Recolonization of a high latitude hard-bottom nearshore community

Early recolonization of sessile organisms and grazer effects on recolonization was examined in a high latitude nearshore rocky environment. A manipulative experiment using cleared boulders and cages was used to determine the time needed for initial recruitment and composition of early recruits. The hypothesis tested was invertebrate grazing is causing slow recolonization. Four years of monitoring cleared boulders resulted in an initial recruitment of primarily barnacles in year 3 on boulders that were caged to exclude grazers. The total percent cover on these boulders after 4 years was less than 2%. Cage control and uncaged boulders showed less recruitment. Uncleared and uncaged control rocks showed insignificant temporal variation. Concurrent observations revealed that macroalgae were reproductive during the study and that space was limiting in this community. This study strongly suggests that perturbations causing removal of hard substrate sessile communities in the Beaufort Sea will result in very slow community recovery.     

Recruitment of the intertidal kelp Lessonia nigrescens Bory in northern Chile: successional constraints and opportunities

The strong 1982/83 El Niño event caused local extinction in populations of the low intertidal kelp Lessonia nigrescens Bory on northern Chilean coasts. The kelp has partially recolonized, but its recovery has apparently been low. This study documents the effect of biological factors that potentially decrease the velocity of kelp recolonization in northern Chile. A removal experiment showed that encrusting coralline algae, which dominate the lower intertidal, significantly reduce the recruitment of L. nigrescens in the presence of herbivores. Epithallium shedding by encrusting corallines is the most probable cause for this inhibition process. On the other hand, grazing on encrusting corallines by the chiton Enoplochiton niger (Barnes), a large-sized herbivore (10–20 cm length), could also affect kelp recruitment. The intestinal content of E. niger, where encrusting corallines are the main item (84.2%), revealed the presence of L. nigrescens. Among the natural substrata on which kelp recruits, measured at five different localities, the coarsely-branched alga Corallina officinalis (Decaisne) Kützing reached the highest frequency, despite its extremely low cover (< 1%) in the field. This suggests that settlement on turfs of C. officinalis allows the kelp to escape from herbivory, thus facilitating its recruitment. The roles of timing of kelp recruitment and seasonal grazing are also discussed.     

Competitive dominance among sessile marine organisms in a high Arctic boulder community

In most hard substrate environments, space is a limiting resource for sessile organisms. Competition for space is often high and is a structuring force within the community. In the Beaufort Sea’s Boulder Patch, crustose coralline red algae are major space occupiers. This research determined if coralline algae were competitively dominant over other sessile organisms. To test this hypothesis, overgrowth was documented in terms of “winners” and “losers” on the contact borders between different species. Crustose corallines occurred in over 80% of the observed interactions but were only winners in approximately half of them. Most frequently, bryozoans, tunicates, and sponges were superior competitors over crustose corallines, while at the same time these invertebrate groups were among the least abundant space occupiers.     

Sub-rubble communities of Curaçao and Bonaire coral reefs

The distribution and abundance of sessile organisms under coral rubble has been studied at Bonaire and Curaçao, Netherlands Antilles. Species richness under rubble is extremely high with at least 367 species of which sponges, tunicates and bryozoans are the most important. Shallow sub-rubble communities can be considered refuges as the majority of these species are crypt-obligate. Sub-rubble communities may also have a preserve function for sponges, but do not harbour enough corals to ensure a quick coral recolonization of the reef surface after a major disaster. Cryptic community composition is affected by depth and pollution, and differs substantially between the two neighbouring islands, possibly as a result of different bottom characteristics. Biomass of the sub-rubble communities may contribute considerably to total reef biomass. Diversity varies inversely with increased depth and increased rubble size, possibly indicating abiotic control (e.g. physical disturbance by wave action and reef slope substrate collapse).     

Early Cambrian coelobiontic communities in tectonically unstable crevices developed in Neoproterozoic andesites, Ossa-Morena, southern Spain

A well-preserved biota of Lower Cambrian cavity-dwelling organisms is recorded within fissures in Neoproterozoic andesites in Ossa-Morena (southern Spain). The cavities are unique among described Lower Cambrian coelobiontic communities due to the igneous character of the host rock. Coelobiontic habitat was episodically enlarged by synsedimentary tectonic fracturing reflecting polyphase infill of recurrent facies. The pioneer coelobiontic biota was diverse, and consisted of encrusting stromatolites and thromboids (dominated by Epiphyton and Renalcis), attached to walls and ceilings of the cavities, associated with archaeocyaths. Sponge spicules and chancelloriid sclerites occur as dense clusters indicating in situ growth, death and decay of spiculate sponges and coeloscleritophorans. Other organisms, such as echinoderms, trilobites and brachiopods, are also found within the cavities as reworked skeletons, indicating that they were washed in from the overlying, open seafloor. The main feature of the coelobiontic biota is the dominance of a sessile, chemosynthetic and filter-feeding epibenthos, composed of microbial communities, archaeocyaths, spiculate sponges (demosponges and rarer hexactinellides) and coeloscleritophorans.     

The relation of food availability and food preferences to the field diet of an echinoid Evechinus chloroticus (Valenciennes)

Echinoids are important in causing disturbance in many marine communities yet little is known of their foraging behaviour in the field. This work investigates the field diet of an echinoid species, Evechinus chloroticus (Valenciennes), in relation to food availability and food preferences. E. chloroticus is a generalist grazer occurring abundantly in a variety of algal and encrusting communities dominated by sponges. In the field, urchins grazed the sponge species according to abundance. In the laboratory, food preferences were demonstrated for two populations of E. chloroticus when presented with 24 species of sponge. One population of urchins came from an intertidal reef where sponges were rarely encountered and the other from a sublittoral habitat where the encrusting community was dominated by sponges. The sponges used as food in the preference experiments came from two different areas: one where urchins were almost absent, the other (from which the second group of urchins were taken) where urchins were abundant and were observed to feed upon sponges. When presented with sponge species unfamiliar to both urchin groups, urchins ranked sponge species similarly. Food preferences did not relate to the diet of the urchins in the field. Reasons for the difference between experimental results and field observations are suggested.     

The responses of a community to disturbance: The importance of successional age and species' life histories

Summary The responses of different successional stages of a temperate intertidal algal community to disturbance were investigated with a field experiment. The experiment was conducted in a low intertidal boulder field in southern California. In this habitat, the top surfaces of boulders are covered with algae. The composition of the assemblage on any particular boulder depends on the length of time since it was last overturned by wave action. When a boulder is overturned, the algae on what was formerly the top surface, are killed in whole or part by a combination of sea urchin grazing, anoxia, light levels below compensation intensity, and mechanical damage caused by crushing or abrasion. The length of time that a boulder remains overturned and the local abundance of sea urchins determines the intensity of the disturbance. When the boulder is righted, recolonization begins either by vegetative regrowth of survivors and/or by spores from outside.Using a three-factorial design, this natural form of disturbance was experimentally mimicked and the responses of three different successional stages of the algal community monitored. Boulders in each successional category were overturned for periods of 17, 27 and 54 days in areas with and without sea urchins, then righted. Two aspects of community response to perturbation were evaluated. These were (1) the assemblage's ability to resist change and (2) its ability, if altered, to adjust to some semblance of its original state. The resistance of each assemblage and of its component species to change was measured by the percent decrease in algal cover and by the decline in percent similarity of the community to its original composition. The recovery rate of each assemblage and of the cover lost by each species during the first 35 days following a disturbance was measured by the rate of increase in percent similarity to the original composition and the percent reestablishment of lost cover.The experimental evidence demonstrates that the successional stages of the producer level of an intertidal algal community differ significantly in their responses to disturbance. Early successional communities suffer more damage from a given level of perturbation but recover more quickly than either middle or late successional communities. Damage to any particular assemblage of algae, irrespective of successional age, is more extensive and recovery slower, the longer the boulder is overturned and/or sea urchins are present. Several thresholds in these responses were also identified.Differences in community responses and non-linearities in these responses were attributable to the life history characteristics of the component species rather than emergent properties of the assemblage. These characteristics have evolved in response to a variety of recurrent natural disturbances. This interpretation is in agreement with recent critical reevaluations of the trends and mechanisms of successional change in natural communities.     

Impact of iceberg scouring on macrobenthic communities in the high-Antarctic Weddell Sea

UW-video guided multibox corer sampling in- and outside iceberg scours provided quantitative macrozoobenthos samples for analyses of effects of grounding icebergs on infaunal benthic communities. These studies were performed on the southeastern Weddell Sea shelf off Kapp Norvegia and Austasen. Based on the UW-video sequences, stations were grouped a priori into two different "disturbance categories" and into undisturbed areas.Average biomass of major taxa in the cores of undisturbed areas was significantly higher (14,716.5 g wet weight m^-2) than in old (405.3 g w. wt. m^-2) or in young scour marks (9.2 g w. wt. m^-2). The habitat taxon richness, too, was highest in undisturbed areas (on average, 11.8 taxonomic units occurred per core), decreased in old scour marks (9.0) and was lowest in young scours (6.8). In undisturbed areas, a higher developed community structure was reflected by a greater variety of taxonomic groups, some of which were principally absent in scours.In young scours, the number of taxa was significantly reduced. Motile forms such as echinoderms and crustaceans, mainly amphipods, and juvenile polychaetes, in particular pioneering species of the family Spionidae, started the recolonization of the devastated areas. Burrowing organisms occurred in older scours where the initially overcompacted sediment had softened. In the course of the re-establishment of macrofaunal communities (after some years/decades), the faunal composition is expected to change towards a "normal" dominance of suspension-feeding organisms, mainly sponges and bryozoans, being typical for wide areas on the southeastern Weddell Sea shelf.A more detailed taxonomical approach, using 167 polychaete species as representatives of the macrozoobenthos, also revealed significant differences between undisturbed and affected areas. The mean abundances (784, 389, and 242 ind. m^-2, respectively), as well as habitat species richness (11.6, 5.5, and 3.1, respectively), decreased from undisturbed areas to old to young iceberg scours. Similarly, a large variety of motile and sessile forms was encountered among the polychaetes at undisturbed sites, whereas in scours, the polychaete fauna was impoverished in terms of species richness, abundance and variety of feeding types and lifestyles.     

Newly dominant benthic invertebrates reshape competitive networks on contemporary Caribbean reefs

Competition is a fundamental process structuring ecological communities. On coral reefs, space is a highly contested resource and the outcomes of spatial competition can dictate community composition. In the Caribbean, reefs are increasingly dominated by non-scleractinian species like sponges, gorgonians, and zoanthids, yet there is a paucity of data on interactions between these increasingly common organisms and historically dominant corals. Here, we investigated interactions among these groups of sessile benthic invertebrates to better understand the role of spatial competition in shaping benthic communities on Caribbean reefs. We coupled surveys of competitive interactions on the reef with a common garden competition experiment to determine the frequency and outcome of interference competition among eight focal species. We found that competitive interactions were pervasive on Florida reefs, with 60% of sessile benthic invertebrates interacting with at least one other invertebrate. Increasingly common non-scleractinian species were some of the most abundant taxa and consistently outcompeted the contemporarily common scleractinian species Porites porites and Siderastrea siderea. The encrusting gorgonian, Erythropodium caribaeorum, was the most aggressive species, reducing the live area of its competitors on average 42% ± 7.04 (SE) over the course of 5 months. Surprisingly, the most aggressive species declined in size when competing, while some less aggressive species were able to increase or maintain area, suggesting a trade-off between aggressiveness and growth. Our findings suggest that competition among sessile invertebrates is likely to remain an important process in structuring coral reefs, but that the optimal strategies for maintaining space on the benthos may change. Importantly, many non-scleractinian species that now dominate reefs appear to be superior competitors, potentially increasing the stress on corals on contemporary reefs.

     

Temporal vegetation dynamics and recolonization mechanisms on different-sized soil disturbances in tallgrass prairie

Assessing the various mechanisms by which plants revegetate disturbances is important for understanding the effects of disturbances on plant population dynamics, plant community structure, community assembly processes, and ecosystem function. We initiated a 2-yr experiment examining temporal vegetation dynamics and mechanisms of recolonization on different-sized soil disturbances created to simulate pocket gopher mounds in North American tallgrass prairie. Treatments were designed to assess potential contributions of the seed rain, soil seed bank, clonal propagation from the edges of a soil mound, and regrowth of buried plants. Small mounds were more rapidly recolonized than large mounds. Vegetative regrowth strategies were the dominant recolonization mechanisms, while the seed rain was considerably less important in maintaining the diversity of forbs and annuals than previously believed. All recolonization mechanisms influenced plant succession, but stem densities and plant mass on soil mounds remained significantly lower than undisturbed controls after two growing seasons. Because natural pocket gopher mounds are indistinguishable from undisturbed areas after two seasons, these results suggest that multiple modes of recruitment concurrently, albeit differentially, contribute to the recolonization of soil disturbances and influence tallgrass prairie plant community structure and successional dynamics.     

Acidification effects on biofouling communities: winners and losers

How ocean acidification affects marine life is a major concern for science and society. However, its impacts on encrusting biofouling communities, that are both the initial colonizers of hard substrata and of great economic importance, are almost unknown. We showed that community composition changed significantly, from 92% spirorbids, 3% ascidians and 4% sponges initially to 47% spirorbids, 23% ascidians and 29% sponges after 100 days in acidified conditions (pH 7.7). In low pH, numbers of the spirorbid Neodexiospira pseudocorrugata were reduced ×5 compared to controls. The two ascidians present behaved differently with Aplidium sp. decreasing ×10 in pH 7.7, whereas Molgula sp. numbers were ×4 higher in low pH than controls. Calcareous sponge (Leucosolenia sp.) numbers increased ×2.5 in pH 7.7 over controls. The diatom and filamentous algal community was also more poorly developed in the low pH treatments compared to controls. Colonization of new surfaces likewise showed large decreases in spirorbid numbers, but numbers of sponges and Molgula sp. increased. Spirorbid losses appeared due to both recruitment failure and loss of existing tubes. Spirorbid tubes are comprised of a loose prismatic fabric of calcite crystals. Loss of tube materials appeared due to changes in the binding matrix and not crystal dissolution, as SEM analyses showed crystal surfaces were not pitted or dissolved in low pH conditions. Biofouling communities face dramatic future changes with reductions in groups with hard exposed exoskeletons and domination by soft‐bodied ascidians and sponges.     

Vegetation succession after fire in sclerophyll woodland communities in south-eastern Australia

The dynamics of vegetation regeneration after burning were examined in three dry sclerophyll communities near Canberra, in south-eastern Australia. Changes in seedling and regrowth populations were followed in permanent quadrats during the first two years after burning, compared with both the preburn vegetation and population changes over the same period in adjacent, unburnt plots. All species represented either by living plants in the tree and shrub strata and/or by seed in the soil and litter prior to burning regenerated during the first year after the fire treatments. No new species invaded the areas after burning. Species varied in their regenerative strategy and recovered after the fires either by germination of seed residual in the soil and ash or released from trees after burning, by regrowth from surviving vegetative organs, or by a combination of germination and regrowth. Both seedling input and the vegetative recovery of populations were higher during the first than second year after burning. The vegetative multiplication and seed gertnination of many species were stimulated by fire. It was concluded that the regeneration of the communities studied, as well as the post-fire recovery of numerous different communities reported in the literature, closely resembled an initial floristic composition model. It is considered that the process of vegetation redevelopment after a disturbance (i.e. secondary succession) will be influenced greatly by the species composition at the time of disturbance, and by the type of disturbing agent. A single model would not be expected to adequately describe secondary succession following disturbance by agents imposing different stress conditions on a community.     

Temporal kinetics of fluoride accumulation: from fetal to adult deer

In June 2011, a volcano deposited about 100 million tons of tephra over parts of Chile and over 36 million ha of Argentina. Initially, fluoride was considered irrelevant; however, recently wild deer exhibited strong fluorosis, with fluoride level increasing 38-fold among severely affected deer. Whereas mothers averaged 2,151 ppm, their late-term fetuses had only 19.8 ppm, indicating a barrier to fluoride transport in utero. Levels among four age classes increased significantly, at a rate of about 1,000 ppm/year. The temporal kinetics of accumulation suggests that sources of available fluoride are highly effective. Thus, compared to prior background levels (63 ppm in adults) and to fetuses starting at about 20 ppm, 1-year-old calves averaged 1,035 ppm (maximum 1,830 ppm), 2-year olds averaged 2,151 ppm (maximum 2,513 ppm), and older deer averaged 2,806 ppm (maximum 5,175 ppm). As osteofluorosis occurs in deer with >4,000 ppm, accumulation of 1,000 ppm/year would result in adults reaching levels causing osteopathology in 1–2 years. Importantly, impacts may be further exacerbated by regional iodine and selenium deficiencies. Iodine deficiency may increase incidences of dental fluorosis and severity of damages, while selenium deficiency impacts iodine metabolism. Fluorosis will affect population dynamics, morbidity, predation susceptibility, and other ecosystem components like scavenger and plant communities.     

Effects of disturbance on plant regrowth along snow pack gradients in alpine habitats

Human disturbance in alpine habitats is expected to increase, and improved knowledge of short-term recovery after disturbance events is necessary to interpret vegetation responses and formulate planning and mitigation efforts. The ability of a plant community to return to its original state after a disturbance (community resilience) depends on species composition and environmental conditions. The aim of this study is to analyze initial short-term effects of disturbance in alpine plant communities in contrasting climates (oceanic vs. continental; central Norway). We used a nested block-design to examine vegetative regrowth and seedling recruitment after experimental perturbation. Three plant community types along the snow pack gradient were exposed to (1) no disturbance, (2) clipping, and (3) clipping and uprooting. Slow vegetative regrowth and low seedling establishment rates were found in dry alpine ridges and late-melting oceanic snowbed communities. Leeside habitats with intermediate snow conditions were found more resilient. The difference was related to growth form and species diversity. Woody species, which dominated in ridges and oceanic snowbeds, showed the most negative response to disturbance. Species-rich plant communities dominated by graminoids and herbs showed higher rates of regrowth. Species richness seems to cause resilience to the plant communities through higher response diversity. Plant communities at the extreme ends of abiotic gradients, ridges and late-melting snowbeds, will be most sensitive to both disturbance and environmental change. In an up-scaled human-used landscape disturbance effects will be amplified and further limit recovery to a pre-disturbance state.     

Long-term legacies and partial recovery of mycorrhizal communities after invasive plant removal

Invasive plants can have strong impacts on native communities, which have prompted intense efforts at invasive removal. However, relatively little is known about how native communities will reassemble after a dominant invader has been removed from the system. Legacy effects of invasive plants on soil microbial communities may alter native plant community reassembly long after the invader is gone. Here we found that arbuscular mycorrhizal fungal (AMF) communities have shown some recovery in experimental plots following 6 years of removal of the invasive Alliaria petiolata (garlic mustard, a species known to degrade AMF communities) in terms of taxonomic richness and community composition. However, despite this recovery, the density of A. petiolata at the beginning of the experiment (in 2004) still correlated with lower AMF richness and altered community composition after 6 years of annual weeding, suggesting long-term legacies of dense A. petiolata infestations. Because native plant and mycorrhizal fungal communities may show interdependence, reassembly of one community may be limited by the reassembly of the other. Restoration may be more effective if practices address both communities simultaneously.     

Biodiversity of <Emphasis Type="Italic">Spongosorites coralliophaga</Emphasis> (Stephens, 1915) on coral rubble at two contrasting cold-water coral reef settings

Cold-water coral reefs (CWRs) in the northeast Atlantic harbor diverse sponge communities. Knowledge of deep-sea sponge ecology is limited and this leaves us with a fragmented understanding of the ecological roles that sponges play in CWR ecosystems. We present the first study of faunal biodiversity associated with the massive demosponge Spongosorites coralliophaga (Stephens, 1915) that typically colonizes coral debris fields of CWRs. Our study focused on the sessile fauna inhabiting sponges mixed with coral rubble at two contrasting settings in the northeast Atlantic: the shallow inshore (120–190 m water depth) Mingulay Reef Complex (MRC) and the deep offshore (500–1200 m) Logachev Mound (LM) coral province. MRC is dominated by the scleractinian Lophelia pertusa, while LM is dominated by L. pertusa and Madrepora oculata. Nine sponge–coral rubble associations were collected from MRC and four from LM. Measurements of abundance, species richness, diversity, evenness, dry biomass, and composition of sessile fauna on sponge and coral rubble microhabitats were undertaken. Differences in community composition between the two regions were mainly a response to changes in fauna with depth. Fauna composition was also different between sponge and coral rubble within each region. Infauna constituted a minor component of the sponge-associated fauna in MRC but had a higher contribution in LM. Sponge and coral rubble sessile fauna in both regions was mainly composed of cnidarians and molluscs, similarly to some previous studies. Sponges’ outer surfaces at MRC were colonized by a species-rich community with high abundance and biomass suggesting that S. coralliophaga at MRC acts as a settlement surface for various organisms but such a role is not the case at LM. This difference in the role of S. coralliophaga as a biological structure is probably related to differences in fauna composition with depth, bottom current speed, and the quantity/quality of food supplied to the benthos.     

Contrasting rates of coral recovery and reassembly in coral communities on the Great Barrier Reef

Changes in the relative abundances of coral taxa during recovery from disturbance may cause shifts in essential ecological processes on coral reefs. Coral cover can return to pre-disturbance levels (coral recovery) without the assemblage returning to its previous composition (i.e., without reassembly). The processes underlying such changes are not well understood due to a scarcity of long-term studies with sufficient taxonomic resolution. We assessed the trajectories and time frames for coral recovery and reassembly of coral communities following disturbances, using modeled trajectories based on data from a broad spatial and temporal monitoring program. We studied coral communities at six reefs that suffered substantial coral loss and subsequently regained at least 50 % of their pre-disturbance coral cover. Five of the six communities regained their coral cover and the rates were remarkably consistent, taking 7–10 years. Four of the six communities reassembled to their pre-disturbance composition in 8–13 years. The coral communities at three of the reefs both regained coral cover and reassembled ten years. The trajectories of two communities suggested that they were unlikely to reassemble and the remaining community did not regain pre-disturbance coral cover. The communities that regained coral cover and reassembled had high relative abundance of tabulate Acropora spp. Coral communities of this composition appear likely to persist in a regime of pulse disturbances at intervals of ten years or more. Communities that failed to either regain coral cover or reassemble were in near-shore locations and had high relative abundance of Porites spp. and soft corals. Under current disturbance regimes, these communities are unlikely to re-establish their pre-disturbance community composition.     

Hypothesized Kinetic Models for Describing the Growth of Globular and Encrusting Demosponges

The marine sponges Dysidea avara and Chondrosia reniformis (globular forms) were cultured in the laboratory on a diet of viable Phaeodactylum tricornutum cells and dissolved nutrients (algae and fish powders). Our growth data were combined with literature data for Pseudosuberites andrewsi (a globular sponge) and for the encrusting sponges Oscarella lobularis, Hemimycale columella, and Crambe crambe. The suitability of three growth models—linear, exponential, and radial accretive—for describing the growth of globular and encrusting sponges was assessed. Radial accretive growth was determined to be the best model to describe growth of both encrusting and globular sponges. Average growth rates of 0.051 ± 0.016 and 0.019 ± 0.003 mm/day (calculated as the increase of the radius of the sponge per day) were obtained experimentally for D. avara and C. reniformis, respectively.     

Disturbance, colonization and development of Antarctic benthic communities

A decade has yielded much progress in understanding polar disturbance and community recovery-mainly through quantifying ice scour rates, other disturbance levels, larval abundance and diversity, colonization rates and response of benthos to predicted climate change. The continental shelf around Antarctica is clearly subject to massive disturbance, but remarkably across so many scales. In summer, millions of icebergs from sizes smaller than cars to larger than countries ground out and gouge the sea floor and crush the benthic communities there, while the highest wind speeds create the highest waves to pound the coast. In winter, the calm associated with the sea surface freezing creates the clearest marine water in the world. But in winter, an ice foot encases coastal life and anchor ice rips benthos from the sea floor. Over tens and hundreds of thousands of years, glaciations have done the same on continental scales-ice sheets have bulldozed the seabed and the zoobenthos to edge of shelves. We detail and rank modern disturbance levels (from most to least): ice; asteroid impacts; sediment instability; wind/wave action; pollution; UV irradiation; volcanism; trawling; non-indigenous species; freshwater inundation; and temperature stress. Benthic organisms have had to recolonize local scourings and continental shelves repeatedly, yet a decade of studies have demonstrated that they have (compared with lower latitudes) slow tempos of reproduction, colonization and growth. Despite massive disturbance levels and slow recolonization potential, the Antarctic shelf has a much richer fauna than would be expected for its area. Now, West Antarctica is among the fastest warming regions and its organisms face new rapid changes. In the next century, temperature stress and non-indigenous species will drastically rise to become dominant disturbances to the Antarctic life. Here, we describe the potential for benthic organisms to respond to disturbance, focusing particularly on what we know now that we did not a decade ago.