Net community production, Thorium-234 activity and particle flux during the Southern Ocean iron fertilization experiment LOHAFEX

A closed eddy core in the Subantarctic Atlantic Ocean was fertilized twice with two tons of iron (as FeSO4), and the 300 km**2 fertilized patch was studied for 39 days to test whether fertilization enhances downward particle flux into the deep ocean. Chlorophyll a and primary productivity doubled after fertilization, and photosynthetic quantum yield (FV/FM) increased from 0.33 to >0.40. Silicic acid (<2 µmol/L) limited diatoms, which contributed <10% of phytoplankton biomass. Copepods exerted high grazing pressure. This is the first study of particle flux out of an artificially fertilized bloom with very low diatom biomass. Net community production (NCP) inside the patch, estimated from O2:Ar ratios, averaged 21 mmol POC/m**2/d, probably ±20%. 234Th profiles implied constant export of ~6.3 mmol POC/m**2/d in the patch, similar to unfertilized waters. The difference between NCP and 234Th-derived export partly accumulated in the mixed layer and was partly remineralized between the mixed layer and 100 m. Neutrally buoyant sediment traps at 200 and 450 m inside and outside the patch caught mostly <1.1 mmol POC/m**2/d, predominantly of fecal origin; flux did not increase upon fertilization. Our data thus indicate intense flux attenuation between 100 and 200 m, and probably between the mixed layer and 100 m. We attribute the lack of fertilization-induced export to silicon limitation of diatoms and reprocessing of sinking particles by detritus feeders. Our data are consistent with the view that nitrate-rich but silicate-deficient waters are not poised for enhanced particle export upon iron addition.

Carbon, nitrogen and lipid content and egg and pellet production of female C. hyperboreus in situ and during incubation experiments

The effects of temperature and food availability on feeding and egg production of the Arctic copepod Calanus hyperboreus were investigated in Disko Bay, western Greenland, from winter to spring 2009. The abundance of females in the near bottom layer and the egg production of C. hyperboreus prior to the spring bloom document that reproduction relies on lipid stores. The maximum in situ egg production (± SE) of 54 ± 8 eggs female/d was recorded in mid-February at chlorophyll a concentrations below 0.1 µg/l, whereas no egg production was observed in mid-April when the spring bloom developed. After reproduction, the females migrated to the surface layer to exploit the bloom and refill their lipid stores. In 2 laboratory experiments, initiated before and during the spring bloom, mature females were kept with and without food at 5 different temperatures ranging from 0 to 10°C and the fecal pellet and egg production were monitored. Food had a clear effect on fecal pellet production but no effect on egg production, while temperature did not have an effect on egg or fecal pellet production in any of the experiments. Analyses of carbon and lipid content of the females before and after the experiments did not reflect any effect of food or temperature in the pre-bloom experiment, whereas in the bloom experiment a clear positive effect of food was detected in female biochemical profiles. The lack of a temperature response suggests a future warmer ocean could be unfavorable for C. hyperboreus compared to smaller Calanus spp. which are reported to exploit minor temperature elevations for increased egg production.

Spatial modeling of data with excessive zeros applied to reindeer pellet-group counts

We analyze a real data set pertaining to reindeer fecal pellet-group counts obtained from a survey conducted in a forest area in northern Sweden. In the data set, over 70% of counts are zeros, and there is high spatial correlation. We use conditionally autoregressive random effects for modeling of spatial correlation in a Poisson generalized linear mixed model (GLMM), quasi-Poisson hierarchical generalized linear model (HGLM), zero-inflated Poisson (ZIP), and hurdle models. The quasi-Poisson HGLM allows for both under- and overdispersion with excessive zeros, while the ZIP and hurdle models allow only for overdispersion. In analyzing the real data set, we see that the quasi-Poisson HGLMs can perform better than the other commonly used models, for example, ordinary Poisson HGLMs, spatial ZIP, and spatial hurdle models, and that the underdispersed Poisson HGLMs with spatial correlation fit the reindeer data best. We develop R codes for fitting these models using a unified algorithm for the HGLMs. Spatial count response with an extremely high proportion of zeros, and underdispersion can be successfully modeled using the quasi-Poisson HGLM with spatial random effects.

Spatial and temporal patterns in ungulate-ecosystem interactions

Ungulates are important components of a variety of ecosystems worldwide. This dissertation integrates aspects of ungulate and forest ecology to increase our understanding of how they work together in ways that are of interest to natural resource managers, educators, and those who are simply curious about nature. Although animal ecology and ecosystem ecology are often studied separately, one of the general goals of this dissertation is to examine how they interact across spatial and temporal scales. Forest ecosystems are heterogeneous across a range of scales. Spatial and temporal habitat use patterns of forest ungulates tend to be congregated in patches where food and/or cover are readily available. Ungulates interact with ecosystem processes by selectively foraging on plants and excreting waste products in concentrated patches. Positive feedbacks may develop where these activities increase the value of habitat through soil fertilization or the alteration of plant chemistry and architecture. Heterogeneity in ecosystem processes and plant community structure, observed at both stand and local scales, may be the integrated outcome of feedbacks between ungulate behavior and abiotic resource gradients. The first chapter of this dissertation briefly discusses pertinent background information on ungulate ecology, with a focus on white-tailed deer (Odocoileus virginianus) in the Upper Great Lakes region and moose (Alces acles) in Isle Royale National Park, Michigan, USA. The second chapter demonstrates why ecological context is important for studying ungulate ecology in forest ecosystems. Excluding deer from eastern hemlock (Tsuga canadensis) stands, which deer use primarily as winter cover, resulted in less spatial complexity in soil reactive nitrogen and greater complexity in diffuse light compared to unfenced stands. The spatial patterning of herbaceous-layer cover was more similar to nitrogen where deer were present, and was a combination of nitrogen and light within deer exclosures. This relationship depends on the seasonal timing of deer habitat use because deer fertilize the soil during winter, but leave during the growing season. The third chapter draws upon an eight-year, 39-stand data set of deer fecal pellet counts in hemlock stands to estimate the amount of nitrogen that deer are depositing in hemlock stands each winter. In stands of high winter deer use, deer-excreted nitrogen inputs consistently exceeded those of atmospheric deposition at the stand scale. At the neighborhood scale, deer-excreted nitrogen was often in excess of atmospheric deposition due to the patchy distribution of deer habitat use. Spatial patterns in habitat use were consistent over the eight-year study at both stand and neighborhood scales. The fourth chapter explores how foraging selectivity by moose interacts with an abiotic resource gradient to influence forest structure and composition. Soil depth on Isle Royale varies from east to west according to glacial history. Fir saplings growing in deeper soils on the west side are generally more palatable forage for moose (lower foliar C:N) than those growing in shallower soils on the east side. Therefore, saplings growing in better conditions are less likely to reach the canopy due to moose browsing, and fir is a smaller overstory component on the west side. Lastly, chapter five focuses on issues surrounding eastern hemlock regeneration failure, which is a habitat type that is important to many wildlife species. Increasing hemlock on the landscape is complicated by several factors including disturbance regime and climate change, in addition to the influence of deer.

(Table 2) Organic residue at DSDP Hole 76-534A

The kind, sedimentation rate, and diagenesis of organic particles delivered to the North Atlantic seafloor during the Middle Jurassic-Early Cretaceous were responsible for the presence of carbonaceous sediments in Hole 534A. Organic-rich black clays formed from the rapid supply of organic matter; this organic matter was composed of either abundant, well-preserved, and poorly sorted particles of land plants deposited in clays and silty clays within terrigenous turbiditic sequences (tracheal facies) or abundant amorphous debris (xenomorphic facies) generated through the digestive tracts of marine zooplankton and sedimented as fecal pellets. Evidence for the fecal-pellet origin of xenomorphic debris is illustrated. Black clays were also produced in sediments containing less organic matter as a result of the black color of carbonized particles composing all or most of the residues (micrinitic facies). Slowly sedimented hematitic Aptian clays contain very little carbonized, organic debris that survived diagenetic oxidation. In the red calcareous clay sequence of the Late Jurassic, larger amounts of this oxidized debris turned several clay layers black or blackish red. Carbonized debris also dominates the residues recovered in interbedded black and green Albian clays. Carbonization of organic matter in these sediments either turned them black or provided the diagenetic environment for reduced iron. Carbonized debris is also appreciable in burrow-mottled black-green Kimmeridgian clay. The study of Hole 534A organic matter indicates that during the middle Callovian there was a rapid supply of terrigenous organic matter, followed by a late Callovian episode of rapidly supplied xenomorphic debris deposited as fecal pellets. The Late Jurassic-Berriasian was a time of slower sedimentation of organic matter, primarily of a marine dinoflagellate flora in a poorly preserved xenomorphic facies variously affected by diagenetic oxidation. Several intervals of carbonized tracheal tissue in the Oxfordian and Kimmeridgian suggest episodes of oxidized terrigenous matter. The same sequence of Callovian organic events is evident in much of the Early Cretaceous

(Appendix A) Organic facies at DSDP Hole 93-603B

The Palynology of two sections recovered during Leg 93 drilling by the Deep Sea Drilling Project in the continental rise along the western margin of the North Atlantic is reported. In Hole 603B at Site 603, the dinoflagellate stratigraphy indicates that the interval from Cores 603B-82 to 603B-26 ranges in age from late Berriasian to Santonian. The BlakeBahama Formation ranges from late Berriasian to Aptian. The Hatteras Formation ranges from Aptian to Cenomanian, although the uppermost part may be Turonian. Dinoflagellate evidence from the middle part of the Plantagenet Formation indicates an age from late Coniacian or early Santonian to Santonian within the interval of Cores 603B-28 to 603B-26. Magnetic polarity evidence of the stratigraphy of the Early Cretaceous for the western North Atlantic indicates a reliable correlation with the dinoflagellate zonation. The stratigraphic sequence of palynologically defined organic facies in carbonaceous claystone lithologies in Hole 603B shows that organic stratigraphic units consisting predominantly of fecal-pellet-derived, pelagic organic matter (xenomorphic facies) alternate with units consisting predominantly of terrigenous organic matter (tracheal and exinitic facies), corresponding to that described from other sites in the North Atlantic. A terrigenous organic facies is identified for the first time from the Plantagenet Formation. The claystone organic facies and major lithofacies are closely correlated. The tracheal and exinitic facies occur in carbonaceous terrigenous claystones and claystone turbidites associated with sandstone/siltstone terrigenous turbidites. The xenomorphic facies occurs in claystones within pelagic limestones lacking any turbidites, and in blackish, noncalcareous claystones which correlate in age with the marine-carbon-rich sapropels which are widespread in the North Atlantic Cenomanian. This facies also occurs with an admixture of terrigenous organic particles in the Blake-Bahama Formation, but the mixture is consistent with the submarine fan setting of this interval. The concentration of refractory organic matter (carbonized particles) in the micrinitic and carbonized tracheal facies is considered to be the result, at least in part, of the oxidation of sediment buried below a surface slowly accumulating pelagic clays below the carbonate compensation depth. The progressive increase in number of dinoflagellate species per stage through the Early Cretaceous (except for the late Barremian-Aptian) may have resulted indirectly from the generally progressive rise in global sea level during this time. At Site 605, the dinoflagellate stratigraphy across the Cretaceous/Tertiary boundary is remarkably close to that published from the Maestrichtian and Danian of Denmark. The Maestrichtian/Danian boundary is placed precisely within Section 605-66-1 by dinoflagellate evidence, agreeing with that predicted by other microfossils. The new dinoflagellate-cyst-based genus, Pierceites and its new species P. schizocystis, and the new combination P. ( = Trithyrodinium) pentagonum (May) are proposed. Diacanthum hollisteri Habib, type species of Diacanthum, is emended to accommodat e cysts with the archeopyle formulas P3'', 2P2''-3'', 2P3''-4'', and 3P2''-3''-4''.

Rates of egg production, grazing and mortatlity of Calanus finmarchicus measured experimentally in the North Atlantic during the Maria S. Merian cruise MSM26, spring 2013

An incubation experiment at five different temperatures was used to assess the potential for adaptation of Calanus finmarchicus to future warming of the ocean. During a short term (3 h) and long term (6 day) exposure of individual females to a gradient of temperature stress, egg production and fecal pellet production were monitored to indicate secondary production and grazing rates. A longer term (10 day) exposure to elevated temperatures followed by a return to ambient sea temperatures was used to assess the potential recovery of individuals exposed to temperature stress. Females were picked out from WP2 net samples and acclimatised in 2 L bottles of GFF filtered seawater with Thalassiosira weissflogii as prey for >48 h at ambient SST. Experimental bottles were filled with filtered seawater (GFF filtered from non-toxic seawater supply) and acclimated to experimental temperature overnight (0, 5, 10, 15 and 20 °C). Individual females were transferred into bottles using forceps and the bottles were inoculated with T. weissflogii to a final concentration of 5 µg chl L-1. Bottles were then placed into water baths and incubated for 3h or 6 d, and monitored for egg and fecal pellet production rates. A 10 day exposure experiment was used to test the potential for recovery from temperature stress, by returning females incubated at 5, 10, 15 and 20 °C back to 10 °C for 24 h and counting egg and fecal pellet production.