Nuclear markers reveal unexpected genetic variation and a Congolese-Nilotic origin of the Lake Victoria cichlid species flock

Phylogenetic analyses based on mitochondrial (mt) DNA have indicated that the cichlid species flock of the Lake Victoria region is derived from a single ancestral species found in East African rivers, closely related to the ancestor of the Lake Malawi cichlid species flock. The Lake Victoria flock contains ten times less mtDNA variation than the Lake Malawi radiation, consistent with current estimates of the ages of the lakes. We present results of a phylogenetic investigation using nuclear (amplified fragment length polymorphism) markers and a wider coverage of riverine haplochromines. We demonstrate that the Lake Victoria–Edward flock is derived from the morphologically and ecologically diverse cichlid genus Thoracochromis from the Congo and Nile, rather than from the phenotypically conservative East African Astatotilapia. This implies that the ability to express much of the morphological diversity found in the species flock may by far pre–date the origin of the flock. Our data indicate that the nuclear diversity of the Lake Victoria–Edward species flock is similar to that of the Lake Malawi flock, indicating that the genetic diversity is considerably older than the 15 000 years that have passed since the lake began to refill. Most of this variation is manifested in trans–species polymorphisms, indicating very recent cladogenesis from a genetically very diverse founder stock. Our data do not confirm strict monophyly of either of the species flocks, but raise the possibility that these flocks have arisen from hybrid swarms.

Persistence of Artemisia steppe in the Tangra Yumco Basin, west-central Tibet, China: despite or in consequence of Holocene lake-level changes?

The closed Tangra Yumco Basin underwent the strongest Quaternary lake-level changes so far recorded on the Tibetan Plateau. It was hitherto unknown what effect this had on local Holocene vegetation development. A 3.6-m sediment core from a recessional lake terrace at 4,700 m a.s.l., 160 m above the present lake level of Tangra Yumco, was studied to reconstruct Holocene flooding phases (sedimentology and ostracod analyses), vegetation dynamics and human influence (palynology, charcoal and coprophilous fungi analyses). Peat at the base of the profile proves lake level was below 4,700 m a.s.l. during the Pleistocene/Holocene transition. A deep-lake phase started after 11 cal ka BP, but the ostracod record indicates the level was not higher than similar to 4,720 m a.s.l. (180 m above present) and decreased gradually after the early Holocene maximum. Additional sediment ages from the basin suggest recession of Tangra Yumco from the coring site after 2.6 cal ka BP, with a shallow local lake persisting at the site until similar to 1 cal ka BP. The final peat formation indicates drier conditions thereafter. Persistence of Artemisia steppe during the Holocene lake high-stand resembles palynological records from west Tibet that indicate early Holocene aridity, in spite of high lake levels that may have resulted from meltwater input. Yet pollen assemblages indicate humidity closer to that of present potential forest areas near Lhasa, with 500-600 mm annual precipitation. Thus, the early mid-Holocene humidity was sufficient to sustain at least juniper forest, but Artemisia dominance persisted as a consequence of a combination of environmental disturbances such as (1) strong early Holocene climate fluctuations, (2) inundation of habitats suitable for forest, (3) extensive water surfaces that served as barriers to terrestrial diaspore transport from refuge areas, (4) strong erosion that denuded the non-flooded upper slopes and (5) increasing human influence since the late glacial.

Glacial forcing of central Indonesian hydroclimate since 60,000 y B.P.

The Indo-Pacific warm pool houses the largest zone of deep atmospheric convection on Earth and plays a critical role in global climate variations. Despite the region’s importance, changes in Indo-Pacific hydroclimate on orbital timescales remain poorly constrained. Here we present high-resolution geochemical records of surface runoff and vegetation from sediment cores fromLake Towuti, on the island of Sulawesi in central Indonesia, that continuously span the past 60,000 y.We show that wet conditions and rainforest ecosystems on Sulawesi present during marine isotope stage 3 (MIS3) and the Holocene were interrupted by severe drying between ∼33,000 and 16,000 y B.P. when Northern Hemisphere ice sheets expanded and global temperatures cooled. Our record reveals little direct influence of precessional orbital forcing on regional climate, and the similarity between MIS3 and Holocene climates observed in Lake Towuti suggests that exposure of the Sunda Shelf has a weaker influence on regional hydroclimate and terrestrial ecosystems than suggested previously. We infer that hydrological variability in this part of Indonesia varies strongly in response to high-latitude climate forcing, likely through reorganizations of the monsoons and the position of the intertropical convergence zone. These findings suggest an important role for the tropical western Pacific in amplifying glacial–interglacial climate variability.

Response of chironomid assemblages to environmental change during the early Late-glacial at Gerzensee, Switzerland

Prior to ca. 14,660 yr BP, during the early Late-glacial (Oldest Dryas), larval assemblages of Chironomidae (Insecta: Diptera) in Gerzensee, Switzerland, were dominated by cold stenothermic taxa as well as by taxa typical of subalpine lakes today. This was the coldest period of the entire sequence. After ca. 14,660 yr BP, in the Late Glacial Interstadial (Bølling–Allerød), a temperature increase is recorded by a sharp rise in the oxygen-isotope ratio in lake marl and by an increase in the organic-matter content of the sediments. Changes in the chironomid fauna then are consistent with rising temperatures. This warming trend is interrupted between 14,070 and 13,940 yr BP, coinciding with the GI-1d cold oscillation, but the change in the chironomid assemblage is more consistent with a response to increasing lake depth and density of aquatic macrophytes than falling temperature. A rise in cold-adapted chironomid taxa between 13,840 and 13,710 yr BP suggests that summer air temperatures may have declined. Changes in the chironomid assemblage after 13,710 yr BP suggest a decline in submerged macrophytes coupled with a rise in lake productivity and summer temperature, although the latter is not reflected in the oxygen-isotope record. This suggests that there may have been increasing seasonality during this period when summer temperatures were rising, driven by rising summer insolation, and winters becoming cooler, which is largely reflected in the oxygen-isotope record. A decline in thermophilic chironomids and a rise in cold-adapted taxa after 13,180 yr BP suggest a response to cooling at the beginning of the Gerzensee Oscillation.

Response of subfossil Cladocera in Gerzensee (Swiss Plateau) to early Late Glacial environmental change

Sub-fossil Cladocera were studied in a core from Gerzensee (Swiss Plateau) for the late-glacial periods of Oldest Dryas, Bølling, and Allerød. Cladocera assemblages were dominated by cold-tolerant littoral taxa Chydorus sphaericus, Acroperus harpae, Alonella nana, Alona affinis, and Alonella excisa. The rapid warming at the beginning of the Bølling (GI-1e) ca. 14,650 yr before present (BP: before AD 1950) was indicated by an abrupt 2‰ shift in carbonate δ18O and a clear change in pollen assemblages. Cladocera assemblages, in contrast, changed more gradually. C. sphaericus and A. harpae are the most cold-tolerant, and their abundance was highest in the earliest part of the record. Only 150–200 years after the beginning of the Bølling warming we observed an increase in less cold-tolerant A. excisa and A. affinis. The establishment of Alona guttata, A. guttata var. tuberculata, and Pleuroxus unicatus was delayed by ca. 350, 770, and 800 years respectively after the onset of the Bølling. The development of the Cladocera assemblages suggests increasing water temperatures during the Bølling/Allerød, which agrees with the interpretation by von Grafenstein et al. (2013-this issue) that decreasing δ18O values in carbonates in this period reflect increasing summer water temperatures at the sediment–water interface. Other processes also affected the Cladocera community, including the development and diversification of aquatic vegetation favourable for Cladocera. The record is clearly dominated by Chydoridae, as expected for a littoral core. Yet, the planktonic Eubosmina-group occurred throughout the core, with the exception of a period at ca. 13,760–13,420 yr BP. Lake levels reconstructed for this period are relatively low, indicating that the littoral location might have become too shallow for Eubosmina in that period.

Antarctic Glacial History Since the Last Glacial Maximum: An Overview of the Record on Land

This overview examines available circum-Antarctic glacial history archives on land, related to developments after the Last Glacial Maximum (LGM). It considers the glacial-stratigraphic and morphologic records and also biostratigraphical information from moss banks, lake sediments and penguin rookeries, with some reference to relevant glacial marine records. It is concluded that Holocene environmental development in Antarctica differed from that in the Northern Hemisphere. The initial deglaciation of the shelf areas surrounding Antarctica took place before 10000 C-14 yrs before present(sp), and was controlled by rising global sea level. This was followed by the deglaciation of some presently ice-free inner shelf and land areas between 10000 and 8000 yr sp. Continued deglaciation occurred gradually between 8000 yr sp and 5000 yr sp. Mid-Holocene glacial readvances are recorded from various sites around Antarctica. There are strong indications of a circum-Antarctic climate warmer than today 4700-2000 yr sp. The best dated records from the Antarctic Peninsula and coastal Victoria Land suggest climatic optimums there from 4000-3000 yr sp and 3600-2600 yr sp, respectively. Thereafter Neoglacial readvances are recorded. Relatively limited glacial expansions in Antarctica during the past few hundred years correlate with the Little Ice Age in the Northern Hemisphere.

Lake surface water temperatures of European Alpine lakes (1989–2013) based on the Advanced Very High Resolution Radiometer (AVHRR) 1 km data set

Lake water temperature (LWT) is an important driver of lake ecosystems and it has been identified as an indicator of climate change. Consequently, the Global Climate Observing System (GCOS) lists LWT as an essential climate variable. Although for some European lakes long in situ time series of LWT do exist, many lakes are not observed or only on a non-regular basis making these observations insufficient for climate monitoring. Satellite data can provide the information needed. However, only few satellite sensors offer the possibility to analyse time series which cover 25 years or more. The Advanced Very High Resolution Radiometer (AVHRR) is among these and has been flown as a heritage instrument for almost 35 years. It will be carried on for at least ten more years, offering a unique opportunity for satellite-based climate studies. Herein we present a satellite-based lake surface water temperature (LSWT) data set for European water bodies in or near the Alps based on the extensive AVHRR 1 km data record (1989–2013) of the Remote Sensing Research Group at the University of Bern. It has been compiled out of AVHRR/2 (NOAA-07, -09, -11, -14) and AVHRR/3 (NOAA-16, -17, -18, -19 and MetOp-A) data. The high accuracy needed for climate related studies requires careful pre-processing and consideration of the atmospheric state. The LSWT retrieval is based on a simulation-based scheme making use of the Radiative Transfer for TOVS (RTTOV) Version 10 together with ERA-interim reanalysis data from the European Centre for Medium-range Weather Forecasts. The resulting LSWTs were extensively compared with in situ measurements from lakes with various sizes between 14 and 580 km2 and the resulting biases and RMSEs were found to be within the range of −0.5 to 0.6 K and 1.0 to 1.6 K, respectively. The upper limits of the reported errors could be rather attributed to uncertainties in the data comparison between in situ and satellite observations than inaccuracies of the satellite retrieval. An inter-comparison with the standard Moderate-resolution Imaging Spectroradiometer (MODIS) Land Surface Temperature product exhibits RMSEs and biases in the range of 0.6 to 0.9 and −0.5 to 0.2 K, respectively. The cross-platform consistency of the retrieval was found to be within ~ 0.3 K. For one lake, the satellite-derived trend was compared with the trend of in situ measurements and both were found to be similar. Thus, orbital drift is not causing artificial temperature trends in the data set. A comparison with LSWT derived through global sea surface temperature (SST) algorithms shows lower RMSEs and biases for the simulation-based approach. A running project will apply the developed method to retrieve LSWT for all of Europe to derive the climate signal of the last 30 years. The data are available at doi:10.1594/PANGAEA.831007.

The stable carbon isotopic composition of Daphnia ephippia in small, temperate lakes reflects in-lake methane availability

Daphnia can ingest methane-oxidizing bacteria and incorporate methanogenic carbon into their biomass, leading to low stable carbon isotope ratios (expressed as δ13C values) of their tissue. Therefore, δ13C analysis of Daphnia resting eggs (ephippia) in lake sediment records can potentially be used to reconstruct past in-lake availability of methane (CH4). However, detailed multilake studies demonstrating that δ13C values of recently deposited Daphnia ephippia (δ13Cephippia) are systematically related to in-lake CH4 concentrations (CH4aq) are still missing. We measured δ13Cephippia from surface sediments of 15 small lakes in Europe, and compared these values with late-summer CH4aq. δ13Cephippia ranged from −51.6‰ to −25.9‰, and was strongly correlated with CH4aq in the surface water and above the sediment (r −0.73 and −0.77, respectively), whereas a negative rather than the expected positive correlation was found with δ13C values of carbon dioxide (CO2) (r −0.54), and no correlation was observed with CO2aq. At eight sites, offsets between δ13 CCO2 and δ13Cephippia exceeded offsets between δ13 CCO2 and δ13Calgae reported in literature. δ13Cephippia was positively correlated with δ13C values of sedimentary organic matter (r 0.54), but up to 20.7‰ lower in all except one of the lakes (average −6.1‰). We conclude that incorporation of methanogenic carbon prior to ephippia formation must have been widespread by Daphnia in our study lakes, especially those with high CH4aq. Our results suggest a systematic relationship between δ13Cephippia values and CH4aq in small temperate lakes, and that δ13Cephippia analysis on sediment records may provide insights into past changes in in-lake CH4aq.

Coupled human and natural system dynamics as key to the sustainability of Lake Victoria's ecosystem services

East Africa’s Lake Victoria provides resources and services to millions of people on the lake’s shores and abroad. In particular, the lake’s fisheries are an important source of protein, employment, and international economic connections for the whole region. Nonetheless, stock dynamics are poorly understood and currently unpredictable. Furthermore, fishery dynamics are intricately connected to other supporting services of the lake as well as to lakeshore societies and economies. Much research has been carried out piecemeal on different aspects of Lake Victoria’s system; e.g., societies, biodiversity, fisheries, and eutrophication. However, to disentangle drivers and dynamics of change in this complex system, we need to put these pieces together and analyze the system as a whole. We did so by first building a qualitative model of the lake’s social-ecological system. We then investigated the model system through a qualitative loop analysis, and finally examined effects of changes on the system state and structure. The model and its contextual analysis allowed us to investigate system-wide chain reactions resulting from disturbances. Importantly, we built a tool that can be used to analyze the cascading effects of management options and establish the requirements for their success. We found that high connectedness of the system at the exploitation level, through fisheries having multiple target stocks, can increase the stocks’ vulnerability to exploitation but reduce society’s vulnerability to variability in individual stocks. We describe how there are multiple pathways to any change in the system, which makes it difficult to identify the root cause of changes but also broadens the management toolkit. Also, we illustrate how nutrient enrichment is not a self-regulating process, and that explicit management is necessary to halt or reverse eutrophication. This model is simple and usable to assess system-wide effects of management policies, and can serve as a paving stone for future quantitative analyses of system dynamics at local scales.

Divergent Macroparasite Infections in Parapatric Swiss Lake-Stream Pairs of Threespine Stickleback (Gasterosteus aculeatus)

Spatial heterogeneity in diversity and intensity of parasitism is a typical feature of most host-parasite interactions, but understanding of the evolutionary implications of such variation is limited. One possible outcome of infection heterogeneities is parasite-mediated divergent selection between host populations, ecotypes or species which may facilitate the process of ecological speciation. However, very few studies have described infections in population-pairs along the speciation continuum from low to moderate or high degree of genetic differentiation that would address the possibility of parasite-mediated divergent selection in the early stages of the speciation process. Here we provide an example of divergent parasitism in freshwater fish ecotypes by examining macroparasite infections in threespine stickleback (Gasterosteus aculeatus) of four Swiss lake systems each harbouring parapatric lake-stream ecotype pairs. We demonstrate significant differences in infections within and between the pairs that are driven particularly by the parasite taxa transmitted to fish from benthic invertebrates. The magnitude of the differences tended to correlate positively with the extent of neutral genetic differentiation between the parapatric lake and stream populations of stickleback, whereas no such correlation was found among allopatric populations from similar or contrasting habitats. This suggests that genetic differentiation is unrelated to the magnitude of parasite infection contrasts when gene flow is constrained by geographical barriers while in the absence of physical barriers, genetic differentiation and the magnitude of differences in infections tend to be positively correlated.

Estimating population and livestock density of mobile pastoralists and sedentary settlements in the southeastern Lake Chad area

Mobile pastoralists provide major contributions to the gross domestic product in Chad, but little information is available regarding their demography. The Lake Chad area population is increasing, resulting in competition for scarce land and water resources. For the first time, the density of people and animals from mobile and sedentary populations was assessed using randomly defined sampling areas. Four sampling rounds were conducted over two years in the same areas to show population density dynamics. We identified 42 villages of sedentary communities in the sampling zones; 11 (in 2010) and 16 (in 2011) mobile pastoralist camps at the beginning of the dry season and 34 (in 2011) and 30 (in 2012) camps at the end of the dry season. A mean of 64.0 people per km2 (95% confidence interval, 20.3-107.8) were estimated to live in sedentary villages. In the mobile communities, we found 5.9 people per km2 at the beginning and 17.5 people per km2 at the end of the dry season. We recorded per km2 on average 21.0 cattle and 31.6 small ruminants in the sedentary villages and 66.1 cattle and 102.5 small ruminants in the mobile communities, which amounts to a mean of 86.6 tropical livestock units during the dry season. These numbers exceed, by up to five times, the published carrying capacities for similar Sahelian zones. Our results underline the need for a new institutional framework. Improved land use management must equally consider the needs of mobile communities and sedentary populations.

Local adaptations in populations of rock-dwelling haplochromines (Pisces:Cichlidae) from southern Lake Victoria

The radiation of cichlid fishes in the African great lakes is often described as adaptive, because, at a superficial level, cichlid fishes seem adapted to the ecological niches they occupy. However, adaptiveness has rarely been studied. We investigated to what extent island populations of three species of the rock-dwelling genus Neochromis, endemic to Lake Victoria, are adapted anatomically to exploit locally abundant resources. Specifically, we asked whether different resource environments were reflected in differences in the feeding apparatus, both within species and between species. In populations of two specialized biters, the algae scrapers N. rufocaudalis and N. omnicaeruleus, the biting force of the lower jaw increased with increasing amount of items that require biting in the diet. N. greenwoodi is a less specialized biter; we found differences between two populations in the hyoid position and in the premaxilla that enhance suction feeding. These adaptations were related to the amount of items requiring suction. Comparing across three sympatric pairs of species, in each case different diets were reflected in differences in anatomy.

Patterns of the Remnant Cichlid Fauna in Southern Lake Victoria. Patrones de la Fauna de Ciclidos Remanentes en el Sur del Lago Victoria

During the years 1984–1987 Lake Victoria in East Africa experienced what is probably the largest mass extinction of contemporary vertebrates. Within a decade about 200 endemic species of haplochromine cichlids disappeared. The extinctions that occurred in the 1980s have been documented predominantly on species of offshore and sub-littoral waters in the Mwanza Gulf of southern Lake Victoria. Although the littoral fauna of southern Lake Victoria had not been examined in detail, their diversity seemed less affected by the changes in the ecosystem. We give results of the first comprehensive inventory of the littoral cichlid fauna in southern Lake Victoria and discuss its conservation status. We also report on new developments in the sub-littoral fauna after 1990. More than 50 littoral and 15 sub-littoral stations were sampled between the years 1991 to 1995. Of the littoral stations, 34 were sampled for the first time. One hundred sixty three species of haplochromines were collected. Of these, 102 species were previously unknown. About two thirds of them live in rocky areas that were sampled for the first time. Littoral rocky habitats harbored the highest diversity. Since 1990, however, 13 more species disappeared from established sampling stations in littoral habitats. Fishing practices, spreading of exotic fishes, water hyacinth, and eutrophication are considered important threats to the littoral fauna. Also in the upper sub-littoral the number of species declined further. On deeper sub-littoral mud bottoms individual and species numbers increased again, although they are nowhere close to those found before the Nile perch (Lates niloticus) upsurge. This fauna differs from the well studied pre-Nile perch fauna. At well-established sampling stations, the sub-littoral zone is dominated by previously unknown species, and some known species have performed dramatic habitat shifts.

The 1985 catastrophic drainage of a moraine-dammed lake, Khumbu Himal, Nepal: cause and consequences

On 4 August 1985 Dig Tsho, a moraine-dammed glacial lake in the Khumbu area of eastern Nepal, burst above Thame. For the region close to the origin of the outbreak the consequences were catastrophic. The destruction of a newly built hydroelectricp ower plant, 14 bridges, about 30 houses, and many hectares of valuable arable land, as well as a heavily damaged trail network, resulted from 5 million m3 of water plummetting down the Bhote Kosi and Dudh Kosi valleys. The breaching of the moraine was triggered by wave action following an ice avalanche of 150,000 m3 into the lake. The surge had a peak discharge of 1,600 m3/sec; 3 million m3 of debris were moved within a distance of less than 40 km. However, only 10-15 percent of the material left the region as suspended load. The potential hazard of glacial lakes persists and increases. A hazard assessment including an identificationo f source areas and subsequent monitoring of glacial lakes is proposed. It should be incorporated into any development concept for the Himalayan region.

Depositional modes and lake-level variability at Lake Towuti, Indonesia, during the past ~29 kyr BP

Lake Towuti (2.5°S, 121.5°E) is a long-lived, tectonic lake located on the Island of Sulawesi, Indonesia, and in the center of the Indo-Pacific warm pool (IPWP). Lake Towuti is connected with upstream lakes Matano and Mahalona through the Mahalona River, which constitutes the largest inlet to the lake. The Mahalona River Delta is prograding into Lake Towuti’s deep northern basin thus exerting significant control on depositional processes in the basin. We combine high-resolution seismic reflection and sedimentological datasets from a 19.8-m-long sediment piston core from the distal edge of this delta to characterize fluctuations in deltaic sedimentation during the past ~29 kyr BP and their relation to climatic change. Our datasets reveal that, in the present, sedimentation is strongly influenced by deposition of laterally transported sediments sourced from the Mahalona River Delta. Variations in the amount of laterally transported sediments, as expressed by coarse fraction amounts in pelagic muds and turbidite recurrence rates and cumulative thicknesses, are primarily a function of lake-level induced delta slope instability and delta progradation into the basin. We infer lowest lake-levels between ~29 and 16, a gradual lake level rise between ~16 and 11, and high lake-levels between ~11 and 0 kyr BP. Periods of highest turbidite deposition, ~26 to 24 and ~18 to 16 kyr BP coincide with Heinrich events 2 and 1, respectively. Our lake-level reconstruction therefore supports previous observations based on geochemical hydroclimate proxies of a very dry last glacial and a wet Holocene in the region, and provides new evidence of millennial-scale variations in moisture balance in the IPWP.