Status of Divisions of the International Geologic Time Scale

Each chronostratigraphic unit of the International Geologic Time Scale will be defined at its base by a Global Stratotype Section and Point (GSSP) or Global Standard Stratigraphic Age (GSSA). Nearly 50 GSSPs and 10 GSSAs have now been ratified. Ideally, the GSSP coincides with events having a global correlation potential. The international stage divisions of some systems, such as the Jurassic or Neogene, are similar to traditional usage in European geology. However, in order to utilize global correlation horizons, the international stage divisions of other systems, such as the Ordovician or Permian, have required assembling new stage nomenclatures or hybrids of different regional stages. A reference table by the International Commission on Stratigraphy itemizes the current or potential GSSP and GSSA definitions of all international geologic time units.     

Fossil ovipositions of dragonflies: Review and interpretation

The paper deals with the stratigraphic range and morphology of fossil formations on plants interpreted as insect ovipositions. Our analysis of the insect fossil record has shown that the endophytic ovipositions probably belong to the Kennedyina and Triadophlebiina (in the Paleozoic and Lower Mesozoic) and to the Calopterygina (in the Upper Mesozoic and Cenozoic).     

The post‐Palaeozoic fossil record of drilling predation on lingulide brachiopods

Research on drilling predation, one of the most studied biological interactions in the fossil record, has been concentrated on prey with calcareous skeletons (e.g. molluscs, echinoids, rhynchonelliform brachiopods). Based on a compilation of literature sources and surveys of paleontological collections of the Florida Museum of Natural History and the National Museum of Natural History, we provide a tentative evaluation of the post‐Palaeozoic history of drilling predation on the organophosphatic brachiopods of the family Lingulidae. Despite temporal, geographical and methodological limitations of the data assembled here, the results indicate that lingulide brachiopods have been subject to drilling predation since at least the Eocene. Variation in drilling frequencies at the locality level suggests that lingulides may occasionally experience somewhat elevated predation pressures from drilling organisms. Overall, however, drilling predation on lingulide brachiopods has been infrequent in the Cenozoic and may have been absent in the Mesozoic. The Mesozoic‐to‐Cenozoic increase in drilling frequencies on lingulides is similar to the trends observed in other marine benthic invertebrates and consistent with the hypothesis that predation pressures increased through time in marine ecosystems.     

Norian-Rhaetian reefs in Argolis Peninsula,Greece

Summary Upper Triassic to Lower Jurassic shallow-water carbonate sequences of the ‘Pantokrator limestones’ are widely distributed in the Argolis Peninsula, southern Greece. Within this sequence are some reef or reefal structures. In the Mavrovouni Mountains, near Sarmeika, 6 km SE of the ancient theatre of Epidavros (Argolis Peninsula), a Norian-Rhaetian reef complex has been identified. This is the first well-documented Norian-Rhaetian reef in Greece. The main reef builders are coralline sponges (‘sphinctozoans,’ ‘inozoans’, and sclerosponges), followed by dendroid, cerioid, and solitary corals, and algae. The reef type corresponds to a ‘sponge-coral reef’.     

Parrotfish bioerosion on Egyptian Red Sea reefs

Parrotfishes (family Scaridae) are important agents in marine bioerosion. Here, the feeding ecology of seven species of parrotfishes was studied on Egyptian Red Sea reefs. The most abundant species on both the reef flat and slope was Chlorurus sordidus. In contrast, C. gibbus had the lowest abundance on the reef flat, and Cetoscarus bicolor was the least abundant species on the reef slope. Scarus niger exhibited the highest feeding rate (98.9 bites 5 min^− 1), followed by C. sordidus (76.5 bites 5 min^− 1), whereas the rates for C. bicolor and C. gibbus were low (29.4 and 31.9 bites 5 min^− 1, respectively). The daily feeding patterns of all seven species showed agreement in that activity was relatively constant over the day, with highest values in the early afternoon (1400 h) and a steady decrease until 1800 h. C. sordidus was more similar to S. niger and S. ghobban in showing somewhat higher activities in the morning (0800 h) followed by a slight decrease until noon. The average bite volumes of C. gibbus and C. bicolor were high (0.114 and 0.110 cm³, respectively), whereas S. niger had the lowest average value (0.002 cm³). Based on their feeding intensity, C. gibbus, S. ghobban and C. bicolor have high bioerosion rates on the Egyptian Red Sea reefs. Overall, S. ghobban is the most important bioeroder because it is more abundant than the other two species. All parrotfish species fed on dead coral and hard substrates which are rich in algae, but C. gibbus, C. bicolor and S. ghobban also fed on live coral on both reef zones; C. sordidus avoided live coral. The fresh scars on live coral were bigger than on dead coral because the three large parrotfish (C. gibbus, C. bicolor and S. ghobban) fed mainly on live corals.     

Modern and fossil traces in terrestrial lithic substrates

A number of biogenic processes leads to the formation of distinctive traces in terrestrial lithic substrates. These include: burrowing by vertebrates in moderately lithified rocks; scraping by mammals; smoothing and polishing of limestone surfaces by the locomotion of mammals; excavation by bees, wasps, and ants producing nesting and dwelling tunnels; dissolution of limestone surfaces by terrestrial snails; endolithic activity of fungi, algae, and lichens on subaerial rock surfaces; root corrosion; etc. Processes of biochemical weathering, biophysical erosion, and enlargement of cracks and fissures by the pressure of plant roots do not leave distinctive traces and therefore lie outside the ichnological realm. The fossil preservation of terrestrial bioerosional traces is expected to be uncommon. Nevertheless, various possible means of preservation must be considered, such as by rapid burial by volcanic material, by fluvial sediments, by travertine or tufa, by loess, “conservation”; in caves, case hardening of surfaces of porous rocks, and preservation of subsoil traces below fossil soils.     

Bioerosion in Late Permian Rugosa from reefal blocks (Hawasina Complex,Oman Mountains): implications for reef degradation

Summary Rugose corals are known from allochthonous Late Permian reefal blocks of the A1 Jil and Ba’id Formation (Hawasina Complex), Oman Mountains. In contrast to many Late Permian Rugosa found elsewhere in the Tethys, they occurred in sponge reefs and contributed to reef construction. The waagenophyllid warm water coral fauna is moderately diverse comprising cerioid, thamnasterioid, and fasciculate taxa. In contrast to sponges, chaetetids, and low-growing reefbuilders, the corals secreted diagenetically stable, most probably Mg-calcitic skeletons. Borings in coral skeletons are consequently well preserved providing important data for the interpretation of reef destructive processes. Thin-section analysis revealed three taxa of infaunal borers includingEntobia Bronn 1837, uncertain thallophyte borings, and borings of unknown bioeroders. Macroborers were more important than microborers, because of the dominance of clionid sponges. Good evidence exists also for the occurrence of two types of undetermined grazers which destroyed the coral surfaces. The amount and distribution of bioerosions is variable among different coral taxa. The fasciculate coralPraewentzelella regulare Flügel 1995 was the favorate substrate. Up to 33% of the calices were bored. Dendroid and compound corals were bored subordinately. Bioerosion of these colonies does not exceed 2%. There is good evidence for substrate preference amongst the borers. Major controlling factors affecting borer distribution are believed to be variations of skeletal density and gross morphology. The borer assemblage could not limit reef accretion significantly. Factors controlling boring activity might have been quality of substrate, sedimentation rate, rapid incrustation of substrates, and competition for food with reef constructors including sponges, chaetetids, and rugose corals.     

Ordovician reefs of the Ural Mountains,Russia: A review

Summary The Upper Ordovician reefs of the Urals were formed at a subsiding shelf-margin during an early Late Ashgillian (Sur’ya time interval) regressive phase. Reefs of this age were studied in detail from the western slope of the Northern, the Subpolar and the Polar Urals with respect to lithofacies, biotic composition and paleogeographical patterns. The thickness of the reefs varies between 100 and 500 m. The backreef areas are characterized by lagoons with increased salinity and sabkha development. Microbial associations and a diverse algal flora (Cyanophyta, green and red algae and alga incertae sedis) are the main constituents of reefal boundstones. Tabulate and rugose corals, heliolitids, calcareous sponge-like fossils, bryozoans and problematic hydroids were also part of the reef communities. Each reef exhibits a characteristic framework-building association. Reef development was terminated by a rapid and abrupt sea-level rise at the end of the middle Upper Ashgillian connected with the global Late Ordovician glaciation.     

Early Devonian reefs at Reefton, New Zealand: guilds, origin and paleogeographic significance

Study of 35 systematically collected and 10 semi‐random samples (about 100 cm 2 each) from one outcrop of the Lankey Limestone (Emsian) near Reefton, New Zealand indicates that it is a reef framestone built by tabulate (4? spp.) and clonal rugose (one sp.) corals, two species of stromatoporoids, a few bryozoans (2? spp.) and crinoids. The guild structure of the reef community and data on skeleton orientation and growth direction further indicate that there was relatively minor biostratonomic alteration of the original community. Skeletons are either in growth position or tipped, toppled or even overturned, but they have remained in situ. Other clone‐rich Lankey Limestone outcrops in the Reefton area support the notion of an upper shelf reef system and stratigraphic and sedimentologic data suggest that it was located near the Gondwana margin at a paleolatitude of about 35°S.     

The origin of High-Mg-Calcite microbialites in cryptic habitats of Caribbean coral reefs—Their dependence on light and turbulence

Summary In coral reefs of St. Croix, U.S.V.I., microbialites are found in weakly illuminated cryptic habitats. They form crusts of several centimeter thickness on coralgal framework. A high energy environment causes a thrombolitic texture whereas under low energy conditions stromatolitic crusts are developed. High-Mg-Calcite crystallizes in an organic matrix and silt-sized particles are trapped on an organic mucus; both processes contribute to the accretion of microbialites. A sessile benthic community dominated by foraminifera uses microbialites as a hard substrate to grow.     

Paleocene reefs on the Maiella Platform Margin, Italy: An example of the effects of the cretaceous/tertiary boundary events on reefs and carbonate platforms

Summary Reef facies, reef types and their biotic associations in the Maiella platform margin (central Italy) provide qualitative evidence for a significant reef decline across the Cretaceous/Tertiary (K/T) boundary, and indicate two phases of reef recovery during the Paleocene. Rudists dominated the reef community until the latest Cretaceous. A significant sea-level fall around the time of the K/T boundary is documented by a truncation surface associated with emersion. During sea-level highstands in the Danian to Early Thanetian and, more extensively, during the Late Thanetian, coral-algal patch-reefs grew along the platform margin and top. Already in the Danian to Early Thanetian, the reef communities were more diverse and the constructional types more evolved than previously known from this time. Differences between the Danian to Early Thanetian coral association, the Late Thanetian association, and Late Cretaceous coral faunas may have ecological or evolutionary causes. Repeated emergence produced a complex diagenetic history in the Danian to Lower Thanetian limestones. All Paleocene reefs were displaced by gravitative redeposition. Coral-algal reefs are less important in the Early to mid Eocene, when alveolinid foraminifera dominated on the Maiella shelf. Reefs on the Maiella platform diversified and attained large sizes in the Late Eocene to Early Oligocene, as known from other Mediterranean platforms. The external controls on the Late Cretaceous to Oligocene evolution and demise of reef communities that are most easily demonstrated with our data are sealevel fluctuations and climate change. We propose that the change in reef biota and reef types across the K/T boundary and during the Early Tertiary were important causes of the parallel changes in platform growth style.     

Diversity and extinction in the Cenozoic history of Caribbean reefs

Occurrences of reef corals are examined at Caribbean fossil localities to determine how biodiversity has changed within the region over the past 50 million years. Analyses of 294 species (66 genera) at 58 fossil localities show that Caribbean generic diversity rose to 44 between 50–22 Ma, ranged from 32–39 between 22–2 Ma, and dropped to 25 afterwards. Regional species diversity was high at 40–36 Ma, 28–22 Ma, and 5–2 Ma. Origination rates were elevated throughout each high diversity interval, but extinction was concentrated near the end of each interval. Regional highs of origination and extinction, therefore, differed in timing and duration, causing the observed regional diversity increases during the three remarkably long intervals of turnover. Highs of generic origination decreased in magnitude as immigration from the Mediterranean ceased, but speciation highs increased in association with emergence of the Central American isthmus. Peaks of extinction coincided with regional changes in climate and oceanic circulation. Maximum species diversities within assemblages increased to 40–60 between 50–36 Ma, and have remained relatively constant ever since. Assemblage compositions differed among localities having similar ages and environments, suggesting that the timing and pattern of turnover varied across the region. Stable diversities but variable compositions within assemblages suggest that dispersal and recruitment influenced the pattern of faunal change during turnover. Accepted: 22 August 1999     

The short-term influence of herbivory near patch reefs varies between seagrass species

The coexistence of multiple species within a trophic level can be regulated by consumer preferences and nutrient supply, but the influence of these factors on the co-occurrence of seagrass species is not well understood. We examined the biomass and density responses of two seagrass species in the Florida Keys Reef Tract to grazing pressure near patch reefs, and evaluated how nutrient enrichment impacted herbivory dynamics. We transplanted Halodule wrightii (shoalgrass) sprigs into caged and uncaged plots in a Thalassia testudinum (turtlegrass) bed near a patch reef. Nutrients (N and P) were added to half of the experimental plots. We recorded changes in seagrass shoot density, and after three months, we measured above- and belowground biomass and tissue nutrient content of both species. Herbivory immediately and strongly impacted H. wrightii. Within six days of transplantation, herbivory reduced the density of uncaged H. wrightii by over 80%, resulting in a decrease in above- and belowground biomass of nearly an order of magnitude. T. testudinum shoot density and belowground biomass were not affected by herbivory, but aboveground biomass and leaf surface area were higher within cages, suggesting that although herbivory influenced both seagrass species, T. testudinum was more resistant to herbivory pressure than H. wrightii. Nutrient addition did not alter herbivory rates or the biomass of either species over the short-term duration of this study. In both species, nutrient addition had little effect on the tissue nutrient content of seagrass leaves, and N:P was near the 30:1 threshold that suggested a balance between N and P. The different impacts of grazing on these two seagrass species suggest that herbivory may be an important regulator of the distribution of multiple seagrass species near herbivore refuges like patch reefs in the Caribbean.     

中国中生代昆虫化石研究新进展

回顾了1990年以来中国中生代昆虫化石研究的新进展,简述了我国学者在古昆虫分类学、昆虫内部形态学、昆虫系统学、古生态学、昆虫生物地层学、昆虫区系的历史演变、生物古地理学、昆虫与植物的关系等8个方面取得的突出贡献,并给出了我国发现的世界最低层位昆虫科级类群表。     

Spatial variation in sea urchins, fish predators, and bioerosion rates on coral reefs of Belize

Although sea urchins are critical for controlling macroalgae on heavily fished coral reefs, high densities threaten reefs, as urchins are also prodigous bioeroders. This study examined urchin population characteristics, bioerosion rates, their fish predators (Labridae), and potential competitors (Scaridae) on unprotected reefs and a reef within a marine protected area (MPA) in the lagoonal regions off Belize. Urchin density (<1 m⁻²) and bioerosion rates (∼0.2 kg CaCO₃ m⁻² year⁻¹) were lowest and members of the Labridae were the highest (∼20 fish 200 m⁻³) within the MPA, while several unprotected reefs had higher (∼18–40 m⁻²) urchin densities, lower Labridae abundances (1–3 fish 200 m⁻³), and bioerosion rates ranging from ∼0.3–2.6 kg CaCO₃ m⁻² year⁻¹. Urchin abundances were inversely related to Labridae (wrasses and hogfish) densities; however, on reef ridges, low algal cover (∼15%), small urchin size (∼14 mm), and low proportion of organic material in urchin guts suggested food limitation. Both top–down (predation) and bottom–up factors (food limitation) likely contribute to the control of urchins, predominantly Echinometra viridis, off Belize, thereby potentially diminishing the negative impacts of bioerosion activities by urchins.     

Boring sponges,an increasing threat for coral reefs affected by bleaching events

Coral bleaching is a stress response of corals induced by a variety of factors, but these events have become more frequent and intense in response to recent climate‐change‐related temperature anomalies. We tested the hypothesis that coral reefs affected by bleaching events are currently heavily infested by boring sponges, which are playing a significant role in the destruction of their physical structure. Seventeen reefs that cover the entire distributional range of corals along the Mexican Pacific coast were studied between 2005/2006, and later between 2009/2010. Most of these coral reefs were previously impacted by bleaching events, which resulted in coral mortalities. Sponge abundance and species richness was used as an indicator of bioerosion, and coral cover was used to describe the present condition of coral reefs. Coral reefs are currently highly invaded (46% of the samples examined) by a very high diversity of boring sponges (20 species); being the coral reef framework the substrate most invaded (56%) followed by the rubbles (45%), and the living colonies (36%). The results also indicated that boring sponges are promoting the dislodgment of live colonies and large fragments from the framework. In summary, the eastern coral reefs affected by bleaching phenomena, mainly provoked by El Niño, present a high diversity and abundance of boring sponges, which are weakening the union of the colony with the reef framework and promoting their dislodgment. These phenomena will probably become even more intense and severe, as temperatures are projected to continue to rise under the scenarios for future climate change, which could place many eastern coral reefs beyond their survival threshold.     

The origin of Jurassic reefs: Current research developments and results

Summary In order to elucidate the control of local, regional and global factors on occurrence, distribution and character of Jurassic reefs, reefal settings of Mid and Late Jurassic age from southwestern Germany, Iberia and Romania were compared in terms of their sedimentological (including diagenetic), palaeoecological, architectural, stratigraphic and sequential aspects. Upper Jurassic reefs of southern Germany are dominated by siliceous sponge—microbial crust automicritic to allomicritic mounds. During the Oxfordian these form small to large buildups, whereas during the Kimmeridgian they more frequently are but marginal parts of large grain-dominated massive buildups. Diagenesis of sponge facies is largely governed by the original composition and fabric of sediments. The latest Kimmeridgian and Tithonian spongiolite development is locally accompanied by coral facies, forming large reefs on spongiolitic topographic elevations or, more frequently, small meadows and patch reefs within bioclastic to oolitic shoal and apron sediments. New biostratigraphic results indicate a narrower time gap between Swabian and Franconian coral development than previously thought. Palynostratigraphy and mineralostratigraphy partly allow good stratigraphic resolution also in spongiolitic buildups, and even in dolomitised massive limestones. Spongiolite development of the Bajocian and Oxfordian of eastern Spain shares many similarities. They are both dominated by extensive biostromal development which is related to hardground formation during flooding events. The Upper Jurassic siliceous sponge facies from Portugal is more localised, though more differentiated, comprising biostromal, mudmound and sponge-thrombolite as well as frequent mixed coral-sponge facies. The Iberian Upper Jurassic coral facies includes a great variety of coral reef and platform types, a pattern which together with the analysis of coral associations reflects the great variability of reefal environments. Microbial reefs ranging from coralrich to siliceous sponge-bearing to pure thrombolites frequently developed at different water depths. Reef corals even thrived within terrigeneous settings. In eastern Romania, small coral reefs of various types as well as larger siliceous sponge-microbial crust mounds grew contemporaneously during the Oxfordian, occupying different bathymetric positions on a homoclinal ramp. Application of sequence stratigraphic concepts demonstrates that onset or, in other cases, maximum development of reef growth is related to sea level rise (transgressions and early highstand) which caused a reduction in allochthonous sedimentation. The connection of reef development with low background sedimentation is corroborated by the richness of reefs in encrusting organisms, borers and microbial crusts. Microbial crusts and other automicrites can largely contribute to the formation of reef rock during allosedimentary hiatuses. However, many reefs could cope with variable, though reduced, rates of background sedimentation. This is reflected by differences in faunal diversities and the partial dominance of morphologically adapted forms. Besides corals, some sponges and associated brachiopods show distinct morphologies reflecting sedimentation rate and substrate consistency. Bathymetry is another important factor in the determination of reefal composition. Not only a generally deeper position of siliceous sponge facies relative to coral facies, but also further bathymetric differentiation within both facies groups is reflected by changes in the composition, diversity and, partly, morphology of sponges, corals, cementing bivalves and microencrusters. Criteria such as authigenic glauconite, dysaerobic epibentic bivalves,Chondrites burrows or framboidal pyrite in the surrounding sediments of many Upper Jurassic thrombolitic buildups suggest that oxygen depletion excluded higher reefal metazoans in many of these reefs. Their position within shallowing-upwards successions and associated fauna from aerated settings show that thrombolitic reefs occurred over a broad bathymetric area, from moderately shallow to deep water. Increases in the alkalinity of sea water possibly enhanced calcification. Reefs were much more common during the Late Jurassic than during the older parts of this period. Particularly the differences between the Mid and Late Jurassic frequencies of reefs can be largely explained by a wider availability of suitable reef habitats provided by the general sea level rise, rather than by an evolutionary radiation of reef biota. The scarcity of siliceous sponge reefs on the tectonically more active southern Tethyan margin as well as in the Lusitanian Basin of west-central Portugal reflects the scarcity of suitable mid to outer ramp niches. Coral reefs occurred in a larger variety of structural settings. Upper Jurassic coral reefs partly grew in high latitudinal areas suggesting an equilibrated climate. This appears to be an effect of the buffering capacity of high sea level. These feedback effects of high sea level also may have reduced oceanic circulation particularly during flooding events of third and higher order, which gave rise to the development of black shales and dysaerobic thrombolite reefs. Hence, the interplay of local, regional and global factors caused Jurassic reefs to be more differentiated than modern ones, including near-actualistic coral reefs as well as non-actualistic sponge and microbial reefs.     

Significance of stromatoporoids in Jurassic reefs and carbonate platforms—concepts and implications

Although many case studies describe stromatoporoid-rich Jurassic reefs, there are only few reliable data as to their distribution pattern. This is in part due to a largely taxonomic and systematic focus on the enigmatic stromatoporoids which now are interpreted as a polyphyletic informal group of demosponges by most specialists. The common co-occurrence of Jurassic scleractinian corals and stromatoporoids might, at first hand, point to very similar environmental demands of both organismic groups, but autecological considerations as well as evaluation of stromatoporoid distribution patterns should allow for a much more refined interpretation. This study concludes that Jurassic corals and stromatoporoids show a relatively broad overlap of environmental demands but their maximum ecological tolerances appear to differ considerably. Jurassic corals were dominating in mesotrophic to mildly oligotrophic, slightly deeper settings, where they largely outcompeted stromatoporoids. On the other hand, stromatoporoid growth was particularly favoured in very shallow water, strongly abrasive, high-energy settings as well as in possibly overheated waters. Many taxa and growth forms were very tolerant towards frequent reworking and redistribution, a feature which is compatible with the sponge nature of the stromatoporoids. As such, stromatoporoid facies may be common in low-accommodation regimes, giving rise to frequent “shelf shaving” and redistribution across wide shelf areas. The mixed coral-stromatoporoid reefs from the margins of isolated Intra-Tethys platforms are interpreted to be indicative of oligotrophic normal marine waters. This is corroborated by statistical cluster analysis of stromatoporoid taxa from representative areas. In addition, Arabian stromatoporoid occurrences might have been adapted to overheated and slightly hypersaline waters. There also are a few exceptional stromatoporoid taxa which might have had environmental tolerances different from the bulk tolerances of other Jurassic stromatoporoids. Part of our interpretations are preliminary and should stimulate further research. However, the present results already help explain the observed compositional differences between Jurassic North Tethys/North Atlantic, Intra-Tethys, and South Tethys shallow-water reefs and platforms.