The successional dynamics of lightning-initiated canopy gaps in the mangrove forests of Shark River, Everglades National Park, USA

Gap succession is a significant determinant of structure and development in most forest communities. Lightning strikes are an important source of canopy gaps in the mangrove forest of Everglades National Park. I investigated the successional dynamics of lightning-initiated canopy gaps to determine their influence on forest stand structure of the mixed mangrove forests ( Rhizophora mangle, Laguncularia racemosa, and Avicennia germinans ) of the Shark River. I measured gap size, gap shape, light environment, soil characteristics, woody debris, and fiddler crab abundance. I additionally measured the vegetative composition in a chronosequences of gap successional stages (new, recruiting, and growing gaps). I recorded survivorship, recruitment, growth and soil elevation dynamics within a subset of new and growing gaps. I determined the relationship between intact forest soil elevation and site hydrology in order to interpret the effects of lightning disturbance on soil elevation dynamics. ^ Gap size averaged 289 ± 20 m2 (± 1SE) and light transmittance decreased exponentially as gaps filled with saplings. Fine woody debris was highest in recruiting gaps. Soil strength was lower in the gaps than in the forest. The abundance of large and medium fiddler crab burrows increased linearly with total seedling abundance. Soil surface elevation declined in newly formed lightning gaps; this loss was due to a combination of superficial erosion (8.5 mm) and subsidence (60.9 mm). A distinct two-cohort recruitment pattern was evident in the seedling/sapling surveys, suggesting a partitioning of the succession between individuals present before and after lightning strike. In new gaps, the seedling recruitment rate was twice as high as in forest and the sapling population increased. At the growing gap stage, R. mangle seedling mortality was 10 times greater and sapling mortality was 13 times greater than recruitment. Growing gaps had reduced seedling stem elongation, sapling growth and adult growth. However, a few individuals (R. mangle saplings) were able to recruit into the adult life stage. In conclusion, the high density of R. mangle seedlings and saplings imply that lightning strike disturbances in these mangrove forests favor their recruitment over that of A. germinans and L. racemosa. ^

Sedimentology of cores from the Iberian margin

Bioturbation in marine sediments has basically two aspects of interest for palaeo-environmental studies. First, the traces left by the burrowing organisms reflect the prevailing environmental conditions at the seafloor and thus can be used to reconstruct the ecologic and palaeoceanographic situation. Traces have the advantage over other proxies of practically always being preserved in situ. Secondly, for high- resolution stratigraphy, bioturbation is a nuisance due to the stirring and mixing processes that destroy the stratigraphic record. In order to evaluate the applicability of biogenic traces as palaeoenvironmental indicators, a number of gravity cores from the Portuguese continental slope, covering the period from the last glacial to the present were investigated through X-ray radiographs. In addition, physical and chemical parameters were determined to define the environmental niche in each core interval. A number of traces could be recognized, the most important being: Thalassinoides, Planolites, Zoophycos, Chondrites, Scolicia, Palaeophycus, Phycosiphon and the generally pyritized traces Trichichnus and Mycellia. The shifts between the different ichnofabrics agree strikingly well with the variations in ocean circulation caused by the changing climate. On the upper and middle slope, variations in current intensity and oxygenation of the Mediterranean Outflow Water were responsible for shifts in the ichnofabric. Larger traces such as Planolites and Thalassinoides dominated in coarse, well oxygenated intervals, while small traces such as Chondrites and Trichichnus dominated in fine grained, poorly oxygenated intervals. In contrast, on the lower slope where calm steady sedimentation conditions prevail, changes in sedimentation rate and nutrient flux have controlled variations in the distribution of larger traces such as Planolites, Thalassinoides, and Palaeophycus. Additionally, distinct layers of abundant Chondrites correspond to Heinrich events 1, 2, and 4, and are interpreted as a response to incursions of nutrient rich, oxygen depleted Antarctic waters during phases of reduced thermohaline circulation. The results clearly show that not one single factor but a combination of several factors is necessary to explain the changes in ichnofabric. Furthermore, large variations in the extent and type of bioturbation and tiering between different settings clearly show that a more detailed knowledge of the factors governing bioturbation is necessary if we shall fully comprehend how proxy records are disturbed. A first attempt to automatize a part of the recognition and quantification of the ichnofabric was performed using the DIAna image analysis program on digitized X-ray radiographs. The results show that enhanced abundance of pyritized microburrows appears to be coupled to organic rich sediments deposited under dysoxic conditions. Coarse grained sediments inhibit the formation of pyritized burrows. However, the smallest changes in program settings controlling the grey scale threshold and the sensitivity resulted in large shifts in the number of detected burrows. Therefore, this method can only be considered to be semi-quantitative. Through AMS-^C dating of sample pairs from the Zoophycos spreiten and the surrounding host sediment, age reversals of up to 3,320 years could be demonstrated for the first time. The spreiten material is always several thousands of years younger than the surrounding host sediment. Together with detailed X-ray radiograph studies this shows that the trace maker collects the material on the seafloor, and then transports it downwards up to more than one meter in to the underlying sediment where it is deposited in distinct structures termed spreiten. This clearly shows that age reversals of several thousands of years can be expected whenever Zoophycos is unknowingly sampled. These results also render the hitherto proposed ethological models proposed for Zoophycos as largely implausible. Therefore, a combination of detritus feeding, short time caching, and hibernation possibly combined also with gardening, is suggested here as an explanation for this complicated burrow.

Ichnology and palaeobiology of Phoebichnus trochoides and Schaubcylindrichnus heberti comb. nov.

The three-dimensional reconstructions of Phoebichnus trochoides and Schaubcylindrichnus (Palaeophycus) heberti created as part of this thesis allow us to fully understand and characterize the three-dimensional morphology and palaeobiology of these common taxa. Three-dimensional reconstructions demonstrate that P. trochoides is a large stellate burrow composed of numerous long galleries produced by a deposit feeding organism. This study reports for the first time that the central zone is composed of stacked disk-shaped layers of highly bioturbated sediment, the radial burrows are composed of a sand-rich lining of pelleted annuli surrounding an active sand-rich fill, and the presence of subtle conical features above the radial galleries that are inferred to result from collapse cone feeding. Reconstructions of heberti demonstrate that the thick walled burrows are composed of sand-rich annular rings, are a broad U-shape, and may be either clustered or isolated. Our observations show that the morphology of heberti is inconsistent with the generic diagnosis of Palaeophycus, but is morphologically comparable to Schaubcylindrichnus, and is herein synonymised with Schaubcylindrichnus to create S. heberti comb. nov. The three-dimensional reconstructions have revealed a number of hitherto unknown morphological elements to both taxa which has facilitated new interpretations of the trace-makers behaviour. The data improves the taxonomic understanding of both P. trochoides and S. heberti which require significant taxonomic change and emendation of diagnoses at the species and genus level.

Three-dimensional tomographic reconstruction of an exceptionally well preserved ichnological assemblage from the Stainmore Foundation, Carboniferous, UK

This Ph.D. thesis addresses current issues with ichnotaxonomic practice, and characterizes an exceptionally well preserved ichnological assemblage from the Carboniferous Stainmore Formation, Northumberland, United Kingdom. Samples were collected from closely localized float representative of various units throughout the succession, which was deposited in a storm-dominated marine shoreface. Three dominant ichnotaxa were selected for three-dimensional morphological analysis due to their complicated morphology and/or unclear taxonomic status: 1) Dactyloidites jordii isp. nov.; 2) Beaconites capronus, and; 3) Neoeione moniliformis comb. nov. Using serial grinding and photography, these ichnotaxa were ground and modelled in true colour. High-resolution models of three taxa produced in this study are the basis of the first complete three-dimensional consideration of the traces, and forms the basis for refined palaeobiological and ethological analysis of these taxa. Dactyloidites jordii isp. nov. is a stellate to palmate burrow composed of numerous long, narrow rays that exhibit three orders of branching arranged into tiered galleries radiating from a central shaft. It is considered to be the feeding structure produced by a vermiform organism. Beaconites capronus is a winding trace with distinctly chevron-shaped, meniscate backfill demonstrated herein to backfill the vertical shafts associated with its burrows in a comparable fashion to the horizontal portion of the burrow. This lack of a surface connection would result in the trace making organism being exposed to low-oxygen porewater. Coping with this porewater dysoxia could be approached by burrowing organisms in a number of ways: 1) revisiting the sediment-water interface; 2) creating periodic shafts; or 3) employing anaerobic metabolism. Neoeione moniliformis was originally introduced as Eione moniliformis, however, the genus Eione Tate, 1859 is a junior homonym of Eione Rafinesque, 1814. This led to the transfer of Eione moniliformis to Parataenidium. Through careful examination and three-dimensional characterization of topotypes, the transfer to Parataenidium moniliformis is demonstrated herein to be problematic, as Parataenidium refers to primarily horizontal burrows with two distinct layers and Eione moniliformis is composed of one distinct level. As such, the new ichnogenus Neoeione is created to accommodate Neoeione moniliformis.

Georeferenced standardized apparent wave power along the coastline of Cabrera National Park for analysis of essential habitats of littoral species

Wind- induced exposure is one of the major forces shaping the geomorphology and biota in coastal areas. The effect of wave exposure on littoral biota is well known in marine environments (Ekebon et al., 2003; Burrows et al., 2008). In the Cabrera Archipelago National Park wave exposure has demostrated to have an effect on the spatial distribution of different stages of E.marginatus (Alvarez et al., 2010). Standarized average wave exposures during 2008 along the Cabrera Archipelago National park coast line were calculated to be applied in studies of littoral species distribution within the archipelago. Average wave exposure (or apparent wave power) was calculated for points located 50 m equidistant on the coastline following the EXA methodology (EXposure estimates for fragmented Archipelagos) (Ekebon et al., 2003). The average wave exposures were standardized from 1 to 100 (minimum and maximum in the area), showing coastal areas with different levels of mea wave exposure during the year. Input wind data (direction and intensity) from 2008 was registered at the Cabrera mooring located north of Cabrera Archipelago. Data were provided by IMEDEA (CSIC-UIB, TMMOS http://www.imedea.uib-csic.es/tmoos/boyas/). This cartography has been developed under the framework of the project EPIMHAR, funded by the National Park's Network (Spanish Ministry of Environment, Maritime and Rural Affairs, reference: 012/2007 ). Part of this work has been developed under the research programs funded by "Fons de Garantia Agrària i Pesquera de les Illes Balears (FOGAIBA)".

Geochemical analysis of two sediment cores from the southern Mendeleev Ridge

We present results of an inorganic geochemical pore water and sediment study conducted on Quaternary sediments from the western Arctic Ocean. The sediment cores were recovered in 2008 from the southern Mendeleev Ridge during RV Polarstern Expedition ARK-XXIII/3. With respect to sediment sources and depositional processes, peaks in Ca/Al, Mg/Al, Sr/Al and Sr/Mg indicate enhanced input of both ice-rafted (mainly dolomite) and biogenic carbonate during deglacial warming phases. Distinct and repetitive brown layers enriched in Mn (oxyhydr)oxides occur mostly in association with these carbonate-rich intervals. For the first time, we show that the brown layers are also consistently enriched in scavenged trace metals Co, Cu, Mo and Ni. The bioturbation patterns of the brown layers, specifically well-defined brown burrows into the underlying sediments, support formation close to the sediment-water interface. The Mn and trace metal enrichments were probably initiated under warmer climate conditions. Both river runoff and melting sea ice delivered trace metals to the Arctic Ocean, but also enhanced seasonal productivity and organic matter export to the sea floor. As Mn (oxyhydr)oxides and scavenged trace metals were deposited at the sea floor, a co-occurring organic matter "pulse" triggered intense diagenetic Mn cycling at the sediment-water interface. These processes resulted in the formation of Mn and trace metal enrichments, but almost complete organic matter degradation. As warmer conditions ceased, reduced riverine runoff and/or a solid sea ice cover terminated the input of riverine trace metal and fresh organic matter, and greyish-yellowish sediments poor in Mn and trace metals were deposited. Oxygen depletion of Arctic bottom waters as potential cause for the lack of Mn enrichments during glacial intervals is highly improbable. While the original composition and texture of the brown layers resulted from specific climatic conditions (including transient Mn redox cycling at the sediment-water interface), pore water data show that early diagenetic Mn redistribution is still affecting the organic-poor sediments in several meters depth. Given persistent steady state diagenetic conditions, purely authigenic Mn-rich brown layers may form, while others may completely vanish. The degree of diagenetic Mn redistribution largely depends on the depositional environment within the Arctic Ocean, the availability of Mn and organic matter, and seems to be recorded by the Co/Mo ratios of single Mn-rich layers. We conclude that brown Arctic sediment layers are not necessarily synchronous features, and correlating them across different parts of the Arctic Ocean without additional age control is not recommended.

Some Þing To Talk About : Assemblies in the Íslendingasögur

Date of Acceptance: 16/06/2015

Some Þing To Talk About : Assemblies in the Íslendingasögur

Date of Acceptance: 16/06/2015

(Table1) Frequency of well defined bioturbation in sediments of the Eckernförder Bay, Western Baltic Sea

The lamination and burrowing patterns in 17 box cores were analyzed with the aid of X-ray photographs and thin sections. A standardized method of log plotting made statistical analysis of the data possible. Several 'structure types' were established, although it was realized that the boundaries are purely arbitrary divisions in what can sometimes be a continuous sequence. In the transition zone between marginal sand facies and fine-grained basin facies, muddy sediment is found which contains particularly well differentiated, alternating laminae. This zone is also characterized by layers rich in plant remains. The alternation of laminae shows a high degree of statistical scattering. Even though a small degree of cyclic periodicity could be defined, it was impossible to correlate individual layers from core to core across the bay. However, through a statistical handling of the plots, zones could be separated on the basis of the number of sand layers they contained. These more or minder sandy zones clarified the bottom reflections seen in the records of the echograph from the area. The manner of facies change across the bay, suggests that no strong bottom currents are effective in the Eckernförde Bay. The marked asymmetry between the north and south flanks of the profile can be attributed to the stronger action of waves on the more exposed areas. Grain size analyses were made from the more homogeneous units found in a core from the transition-facies zone. The results indicate that the most pronounced differences between layers appear in the silt range, and although the differences are slight, they are statistically significant. Layers rich in plant remains were wet-sieved in order to separate the plant detritus. This was than analyzed in a sediment settling balance and found to be hydrodynamically equivalent to a well-sorted, finegrained sand. A special, rhythmic cross-bedding type with dimensions in the millimeter range, has been named 'Crypto-cross-lamination' and is thought to represent rapid sedimentation in an area where only very weak bottom currents are present. It is found only in the deepest part of the basin. Relatively large sand grains, scattered within layers of clayey-silty matrix, seem to be transported by flotation. Thin section examination showed that the inner part of Eckernförder Bay carbonate grains (e. g. Foraminifera shells) were preserved throughout the cores, while in the outer part of the bay they were not present. Well defined tracks and burrows are relatively rare in all of the facies in comparision to the generally strongly developed deformation burrowing. The application of special measures for the deformation burrowing allowed to plot their intensity in profile for each core. A degree of regularity could be found in these burrowing intensity plots, with higher values appearing in the sandy facies, but with no clear differences between sand and silt layers in the transition facies. Small sections in the profiles of the deepest part of the bay show no bioturbation at all.

(Table T1) Lithologic and ichnologic description of ODP Hole 174-1073A

When examined in their sedimentologic and stratigraphic context, ichnofabrics and component ichnofossils can help decipher paleoceanography and sea-level histories from marine deposits (Savrda, 1995, http://www.jstor.org/stable/3515095). Thus far, applications of ichnology in paleoceanographic investigations have been restricted to slowly deposited, predominantly biogenic sediments and/or strata deposited in oxygen-deficient, tectonically active basins. Moreover, ichnologic applications in sequence stratigraphic studies largely have been restricted to strata deposited in relatively shallow-water shelf or foreland basin settings. The limits of previous studies provided impetus for detailed postcruise examination of Quaternary deposits recovered at Ocean Drilling Program (ODP) Site 1073 on the New Jersey margin. These deposits provide the opportunity to assess the sedimentary and ichnofabric record of glacio-eustatic cycles in a passive continental slope setting characterized by relatively rapid accumulation of siliciclastic sediments in an area not far removed from the Laurentide ice margin. The primary purpose of this data report is to present basic sedimentologic and ichnologic observations made at the decimeter scale throughout the Quaternary sequence from Site 1073. Data analysis and interpretation in the context of climate and sea-level histories, as inferred from oxygen isotopic, palynologic, and seismic studies, are ongoing and will be presented in subsequent papers prepared for open literature (e.g., Savrda et al., in press).

A hierarchical approach to defining marine heatwaves

Marine heatwaves (MHWs) have been observed around the world and are expected to increase in intensity and frequency under anthropogenic climate change. A variety of impacts have been associated with these anomalous events, including shifts in species ranges, local extinctions and economic impacts on seafood industries through declines in important fishery species and impacts on aquaculture. Extreme temperatures are increasingly seen as important influences on biological systems, yet a consistent definition of MHWs does not exist. A clear definition will facilitate retrospective comparisons between MHWs, enabling the synthesis and a mechanistic understanding of the role of MHWs in marine ecosystems. Building on research into atmospheric heatwaves, we propose both a general and specific definition for MHWs, based on a hierarchy of metrics that allow for different data sets to be used in identifying MHWs. We generally define a MHW as a prolonged discrete anomalously warm water event that can be described by its duration, intensity, rate of evolution, and spatial extent. Specifically, we consider an anomalously warm event to be a MHW if it lasts for five or more days, with temperatures warmer than the 90th percentile based on a 30-year historical baseline period. This structure provides flexibility with regard to the description of MHWs and transparency in communicating MHWs to a general audience. The use of these metrics is illustrated for three 21st century MHWs; the northern Mediterranean event in 2003, the Western Australia ‘Ningaloo Niño’ in 2011, and the northwest Atlantic event in 2012. We recommend a specific quantitative definition for MHWs to facilitate global comparisons and to advance our understanding of these phenomena.

A hierarchical approach to defining marine heatwaves

Marine heatwaves (MHWs) have been observed around the world and are expected to increase in intensity and frequency under anthropogenic climate change. A variety of impacts have been associated with these anomalous events, including shifts in species ranges, local extinctions and economic impacts on seafood industries through declines in important fishery species and impacts on aquaculture. Extreme temperatures are increasingly seen as important influences on biological systems, yet a consistent definition of MHWs does not exist. A clear definition will facilitate retrospective comparisons between MHWs, enabling the synthesis and a mechanistic understanding of the role of MHWs in marine ecosystems. Building on research into atmospheric heatwaves, we propose both a general and specific definition for MHWs, based on a hierarchy of metrics that allow for different data sets to be used in identifying MHWs. We generally define a MHW as a prolonged discrete anomalously warm water event that can be described by its duration, intensity, rate of evolution, and spatial extent. Specifically, we consider an anomalously warm event to be a MHW if it lasts for five or more days, with temperatures warmer than the 90th percentile based on a 30-year historical baseline period. This structure provides flexibility with regard to the description of MHWs and transparency in communicating MHWs to a general audience. The use of these metrics is illustrated for three 21st century MHWs; the northern Mediterranean event in 2003, the Western Australia ‘Ningaloo Niño’ in 2011, and the northwest Atlantic event in 2012. We recommend a specific quantitative definition for MHWs to facilitate global comparisons and to advance our understanding of these phenomena.

Historical comparisons reveal multiple drivers of decadal change of an ecosystem engineer at the range edge

Biogenic reefs are important for habitat provision and coastal protection. Long-term datasets on the distribution and abundance of Sabellaria alveolata (L.) are available from Britain. The aim of this study was to combine historical records and contemporary data to (1) describe spatiotemporal variation in winter temperatures, (2) document short-term and long-term changes in the distribution and abundance of S. alveolata and discuss these changes in relation to extreme weather events and recent warming, and (3) assess the potential for artificial coastal defense structures to function as habitat for S. alveolata. A semi-quantitative abundance scale (ACFOR) was used to compare broadscale, long-term and interannual abundance of S. alveolata near its range edge in NW Britain. S. alveolata disappeared from the North Wales and Wirral coastlines where it had been abundant prior to the cold winter of 1962/1963. Population declines were also observed following the recent cold winters of 2009/2010 and 2010/2011. Extensive surveys in 2004 and 2012 revealed that S. alveolata had recolonized locations from which it had previously disappeared. Furthermore, it had increased in abundance at many locations, possibly in response to recent warming. S. alveolata was recorded on the majority of artificial coastal defense structures surveyed, suggesting that the proliferation of artificial coastal defense structures along this stretch of coastline may have enabled S. alveolata to spread across stretches of unsuitable natural habitat. Long-term and broadscale contextual monitoring is essential for monitoring responses of organisms to climate change. Historical data and gray literature can be invaluable sources of information. Our results support the theory that Lusitanian species are responding positively to climate warming but also that short-term extreme weather events can have potentially devastating widespread and lasting effects on organisms. Furthermore, the proliferation of coastal defense structures has implications for phylogeography, population genetics, and connectivity of coastal populations.

Historical comparisons reveal multiple drivers of decadal change of an ecosystem engineer at the range edge

Biogenic reefs are important for habitat provision and coastal protection. Long-term datasets on the distribution and abundance of Sabellaria alveolata (L.) are available from Britain. The aim of this study was to combine historical records and contemporary data to (1) describe spatiotemporal variation in winter temperatures, (2) document short-term and long-term changes in the distribution and abundance of S. alveolata and discuss these changes in relation to extreme weather events and recent warming, and (3) assess the potential for artificial coastal defense structures to function as habitat for S. alveolata. A semi-quantitative abundance scale (ACFOR) was used to compare broadscale, long-term and interannual abundance of S. alveolata near its range edge in NW Britain. S. alveolata disappeared from the North Wales and Wirral coastlines where it had been abundant prior to the cold winter of 1962/1963. Population declines were also observed following the recent cold winters of 2009/2010 and 2010/2011. Extensive surveys in 2004 and 2012 revealed that S. alveolata had recolonized locations from which it had previously disappeared. Furthermore, it had increased in abundance at many locations, possibly in response to recent warming. S. alveolata was recorded on the majority of artificial coastal defense structures surveyed, suggesting that the proliferation of artificial coastal defense structures along this stretch of coastline may have enabled S. alveolata to spread across stretches of unsuitable natural habitat. Long-term and broadscale contextual monitoring is essential for monitoring responses of organisms to climate change. Historical data and gray literature can be invaluable sources of information. Our results support the theory that Lusitanian species are responding positively to climate warming but also that short-term extreme weather events can have potentially devastating widespread and lasting effects on organisms. Furthermore, the proliferation of coastal defense structures has implications for phylogeography, population genetics, and connectivity of coastal populations.

Facing the future: the importance of substratum features for ecological engineering of artificial habitats in the rocky intertidal

Coastal defences are proliferating in response to climate change, leading to the creation of more vertical substrata. Efforts are being made to mitigate their impacts and create novel habitats to promote biodiversity. Little is known about the effect of aspect (i.e. north–south directionality) and inclination on intertidal biodiversity in artificial habitats. Artificial and natural habitats were compared to assess the role of aspect and substratum inclination in determining patterns of biodiversity at two tidal heights (high and mid). We also compared grazing activity between north- and south-facing surfaces in natural habitats to examine the potential for differential grazing pressure to affect community structure and functioning. Results were variable but some clear patterns emerged. Inclination had no effect on biodiversity or abundance. There was a general trend towards greater taxon richness and abundance on north-facing than south-facing substrata in natural and artificial habitats. On natural shores, the abundance and grazing activity of ‘southern’ limpets (i.e. Patella depressa) was greater on south-facing than north-facing substrata, with possible implications for further range-expansion. These results highlight the importance of incorporating shaded habitats in the construction of artificial habitats. These habitats may represent an important refuge from grazing pressure and thermal and desiccation stress in a warming climate.