Sub-fossil Chironomidae from East Africa. 1. Tanypodinae and Orthocladiinae

This paper and a companion article present illustrated guides to the identification of sub-fossil chironomid larvae (Insecta: Diptera: Chironomidae) preserved in the sediments of low- and mid-elevation lakes in East Africa. They are based on analysis of surface-sediment death assemblages from 61 lakes located in the humid to semi-arid environments of equatorial East Africa (Uganda, Kenya, Tanzania), supplemented with similar surface-sediment samples from 12 lakes in the Horn of Africa (Ethiopia), and sub-recent core samples from six lakes in Kenya and two in Uganda. We analyzed about 11,000 specimens and identified 98.4% of these to species, species group, genus, or tribe level depending on current -taxonomic knowledge of the various genera considered and the taxonomic resolution of preserved diagnostic features. We distinguished 90 different sub-fossil morphotypes, of which 16 are Tanypodinae, 19 are Orthocladiinae, and 55 are Chironominae. Diagnostic characters distinguishing these morphotypes from each other resemble differences at the species level in the better-known Holarctic fauna, hence we consider most of our morphotypes equivalent to morphological species or groups of closely related species. In this paper we focus on the Tanypodinae and Orthocladiinae, with special attention to the high taxon richness among the Pentaneurini. Patterns of cephalic setation were found to facilitate identification of Tanypodinae both at the genus and species level, and contributed to improved taxonomic resolution in sub-fossil East African material. High taxon richness and numerical abundance of the Orthocladiinae in our study lakes indicates that a considerable number of African Orthocladiinae is adapted to warm standing-water environments.     

Subfossil Chironomidae from Lake Tanganyika, East Africa. 2. Chironominae (Chironomini and Tanytarsini)

Analysis of subfossil remains of larval Chironomidae in 38 surface-sediment samples from between 53 and 189 meter depth in Lake Tanganyika (East Africa) yielded 77 morphotypes, among which 7 Tanypodinae, 19 Orthocladiinae, and 51 Chironominae. Character-state differences between these morphotypes resemble differences at the species level in the better-known Holarctic fauna, hence we consider most of our Lake Tanganyika morphotypes equivalent to morphological species. Individual morphotypes were identified to species, genus, or tribe level depending on current alpha-taxonomic knowledge on the larvae of the group concerned, and the taxonomic resolution of preserved diagnostic features. This paper presents taxon diagnoses and an illustrated key to the Chironominae (Chironomini and Tanytarsini) in this collection. As the living chironomid fauna of Lake Tanganyika has never been comprehensively studied, we also briefly discuss faunistic aspects, and the ecology of the recovered species and genera in relation to benthic habitat diversity in Lake Tanganyika.     

Subfossil Chironomidae from Lake Tanganyika, East Africa 1. Tanypodinae and Orthocladiinae

Analysis of subfossil remains of larval Chironomidae in 38 surface-sediment samples from between 53 and 189 m depth in Lake Tanganyika (East Africa) yielded 77 different morphotypes, including 7 Tanypodinae, 19 Orthocladiinae, and 51 Chironominae. Character-state differences between these morphotypes resemble differences at the species level in the better-known Holarctic fauna, hence we consider most of our Lake Tanganyika morphotypes equivalent to morphological species. Individual morphotypes were identified to species, genus, or tribe level depending on current alpha-taxonomic knowledge on the larvae of the group concerned, and the taxonomic resolution of preserved diagnostic features. This paper presents taxon diagnoses and an illustrated key to the Tanypodinae and Orthocladiinae in this Lake Tanganyika collection, with the aim to promote consistency of identification in studies using African Chironomidae as biological indicators of natural and anthropogenic environmental change in lacustrine ecosystems.     

Paleolimnology of Lake Tanganyika, East Africa, over the past 100 kyr

New sediment core data from a unique slow-sedimentation rate site in Lake Tanganyika contain a much longer and continuous record of limnological response to climate change than have been previously observed in equatorial regions of central Africa. The new core site was first located through an extensive seismic reflection survey over the Kavala Island Ridge (KIR), a sedimented basement high that separates the Kigoma and Kalemie Basins in Lake Tanganyika.Proxy analyses of paleoclimate response carried out on core T97-52V include paleomagnetic and index properties, TOC and isotopic analyses of organic carbon, and diatom and biogenic silica analyses. A robust age model based on 11 radiocarbon (AMS) dates indicates a linear, continuous sedimentation rate nearly an order of magnitude slower here compared to other core sites around the lake. This age model indicates continuous sedimentation over the past 79 k yr, and a basal age in excess of 100 k yr.The results of the proxy analyses for the past 20 k yr are comparable to previous studies focused on that interval in Lake Tanganyika, and show that the lake was about 350 m lower than present at the Last Glacial Maximum (LGM). Repetitive peaks in TOC and corresponding drops in ¹³C over the past 79 k yr indicate periods of high productivity and mixing above the T97-52V core site, probably due to cooler and perhaps windier conditions. From 80 through 58 k yr the ¹³C values are relatively negative (–26 to –28 l) suggesting predominance of algal contributions to bottom sediments at this site during this time. Following this interval there is a shift to higher values of ¹³C, indicating a possible shift to C-4 pathway-dominated grassland-type vegetation in the catchment, and indicating cooler, dryer conditions from 55 k yr through the LGM. Two seismic sequence boundaries are observed at shallow stratigraphic levels in the seismic reflection data, and the upper boundary correlates to a major discontinuity near the base of T97-52V. We interpret these discontinuities to reflect major, prolonged drops in lake level below the core site (393 m), with the lower boundary correlating to marine oxygen isotope Stage 6. This suggests that the previous glacial period was considerably cooler and more arid in the equatorial tropics than was the last glacial period.     

Spatial variability of chironomid death assemblages in the surface sediments of a fluctuating tropical lake (Lake Naivasha,Kenya)

Studies addressing within-lake variability of fossil chironomid assemblages are very few, and all deal with hydrologically stable temperate lakes where the question of spatial integration mostly relates to the mixing of faunal assemblages associated with shallow, warm-water habitat and those associated with deeper, cold-water habitat. Here we study within-lake variability of surface-sediment chironomid assemblages in the fairly large (∼100–170 km² since 1983) and shallow (Z _max = 5–8 m) fluctuating tropical lake basin of Lake Naivasha, Kenya, and compare the patterns observed with those in two smaller adjacent basins, one similarly shallow (Lake Oloidien, 5.1–5.7 km2, 5–8 m), the other deep and stratified (Crescent Island Crater, 1.9 km², 14–17 m). Chironomid assemblages were analysed in core-top samples and surface sediments along inshore to offshore transects, and how well individual samples represented the total (basin-wide mean) subfossil assemblage was considered both in terms of taxon richness and taxon percent composition. Within-lake variability of subfossil chironomid concentrations (with generally higher absolute values in nearshore samples) could be explained by effects of sediment winnowing and focusing, whereas between-lake variability reflected their relative susceptibility to wind-driven sediment disturbance or bottom anoxia. In all study lakes, but most significantly in lakes Naivasha and Oloidien, species distribution in the subfossil chironomid assemblages showed a strong nearshore to offshore gradient, which in these shallow lakes, reflects the dominant control of substrate and food quality on species distribution in the living community. Particularly in the larger basins, nearshore samples better represented the total lake assemblage than offshore samples, because the former always contained a component of mud-dwelling species whereas the latter often lacked a component of macrophyte-dwelling species. Our results show that although sedimentation dynamics in the shallow, wind-stressed Lake Naivasha is dominated by frequent resuspension and random sediment redistribution, the near- to offshore gradient in chironomid habitat remains imprinted on subfossil assemblages. We conclude that also in shallow fluctuating lakes, given sufficient size, incomplete pre-burial spatial integration of habitat-specific chironomid assemblages can be exploited for within-lake calibration of environmental gradients.     

Mass occurance and sporadic distribution of Corynocera ambigua Zetterstedt (Diptera, Chironomidae) in Danish lakes. Neo- and palaeolimnological records

The chironomid Corynocera ambigua (Tanytarsini) is commonly reported as a cold-stenothermal species living in shallow lakes in arctic and subarctic regions. In palaeoecological studies of temperate lakes, larval remains of C. ambigua are usually found in late-glacial sediments from the Allerød and Younger Dryas periods, and often in association with subfossil Chara oospores. During a surface sampling program of chironomid head-capsules in 41 Danish temperate lakes, C. ambigua was found to comprise 25% of the chironomid assemblages in two lakes, and was sporadically found in 8 other lakes (0.5-10%). A 70 cm palaeo-stratigraphy from the shallow (max depth 1.2 m) and eutrophic (total phosphorus = 150 g P l^-1) Lake Stigsholm showed that C. ambigua has been abundant in the last 4-5 centuries. At a sediment level of 25 cm (~year 1925, ²¹⁰Pb dating), C. ambigua began to decrease in frequency while Chironomus plumosus, Procladius sp., Cladotanytarsus gr. mancus and Tanytarsus spp. increased, suggesting an increased nutrient loading and an approach to eutrophic conditions. In 1995 C. ambigua was still very abundant in Lake Stigsholm but in early March 1997 no living larvae were found. An extremely heavy growth of Elodea, Enteromorpha and filamentous algae in the summers of 1995 and 1996, with following degradation in the fall, might have influenced the invertebrate population dynamics. No significant distinguishing characteristics were found for the lakes supporting C. ambigua. Its occurrence in warm (~20°C) Danish lakes brings into question the perception of the species as being cold-stenothermal.     

Application of the USLE in a Savannah Environment: Comparative Experiences from East and West Africa

The use of empirical models for predicting erosion hazard in Africa is widespread. One of the most commonly used models is the Universal Soil Loss Equation (USLE). This paper explores the applicability of the USLE in the East and West African Savannah environments, using both primary and secondary data. The results show that the USLE can be applied to Savannah conditions if some level of instrumentation is provided to enable model calibration and validation. The rainfall erosivity factor (R) is one variable which is calculated differently in both the East and the West African Savannah, reflecting differences in the storm characteristics between the two regions. The soil erodibility factor (K) is applied to cultivated soils with relatively high organic matter but does not adequately predict soil loss if the soils have large quantities of stone cover or if they exhibit surface sealing properties. The topographic factors (LS) are transferable, but in some parts of the Savannah, very steep slopes are cultivated, limiting the accuracy of the LS factor. The crop and management factor (C) is applicable for single stand crops, but more studies are required to develop C factors for multiple cropping and rangelands as these are more prevalent in the Savannah. The conservation practices factors (P) as described in the USLE routines do not adequately cater for soil conservation structures found in Africa, such as trash lines and stone lines. Thus, local indices need to be developed. In general, the USLE tends to underestimate soil erosion in East Africa while overestimating it in the West African Savannah. User‐friendly Geographical Information Systems (GIS) applications that depict the uncertainty of model estimates should be explored in the future.     

A protocol for preparing subfossil chironomid head capsules (Diptera: Chironomidae) for stable isotope analysis in paleoclimate reconstruction and considerations of contamination sources

Several techniques are available to examine the isotopic composition of historic lake waters, providing data that can subsequently be used to examine environmental changes. A recently-developed technique is the stable oxygen isotope analysis of subfossil chironomid (Diptera: Chironomidae) head capsules (mostly chitin) preserved in lake sediments. This technique involves a high Temperature Conversion Elemental Analyzer (TC/EA), which has been a relatively recent addition to the suite of online peripherals for analyzing the stable isotopic composition of organic samples. The highly precise and accurate ¹⁸O/¹⁶O and D/H measurements obtainable using the TC/EA with samples in the microgram range make this instrumentation suitable for studying geochemical and biological processes. Preparation of organic samples for isotopic analysis typically requires first weighing each sample into silver/tin capsules. These capsules can introduce oxygen and hydrogen contamination (a “blank effect”), which is especially problematic for analysis of small organic samples (e.g. less than 100 μg). Here we tested tin and silver capsules from two manufacturers and a range of sizes to assess contamination to small organic samples on the TC/EA. We also assessed how a method for cleaning silver capsules affected our analysis of commercial chitin. In general, capsules made of silver have less detectible oxygen than those made of tin, and capsules from the two manufacturers varied in their detectible oxygen. There was no detectable H contamination from silver capsules. In addition to our empirical findings, we present a model demonstrating the influence that contaminant oxygen can have on the δ¹⁸O of small organic samples. Sample mass becomes an important issue for such analyses. In light of our findings, we recommend a minimum sample mass ≥50 μg (approximately 120 whole chironomid head capsules) on a TC/EA-IRMS (Delta^plus XP system). Finally, we present a detailed protocol for preparing and transferring chironomid head capsules into silver capsules that minimizes the influence of contaminant oxygen. This protocol provides the paleo-community with another potential method for reconstructing paleoenvironments.     

Quaternary and extant euryhaline Lamprothamnium Groves (Charales) from Australia: Gyrogonite morphology and paleolimnological significance

Charophytes are very common in Australian modern and Quaternary waterbodies, and are quite commonly incorrectly reported as “Chara” sp. or Lamprothamnium papulosum (Wallroth) Groves. This paper is the first attempt at the identification of the widespread euryhaline genus Lamprothamnium in Australia, and its use as a paleoenvironmental indicator. Lamprothamnium is distributed worldwide in all continents, except north and central America. The Australian environment, characterized by increasing aridity during the last 500 ka, has an abundance of saline lakes. We sampled 30 modern lakes and identified extant Lamprothamnium macropogon (A. Braun) Ophel and Lamprothamnium succinctum (A. Braun in Ascherson) Wood. Fossil gyrogonites, from lacustrine sediments ∼65 ka old from Madigan Gulf, Lake Eyre, were identified as Lamprothamnium williamsii sp. nov. We applied statistical analysis (analysis of variance, ANOVA) to the morphometry of the gyrogonites from one fossil and three living Lamprothamnium populations. The ANOVA test suggests all the populations are different, including two separate populations of extant L. macropogon, interpreted in this case as the expression of ecophenotypic variability. Lamprothamnium is a useful paleoenvironmental indicator because it indicates a non-marine environment with varying salinity ranging from fresh (usually 2–3 g l⁻¹) to 70 g l⁻¹, and water bodies holding water for at least 70 consecutive days. Collectively, these parameters provide important information in the study of ephemeral habitats.     

Solar and anthropogenic imprints on Lake Masoko (southern Tanzania) during the last 500 years

The Masoko crater-lake in southern Tanzania provides a continuous record of environmental changes covering the last 500 years. Multi-proxy studies were performed on a 52 cm sediment core retrieved from the deepest part of the lake. Magnetic, organic carbon, geochemical proxies and pollen assemblages indicate a dry climate during the ‘Little Ice Age’ (AD 1550–1850), confirming that the LIA in eastern Africa resulted in marked and synchronous hydrological changes. However, the direction of response varies between different African lakes (low versus high lake-levels), indicating strong regional contrasts that prevent the clear identification of climate trends over eastern Africa at this time. Inferred changes in Masoko lake-levels closely resemble the record of solar activity cycles, indicating a possible control of solar activity on the climate in this area. This observation supports previous results from East African lakes, and extends this relationship southward. Finally, anthropogenic impact is observed in the Masoko sediments during the last 60 years, suggesting that human disturbance significantly affected this remote basin during colonial and post-colonial times.     

A 40,000-year record of environmental change from ancient Lake Ohrid (Albania and Macedonia)

Lake Ohrid is considered to be of Pliocene origin and is the oldest extant lake in Europe. A 1,075-cm-long sediment core was recovered from the southeastern part of the lake, from a water depth of 105 m. The core was investigated using geophysical, granulometric, biogeochemical, diatom, ostracod, and pollen analyses. Tephrochronology and AMS radiocarbon dating of plant macrofossils reveals that the sediment sequence spans the past ca. 39,500 years and features a hiatus between ca. 14,600 and 9,400 cal. year BP. The Pleistocene sequence indicates relatively stable and cold conditions, with steppe vegetation in the catchment, at least partial winter ice-cover of the lake, and oxygenated bottom waters at the coring site. The Holocene sequence indicates that the catchment vegetation had changed to forest dominated by pine and summer-green oak. Several of the proxies suggest the impact of abrupt climate oscillations such as the 8.2 or 4.0 ka event. The observed changes, however, cannot be related clearly to a change in temperature or humidity. Human impact started about 5,000 cal. year BP and increased significantly during the past 2,400 years. Water column mixing conditions, inflow from subaquatic springs, and human impact are the most important parameters influencing internal lake processes, notably affecting the composition and characteristics of the sediments.     

Diatoms and sockeye salmon (Oncorhynchus nerka) population dynamics: Reconstructions of salmon-derived nutrients over the past 2,200 years in two lakes from Kodiak Island, Alaska

The return of hundreds to millions of adult sockeye salmon (Oncorhynchus nerka), which have returned from the ocean to their natal nursery lake environment to spawn, can result in significant nutrient loading. By analyzing sedimentary diatom assemblages from nursery lakes, we demonstrated that a salmon-derived nutrient signal could be traced over time and be used to infer past sockeye salmon population dynamics. We conducted a 2,200 year paleolimnological study of two Alaskan sockeye salmon nursery lakes, Karluk and Frazer lakes. The two lakes are very similar, except that sockeye salmon were only introduced into Frazer Lake in 1951 (first spawners returned in 1956). In both lakes we found a strong correspondence between diatom assemblages and the number of adult salmon spawners recorded in the historical data (40 and 70 years for Frazer and Karluk lakes, respectively). Given this robust relationship, we then used our analyses of diatoms from Karluk Lake over the past 2,200 years to gain insight into salmon-derived nutrient loading changes (which are directly related to the number of sockeye salmon spawners). The diatom record from Karluk Lake recorded dramatic species changes on both decadal and century timescales, and was strongly correlated with an independent indicator of sockeye salmon abundances, ¹⁵N. Together, these data suggest pronounced variability in sockeye salmon abundances at Karluk Lake over the past 2,200 years. The direct impacts of regional environmental variability were not likely responsible for the patterns apparent in Karluk Lake, as the diatom and ¹⁵N profiles from Frazer Lake were relatively stable prior to the introduction of sockeye salmon. Application of total phosphorus transfer functions to the Karluk and Frazer lakes' diatom records revealed that sockeye salmon carcasses substantially increased the trophic status in these lakes, which has important implications for the health of juvenile salmon that rear in nursery lakes. Overall, this paper illustrates the potential use of diatoms in reconstructing past sockeye salmon population dynamics, which in turn can lead to a greater understanding of the mechanisms influencing abundances of sockeye salmon.     

Holocene Environmental Changes of Lake Geneva (Lac Léman) from Stable Isotopes (δ13C, δ18O) and Trace Element Records of Ostracod and Gastropod Carbonates

Stable isotopes and trace-element contents of ostracod (Candona neglecta) valves mostly from the Holocene portion of two assembled cores from Petit Lac (Lake Geneva, Switzerland-France) were analysed in order to depict the geochemical record of post-glacial environmental changes of this lake. Additional stable isotope and trace element data from the gastropod Bithynia tentaculata (shells and opercula) from some intervals of these cores, as well as previous data from bulk carbonate from the lower part of the studied intervals were also considered. Mg/Ca and Sr/Ca molar ratios for the Holocene lake water have been estimated from evaluations of the partitioning coefficients for Mg and Sr for C. neglecta and B. tentaculata taking into account the modern-lake water composition. This study shows an overall gentle trend to higher δ¹⁸O values in C. neglecta valves from the Boreal interval (mean −8.44‰) to the upper part of the core (mean −8.11‰). This trend is superimposed to higher frequency oscillations of stable isotope values and trace element ratios, especially through the upper Older Atlantic and the Subboreal. The overall isotopic oxygen trend includes several shifts in δ¹⁸O of about 1‰. These shifts are interpreted as major regional-global climate changes that have also been observed in other coeval δ¹⁸O and pollen records which reflect the Holocene climate variability in other European basins. Especially well-defined peaks in some episodes like Older Atlantic (~8200 yr BP), Younger Atlantic – Subboreal transition (~5600 yr BP) and early Subatlantic (~ 2500 yr BP) correspond to well-recognized events in globally-distributed records. Some of these shifts are correlated with pulses in the lake-level curve of the Lake Geneva. An erratum to this article is available at .     

Establishment of tree-ring chronology and climatic response of Tibetan juniper (S. tibetica) in south Tibet, western China

A tree-ring width chronology of 442 years(1567-2008) was developed from Tibetan junipers(S.tibetica) derived from south Tibet in western China.Three versions of chronology were produced according to standard dendrochronological techniques.The correlation and response analysis displays a high correlation between the standard tree ring-width chronology and observed annual mean precipitation series during the period 1961-2008.Based on a linear regression model,an annual(prior August to current July) precipitation for the past 229 years was reconstructed.This is the first well-calibrated precipitation reconstruction for the Nanggarze region,south Tibet.The results show that relatively wet years with above-average precipitation occurred in 1780-1807,1854-1866,1886-1898,1904-1949,1967-1981 and 2000-2008,whereas relatively dry years with below-average precipitation prevailed during 1808-1853,1867-1885,1899-1903,1950-1966 and 1982-1999.Common dry/wet periods during 1890s,1910s,1940s-1960s and 1980s were also identified from other moisture reconstructions of nearby regions,indicating a synchronous climatic variation in south Tibet.Abrupt change beginning in 1888 was detected,revealing a transition from wet to dry conditions in south Tibet.Power spectrum analysis reveals significant cycles of 28-year,5.5-5.6-year and 3.3-year during the past 200 years.     

1960—2017年太湖流域不同等级降水时空特征

基于太湖流域1960—2017年逐日降水数据,运用Mann-Kendall非参数检验法、R/S分析等方法,分析太湖流域不同等级降水的时空变化特征,并探讨了不同等级降水对年降水的影响。结果表明：1）近60年来,流域小雨发生率最高,为73.55%;年总降水量中,中雨量所占比例最大,为32.05%。小雨发生率呈显著减少趋势,暴雨贡献率呈显著增加趋势。2）太湖流域大雨、暴雨的降水量和降水日数都呈显著增加,小雨日数显著减少,小雨强度、年总降水强度显著增强。3）不同等级降水变化趋势的空间分布存在明显差异。小雨日数与年总降水日数,以及小雨强度与年总降水强度的变化趋势空间格局相一致。中雨日数、大雨日数、暴雨日数变化趋势的空间分布与其对应的降水量变化趋势的空间格局相似。4）R/S分析结果显示,小雨、暴雨、年总降水相关指标（小雨量除外）都表现出较强的持续性,未来变化趋势与过去相一致。5）近60年来,太湖流域年总降水量、降水日数、年总降水强度的变化,分别受中雨量、小雨日数、暴雨量的影响较大。在旱年流域年降水量偏少受大雨量减少的影响较大,而涝年年降水量偏多受暴雨增加的影响较大。     

Lake Shira, a Siberian salt lake: ecosystem structure and function. 2. Brachionus plicatilis (Rotatoria) growth and development

The growth and development of Brachionus plicatilis (Müller), a rotifer and one of the most abundant zooplankton species in Lake Shira, a saline lake at low temperature in the Republic of Khakassia, was studied in cultures at different trophic conditions. Growth indices were evaluated at natural (lake) concentrations of bacteria, Chlorella vulgaris, a microalga, and Candida utilis, a yeast. The negative effect on rotifer reproduction in the algal-bacterial community by blue-green algae was demonstrated. The ratio of ontogenetic periods was shown to be in accord with species and food concentration. Base values required to calculate B. plicatilis population simulation model parameters were defined.