ENCORE: The effect of nutrient enrichment on coral reefs. Synthesis of results and conclusions

Coral reef degradation resulting from nutrient enrichment of coastal waters is of increasing global concern. Although effects of nutrients on coral reef organisms have been demonstrated in the laboratory, there is little direct evidence of nutrient effects on coral reef biota in situ. The ENCORE experiment investigated responses of coral reef organisms and processes to controlled additions of dissolved inorganic nitrogen (N) and/or phosphorus (P) on an offshore reef(One Tree Island) at the southern end of the Great Barrier Reef, Australia. A multi-disciplinary team assessed a variety of factors focusing on nutrient dynamics and biotic responses. A controlled and replicated experiment was conducted over two years using twelve small patch reefs ponded at low tide by a coral rim. Treatments included three control reefs (no nutrient addition) and three + N reefs (NH4Cl added), three + P reefs (KH2PO4 added), and three + N + P reefs. Nutrients were added as pulses at each low tide (ca twice per day) by remotely operated units. There were two phases of nutrient additions. During the initial, low-loading phase of the experiment nutrient pulses (mean dose = 11.5 muM NH4+; 2.3 muM PO4-3) rapidly declined, reaching near-background levels (mean = 0.9 muM NH4+; 0.5 muM PO4-3) within 2-3 h. A variety of biotic processes, assessed over a year during this initial nutrient loading phase, were not significantly affected, with the exception of coral reproduction, which was affected in all nutrient treatments. In Acropora longicyathus and A. aspera, fewer successfully developed embryos were formed, and in A. longicyathus fertilization rates and lipid levels decreased. In the second, high-loading, phase of ENCORE an increased nutrient dosage (mean dose = 36.2 muM NH4+; 5.1 muM PO4-3 declining to means of 11.3 muM NH4+ and 2.4 muM PO4-3 at the end of low tide) was used for a further year, and a variety of significant biotic responses occurred. Encrusting algae incorporated virtually none of the added nutrients. Organisms containing endosymbiotic zooxanthellae (corals and giant clams) assimilated dissolved nutrients rapidly and were responsive to added nutrients. Coral mortality, not detected during the initial low-loading phase, became evident with increased nutrient dosage, particularly in Pocillopora damicornis. Nitrogen additions stunted coral growth, and phosphorus additions had a variable effect. Coral calcification rate and linear extension increased in the presence of added phosphorus but skeletal density was reduced, making corals more susceptible to breakage. Settlement of all coral larvae was reduced in nitrogen treatments, yet settlement of larvae from brooded species was enhanced in phosphorus treatments. Recruitment of stomatopods, benthic crustaceans living in coral rubble, was reduced in nitrogen and nitrogen plus phosphorus treatments. Grazing rates and reproductive effort of various fish species were not affected by the nutrient treatments. Microbial nitrogen transformations in sediments,were responsive to nutrient loading with nitrogen fixation significantly increased in phosphorus treatments and denitrification increased in all treatments to which nitrogen had been added. Rates of bioerosion and grazing showed no significant effects of added nutrients, ENCORE has shown that reef organisms and processes investigated ill situ were impacted by elevated nutrients. Impacts mere dependent on dose level, whether nitrogen and/or phosphorus mere elevated and were often species-specific. The impacts were generally sub-lethal and subtle and the treated reefs at the end of the experiment mere visually similar to control reefs. Rapid nutrient uptake indicates that nutrient concentrations alone are not adequate to assess nutrient condition of reefs. Sensitive and quantifiable biological indicators need to be developed for coral reef ecosystems. The potential bioindicators identified in ENCORE should be tested in future research on coral reef/nutrient interactions. Synergistic and cumulative effects of elevated nutrients and other environmental parameters, comparative studies of intact vs. disturbed reefs, offshore vs, inshore reefs, or the ability of a nutrient-stressed reef to respond to natural disturbances require elucidation. An expanded understanding of coral reef responses to anthropogenic impacts is necessary, particularly regarding the subtle, sub-lethal effects detected in the ENCORE studies. (C) 2001 Published by Elsevier Science Ltd.

Diel Patterns of Patch Reef Community Dynamics in a Caribbean Back-Reef System

Back-reef seascapes represent critical habitat for juvenile and adult fishes. Patch reef, seagrass, and mangrove habitats form a heterogeneous mosaic, often linked by species that use reefs as structure during the day and make foraging migrations into soft-bottom habitat at night. Artificial reefs are used to model natural patch reefs, however may not function equivalently as fish habitat. To study the relative value of natural and artificial patch reefs as fish habitat, these communities in the Sea of Abaco, Bahamas were compared using roving diver surveys and time-lapse photography. Diel turnover in fish abundance, recorded with time-lapse photography and illuminated by infrared light, was quantified across midday, dusk, and night periods to explore possible effects of reef type (artificial vs. natural) on these patterns. Diurnal communities on natural reefs exhibited greater fish abundance, species richness, and functional diversity compared to artificial reefs. Furthermore, both types of reef communities exhibited a significant shift across the diel period, characterized by a decline in total fish density at night, especially for grunts (Haemulidae). Cross-habitat foraging migrations by diurnal or nocturnal species, such as haemulids, are likely central drivers of this twilight turnover and can represent important energy and nutrient subsidies. Time-lapse surveys provided more consistent measures of reef fish assemblages for the smaller artificial reef habitats, yet underestimated abundance of certain taxa and species richness on larger patch habitats when compared to the roving diver surveys. Time-lapse photography complemented with infrared light represent a valuable non-invasive approach to studying behavior of focal species and their fine-scale temporal dynamics in shallow-reef communities.

Nutrient-induced perturbations to delta C-13 and delta N-15 in symbiotic dinoilagellates and their coral hosts

Inorganic nutrients play a critical role in determining benthic community structure in tropical seas. This study examined the impact of adding inorganic nutrients (ammonium and phosphate) on the isotopic composition of 2 reef-building corals, Pocillopora damicornis and Heliofungia actiniformis, on the southern Great Barrier Reef. The addition of elevated nutrients to patch reefs that pond at low tide did not perturb the C:N ratio of either species or their symbiotic dinoflagellates. The C:N ratios were significantly higher in material extracted from the skeleton (14.8 +/- 1.50 and 10.8 +/- 1.42) than either host (7.6 +/- 0.87 and 6.0 +/- 0.71) or symbiotic dinoflagellates (5.7 +/- 0.48 and 6.9 +/- 0.66) (P. damicornis and H. actiniformis respectively; 95 confidence intervals). The ratio of acquired N to background N suggests that the added dissolved inorganic nitrogen (DIN) accounted for 50 to 100% of total nitrogen within the tissues of P. damicornis and H. actiniformis at the end of the experiment. The addition of the isotopically depleted nutrients (delta(15) N = 0parts per thousand) to patch reefs significantly decreased delta(15)N from control values of between 3 and 4 to values to below 1 in the case of all compartments, while delta(13)C values were relatively unresponsive to nutrient treatments. These findings suggest that coral delta(15)N has the potential to provide a historical record of the delta(15)N of dissolved nitrogen surrounding reef-building corals and their symbiotic dinoflagellates.

Cleaner fish Labroides dimidiatus reduce 'temporary" parasitic corallanid isopods on the coral reef fish Hemigymnus melapterus

To determine if cleaners affect 'temporary' parasitic corallanid isopods (Argathona macronema) on fish, we used caged fish Hemigymnus meldpterus (Labridae) on 5 patch reefs on Lizard Island, Great Barrier Reef, and removed all cleaner fish Labroides dimidiatus (Labridae) from 3 of the reefs, In a short-term experiment, fish were sampled after 12 or 24 h, at dawn and sunset respectively, and in a long-term experiment they were sampled after 12 d at sunset. Isopod prevalence, abundance and size were measured. In the short-term experiment, on reefs without cleaners the prevalence of A. macronema was higher after 24 h than after 12 h while on reefs with cleaners, prevalence was low at all times, Although the abundance of A, macronema did not vary after 12 and 24 h, when combined over the 24 h, the effect of cleaners was significant with only 2 % of all the A. macronema found on reefs with cleaners. Cleaners had no effect on the size frequency distribution of A. macronema in the short-term experiment, most likely because fish had so few isopods on reef with cleaners. In the longer-term experiment, the effects of cleaners on isopod prevalence and abundance were less clear. Their effect on isopod size was, however, significant with smaller parasites on reefs without cleaners. The reduction of isopod prevalence and abundance by cleaner fish over a period of hours may explain why these A, macronema are rare on wild fish. Our findings support the idea that cleaning is beneficial to clients and has important implications for the control of parasites of fish farmed in cages,

Evolution of the Pelagonian carbonate platform complex and the adjacent oceanic realm in response to plate tectonic forcing (Late Triassic and Jurassic), Evvoia, Greece

The Late Triassic and Jurassic platform and the oceanic complexes in Evvoia, Greece, share a complementary plate-tectonic evolution. Shallow marine carbonate deposition responded to changing rates of subsidence and uplift, whilst the adjacent ocean underwent spreading, and then convergence, collision and finally obduction over the platform complex. Late Triassic ocean spreading correlated with platform subsidence and the formation of a long-persisting peritidal passive-margin platform. Incipient drowning occurred from the Sinemurian to the late Middle Jurassic. This subsidence correlated with intra-oceanic subduction and plate convergence that led to supra-subduction calc-alkaline magmatism and the formation of a primitive volcanic arc. During the Middle Jurassic, plate collision caused arc uplift above the carbonate compensation depth (CCD) in the oceanic realm, and related thrust-faulting, on the platform, led to sub-aerial exposures. Patch-reefs developed there during the Late Oxfordian to Kimmeridgian. Advanced oceanic nappe-loading caused platform drowning below the CCD during the Tithonian, which is documented by intercalations of reefal turbidites with non-carbonate radiolarites. Radiolarites and bypass-turbidites, consisting of siliciclastic greywacke, terminate the platform succession beneath the emplaced oceanic nappe during late Tithonian to Valanginian time.

Non-random reef use by fishes at two dominant zones in a tropical, algal-dominated coastal reef

Habitat use and the processes which determine fish distribution were evaluated at the reef flat and reef crest zones of a tropical, algal-dominated reef. Our comparisons indicated significant differences in the majority of the evaluated environmental characteristics between zones. Also, significant differences in the abundances of twelve, from thirteen analyzed species, were observed within and between-sites. According to null models, non-random patterns of species co-occurrences were significant, suggesting that fish guilds in both zones were non-randomly structured. Unexpectedly, structural complexity negatively affected overall species richness, but had a major positive influence on highly site-attached species such as a damselfish. Depth and substrate composition, particularly macroalgae cover, were positive determinants for the fish assemblage structure in the studied reef, prevailing over factors such as structural complexity and live coral cover. Our results are conflicting with other studies carried out in coral-dominated reefs of the Caribbean and Pacific, therefore supporting the idea that the factors which may potentially influence reef fish composition are highly site-dependent and variable.

Mapeamento de recifes no Rio Grande do Norte: Touros e Macau/RN

Studies on the Brazilian continental shelf were developed between Touros and Macau-RN (NE Brazil) with the aim to mapping carbonate buildups, and especially, reef ecosystems. Remote sensing, submarine movies, bathymetric survey and diving were the methods used. This paper will focus on three main aspects of the Rio Grande do Norte reefs: 1) characterization of the coral fauna; 2) the classification and, 3) the distribution of the main northriograndense carbonate buildups. Reefs environments are found predominantly on the inner shelf adjacent to Touros, at depths shallower than 10 m. These reefs generally show NW-SE orientation parallel to the coastline and constitute groupings of knolls and patch reefs. A prominent carbonate buildup, where the corals are almost absents and algae incrustation is thin, occurs in average depths of 25 m, along the northern part of Rio Grande do Norte State and is classified as sandstone bank. Others sandstone banks, with smaller lengths and heights were mapped around 10 m depth. They should be related to an ancient coastline; however the corresponding litostratigraphic unit was not defined. This work is a contribution to the mapping of coral reef, in particular, and carbonate buildups, in general, on the Rio Grande do Norte coast

Caracterização Morfodinâmica e Sedimentologia da Plataforma Continental Rasa na APA Estadual dos Recifes de Corais - RN

This study had to aimed to characterize the sediments of shallow continental shelf and realize the mapping of features visible for satellite images by using remote sensing techniques, digital image processing and analysis of bathymetry between Maxaranguape and Touros - RN. The study s area is located in the continental shallow shelf of Rio Grande do Norte, Brazil, and is part of the Environmental Protection Area (APA) of Coral Reefs. A total of 1186 sediment samples were collected using a dredge type van veen and positioning of the vessel was made out with the aid of a Garmin 520s. The samples were treated In the laboratory to analyze particle size of the sediment, concentration of calcium carbonate and biogenic composition. The digital images from the Landsat-5 TM were used to mapping of features. This stage was used the band 1 (0,45-1,52 μm) where the image were georeferenced, and then adjusting the histogram, giving a better view of feature bottom and contacts between different types of bottom. The results obtained from analysis of the sediment showed that the sediments of the continental shelf east of RN have a dominance of carbonate facies and a sand-gravelly bottom because the region is dominated by biogenic sediments, that are made mainly of calcareous algae. The bedform types identified and morphological features found were validated by bathymetric data and sediment samples examined. From the results obtained a division for the shelf under study is suggested, these regions being subdivided, in well characterized: (1) Turbid Zone, (2) Coral Patch Reefs Zone, (3) Mixed Sediments Carbonates Zone, ( 4) Algae Fouling Zone, (5) Alignment Rocky Zone, (6) Sand Waves Field (7) Deposit siliciclastic sands

Masking environmental feedback : Misfits between institutions and ecosystems in Belize and Thailand

The thesis analyses relationships between ecological and social systems in the context of coastal ecosystems. It examines human impacts from resource extraction and addresses management and governance behind resource exploitation. The main premises are that a lack of ecological knowledge leads to poor ecosystem management and that the dichotomy between social and natural systems is an artificial one. The thesis illustrates the importance of basing resource management on the ecological conditions of the resource and its ecosystem. It also demonstrates the necessity of accounting for the human dimension in ecosystem management and the challenges of organising human actions for sustainable use of ecosystem services in the face of economic incentives that push users towards short-term extraction. Many Caribbean coral reefs have undergone a shift from coral to macroalgal domination. An experiment on Glovers Reef Atoll in Belize manually cleared patch reefs in a no-take zone and a fished zone (Papers I and II). The study hypothesised that overfishing has reduced herbivorous fish populations that control macroalgae growth. Overall, management had no significant effect on fish abundance and the impacts of the algal reduction were short-lived. This illustrated that the benefits of setting aside marine reserves in impacted environments should not be taken for granted. Papers III and IV studied the development of the lobster and conch fisheries in Belize, and the shrimp farming industry in Thailand respectively. These studies found that environmental feedback can be masked to give the impression of resource abundance through sequential exploitation. In both cases inadequate property rights contributed to this unsustainable resource use. The final paper (V) compared the responses to changes in the resource by the lobster fisheries in Belize and Maine in terms of institutions, organisations and their role in management. In contrast to Maine’s, the Belize system seems to lack social mechanisms for responding effectively to environmental feedback. The results illustrate the importance of organisational and institutional diversity that incorporate ecological knowledge, respond to ecosystem feedback and provide a social context for learning from and adapting to change.

Growth rates of late Miocene corals from Crete (Greece)

Modern scleractinian corals are classical components of marine shallow warm water ecosystems. Their occurrence and diversity patterns in the geological record have been widely used to infer past climates and environmental conditions. Coral skeletal composition data reflecting the nature of the coral environment are often affected by diagenetic alteration. Ghost structures of annual growth rhythms are, however, often well preserved in the transformed skeleton. We show that these relicts represent a valuable source of information on growth conditions of fossil corals. Annual growth bands were measured in massive hemispherical Porites of late Miocene age from the island of Crete (Greece) that were found in patch reefs and level bottom associations of attached mixed clastic environments as well as isolated carbonate environments. The Miocene corals grew slowly, about 2-4 mm/yr, compatible with present-day Porites from high-latitude reefs. Slow annual growth of the Miocene corals is in good agreement with the position of Crete at the margin of the Miocene reef belt. Within a given time slice, extension rates were lowest in level bottom environments and highest in attached inshore reef systems. Because sea surface temperatures (SSTs) can be expected to be uniform within a time slice, spatial variations in extension rates must reflect local variations in light levels (low in the level bottom communities) and nutrients (high in the attached reef systems). During the late Miocene (Tortonian-early Messinian), maximum linear extension rates remained remarkably constant within seven chronostratigraphic units, and if the relationship of SSTs and annual growth rates observed for modern massive Indo-Pacific Porites spp. applies to the Neogene, minimum (winter) SSTs were 20°-21°C. Although our paleoclimatic record has a low resolution, it fits the trends revealed by global data sets. In the near future we expect this new and easy to use Porites thermometer to add important new information to our understanding of Neogene climate.

Carbonate system data on the Molokai reef flat

The severity of the impact of elevated atmospheric pCO2 to coral reef ecosystems depends, in part, on how seawater pCO2 affects the balance between calcification and dissolution of carbonate sediments. Presently, there are insufficient published data that relate concentrations of pCO2 and CO3**2- to in situ rates of reef calcification in natural settings to accurately predict the impact of elevated atmospheric pCO2 on calcification and dissolution processes. Rates of net calcification and dissolution, CO3**2- concentrations, and pCO2 were measured, in situ, on patch reefs, bare sand, and coral rubble on the Molokai reef flat in Hawaii. Rates of calcification ranged from 0.03 to 2.30 mmol CaCO3/m**2/h and dissolution ranged from -0.05 to -3.3 mmol CaCO3/m**2/h. Calcification and dissolution varied diurnally with net calcification primarily occurring during the day and net dissolution occurring at night. These data were used to calculate threshold values for pCO2 and CO3**2- at which rates of calcification and dissolution are equivalent. Results indicate that calcification and dissolution are linearly correlated with both CO3**2- and pCO2. Threshold pCO2 and CO3**2- values for individual substrate types showed considerable variation. The average pCO2 threshold value for all substrate types was 654±195 µatm and ranged from 467 to 1003 µatm. The average CO3**2- threshold value was 152±24 µmol/kg, ranging from 113 to 184 µmol/kg. Ambient seawater measurements of pCO2 and CO3**2- indicate that CO3**2- and pCO2 threshold values for all substrate types were both exceeded, simultaneously, 13% of the time at present day atmospheric pCO2 concentrations. It is predicted that atmospheric pCO2 will exceed the average pCO2 threshold value for calcification and dissolution on the Molokai reef flat by the year 2100.

Seawater carbonate chemistry and community calcification during a Molokai reef (Hawaii) study 2006

The severity of the impact of elevated atmospheric pCO2 to coral reef ecosystems depends, in part, on how seawater pCO2 affects the balance between calcification and dissolution of carbonate sediments. Presently, there are insufficient published data that relate concentrations of pCO2 and CO3 to in situ rates of reef calcification in natural settings to accurately predict the impact of elevated atmospheric pCO2 on calcification and dissolution processes. Rates of net calcification and dissolution, CO3 concentrations, and pCO2 were measured, in situ, on patch reefs, bare sand, and coral rubble on the Molokai reef flat in Hawaii. Rates of calcification ranged from 0.03 to 2.30 mmol CaCO3 m**-2 h**-1 and dissolution ranged from -0.05 to -3.3 mmol CaCO3 m**-2 h**-1. Calcification and dissolution varied diurnally with net calcification primarily occurring during the day and net dissolution occurring at night. These data were used to calculate threshold values for pCO2 and CO3 at which rates of calcification and dissolution are equivalent. Results indicate that calcification and dissolution are linearly correlated with both CO3 and pCO2. Threshold pCO2 and CO3 values for individual substrate types showed considerable variation. The average pCO2 threshold value for all substrate types was 654±195 µatm and ranged from 467 to 1003 µatm. The average CO3 threshold value was 152±24 µmol/kg, ranging from 113 to 184 µmol/kg. Ambient seawater measurements of pCO2 and CO3 indicate that CO3 and pCO2 threshold values for all substrate types were both exceeded, simultaneously, 13% of the time at present day atmospheric pCO2 concentrations. It is predicted that atmospheric pCO2 will exceed the average pCO2 threshold value for calcification and dissolution on the Molokai reef flat by the year 2100.

Paleoecology, Diagenesis and Biological Affinity of Tetradiids in the Upper Ordovician (Sandbian) Black River Group of the Kingston Area, Ontario, Canada

Tetradiids are a group of colonial, tubular fossils that occur globally in Middle to Upper Ordovician strata. Tetradiids were first described as a type of tabulate coral; however, based on their four-fold symmetry, division, and presence of a central-sparry canal, they were recently reinterpreted as a florideophyte rhodophyte algae, a reinterpretation that is tested in this thesis. This study focused on understanding the affinity and taphonomy of this order of fossil. Research was conducted by stratigraphic and petrographic analyses of the Black River Group in the Kingston, Ontario region. Tetradiid occurrences were divided into fragment or colonial, with three morphologies of tetradiids described (Tetradium, Phytopsis and Paratetradium). Morphology is specific to depositional environment, with compact Tetradium consistently within ooid grainstones and open branching Phytopsis and chained Paratetradium consistently within mudstones. Two types of patch reefs were recognized: a Paratetradium bioherm, and a Paratetradium, Phytopsis, stromatolite bioherm. The presence of bioherms implies that tetradiids were capable of hypercalcifying. Preservation styles of tetradiids were investigated, and were compared to brachiopods, echinoderms, mollusks, and ooids. Tetradiids were preferentially preserved as molds and demonstrated complete dissolution of skeletal material. Rare specimens, however, demonstrated preserved horizontal partitions, central plates, and a double wall. Skeletal molds were filled with either calcite spar, mud or encrusted by a cryptomicrobial colony. Both calcitic and aragonitic ooids were discovered. The co-occurrence of aragonitic ooids, aragonitic crytodontids, and the evolution of aragonitic, hypercalcifying tetradiids is interpreted as representing the geochemical favoring of aragonite and HMC in a time of global calcite seas. The geochemical favoring of aragonite is interpreted to be independent to global Mg: Ca ratios, but was the result of increased saturation levels and temperature driven by high atmospheric pCO2. Based on the presence of epitheca, tabulae, septa, and the commonality of growth forms, tetradiids are interpreted as an order of Cnidaria. The evolution of an aragonitic skeleton in tetradiids is interpreted to be the result of de novo acquisition of a skeleton from an unmineralized clade.

Long-term stasis in ecological assemblages: Evidence from the fossil record

Studies of plant and animal assemblages from both the terrestrial and the marine fossil records reveal persistence for extensive periods of geological time, sometimes millions of years. Persistence does not require lack of change or the absence of variation from one occurrence of the assemblage to the next in geological time. It does, however, imply that assemblage composition is bounded and that variation occurs within those bounds. The principal cause for these patterns appears to be species-, and perhaps clade-level, environmental fidelity that results in long-term tracking of physical conditions. Other factors that influence persistent recurrence of assemblages are historical, biogeographic effects, the law of large numbers, niche differentiation, and biotic interactions. Much research needs to be done in this area, and greater uniformity is needed in the approaches to studying the problem. However, great potential also exists for enhanced interaction between paleoecology and neoecology in understanding spatiotemporal complexity of ecological dynamics.

Recent and Miocene carbonate sediments from Indonesia. Available in 2 volumes.

Mapping and sediment sampling in reefs of the Pulau Seribu group (southwest Java Sea) shows the existence of ten physiographic zones and subzones represented by seven lithofacies. Reefs in the northern part of the archipelago are smaller, more closely spaced and morphologically sim pler than those in the south. This pattern is attributed to differences in subsidence rate. A th reedimensional model is proposed for the evo lution of these reefs but borehole data are requi red to test this model. Miocene limestones are described in detail from hydrocarbon reservoirs in the Batu Raja Formation of the same area. Brief comparisons a re made with surface outcrops of approximately coeval carbonate developments. The lithofacies developed within these limestones reflect variations in hydrodynam ic regime and basement topography . Ele\le.n diagenetic processes affected the Batu Raja limestones and the dist ribution of these is primarily related to sealevel fluctuations. Early diagenesis was marine and characterised by micritisation and preCipitation of fibrous and bladed cements. Dolomitisat ion occurred in the mixed- water zone and its variable intensity is attributed to the configuration of the carbonate body relative to this zone. Subsequently the limestones were subjected to freshwater phreatic zone diagenesis resulting in dissolution and cementation, and a t a late stage underwent burial compaction. Secondary porosity, which \ar9e1.y determines the suitability of these limestones as hydrocarbon reserVOirs, is a function of the variable intensity of dissolution and cementation, burial compaction, dolomitisation and possibly micrite neomorphism. The sedimentary processes that generated the Batu Raja buildups are inferred f rom comparisons with the Pulau Seribu and other Recent analogues. The contrasting pinnacle form of the Pulau Seribu patch reefs compared with the low relief of the Batu Raja buUdups results from differences in the initial substrate topography and subsequent subsidence rate