and an increase in the number of symbiotic species when studies of nocturnal crinoid associates and sibling species among decapods are included. Our study revealed a rather rich crinoid fauna in the bay compared to other areas of the Indo-West Pacific, and the highest species richness of crinoid associates known from anywhere in the World Ocean.     

New data from Oman indicate benthic high biomass productivity coupled with low taxonomic diversity in the aftermath of the Permian–Triassic Boundary mass extinction

A new Early Triassic marine fauna is described from an exotic block (olistolith) from the Ad Daffah conglomerate in eastern Oman (Batain), which provides new insights into the ecology and diversity during the early aftermath of the Permian–Triassic Boundary mass extinction. Based on conodont quantitative biochronology, we assign a middle Griesbachian age to the upper part of this boulder. It was derived from an offshore seamount and yielded both nektonic and benthic faunas, including conodonts, ammonoids, gastropods and crinoid ossicles in mass abundance. This demonstrates that despite the stratigraphically near extinction at the Permian–Triassic Boundary, Crinoidea produced enough biomass to form crinoidal limestone as early as middle Griesbachian time. Baudicrinus, previously placed in Dadocrinidae, is now placed in Holocrinidae; therefore, Dadocrinidae are absent in the Early Triassic, and Holocrinidae remains the most basal crown‐group articulates, originating during the middle Griesbachian in the Tethyan Realm. Abundant gastropods assigned to Naticopsis reached a shell size larger than 20 mm and provide another example against any generalized Lilliput effect during the Griesbachian. Whereas the benthic biomass was as high as to allow the resumption of small carbonate factories, the taxonomic diversity of the benthos remained low compared to post‐Early Triassic times. This slow benthic taxonomic recovery is here attributed to low competition within impoverished post‐extinction faunas.     

Haemal and coelomic circulatory systems in the arms and pinnules ofFlorometra serratissima (Echinodermata: Crinoidea)

Summary The haemal and coelomic circulatory systems in arms and pinnules of a stalkless crinoid are described by transmission electron microscopy, and the coelomic topography is revealed by scanning electron microscopy of corrosion casts and peritoneal surfaces. In addition, the route of the coelomic circulation in the living crinoid is shown by injection of carmine particles, and sites of peritoneal phagocytosis are demonstrated by injection of latex beads. The most important morphological findings are: the controversial hyponeural circulation is haemal and not coelomic; peritoneal ciliation is general and not limited to the cells of the ciliated pits; and occur smooth muscle cells occur below the peritoneum. Carmine particles injected into the central body coelom rapidly travel outward toward the arm and pinnule tips via the aboral canals; the particles return to the central body via the subtentacular canals. Latex beads injected intracoelomically are taken up by peritoneal cells throughout the subtentacular, genital and aboral canals. The possible functions of the haemal and coelomic circulatory systems of crinoids are discussed.     

Small pits associated with renilla (Cnidaria: Pennatulacea) on a modern low‐energy sandflat,Kiawah Island,South Carolina,U.S.A.

Small, shallow circular pits containing the Atlantic sea pansy, Renilla reniformis, were fairly common at one locality on the lower foreshore of Kiawah Island in June, 1997. Many of the pits were positioned on the stoss side of sand ripples; the organisms were oriented with their bilateral plane of symmetry essentially parallel to ripple crests. If preserved as concave epireliefs or convex hyporeliefs on bedding surfaces, such pits would be difficult not only to attribute to a trace producer or traditional ethologic category, but to identify as trace fossils in the first place.     

An alternative mechanism of clathrin-coated pit closure revealed by ion conductance microscopy

Current knowledge of the structural changes taking place during clathrin-mediated endocytosis is largely based on electron microscopy images of fixed preparations and x-ray crystallography data of purified proteins. In this paper, we describe a study of clathrin-coated pit dynamics in living cells using ion conductance microscopy to directly image the changes in pit shape, combined with simultaneous confocal microscopy to follow molecule-specific fluorescence. We find that 70% of pits closed with the formation of a protrusion that grew on one side of the pit, covered the entire pit, and then disappeared together with pit-associated clathrin-enhanced green fluorescent protein (EGFP) and actin-binding protein-EGFP (Abp1-EGFP) fluorescence. This was in contrast to conventionally closing pits that closed and cleaved from flat membrane sheets and lacked accompanying Abp1-EGFP fluorescence. Scission of both types of pits was found to be dynamin-2 dependent. This technique now enables direct spatial and temporal correlation between functional molecule-specific fluorescence and structural information to follow key biological processes at cell surfaces.     

Ultrastructure of the mycangium of the southern pine beetle,Dendroctonus frontalis (Coleoptera: Curculionidae,Scolytinae): complex morphology for complex interactions

Yuceer, C, Hsu, C.‐Y., Erbilgin, N and Klepzig, K.D. 2011. Ultrastructure of the mycangium of the southern pine beetle, Dendroctonus frontalis (Coleoptera: Curculionidae, Scolytinae): complex morphology for complex interactions. —Acta Zoologica (Stockholm) 92 : 216–224. The southern pine beetle (SPB) (Dendroctonus frontalis Zimmermann) is the most economically important pest of southern pine forests. Beetles carry fungal cells within specialised cuticular structures, called mycangia. Little is known about the mycangia ultrastructure or function. We used cryo‐fracturing and scanning electron microscopy to examine the ultrastructural features of SPB mycangia and surrounding tissues. Mycangia, one on each side of anterior portion of the prothorax, are terminated on the dorsal side at a ‘mycangial bridge’. This sclerotised mycangial bridge does not appear to provide a passage between the two mycangia, suggesting that each mycangium functions independently. Mycangia are surrounded by abundant tracheoles connecting the structures to the outside via openings within the prothorax. Previously unknown pits overlying the mycangial gland cells were also observed in both the inner wall and anterior fold of prothorax. We hypothesise that these openings and pits may play roles in determining which fungi enter, and grow within, the mycangium.     

ANATOMICALLY PRESERVED GLOSSOPTERIS STEMS WITH ATTACHED LEAVES FROM THE CENTRAL TRANSANTARCTIC MOUNTAINS,ANTARCTICA

Stems and buds of Glossopteris skaarensis Pigg and buds of G. schopfii Pigg from the Permian Skaar Ridge locality in the central Transantarctic Mountains, Antarctica demonstrate the first anatomically preserved glossopterids known with stem/leaf attachment. Stems of G. skaarensis are 1–12 mm in diameter ( = 3.1 mm) with a broad pith, poorly defined primary xylem, and a zone of secondary xylem up to 6 mm thick. Pycnoxylic wood conforming to Araucarioxylon Kraus is composed of tracheids with uni- to biseriate oval to hexagonal bordered pits on radial walls, uniseriate rays one to a few cells high, and cupressoid to taxodioid cross-field pitting. Stems have a narrow zone of secondary phloem, aerenchymatous cortex with scattered sclereids, and sometimes a narrow periderm. Two wedge-shaped leaf traces each bifurcate to form four strands in the base of each petiole. Small axillary branches are vascularized by double branch traces that fuse at the margin of the main axis. Buds of G. skaarensis have leaves with narrow lateral laminae and a thickened midrib containing a wide lacuna, delicate vascular strands, and a prominent hypodermis. In contrast, buds of G. schopfii have uniformly thick leaves with prominent, circular vascular bundle sheaths. These anatomical details are used to reconstruct individual types of glossopterid plants, providing new information toward understanding the ecology and evolution of this important group of Permian seed plants.     

Aspects of crinoid palaeontology of the North Esk Inlier,Scotland (Silurian,Llandovery, Telychian)

Abstract: The described fauna of well‐preserved Llandovery (Telychian) echinoderms from the North Esk Inlier, including six crinoids, one echinoid and seven starfish species, is mainly allochthonous. Most of these taxa are known only from starfish beds, channel fill deposits probably representing submarine mass flows and preserving a biota probably derived from elsewhere, presumably shallower water. Only one crinoid species, Pisocrinus cf. campana Miller, is recognized as a common fossil away from the starfish beds and is a biostratigraphic marker for the base of the Wether Law Linn Formation, forming part of the Skenidioides‐Cyrtia Association. Crinoid columnals preserved perpendicular to bedding (that is, in putative life position) in Lamont’s bivalve bed, Deerhope Formation, are tentatively interpreted as being in situ by comparison with a similar occurrence in the Silurian of Arisaig, Nova Scotia. Two new species of crinoid are described, the cladid Dendrocrinus? sp. and the columnal morphospecies Pentagonocyclicus (col.) lamonti sp. nov.     

内蒙古早二叠世早期南洋杉型木属一新种

报道了一种具混合型纹孔（互列式纹孔和对列式纹孔）的化石本。化石木产自内蒙古乌达矿区老石旦矿附近太原组上部,地质时代为早二叠世早期。经比较,确认为南洋杉型木属（Ａｒａｕｃａｒｉｏｘｙｌｏｎ Ｋｒａｕｓ）一种新;老石旦南洋杉型木（Ａｒａｕｃａｒｉｏｘｙｌｏｎ ｌａｏｓｈｉｄａｎｅｎｓｅ ｓｐ．ｎｏｖ．）．根据混合型纹孔的存在及其他特征,认为新种可能代表了一种原始的松杉类植物的木材化石。主要特征：仅保存