In deep water: a crinoid–brachiopod association in the Upper Oligocene of Antigua,West Indies

Extant brachiopods and stalked crinoids are found together in the deeper waters of the Caribbean Sea. Analogous brachiopod/crinoid associations have been reported from diverse palaeoenvironments in the Neogene of the region. Studied examples include the Pleistocene of Jamaica (deeper water fore reef), and the Miocene of Jamaica (island slope chalks), Barbados (accretionary prism) and Carriacou (turbiditic siliciclastic shelf). Comparison with analogous modern environments indicates deposition in 150+m water depth. This association has now been extended back into the Late Oligocene. Crinoids and brachiopods both occur in the Antigua Formation of Antigua; both occur high in the formation, implying deeper water in this retrograde succession. They have received little attention from systematists, although the brachiopods Cistellarcula dubia Cooper and Tichosina foresti Cooper have previously been described from the Antigua Formation; to these, we add Cistellarcula sp., Argyrotheca sp. and Tichosina sp. At Half Moon Bay in southeast Antigua, high in the Antigua Formation, we have found columnals of isocrinid crinoids (cf. Isocrinus sp.) associated with rare brachiopods (Terebratulina sp.) in island slope deposits. These taxa provide independent evidence for the deeper water aspect of this part of the Antigua Formation, in beds that also yield large, thin‐walled fossil sponges.     

Epi- and endobiontic organisms on Late Jurassic crinoid columns from the Negev Desert, Israel: Implications for co-evolution

Columns of the articulate crinoids Millericrinus and Apiocrinites from the Upper Jurassic (Upper Callovian) Zohar and Matmor formations of the Negev Desert of Israel display abundant encrusting organisms of about ten species, as well as diverse trace fossils produced by endobionts. Pluricolumnals were colonized by epi- and endobiontic organisms both during life and post-mortem. Skeletonized encrusting organisms include abundant ostreid bivalves (which evidently colonized both live and dead crinoid columnals), two types of serpulid worms, encrusting foraminifera, three species of bryozoans, and small encrusting sclerosponges. Several types of borings are present: Trypanites (possibly produced by sipunculids), Gastrochaenolites (crypts of boring lithophagid bivalves), elliptical barnacle? borings, and channel-like annelid? borings. In addition, approximately 16% of the pluricolumnals display circular parabolic embedment pits assignable to the ichnogenus Tremichnus. They are associated with substantial deformation of the containing columnals and were probably the work of host-specific ectoparasitic organisms. Discovery of Tremichnus on Jurassic crinoids extends the range of this trace by almost 100 million years, providing evidence for one of the longest-ranging host-parasite interactions documented thus far (over 200 million years). The relationship of epibionts to the Jurassic crinoids thus ranged from simple utilization of dead hard substrate to probable opportunistic commensalism in forms that colonized the live upright stems, as in some oysters, through host-specific parasitism in the case of Tremichnus.     

Astogenese vonAulopora cf.enodis Klaamann 1966Visby-Mergel, Silur von Gotland

This paper deals with bifurcation angle of a reptant, dichotomously branchingAulopora cf.enodis colony (Silurian, Gotland). It can be demonstrated, that during early astogenetic growth the branching angle between offsetting corallites decreases rapidly from 165° to 100° averagely during later growth stages. This growth pattern resembles development of similar dichotomous branching uniseral bryozoans, e.g. PaleozoicCorynotrypa species or MesozoicStomatopora species.     

Rare bryozoans (Stenolaemata) with bilateral colonies from the Jurassic and Cretaceous of the East European Platform

New bryozoans characterized by a bilateral vertical colonies colonies—Cardioecia refuga sp. nov. (Tubuliporida), Elea lyapini sp. nov., E. troshkovensis sp. nov., and E. taylori sp. nov. (Melicerititida) from Middle Jurassic (Middle Callovian) of the Moscow region (Russia), and Biforicula legitima sp. nov. (Melicerititida) from the Upper Cretaceous (Lower Campanian) of the Southern Donets Basin (Ukraine)—are described. All species described in this paper belong to genera that have been recorded for the first time in the East European Platform. Some morphological structures in the colonies of these bryozoans and some distinctive features of the order Melicerititida that emphasize the difference of this order from bryozoans of other orders of the class Stenolaemata are examined.     

Phylogeny and life history evolution of Prodoxus yucca moths (Lepidoptera: Prodoxidae)

Abstract. Yucca moths (Lep., Prodoxidae) are well‐known for their obligate pollination mutualism with yuccas. In addition to the pollinators, yuccas also host many non‐pollinating yucca moths. Here the genus Prodoxus, the non‐pollinating sister group of the pollinators, is revised using morphological and molecular data, their phylogenetic relationships are analysed, and the evolution of host tissue specialization explored. Twenty‐two species are recognized, including nine new species: Prodoxus gypsicolor sp.n. , P. sonorensis sp.n. , P. carnerosanellus sp.n. , P. tamaulipellus sp.n. , P. weethumpi sp.n. , P. tehuacanensis sp.n. , P. californicus sp.n. , P. mapimiensis sp.n. and P. atascosanellus sp.n. Prodoxus y‐inversus Riley, P. coloradensis Riley and P. sordidus Riley are redescribed. The genus Agavenema is synonymized with Prodoxus. Phylogenetic analyses indicated that stalk‐feeding is basal within the group, that there are three separate origins of fruit‐feeding, and one origin of leaf‐mining from a stalk‐feeding ancestor. Although species with different feeding habits often coexist within hosts, the analyses suggest that ecological specialization and diversification within a host only may have occurred within one or possibly two hosts.     

Late Famennian gastropoda from south-west England

The gastropod fauna of the Upper Devonian Baggy and Pilton formations in south‐west England is revised and includes some 30 taxa. The topmost part of the Upper Famennian succession in Devon is represented by clastic near‐shore and shallow shelf sediments, indicating a short‐term transgressive phase (‘Strunian Transgression’). The sequence yields a highly diverse fauna dominated by brachiopods and ostracodes, locally supplemented by crinoids, bryozoans, trilobites and molluscs. The taxa ‘Patellostium’britannicum sp. nov., Angyomphalus (Angyomphalus) junius sp. nov. and Dictyotomaria eurocapillaria sp. nov. are erected; a junior homonym is replaced by Macrochilina? piltonensis nom. nov. The gastropod fauna displays an independent character, where latest Devonian faunal elements overlap with Late Palaeozoic taxa expressing a transition similar to that of the bivalves, brachiopods, echinoderms and corals, without a sharp faunal break at the Devonian/Carboniferous boundary. Apart from the Caenogastropoda, all subclasses of gastropods are represented. Members of the bellerophontoids, pleurotomarioids and loxonematoids are most abundant, followed by murchisonioids, naticimorphs, euomphalomorphs and platyceratoids. The various gastropod groups represent different ecological demands and trophic categories, and together with the accompanying fauna indicate that nearly all habitats and niches were occupied in the shallow South Laurussian Shelf.     

UNICELLULAR CYANOBIONTS IN OPEN OCEAN DINOFLAGELLATES,RADIOLARIANS, AND TINTINNIDS: ULTRASTRUCTURAL CHARACTERIZATION AND IMMUNO‐LOCALIZATION OF PHYCOERYTHRIN AND NITROGENASE1

Cyanobacterial symbionts (cyanobionts) have been identified forming associations with various open ocean eukaryotic host genera, including two dinophysoid genera, Histioneis sp. and Ornithocercus sp., two radiolarians, Spongastaurus and Dictyocoryne truncatum, sp., and a tintinnid, Codonella sp. The TEM analysis revealed that single individual hosts were closely associated with one to two different cyanobacterial morphotypes (cyanobionts) and two hosts had in addition to cyanobionts, one to two bacterial cell types. Eleven significantly (P<0.01) different cell types were identified as cyanobionts, with cell diameters ranging 0.5±0.38–3.7±0.66 μm. Using immunogold‐labeling techniques coupled to the TEM, four of the five cell types contained phycoerythrin (PE) at high levels (>71±28 gold particles·μm^?2). Immunolabeling‐TEM using nitrogenase antisera demonstrated a significant (P<0.01) nitrogenase content in cell type four cyanobionts of Histioneis sp. host 1 (39±34 gold particles·μm^?2). The cyanobionts of the radiolarians were of a cell diameter (0.5–0.8 μm) and showed ultrastructural characters (peripheral thylakoids) reminiscent of Prochlorococcus sp. Also, an open ocean tintinnid, Codonella sp., was shown to contain cyanobacteria as symbionts for the first time. In all cyanobionts, glycogen storage was obvious, no cellular degradation was visible, cells were observed in the process of cellular division, and antisera localization was apparent. These observations suggest that the relationship between host eukaryote and cyanobacteria is an active one, and likely symbiotic.     

Hermit crab biocoenoses: a worldwide review of the diversity and natural history of hermit crab associates

The symbiotic associates of hermit crabs (excluding parasites and flora) are reviewed worldwide. The review includes species found on the shells occupied by hermit crabs (epibiotic species), species boring into these shells (endolithic species), species living within the lumen of the shell (either free-living or attached to the shell), species attached to the hermit crabs themselves, and hypersymbionts. In total over 550 invertebrates, from 16 phyla are found associated with over 180 species of hermit crabs. Among these associates, 114 appear to be obligate commensals of hermit crabs, 215 are facultative commensals, and 232 are incidental associates. The taxa exhibiting the highest number of associates are arthropods (126), polychaetes (105), and cnidarians (100). The communities of species associated with Dardanus arrosor, Paguristes eremita, Pagurus bernhardus, Pagurus cuanensis, and Pagurus longicarpus are the best studied and harbor the most diverse assemblages of species. While trends in biodiversity of hermit crab assemblages do not follow predicted patterns (e.g., hermit crabs within the Indo-West Pacific do not harbor more species than those from temperate regions), this is suggested to reflect a lack of sampling rather than a true representation of the number of associates. Hermit crabs date to at least the Cretaceous and provided a niche for a number of groups (e.g., hydractinians, bryozoans, polydorids), which were already associates of living gastropods. Apparently hermit crab shells initially supplied a substrate for settlement and then these symbiotic relationships were reinforced by enhanced feeding of symbionts through the activity of the hosts. Through their use and recycling of gastropods shells, hermit crabs are important allogenic ecosystem engineers in marine habitats from the intertidal to the deep sea. Hermit crabs benefit from some symbionts, particularly cnidarians and bryozoans, through extension of shell apertures (alleviating need to switch into new shells) and by providing protection from predators. However, hermit crabs are also negatively impacted (e.g., decreased reproductive success, increased predation) by some symbionts and a review of egg predators is provided. Thus, the symbiotic relationships between hermit crabs and many associates are difficult to characterize and often exhibit temporal changes depending on environmental and biological factors. Research on the biology of these symbionts and the costs/benefits of their associations with hermit crabs are analyzed. While some associates (e.g., Hydractinia spp.) have been studied in considerable detail, for most associations little is known in terms of the impacts of symbionts on hosts, and future experimental studies on the multitude of relationships are suggested.     

Host‐specific acrothoracid barnacles on Middle Devonian platyceratid gastropods

Borings, attributed to acrothoracic barnacles, occur on the platyceratid gastropod Naticonema lineatum (Conrad) from the Middle Devonian Hamilton Group of western New York and rarely in specimens as old as the Early Devonian. These latter are the oldest known acrothoracid borings are in the fossil record. The borings are consistently developed as laterally compressed, inequilateral pouches exclusively on these gastropods, commonly as dense infestations.

Naticonema shells yielding borings typically occur associated with partially articulated qrinoid remains, and they are sometimes found attached to crinoids in a manner similar to coprophagous Platyceras. In addition to barnacle borings, Naticonema shells often bear thin encrustations of bryozoans which are usually perforated by these borings but sometimes also overgrow them. Barnacles bored live hosts; gastropods prevented shell penetration by producing cyst‐like secondary secretions of calcite beneath acrothoracid boreholes.

The relative antiquity of these borings and their association with coprophagous platyceratids makes their discovery particularly significant in revealing aspects of the early ecology of barnacles. Attachment to the host commensal gastropods was one of the first successful life modes of these crustaceans prior to their later diversification to other habitats. Mississippian and Pennsylvanian occurrences of similarly bored gastropods demonstrate continuity of the barnacle‐gastropod‐crinoid ecological association from the Middle to Late Paleozoic.     

Systematic re‐appraisal of the gall‐usurping wasp genus Synophrus Hartig, 1843 (Hymenoptera: Cynipidae: Synergini)

Several unanswered questions remain regarding the taxonomy and phylogeny of inquiline gallwasps (Cynipidae: Synergini), obligate inhabitants of plant galls induced primarily by other gallwasps (Cynipidae: Cynipini and Diplolepidini). Here we use morphological and molecular data to revise the inquiline genus Synophrus, members of which are notable for extensively modifying the structure of galls induced by oak gallwasp hosts on oaks in the section Cerris of Quercus subgenus Quercus in the Western Palaearctic. Previous taxonomic treatments have recognized three Western Palaearctic species of Synophrus: S. pilulae, S. politus and S. olivieri. Our results support the establishment of four additional Western Palaearctic species: Synophrus hungaricus sp.n. , S. libani sp.n. , S. syriacus sp.n. and S. hispanicus sp.n. We describe and diagnose these new taxa, analyse their phylogenetic relationships, and show that Synophrus inquilines are able to impose their own gall phenotypes on those of their hosts. We provide an updated key to Synophrus.     

Cyrtocrinids (Echinodermata,Crinoidea) from Upper Jurassic Štramberk‐type limestones in southern Poland

Abstract: A systematic account of highly diverse cyrtocrinid faunules from Upper Jurassic strata of ?tramberk type (Oxfordian–Tithonian) in southern Poland (Polish Carpathians) is presented. Fourteen taxa (Phyllocrinus malbosianus, Ph. stellaris, Ph. sp., Psalidocrinus armatus, Sclerocrinus compressus, S. polonicus sp. nov., Hemicrinus aff. kabanovi, Ancepsicrinus parvus gen. et sp. nov., Tetracrinus baumilleri sp. nov., Eugeniacrinites alexandrowiczi, E. cf. moravicus, E. sp., Eudesicrinus gluchowskii sp. nov. and Hemibrachiocrinus tithonicus sp. nov. are described and illustrated. Representatives of the genus Eudesicrinus, previously recorded only from the Lower Jurassic, are here shown to extend into the uppermost Jurassic. Other cyrtocrinids considered are common in Jurassic/Cretaceous strata across Europe. In the present faunules, isocrinid (Isocrinida), comatulid (Comatulida) and roveacrinid (Roveacrinida sensu Rasmussen, inclusive of Saccocoma) crinoids are associated.     

Use of Alternative Food Plants Exclusively by Adult Male Froghoppers (Homoptera: Cercopidae)1

Male-specific associations of adult froghoppers (Homoptera: Cercopidae) with alternative food plant hosts is reported. Four accounts from the field and one from the literature are presented: Iphirhina quota on Bourreria costaricensis (Boraginaceae), Huaina inca on Solanaceae sp., Prosapia simulans and Prosapia sp. (near P. bicincta) on Ilex haberi (Aquifoliaceae), and Prosapia bicincta on Ilex cornuta and Ilex opaca. In three cases host use is limited to adults while in two cases use is limited to only adult males. In four cases adults feed on plants that differ taxonomically, structurally and ecologically from hosts normally utilized by all life stages. These associations are not considered aggregations per se because froghoppers are not clustered and because these associations cannot be explained by the usual criteria supporting the adaptiveness of insect aggregations. Indirect evidence suggests that these unusual hosts may be unavailable to nymphs and not sought by females. This passive lack of opportunity by nymphs and females, in tandem with an active selection/attraction by males, is a plausible explanation for the existence of male-specific hosts.     

Early carboniferous bryozoans from the Kodinka section, Middle Urals

An assemblage of Viséan-Serpukhovian bryozoans from the vicinity of the village of Kodinka (Sverdlovsk Region, Middle Urals) is studied for the first time. This assemblage is represented by two new species, Primorella kodinkensis sp. nov. and Heloclema uralicum sp. nov.; already known species Heloclema magnificum Gor. and Nikiforopora concentrica (Nikif.); and three morphotypes Fistulamina sp., Penniretepora sp., and Polyporella sp. Some specific features in the distribution of bryozoans of the Middle Urals at the beginning of the Carboniferous are discussed.     

Family coelotubuliporidae fam. nov. and morphological parallelisms in the evolution of bryozoans

Morphologically complex bryozoans from the Upper Devonian and Lower Carboniferous of Eurasia that some research workers include in the order Rhabdomesida and others place in the order Trepostomata are studied. The axial zooecium is demonstrated to be of zooidal nature. A new family, Coelotubuliporidae fam. nov., is established and its position in the order Trepostomata is substantiated. It includes two genera (one new), 11 species (1 new), and four subspecies. Diagnoses of a new family, Coelotubuliporidae, the genus Coelotubulipora Yang, Hu et Xia, 1988, and a new genus, Dunaevella Gorjunova gen. nov., are presented. The following species are described: Coelotubulipora irinae Gorjunova, sp. nov., Dunaevella shishovae (Schulga-Nesterenko, 1955), and D. peristomata (Schulga-Nesterenko, 1955). The emergence of axial zooecia in representatives of the order Trepostomata is considered as one of the evolutionary ways of bryozoans due to morphological parallelisms. It is shown that data presented on the stratigraphic range and paleogeography of the family Coelotubuliporidae are of great significance for solving the problems of biostratigraphy and correlation of deposits of the Upper Devonian and Lower Carboniferous of Eurasia.     

Upper Permian (Murghabian) rugose corals from Oman (Ba’id area,Saih Hatat): Community structure and contributions to reefbuilding processes

Summary Coral-dominated communties are rare in Upper Permian reefs. The study of Murghabian rugose and tabulate corals from allochtonous carbonates (‘Oman exotics’) of the Hawasina Complex and autochthonous carbonates of the Saih Hatat area/Arabian Platform (Oman) provides evidence for a significant contribution of rugose corals to the formation of Late Permian reefs. The corals are described with respect to taxonomy, microfacies and community structure. 8 genera and 7 species were recognized.Monothecalis minor n.sp.,Praewentzelella regulare n.sp. andWentzelella katoi magna n.ssp. are new. The corals represent three communities: (1)Praewentzelella community (Hawasina Complex), (2) cerioid coral community (Hawasina Complex), and (3)Waagenophyllum community (Hawasina Complex and Saih hatat). The corals from the Hawasina Complex and the Saih Hatat flourished in significantly different environments: Rugosa from the Hawasina Complex are representatives of reefs, whereas their counterparts from the Saih Hatat lived in level-bottom communities. Coral-bearing reefal boundstones are characterized by a diverse assemblage of sphinctozoans, inozoans, chaetetids, bryozoans, crinoids,Tubiphytes, Archaeolithoporella and algae. These communities produced bafflestones or framestones and were part of a sponge reef complex. The level-bottom community of the Saih Hatat is low-diverse only comprising rugose and tabulate corals. These of isolated colonies locally acted as bafflers.     

Cyrtocrinids (Cyrtocrinida,Crinoidea) and other associated crinoids from the Jurassic (Kimmeridgian–Tithonian)–Cretaceous (Berriasian–Barremian) of the Carpathian Foredeep basement (western Ukraine)

Upper Jurassic (Kimmeridgian–Tithonian) and Cretaceous (Berriasian–Barremian) strata of the Ukrainian part of the Carpathian Foredeep basement are rich, at least locally, in crinoid remains. Crinoids belonging to cyrtocrinids (Cyrtocrinida) are represented by whole cups, isolated remains of disarticulated cups, brachial plates and columnals. They are assigned to the following taxa: Cyrtocrinida indet., Eugeniacrinites cf. cariophilites (von Schlotheim), Lonchocrinus sp., Phyllocrinus stellaris Zaręczny, Ascidicrinus pentagonus (Jaekel), Gammarocrinites sp., Psalidocrinus armatus (Zittel), Psalidocrinus sp., and Hemibrachiocrinidae gen. indet. Cyrtocrinids are associated with other stalked (isocrinids, Isocrinida and millericrinids, Millericrinida) and stemless (saccocomids, Roveacrinida) crinoids. Columnals, pluricolumnals, brachial plates, and cirrals of isocrinids are assigned to Balanocrinus sp., Isocrinina fam. et subfam. indet., and columnals of millericrinids to Millericrinida indet. Free-living roveacrinids are assigned to Saccocoma sp. and Crassicoma sp. Knowledge on Jurassic and Cretaceous crinoids formerly described from Ukraine is discussed. Although majority of crinoids described herein seems to be allochthonous, autochthonous forms were also found with certainty in some intervals. These include some cyrtocrinids, which dominate in shallow-water environments of the Ukrainian Carpathian Foredeep basement. Isocrinids are also common in this shallow marine environment, whereas sessile saccocomids are assigned to low-energy, mud-supported bottom, open marine, outer-platform/upper slope, and relatively deep environments.     

Host-attacking behavior of a eulophid parasite,Kratochviliana sp., to the leaf mining host,Phytomyza ranunculi (Diptera: Agromyzidae) during its stay on the leaf

The present paper studies how the female parasite of Kratochviliana sp. visits and attacks its host larvae of Ranunculus leaf mining fly, P. ranunculi at a single leaf visit. The parasite visited its hosts at random on the leaf. The frequency of host visits was independent of the host density and the proportion of hosts survived from the parasite attack, in a leaf and its distribution was expressed as a single straight line. It almost always attacked living hosts at the first host visit after isolated from them for one day but with the rate of about 0.5 at the subsequent visits. In consequence, the relationships of the number of host attacks and killed hosts to the host density drew satulated curves in each. A model of host attack by this parasite at its single leaf visit was formulated by modifyingBakker et al.'s model (1972) basing upon these observations and the attack avoidance by the parasite to already attacked hosts previously reported.     

Soft-bottom assemblages of Paterson Inlet,Stewart Island

Abstract

The soft-bottom biota of subtidal areas of Paterson Inlet, Stewart Island, is described on the basis of data collected during cruises (1977 and 1979) in search of Braichiopoda. Two associations are recognised within the areas sampled, and their structure is compared with that of soft-bottom biotas elsewhere in New Zealand. The widespread occurrence of patch-forming bryozoans (especially Cinctipora elegans) on sandy gravels on the southern New Zealand shelf IS recognised.     

A TOURNAISIAN BRACHIOPOD FAUNA FROM SOUTH-EAST WALES

Abstract:  An exceptionally preserved fauna within dolomites of the Friars Point Limestone Formation includes the most diverse brachiopod assemblage yet described from the Tournaisian of the British Isles, and the first from Wales. Each of the 16 brachiopod genera includes a single species, of which four are new ( Schellwienella cheuma , Schuchertella subcrona , Composita ptygmation , Fusella extrata ). Associated fossils are corals (one species), bryozoans (two species) and crinoids (one species). Spiriferoideans and schizophorides are numerically dominant, indicative of level-bottom, inner mid-ramp biotopes. Biogeographical comparisons reflect the cosmopolitanism of early Carboniferous brachiopod generic assemblages. Taxonomic comparisons involve selection of lectotypes for Syringothyris exoleta North, 1920, Syringothyris cyrtorhyncha North, 1920, Tylothyris laminosa beta North, 1920, and Tylothyris laminosa gamma North, 1920.     

Lower Permian Bryozoa from Ellesmere Island (Canada)

A collection of fossils sampled during the 1898–1902 expedition of theFram to the Canadian Arctic Islands includes abundant bryozoans from the Lower Permian (Artinskian) Great Bear Cape Formation of Ellesmere Island. From this material a new genus with one new species —Nansenopora peculiaris n. gen., n. sp. — as well as three new species —Streblotrypella arctica n. sp.,Phragmophera patricki n. sp. andKallodictyon spinatum n. sp. — are described. Furthermore, the speciesUlrichotrypa ramulosa Bassler, 1929 is reported for the first time from the Lower Permian of the Arctic region.