Seed dispersal by the lek-forming white-bearded manakin (Manacus manacus, Pipridae) in the Brazilian Atlantic forest

The movement patterns of males, females and juveniles of lekking species often differ due to differences in the commitment to lek activities, which may lead to differences in the spatial distribution and dispersal distances of seeds they eat. By sampling seeds in three lek and non-lek areas of the white-bearded manakin (Manacus manacus), we tested whether this lekking species increased the abundance and species richness of seeds in lek areas and, at a finer scale, in 21 displaying courts within lek areas. Combining data on seed defecation or regurgitation rates by free-ranging individuals, the number of seeds in droppings or regurgitations of mist-netted birds, and the distances travelled by birds equipped with radio-transmitters, we estimated the potential spatial distribution of seeds generated by six resident males and six females or juveniles during the morning peak of lek activity and when lek activity decreased in the afternoon. There was no difference in the species richness (46 and 44 morphospecies, respectively) and abundance of seeds (15.4 ± 7.3 seeds and 14.0 ± 1.1 seeds, respectively) between lek and non-lek areas. Within leks both parameters increased in courts (45 spp., 17.6 ± 14 seeds) compared with non-court sites (22 spp., 1.9 ± 1.8 seeds), likely as a consequence of the longer time spent by resident males in perches in or near display courts. Distances moved by juveniles and females per 60-min period (183 ± 272 m) were greater than resident males (42.6 ± 22.0 m) in the mornings, while the opposite happened in the afternoons (55.2 ± 40.7 m and 157 ± 105 m, respectively). We conclude that the spatial aggregation of seeds in lek areas of M. manacus occurs at the court level, and the spatial distribution of deposited seeds varies with manakin lekking status and the daily period of foraging. © Cambridge University Press 2013.

Landscape Diversity Influences Dispersal and Establishment of Pest with Complex Nutritional Ecology

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Model complexity affects transient population dynamics following a dispersal event: A case study with pea aphids

Stage-structured population models predict transient population dynamics if the population deviates from the stable stage distribution. Ecologists’ interest in transient dynamics is growing because populations regularly deviate from the stable stage distribution, which can lead to transient dynamics that differ significantly from the stable stage dynamics. Because the structure of a population matrix (i.e., the number of life-history stages) can influence the predicted scale of the deviation, we explored the effect of matrix size on predicted transient dynamics and the resulting amplification of population size. First, we experimentally measured the transition rates between the different life-history stages and the adult fecundity and survival of the aphid, Acythosiphon pisum. Second, we used these data to parameterize models with different numbers of stages. Third, we compared model predictions with empirically measured transient population growth following the introduction of a single adult aphid. We find that the models with the largest number of life-history stages predicted the largest transient population growth rates, but in all models there was a considerable discrepancy between predicted and empirically measured transient peaks and a dramatic underestimation of final population sizes. For instance, the mean population size after 20 days was 2394 aphids compared to the highest predicted population size of 531 aphids; the predicted asymptotic growth rate (λmax) was consistent with the experiments. Possible explanations for this discrepancy are discussed. Includes 4 supplemental files.

Contributions of demography and dispersal parameters to the spatial spread of a stage-structured insect invasion

Stage-structured models that integrate demography and dispersal can be used to identify points in the life cycle with large effects on rates of population spatial spread, information that is vital in the development of containment strategies for invasive species. Current challenges in the application of these tools include: (1) accounting for large uncertainty in model parameters, which may violate assumptions of ‘‘local’’ perturbation metrics such as sensitivities and elasticities, and (2) forecasting not only asymptotic rates of spatial spread, as is usually done, but also transient spatial dynamics in the early stages of invasion. We developed an invasion model for the Diaprepes root weevil (DRW; Diaprepes abbreviatus [Coleoptera: Curculionidae]), a generalist herbivore that has invaded citrus-growing regions of the United States. We synthesized data on DRW demography and dispersal and generated predictions for asymptotic and transient peak invasion speeds, accounting for parameter uncertainty. We quantified the contributions of each parameter toward invasion speed using a ‘‘global’’ perturbation analysis, and we contrasted parameter contributions during the transient and asymptotic phases. We found that the asymptotic invasion speed was 0.02–0.028 km/week, although the transient peak invasion speed (0.03– 0.045 km/week) was significantly greater. Both asymptotic and transient invasions speeds were most responsive to weevil dispersal distances. However, demographic parameters that had large effects on asymptotic speed (e.g., survival of early-instar larvae) had little effect on transient speed. Comparison of the global analysis with lower-level elasticities indicated that local perturbation analysis would have generated unreliable predictions for the responsiveness of invasion speed to underlying parameters. Observed range expansion in southern Florida (1992–2006) was significantly lower than the invasion speed predicted by the model. Possible causes of this mismatch include overestimation of dispersal distances, demographic rates, and spatiotemporal variation in parameter values. This study demonstrates that, when parameter uncertainty is large, as is often the case, global perturbation analyses are needed to identify which points in the life cycle should be targets of management. Our results also suggest that effective strategies for reducing spread during the asymptotic phase may have little effect during the transient phase. Includes Appendix.

The evolution of mating rates in Pieris napi

In the green-veined white butterfly (Pieris napi), females obtain direct fitness benefits from mating multiply and studies have shown that fitness increases seemingly monotonically with number of matings. The reason is that at mating males transfer a large nutritious gift (a so called nuptial gift) to the females that the females use to increase both their fecundity and lifespan. In addition, if exposed to poor food conditions as larvae, females mature at a smaller size compared to males. Accordingly, it was suggested that smaller females could compensate for their size through nuptial feeding by, for instance, mating more frequently. We did not find any support for that hypothesis. On the contrary, larger females remated sooner and had a higher lifetime number of matings. Neither were smaller females able to compensate in any other way, because singly mated females and multiply mated females suffered to the same extent from their smaller size. This thesis also shows that despite the positive relationship between fitness and number of matings, there is a large variation in female mating frequency in wild populations and about every second female mates only once or twice. This variation is not dependent on how often females get courted by males, because female mating frequency was shown not to be affected by male courtship intensity. Hence, the reason for the low mating frequency could either be that males have evolved the ability to manipulate females to mate at a suboptimal rate as a measure of protection against sperm competition, or alternatively, that female mating rate is suppressed by some costs. Using two selection lines, artificially selected for either a high or a low mating rate, we showed that the variation in mating rate was mainly a female trait because which line the females were from affected their mating rate whereas which line the male was from did not. This implies that females mate at a low rate due to hidden costs or due to constraints. The same study also showed that females with a high "intrinsic" mating rate lived shorter, but only when denied remating. This led us to test the hypothesis that the cost females face is to have the ability to mate at a high rate but the cost is only paid when remating opportunities are scarce. However, we found no support for such an idea, because females with a high intrinsic mating rate held in a cold environment where the butterflies were prevented from flying and feeding did not live shorter. Neither was there an effect of a female’s mating rate on her ability to quickly break down and convert male nutrient gifts into egg material. Female mating rate did, on the other hand, affect dispersal tendency, with low mating rate females being more inclined to fly between different habitats. The underlying reason for this is still to be explored.

Influence of pollination and seed dispersal on the genetic population structure of two Commiphora species in Madagascar and South Africa

Pollination and seed dispersal are important ecological processes for the regeneration of plant populations and both vectors for gene exchange between plant populations. For my thesis, I studied the pollination ecology of the South African tree Commiphora harveyi (Burseraceae) and compared it with C. guillauminii from Madagascar. Both species have low visitation rates and a low number of pollinating insect species, resulting in a low fruit set. While their pollination ecology is very similar, they differ in their seed dispersal with a low seed dispersal rate in the Malagasy and a high seed dispersal rate in the South African species. This should be reflected in a stronger genetic differentiation among populations in the Malagasy than in the South African species. My results, based on AFLP markers, contradict these expectations, the overall differentiation was lower in the Malagasy (FST = 0.05) than in the South African species (FST = 0.16). However, at a smaller spatial scale (below 3 km), the Malagasy species was genetically more strongly differentiated than the South African species, which was reflected by the high inter-population variance within the sample site (C. guillauminii: 72.2 - 85.5 %; C. harveyi: 8.4 - 14.5 %). This strong differentiation could arise from limited gene flow, which was confirmed by spatial autocorrelation analyses. The shape of the autocorrelogram suggested that gene exchange between individuals occurred only up to 3 km in the Malagasy species, whereas up to 30 km in the South African species. These results on the genetic structure correspond to the expectations based on seed dispersal data. Thus, seed dispersal seems to be a key factor for the genetic structure in plant populations on a local scale.

Frugivores, seed dispersal and tree regeneration along a human disturbance gradient in East African tropical rainforests

Samenausbreitung und Regeneration von Bäumen sind wichtig für den langfristigen Bestand von Baum- und Frugivorengemeinschaften in tropischen Regenwäldern. Zunehmende Rohdung und Degradation gefährden den Ablauf dieser mutualistischen Prozesse in diesem Ökosystem. Um den Einfluss von kleinräumiger menschlicher Störung auf die Frugivorengemeinschaft und die zentralen Ökosystemprozesse Samenausbreitung und Regeneration zu erforschen, habe ich 1) die Frugivorengemeinschaft und die Samenausbreitungsrate von Celtis durandii (Ulmaceae) und 2) den Zusammenhang zwischen Baumarten mit fleischigen Früchten, Frugivoren und der Etablierung von Keimlingen dieser Baumarten in unterschiedlich stark gestörten Flächen dreier ostafrikanischer tropischer Regenwälder untersucht. Insgesamt konnte ich 40 frugivore Vogel- und Primatenarten in den drei untersuchten Waldgebieten nachweisen. Auf gering gestörten Flächen wurden mehr Frugivore als auf stark gestörten Flächen aufgenommen. Auch die Beobachtungen an C. durandii ergaben mehr frugivore Besucher in Bäumen auf gering gestörten als auf stark gestörten Flächen. Dies führte zu einer marginal signifikant höheren Samenausbreitungsrate auf den gering gestörten Flächen. Diese Ergebnisse waren auf regionaler Ebene in allen drei untersuchten Wäldern konsistent. Dies zeigt, dass kleinräumige Störung einen umfassenderen negativen Einfluss auf Frugivore und ihre Funktion als Samenausbreiter hat als zuvor angenommen. Bei der Vegetationserfassung nahm ich 131 verschiedene Baumarten mit fleischigen Früchten in den drei Regenwäldern auf. Kleinräumige menschliche Störung erhöhte den Artenreichtum dieser Baumarten marginal signifikant, hatte jedoch keinen direkten Einfluss auf die Frugivorendichte und den Artenreichtum von Keimlingen dieser Baumarten. Der Artenreichtum von Baumarten mit fleischigen Früchten zeigte einen marginal signifikant positiven Einfluss auf die Frugivorendichte, allerdings nicht auf die Keimlinge. Allerdings führte die Dichte der Frugivoren zu signifikant erhöhtem Artenreichtum der Keimlinge. Folglich scheint kleinräumige Störung die Keimlingsetablierung indirekt durch erhöhten Baumartenreichtum und erhöhte Frugivorendichte zu beeinflussen. Die Frugivorendichte hatte einen größeren Einfluss auf die Waldregeneration als kleinräumige Störung und Baumartenreichtum. Demnach scheint kleinräumige menschliche Störung sowohl positive als auch negative Effekte auf Samenausbreitung und Regeneration zu haben. Somit sind weitere Studien notwendig, die den Einfluss von kleinräumiger menschlicher Störung auf Mutualismen tropischer Regenwälder aufklären.

Delayed dispersal as a potential route to cooperative breeding in ambrosia beetles

Abstract  Xyleborini are a species-rich tribe of ambrosia beetles, which are haplodiploid and typically mate among siblings within their natal brood chamber. Several characteristics of this tribe would predict the evolution of higher levels of sociality: high genetic relatedness within galleries due to inbreeding, high costs of dispersal and the potential benefit of cooperation in brood care within the natal gallery (e.g. by fungus gardening, gallery extension, offspring feeding and cleaning). However, information on the social system of these beetles is very limited. We examined the potential for cooperative breeding in Xyleborinus saxeseni by monitoring dispersal in relation to brood size and composition. Results show that adult female offspring delay dispersal despite dispersal opportunities, and apparently some females never disperse. The femalesâ?? decision to stay seems to depend on the presence of eggs and dependent siblings. We found no indication that female offspring reproduce in their natal gallery, as colonies with many mature daughters do not contain more eggs than those with few or no daughters. There is a significant positive relationship between the number of females present and the number of dependent siblings (but not eggs), which suggests that cooperative brood care of female offspring raises colony productivity by improving survival rates of immatures. Our results suggest that cooperative breeding is likely to occur in X. saxeseni and possibly other xyleborine species. We argue that a closer look at sociality within this tribe may yield important information on the factors determining the evolution of cooperative breeding and advanced social organization.

Gastropod Seed Dispersal: An Invasive Slug Destroys Far More Seeds in Its Gut than Native Gastropods

Seed dispersal is one of the most important mechanisms shaping biodiversity, and animals are one of the key dispersal vectors. Animal seed dispersal can directly or indirectly be altered by invasive organisms through the establishment of new or the disruption of existing seed dispersal interactions. So far it is known for a few gastropod species that they ingest and defecate viable plant seeds and consequently act as seed dispersers, referred to as gastropodochory. In a multi-species experiment, consisting of five different plant species and four different gastropod species, we tested with a fully crossed design whether gastropodochory is a general mechanism across native gastropod species, and whether it is altered by the invasive alien slug species Arion lusitanicus. Specifically, we hypothesized that a) native gastropod species consume the seeds from all tested plant species in equal numbers (have no preference), b) the voracious invasive alien slug A. lusitanicus – similarly to its herbivore behaviour – consumes a higher amount of seeds than native gastropods, and that c) seed viability is equal among different gastropod species after gut passage. As expected all tested gastropod species consumed all tested plant species. Against our expectation there was a difference in the amount of consumed seeds, with the largest and native mollusk Helix pomatia consuming most seeds, followed by the invasive slug and the other gastropods. Seed damage and germination rates did not differ after gut passage through different native species, but seed damage was significantly higher after gut passage through the invasive slug A. lusitanicus, and their germination rates were significantly reduced.

Dispersal and microsite limitation in an abandoned calcareous grassland of the southern prealps

Dispersal limitation is often involved when the species composition of a dry abandoned grassland shows a slow response to resumed regular mowing. A seed-addition experiment, using 32 species which do not belong to the local species pool, was performed on Monte San Giorgio (southern Switzerland) to test whether the low recruitment success was due to dispersal limitation or due to unfavourable microsite conditions. In October 1997, 20 species were individually sown in six 3 × 4 m blocks of a 2 × 2 factorial “partial” split-plot design with treatments of abandonment vs. mowing and undisturbed vs. root-removed soil, this last being applied in small naturally-degradable pots. Moreover, 12 species were sown only in the treatments on undisturbed soil. Seedlings of sown and spontaneously germinating seeds were observed on 16 occasions over one 12-month period. Seeds of 31 out of the 32 species germinated. Twenty-four species showed germination rates higher than 5% and different seasonal germination patterns. Established vegetation, especially the tussocks ofMolinia arundinacea, reduced the quality of microsites for germination. Whereas a few species germinated better under the litter ofMolinia arundinacea, many more germinated better under the more variable microsite conditions of a mown grassland. Only a few seedlings of 25 species out of the 31 germinated species survived until October 1998. Seedling survival was negatively affected by litter, unfavourable weather conditions (frost and dry periods followed by heavy rains) and herbivory (slugs and grasshoppers). Tussocks ofMolinia arundinacea, however, tended to protect seedlings. The poor establishment success of “new” species observed in abandoned meadows on Monte San Giorgio after resumed mowing is due to dispersal and microsite limitations.

Can seed production and restricted dispersal limit recruitment in Pinus pinaster Aiton from the Spanish Northern Plateau?

Natural regeneration faces increasing difficulties in dry forests from the Mediterranean basin, including for normally well-regenerating species such as maritime pine (Pinus pinaster Aiton). In this paper, we studied female fertility, seed dispersal and spread rates in P. pinaster from the Spanish Northern Plateau, where natural regeneration failure is a main concern for forest managers. For this purpose we periodically collected data from seed traps and trees located at two core locations across several years. We found significant variation in interannual cone production, with the best seed trees being the same across years. In addition, we found highly skewed distributions of female reproductive effort and large fertility differences across stands located few kilometres away. Annual seed dispersal kernels fitted lognormal or 2Dt models depending on the stand analysed, with median dispersal distances between 14 and 25 m. Kernels fitted for maximum dispersal periods showed an outstanding intraseasonal variation of median dispersal distances, from 10 to 54 m, in association to variable patterns of rainfall and maximum wind speed. The amount of seed produced appeared to be enough to guarantee the natural regeneration of the stands during the typical 20-year regeneration period. Colonisation simulations concluded that Mediterranean maritime pine has a notable dispersion capacity, which is strongly influenced by levels of fecundity and, especially, by the number and frequency of long-distance dispersal events. The latter play a key role in tree dispersion processes through enlarging the occupied area and fostering the invasion of abandoned crop land.

Rates of spread of marine pathogens

Epidemics of marine pathogens can spread at extremely rapid rates. For example, herpes virus spread through pilchard populations in Australia at a rate in excess of 10 000 km year(-1), and morbillivirus infections in seals and dolphins have spread at more than 3000 km year(-1). In terrestrial environments, only the epidemics of myxomatosis and calicivirus in Australian rabbits and West Nile Virus in birds in North America have rates of spread in excess of 1000 km year(-1). The rapid rates of spread of these epidemics has been attributed to flying insect vectors, but flying vectors have not been proposed for any marine pathogen. The most likely explanation for the relatively rapid spread of marine pathogens is the lack of barriers to dispersal in some parts of the ocean, and the potential for long-term survival of pathogens outside the host. These findings caution that pathogens may pose a particularly severe problem in the ocean. There is a need to develop epidemic models capable of generating these high rates of spread and obtain more estimates of disease spread rate.

An investigation into effects of long-distance seed dispersal on organelle population genetic structure and colonization rate: a model analysis

A simulation-based modelling approach is used to examine the effects of stratified seed dispersal (representing the distribution of the majority of dispersal around the maternal parent and also rare long-distance dispersal) on the genetic structure of maternally inherited genomes and the colonization rate of expanding plant populations. The model is parameterized to approximate postglacial oak colonization in the UK, but is relevant to plant populations that exhibit stratified seed dispersal. The modelling approach considers the colonization of individual plants over a large area (three 500 km x 10 km rolled transects are used to approximate a 500 km x 300 km area). Our approach shows how the interaction of plant population dynamics with stratified dispersal can result in a spatially patchy haplotype structure. We show that while both colonization speeds and the resulting genetic structure are influenced by the characteristics of the dispersal kernel, they are robust to changes in the periodicity of long-distance events, provided the average number of long-distance dispersal events remains constant. We also consider the effects of additional physical and environmental mechanisms on plant colonization. Results show significant changes in genetic structure when the initial colonization of different haplotypes is staggered over time and when a barrier to colonization is introduced. Environmental influences on survivorship and fecundity affect both the genetic structure and the speed of colonization. The importance of these mechanisms in relation to the postglacial spread and genetic structure of oak in the UK is discussed.

Using data from an encounter sampler to model fish dispersal

A method to estimate speed of free-ranging fishes using a passive sampling device is described and illustrated with data from the Everglades, U.S.A. Catch per unit effort (CPUE) from minnow traps embedded in drift fences was treated as an encounter rate and used to estimate speed, when combined with an independent estimate of density obtained by use of throw traps that enclose 1 m2 of marsh habitat. Underwater video was used to evaluate capture efficiency and species-specific bias of minnow traps and two sampling studies were used to estimate trap saturation and diel-movement patterns; these results were used to optimize sampling and derive correction factors to adjust species-specific encounter rates for bias and capture efficiency. Sailfin mollies Poecilia latipinna displayed a high frequency of escape from traps, whereas eastern mosquitofish Gambusia holbrooki were most likely to avoid a trap once they encountered it; dollar sunfish Lepomis marginatus were least likely to avoid the trap once they encountered it or to escape once they were captured. Length of sampling and time of day affected CPUE; fishes generally had a very low retention rate over a 24 h sample time and only the Everglades pygmy sunfish Elassoma evergladei were commonly captured at night. Dispersal speed of fishes in the Florida Everglades, U.S.A., was shown to vary seasonally and among species, ranging from 0· 05 to 0· 15 m s−1 for small poeciliids and fundulids to 0· 1 to 1· 8 m s−1 for L. marginatus. Speed was generally highest late in the wet season and lowest in the dry season, possibly tied to dispersal behaviours linked to finding and remaining in dry-season refuges. These speed estimates can be used to estimate the diffusive movement rate, which is commonly employed in spatial ecological models.

Age models, accumulation rates, and grain size distributions of sediment cores from the northern subtropical Atlantic

The terrigenous sediment proportion of the deep sea sediments from off Northwest Africa has been studied in order to distinguish between the aeolian and the fluvial sediment supply. The present and fossil Saharan dust trajectories were recognized from the distribution patterns of the aeolian sediment. The following timeslices have been investigated: Present, 6,000, 12,000 and 18,000 y. B. P. Furthermore, the quantity of dust deposited off the Saharan coast has been estimated. For this purpose, 80 surface sediment samples and 34 sediment cores have been analysed. The stratigraphy of the cores has been achieved from oxygen isotopic curves, 14C-dating, foraminiferal transfer temperatures, and carbonate contents. Silt sized biogenic opal generally accounts for less than 2 % of the total insoluble sediment proportion. Only under productive upwelling waters and off river mouths, the opal proportion exceeds 2 % significantly. The modern terrigenous sediment from off the Saharan coast is generally characterized by intensely stained quartz grains. They indicate an origin from southern Saharan and Sahelian laterites, and a zonal aeolian transport in midtropospheric levels, between 1.5 an 5.5 km, by 'Harmattan' Winds. The dust particles follow large outbreaks of Saharan air across the African coast between 15° and 21° N. Their trajectories are centered at about 18° N and continue further into a clockwise gyre situated south of the Canary Islands. This course is indicated by a sickle-shaped tongue of coarser grain sizes in the deep-sea sediment. Such loess-sized terrigenous particles only settle within a zone extending to 700 km offshore. Fine silt and clay sized particles, with grain sizes smaller than 10- 15 µm, drift still further west and can be traced up to more than 4,000 km distance from their source areas. Additional terrigenous silt which is poor in stained quartz occurs within a narrow zone off the western Sahara between 20° and 27° N only. It depicts the present dust supply by the trade winds close to the surface. The dust load originates from the northwestern Sahara, the Atlas Mountains and coastal areas, which contain a particularly low amount of stained quartz. The distribution pattern of these pale quartz sediments reveals a SSW-dispersal of dust being consistent with the present trade wind direction from the NNE. In comparison to the sediments from off the Sahara and the deeper subtropical Atlantic, the sediments off river mouths, in particular off the Senegal river, are characterized by an additional input of fine grained terrigenous particles (< 6 µm). This is due to fluvial suspension load. The fluvial discharge leads to a relative excess of fine grained particles and is observed in a correlation diagram of the modal grain sizes of terrigenous silt with the proportion of fine fraction (< 6 µm). The aeolian sediment contribution by the Harmattan Winds strongly decreased during the Climatic Optimum at 6,000 y. B. P. The dust discharge of the trade winds is hardly detectable in the deep-sea sediments. This probably indicates a weakened atmospheric circulation. In contrast, the fluvial sediment supply reached a maximum, and can be traced to beyond Cape Blanc. Thus, the Saharan climate was more humid at 6,000 y B. P. A latitudinal shift of the Harmattan driven dust outbreaks cannot be observed. Also during the Glacial, 18,000 y. B. P., Harmattan dust transport crossed the African coast at latitudes of 15°-20° N. Its sediment load increased intensively, and markedly coarser grains spread further into the Atlantic Ocean. An expanded zone of pale-quart sediments indicates an enhanced dust supply by the trade winds blowing from the NE. No synglacial fluvial sediment contribution can be recognized between 12° and 30° N. This indicates a dry glacial climate and a strengthened stmospheric circulation over the Sahelian and Saharan region. The climatic transition pahes, at 12, 000 y. B. P., between the last Glacial and the Intergalcial, which is compareable to the Alerod in Europe, is characterized by an intermediate supply of terrigenous particles. The Harmattan dust transport wa weaker than during the Glacial. The northeasterly trade winds were still intensive. River supply reached a first postglacial maximum seaward of the Senegal river mouth. This indicates increasing humidity over the southern Sahara and a weaker atmospheric circulation as compared to the glacial. The accumulation rates of the terrigenous silt proportion (> 6 µm) decrcase exponentially with increasing distance from the Saharan coast. Those of the terrigenous fine fraction (< 6 µm) follow the same trend and show almost similar gradients. Accordingly, also the terrigenous fine fraction is believed to result predominantly from aeolian transport. In the Atlantic deep-sea sediments, the annual terrigenous sediment accumulation has fluctuated, from about 60 million tons p. a. during the Late Glacial (13,500-18,000 y. B. P, aeolian supply only) to about 33 million tons p. a. during the Holocene Climatic Optimum (6,000-9,000 y. B. P, mainly fluvial supply), when the river supply has reached a maximum, and to about 45 million tons p. a. during the last 4,000 years B. P. (fluvial supply only south of 18° N).