Global change and marine communities: alien species and climate change

Anthropogenic influences on the biosphere since the advent of the industrial age are increasingly causing global changes. Climatic change and the rising concentration of greenhouse gases in the atmosphere are ranking high in scientific and public agendas, and other components of global change are also frequently addressed, among which are the introductions of non indigenous species (NIS) in biogeographic regions well separated from the donor region, often followed by spectacular invasions. In the marine environment, both climatic change and spread of alien species have been studied extensively; this review is aimed at examining the main responses of ecosystems to climatic change, taking into account the increasing importance of biological invasions. Some general principles on NIS introductions in the marine environment are recalled, such as the importance of propagule pressure and of development stages during the time course of an invasion. Climatic change is known to affect many ecological properties; it interacts also with NIS in many possible ways. Direct (proximate) effects on individuals and populations of altered physical-chemical conditions are distinguished from indirect effects on emergent properties (species distribution, diversity, and production). Climatically driven changes may affect both local dispersal mechanisms, due to the alteration of current patterns, and competitive interactions between NIS and native species, due to the onset of new thermal optima and/or different carbonate chemistry. As well as latitudinal range expansions of species correlated with changing temperature conditions, and effects on species richness and the correlated extinction of native species, some invasions may provoke multiple effects which involve overall ecosystem functioning (material flow between trophic groups, primary production, relative extent of organic material decomposition, extent of benthic-pelagic coupling). Some examples are given, including a special mention of the situation of the Mediterranean Sea, where so many species have been introduced recently, and where some have spread in very large quantities. An increasing effort by marine scientists is required, not only to monitor the state of the environment, but also to help predicting future changes and finding ways to mitigate or manage them.     

A preliminary analysis of fishery resource exhaustion in the context of biodiversity decline

Marine biodiversity in almost all oceans is being threatened at the genetic, species, and ecosystem levels. The marine ecosystem is being degraded and the extinction rate of marine organisms has accelerated. In this paper, the potential causes of fishery resource exhaustion in the East China Sea are analyzed, including the change in the stoichiometric composition of seawater with regard to the concentrations of N and P, toxic effects of marine pollution, marine habitat destruction, increased seawater temperatures caused by climate warming, ocean acidification, pressure from overfishing, and the spread of marine pathogenic bacteria. It is believed that the factors mentioned above have significant impact on the exhaustion of fishery resources in the East China Sea. However, considering the cumulative, synergistic, and superimposed effects as well as the amplification effects resulting from their interactions, the actual risk of ecological extinction of marine organisms might be even more severe than that previously estimated. Hence, ecosystem management and research focused on a single risk factor or influencing factor is not enough to prevent marine ecosystem degradation and fishery resource exhaustion. A comprehensive, systematic, effective, and ecosystem-based management policy is imperative for healthy and sustainable fishery development in the East China Sea.     

Indirect effects of asymmetrical competition among top predators determine spatial patterns of predation risk for prey

Asymmetrical interspecific competition among top predators can indirectly affect the predation risk for their prey by altering the abundance, diet, and habitat use of inferior competitors. However, the indirect effects of such biological interactions are poorly known because of the difficulties in measuring predation risk in nature. We addressed this issue by assessing the effect of asymmetrical competition among brook trout (Salvelinus fontinalis) and two superior non-piscivorous competitors, creek chub (Semotilus atromaculatus) and white sucker (Catostomus commersonii), on the predation risk of a brook trout prey, northern redbelly dace (Chrosomus eos). We determined the spatio-temporal patterns of relative predation risk of dace with tethering experiments in 11 lakes containing either only brook trout and dace (n = 5), or brook trout, dace, chub, and sucker (n = 6). The diel pattern of the relative predation risk and the overall relative predation risk of dace were not significantly different in lakes with or without brook trout competitors. However, we observed a significant shift in the relative predation risk from the lower pelagic to the upper pelagic and littoral zones in the presence of brook trout competitors. This study highlights the fact that the outcome of interactions can vary in space and that care should be used when extrapolating the results of small-scale experiments or coarse-scale estimates to the whole ecosystem.     

A web-based system for public–private sector collaborative ecosystem management

 To preserve biodiversity over centuries, ecosystem management will need to be accepted and practiced by individuals from a broad spectrum of society's strata. Also, management decisions will need to be based on reliable judgments of the cause and effect relationships that govern an ecosystem's dynamics. This article describes an extant, web-based ecosystem management system (EMS) that allows (a) wide participation in ecosystem assessment and policy impact predictions, (b) convenient construction of probabilistic models of ecosystem processes through an influence diagram, and (c) automatic creation of ecosystem assessment reports. For illustration, the system is used to first model the cheetah population in Kenya, and then to assess the impact on this population of different management options. The influence diagram used herein extends standard influence diagram theory to allow representation of variables governed by stochastic differential equations, birth–death processes, and other nongaussian, continuous probability distributions. For many ecosystems, data sets on ecosystem health indicators can be incomplete, small, and contain unknown measurement errors. Some amount of knowledge of an ecosystem's dynamics however, may exist in the form of expert opinion derived from ecological theory. The proposed EMS uses a nonbayesian parameter estimation method, called consistency analysis that finds parameter estimates such that the fitted ecosystem model is as faithful as possible to both the available data and the collected body of expert opinion. For illustration, consistency analysis is used to estimate the cheetah viability influence diagram using all known cheetah surveys in the country of Kenya plus current understanding of factors such as habitat and prey availability that affect cheetah population dynamics.     

Can soil piping impact environment and society? Identifying new research gaps

Soil piping is a widespread, although often overlooked land degradation process. So far, subsurface soil erosion studies have been focused on the importance of soil piping in hydrological and geomorphological processes, and factors controlling piping processes. Nowadays, the environmental changes being caused by the Anthropocene have clearly demonstrated that society depends on soil more than ever before, so the traditional studies of soil erosion processes need to be redefined. In that sense, geomorphologists face to overcome new piping-related problems. In this article we identify new possible areas of research: (i) soil pipes and pipe collapses (PCs) as natural hazards, (ii) role of soil piping in carbon cycle, (iii) soil pipes and PCs and their relationships with biodiversity, and (iv) piping-affected areas as geodiversity sites. Only better recognition of natural hazards driven by soil piping, such as land subsidence and degradation, landslides, flooding and off-site sediment effects may result in better prevention and control measures in piping-affected areas. Moreover, in the context of Global Change the role of soil piping in carbon cycle should be raised. Land-use and land-cover changes, as well as climate change may affect piping dynamics in different morphoclimatic regions and soil loss due to piping may lead to carbon loss. Soil pipes and PCs are closely interlinked with biodiversity, both positively and negatively. Piping erosion may directly and indirectly destroy vegetation and animals, although in some cases piping erosion may create new habitats and provide favourable conditions for some species. However, soil piping is not only an environmental and societal problem, but it may also contribute to the world geodiversity, which is clearly observed in badland sites. Piping erosion may have a significant impact on environment and society, thus further research with new questions is essential to provide knowledge for sustainable development.     

古湖沼学研究揭示湖泊生态系统服务变化的过程

当今在气候变化和人类活动等多重压力影响下,作为地表过程重要组成部分的湖泊生态系统正遭受着巨大威胁,其服务功能急剧退化.如何客观地评价湖泊生态系统服务的现状,并科学地预测其发展趋势,是当前亟待解决的问题.开展长期生态系统服务变化过程与机理的分析,有助于未来生态系统服务变化趋势的预测.然而,现有的观测数据往往时间较短(通常小于50年).连续的湖泊沉积记录为研究生态系统服务变化的长期过程提供了可能.本文结合前人研究成果,列举了可反映湖泊生态系统服务变化的一些古湖沼学指标,依据这些指标相对明确的生态和环境指示意义,将其与各项服务关联起来.最后,结合巢湖实例分析来说明这些指标在评估湖泊生态系统服务方面的具体应用,研究表明当今巢湖生态系统服务供应能力的持续增加是以调节服务的丧失为代价的.尽管目前的研究尚处于起步阶段,但古湖沼学手段无疑为今后湖泊生态系统服务历史状况的评估提供了途径,为古湖沼学的应用提供了一种新的思路,并为今后生态系统的保护和可持续利用提供重要的决策依据.     

Biological and hydromorphological integrity of the small urban stream

Biological and hydromorphological integrity of five reaches of the small urban stream were assessed. Because macroinvertebrate communities respond to both organic pollution and habitat change, impacts of both measures can be hardly separated. In our study on the urbanized small stream, an impact of organic pollution was excluded as all five sampling sites were assessed as moderately polluted. On the other hand differences in morphological degradation of banks and channel of selected sites enabled us to relate hydromorphological stress and biotic metrics and taxa. Physical habitat quality was assessed using River habitat survey (RHS) methodology. A downstream-upstream gradient of physical habitat degradation was observed and related to the macroinvertebrate community characteristics. Similarity analyses and biotic metrics were calculated and correlated with results of the RHS analyses. Composition of the macroinvertebrate assemblages did not follow the longitudinal pattern of habitat modification observed by the RHS analysis. However, some metrics corresponded well. Percentage of detritivores, percentage of Caenis luctuosa, number of individuals, percentage of EPT individuals were best predictors of changes in the physical habitat quality. However, the metric percentage of EPT individuals was negatively correlated to the habitat degradation, what is in contradiction with results from studies of other authors.     

Forests and water in South America

South America is experiencing rapid change in forest cover, of both native and planted forest. Forest cover loss is primarily attributable to fire, logging, and conversion of native forest to agriculture, pasture, and forest plantations, and types of change vary within and among the many diverse types of forests in South America. Major changes in forest cover and growing policy concerns underscore an urgent need for research on sustainable forest management and water ecosystem services in South America. Differences in land ownership and management objectives create trade‐offs between wood production and water ecosystem services from forests. Work is needed to quantify how forest change and management affect ecosystem services, such as wood production versus water provision. Current scientific understanding of forest management effects on water ecosystem services in South America has important limitations, including a scarcity of long‐term records and few long‐term integrated watershed studies. Industry, government, universities, and local communities should collaborate on integrated applied studies of forests and water. Data archiving and publically available data are required. The creation of national networks and a multi‐country South America network to identify and implement common water research protocols, share results, and explore their implications would promote common and well‐supported policies. Hydrologists working in South America are well placed to tackle the challenges and opportunities for collaborative research that will maintain the intrinsic values and water ecosystem services provided by South America's forests.     

Long-term coexistence of non-indigenous species in aquaculture facilities

Non-indigenous species (NIS) are a growing problem globally and, in the sea, aquaculture activities are critical vectors for their introduction. Aquaculture introduces NIS, intentionally or unintentionally, and can provide substratum for the establishment of other NIS. Little is known about the co-occurrence of NIS over long periods and we document the coexistence over decades of a farmed NIS (a mussel) with an accidently introduced species (an ascidian). Both are widespread and cause serious fouling problems worldwide. We found partial habitat segregation across depth and the position of rafts within the studied farm, which suggests competitive exclusion of the mussel in dark, sheltered areas and physiological exclusion of the ascidian elsewhere. Both species exhibit massive self-recruitment, with negative effects on the industry, but critically the introduction of NIS through aquaculture facilities also has strong detrimental effects on the natural environment.     

Implementing environmental flows in integrated water resources management and the ecosystem approach

Abstract

In many of the world’s river basins, the water resources are over-allocated and/or highly modified, access to good quality water is limited or competitive and aquatic ecosystems are degraded. The decline in aquatic ecosystems can impact on human well-being by reducing the ecosystem services provided by healthy rivers, wetlands and floodplains. Basin water resources management requires the determination of water allocation among competing stakeholders including the environment, social needs and economic development. Traditionally, this determination occurred on a volumetric basis to meet basin productivity goals. However, it is difficult to address environmental goals in such a framework, because environmental condition is rarely considered in productivity goals, and short-term variations in river flow may be the most important driver of aquatic ecosystem health. Manipulation of flows to achieve desired outcomes for public supply, food and energy has been implemented for many years. More recently, manipulating flows to achieve ecological outcomes has been proposed. However, the complexity of determining the required flow regimes and the interdependencies between stakeholder outcomes has restricted the implementation of environmental flows as a core component of Integrated Water Resources Management (IWRM). We demonstrate through case studies of the Rhône and Thames river basins in Europe, the Colorado River basin in North America and the Murray-Darling basin in Australia the limitations of traditional environmental flow strategies in integrated water resources management. An alternative ecosystem approach can provide a framework for implementation of environmental flows in basin water resources management, as demonstrated by management of the Pangani River basin in Africa. An ecosystem approach in IWRM leads to management for agreed triple-bottom-line outcomes, rather than productivity or ecological outcomes alone. We recommend that environmental flow management should take on the principles of an ecosystem approach and form an integral part of IWRM.

Editor D. Koutsoyiannis

Citation Overton, I.C., Smith, D.M., Dalton J., Barchiesi S., Acreman M.C., Stromberg, J.C., and Kirby, J.M., 2014. Implementing environmental flows in integrated water resources management and the ecosystem approach. Hydrological Sciences Journal, 59 (3–4), 860–877.     

Alteration of physico-chemical and microbial properties in freshwater substrates by burrowing invertebrates

The hyporheic interstitial provides habitat for many different organisms – from bacteria to burrowing invertebrates. Due to their burrowing and sediment reworking behaviour, these ecosystem engineers have the potential to affect hyporheic processes such as respiration and nutrient cycling. However, there is a lack of studies that characterize the interactions between bioturbators, physico-chemical habitat properties and microbial communities in freshwater substrates. In a standardized laboratory experiment, we investigated the effects of three functionally different bioturbators, duck mussels (Anodonta anatina, Linnaeus 1758), mayfly nymphs (Ephemera danica, Müller 1764) and tubificid worms (Tubifex tubifex, Müller 1774), on the physico-chemical conditions and bacterial communities in hyporheic substrates. We hypothesized that different invertebrates distinctly alter habitat conditions and thus microbial community composition, depending on the depth and the manner of burrowing. A. anatina and E. danica caused an increase in interstitial oxygen concentration, whereas strong declines in oxygen concentration and redox potential were detected in the T. tubifex treatment. These effects on physico-chemical habitat properties were even detectable in open water. Mussels and tubificid worms also significantly influenced the composition of bacterial communities in the hyporheic zone. A loss or replacement of bioturbators in stream ecosystems due to anthropogenic habitat alterations is expected to result in shifts in microbial community compositions, with effects on nutrient fluxes, pollutant degradation and benthic food webs. An understanding of the effects of functionally different native and invasive bioturbators is crucial to predict changes in stream ecosystem functioning.     

受压水生物群落生态系统目标函数的变化

The changes of biomass, exergy and structural exergy under the action of various chemical pollution (addition of nutrient compounds, pesticides, chlorinated phenols, oil, heavy metal ions, acidification etc.) are analysed for 50 experimental works with model aquatic ecosystems -microcosms, mesocosms and experimental ponds.Structural exergy is shown to remain at constant level or to increase when the allochtonous compounds can be metabolised by ecosystem, or when the ecosystem can adapt itself to the input of toxicant by structural changes. When the substance is too conservative, too toxic or/and is coming in too high concentrations, structural exergy is decreasing, demonstrating the inability of ecosystem to adapt to this influence and irreversibility of changes in ecosystem.The possibility to use such parameter as structural exergy for estimation of ecosystem state and its changes under various external influences is demonstrated. It reflects the state of ecosystem and can indicate the degree of ecosystem adaptation, decreasing when important for ecosystem functioning components are eliminated.In experiments with mathematical model of lake Baikal ecosystem exergy content is shown to increase after addition of nutrients and to decrease after addition of phenolic compounds, reflecting the general shifts in ecosystem.     

杂食性鱼类对浅水水体底栖—浮游生境耦合作用的影响:微综述

浅水水体存在着强烈的底栖-浮游生境耦合作用,耦合的结果决定着水生态系统关键特征.在缺少大型水生植物的浅水系统中,底栖藻类和浮游藻类对光照和营养盐的竞争是底栖-浮游生境耦合最为重要的生态过程之一,但该过程受到杂食性鱼类的影响.本文以浅水水体的底栖-浮游生境耦合作用为切入点,综述了杂食性鱼类对浅水水体底栖-浮游生境耦合作用的影响过程及机理.一般而言,杂食性鱼类有利于提高水层营养盐浓度,促进浮游藻类生长,降低水体透明度,不利于底栖藻类竞争,从而加速水体富营养化.但不同种类的杂食性鱼类(如底栖杂食性鱼类、偏植物性饵料的杂食性鱼类、偏动物性饵料的杂食性鱼类以及小型杂食性鱼类)因食性差异,对底栖-浮游生境耦合的影响机理不同,产生的生态环境效应各异;即便同一种杂食性鱼类也可因发育阶段不同对底栖-浮游生境耦合产生不同的影响.在人类活动、全球变暖以及富营养化等多重因子胁迫下杂食性鱼类在鱼类群落结构中的比例上升,因此,杂食性鱼类对水生态环境产生的影响深远、复杂,值得持续关注.     

The impacts of climate extremes on the terrestrial carbon cycle:A review

The increased frequency of climate extremes in recent years has profoundly affected terrestrial ecosystem functions and the welfare of human society. The carbon cycle is a key process of terrestrial ecosystem changes. Therefore, a better understanding and assessment of the impacts of climate extremes on the terrestrial carbon cycle could provide an important scientific basis to facilitate the mitigation and adaption of our society to climate change. In this paper, we systematically review the impacts of climate extremes(e.g. drought, extreme precipitation, extreme hot and extreme cold) on terrestrial ecosystems and their mechanisms. Existing studies have suggested that drought is one of the most important stressors on the terrestrial carbon sink, and that it can inhibit both ecosystem productivity and respiration. Because ecosystem productivity is usually more sensitive to drought than respiration, drought can significantly reduce the strength of terrestrial ecosystem carbon sinks and even turn them into carbon sources. Large inter-model variations have been found in the simulations of drought-induced changes in the carbon cycle, suggesting the existence of a large gap in current understanding of the mechanisms behind the responses of ecosystem carbon balance to drought, especially for tropical vegetation. The effects of extreme precipitation on the carbon cycle vary across different regions. In general, extreme precipitation enhances carbon accumulation in arid ecosystems, but restrains carbon sequestration in moist ecosystems. However, current knowledge on the indirect effects of extreme precipitation on the carbon cycle through regulating processes such as soil carbon lateral transportation and nutrient loss is still limited. This knowledge gap has caused large uncertainties in assessing the total carbon cycle impact of extreme precipitation. Extreme hot and extreme cold can affect the terrestrial carbon cycle through various ecosystem processes. Note that the severity of such climate extremes depends greatly on their timing, which needs to be investigated thoroughly in future studies. In light of current knowledge and gaps in the understanding of how extreme climates affect the terrestrial carbon cycle, we strongly recommend that future studies should place more attention on the long-term impacts and on the driving mechanisms at different time scales.Studies based on multi-source data, methods and across multiple spatial-temporal scales, are also necessary to better characterize the response of terrestrial ecosystems to climate extremes.     

Assessment of flow–ecology relationships for environmental flow standards: a synthesis focused on the southeast USA

ABSTRACT

Environmental flow standards are a management tool that can help to protect the ecosystem services sustained by rivers. Although environmental flow requirements can be assessed using a variety of methods, most of these methods require establishing relationships between flow and habitat of species of concern. Here, we conducted a synthesis of past flow–ecology studies in the southeast USA. For each state or interstate river basin, we used the published data to determine the flow metrics that resulted in the greatest changes in ecological metrics, and the ecological metrics that were most sensitive to hydrologic alteration. The flow metrics that were most important in preserving ecological metrics were high-flow duration and frequency, 3-day maximum and minimum, and number of reversals. The ecological metrics most sensitive to hydrologic alteration were mostly related to presence or absence of key indicator species.     

Global desertification: Drivers and feedbacks

Desertification is a change in soil properties, vegetation or climate, which results in a persistent loss of ecosystem services that are fundamental to sustaining life. Desertification affects large dryland areas around the world and is a major cause of stress in human societies. Here we review recent research on the drivers, feedbacks, and impacts of desertification. A multidisciplinary approach to understanding the drivers and feedbacks of global desertification is motivated by our increasing need to improve global food production and to sustainably manage ecosystems in the context of climate change. Classic desertification theories look at this process as a transition between stable states in bistable ecosystem dynamics. Climate change (i.e., aridification) and land use dynamics are the major drivers of an ecosystem shift to a “desertified” (or “degraded”) state. This shift is typically sustained by positive feedbacks, which stabilize the system in the new state. Desertification feedbacks may involve land degradation processes (e.g., nutrient loss or salinization), changes in rainfall regime resulting from land-atmosphere interactions (e.g., precipitation recycling, dust emissions), or changes in plant community composition (e.g., shrub encroachment, decrease in vegetation cover). We analyze each of these feedback mechanisms and discuss their possible enhancement by interactions with socio-economic drivers. Large scale effects of desertification include the emigration of “environmental refugees” displaced from degraded areas, climatic changes, and the alteration of global biogeochemical cycles resulting from the emission and long-range transport of fine mineral dust. Recent research has identified some possible early warning signs of desertification, which can be used as indicators of resilience loss and imminent shift to desert-like conditions. We conclude with a brief discussion on some desertification control strategies implemented in different regions around the world.     

近十年青海湖水生态系统生产总值变化及其影响因素

受气候变化和人类活动影响,湖泊水生态系统及其服务功能发生改变,研究湖泊水生态系统生产总值动态变化及其影响因素,对于维护湖泊健康生命、实现湖泊功能永续利用具有重要意义。选择位于气候敏感区和生态脆弱区的青海湖作为研究对象,根据青海湖水生态系统特征及当前保护状况,筛选调节服务及文化服务2类8个评估指标,构建湖泊水生态系统生产总值指标体系,核算2010—2020年青海湖水生态系统生产总值,分析其变化趋势及主要影响因素。结果表明,2010—2020年青海湖水生态系统生产总值总体呈波动上升趋势,变化范围为6903.47亿～7848.55亿元;调节服务是青海湖水生态系统主要的服务类型,占比高达91%。近十年,气候调节和水质净化价值有所减少,其他服务功能价值均呈增加趋势。水源涵养价值增加最多,增长760.70亿元;气候调节价值下降最多,减少658.59亿元。偏最小二乘回归分析表明,水温、水位是影响青海湖水生态系统生产总值的主要因素。气候变化影响下,水温升高引起初级生产力增加及鱼类数量增长,同时近年来水体矿化度下降有利于水生生物生长,提高了固碳释氧和物种保育价值。水位与水面面积增加引起水源涵养、洪水调蓄...     

Reviews on land use change induced effects on regional hydrological ecosystem services for integrated water resources management

This paper proposed to provide valuable information for integrated water resources management through evaluating the research on the interaction mechanism among land use changes, regional hydrological ecosystem services and human well-being. Firstly, the driving mechanism of land use and land cover changes was introduced in this paper. Secondly, the overview of the interaction mechanism among land use and land cover changes, regional hydrological ecosystem services and human well-being was given. Based on the meta-analysis, land use changes have a profound influence on regional hydrological ecosystem services, and the variation of hydrological ecosystem could benefit or impair human well-being. Finally, two suggestions were emphasized for managers or policy makers for the future integrated water resources management: (1) Proper land use makes for the water resource management; (2) Blindly pursuing the provisioning services weakens other services of hydrological ecosystem.     

Emergent geomorphic–vegetation interactions on a subalpine alluvial fan

Following perturbation, an ecosystem (flora, fauna, soil) should evolve as a function of time at a rate conditioned by external variables (relief, climate, geology). More recently, biogeomorphologists have focused upon the notion of co‐development of geomorphic processes with ecosystems over very short through to very long (evolutionary) timescales. Alpine environments have been a particular focus of this co‐development. However, work in this field has tended to adopt a simplified view of the relationship between perturbation and succession, including: how the landform and ecosystem itself conditions the impact of a perturbation to create a complex spatial response impact; and how perturbations are not simply ecosystem destroyers but can be a significant source of ecosystem resources. What this means is that at the within landform scale, there may well be a complex and dynamic topographic and sedimentological template that co‐develops with soil, flora and fauna. Here, we present and test a conceptual model of this template for a subalpine alluvial fan. We combine detailed floristic inventory with soil inventory, determination of edaphic variables and analysis of historical aerial imagery. Spatial variation in the probability of perturbation of sites on the fan surface was associated with down fan variability in the across‐fan distribution of fan ages, fan surface channel characteristics and fan surface sedimentology. Floristic survey confirmed that these edaphic factors distinguished site floristic richness and plant communities up until the point that the soil–vegetation system was sufficiently developed to sustain plant communities regardless of edaphic conditions. Thus, the primary explanatory variable was the estimated age of each site, which could be tied back into perturbation history and its spatial expression due to the geometry of the fan: distinct plant communities were emergent both across fan and down fan, a distribution maintained by the way in which the fan dissipates potentially perturbing events. Copyright © 2015 John Wiley & Sons, Ltd.