Palaeoecology and evolution of Mesozoic salinity-controlled benthic macroinvertebrate associations

Salinity-controlled benthic macroinvertebrate associations are typical of many Mesozoic marginally marine environments. They can be recognized by abiotic criteria (e.g., environmental setting, specific autigenic minerals), by biotic criteria (faunal composition, diversity, shell morphology, size-frequency histograms, taphonomic features, associated micro fauna and microflora), and by isotope geochemistry of shells. Although salinity-controlled associations must have been widespread in the European German Triassic, very little is known about their ecology. They appear to have been dominated by the bivalve Unionites and the brachiopod Lingula. In the Jurassic, brackish-water associations are characterized by bivalves, in particular neomiodontids, corbulids, mytilids, bakevelliids, isognomonids, and oysters. In the Cretaceous, in addition, corbiculid bivalves and gastropods become increasingly abundant. Salinity-controlled benthic macroinvertebrate associations can be used to reconstruct salinity regimes of ancient environments, but emphasis should be placed on an integrated sedimentological and ecological approach, as salinity is rarely the only parameter influencing faunal composition and diversity. Although the species composition of salinity-controlled benthic associations changes distinctly through time, the composition of morphotypes remains surprisingly constant throughout the Mesozoic and up to the Recent, evidence of a conservative evolution of benthic faunas within marginal marine high-stress environments. □ Salinity, benthic associations, palaeoecology, Mesozoic.     

Bivalve ecology in a Silurian shelf environment

The Ludlow Series of the Welsh Borderland represents a gradient from quiet. deep shelf muds to shallow, storm-deposited silts. A fauna of 44 bivalve species shows a pattern of overlapping environmental ranges which span the full extent of the shelf gradient. Bivalve abundance is highest at the extremes of this gradient, where endobyssate forms are common in storm-deposited silts, and nuculoids locally dominate the fauna of the deep shelf muds. Species diversity of bivalves, however, is highest in low-stress. mid-shelf environments, where bivalves of small population sizes comprise a minimal proportion of the benthic fauna. Niche sizes, species diversity. and total environmental range of the bivalves is comparable to that of Ludlow brachiopods, even though the latter group is far more abundant as individuals.     

Early Jurassic shell beds from marginal marine environments in southern Mendoza, Argentina

External estuarine facies in Late Sinemurian beds from the base of the Puesto Araya Formation in the Atuel river region, Mendoza Province, Argentina, contain concentrations of a new species of Cardinioides (a shallow burrowing suspension-feeding Pachycardiidae bivalve) and a low diversity benthic fauna. These bivalves appear in tidal inlet facies of a wave-dominated estuary (environment similar to lagoon-barrier island systems), forming bioclastic lags at the channel bases. The taphonomic attributes (such as shell articulation, degree of fragmentation and abrasion, bioclast size, orientation) change from North to South, from monospecific concentrations of nearly complete large Cardinioides shells with low degree of fragmentation and abrasion, through beds with smaller Cardinioides shells associated with a few bivalves from marine environments, to only isolated fragmented specimens associated with a fully marine biota including not only other bivalves but also brachiopods and ammonoids. This, together with different palaeoecologic, sedimentologic and stratigraphic attributes at the studied localities are evidence of southwards changing environmental conditions from marginal marine (brackish) to fully marine on the eastern margin of the Neuquén basin at that time. From detailed analysis of taphonomic features, size distribution of shells, and the associated fauna at the different localities, the new species is regarded as a brackish water endemic, possibly a low salinity euryhaline species, which inhabited well oxygenated waters. Salinity, turbulence levels and food supply were the main limiting factors in the distribution of the species.     

The impact of an introduced bivalve (Corbicula fluminea) on the benthos of a sandy stream

1. The effect of an exotic, burrowing bivalve ( Corbicula fluminea ) on the benthic fauna of a sandy-bottomed stream was assessed by field and laboratory experiments. Corbicula differs from other freshwater, non-native bivalves in that it both filter- and pedal-feeds and thus has the potential to influence the streambed community in different ways.
2. In the field, cages were used to vary the abundance of Corbicula in the streambed. Increasing abundance of Corbicula was negatively associated with the abundance of benthic bacteria and flagellates but had no apparent effect on other protists or meiofauna.
3. In the laboratory, we compared the effect of Corbicula on the benthic community when the bivalves were able both to filter- and pedal-feed with that when they were able only to filter-feed. Bivalves restricted to filter-feeding were placed in sediment lacking benthic fauna and organic matter, then the benthos in nearby natural sediment was compared with the community present when Corbicula was able to move freely through the sediment and both to filter- and pedal-feed. Corbicula able to pedal-feed were again associated with a decreased abundance of benthic flagellates and bacteria, as well as diatoms.     

Diversity and density characteristics of Pliensbachian-Toarcian molluscan and brachiopod faunas of the North Atlantic margins

Pliensbachian and Toarcian molluscs and brachiopods of Morocco, Iberia, England, and Greenland are analysed in terms of diversity and density. With regard to local variations, Substrate and food supply are thought to be the major factors controlling the predominance of suspension-feeding bivalves in shallow water deposits while the greater physical stability of deeper water favoured the ammonites; high diversity of the gastropods relates to firm substrates. On a regional scale, both bivalve density and diversity tend to increase from the Tethyan to the Boreal redm, while there is a Variable tendency for a northward diversity reduction in other groups. The bivalve data are incompatible with either temperature or salinity control of faunal provinciality. To provide a better account of this, a model is developed which lays emphasis on the physical instability of a boreal inland sea as compared with the Tethys. Some broader implications for endemism in Jurassic faunas are briefly discussed.     

The functional role of burrowing bivalves in freshwater ecosystems

1. Freshwater systems are losing biodiversity at a rapid rate, yet we know little about the functional role of most of this biodiversity. The ecosystem roles of freshwater burrowing bivalves have been particularly understudied. Here we summarize what is known about the functional role of burrowing bivalves in the orders Unionoida and Veneroida in lakes and streams globally. 2. Bivalves filter phytoplankton, bacteria and particulate organic matter from the water column. Corbicula and sphaeriids also remove organic matter from the sediment by deposit feeding, as may some unionids. Filtration rate varies with bivalve species and size, temperature, particle size and concentration, and flow regime. 3. Bivalves affect nutrient dynamics in freshwater systems, through excretion as well as biodeposition of faeces and pseudofaeces. Excretion rates are both size and species dependent, are influenced by reproductive stage, and vary greatly with temperature and food availability. 4. Bioturbation of sediments through bivalve movements increases sediment water and oxygen content and releases nutrients from the sediment to the water column. The physical presence of bivalve shells creates habitat for epiphytic and epizoic organisms, and stabilizes sediment and provides refugia for benthic fauna. Biodeposition of faeces and pseudofaeces can alter the composition of benthic communities. 5. There is conflicting evidence concerning the role of resource limitation in structuring bivalve communities. Control by bivalves of primary production is most likely when their biomass is large relative to the water volume and where hydrologic residence time is long. Future studies should consider exactly what bivalves feed upon, whether feeding varies seasonally and with habitat, and whether significant overlap in diet occurs. In particular, we need a clearer picture of the importance of suspension versus deposit feeding and the potential advantages and tradeoffs between these two feeding modes. 6. In North America, native burrowing bivalves (Unionidae) are declining at a catastrophic rate. This significant loss of benthic biomass, coupled with the invasion of an exotic burrowing bivalve (Corbicula), may result in large alterations of ecosystem processes and functions.     

侏罗纪双壳类动物群的分布与古气候和古地理因素

生物群的地理分布受各种生物和环境因素的影响,对于不同地史晚期以及不同的生物类型,控制生物群分布的主导因素也不同的。就侏罗纪古大西洋双壳类动物群而言古地理和古气候条件是影响其分布的主要环境因素。     

Middle and Upper Jurassic bivalve associations from the Iberian Range (Spain)

The four sections richest in bivalves from the Middle and Upper Jurassic of the Iberian Range (Spain) were selected for a quantitative palaeoecological analysis of the bivalve fraction of the macrobenthos. Five bivalve associations and two assemblages were recognized with the help of a Q-mode hierarchical cluster analysis (Ward method). The main environmental factors controlling bivalve associations are thought to be substrate, water energy and distribution of organic matter. The bivalves exhibit a distinct spatial and temporal distribution pattern within the Aragonian Branch of the Iberian Range. Four of the bivalve associations occur in the Upper Oxfordian (Sot de Chera Fm) and one association in the Lower Callovian (Chelva Fm). In the Sot de Chera and Loriguilla formations, the abundance of bivalves decreases from NW to SE i.e., from relatively close to the shore line towards the distal-most part of the carbonate platform. In the Chelva Fm, bivalves are abundant in the Ariño region, interpreted as a palaeogeographic high. The spatial distribution of bivalves might have been largely controlled by the availability of nutrients.     

Paläobiogeographie jurassischer Muschelfaunen: Beziehung zwischen Süd- und Nordrand der Tethys

Similarities of mid-Jurassic bivalve faunas between the European and the Ethiopian faunal province are very high at the genus-level. At the species-level, however, it is shown that during the Bathonian and Callovian 35% of the bivalves occurring in the Ethiopian faunal province are restricted to this province. In the region of Kachchh (W-India) in the same time-interval 25% of all bivalves are endemic. In the Ethiopian faunal province a clear tendency of increasing endemism from the Bathonian to the Tithonian/Lower Cretaceous at the genus-level and, even more obviously, at the species-level exists. Endemism and provincialism are most marked within the orders Arcoida, Trigonioida, and Nuculoida. The degree of endemism is lower within the Veneroida, but still very high. The orders Mytiloida, Pterioida, and Pholadomyoida hold the largest portion of cosmopolitan species. The rise of endemism and provincialism in Kachchh and m the Ethiopian faunal province from the Bathonian onwards can be explained only partly by the increasing broadening of the Tethys and its effect as an oceanic barrier. The steep increase of endemism in the Upper Jurassic of Kachchh is essentially caused by a radiation within the astartids and trigoniids, accompanied by a reduction of facies-types, due to a regional regression. The very southerly palaeogeographic position of India, the opening of the ‘South African Seaway’, and a change in the marine current system in the uppermost Jurassic led to an increasing differentiation of the Ethiopian faunal province in an ‘Ethiopian-Tethyan’ subprovince to the north and an ‘Ethiopian-Austral’ subprovince to the south. A migration of bivalves in mid-Jurassic times can be reconstructed along the southern margin of the Tethys mainly from east to west. On the other hand, an easternward migration of bivalves along the northern margin of the Tethys from Europe to China and Japan can be documented especially in the Upper Jurassic. This corroberates the existence of a clock-wise marine current system in the northern hemisphere in the Jurassic. The distribution patterns of bivalves in Kachchh and the Ethiopian faunal province are essentially characterized by ‘migration’ of bivalves. The opening of the ‘Hispanic Corridor’ in the Pliensbachian gave way to the immigration of East Pacific bivalves via the western Tethys as far as Kachchh and Madagascar. The dispersal ofPisotrigonia, Seebachia, Tendagurium, andMegacucullaea in the uppermost Jurassic/lowermost Cretaceous from Kachchh and East-Africa respectively to South-Africa and South-America documents the establishment of a ‘South-African Seaway’ and favours migration. However, ‘migration’ and ‘vicariance’ do not exclude each other. On the contrary, both are important mechanisms for creating distributional patterns of bivalves, although within different geological dimensions. Vicariance events produce faunal provinces which last for a long time and within this time-interval, migration seems to be the more important mechanism affecting palaeobiogeographic distribution of bivalves. There is no evidence that the distribution patterns of bivalves in Kachchh and in the Ethiopian faunal province are governed by eustatic sea-level changes. The dominating factors have been a change of the palaeogeographic constellation as a consequence of the break-up of Gondwana, and the local facies distribution. The number of bivalve species known from Europe is much larger than the number of species of the Ethiopian faunal province. A comparison of rarefaction curves of associations, however, shows that this is not a primary feature, but is a consequence of a greater number of different facies types and is due to a far more intense collecting activity in Europe. The bivalves of the Spiti Shales are unequivocal Ethiopian-Tethyan in character. The composition of the fauna indicates the deposition on the deeper shelf of the southern margin of the Tethys. All occurring ‘European’ faunal elements are not significant because of their more or less cosmopolitan distribution.     

Palaeoenvironmental changes across the Permian/Triassic boundary at Shangsi (N. Sichuan,China)

The section at Shangsi in Sichuan contains one of the most detailed and best records of events during the Permian/Triassic (P/T) mass extinction. Continuous deep water deposition is only punctuated by a minor shallowing in the late Changxingian. The micritic mudstones and wackestones of the Changxingian Dalong Formation contain abundant ammonoids and radiolaria and diverse and common benthic taxa (mostly bivalves and brachiopods) in a thoroughly bioturbated sediment. The presence of a well developed tiered burrow profile is just one line of evidence for a fully oxygenated water column in the late Permian. The faunal crisis occurs in the top few decimetres of the Dalong Formation and severely affected all groups (benthos, nekton and plankton). The extinction coincides precisely with the development of anaerobic and dysaerobic facies. The basal Triassic sediments of the Feixanguan Formation are thinnly‐bedded or laminated silicic marls and contain pyrite and several levels of elevated organic carbon concentrations. The fauna is restricted to rare ammonoids and a few bedding planes covered in Claraia. The presence of abundant coccoid cyanobacteria in these sediments may indicate an unusually simple trophic web in the early Triassic seas as these picoautotrophs are normally grazed by zooplankton, they are rarely directly incorporated into seafloor sediment. The recent discovery of black shales in P/T pelagic sediments of Japan indicates that the anoxic event also affected deep ocean waters and further strengthens the link between extinction and anoxia.     

Benthic palaeoecology of Danian deep-shelf bryozoan mounds in the Danish Basin

Early Danian cool-water bryozoan mounds exposed in the coastal cliff Stevns Klint in Denmark were formed shortly after the Cretaceous–Tertiary mass extinction. They represent a relatively deep-water, highly diverse benthic ecosystem within the epeiric seaway that covered the Danish Basin. The mounds are 50–110 m long and reached a height of about 5–10 m above the seafloor; they are asymmetrical with a steep southern and a gentle northern flank, and were dominated by small suspension feeders. The benthic elements generally occur as fragments set in a carbonate mud matrix. The main skeletal contributors are delicate branching bryozoans with minor contributions of bryozoan sheets, and nodular/arborescent bryozoans. Locally abundant octocorals occur on the mound crests and upper parts of the steep flanks. Echinoids are present in minor amounts, but are locally abundant. Serpulids, crinoids, asteroids, brachiopods, bivalves, massive calcareous sponges, and benthic foraminifers are generally minor contributors to the benthic mound fauna. Influx of planktonic foraminifers, coccoliths and other planktonic organisms was high and was probably a major source of nutrient supply to the mainly suspension-feeding benthic fauna.

The faunal association reflects a relatively low energy environment with a high, possibly seasonal influx of particulate nutrients. The best growth conditions with respect to nutrient influx were on the mound crest and upper steep flank reflected by the diverse and relatively largest benthic faunal elements. Periodic reworking and winnowing occurred across the entire mound structure but most prominent on the gentle northern flanks limiting the benthic growth and notably the colony density and size of delicate branching bryozoans. Vagile benthic faunas were also adapted to different areas on the mound. Irregular echinoids preferred the intermound areas within fine-grained wackestone–packstone facies where they ploughed through the sediment, whereas regular echinoids were epifaunal and preferred the upper parts of the mounds, possibly feeding mainly on bryozoans. Skeletons of both groups became concentrated at the toe of the steep flanks and in the intermound areas by physical reworking during major storms.

Changes in faunal composition on the mound crests occurred rhythmically on both small and large scale during mound growth. Rhythmically recurring faunal assemblages reflect alternating hydrodynamic conditions on the seafloor with respect to nutrient influx and energy, which probably were linked to short-term seasonal and long-term climatic variations; the long-term alternation may be within the Milankovitch frequency band. Blooming events of bryozoan sheets resulted from relatively short periods with large amounts of available food and suitable substrate. Successful colonisation by octocorals on the other hand reflected longer-term favourable conditions on the mounds possibly associated with overall higher energy levels.

A possible Pleistocene analogue to the bryozoan-dominated Danian mounds occurs at the shelf-slope break of the Great Australian Bight. Both of these cool-water mound systems deviate from most other biogenic mounds known from the fossil record in their non-cemented nature, regular geometry and a lack of core and flank facies.     

Ecological interactions on macroevolutionary time scales: clams and brachiopods are more than ships that pass in the night

Competition among organisms has ecological and evolutionary consequences. However, whether the consequences of competition are manifested and measureable on macroevolutionary time scales is equivocal. Marine bivalves and brachiopods have overlapping niches such that competition for food and space may occur. Moreover, there is a long‐standing debate over whether bivalves outcompeted brachiopods evolutionarily, because brachiopod diversity declined through time while bivalve diversity increased. To answer this question, we estimate the origination and extinction dynamics of fossil marine bivalve and brachiopod genera from the Ordovician through to the Recent while simultaneously accounting for incomplete sampling. Then, using stochastic differential equations, we assess statistical relationships among diversification and sampling dynamics of brachiopods and bivalves and five paleoenvironmental proxies. None of these potential environmental drivers had any detectable influence on brachiopod or bivalve diversification. In contrast, elevated bivalve extinction rates causally increased brachiopod origination rates, suggesting that bivalves have suppressed brachiopod evolution.     

Marine paleoecology in the Upper Devonian of New York

In the clastic Genesee Group of the Catskill delta, lateral changes of the fauna are believed to reflect onshore-offshore physicochemical gradients. A shoreward increase of infauna is interpreted as adaptation to increased environmental stress. Free immobile taxa were concentrated offshore, while vagile forms, presumably able to cope with shifting substrata, are dominant nearshore. A shorewards replacement of brachiopods by bivalves reflects the eurytopy and infaunal habits of the bivalves.
In Gencsee time, progradation was first rapid, then slow. The sequence is reversed in the superjacent Sonyea Group and the accompanying reversal of faunal patterns is strong evidence of faunal control by the rate of progradation. This indicates the hazardous nature of attempts to trace 'community evolutioneditor' using only a few studies from each period.     

背角无齿蚌(Anodonta woodiana)对浅水系统底栖和浮游藻类竞争关系的影响

底栖藻类和浮游藻类之间的竞争关系对浅水生态系统的结构、功能具有重要的影响,双壳类可通过滤食控制浮游藻类,从而改变底栖藻类与浮游藻类之间的竞争结果。论文通过比较放养背角无齿蚌(Anodonta woodiana)(蚌处理组)与不放养背角无齿蚌(对照组)系统中底栖藻类、浮游藻类的生物量和优势种等的变化,研究了滤食性双壳类对底栖藻类和浮游藻类间竞争的影响。结果表明,背角无齿蚌可显著降低浮游藻类生物量,提高水体透明度和沉积物表面光照条件,从而显著提高底栖藻类的生物量;背角无齿蚌也改变了浮游藻类的优势种,使优势种由蓝藻转变成硅藻。因此,滤食性双壳类有利于促进浅水生态系统从混水态向清水态转变,本研究结果对富营养化浅水湖泊修复与管理具有一定的参考意义。     

Sedimentology,taphonomy, and palaeoecology of a laminated plattenkalk from the Kimmeridgian of the northern Franconian Alb (southern Germany)

At Wattendorf in the northern Franconian Alb, southern Germany, centimetre- to decimetre-thick packages of finely laminated limestones (plattenkalk) occur intercalated between well bedded graded grainstones and rudstones that blanket a relief produced by now dolomitized microbialite-sponge reefs. These beds reach their greatest thickness in depressions between topographic highs and thin towards, and finally disappear on, the crests. The early Late Kimmeridgian graded packstone–bindstone alternations represent the earliest plattenkalk occurrence in southern Germany. The undisturbed lamination of the sediment strongly points to oxygen-free conditions on the seafloor and within the sediment, inimical to higher forms of life. The plattenkalk contains a diverse biota of benthic and nektonic organisms. Excavation of a 13 cm thick plattenkalk unit across an area of 80 m² produced 3500 fossils, which, with the exception of the bivalve Aulacomyella, exhibit a random stratigraphic distribution. Two-thirds of the individuals had a benthic mode of life attached to hard substrate. This seems to contradict the evidence of oxygen-free conditions on the sea floor, such as undisturbed lamination, presence of articulated skeletons, and preservation of soft parts. However, palaeoecological and taphonomic analyses indicate that the benthic faunal elements are allochthonous having settled out of suspension and thus must have been derived from hard substrate areas of neighbouring topographic highs. Solely the bivalve Aulacomyella, which occurs concentrated on a single bedding plane, may have colonised the plattenkalk depression during brief periods of oxygenation. Alternatively, a pseudoplanktonic mode of life, attached to floating sea weed, is envisaged for the bivalve. The formation of the plattenkalk is related to local and global factors: a drop in relative sea level caused the death of the microbialite-sponge reefs. Restricted circulation in depressions between the crests of the former reefs led to anoxic conditions and the formation of finely laminated sediments stabilised by microbial mats. The small depressions received sediment and skeletal elements of benthic organisms that were swept off neighbouring topographic highs by storms and that entered the depressions partly as turbidity currents, partly as suspension clouds. A rise in the relative sea level finally terminated the restricted circulation in the depressions and closed the taphonomic window that led to the preservation of the plattenkalk biota.     

An ecologically mixed brachiopod fauna from Changhsingian deep-water basin of South China: consequence of end-Permian global warming

The Late Permian Shaiwa Group of the Ziyun area of Guizhou, South China is a deep-water facies succession characterized by deep-water assemblages of pelagic radiolarians, foraminifers, bivalves, ammonoids and brachiopods. Here we report 20 brachiopod species in 18 genera from the uppermost Shaiwa Group. This brachiopod fauna is latest Changhsingian in age and dominated by productides. The palaeoecologic and taphonomic analysis reveals that the brachiopod fauna is preserved in situ. The attachment modes and substratum preference demonstrate that the Shaiwa brachiopod fauna comprises admixed elements of deep-water and shallow-water assemblages. The presence of the shallow-water brachiopods in the Shaiwa faunas indicates the involuntary settlement of shallow-water brachiopods. The stressed ecologic pressure, triggered by warming surface waters, restricted ecospace and short food sources, may have forced some shallow-water elements to move to hospitable deep-water settings and others to modify their habiting behaviours and exploit new ecospace in deep-water environments. We infer that the end-Permian global warming and subsequent transgression event may have accounted for the stressed environmental pressure in the shallow-water communities prior to the end-Permian mass extinction.     

Rare but large bivalves alter benthic respiration and nutrient recycling in riverine sediments

Bioturbation studies have generally analyzed small and abundant organisms while the contribution to the benthic metabolism by rare, large macrofauna has received little attention. We hypothesize that large, sporadic bivalves may represent a hot spot for benthic processes due to a combination of direct and indirect effects as their metabolic and bioturbation activities. Intact riverine sediments with and without individuals of the bivalve Sinanodonta woodiana were collected in a reach with transparent water, where the occurrence of the mollusk was clearly visible. The bivalve metabolism and its effects on sedimentary fluxes of dissolved gas and nutrients were measured via laboratory incubations of intact cores under controlled conditions. S. woodiana contributed significantly to O₂ and TCO₂ benthic fluxes through its respiration and to \({\text{NH}}_{4}^{ + }\), SRP and SiO₂ regeneration via its excretion. The bivalve significantly stimulated also microbial denitrification and determined a large efflux of CH₄, likely due a combination of bioturbation and biodeposition activities or to anaerobic metabolism within the mollusk gut. This study demonstrates that a few, large individuals of this bivalve produce significant effects on aerobic and anaerobic benthic metabolism and nutrient mobilization. Random sediment sampling in turbid waters seldom catches these important effects due to low densities of large fauna.     

Signs of predation in the Middle Jurassic of south-central Poland: evidence from echinoderm taphonomy

Distinct faunal aggregates are described from the Middle Jurassic (uppermost Bajocian/lowermost Bathonian and Middle Bathonian) clay deposits of Częstochowa area, south-central Poland. These aggregates are composed of molluscs (scaphopods, gastropods, bivalves, ammonites and belemnites), articulate brachiopods and echinoderms (asteroids, crinoids and echinoids). A large percentage of the fossils, especially bivalves, are fragmented, but some fossils are complete. Although most of the fossils are crushed and fragmented, they are still identifiable to at least the genus level. Thorough statistical analysis of taphonomic features indicates that the preservation of asteroid marginal plates is distinct from the ossicles derived from the host clays. The high frequency of bite marks and the good state of preservation suggest that the accumulations are the products of predation activities and most probably are the effect of regurgitation. Taking into account the rich and diverse fauna, the predator was a bottom-feeding generalist. Possible predators include palaeospinacid sharks, a tooth of which was collected from the same bedding surface, but not associated with regurgitated remains. Although the bite marks on the asteroid ossicles point to sharks as potential producers of regurgitates, other vertebrates, like durophagous pycnodontiform fish, cannot be excluded.     

Neogene paleoceanographic events recorded in an active-margin setting: Humboldt basin, California

Recognition of North Pacific paleoceanographic events in the marginal Humboldt (Eel River) basin of northern California enables correlation of stratigraphic sections and development of a chronostratigraphy. Paleoclimatically related coiling shifts in Neogloboquadrina pachyderma (Ehrenberg) and benthic foraminiferal datums form the basis of the chronostratigraphy. Benthic foraminiferal datums are defined by the occurrence of selected benthic species and abundance maxima of benthic biofacies. The compiled chronostratigraphy is used to refine reconstructions of the depositional history of Humboldt basin. Paleoceanographic events, recognized by the distribution of benthic foraminiferal biofacies, are used to infer paleoceanographic history along the northeastern Pacific margin.

The similarity in coiling curves of N. pachyderma from the marine sequence at DSDP Site 173 and the coastal Centerville Beach section of Humboldt basin and at other independently dated sites along the northeastern Pacific margin demonstrates that matching records of climatic oscillations is a reliable method of correlating marine sequences. Benthic fauna from the Centerville Beach section vary in phase with climatically related coiling shifts in N. pachyderma. In particular these data show an increase in displaced neritic fauna during inferred warm intervals and resurgence of deeper bathyal fauna during inferred cool events. Similar data are observed from the inland Eel River section, demonstrating that benthic foraminiferal trends recognized at Centerville Beach can be identified elsewhere in Humboldt basin. This in-phase benthic response to climatic fluctuations probably results from changes in vertical depth range of many benthic species in response to paleoclimatically related vertical changes in water-mass position.

Depositional histories reconstructed for two key sites in southern Humboldt basin indicate low rates of sediment accumulation during early basin filling with hemipelagic sediments. Initiation of turbidite sedimentation in the early Pliocene resulted in a sharp increase in rate of sediment accumulation. This increase in rate of sediment accumulation is partially a response to tectonic uplift in the northern Coast Ranges and may be an effect of realignment of motion between the Pacific and North American plates at about this time. The inland site shoaled more rapidly during turbidite sedimentation as a result of a higher rate of sediment accumulation. The rate of sediment accumulation increased again at this site in the late Pliocene during deposition of shelf and nearshore facies. The Eel River region subsided concurrent with deposition of these shallow-water deposits.