Natural patterns and anthropogenic disturbance in north Adriatic marine benthic assemblages: descriptive and methodological studies

Marine soft bottom systems show a high variability across multiple spatial and temporal scales. Both natural and anthropogenic sources of disturbance act together in affecting benthic sedimentary characteristics and species distribution. The description of such spatial variability is required to understand the ecological processes behind them. However, in order to have a better estimate of spatial patterns, methods that take into account the complexity of the sedimentary system are required. This PhD thesis aims to give a significant contribution both in improving the methodological approaches to the study of biological variability in soft bottom habitats and in increasing the knowledge of the effect that different process (both natural and anthropogenic) could have on the benthic communities of a large area in the North Adriatic Sea. Beta diversity is a measure of the variability in species composition, and Whittaker’s index has become the most widely used measure of beta-diversity. However, application of the Whittaker index to soft bottom assemblages of the Adriatic Sea highlighted its sensitivity to rare species (species recorded in a single sample). This over-weighting of rare species induces biased estimates of the heterogeneity, thus it becomes difficult to compare assemblages containing a high proportion of rare species. In benthic communities, the unusual large number of rare species is frequently attributed to a combination of sampling errors and insufficient sampling effort. In order to reduce the influence of rare species on the measure of beta diversity, I have developed an alternative index based on simple probabilistic considerations. It turns out that this probability index is an ordinary Michaelis-Menten transformation of Whittaker's index but behaves more favourably when species heterogeneity increases. The suggested index therefore seems appropriate when comparing patterns of complexity in marine benthic assemblages. Although the new index makes an important contribution to the study of biodiversity in sedimentary environment, it remains to be seen which processes, and at what scales, influence benthic patterns. The ability to predict the effects of ecological phenomena on benthic fauna highly depends on both spatial and temporal scales of variation. Once defined, implicitly or explicitly, these scales influence the questions asked, the methodological approaches and the interpretation of results. Problem often arise when representative samples are not taken and results are over-generalized, as can happen when results from small-scale experiments are used for resource planning and management. Such issues, although globally recognized, are far from been resolved in the North Adriatic Sea. This area is potentially affected by both natural (e.g. river inflow, eutrophication) and anthropogenic (e.g. gas extraction, fish-trawling) sources of disturbance. Although few studies in this area aimed at understanding which of these processes mainly affect macrobenthos, these have been conducted at a small spatial scale, as they were designated to examine local changes in benthic communities or particular species. However, in order to better describe all the putative processes occurring in the entire area, a high sampling effort performed at a large spatial scale is required. The sedimentary environment of the western part of the Adriatic Sea was extensively studied in this thesis. I have described, in detail, spatial patterns both in terms of sedimentary characteristics and macrobenthic organisms and have suggested putative processes (natural or of human origin) that might affect the benthic environment of the entire area. In particular I have examined the effect of off shore gas platforms on benthic diversity and tested their effect over a background of natural spatial variability. The results obtained suggest that natural processes in the North Adriatic such as river outflow and euthrophication show an inter-annual variability that might have important consequences on benthic assemblages, affecting for example their spatial pattern moving away from the coast and along a North to South gradient. Depth-related factors, such as food supply, light, temperature and salinity play an important role in explaining large scale benthic spatial variability (i.e., affecting both the abundance patterns and beta diversity). Nonetheless, more locally, effects probably related to an organic enrichment or pollution from Po river input has been observed. All these processes, together with few human-induced sources of variability (e.g. fishing disturbance), have a higher effect on macrofauna distribution than any effect related to the presence of gas platforms. The main effect of gas platforms is restricted mainly to small spatial scales and related to a change in habitat complexity due to a natural dislodgement or structure cleaning of mussels that colonize their legs. The accumulation of mussels on the sediment reasonably affects benthic infauna composition. All the components of the study presented in this thesis highlight the need to carefully consider methodological aspects related to the study of sedimentary habitats. With particular regards to the North Adriatic Sea, a multi-scale analysis along natural and anthopogenic gradients was useful for detecting the influence of all the processes affecting the sedimentary environment. In the future, applying a similar approach may lead to an unambiguous assessment of the state of the benthic community in the North Adriatic Sea. Such assessment may be useful in understanding if any anthropogenic source of disturbance has a negative effect on the marine environment, and if so, planning sustainable strategies for a proper management of the affected area.

Studio degli eventi erosivi in quattro aree campione della costa emiliano-romagnola

Il progetto di tesi ha analizzato i processi erosivi in atto in quattro aree della costa emilianoromagnola settentrionale, situate davanti a importanti foci fluviali: Volano, Reno, F. Uniti e Savio. Il lavoro di tesi si colloca all’interno di un progetto più ampio, che prevede la collaborazione tra la Regione Emilia Romagna, Servizio Geologico, Sismico e dei Suoli e l’Istituto di Scienze Marine ISMAR-CNR, sede di Bologna. Quest’ultimo ha acquisito, durante la campagna ERO2010, 210 km di profili sismici ad alta risoluzione in un’area sotto-costa, posta tra i 2 e gli 8 metri di profondità. Il lavoro di tesi ha usufruito di una nuova strategia di analisi, la sismica ad alta risoluzione (Chirp sonar), che ha permesso di identificare ed esaminare l’architettura geologica riconoscibile nei profili ed approfondire la conoscenza dell’assetto sub-superficiale dei depositi, ampliando le conoscenze di base riguardanti la dinamica dei litorali. L’interpretazione dei dati disponibili è avvenuta seguendo differenti fasi di studio: la prima, più conoscitiva, ha previsto l’identificazione delle evidenze di erosione nelle aree in esame mediante l’analisi della variazione delle linee di riva, l’osservazione delle opere poste a difesa del litorale e lo studio dell’evoluzione delle principali foci. Nella fase successiva le facies identificate nei profili sismici sono state interpretate in base alle loro caratteristiche geometriche ed acustiche, identificando le principali strutture presenti e interpretando, sulla base delle informazioni storiche apprese e delle conoscenze geologiche a disposizione, i corpi sedimentari riconosciuti. I nuovi profili Chirp sonar hanno consentito la ricostruzione geologica mediante la correlazione dei dati a mare (database ISMAR-CNR, Bologna, Carta geologica dei mari italiani 1:250.000) con quelli disponibili a terra, quali Carta dell’evoluzione storica dei cordoni costieri (Servizio Geologico e Sismico dei Suoli, Bo) e Carta geologica 1:50.000 (Servizio Geologico d’Italia e Progetto CARG). La conoscenza dei termini naturali e antropici dello stato fisico dei sistemi costieri è il presupposto necessario per l'esecuzione di studi ambientali atti a una corretta gestione integrata della costa. L’analisi approfondita della geologia superficiale fornisce un’opportunità per migliorare il processo decisionale nella gestione dei litorali e nella scelta degli interventi da attuare sulla costa, che devono essere fatti consapevolmente considerando l’assetto geologico e prevedendo una strategia di manutenzione della costa a medio termine. Un singolo intervento di ripascimento produce effetti di breve durata e non sufficienti a sanare il problema e a mitigare il rischio costiero. Nei tratti costieri scarsamente alimentati, soggetti a persistenti fenomeni erosivi, occorre, pertanto, mettere in atto ripetuti interventi di ripascimento accompagnati da un idoneo piano di monitoraggio.

Benthic fauna at station FINO 1, 2005-2007

In der Nordsee wurden auf der Forschungsplattform FINO 1 Felduntersuchungen durchgeführt, um spezielle Fragen zu möglichen Auswirkungen von Offshore-Windenergieparks auf die marine Umwelt zu beantworten. Der Fokus war dabei auf die Konsequenzen für die Lebensgemeinschaft am Meeresboden gerichtet. Es wurden die benthosökologischen Prozesse im Nahbereich der Piles sowie die mittelfristige Entwicklung der Aufwuchsfauna auf der künstlichen Unterwasserstruktur dokumentiert. Die Ansammlung pelagischer Fischen um die Plattform und der Export organischen Materials von der Plattform wurden quantifiziert. Die räumliche Ausdehnung und die Erheblichkeit von Auswirkungen auf die Lebensgemeinschaften des Meeresbodens wurden anhand mathematischer Modellierung abgeschätzt. Zusätzlich wurde die Anwendbarkeit der elektrochemischen Akretionstechnologie zur Schaffung naturnaher Kalksubstrate in der Nordsee getestet und geeignete Parameter für eine erfolgreiche Umsetzung unter Nordseebedingungen ermittelt. Die auch 4,5 Jahre nach Errichtung der Plattform noch ansteigende Artenzahl der Aufwuchsfauna lässt darauf schließen, dass der Sukzessionsprozess noch nicht abgeschlossen ist. Die stark vertikal zonierte Aufwuchsfauna auf der Unterwasserkonstruktion erreicht eine Masse von ca. 5 Tonnen mit ausgeprägten saisonalen Schwankungen. Anhand von echoakustischen Untersuchungen wurden saisonal auftretende Ansammlungen pelagischer Fische um die Plattform dokumentiert. Der Nahbereich der Plattform unterschied sich durch eine Schillauflage und eine räumlich und zeitlich sehr variable Sediment- und Bodenfaunazusammensetzung deutlich von einem Referenzgebiet in 200-400 m Abstand von der Plattform. Eine konzentrische Zonierung mit unterschiedlich stark ausgeprägten Veränderungen der Bodenfauna lässt auf komplexe Veränderung des gesamten lokalen Nahrungsgefüges im Nahbereich der Plattform schließen. Anhand einer Modellierung konnte der Materialexport in die umgebenden Weichbodenbereiche für einzelne Piles und einen hypothetischen Windpark abgeschätzt werden. Die lokale Ausbildung einer hohen Biomasse auf der Unterwasserkonstruktion von WEA sowie der Export mit anschließender Sedimentation lassen zumindest lokal einen erheblichen Einfluss auf Stoff- und Energieflüsse erwarten.

Abundance of mesozooplankton in the western part of the Black Sea in 1995 during the Cooperative Marine Science Program for the Black Sea (CoMSBlack)

The "CoMSBlack-95" dataset is based on samples collected in the summer of 1995. The whole dataset is composed of 81 samples (28 stations) with data of zooplankton species composition, abundance and biomass. Samples were collected in discrete layers 0-10, 0-20, 0-50, 10-25, 25-50, 50-100 and from bottom up to the surface at depths depending on water column stratification and the thermocline depth. Zooplankton samples were collected with vertical closing Juday net,diameter - 36 cm, mesh size 150 µm. Tows were performed from surface down to bottom meters depths in discrete layers. Samples were preserved by a 4% formaldehyde sea water buffered solution. Sampling volume was estimated by multiplying the mouth area with the wire length. Mesozooplankton abundance: The collected material was analysed using the method of Domov (1959). Samples were brought to volume of 25-30 ml depending upon zooplankton density and mixed intensively until all organisms were distributed randomly in the sample volume. After that 5 ml of sample was taken and poured in the counting chamber which is a rectangle form for taxomomic identification and count. Large (> 1 mm body length) and not abundant species were calculated in whole sample. Counting and measuring of organisms were made in the Dimov chamber under the stereomicroscope to the lowest taxon possible. Taxonomic identification was done at the Institute of Oceanology by Asen Konsulov and Lyudmila Kamburska using the relevant taxonomic literature (Mordukhay-Boltovskoy, F.D. (Ed.). 1968, 1969,1972). Taxon-specific abundance: The collected material was analysed using the method of Domov (1959). Samples were brought to volume of 25-30 ml depending upon zooplankton density and mixed intensively until all organisms were distributed randomly in the sample volume. After that 5 ml of sample was taken and poured in the counting chamber which is a rectangle form for taxomomic identification and count. Copepods and Cladoceras were identified and enumerated; the other mesozooplankters were identified and enumerated at higher taxonomic level (commonly named as mesozooplankton groups). Large (> 1 mm body length) and not abundant species were calculated in whole sample. Counting and measuring of organisms were made in the Dimov chamber under the stereomicroscope to the lowest taxon possible. Taxonomic identification was done at the Institute of Oceanology by Asen Konsulov and Lyudmila Kamburska using the relevant taxonomic literature (Mordukhay-Boltovskoy, F.D. (Ed.). 1968, 1969,1972).

Benthic foraminiferal composition at the inner wall of the Middle America Trench, Costa Rica

The sediments of Deep Sea Drilling Project Site 565 and University of Texas Marine Science Institute Cores IG-24-7-38 to -42 taken on the landward slope of the Middle America Trench exhibit characteristics of material subject to reworking during downslope mass flow. These characteristics include a generally homogeneous texture, lack of sedimentary structures, pervasive presence of a penetrative scaly fabric, and presence of transported benthic foraminifers. Although these features occur throughout the sediments examined, trends in bulk density, porosity, and water content, and abrupt shifts in these index physical properties and in sediment magnetic properties at Site 565 indicate that downslope sediment creep is presently most active in the upper 45 to 50 m of sediment. It cannot be determined whether progressive dewatering of sediment has brought the material at this depth to a plastic limit at which sediment can no longer flow (thus resulting in its accretion to the underlying sediments) or whether this depth represents a surface along which slumping has occurred. We suspect both are true in part, that is, that mass movements and downslope reworking accumulate sediments in a mobile layer of material that is self-limiting in thickness.

Ostracode fauna of sediment core GIK23414-9 in the northeast Atlantic

Sediment core M23414 from the Rockall Plateau (North Atlantic) covering the last two climatic cycles, marine isotope stages (MIS) 7 to 1, was investigated for glacial-interglacial variations in the deep-sea benthic ostracode fauna. A highly diversified ostracode fauna including 98 species was found. Two climate-related assemblages were identified, associated with interglacial and peak glacial periods, respectively. The 'interglacial' group occurs during the end of MIS 7, 5 and 1 and is composed of the genera Henryhowella, Pelecocythere, Echinocythereis, Cytherella, Bradleya, Aversovalva and Eucytherura. The 'glacial' group consists of the genera Acetabulastoma (which is known as 'sea ice indicator' in the modern Arctic Ocean), Polycope, Bythoceratina, ?Rhombobythere, and some species possibly belonging to the genus Pseudocythere and is found during MIS 6, 4 and 2. These longer-term variations within the ostracode fauna seem related to the particular glacial and interglacial climate conditions that affected both deep-water production as well as primary production in the surface waters. However, a detailed comparison of ostracode abundances with the occurrence of events marked by increased ice-rafted debris reveals also much shorter-term climate related changes in the ostracode fauna. Thus, the temporal fluctuations within ostracode assemblages reflect long- and short-term alterations of the deep-sea environment that are clearly linked to climate changes.

Radiolarian chronology and fauna across the middle/late Eocene boundary at ODP Hole 171-1052A

Quantitative radiolarian assemblage analysis has been conducted on middle and upper Eocene sediments (Zones RP16 to RP18) from Ocean Drilling Program Site 1052 in order to establish the radiolarian magnetobiochronology and determine the nature of the faunal turnover across the middle/late Eocene boundary in the western North Atlantic Ocean. We recognize and calibrate forty-five radiolarian bioevents to the magneto- and cyclo-stratigraphy from Site 1052 to enhance the biochronologic resolution for the middle and late Eocene. Our data is compared to sites in the equatorial Pacific (Leg 199) to access the diachrony of biostratigraphic events. Eleven bioevents are good biostratigraphic markers for tropical/subtropical locations (south of 30°N). The primary markers (lowest occurrences of Cryptocarpium azyx and Calocyclas bandyca) which are tropical zonal boundary markers for Zones RP17 and RP18 provide robust biohorizons for correlation and age determination from the low to middle latitudes and between the Atlantic and Pacific Oceans. Some other radiolarian bioevents are highly diachronous (<1 million years) between oceanic basins. A significant faunal turnover of radiolarians is recognized within Chron C17n.3n (37.7 Ma) where 13 radiolarian species disappear rapidly in less than 100 kyr and 4 new species originate. The radiolarian faunal turnover coincides with a major extinction in planktonic foraminifera. We name the turnover phase, the Middle/Late Eocene Turnover (MLET). Assemblage analysis reveals the MLET to be associated with a decrease in low-mid latitude taxa and increase in cosmopolitan taxa and radiolarian accumulation rates. The MLET might be related to increased biological productivity rather than to surface-water cooling.

Radiolarian fauna of sediment core MD96-2089 off Lüderitz, Namibia

The study of radiolarian assemblages from Core MD 962086 provides new information on the variability in the upwelling intensity and origin of upwelled water masses over the past 350 ky in one of the major filamentous regions of the Benguela Upwelling System (BUS), located off Lüderitz, Namibia. The use of key radiolarian species to trace the source of upwelled waters, and the use of a radiolarian-based upwelling index (URI) to reconstruct the upwelling intensity represent the first use of radiolarians for paleoceanographic reconstructions in the BUS. These radiolarian-based proxies indicate strongest upwelling during Marine Isotope Stages (MIS) 3, 5, and 8, which compares well with other studies. While during MIS 3 and 8, the radiolarian-based proxies indicate the influx of waters of Southern Ocean origin, they also point to the increased influence of tropical waters during the lower portion of MIS 5. During MIS 2, 4 and 6 the radiolarian assemblages indicate generally lower upwelling intensities, although this signal is complicated by the increased occurrence of organic carbon in the sediments during these intervals. During MIS 2 there appears to be less of an input of Southern Ocean waters to the BUS, although during the also glacial MIS 4 and 6, there is evidence for an increased influence of cold Antarctic waters. The comparison of the results from Core MD 962086 with other studies in the BUS area indicates a non-uniform pattern of upwelling intensity and advection of cold, southern waters into this system during MIS 2. Weaker upwelling signaled by the radiolarian-based proxy in MIS 4 is in contrast to other studies that indicate higher productivity during this time period. In general, the data show that there is a strong spatiotemporal complexity in upwelling intensity in the BUS and that the advection of water into it is not strongly tied to glacial-interglacial variations in climate.

Diversity of planktonic foraminifer fauna found in 18 deep-sea cores from the North Atlantic

Greenland stadial/interstadial cycles are known to affect the North Atlantic's hydrography and overturning circulation and to cause ecological changes on land (e.g., vegetation). Hardly any information, directly expressed as diversity indices, however, exists on the impacts of these millennial-scale variations on the marine flora and fauna. We calculated three diversity indices (species richness, Shannon diversity index, Hurlbert's probability of interspecific encounter) for the planktonic foraminifer fauna found in 18 deep-sea cores covering a time span back to 60 ka. Clear differences in diversity response to the abrupt climate change can be observed and some records can be grouped accordingly. Core SO82-05 from the southern section of the subpolar gyre, the cores along the British margin and core MD04-2845 in the Bay of Biscay show two modes of diversity distribution, with reduced diversity (uneven fauna) during cold phases and the reverse (even fauna) during warm phases. Along the Iberian margin high species diversity prevailed throughout most of the glacial period. The exceptions were the Heinrich stadials when the fauna abruptly shifted from an even to an uneven or less even fauna. Diversity changes were often abrupt, but revealed a high resilience of the planktonic foraminifer faunas. The subtropical gyre waters seem to buffer the climatic effects of the Heinrich events and Greenland Stadials allowing for a quick recovery of the fauna after such an event. The current work clearly shows that planktonic foraminifer faunas quickly adapt to climate change, albeit with a reduced diversity.

Sedimentology, planktonic foraminifera distribution and stable isotope composition during marine isotope stage 3 of ODP Site 172-1060 in the West Atlantic

We have studied Ocean Drilling Program Site 1060 on the Blake Outer Ridge, which lies beneath the Gulf Stream. We focus on marine isotope stage 3, 60-25 thousand years before present (ka). Sea surface temperatures (SSTs) inferred both from foraminiferal fauna and alkenone ratios, as well as counts of iceberg melt-out debris and benthic stable isotope analyses, enable our record to be interpreted in terms of regional hydrographic changes as well as changing thermohaline circulation (THC). The observed SST record is consistent with the air temperature record from the Greenland ice cores. However, Site 1060 exhibits important differences in detail compared with the ice core record, and when compared to other sites within the North Atlantic, significant longitudinal differences emerge. At Site 1060 in the western Atlantic, all Greenland stadials (GS) whether associated with Heinrich events (HEs) or not, show a similar small amplitude of cooling; mean faunal-based SSTaug during GS is only 1.5°C colder than during Greenland interstadials (GIS). In addition, during GS the coldest SSTs are limited to apparently brief events. This is in contrast to several eastern Atlantic sites where HE stadials exhibit coolings that are enhanced by 2°C compared to other GS and where cold conditions are not restricted to cold pulses but cover 2 ka-long intervals. Furthermore, Site 1060 SSTs remained warm right through each interstadial, in contrast to the sustained and uniform cooling trend through interstadials that is consistently observed in Greenland, indicated by measurements of delta18O in ice.

Triassic marine ostracodes of the Australian Margin

Sixty-five species of benthic ostracodes have been discovered in the Triassic sediments of Ocean Drilling Program Leg 122, drilled on the northwestern margin of Australia. Known species were found in the samples studied from the upper Norian-Rhaetian at Holes 759B and 760B and from the Rhaetian at Holes 761C, 764A, and 764B. A large part of material of the recovered ostracodes belong to taxa that are related to ostracodes described in the Tethyan province. Seven species are known from northwestern Europe and five from Iran. Ogmoconcha and Rhombocythere, which are stratigraphically important genera in northern Europe, extend into the Tethyan province. Species of Ogmoconcha are present in Holes 760B, 764A, and 764B. The highly ornate baidiids of the Alpine Tethyan province are dominant in Hole 761C. The affinities with the fauna of Iran noted by Kristan-Tollmann are corroborated by the presence of the genera Mostlerella, Hiatobairdia, and other bairdiids.

Distribution of marine litter at bathyal and abyssal depths in the Mediterranean Sea

The distribution, type and quantity of marine litter accumulated on the bathyal and abyssal Mediterranean seafloor has been studied in the framework of the Spanish national projects PROMETEO and DOS MARES and the ESF-EuroDEEP project BIOFUN. Litter was collected with an otter trawl and Agassiz trawl while sampling for megafauna on the Blanes canyon and adjacent slope (Catalan margin, north-western Mediterranean) between 900 and 2700 m depth, and on the western, central and eastern Mediterranean basins at 1200, 2000 and 3000 m depth. All litter was sorted into 8 categories (hard plastic, soft plastic, glass, metal, clinker, fabric, longlines and fishing nets) and weighed. The distribution of litter was analysed in relation to depth, geographic area and natural (bathymetry, currents and rivers) and anthropogenic (population density and shipping routes) processes. The most abundant litter types were plastic, glass, metal and clinker. Lost or discarded fishing gear was also commonly found. On the Catalan margin, although the data indicated an accumulation of litter with increasing depth, mean weight was not significantly different between depths or between the open slope and the canyon. We propose that litter accumulated in the canyon, with high proportions of plastics, has predominantly a coastal origin, while litter collected on the open slope, dominated by heavy litter, is mostly ship-originated, especially at sites under major shipping routes. Along the trans-Mediterranean transect, although a higher amount of litter seemed to be found on the Western Mediterranean, differences of mean weight were not significant between the 3 geographic areas and the 3 depths. Here, the shallower sites, also closer to the coast, had a higher proportion of plastics than the deeper sites, which had a higher proportion of heavy litter and were often affected by shipping routes. The weight of litter was also compared to biomass of megafauna from the same samples. On the Blanes slope, the biomass of megafauna was significantly higher than the weight of litter between 900 and 2000 m depth and no significant differences were found at 2250 and 2700 m depth. Along the trans-Mediterranean transect, no significant differences were found between biomass and litter weight at all sites except in two sites: the Central Mediterranean at 1200 m depth, where biomass was higher than litter weight, and the Eastern Mediterranean at 1200 m depth, where litter weight was higher than biomass. The results are discussed in the framework of knowledge on marine litter accumulation, its potential impact on the habitat and fauna and the legislation addressing these issues.

Composition of fossil manganese nodules from Costa Rica

Horizons of several types of Upper Jurassic to Lower Cretaceous manganese nodules occur locally in sequences of radiolarian cherts within the Nicoya Ophiolite Complex (NW Costa Rica). Field studies, X-ray diffraction analysis, petrographic, chemical and experimental studies give evidence of a sedimentary, early diagenetic origin of the nodules, in contrast to earlier suggestions. Smooth, discoidal, compact and very dense nodules with diameters of some mm to 9 cm dominate. They are characterized by braunite, hollandite, pyrolusite and quartz as well as 39-61% Mn, 0.9-1.6% Fe, 5-26% SiO2, 1.3-1.9% Al2O3, 1.5-3.0% Ba, 460-5400 ppm Cu, 85-340 ppm Ni and 40-130 ppm Co, among others. It is suggested that the original mineralogy (todorokite?) was altered during thermometamorphic (braunite) and hydrothermal (hollandite. pyrolusite) events. Petrographic similarities between the fossil nodules and modern deep-sea nodules are striking. Using standard hydrothermal techniques in an experimental study it is shown that under special conditions, braunite can be produced from modern nodule material.