How Population Decline Can Impact Genetic Diversity: a Case Study of Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis>) in Morro Bay,California

Seagrass populations are in decline worldwide. Eelgrass (Zostera marina L.), one of California’s native seagrasses, is no exception to this trend. In the last 8 years, the estuary in Morro Bay, California, has lost 95% of its eelgrass. Population bottlenecks like this one often result in severe reductions in genetic diversity; however, this is not always the case. The decline of eelgrass in Morro Bay provides an opportunity to better understand the effects of population decline on population genetics. Furthermore, the failure of recent restoration efforts necessitates a better understanding of the genetic underpinnings of the population. Previous research on eelgrass in California has demonstrated a link between population genetic diversity and eelgrass bed health, ecosystem functioning, and resilience to disturbance and extreme climatic events. The genetic diversity and population structure of Morro Bay eelgrass have not been assessed until this study. We also compare Morro Bay eelgrass to Bodega Bay eelgrass in Northern California. We conducted fragment length analysis of nine microsatellite loci on 133 Morro Bay samples, and 20 Bodega Bay samples. We found no population differentiation between the remaining beds in Morro Bay and no difference among samples growing at different tidal depths. Comparisons with Bodega Bay revealed that Morro Bay eelgrass contains three first-generation migrants from the north, but Morro Bay remains considerably genetically differentiated from Bodega Bay. Despite the precipitous loss of eelgrass in Morro Bay between 2008 and 2017, genetic diversity remains relatively high and comparable to other populations on the west coast.     

Impacts of Varying Estuarine Temperature and Light Conditions on <Emphasis Type="Italic">Zostera marina</Emphasis> (Eelgrass) and its Interactions With <Emphasis Type="Italic">Ruppia maritima</Emphasis> (Widgeongrass)

Seagrass populations have been declining globally, with changes attributed to anthropogenic stresses and, more recently, negative effects of global climate change. We examined the distribution of Zostera marina (eelgrass) dominated beds in the York River, Chesapeake Bay, VA over an 8-year time period. Using a temperature-dependent light model, declines in upriver areas were associated with higher light attenuation, resulting in lower light availability relative to compensating light requirements of Z. marina compared with downriver areas. An inverse relationship was observed between SAV growth and temperature with a change between net bed cover increases and decreases for the period of 2004–2011 observed at approximately 23 °C. Z. marina-dominated beds in the lower river have been recovering from a die-off event in 2005 and experienced another near complete decline in 2010, losing an average of 97 % of coverage of Z. marina from June to October. These 2010 declines were attributed to an early summer heat event in which daily mean water temperatures increased from 25 to 30 °C over a 2-week time period, considerably higher than previous years when complete die-offs were not observed. Z. marina recovery from this event was minimal, while Ruppia maritima (widgeongrass) expanded its abundance. Water temperatures are projected to continue to increase in the Chesapeake Bay and elsewhere. These results suggest that short-term exposures to rapidly increasing temperatures by 4–5 °C above normal during summer months can result in widespread diebacks that may lead to Z. marina extirpation from historically vegetated areas, with the potential replacement by other species.     

Light Requirements for Growth and Survival of Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) in Pacific Northwest (USA) Estuaries

We developed light requirements for eelgrass in the Pacific Northwest, USA, to evaluate the effects of short- and long-term reductions in irradiance reaching eelgrass, especially related to turbidity and overwater structures. Photosynthesis-irradiance experiments and depth distribution field studies indicated that eelgrass productivity was maximum at a photosynthetic photon flux density (PPFD) of about 350–550 μmol quanta m⁻² s⁻¹. Winter plants had approximately threefold greater net apparent primary productivity rate at the same irradiance as summer plants. Growth studies using artificial shading as well as field monitoring of light and eelgrass growth indicated that long-term survival required at least 3 mol quanta m⁻² day⁻¹ on average during spring and summer (i.e., May-September), and that growth was saturated above about 7 mol quanta m⁻² day⁻¹. We conclude that non-light-limited growth of eelgrass in the Pacific Northwest requires an average of at least 7 mol quanta m⁻² day⁻¹ during spring and summer and that long-term survival requires a minimum average of 3 mol quanta m⁻² day⁻¹.     

Twelve-Year Mapping and Change Analysis of Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis>) Areal Abundance in Massachusetts (USA) Identifies Statewide Declines

In 1994, 1995, and 1996, seagrasses in 46 of the 89 coastal embayments and portions of seven open-water near-shore areas in Massachusetts were mapped with a combination of aerial photography, digital imagery, and ground truth verification. In the open-water areas, 9,477.31 ha of seagrass were identified, slightly more than twice the 4,846.2 ha detected in the 46 coastal embayments. A subset of the 46 embayments, including all regions of the state were remapped in 2000, 2001, and 2002 and again in 2006 and 2007. We detected a wide range of changes from increases as high as 29% y⁻¹ in Boston Harbor to declines as large as −33% y⁻¹ in Salem Harbor. One embayment, Waquoit Bay, lost all of its seagrass during the mapping period. For the 12-year change analysis representing all geographic regions of the state, only three embayments exhibited increases in seagrass coverage while 30 of the original 46 embayments showed some indication of decline. For the decadal period, rates of decline in the individual embayments ranged from −0.06% y⁻¹ to as high as −14.81% y⁻¹. The median rate of decline by region ranged from −2.21% y⁻¹ to −3.51% y⁻¹ and was slightly less than the recently reported global rate of decline for seagrasses (−3.7% y⁻¹). Accounting for the gains in three of the embayments, 755.16 ha (20.6%) of seagrass area originally detected was lost during the mapping interval. The results affirm that previously reported losses in a few embayments were symptomatic of more widespread seagrass declines in Massachusetts. State and Federal programs designed to improve environmental quality for conservation and restoration of seagrasses in Massachusetts should continue to be a priority for coastal managers.     

Restoring Resiliency: Case Studies from Pacific Northwest Estuarine Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) Ecosystems

An objective of many ecological restoration projects is to establish resilience to disturbances. Eelgrass (Zostera marina L.) represents a useful model to evaluate resilience because the plant community is dominated by one species and the estuarine environment is dynamic. Our studies of planted and reference plots used shoot density monitoring data from three projects spanning 3 to 12 years. Data show that eelgrass can recover from major shifts in pond position and shape on sandflats, as well as natural disturbances causing >20-fold change in density. However, cumulative effects of multiple stressors on unestablished plantings suggest algal blooms of unusual magnitude can tip normally marginal conditions to unfavorable. Thus, potential resilience appears to depend on landscape conditions. A dynamic equilibrium was evinced in even the deepest, lowest-density plantings, probably associated with light-mediated carrying capacity and vegetative belowground production characteristic of the Pacific Northwest. We recommend eight resilience-related planning elements to reduce uncertainties in eelgrass restoration.     

Large-Scale Differences in Community Structure and Ecosystem Services of Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis>) Beds Across Three Regions in Eastern Canada

Eelgrass (Zostera marina) forms extensive beds in temperate coastal and estuarine environments worldwide and provides important ecosystem services, including habitat for a wide range of species as well as nutrient cycling and carbon storage. However, little is known about how eelgrass ecosystem structure and services differ naturally among regions. Using large-scale field surveys, we examined differences in eelgrass bed structure, carbon and nitrogen storage, community composition, and habitat services across three distinct regions in Eastern Canada. We focused on eelgrass beds with low anthropogenic impacts to compare natural differences. In addition, we analyzed the relationships of eelgrass bed structure with environmental conditions, and species composition with bed structure and environmental conditions, to elucidate potential drivers of observed differences. Our results indicate that regional differences in eelgrass bed structure were weakly correlated with water column properties, whereas differences in carbon and nitrogen storage were mainly driven by differences in eelgrass biomass. There were distinct regional differences in species composition and diversity, which were particularly linked to temperature, as well as eelgrass bed structure indicating differences in habitat provision. Our results highlight natural regional differences in ecosystem structure and services which could inform spatial management and conservation strategies for eelgrass beds.     

Regional-Scale Differences in Eutrophication Effects on Eelgrass-Associated (<Emphasis Type="Italic">Zostera marina</Emphasis>) Macrofauna

Eutrophication has caused strong shifts from perennial seagrass to opportunistic macroalgae and phytoplankton in many coastal ecosystems worldwide, yet responses of the primary-producer assemblage can vary with regional environmental and nutrient-loading conditions. The wider consequences of this variable primary-producer response on the associated animal community are little known. We used large-scale field surveys across 12 study sites with low or high eutrophication levels in two geographic provinces in Atlantic Canada to examine region-specific responses of macrofauna associated with eelgrass beds. In both regions, abundances of all groups increased with eutrophication, but species richness of mobile fishes and invertebrates decreased. Generally, filter feeders, epibenthic detritivores and some herbivores increased, while more hypoxia sensitive species declined. Small fishes and invertebrate predators increased with eutrophication mirrored by decreases in their prey. Despite similar general trends, our results show distinct shifts in species composition in each geographic region associated with differences in food availability and predation refuge offered by phytoplankton and opportunistic epiphytic or benthic macroalgae as well as tolerance to an increasingly hostile physico-chemical environment. So far, the continued persistence of eelgrass beds at our “highly” eutrophied sites indicates intermediate eutrophication levels with short-term benefits for some species. However, the loss of sensitive species and decrease in species richness highlight that eutrophication has already changed seagrass ecosystems in Atlantic Canada. Our work suggests that mitigating these changes will require regional-scale management.     

Restored Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) as a Refuge for Epifaunal Biodiversity in Mid-Western Atlantic Coastal Bays

As nearshore ecosystems are increasingly degraded by human activities, active restoration is a critical strategy in ensuring the continued provision of goods and services by coastal habitats. After being absent for nearly six decades, over 1800 ha of the foundational species eelgrass (Zostera marina L.) has been successfully re-established in the coastal bays of the mid-western Atlantic, USA, but nothing is known about the recovery of associated animal communities in this region. Here, we determine the patterns and drivers of functional recovery in epifaunal invertebrates associated with the restored eelgrass habitat from 2001 to 2013. After less than a decade, the invertebrate community in the restored bed was richer, more even, and exhibited greater variation in functional traits than a nearby reference bed. Analysis of a suite of environmental and physical variables using random forests revealed these differences were primarily due to the increasing area and density of eelgrass, a direct consequence of ongoing restoration efforts. Based on analysis of functional traits, we propose that the rapid life histories of constituent organisms may have played a key role in their successful recovery. We also speculate that diverse epifaunal communities may have contributed to the restoration success through a well-described mutualism with eelgrass. Given that restored eelgrass now make up 32 % of total seagrass cover in the mid-Atlantic coastal bays, this restoration may conserve regional biodiversity by providing new and pristine habitat, particularly given the general decline of existing eelgrass in this region.     

Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) in the Chesapeake Bay Region of Mid-Atlantic Coast of the USA: Challenges in Conservation and Restoration

Decreases in seagrass abundance reported from numerous locations around the world suggest that seagrass are facing a global crisis. Declining water quality has been identified as the leading cause for most losses. Increased public awareness is leading to expanded efforts for conservation and restoration. Here, we report on abundance patterns and environmental issues facing eelgrass (Zostera marina), the dominant seagrass species in the Chesapeake Bay region in the mid-Atlantic coast of the USA, and describe efforts to promote its protection and restoration. Eelgrass beds in Chesapeake Bay and Chincoteague Bay, which had started to recover from earlier diebacks, have shown a downward trend in the last 5–10 years, while eelgrass beds in the Virginia coastal bays have substantially increased in abundance during this same time period. Declining water quality appears to be the primary reason for the decreased abundance, but a recent baywide dieback in 2005 was associated with higher than usual summer water temperatures along with poor water clarity. The success of eelgrass in the Virginia coastal bays has been attributed, in part, to slightly cooler water due to their proximity to the Atlantic Ocean. A number of policies and regulations have been adopted in this region since 1983 aimed at protecting and restoring both habitat and water quality. Eelgrass abundance is now one of the criteria for assessing attainment of water clarity goals in this region. Numerous transplant projects have been aimed at restoring eelgrass but most have not succeeded beyond 1 to 2 years. A notable exception is the large-scale restoration effort in the Virginia coastal bays, where seeds distributed beginning in 2001 has initiated an expanding recovery process. Our research on eelgrass abundance patterns in the Chesapeake Bay region and the processes contributing to these patterns have provided a scientific background for management strategies for the protection and restoration of eelgrass and insights into the causes of success and failure of restoration efforts that may have applications to other seagrass systems.     

The Bloom-Forming Macroalgae, <Emphasis Type="Italic">Ulva</Emphasis>, Outcompetes the Seagrass, <Emphasis Type="Italic">Zostera marina</Emphasis>, Under High CO<Subscript>2</Subscript> Conditions

While multiple species of macroalgae and seagrass can benefit from elevated CO₂ concentrations, competition between such organisms may influence their ultimate responses. This study reports on experiments performed with a Northwest Atlantic species of the macroalgae, Ulva, and the seagrass, Zostera marina, grown under ambient and elevated levels of pCO₂, and subjected to competition with each other. When grown individually, elevated pCO₂ significantly increased growth rates and productivity of Ulva and Zostera, respectively, beyond control treatments (by threefold and 27%, respectively). For both primary producers, significant declines in tissue δ¹³C signatures suggested that increased growth and productivity were associated with a shift from use of HCO₃^? toward CO₂ use. When grown under higher pCO₂, Zostera experienced significant increases in leaf and rhizome carbon content as well as significant increases in leaf carbon-to-nitrogen ratios, while sediments within which high CO₂ Zostera were grown had a significantly higher organic carbon content. When grown in the presence of Ulva; however, above- and below-ground productivity and tissue nitrogen content of Zostera were significantly lower, revealing an antagonistic interaction between elevated CO₂ and the presence of Ulva. The presence of Zostera had no significant effect on the growth of Ulva. Collectively, this study demonstrates that while Ulva and Zostera can each individually benefit from elevated pCO₂ levels, the ability of Ulva to grow more rapidly and inhibit seagrass productivity under elevated pCO₂, coupled with accumulation of organic C in sediments, may offset the potential benefits for Zostera within high CO₂ environments.     

Successful Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis>) Restoration in a Formerly Eutrophic Estuary (Boston Harbor) Supports the Use of a Multifaceted Watershed Approach to Mitigating Eelgrass Loss

From a watershed perspective, Boston Harbor, MA, USA is an ideal site for eelgrass restoration due to major wastewater improvements. Therefore, by focusing on site selection and transplant methods, high survival and expansion rates were recorded at four large eelgrass-restoration sites planted in Boston Harbor as partial mitigation for a pipeline construction project. Transplanted sites met and exceeded reference and donor bed habitat function after 2 years. Hand planting and seeding in checkerboard-patterned transplant plots were efficient and effective methods for jump-starting eelgrass growth over large areas. Although restoration through planting can be successful, it is highly site specific. Even using a published site-selection model, intensive fieldwork was required to identify sites at fine enough scale to ensure successful planting. Given the effort required to identify scarce potential sites, we recommend that future focus includes alternative mitigation strategies that can more adequately prevent eelgrass loss and address water quality degradation which is the leading cause of dieback, site unsuitability for planting, and lack of natural re-colonization.     

Secondary Production of Macrobenthic Communities in Seagrass (<Emphasis Type="Italic">Zostera marina</Emphasis>, Eelgrass) Beds and Bare Soft Sediments Across Differing Environmental Conditions in Atlantic Canada

In nearshore ecosystems, habitats with emergent structure are often assumed to have higher ecosystem functioning than habitats lacking structure. However, such habitat-specific differences may depend on the surrounding environment. In this study, I examine the robustness of habitat-specific differences in ecosystem functioning for seagrass (Zostera marina) and adjacent bare soft sediments across varying environmental conditions on the Atlantic Coast of Nova Scotia, Canada, using secondary production as a metric. I also examine relationships of community secondary production and faunal structure with measured environmental variables (water depth, temperature, exposure, sediment, and plant properties). Benthic secondary production (invertebrates ≥500 μm) was higher in seagrass compared to bare sediments only at exposed sites with sandy sediments low in organic content, deep and cool water, and high belowground plant biomass. A regression relating community secondary production to the environmental variables explained 56% of the variance, while a constrained ordination explained 16% of the community structure. Important environmental determinants of community production were shoot density, temperature, depth, exposure, sediment organic content, and belowground plant biomass. Community structure was influenced by these variables plus sediment sand content and canopy height. This study shows that habitat-specific differences in secondary production may not be consistent across varying environmental conditions. Furthermore, seagrass beds are not always associated with higher ecosystem functioning than adjacent bare sediment. Both the surrounding environmental conditions and the presence of habitat structure should be considered for optimal management of nearshore ecosystems.     

Feeding Behavior of the Native Mussel <Emphasis Type="Italic">Ischadium recurvum</Emphasis> and the Invasive Mussels <Emphasis Type="Italic">Mytella charruana</Emphasis> and <Emphasis Type="Italic">Perna viridis</Emphasis> in FL,USA, Across a Salinity Gradient

The feeding behavior of three species of mussels, the native Ischadium recurvum and the invasives Mytella charruana and Perna viridis, was studied in an invaded ecosystem in Florida (USA). In situ feeding experiments using the biodeposition method were performed along a salinity gradient in four different locations along the St. Johns River. Water characteristics, such as salinity, temperature, dissolved oxygen, and seston loads, were recorded to identify relationships between these variables and the feeding behavior of the mussels. Feeding behavior of the species varied by study site. Clearance, filtration, organic ingestion, and absorption rates of I. recurvum were negatively affected by salinity. For the invasive mussel, M. charruana, rejection was positively related to salinity while total ingestion, organic ingestion, and absorption rates were positively related to the percentage of organic matter in the seston. For P. viridis, total and organic ingestion rates were negatively affected by salinity but positively affected by total particulate matter. Condition indices for P. viridis and M. charruana were 13.16?±?0.64 and 6.63?±?0.43, respectively, compared to 4.82?±?0.41 for the native species I. recurvum, indicating that these mussels are well adapted to the environmental conditions in the area. This study indicates that the three species have different preferred habitats because of their specific responses to water characteristics. Thus, the invasive mussels will not totally occupy the niche of the native mussel in Florida despite overlapping zones. These data may help identify potential invaded areas and understand the extent of the invasion.     

Depth colonization of eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis>) and macroalgae as determined by water transparency in Danish coastal waters

We present a comparative analysis of lower depth limits for growth of eelgrass, large brown algae and other macroalgae measured by SCUBA-diving along 162 transects in 27 Danish fjords and coastal waters, coupled to 1,400 data series of water chemistry (especially nitrogen) and Secchi depth transparency collected between March and October. Danish coastal waters are heavily eutrophied and characterized by high particle concentrations, turbid water and lack of macrophyte growth in deep water. Median values are 3.6 m for Secchi depth and median lower-depth limits are 4.0 m for eelgrass, 5.3 m for brown algae and 5.0 m for other macroalgae. Depth limits for growth of eelgrass and macroalgae increase linearly with transparency in the coastal waters. The relationships are highly significant (p<10⁻⁶) and transparency accounts for about 60% of the variability of depth limits. Eelgrass extends approximately to half the maximum depth of macroalgae, presumably because of greater respiratory costs to maintain the below-ground rhizomes and roots of eelgrass, which often constitutes half the plant weight. As a reflection of the importance of total nitrogen (TN) in controlling phytoplankton biomass and thus Secchi depth in coastal marine waters, we found that TN could explain 48–73% of the variation in depth limits of eelgrass and macroalgae, according to a multiplicative model (Y=aX^b). As with Secchi depth, the relationship to eelgrass showed a lower slope, reflecting the higher respiratory costs of eelgrass. The models show great sensitivity and a profound quantitative response with proportional effects on Secchi depth and depth limits when total-N concentrations are reduced.     

Sulfate Reduction and Sulfur Cycles at Two Seagrass Beds Inhabited by Cold Affinity <Emphasis Type="Italic">Zostera marina</Emphasis> and Warm Affinity <Emphasis Type="Italic">Halophila nipponica</Emphasis> in Temperate Coastal Waters

To evaluate the impact of invading seagrass on biogeochemical processes associated with sulfur cycles, we investigated the geochemical properties and sulfate reduction rates (SRRs) in sediments inhabited by invasive warm affinity Halophila nipponica and indigenous cold affinity Zostera marina. A more positive relationship between SRR and below-ground biomass (BGB) was observed at the H. nipponica bed (SRR = 0.6809 × BGB ? 4.3162, r ² = 0.9878, p = 0.0006) than at the Z. marina bed (SRR = 0.3470 × BGB ? 4.0341, r ² = 0.7082, p = 0.0357). These results suggested that SR was more stimulated by the dissolved organic carbon (DOC) exuded from the roots of H. nipponica than by the DOC released from the roots of Z. marina. Despite the enhanced SR in spring-summer, the relatively lower proportion (average, 20%) of acid-volatile sulfur (AVS) in total reduced sulfur and the strong correlation between total oxalate-extractable Fe (Fe_(oxal)) and chromium-reducible sulfur (CRS = 0.2321 × total Fe_(oxal) + 1.8180, r ² = 0.3344, p = 0.0076) in the sediments suggested the rapid re-oxidation of sulfide and precipitation of sulfide with Fe. The turnover rate of the AVS at the H. nipponica bed (0.13 day^?1) was 2.5 times lower than that at the Z. marina bed (0.33 day^?1). Together with lower AVS turnover, the stronger correlation of SRR to BGB in the H. nipponica bed suggests that the extension of H. nipponica resulting from the warming of seawater might provoke more sulfide accumulation in coastal sediments.     

Co-occurrence of <Emphasis Type="Italic">Neusticosaurus edwardsii</Emphasis> and <Emphasis Type="Italic">N. peyeri</Emphasis> (Reptilia) in the Lower Meride Limestone (Middle Triassic,Monte San Giorgio)

A newly opened excavation in the Cassina beds of the Lower Meride Limestone (Monte San Giorgio UNESCO World Heritage List, Canton Ticino, Switzerland), has yielded a pachypleurosaurid (Reptilia: Sauropterygia) specimen which is identified as Neusticosaurus peyeri. The resulting co-occurrence of N. peyeri and N. edwardsii, the latter so far regarded as the sole species of the genus present in this horizon, challenges the hypothesis of a single anagenetic lineage in Neusticosaurus species from Monte San Giorgio. In addition, it leads to a reconsideration of the phylogenetic inferences about Neusticosaurus evolution in the Monte San Giorgio area. The stratigraphic distribution of the Neusticosaurus species in the Monte San Giorgio basin is updated on the basis of recent finds.     

Toxicity effects of four typical nanomaterials on the growth of <Emphasis Type="Italic">Escherichia coli</Emphasis>, <Emphasis Type="Italic">Bacillus subtilis</Emphasis> and <Emphasis Type="Italic">Agrobacterium tumefaciens</Emphasis>

To offer an insight into the toxicity of nanomaterials (NM) on the growth of bacteria, Escherichia coli (E. coli), Bacillus subtilis (B. subtilis) and Agrobacterium tumefaciens (A. tumefaciens) were exposed to nano-Au, nano-Ag, nano-Fe and fullerene (C₆₀) in this study. As an effective bactericide, nano-Ag induced high toxicity on these three bacteria; C₆₀ could inhibit their growth; however, B. subtilis and E. coli could recover as exposure time extended. Nano-Au and nano-Fe had hardly any effect on three bacteria. A. tumefaciens showed the lowest resistance and slowest growth rate during exposure. Images obtained by scanning electron microscope (SEM) revealed that nano-Ag could cause damage to the cell structure of three bacteria at 1 μg/mL. Slight damage on E. coli was found when exposed to C₆₀, whereas no obvious physical damage was found after exposure to nano-Au or nano-Fe. It is assumed that surface activities of NM might be responsible for the different toxic effects on these bacteria.     

Life and death of <Emphasis Type="Italic">Saccocoma tenella</Emphasis> (G<Emphasis Type="SmallCaps">oldfuss</Emphasis>)

The morphology of the small stalkless Saccocoma tenella is unique among crinoids. It is characterized by an extremely light skeleton with dish-like lateral wings on the proximal brachials and peculiar paired vertical processes flanking the food grooves of more distal brachials. The arms are heavily branched. The lateral wings obviously were involved in vertical movement. For the vertical processes a “baffle rail” function for arm curling and “snap swimming” has been postulated, with muscles between the processes. However, there is no evidence that the processes were connected by muscles. For food collection a “pulsating funnel” model in the water column is advocated, with the processes serving to collect plankton during upward movement of the arms. The resulting mouth-up position is supported by the biostratinomy. Saccocoma tenella is considered to have been pelagic, a benthic lifestyle is rejected on ecologic and taphonomic grounds. Adorally-curled arms are considered a reaction to hostile environment before death, not a taphonomic artefact.