Luminescence dating methods are widely used to date coastal sediments, while luminescence tracing methods are an upcoming approach to reconstruct coastal sediment pathways. Both methods rely on subaqueous resetting (bleaching) of luminescence signals and would benefit from quantification of this process in the natural coastal environment. We describe the set-up and outcomes of an in situ subaqueous bleaching experiment for luminescence signals of K-feldspar grains in the Dutch Wadden Sea. We deployed a full-day bleaching field experiment with irradiated feldspar samples tethered to a pole at various positions within and above the water column to quantify (1) the bleaching potential, that is, the light intensity and spectrum as a function of time, depth and tidal stage, and (2) the bleaching efficiency, that is, the degree of bleaching of infrared stimulated luminescence (IRSL) and post-infrared IRSL (pIRIR) signals measured at 150, 225 and 290°C after a full day of light exposure above and below water. Our bleaching-potential results show that the strongest subaqueous light attenuation took place during low tide when sediment concentrations are the highest. We also observed stronger attenuation of the ultraviolet part of the spectrum compared to other parts of the spectrum. Our bleaching-efficiency results show that bleaching reduces with depth, that pIRIR signals bleach slower than IRSL signals underwater and that bleaching efficiency reduces with pIRIR measurement temperature. None of the investigated signals were fully reset after 13.5 hours of light exposure, even for subaerially exposed samples. Our work provides the first quantitative data on pIRIR bleaching in a natural subaqueous environment, which is relevant for K-feldspar-based luminescence dating and tracing applications.

The impact of Sacculina carcini infection on the nutritional status of the shore crab Carcinus maenas was investigated in the western Dutch Wadden Sea for a period of 20 months. About 3.3% of the population was sacculinized, i.e. externally infected with S. carcini and only 0.7% presented scars of previous infection. The results of mixed linear models showed that sacculinized and non-sacculinized crabs had similar morphometric condition, while the energy density of parasitized crabs (externa excluded) was significantly reduced by about 4.3% overall, and by up to 5.8% in crabs under 40 mm carapace width. However, when Sacculina externa was included in the energy determinations, the difference in energy density decreased to 1.2%, while total energy content of the pair infected crab-parasite including externa was 30.8% higher than non-sacculinized crabs of similar size. The total energy content of ovigerous females (eggs included) was even higher, near doubling the energy of similar-sized crabs. The same way, total energy content of Sacculina externa was about 4 times lower than total energy of egg mass. The results suggest that the rhizocephalan parasite is efficient in consuming the energy that the host may allocate for growth and maintenance, but require future studies to disentangle the impact of the degree of internal infection and the implications for the dynamics of the population.

The occurrence of 19 cetacean species along the German North Sea coastline as well as the lower reaches of the major rivers discharging into the German Bight is reviewed for the period 1604–2017 based on records of dead animals, either stranded dead or put to death. The harbour porpoise (Phocoena phocoena) is considered the most abundant and only native species in German coastal and riverine waters. Based on written sources its presence can be traced back to at least 1651, although with statistical data only available from 1990. Finds of further 18 species have been documented: white-beaked dolphin (Lagenorhynchus albirostris), bottlenose dolphin (Tursiops truncatus), Atlantic white-sided dolphin (Lagenorhynchus acutus), common dolphin (Delphinus delphis), striped dolphin (Stenella coeruleoalba), Risso's dolphin (Grampus griseus), long-finned pilot whale (Globicephala melas), killer whale (Orcinus orca), beluga whale (Delphinapterus leucas), narwhal (Monodon monoceros), Sowerby's beaked whale (Mesoplodon bidens), northern bottlenose whale (Hyperoodon ampullatus), sperm whale (Physeter macrocephalus), minke whale (Balaenoptera acutorostrata), sei whale (Balaenoptera borealis), fin whale (Balaenoptera physalus), blue whale (Balaenoptera musculus) and humpback whale (Megaptera novaeangliae). This review corrects several false species assignments earlier introduced in literature based on incorrect scientific or ambiguous German vernacular names and recovers lost records of beluga whale, northern bottlenose whale, sperm whale and fin whale.

Accelerator mass spectrometry (AMS) radiocarbon (14C) dating of Cerastoderma edule (Linnaeus 1767) and Mytilus edulis (Linnaeus 1758) shells sampled in AD 1889 near the island of Wangerooge gave a new local correction factor ΔR of −85 ± 17 14C years for the Wadden Sea area. The value is considerably higher than the available scattered data from the North Sea, which were obtained from pre-bomb growth rings of living Arctica islandica (Linnaeus 1767). This can be explained by the incorporation of 14C-depleted terrestrial carbon into the shell material which compensates the intensified exchange of CO2 between atmosphere and shallow coastal water, e.g. by tidal currents. Additionally, two examples of application of the new ΔR value in coastal research give deeper insights into the dynamics of bivalve shell preservation in the Wadden Sea and the need for further research to clarify the Holocene reintroduction of Mya arenaria (Linnaeus 1758) into European waters.

Rising sea levels due to climate change can have severe consequences for coastal populations and ecosystems all around the world. Understanding and projecting sea-level rise is especially important for low-lying countries such as the Netherlands. It is of specific interest for vulnerable ecological and morphodynamic regions, such as the Wadden Sea UNESCO World Heritage region.

Here we provide an overview of sea-level projections for the 21st century for the Wadden Sea region and a condensed review of the scientific data, understanding and uncertainties underpinning the projections. The sea-level projections are formulated in the framework of the geological history of the Wadden Sea region and are based on the regional sea-level projections published in the Fifth Assessment Report of the Intergovernmental Panel on Climate Change (IPCC AR5). These IPCC AR5 projections are compared against updates derived from more recent literature and evaluated for the Wadden Sea region. The projections are further put into perspective by including interannual variability based on long-term tide-gauge records from observing stations at Den Helder and Delfzijl.

We consider three climate scenarios, following the Representative Concentration Pathways (RCPs), as defined in IPCC AR5: the RCP2.6 scenario assumes that greenhouse gas (GHG) emissions decline after 2020; the RCP4.5 scenario assumes that GHG emissions peak at 2040 and decline thereafter; and the RCP8.5 scenario represents a continued rise of GHG emissions throughout the 21st century. For RCP8.5, we also evaluate several scenarios from recent literature where the mass loss in Antarctica accelerates at rates exceeding those presented in IPCC AR5.

For the Dutch Wadden Sea, the IPCC AR5-based projected sea-level rise is 0.07±0.06m for the RCP4.5 scenario for the period 2018–30 (uncertainties representing 5–95%), with the RCP2.6 and RCP8.5 scenarios projecting 0.01m less and more, respectively. The projected rates of sea-level change in 2030 range between 2.6mma−1 for the 5th percentile of the RCP2.6 scenario to 9.1mma−1 for the 95th percentile of the RCP8.5 scenario. For the period 2018–50, the differences between the scenarios increase, with projected changes of 0.16±0.12m for RCP2.6, 0.19±0.11m for RCP4.5 and 0.23±0.12m for RCP8.5. The accompanying rates of change range between 2.3 and 12.4mma−1 in 2050. The differences between the scenarios amplify for the 2018–2100 period, with projected total changes of 0.41±0.25m for RCP2.6, 0.52±0.27m for RCP4.5 and 0.76±0.36m for RCP8.5. The projections for the RCP8.5 scenario are larger than the high-end projections presented in the 2008 Delta Commission Report (0.74m for 1990–2100) when the differences in time period are considered. The sea-level change rates range from 2.2 to 18.3mma−1 for the year 2100.

We also assess the effect of accelerated ice mass loss on the sea-level projections under the RCP8.5 scenario, as recent literature suggests that there may be a larger contribution from Antarctica than presented in IPCC AR5 (potentially exceeding 1m in 2100). Changes in episodic extreme events, such as storm surges, and periodic (tidal) contributions on (sub-)daily timescales, have not been included in these sea-level projections. However, the potential impacts of these processes on sea-level change rates have been assessed in the report.

Ground surface dynamics is one of the processes influencing the future of the Wadden Sea area. Vertical land movement, both subsidence and heave, is a direct contributor to changes in the relative sea level. It is defined as the change of height of the Earth's surface with respect to a vertical datum. In the Netherlands, the Normaal Amsterdams Peil (NAP) is the official height datum, but its realisation via reference benchmarks is not time-dependent. Consequently, NAP benchmarks are not optimal for monitoring physical processes such as land subsidence. However, surface subsidence can be regarded as a differential signal: the vertical motion of one location relative to the vertical motion of another location. In this case, the actual geodetic height datum is superfluous.

In the present paper, we highlight the processes that cause subsidence, with specific focus on the Wadden Sea area. The focus will be toward anthropogenic causes of subsidence, and how to understand them; how to measure and monitor and use these measurements for better characterisation and forecasting; with some details on the activities in the Wadden Sea that are relevant in this respect. This naturally leads to the identification of knowledge gaps and to the formulation of notions for future research.

The Wadden Sea is a unique coastal wetland containing an uninterrupted stretch of tidal flats that span a distance of nearly 500km along the North Sea coast from the Netherlands to Denmark. The development of this system is under pressure of climate change and especially the associated acceleration in sea-level rise (SLR). Sustainable management of the system to ensure safety against flooding of the hinterland, to protect the environmental value and to optimise the economic activities in the area requires predictions of the future morphological development.

The Dutch Wadden Sea has been accreting by importing sediment from the ebb-tidal deltas and the North Sea coasts of the barrier islands. The average accretion rate since 1926 has been higher than that of the local relative SLR. The large sediment imports are predominantly caused by the damming of the Zuiderzee and Lauwerszee rather than due to response to this rise in sea level. The intertidal flats in all tidal basins increased in height to compensate for SLR.

The barrier islands, the ebb-tidal deltas and the tidal basins that comprise tidal channels and flats together form a sediment-sharing system. The residual sediment transport between a tidal basin and its ebb-tidal delta through the tidal inlet is influenced by different processes and mechanisms. In the Dutch Wadden Sea, residual flow, tidal asymmetry and dispersion are dominant. The interaction between tidal channels and tidal flats is governed by both tides and waves. The height of the tidal flats is the result of the balance between sand supply by the tide and resuspension by waves.

At present, long-term modelling for evaluating the effects of accelerated SLR mainly relies on aggregated models. These models are used to evaluate the maximum rates of sediment import into the tidal basins in the Dutch Wadden Sea. These maximum rates are compared to the combined scenarios of SLR and extraction-induced subsidence, in order to explore the future state of the Dutch Wadden Sea.

For the near future, up to 2030, the effect of accelerated SLR will be limited and hardly noticeable. Over the long term, by the year 2100, the effect depends on the SLR scenarios. According to the low-end scenario, there will be hardly any effect due to SLR until 2100, whereas according to the high-end scenario the effect will be noticeable already in 2050.

Tidal inlets and the associated ebb-tidal deltas can significantly impact the coastal sediment budget due to their ability to store or release large quantities of sand. Nearly 300 million m3 (mcm) of sediments were eroded from Texel Inlet's ebb-tidal delta and the adjacent coasts following the closure of the Zuiderzee in 1932. This erosion continues even today as a net loss of 77 mcm was observed between 1986 and 2015. To compensate, over 30 mcm of sand has been placed on the adjacent coastlines since 1990, making maintenance of these beaches the most intensive of the entire Dutch coastal system.

Highly frequent and detailed observations of both the hydrodynamics and morphodynamics of Texel Inlet have resulted in a unique dataset of this largest inlet of the Wadden Sea, providing an opportunity to investigate inlet sediment dynamics under the influence of anthropogenic pressure. By linking detailed measurements of bathymetric change to direct observations of processes we were able to unravel the various components that have contributed to the supply of sediment to the basin, and develop a four-stage conceptual model describing the multi-decadal adaptation of the ebb-tidal delta.

Prior to closure of the Zuiderzee a dynamic equilibrium state (stage 1) existed with a stable ebb-tidal delta. The largest morphological changes occurred in roughly the first 40 years since the closure, and were dominated by the rotation and scouring of large tidal channels and landward retreat of the Noorderhaaks ebb shoal (stage 2; adaptation). Between 1975 and 2001 the general layout of main channels and shoals was stable, but large sediment losses continued to occur (stage 3; equilibrium erosional state). Since 2001, the erosion rates have significantly reduced to 2 mcm a−1 (stage 4; stabilisation).

Twenty-five years of data on ‘Dynamic Preservation’ prove that sand nourishments are well capable of keeping the coastlines adjacent to the Texel Inlet in place. Moreover, the abundant supply of sediment may also have compensated for the sediment losses on the larger scale of the southern part of the ebb-tidal delta, resulting in a recent stabilisation of its volume. This response illustrates the potential benefits of Dynamic Preservation not only for coastline resilience but also on the larger scale of the inlet system. Such knowledge is essential for future preservation, management and maintenance of inlet systems in the scope of climate change and accelerated sea-level rise.

Cores from the salt marshes along the drowned melt-water valley of river Varde Å in the Danish Wadden Sea have been dated and analysed (litho- and biostratigraphically) to reconstruct the Holocene geomorphologic evolution and relative sea level history of the area. The analysed cores cover the total post-glacial transgression, and the reconstructed sea level curve represents the first unbroken curve of this kind from the Danish Wadden Sea, including all phases from the time where sea level first reached the Pleistocene substrate of the area. The sea level has been rising from − 12 m below the present level at c. 8400 cal yr BP, interrupted by two minor drops of < 0.5 m at c. 5500 cal yr BP and 1200 cal yr BP, and one major drop of ∼ 1.5 m at c. 3300 cal yr BP. Sediment deposition has been able to keep pace with sea level rise, and the Holocene sequence consists in most places of clay atop a basal peat unit overlying sand of Weichselian age and glacio-fluvial origin. In its deepest part, the basal peat started to form around 8400 cal yr BP, and reached a thickness of up to 3.5 m. This thickness is about half of the original, when corrected for auto-compaction. The superimposed clay contains small (63–355 μm) red iron stains in the top and bottom units, and foraminifers of the calcareous type in the middle. The fact that iron stains and foraminifers in no cases coexist, but always exclude each other is interpreted as a result of the difference between salt-marsh facies (iron stains) and tidal-flat facies (foraminifers). This represents a novel and easy way to distinguish between these two otherwise often undistinguishable sedimentary facies in the geological record.

Cores and exposed cliff sections in salt marshes around Ho Bugt, a tidal embayment in the northernmost part of the Danish Wadden Sea, were subjected to 14C dating and litho- and biostratigraphical analyses to reconstruct paleoenvironmental changes and to establish a late Holocene relative sea-level history. Four stages in the late Holocene development of Ho Bugt can be identified: (1) groundwater-table rise and growth of basal peat (from at least 2300 BC to AD 0); (2) salt-marsh formation (0 to AD 250); (3) a freshening phase (AD 250 to AD 1600?), culminating in the drying out of the marshes and producing a distinct black horizon followed by an aeolian phase with sand deposition; and (4) renewed salt-marsh deposition (AD 1600? to present). From 16 calibrated AMS radiocarbon ages on fossil plant fragments and 4 calibrated conventional radiocarbon ages on peat, we reconstructed a local relative sea-level history that shows a steady sea-level rise of 4 m since 4000 cal yr BP. Contrary to suggestions made in the literature, the relative sea-level record of Ho Bugt does not contain a late Holocene highstand. Relative sea-level changes at Ho Bugt are controlled by glacio-isostatic subsidence and can be duplicated by a glacial isostatic adjustment model in which no water is added to the world's oceans after ca. 5000 cal yr BP.

Quantifying exchange of particulate matter between coastal and open waters is an important and often unresolved issue. Here, we apply phytoplankton order richness as an innovative marine tracer to identify the geographic position of a coastal exchange zone in the SE North Sea, including its variability in time and space. Previous observations on dynamics of suspended particulate matter accumulation resulted in a hypothesized boundary between coastal waters (including the Wadden Sea) and open North Sea waters, the so-called ‘line-of-no-return’. Our study along two transects (Terschelling, Noordwijk) in the Dutch coastal zone showed seasonality patterns in phytoplankton order richness, both for diatoms and flagellates. The coastal Wadden Sea was found to be clearly different from the open North Sea, implying that seasonality in Wadden Sea phytoplankton is at least partly driven by local environmental conditions. Seasonality in flagellates was found to be more uniform than seasonality in diatoms. Stations in the coastal North Sea to a distance of 10 km (Terschelling) to 20 km (Noordwijk) from the shore appeared to be at the inside of the ‘line-of-no-return’. Our findings indicate that this approach is a useful aid in exploring mixing of particulate matter between coastal and open waters and to study the responses of phytoplankton communities to environmental drivers.

During the Pleistocene, the coastal marine bivalve mollusc Mya arenaria became extinct in northwest Europe. The species remained present in North America. Datings of Mya shells found in northern Denmark and the southern Baltic Sea suggest that repopulation of northwest European coasts already occurred before Columbus’ discovery of America (1492), possibly facilitated by Viking (Norse) settlers at Greenland and northeast North America. In this paper we report on findings of M. arenaria at five locations in the coastal landscape of the Netherlands: polders reclaimed from the Wadden Sea and the former estuaries of Oer-IJ and Old Rhine. The shells from four of these locations also date before 1492 AD.

During an aerial survey on 30 December 2014 in the Danish Wadden Sea, a newborn grey seal pup (Halichoerus grypus) with umbilical cord was observed at a sandbank located north of the island Rømø together with the mother. This is the first report in the scientific literature on grey seals breeding in the Danish Wadden Sea since the species disappeared from the area in the 16th Century.

In this article we study the morphodynamics of the Slufter on the short-term (months) and long-term (years to decades). The Slufter is a small, shallow tidal inlet located on the island of Texel, the Netherlands. A narrow (tens of meters) channel connects the North Sea with a dune valley of 400 ha. This narrow channel is located in between a 400-700 m wide opening in the dunes. Approximately 80% of the basin of the Slufter is located above mean high water level and is flooded only during storms, when a threshold water level is exceeded.

Analysis of historical aerial photographs revealed that the inlet channel migrates about 100 m per year. In the 1970's it migrated to the south, while since 1980 it is migrating to the north. When the channel reached the dunes at the north side of the dune breach the channel was relocated to the south by man. The channel inside the backbarrier basin was less dynamic. It shows a gradual growth and southward migration of a meander on a decadal time scale.

The short-term dynamics of the Slufter were studied during a field campaign in 2008. The campaign aimed at identifying the dominant hydrodynamic processes and morphological change during fair weather conditions and during storm events. During fair weather flow velocities in the main inlet channel were 0.5-0.8 m/s at water depths of 0-1.5 m, slightly ebb-dominant and associated morphological change was small. When water levels were above critical levels during a storm period the hydrodynamics in the main channel drastically changed. The flow in the main channel was highly ebb dominant. Long ebb periods with typical flow velocities of 2 m/s were alternated by much shorter flood periods with typical velocities of 0.5-1 m/s. This resulted in a net outflow of water via the main channel, while we measured a net inflow of water at the beach plain. During the storm period in 2008 we measured a 10 m migration of the channel to the north.

We conclude that the Slufter is a storm-dominated tidal inlet system.

Modelling of the shallow subsurface of the Dutch Wadden Sea is merely based on lithological information extracted from a limited amount of core samples. In order to improve the subsurface model and to provide a better basis for engineering purposes, seismic data have recently been acquired, processed and interpreted. This study focuses on the interpretation of seismic data in a pilot area in the southwestern part of the Dutch Wadden Sea near the Afsluitdijk. In order to acquire a maximum detail of subsurface information in a time-efficient way, multiple types of seismic systems were deployed simultaneously in a ‘one-sweep-survey’, providing information over depth ranges up to 60 m subsurface depth. Data from three seismic systems are presented; a chirp system, a boomer and sparker source in combination with hydrophone streamers. Geological interpretation of the seismic data was made by identifying seismic facies units and subsequently correlating them to geological cross-sections, running parallel to the Afsluitdijk. Geological cross-sections were derived from the existing geological and hydrogeological model and from relatively densely spaced borehole information. Six key reflectors were identified on the seismic data along the Dutch Afsluitdijk that make up four seismic facies units. Results of seismic profiles show good recognition of internal structures in especially Holocene sediments. A clay plug and a shallowing of a channel at the eastern side of the pilot area were interpreted as channel infills resulting from the rather sudden dominance by newer tidal channels to the west, probably coinciding with the opening of the Marsdiep channel. The channel wall deposits observed were interpreted as a turning of the drainage channel after closure of the IJsselmeer. Strong reflections of deeper levels (>15 m below Dutch vertical datum, i.e. N.A.P.) were interpreted as clay/sand interfaces in the Middle-Pleistocene Urk Formation and were more continuous than previously thought. It is concluded that high resolution seismics add valuable information yielding improved understanding of the sedimentary structure of the shallow subsurface, which in turn can be useful for near future engineering works along the Afsluitdijk.

The availability of nearly 100 years of bathymetric measurements allows the analysis of the morphodynamic evolution of the Dutch Wadden Sea under rising sea level and increasing human constraint. The historically observed roll-over mechanisms of landward barrier and coastline retreat cannot be sustained naturally due to numerous erosion control measures that have fixed the tidal basin and barrier dimensions. Nevertheless, the large continuous sedimentation in the tidal basins (nearly 600 million m3), the retained inlets and the similar channel-shoal characteristics of the basins during the observation period indicate that the Wadden Sea is resilient to anthropogenic influence, and can import sediment volumes even larger than those needed to compensate the present rate of sea-level rise. The largest sedimentation occurs in the Western Wadden Sea, where the influence of human intervention is dominant. The large infilling rates in closed-off channels, and along the basin shoreline, rather than a gradual increase in channel flat heights, render it likely that this sedimentation is primarily a response to the closure of the Zuiderzee and not an adaptation to sea-level rise. Most of the sediments were supplied by the ebb-tidal deltas. It is, however, unlikely that the sediment volume needed to reach a new equilibrium morphology in the Western Wadden Sea can be delivered by the remaining ebb-tidal deltas alone.

The survival of the highly endangered, anadromous fish species North Sea houting (Coregonus oxyrinchus) depends on the correct timing of downstream dispersal during its early ontogenetic stages. To date, however, no studies have investigated the ontogenetic differentiation of swimming performance and behaviour, including the potential of habitat complexity to influence dispersal rates. By testing larval and juvenile North Sea houting in a laboratory, we examined (1) swimming performance measured as maximum swimming performance (Umax) and routine swimming speed (Uroutine) and (2) the potential of habitat complexity (i.e., cover providing shade) to influence dispersal behaviour in an indoor stream channel. The Umax and the Uroutine were 9.4 and 4.6cm s-1, respectively, in the larvae [body length (BL) s-1: 7.3 and 3.5, respectively], and 25.2 and 16.3 cm s-1 in the juveniles (BL s-1: 7.0 and 5.2, respectively). We compared laboratory swimming performance data with water speeds in North Sea houting spawning areas in the Danish River Vidaa. Results showed that the water speeds present in 95% and 85% of the water column caused downstream displacement of larvae and juveniles, respectively. However, areas with slow-flowing water near river banks and river beds could function as nursery habitats. Stream channel experiments showed that cover providing shade caused delayed dispersal in both larvae and juveniles, but the larvae dispersed later and spent less time under cover than the juveniles, a finding that implies ontogenetic effects. Finally, the larvae refused to cross an upstream-positioned cover, a behaviour that was not observed in the juveniles. Therefore, habitat complexity may have the potential to influence dispersal behaviour in both larval and juvenile North Sea houting. Overall, we provided the first evidence of ontogenetic differentiation in the North Sea houting. These findings will be valuable for the development and dissemination of science-based conservation strategies.

Common seals Phoca vitulina give birth in the Wadden Sea area during the summer months. We provide evidence that the pupping season has advanced in date in this region. Analysis of stranding dates of recently-born, orphaned pups admitted for rehabilitation, revealed a shift of, on average, 0.88 days per year over the period 1974–2008, yielding a total advance of 26 days. Although the pupping season has become progressively earlier, there were no indications of any negative impact on the weight of the pups, nor was there an increase in the proportion of seals with a lanugo coat. These observations suggest that the most likely explanation for the change in phenology of the pupping season is a corresponding change in the timing of cessation of the period of delayed implantation. It is suggested that shifts in phenology could reflect an adaptive response of the animals to altered local circumstances. The latter may in turn be induced by larger scale phenomena such as food availability or climate change.

Light microscopy and scanning electron microscopy were used to study Amphidinium glabrum sp. nov., a new dinoflagellate species from tidal sand flats in the North German Wadden Sea and from the surface water and the lowermost layer of first-year ice in the European Arctic. Cells are strongly flattened dorso-ventrally, with nearly parallel sides in ventral view. The distal region of the strongly displaced cingulum is not connected with the sulcus. There is no surface striation. The sulcus extends from the antapex, where it forms a large excavation, to the apex and further onto the dorsal side of the epicone. The new species is compared with two morphologically similar dinoflagellates: A. scissum Kofoid & Swezy and A. scissoides Lebour. Characters that distinguish the three taxa are: cell shape and size, shape of epicone and hypocone, cell surface, cingulum displacement, distal region of the cingulum and antapical notch.