Summer Rains and Dry Seasons in the Upper Blue Nile Basin: The Predictability of Half a Century of Past and Future Spatiotemporal Patterns

During the last 100 years the Ethiopian upper Blue Nile Basin (BNB) has undergone major changes in land use, and is now potentially facing changes in climate. Rainfall over BNB supplies over two-thirds of the water to the Nile and supports a large local population living mainly on subsistence agriculture. Regional food security is sensitive to both the amount and timing of rain and is already an important political challenge that will be further complicated if scenarios of climate change are realized. In this study a simple spatial model of the timing and duration of summer rains (Kiremt) and dry season (Bega), and annual rain over the upper BNB was established from observed data between 1952 and 2004. The model was used to explore potential impacts of climate change on these rains, using a down-scaled ECHAM5/MP1-OM scenario between 2050 and 2100. Over the observed period the amount, onset and duration of Kiremt rains and rain-free Bega days have exhibited a consistent spatial pattern. The spatially averaged annual rainfall was 1490 mm of which 93% was Kiremt rain. The average Kiremt rain and number of rainy days was higher in the southwest (322 days) and decreased towards the north (136 days). Under the 2050–2100 scenario, the annual mean rainfall is predicted to increase by 6% and maintain the same spatial pattern as in the past. A larger change in annual rainfall is expected in the southwest (ca. +130 mm) with a gradually smaller change towards the north (ca. +70 mm). Results highlight the need to account for the characteristic spatiotemporal zonation when planning water management and climate adaptation within the upper BNB. The presented simple spatial resolved models of the presence of Kiremt and annual total rainfall could be used as a baseline for such long-term planning.     

Spawning periodicity and annual breeding seasons of Tilapia nilotica (L.) in the White Nile

Two annual breeding seasons are indicated for populations of T. nilotica in the White Nile: a major autumnal season (mid-July–September) coinciding with the annual rainfall and a smaller back-up season in February–March. During these seasons reproducing females constituted 80% and 40% of the total population of mature females compared to 10–12% in the intervening summer period. Enhanced gonadal development, fecundity and GSI values characterized the breeding seasons. Gonadal maturation proceeded uninterruptedly even during the summer and repeated fractional spawning is suggested for this species within and beyond the breeding seasons. Most of the spring breeders were either very young or relatively old fish and most medium-sized females bred in the main autumnal season. Older fish were less fecund (by 70–75%) compared to younger fish. In the latter, fecundity increased progressively with body growth (r² = 0.732; p<0.001); the proportionality was lost in medium-sized fish and in older females (0.9–1.5 kg) fecundity correlated negatively with increased body size.     

The reproductive biology and early development of the African pike,Hepsetus odoe,in the Okavango Delta,Botswana

Synopsis The reproductive biology of the African pike,Hepsetus odoe, was studied over a four-year period in the Okavango Delta, Botswana. Spawning takes place between August and May in different parts of the Delta. Fish in the seasonal swamp and drainage rivers have a truncated spawning season between August and January, while those in the perennial swamp have an extended spawning season from September to May. The variation in spawning season appears to be associated with the degree of environmental perturbation brought about by the annual flood cycle. Male and female pike mature at 140 and 160 mm standard length (SL) respectively. Females predominate in the population and produce an average of 6440 eggs per season and are multiple spawners. The number of eggs per spawning averages about 2630. The eggs are deposited in a foam nest which is guarded by the parents. Newly hatched embryos suspend themselves below the nest via a cement gland and remain in the vicinity of the spawning site until they have reached a relatively advanced stage in their development. The larval period is relatively short which implies that this species undergoes nearly a direct development. Foam nests appear to be a predator avoidance mechanism as well as an adaptation to fluctuating oxygen and water levels. The initiation of the spawning season is more closely associated with water temperature than with the annual flood cycle which appears to be more important in regulating the size of the spawning stock.     

Shifts in the food of Nile tilapia, Oreochromis niloticus (L.) in Lake Victoria, Kenya

Studies of the food of introduced Nile tilapia, Oreochromis niloticus (L.) with respect to size, habitat and season were conducted between November 1998 and October 2000 in Kenyan waters of Lake Victoria. Stomach contents of 1980 specimens collected by demersal trawl and seining were analysed. Nile tilapia originally known to be herbivorous, feeding mostly on algae has diversified its diet to include insects, fish, algae and plant materials. The major diet of fish <5 cm total length was zooplankton whereas bigger fish included a wider range of food items in their diet. There was spatial variation in diet with insects and algae dominating in the gulf and open water habitats respectively. There was no seasonal variation in the food items ingested and diel feeding regime indicated that O. niloticus is a diurnal feeder. The shift in diet could be due to ecological and environmental changes in Lake Victoria, which have been associated with changes in composition and diversity of fish and invertebrate fauna, emergence and dominance of different flora including water hyacinth Eichhornia crassipes (Mart.) Solms‐Laub., and algae communities. The feeding habit of O. niloticus is discussed in the context of changes occurring in the lake.     

西南干旱对哀牢山常绿阔叶林凋落物及叶面积指数的影响

为探讨2010年初西南干旱对这一地区原生植被林冠和凋落物量的影响,以及这一地区凋落物量和气候条件之间的关系,对比研究了哀牢山亚热带常绿阔叶林2010年和一般年份的凋落物特征以及各层的叶面积指数,并分析了凋落物量和气候因子之间的关系.2010年凋落物总量和往年相比无显著差异,但是叶凋落总量、旱季凋落物总量、旱季叶凋落量为历年来最高,其中旱季叶凋落物量比一般年份平均高35.2％ (0.81 t/hm2).而2010年附生苔蘚年凋落量为历年来最低.2010年最旱月的乔木层和灌木层叶面积指数和2005同期相比无显著差异,但是草本层叶面积指数却极显著低于2005年同期.因此,2010年初西南干旱尽管使哀牢山常绿阔叶林旱季落叶增加,但还没有到显著影响冠层叶面积指数的程度.而草本层和附生苔藓的生长则受到了干旱事件的显著影响.此外,哀牢山常绿阔叶林年总凋落量和年降水量显著正相关,而和年均温却不相关,表明该亚热带森林凋落物量主要由降水而非温度决定.     

Fish species and trophic diversity of haplochromine cichlids in the Kyoga satellite lakes (Uganda)

Prior to the 1980s, lakes Kyoga and Victoria previously supported an exceptionally diverse haplochromine fish fauna comprising at least 11 trophic groups. The species and trophic diversity in these lakes decreased when the introduced Nile perch depleted haplochromine stocks. From December 1996 to October 1998, we studied species and trophic diversity of haplochromine fishes in six satellite lakes without Nile perch in the Kyoga basin and compared them with the Kyoga main lake against historical data from Lake Victoria where Nile perch were introduced. Forty‐one species were found in the study area, of which, the Kyoga satellite lakes contributed 37 species in comparison to only 14 from the Kyoga main lake. Analysis of trophic diversity based on 24 species that contained food material revealed seven haplochromine trophic groups (insectivores, peadophages, piscivores, algal eaters, higher plant eaters, molluscivores and detritivores) in the Kyoga satellite lakes in comparison to two trophic groups (insectivores and molluscivores) in the Kyoga main lake. Many of the species and trophic groups of haplochromines depleted by the introduced Nile perch in lakes Kyoga and Victoria still survive in the Kyoga satellite lakes. This is attributed to the absence of Nile perch in those lakes. Nile perch has been prevented from spreading into the satellite lakes by swamp vegetation that separate them from the main lakes. If these swamps prevent Nile perch from spreading into the lakes, it is possible to conserve fish species, especially haplochromines, which are threatened by introduction of Nile perch in the main lakes.     

Aquatic invertebrate fauna of wells in a tropical mountain climate,western Cameroon

The present study, conducted between May 2012 and September 2013, aimed to determine the distribution of groundwater invertebrates in the Bamoun tableland, Cameroon. A total of 216 samples taken from 30 wells in four localities – Foumban, Foumbot, Kouoptamo and Magba – contained 80 invertebrate taxa belonging to Crustacea, Oligochaeta, Arachnida, Insecta, Nematoda, Tardigrada, and Gastropoda. The community was relatively rich, diverse, and was dominated by crustaceans. The distribution of groundwater fauna was influenced mainly by local factors, especially those related to the physical structure of wells such as the presence of a margin, a closing system, and total depth. At the regional scale, elevation and season were the only factors influencing the distribution of fauna. Further research needs to be conducted to highlight the relationship between groundwater invertebrates and human impacts in rural areas.     

Arthropod community of dandelion (Taraxacum officinale) capitula during seed dispersal

Dandelion (Taraxacum officinale) is an abundant Asteraceae species of grassland and vasteland stands. Through the vegetative season each plant produces a number of short-lived inflorescens. Final stage of inflorescence development is seed dispersal that lasts 2 days. Despite its ephemeral persistence, inflorescences at this period host abundant fauna. In 2005–2010 the inflorescences were collected in regular intervals in May–October, at Prague-Ruzyně and selected sites of western Czech Republic. Typical fauna consist of Heteroptera adults and larvae (34 species), phytophagans feeding on receptacle and seeds (mainly Miridae, Pentatomidae, Rhopalidae) and predators consuming thrips (Anthocoridae). The composition of Heteroptera fauna varies through the vegetative season, and its abundance decreases with increasing altitude of the locality. Other fauna are relics of species consuming seeds in the post-flowering phase of inflorescence maturation (mainly Thysanoptera) and occasional visitors with no trophic relationship to dandelion (Coleoptera, Diptera, Hymenoptera, Araneae). The reasons for association of phytophagous Heteroptera and dandelion may be preferences for seed consumption and chemicals present in plant tissues and possibly used for sequestration of aposematic substances.     

The bottom fauna of the rapids of the Black Volta River in Ghana

Summary The increase in abundance and biomass of the bottom fauna in the Black Volta River rapids seems to be closely related to increasing rate of water flow. These differences are due mainly to the increasing numbers of Cheumatopsyche larvae, a web spinning caddisfly. At higher current speeds more food is provided for this omnivorous species, and this seems to determine its density on the substratum. In a very slow current which prevents the use of webs for food catching, this genus is absent.The abundance and standing crop of bottom fauna at the end of dry season can be considered as the highest for the whole year. The presence of several instars of the commonest species guarantees almost continuous emergence and recolonization of the substratum during the dry season. With the onset of heavy rains and floods the communities found in the rapids are disturbed, and eventually completely destroyed. This is because the substrate is carried away by the swift flow of the river. Recolonization can be expected after the rains when the water level is low and the substratum again stabilized.The fauna of rapids is a community which can be expected to be almost completely destroyed when it becomes permanently flooded in any man-made lake. Some organisms, however, such as the larvae of Ecnomus and Cricotopus may establish themselves again in a habitat physically and chemically akin to that of rapids; such a habitat is for example the bark of flooded trees exposed to wave action and surrounded by water well saturated with oxygen.     

Benthic, Drifting and Marginal Macroinvertebrate Assemblages in a Lowland River: Temporal and Spatial Variations and Size Structure

Aquatic macroinvertebrates living in anastomosing lowland rivers use different habitats and respond differently to the hydrological regime. In this paper, the structure and composition of benthic, drifting and marginal macroinvertebrate assemblages are analyzed in the lowland river Ctalamochita (Córdoba, Argentina). The assemblages were studied in an annual cycle; a comparison among the composition of benthos, drift and marginal fauna was carried out; and size structure of the assemblages was characterized. Samples were obtained from two sites: a rural and an urban site. In total 73 taxa of aquatic macroinvertebrates were collected. Benthos was characterized by Chironomidae and Oligochaeta; marginal fauna was mainly constituted by Coleoptera, Heteroptera, Decapoda, the Trichoptera Nectopsyche sp., Ephemeroptera and Odonata. Drifting assemblage was composed by macroinvertebrates from local and remote upstream benthos, and from the marginal zone. Marginal fauna diversity was higher than benthos and drift. Total biomass of the assemblages pooled together was relatively equitably among size classes. Larger size classes consisted of organisms from the marginal zone whereas the smallest ones were composed by benthic and drifting organisms. In the study area there is habitat partitioning in the lateral dimension of the river. Marginal fauna was more diverse due to the asymmetry of transport and deposit processes, which generate a heterogeneous habitat in the bankside. The relation between fine substrate and high current velocity determines an unstable habitat in the central channel, which makes colonization by benthic macroinvertebrates difficult.     

Simulating the East African wildebeest migration patterns using GIS and remote sensing

The Serengeti–Mara ecosystem in East Africa is a spectacular natural heritage endowed with diverse fauna and flora. The presence of the seasonally migrating wildebeest (Connochaetes taurinus) is a major boost for tourism. This migration however has enormous impacts to the ecosystem. Consequently efforts at monitoring the herd's migration trends and patterns remain a challenge to wildlife managers and ecologists in the region. In this paper, the relative influence of vegetation (normalized difference vegetation index), landscape and relief on herds migration routes are investigated and the migration routes simulated using GIS and remote sensing techniques. The results are compared with the annual mean route taken by the herds, as determined by radio tracking over the 1995–1997 period. Green vegetation availability is shown to be the major criterion in route choice. It is also shown that during the dry season phases of the migration (western trek, western corridor), the herd endures complex relief (complexity quantified based on slope and inter‐visibility) in the search for greener grass. During the season of abundance (southern trek), relief becomes critical in making route choices, with herds avoiding difficult terrain, notwithstanding their relatively more abundant vegetation. The method proposed in this paper is viable for rapid prediction of approximate routes for the migrating wildebeest in different climatic conditions.     

雅鲁藏布江流域NDVI变化与风沙化土地演变的耦合关系

运用1982-2010年的两种NDVI数据集（Pathfinder AVHRR和SPOT VEGETATION）,以及1975、1990、2000和2008年4期遥感数据,通过GIS技术、人工目视解译和灰色关联分析方法,研究了雅鲁藏布江流域NDVI变化和风沙化土地演变的耦合关系,结合1957-2007年降水和气温逐日气象资料,探讨了气候变化对其耦合关系的影响。结果表明：（1）流域内1982-2010年NDVI的年际变化总体上呈波动式增长的趋势。NDVI空间分布呈现由下游向中上游逐渐降低的趋势,以米林宽谷最大、马泉河宽谷最小。（2）2008年流域内共有风沙化土地273 697.54hm²,呈现由江源区马泉河宽谷向中下游递减的趋势。1975-2008年流域内风沙化土地呈缓慢增长趋势,以1990-1999年增长率最高,2000-2008年的增长率最小。（3）对于NDVI年变化和植被生长季（7-9月份）的变化,马泉河宽谷受平均气温的影响最大,日喀则宽谷和山南宽谷受年降水量的影响最大;米林宽谷NDVI的年变化受风沙化土地扩展的影响最大,植被生长季变化受年降水量的影响最大。（4）不同宽谷段NDVI与风沙化土地年变化的关联度自下游向中上游呈总体减小的趋势。流域尺度NDVI植被生长季变化主要受平均气温和年降水量的影响,非植被生长季（10月-翌年6月）变化主要受风沙化土地扩展的影响。     

Cladocera from the Sudan: Red Sea Hills,Jebel Marra and valley of the main Nile

Twenty species of Cladocera are reported from the Nile, where lacustrine species dominate, and from Jebel Marra and the Red Sea Hills, where chydorids dominate. The community found in the Red Sea Hills is more typically desertic than that of Jebel Marra, which appears closely related to the fauna of the West and Central African Sahel.     

川西高山林线三种灌木凋落叶分解中的无脊椎动物多样性

以无脊椎动物为主体的土壤动物是影响凋落物分解的重要生物因素,对维持陆地生态系统物质循环和能量流动具有重要作用。高山林线交错带是高山植被垂直带谱中重要的过渡区域,拥有比相邻生态系统更高的生境复杂性和物种多样性。林线上温度波动和冻融循环频率显著高于针叶林,为了了解林线交错带上环境差异对凋落物分解过程中的土壤动物群落结构和多样性的影响,采用凋落物分解袋的方法,于高山生态系统的两个主要时期,即雪被末期和生长季末期,研究了林线主要代表性灌木——高山柳(Salix cupularis)、高山杜鹃(Rhododendron lapponicum)和红毛花楸(Sorbus rufopilosa)凋落叶分解的土壤动物多样性特征。结果表明:凋落物中的无脊椎动物群落多样性及个体、类群密度随物种、海拔梯度和季节而变化,且季节差异对无脊椎动物多样性的影响比物种和海拔梯度更显著。3个因子的交互作用不仅影响土壤动物群落多样性和均匀度,而且影响群落个体密度和类群密度。雪被末期,凋落物中的无脊椎动物多样性指数H、均匀度指数J及丰富度指数D以针叶林最高,优势度指数C以林线最高;生长季节末期的无脊椎动物类群密度和个体密度显著高于雪被末期。总体上,凋落物中的无脊椎动物群落丰富度以生长季末期最高,林线较针叶林丰富。这意味着,未来气候变暖情景下,灌丛密度增加,凋落物输入量增大,可能导致无脊椎动物多样性增加。     

The destruction of an endemic species flock: quantitative data on the decline of the haplochromine cichlids of Lake Victoria

Synopsis The Lake Victoria fish fauna included an endemic cichlid flock of more than 300 species. To boost fisheries, Nile perch (Lates sp.) was introduced into the lake in the 1950s. In the early 1980s an explosive increase of this predator was observed. Simultaneously, catches of haplochromines decreased. This paper describes the species composition of haplochromines in a research area in the Mwanza Gulf of Lake Victoria prior to the Nile perch upsurge. The decline of the haplochromines as a group and the decline of the number of species in various habitats in the Mwanza Gulf was monitored between 1979 and 1990. Of the 123+ species originally caught at a series of sampling stations ca. 80 had disappeared from the catches after 1986. In deepwater regions and in sub-littoral regions haplochromine catches decreased to virtually zero after the Nile perch boom. Haplochromines were still caught in the littoral regions where Nile perch densities were lower. However, a considerable decrease of species occurred in these regions too. It is expected that a remnant of the original haplochromine fauna will survive in the littoral region of the lake. Extrapolation of the data of the Mwanza Gulf to the entire lake would imply that approximately 200 of the 300+ endemic haplochromine species have already disappeared, or are threatened with extinction. Although fishing had an impact on the haplochromine stocks, the main cause of their decline was predation by Nile perch. The speed of decline differed between species and appeared to depend on their abundance and size, and on the degree of habitat overlap with Nile perch. Since the Nile perch upsurge, the food web of Lake Victoria has changed considerably and the total yield of the fishery has increased three to four times. Dramatic declines of native species have also been observed in other lakes as a result of the introduction of alien predators. However, such data concern less speciose communities and, in most cases, the actual process of extinction has not been monitored.     

The impact of the introduction of Nile perch, Lates niloticus (L.) on the fisheries of Lake Victoria

The piscivorous Nile perch was introduced into Lake Victoria some 30 years ago, since when it has completely transformed the fishing industry and the species composition of the fish fauna of the lake. The original multispecies fishery, based mostly on cichlids (haplochromines, tilapias), cyprinids ( Barbus, Labeo, Rastrineobola ) and siluroids ( Bagrus, Clarias, Synodontis, Schilbe ), has changed dramatically to one based on three species: the introduced Nile perch, the cyprinids, Rastrineobola argenrea (Pellegrin), and the introduced Nile tilapia, Oreochromis niloticus (Linnaeus).
Within 25 years of its introduction the Nile perch became ubiquitous and now occurs in virtually every habitat with the exception of swamps and affluent rivers. It has preyed on all other species with profound effects, especially on the stocks of haplochromines. These originally comprised 80% of the total fish biomass in Lake Victoria, but have now decreased to less than 1% offish catches from the Kenyan waters of the lake. The fishermen of Lake Victoria have adjusted to this ecological crisis by using large-meshed nets to catch Nile perch, which has become the most important commercial species. For the first time in the history of Lake Victoria, fish fillets are now being exported to several overseas countries: the fillets are all from Nile perch.     

珠江口及邻近海域大型底栖动物多样性随盐度、水深的变化趋势

利用2006年7-8月(夏季)、2007年4-5月(春季)和2007年10-12月(秋季)珠江丰水期、平水期和枯水期3个航次在南中国海北部珠江口附近海域4条由河口、近岸到深水区调查断面的数据, 研究大型底栖动物多样性由河口-近岸-深水的变化趋势及与环境因子的关系。春季、夏季和秋季分别获得大型底栖动物273、256和148种, 各季节均以环节动物种类最多, 节肢动物次之。大型底栖动物种类数、丰度、生物量和Shannon-Wiener多样性指数均由河口向近岸海域升高, 再由近岸向外海深水区降低。Pielou均匀度深水区最高, 其次为近岸。河口和深水区大型底栖动物k-优势度曲线位于近岸浅水域曲线之上, 表明生物多样性由河口向近岸升高, 而由近岸向深水则降低。大型底栖动物与环境因子Pearson相关性分析表明, 春、秋季大型底栖动物种类数、丰度和生物量与水深呈显著的负相关, 秋季种类多样性指数和均匀度也与水深呈显著的负相关性, 而夏季仅生物量与水深呈显著的负相关; 春、秋季大型底栖动物种类数、生物量、丰度、多样性指数和种类均匀度与盐度的相关性不显著, 但是夏季大型底栖动物种类数、丰度、多样性指数和种类均匀度与盐度呈显著正相关。单位面积(0.2 m²)内, 珠江口及邻近海域大型底栖动物在近岸浅水区较深水区和河口生物多样性高, 且生物量丰富。     

Effects of a Long-Term Disturbance on Arthropods and Vegetation in Subalpine Wetlands: Manifestations of Pack Stock Grazing in Early versus Mid-Season

Conclusions regarding disturbance effects in high elevation or high latitude ecosystems based solely on infrequent, long-term sampling may be misleading, because the long winters may erase severe, short-term impacts at the height of the abbreviated growing season. We separated a) long-term effects of pack stock grazing, manifested in early season prior to stock arrival, from b) additional pack stock grazing effects that might become apparent during annual stock grazing, by use of paired grazed and control wet meadows that we sampled at the beginning and end of subalpine growing seasons. Control meadows had been closed to grazing for at least two decades, and meadow pairs were distributed across Sequoia National Park, California, USA. The study was thus effectively a landscape-scale, long-term manipulation of wetland grazing. We sampled arthropods at these remote sites and collected data on associated vegetation structure. Litter cover and depth, percent bare ground, and soil strength had negative responses to grazing. In contrast, fauna showed little response to grazing, and there were overall negative effects for only three arthropod families. Mid-season and long-term results were generally congruent, and the only indications of lower faunal diversity on mid-season grazed wetlands were trends of lower abundance across morphospecies and lower diversity for canopy fauna across assemblage metrics. Treatment x Season interactions almost absent. Thus impacts on vegetation structure only minimally cascaded into the arthropod assemblage and were not greatly intensified during the annual growing season. Differences between years, which were likely a response to divergent snowfall patterns, were more important than differences between early and mid-season. Reliance on either vegetation or faunal metrics exclusively would have yielded different conclusions; using both flora and fauna served to provide a more integrative view of ecosystem response.     

1956-2009年内蒙古苏尼特左旗荒漠草原的降水格局

弄清全球变化背景下不同地区降水格局的变化对科学理解气候变化及其影响具有重要意义。苏尼特左旗荒漠草原是温带干旱半干旱地区的典型荒漠草原,对气候变化,特别是降水变化非常敏感。利用1956-2009年的日降水资料探讨了苏尼特左旗荒漠草原降水格局的变化规律,以为揭示气候变化的影响机制提供依据。结果表明,该地区年均降水量为191.9 mm,年际变化剧烈,变异系数达26.63%;年均降水日数为63.8 d,变异系数16.9%。生长季降水占全年的85%,但各月变异系数均>50%;降水日数占全年的63%。年和生长季的各月降水以中等降水量、弱降水日数为主,中等强度以上降水事件较少。近50 a来,年和生长季的降水量、降水日数与各等级降水事件均呈下降趋势,年降水减少的原因在于中等降水事件的减少、生长季降水减少的原因在于弱降水事件的减少。年降水减少将影响草地的土壤水分与植物返青;而生长季降水减少将直接影响草地固碳。气候变化背景下年与生长季降水的减少将进一步加剧该地区干旱程度并影响植物的生长发育,从而直接威胁到草地畜牧业的发展。     

Kissinella-Christiania Associations in the early Ashgill Foliomena brachiopod fauna of South China

A new ecological unit within the well-documented late Ordovician brachiopod Foliomena fauna is defined on the basis of assemblages from Northern Cuizhou and southern Sichuan. Southwest China. The Kassinella-Chrisriania Association, from the lower Ashgill Linhsiang and Chientsaokou formations, is dominated by the eponymous genera together with species of mainly Dedzetina, Sericoidea and Kozlowskites Foliomena is rare or absent. To date, the Foliomena fauna has been documented only from deep-water environments however, the new association apparently occupied shallower depths, probably in lowermost Benthic Assemblage Zone 3 and uppermost Benthic Assemblage Zone 4, presumably at the shallow end of the depth range of the Foliomere fauna. The incursion of the Foliomena fauna into relatively shallow-water environments may have been encouraged by abnormally low oxygen levels and sparse nutrients together with persistent soft substrates across this part of the Yangtze Platform during the early Ashgill. *** Ordovician, Ashgill, China, brachiopods. F oliomena fauna.