Phenotypic evolution in a fossil gastropod species lineage: evidence for adaptive radiation?

Detecting speciation in the fossil record is a particularly challenging matter. Palaeontologists are usually confronted with poor preservation and limited knowledge on the palaeoenvironment. Even in the contrary case of adequate preservation and information, the linkage of pattern to process is often obscured by insufficient temporal resolution. Consequently, reliable documentations of speciation in fossils with discussions on underlying mechanisms are rare. Here we present a well-resolved pattern of morphological evolution in a fossil species lineage of the gastropod Melanopsis in the long-lived Lake Pannon. These developments are related to environmental changes, documented by isotope and stratigraphical data. After a long period of stasis, the ancestral species experiences a phenotypic change expressed as shift and expansion of the morphospace. The appearance of several new phenotypes along with changes in the environment is interpreted as adaptive radiation. Lake-level high stands affect distribution and availability of habitats and, as a result of varied functional demands on shell geometry, the distribution of phenotypes. The on-going divergence of the morphospace into two branches argues for increasing reproductive isolation, consistent with the model of ecological speciation. In the latest phase, however, progressively unstable conditions restrict availability of niches, allowing survival of one branch only.

Volcaniclastic sediments of ODP Leg 136 sites

Sediment cores recovered from three holes drilled during Ocean Drilling Program Leg 136 include volcaniclastics probably derived from the Hawaiian islands. The volcaniclastics shallower than 10 meters below seafloor are fresh and are composed of basaltic glass (sideromelane), basaltic fragments (mainly tachylite), plagioclase, olivine, pyroxene, and opaque minerals. Most of these glasses are probably products of hydrovolcanism. Visibly, some of these volcaniclastics are recognized as bedded ash layers having thicknesses that range from 5 to 10 cm. However, many volcaniclastics are disrupted by bioturbation to some degree, and are sometimes totally mixed with ambient brown clays. No visible correlative ash layer among these holes was found. It seems that many ash layers thinner than the bedded layers were disrupted by bioturbation because of the low sedimentation rate of volcaniclastics. The volcaniclastics were probably transported one of two ways: through air fall and pelagic settling or through turbidity-current transport. Other archipelagic apron volcaniclastic sediments of volcanic seamounts suggest that turbidite transport is the favored explanation of origin.

Paleocene radiolarian distribution and biostratigraphy from offshore eastern New Zealand

A 100-m-thick Paleocene sequence of mainly pelagic sediments at ODP Site 1121, on the eastern flanks of the Campbell Plateau, contains few to common radiolarians of relatively low diversity in the lower 40 m (Early to early Late Paleocene) and abundant, diverse radiolarian assemblages in the upper 60 m (mid-Late Paleocene). The 150 taxa recorded from the entire Paleocene interval are thought to under-represent the actual species diversity by at least one half as many morphotypes have not been differentiated below the level of genus. Assemblages in the lower 40 m are similar to those described from onland New Zealand and DSDP Site 208 (northern Lord Howe Rise); they are correlated with South Pacific radiolarian zones RP4 and RP5. Assemblages in the upper 60 m differ from other known Late Paleocene assemblages in the great abundance of plagiacanthids and cycladophorids. Similarities are noted with later Cenozoic cool-water assemblages. This upper interval is correlated with South Pacific zone RP6, as revised herein, based on comparison with faunas from Site 208 and Marlborough, New Zealand. The interval is also correlated with the upper part of North Atlantic zone RP6 (RP6b-c) based on the presence of Aspis velutochlamydosaurus, Plectodiscus circularis and Pterocodon poculum. Other species, such as Buryella tetradica and Buryella pentadica, are valuable for local correlation but exhibit considerable diachroneity between the Pacific, Indian and Atlantic Oceans. An age model for the Paleocene interval at Site 1121, based on well-constrained nannofossil and radiolarian datums, indicates that the rate of compacted sediment accumulation doubles from 15 to 30 mm/ka at the RP5/RP6 zonal boundary. In large part this is due to a sudden and pronounced increase in accumulation rates for all siliceous fossils; radiolarians and larger diatoms increase from <100 to >10 000 specimens/cm2/ka. This apparent increase in biosiliceous productivity is age-equivalent to a mid-Paleocene cooling event (57-59 Ma) identified from global stable isotope records that is associated with the heaviest delta13C values for the entire Cenozoic.

Nannofossil and stable isotope record across the Paleocene/Eocene Thermal Maximum in ODP Hole 183-1135A

The Paleocene/Eocene Thermal Maximum (PETM, ca. 55 Ma) is an abrupt, profound perturbation of climate and the carbon cycle associated with a massive injection of isotopically light carbon into the ocean-atmosphere system. As such, it provides an analogue for understanding the interplay between phytoplankton and climate under modern anthropogenic global-warming conditions. However, the accompanying enhanced dissolution poses uncertainty on the reconstruction of the affected ecology and productivity. We present a high-resolution record of bulk isotopes and nannofossil absolute abundance from Ocean Drilling Program (ODP) Site 1135 on the Kerguelen Plateau, Southern Indian Ocean to quantitatively constrain for the first time the influence of dissolution on paleoecological reconstruction. Our bulk-carbonate isotope record closely resembles that of the classic PETM site at ODP Site 690 on the opposite side of the Antarctic continent, and its correlation with those from ODP Sites 690, 1262 and 1263 records allows recognition of 14 precessional cycles upsection from the onset of the carbon isotopic excursion (CIE). This, together with a full range of common Discoasteraraneus and an abundance crossover between Fasciculithus and Zygrhablithusbijugatus, indicates the presence of the PETM at Site 1135, a poorly known record with calcareous fossils throughout the interval. The strong correlation between the absolute abundances of Chiasmolithus and coccolith assemblages reveals a dominant paleoecological signal in the poorly preserved fossil assemblages, while the influence of dissolution is only strong during the CIE. This suggests that r-selected taxa can preserve faithful ecological information even in the severely-altered assemblages studied here, and therefore provide a strong case for the application of nannofossils to paleoecological studies in better-preserved PETM sections. The inferred nannoplankton productivity drops abruptly at the CIE onset, but rapidly increases after the CIE peak, both of which may be driven by nutrient availability related to ocean stratification and vertical mixing due to changed sea-surface temperatures.

Initial biomass and biomass change of tundra and forest plants in three herbivore treatments in Norway and Sweden

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana, has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (> 0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (< 0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.

Stable organic carbon isotope record and total organic carbon content from the Cenomanian-Turonian boundary

Ocean Drilling Program (ODP) Leg 207 recovered expanded sections of organic-carbon-rich laminated shales on Demerara Rise (western tropical Atlantic). High-resolution organic carbon isotope and total organic carbon (TOC) records are presented, which span the Cenomanian-Turonian boundary interval (CTBI), including the Oceanic Anoxic Event (OAE) 2, from four sites oriented along a NW striking depth transect. These records represent the first high-resolution carbon isotope records across OAE 2 from the South American margin of the tropical Atlantic. Due to the scarcity of age significant fossils, the main purpose of this study was to develop a detailed carbon isotope stratigraphy in order to correlate the CTBI across the depth transect and to tie this to biostratigraphically well-defined sections in the Western Interior Basin (Pueblo, USA), boreal shelf seas (Eastbourne, England), and western Tethys (Oued Mellegue, Tunisia). All four sections studied document a 6 per mil increase of d13Corg values at the base of the CTBI, which is followed by an interval of elevated d13Corg values and a subsequent decrease. Our results supply an important stratigraphic base for subsequent paleoceanographic studies on Late Cenomanian to Early Turonian sediments from Demerara Rise and elsewhere.

Presence-absence species/sample matrix for the measured sections at the area of Maghara, North Sinai, Egypt

As age-diagnostic fossils are rare in the Middle to Upper Jurassic sedimentary succession of Gebel Maghara, North Sinai, Egypt, and in order to ensure maximal stratigraphic resolution, chronostratigraphic boundaries were determined based on quantitative biostratigraphy. A data matrix comprising 231 macrofaunal taxa in 93 samples from four sections has been processed with the Unitary Association (UA) Method. This led to construction of a sequence of 29 UAs (maximal sets of actually or virtually coexisting taxa), which have been grouped into 14 laterally reproducible association zones. The UA method allowed an in-depth analysis of the stratigraphically conflicting taxa, enabled the biostratigraphic subdivision of the studied interval, and also provided stratigraphic correlation among the measured sections and with the Tethyan ammonite zones.

Tab. 1: Meerfelder Maar Drilling B4, stratigraphy and sequence of the lake sediments (pdf 215 kB)

A core from Meerfelder Maar, with a basal age of 29,000 years, provides a continuous sedimentary sequence from Late-Glacial times to the present. It includes the stratigraphical marker of the Laach Pumice Tuff. Sedimentological, geochemical, palynological, palaeobiological, palaeomagnetic and palaeontological analyses permit reconstructions of the history of the lake and its catchment area, and hence of the climate of the region, to be made. The discovery of Middle Oligocene marine, detrital fossils in the maar sediments provides insights into the palaeogeography of the Eifel region during Tertiary times.

Documentation of sediment cores from the Great Meteor Seamount, North Atlantic

Sedimentological and biostratigraphic investigations of 15 cores (total length: 88 m) from the vicinity of Great Meteor seamount (about 30° N, 28° W) showed that the calcareous ooze are asymmetrically distributed around the seamount and vertically differentiated into two intervals. East and west of the seampunt, the upper "A"-interval is characterized by yellowish-brown sediment colors and bioturbation; ash layers and diatoms are restricted to the eastern cores. On both seamount flanks, the sediment of the lower "B"-interval are white and very rich in CaCO3 with a major fine silt (2-16 µ) mode (mainly coccoliths). Lamination, manganese micronodules, Tertiary foraminifera and discoasters, and small limestone and basalt fragments are typical of the "B"-interval of the eastern cores only. The sediments contain abundant displaced material which was reworked from the upper parts of the seamount. The sedimentation around the seamount is strongly influenced by the kind of displaced material and the intensity of its differentiated dispersal: the sedimentation rates are generally higher on the east than on the west flank /e.g. in "B": 0.9 cm/1000 y in the W; 3.1 cm/1000 y in the E), and lower for the "A" than for the "B"-interval. The lamination is explained by the combination of increased sedimentation rates with a strong input of material poor in organic carbon producing a hostile environment for benthic life. The CaCO3 content of the core is highly influenced by the proportion of displaced bigenous carbonate material (mainly coccoliths). The genuine in-situ conditions of the dissolution facies are only reflected by the minimum CaCO3 values of the cores (CCD = about 5,500 m; first bend in dissolution curve = 4,000 m; ACD = about 3,400 m). The preservation of the total foraminiferal association depends on the proportions of in-situ versus displaced specimens. In greater water depths (stronger dissolution), for example, the preservation can be improved by the admixture of relatively well preserved displaced foraminifera. Carbonate cementation and the formation of manganese micronodules are restricted to microenvironments with locally increased organic carbon contents (e.g. pellets; foraminifera). The ash layers consist of redeposited, silicic volcanic glass of trachytic composition and Mio-Pliocene age; possibly, they can be derived from the upper part of the seamount. Siliceous organisms, especially diatoms, are frequent close to the ash layers and probably also redeposited. Their preservation was favoured by the increase of the SiO2 content in the pore water caused by the silicic volcanic glass. The cores were biostraftsraphically subdivided with the aid of planktonic foraminifera and partly alsococcoliths. In most cases, the biostratigraphically determined cold- and warm sections could be correlated from core to core. Almost all cores do not penetrate the Late Pleistocene. All Tertiary fossils are reworked. In general, the warm/cold boundary W2/C2 corresponds with the lithostratigraphic A/B boundray. Benthonic foraminifera indicate the original site deposition of the displaced material (summit plateau or flanks of the seamount). The asymmetric distribution of the sediments around the seamount east and west of the NE-directed antarctic bottom current (AABW) is explained by the distortion of the streamlines by the Coriolis force; by this process the current velocity is increased west of the seamount and decreased east of it. The different proportion of displaced material within the "A" and "B" interval is explained by changes of the intensity of the oceanic circulation. At the time of "B" the flow of the AABW around the seamount was stronger than during "A"; this can be inferred from the presence of characteristic benthonic foraminifera. The increased oceanic circulation implies an enhanced differentiation of the current velocities, and by that, also of the sedimentation rates, and intensifies the winnowed sediment material was transported downslope by turbid layers into the deep-sea, incorporated into the current system of the AABW, and asymmetrically deposited around the seamount.

Miocene dissolution facies and microfacies at Northwest Atlantic Deep Sea Drilling Project Site 93-603

Depth fluctuations of the lysocline and calcite compensation depth (CCD) through time were investigated at Deep Sea Drilling Project Site 603, Leg 93. The CCD fell during the middle Miocene at the onset of the Western Boundary Undercurrent, correlated with seismic Horizon X. Subsequently deposited units show fluctuations of the dissolution curve. Major changes in dissolution facies correspond with lithologic boundaries.

Composition of bioclastic sands, carbonates and pyroclastic rocks of the Great Meteor and Josephine Seamounts, eastern North Atlantic

1. Great Meteor Seamount (GMS) is a very large (24,000 km**3) guyot with a flat summit plateau at 330-275 m; it has a volcanic core, capped by 150-600 m of post-Middle-Miocene carbonate and pyroclastic rocks, and is covered by bioclastic sands. The much smaller Josephine Seamount (JS, summit 170- 500 m w. d.) consists mainly of basalt which is only locally covered by limestones and bioclastic sands. 2. The bioclastic sands are almost free of terrigenous components, and are well sorted, unimodal medium sands. (1) "Recent pelagic sands" are typical of water depths > 600 m (JS) or > 1000 m (GMS). (2) "Sands of mixed relict-recent origin" (10-40% relict) and (3) "relict sands" (> 40% relict) are highly reworked, coarse lag deposits from the upper flanks and summit tops in which recent constituents are mixed with Pleistocene or older relict material. 3. From the carbonate rocks of both seamounts, 12 "microfacies" (MF-)types were distinguished. The 4 major types are: (1) Bio(pel)sparites (MF 1) occur on the summit plateaus and consist of magnesian calcite cementing small pellets and either redeposited planktonic bioclasts or mixed benthonic-planktonic skeletal debris ; (2) Porous biomicrites (MF 2) are typical of the marginal parts of the summit plateaus and contain mostly planktonic foraminifera (and pteropods), sometimes with redeposited bioclasts and/or coated grains; (3) Dense, ferruginous coralline-algal biomicrudites with Amphistegina sp. (MF 3.1), or with tuffaceous components (MF 3.2); (4) Dense, pelagic foraminiferal nannomicrite (MF 4) with scattered siderite rhombs. Corresponding to the proportion and mineralogical composition of the bioclasts and of the (Mgcalcitic) peloids, micrite, and cement, magnesian calcite (13-17 mol-% MgCO3) is much more abundant than low-Mg calcite and aragonite in rock types (1) and (2). Type (3) contains an "intermediate" Mg-calcite (7-9 mol-X), possibly due to an original Mg deficiency or to partial exsolution of Mg during diagenesis. The nannomicrite (4) consists of low-Mg calcite only. 4. Three textural types of volcanic and associated gyroclastic rocks were distinguished: (1) holohyaline, rapidly chilled and granulated lava flows and tuffs (palagonite tuff breccia and hyaloclastic top breccia); (2) tachylitic basalts (less rapidly chilled; with opaque glass); and (3) "slowly" crystallized, holocrystalline alkali olivine basalts. The carbonate in most mixed pyroclastic-carbonate sediments at the basalt contact is of "post-eruptive" origin (micritic crusts etc.); "pre-eruptive" limestone is recrystallized or altered at the basalt contact. A deuteric (?hydrothermal) "mineralX", filling vesicles in basalt and cementing pyroclastic breccias is described for the first time. 5. Origin and development of GMS andJS: From its origin, some 85 m. y. ago, the volcano of GMS remained active until about 10 m. y. B. P. with an average lava discharge of 320 km**3/m. y. The volcanic origin of JS is much younger (?Middle Tertiary), but the volcanic activity ended also about 9 m. y. ago. During L a t e Miocene to Pliocene times both volcanoes were eroded (wave-rounded cobbles). The oldest pyroclastics and carbonates (MF 3.1, 3.2) were originally deposited in shallow-water (?algal reef hardground). The Plio (-Pleisto) cene foraminiferal nannomicrites (MF 4) suggest a meso- to bathypelagic environment along the flanks of GMS. During the Quaternary (?Pleistocene) bioclastic sands were deposited in water depths beyond wave base on the summit tops, repeatedly reworked, and lithified into loosely consolidated biopelsparites and biomicrites (MF 1 and 2; Fig. 15). Intermediate steps were a first intragranular filling by micrite, reworking, oncoidal coating, weak consolidation with Mg-calcite cemented "peloids" in intergranular voids and local compaction of the peloids into cryptocrystalline micrite with interlocking Mg-calcite crystals up to 4p. The submarine lithification process was frequently interrupted by long intervals of nondeposition, dissolution, boring, and later infilling. The limestones were probably never subaerially exposed. Presently, the carbonate rocks undergo biogenic incrustation and partial dissolution into bioclastic sands. The irregular distribution pattern of the sands reflects (a) the patchy distribution of living benthonic organisms, (b) the steady rain of planktonic organism onto the seamount top, (c) the composition of disintegrating subrecent limestones, and (d) the intensity of winnowing and reworking bottom current

Mineralogy and petrographic summary of ODP Leg 198 sites and DSDP Hole 62-463, Shatsky Rise

This petrological study of the lower Aptian Oceanic Anoxic Event (OAE1a) focused on the nature of the organic-rich interval as well as the tuffaceous units above and below it. The volcaniclastic debris deposited just prior to the OAE1a is consistent with reactivation of volcanic centers across the Shatsky Rise, concurrent with volcanism on the Ontong Java Plateau. This reactivation may have been responsible for the sub-OAE1a unconformity. Soon after this volcanic pulse, anomalous amounts of organic matter accumulated on the rise, forming a black shale horizon. The complex textures in the organic-rich intervals suggest a history of periodic anoxia, overprinted by bioturbation. Components include pellets, radiolarians, and fish debris. The presence of carbonate-cemented radiolarite under the OAE1a intervals suggests that there has been large-scale remobilization of carbonate in the system, which in turn may explain the absence of calcareous microfossils in the section. The volcanic debris in the overlying tuffaceous interval differs in that it is significantly epiclastic and glauconitic. It was likely derived from an emergent volcanic edifice.