不同土壤培肥措施对华北高产农田生态系统蚯蚓种群的影响

通过在华北高产农田生态系统进行的土壤培肥试验 ,研究了不同秸秆还田方式和施肥措施对土壤大型动物——蚯蚓的影响。研究结果表明 :该区域高肥力土壤共有 7种蚯蚓 ,其中以正蚓科梯形流蚓为主 ,并同时分布巨蚓科的远盲蚓属、腔蚓属和链胃蚓科的杜拉蚓属的一些广布种。全年平均种群密度为 83.83条 /m2 ,最高种群数量出现在 1998年 8月份 ,其季节变化趋势为夏季 >春季 >秋冬季。在投入同量化肥条件下 ,有机物投入越多 ,蚯蚓越丰富 ,其生物量有以下趋势 :化肥投入 <化肥与麦秸还田 <化肥与玉米秸麦秸全还田 <化肥与玉米秸麦秸还田加施有机肥处理。在有有机物投入的条件下 ,化肥对土壤生物的负面效应不明显 ,不会对土壤中的蚯蚓有较大影响 ;而单施化肥 ,对蚯蚓负面影响较为显著。有机肥的施入可以增加蚯蚓的种群数量 ,随着时间的增加这种趋势越明显     

华北高产农田生态系统中蚯蚓种群次生演替规律

通过对山东省桓台县不同土壤肥力的农田生态系统中蚯蚓种群的调查发现 ,该地区农田生态系统中有 7种蚯蚓 ,高肥力土壤中蚯蚓相对较丰富 ,种群密度可达 83.83条 / m2 ,梯形流蚓为优势种 ;低肥力土壤中蚯蚓种群密度 4 0 .18条 / m2 ,优势种是天锡杜拉蚓。梯形流蚓和湖北远盲蚓在高肥力土壤中的种群密度要明显高于低肥力土壤 ;而其他几种蚯蚓种群数量变化不大 ,随着土壤肥力的演变 ,低肥力土壤中天锡杜拉蚓的优势地位随着土壤肥力的提高逐渐被梯形流蚓所代替。合理的投入特别是农田有机物投入可以加速农田生态系统中生态演替 ;试验中不同处理间蚯蚓的种群生物量有以下趋势 :化肥投入 <化肥与麦秸还田 <化肥与玉米麦秸全还田 <化肥与玉米麦秸全还田以及有机肥的施用。无论在高肥力还是低肥力的土壤上都表现出相同的规律 ,但蚯蚓种群组成并没有明显差异     

华北盐渍化改造区蚯蚓种群次生演替与生产投入的关系——以河北省曲周试区为例

通过对河北省曲周县盐渍化改造区农田生态系统中蚯蚓物种多样性分析,发现蚯蚓种群的次生演替与农田肥料投稿有很好的相关性,合理的投稿特别是有机肥的投入可以加速农田生态系统中生态演替;该地区蝗蚓种群分布总的趋势是：第2代试区＞第1代试区＞第3代试区＞原貌区;不同土壤利用方式蚯蚓的分布和数量各不相同,菜地＞农田、边缘地和果园。     

哀牢山、无量山与苍山蚯蚓物种调查及系统发育与扩散分析

哀牢山、无量山与苍山是云南省地理二级阶梯生物多样性研究的关键地区之一,但鲜有蚯蚓物种记录。为完善云南省蚯蚓多样性、系统发育与演化研究,本研究对哀牢山国家级自然保护区、无量山国家级自然保护区和苍山洱海国家级自然保护区展开蚯蚓物种野外调查、形态分类、分子系统发育与扩散分析。研究共获得蚯蚓物种3科8属27种,其中巨蚓科(Megascolecidae)21种,正蚓科(Lumbricidae)4种,链胃蚓科(Moniligastridae)2种,鉴定了3个新物种——无量山远盲蚓(Amynthas wuliangmontis sp.nov.)、景东远盲蚓(Amynthas jingdongensis sp.nov.)和湖畔远盲蚓(Amynthas lacustris sp.nov.)。我们提取并测定样本的线粒体COⅠ、COⅡ、12S r RNA、16S r RNA和ND1基因序列,该地区物种间上述5基因联合序列的平均遗传距离为19.6%±0.5%,种群阶元遗传距离范围为0~2.7%;其次,构建最大似然系统发育树和贝叶斯系统发育树,阐述了该地区蚯蚓物种的系统发育关系,再次验证了远盲蚓属与腔蚓属均不为单系群的观点;最后,依据祖先分布区域重建结果,并结合之前研究,推测该地区巨蚓科蚯蚓物种的主要扩散趋势为由南至北扩散——自哀劳山、无量山向苍山方向扩散。     

海南岛蚯蚓物种组成及其系统发育分析

海南岛独特的地理位置和复杂的古地理事件使其成为我国生物多样性的热点地区。已有的海南岛蚯蚓资料显示, 该岛的蚯蚓区系十分特殊, 且与大陆地区蚯蚓存在扩散与迁移。然而, 当海南岛与周围大陆断开形成独立岛屿后, 海南岛蚯蚓如何为适应海南岛的环境而逐渐形成现在岛内的分布与区系, 仍然是一个值得研究和探讨的问题。因此, 本研究在海南岛全面调查和采集蚯蚓标本, 整理海南岛地区的蚯蚓物种组成及其地理分布特征, 并联合5个线粒体基因COⅠ、COⅡ、ND1、12S rRNA和16S rRNA构建海南岛蚯蚓的分子系统发育树, 推测其分化时间和祖先分布区域, 探讨海南岛蚯蚓在岛内的分化与扩散过程。研究结果表明: (1)海南岛共有蚯蚓6科9属122种, 巨蚓科蚯蚓为优势科, 且全部为环毛类蚯蚓, 其中103种为海南岛特有种。蚯蚓物种数沿海拔呈先增加后减少的趋势, 在800‒1,000 m最大; (2)海南岛环毛类蚯蚓不同水平的遗传距离与我国巨蚓科蚯蚓的遗传距离区间基本一致。物种水平上, 美丽远盲蚓(Amynthas scitulus*)与纬向远盲蚓(A. zonarius)的基因遗传距离最小。在亚种水平, 保宁腔蚓指名亚种(Metaphire magna magna)和保宁腔蚓小型亚种(M. magna minuscula)的遗传距离均接近整体的物种水平。在种群水平, 等毛远盲蚓(A. homosetus)不同种群间遗传距离均接近整体的亚种水平; (3)海南岛环毛类蚯蚓可划分为7个类群, 其祖先于68.26 Ma开始分化, 可能起源于吊罗山。在新生代, 7个类群均得到较大发展。通过对海南岛蚯蚓组成及系统发育的梳理, 不仅为我国蚯蚓物种多样性研究提供基础资料, 也为岛屿蚯蚓物种系统发育关系分析提供科学参考。     

不同培肥措施对低肥力土壤生态系统蚯蚓种群数量的影响

通过在华北低肥力农田生态系统进行的不同土壤培肥试验 ,研究了不同秸秆还田方式和施肥措施对土壤中蚯蚓种群的影响。研究结果表明 :一年中 5次调查到的蚯蚓属于同一个目——后孔寡毛目 ,3科、6属、7种 ;在 7种蚯蚓中 ,天锡杜拉蚓Drawida gisti是该地区较低肥力土壤中的蚯蚓优势种。蚯蚓种群数量的季节变化趋势为 8月 >9月 >4月 >5月 >11月。在低肥力土壤上 ,单独施用化肥与对照相比可以增加蚯蚓的数量 ,施用化肥对蚯蚓的影响程度依赖于有机物的投入情况 ,没有有机物投入或只有麦秸还田条件下 ,施用化肥会对土壤中蚯蚓的种群数量产生负面影响 ,随着时间的延长 ,这种影响表现得比较明显。在同时用玉米秸和麦秸还田的情况下 ,施用化肥对蚯蚓的种群数量影响较小。有机肥的施入可以增加蚯蚓的种群数量 ,随着时间的延长这种趋势愈为明显。在几种秸秆还田的投入方式中 ,蚯蚓种群数量的大小次序为双倍麦还 >玉米麦秸全还 >麦还。因此在施用化肥的时候应当同时进行有机物投入 ,以保持土壤中良好的生物状况 ,这也是土壤培肥的重要环节。     

西双版纳热带次生林和橡胶林蚯蚓数量分布及其与环境因子的关系

2006年1月,通过样地调查法比较了西双版纳次生林和橡胶林2种不同土地利用方式的蚯蚓数量分布状况及其与环境因子的关系.结果表明:自然林转变为橡胶林后,土壤养分及植物有机碳输入均明显减少,同时蚯蚓群落发生较大的变化:在橡胶林,热带外来种蚯蚓无围海蚓的密度(52.27 ind·m-2)显著高于林地内本地种蚯蚓远环蚓的密度(0.53 ind·m-2),但在次生林,2种蚯蚓的密度和生物量无显著差异;橡胶林本地种蚯蚓远环蚓的密度(0.53 ind·m-2)和生物量(0.32 g·m-2)与次生林(6.93 ind·m-2、7.76 g·m-1)相比呈下降趋势.无围海蚓的幼蚓在其种群中占有较大比例,类似增长型种群年龄结构,而远环蚓则主要以成蚓为主.相关分析表明,橡胶林的细根生物量与无围海蚓的密度呈显著正相关.由预示蚯蚓群落结构的这种变化可能与橡胶林土地利用引起的植被变化、土壤养分下降等因素密切相关,同时外来种蚯蚓与本地种蚯蚓本身的生态适应对策也不可忽视.     

中国东、南部的3个腔蚓属蚯蚓新物种

本文报道了寡毛纲(Oligochaeta)巨蚓科(Megascolecidae)腔蚓属(Metaphire)3个新发现物种,分别是象头山腔蚓(M. xiangtoumontis Dong & Jiang sp. nov.),韩摆渡腔蚓(M. hanbaiduensis Dong & Sun sp. nov.)和长白山腔蚓(M. changbaimontis Dong & Shen sp. nov.)。象头山腔蚓受精囊孔2对,位于7/8 ~ 8/9节间,属于M. insulana物种群。韩摆渡腔蚓受精囊孔3对,位于6/7 ~ 8/9节间,属于M. houlleti物种群。长白山腔蚓受精囊孔2对,位于6/7 ~ 7/8节间,属于M. glandularis物种群。所有新物种均附有形态学描述、图片、与相似物种的形态学比较及与GenBank上亲缘关系相近物种的遗传距离计算分析。以上结果丰富了我国腔蚓属蚯蚓的物种多样性,并首次记录了采集于长白山国家级自然保护区的巨蚓科蚯蚓新物种。     

蚯蚓吞食过程中土壤理化性质与放线菌多样性的变化特征

为探明土壤理化性质与放线菌多样性在蚯蚓吞食过程中的变化特征,将蚯蚓生活土壤、肠道内容物和蚓粪视作蚯蚓吞食前、中、后阶段的3种特殊生境土壤,采用纯培养法分离纯化3阶段中的放线菌;克隆文库法分析3阶段中的放线菌多样性;国标法测定吞食前、中、后土壤的基本理化性质;并利用主成分分析与相关性分析法分析土壤理化性质与放线菌多样性的相关性。结果显示:从生活土壤、肠道内容物和蚓粪中分别获得27株、15株和17株放线菌,形态、培养特征及16S rDNA序列鉴定生活土壤放线菌分为链霉菌属、拟诺卡氏菌属、束丝放线菌属,肠道内容物和蚓粪放线菌均属链霉菌属;放线菌多样性从生活土壤、蚓粪、肠道内容物依次递减。生活土壤文库含40个OTUs,分为11科,未知菌占24%,类诺卡氏菌科是优势菌群;肠道内容物文库含20个OTUs,分为6科,未知菌占3.3%,微杆菌科是优势菌群;蚓粪文库含30个OTUs,分为6科,未知菌占11.7%,链霉菌科是优势菌群。3种土壤全磷含量无显著差异,生活土壤其余理化含量均最低,肠道内容物速效氮含量最高,蚓粪的有机质、全氮、钾及有效磷、钾含量最高。主成分分析和相关性分析显示:蚯蚓吞食过程中,土壤有效磷、全氮、全钾、速效钾和有机质含量对放线菌多样性影响较大,其中全氮、有效磷与放线菌多样性显著负相关,相关系数分别为-0.998、-1,从而为明确蚯蚓、土壤与放线菌的相互关系提供了理论依据。     

四川陆栖寡毛类两新种记述：寡毛纲：链胃蚓科,棘蚓科

记述了采集于四川省酉阳县的链胃科、杜拉属Drawida蚯蚓一新种,双斑杜拉蚓Drawida bimaculata sp.nov.和在四川省汉源县采到的棘蚓科,毛蚓属Plutellus蚯蚓一新种,汉源毛蚓Plutellus hanyuangensis sp.nov.。毛蚓属是我国首次记录。     

Non-native earthworms alter the assembly of a meadow plant community

Non-native earthworms can alter ecosystems by modifying soil structure, depredating seeds and seedlings, and consuming soil organic matter, yet the initial responses of plant communities to earthworm invasions remain poorly understood. We assessed the effect of non-native earthworms on seedling survival during germination and after establishment using six native and six non-native plant species grown from seed in single- and multi-species experimental mesocosms. We examined the extent to which earthworms (1) influenced seedling survival, (2) selectively depredated native versus non-native plants, (3) impacted establishment based on seed size and/or root morphology, and (4) shaped community assembly. The effect of earthworms on seedling survival varied temporally and among species but inconsistently with respect to species origin. Differences in seed/seedling survival translated to changes in community assembly. Earthworms tended to reduce species abundance, richness, evenness, and diversity in multi-species mesocosms and led to the divergence of communities by treatment. In general, species with large seeds and fibrous roots dominated communities with earthworms present, whereas species with small seeds and taproots only persisted in multi-species mesocosms without earthworms. Our findings suggest that earthworms act as ecological filters in the early stages of invasion to shape community composition based on plant morphological traits.

     

Drought stress in rice (Oryza sativa L.) is enhanced in the presence of the compacting earthworm Millsonia anomala

Earthworms increase growth of most plant species through a number of poorly investigated mechanisms. We tested the hypothesis that earthworm modifications of soil structure and the resulting changes in water availability to plants explain this positive effect. Addition of endogeic earthworms Millsonia anomala induced a 40% increase in shoot biomass production and a 13% increase in CO₂ assimilation rate of well watered rice plants grown in pots. Conversely, when plants were subjected to water deficit, presence of earthworms had no effect on shoot biomass production and a negative impact on CO₂ assimilation rate (−21%). Early stomatal closure in presence of earthworms indicated lower water availability. The hypothesis that earthworms improve plant biomass production through soil physical structure modification was thus rejected. Three hypotheses were tested to explain this decrease in water availability: (i) a decrease in soil water retention capacity, (ii) an increase in evaporation from the soil or/and (iii) an increase in plant transpiration. Results showed that earthworms significantly reduced soil water retention capacity by more than 6%, but had no effect on evaporation rate. Water losses through transpiration were greater in the presence of earthworms when the soil was maintained at field capacity, but this was not the case under drought conditions. This experiment showed that the endogeic compacting earthworm M. anomala significantly increased plant photosynthesis by an undetermined mechanism under well-watered conditions. However, photosynthesis was reduced under drought conditions due to reduced soil water retention capacity.     

Interactions of earthworms with Atrazine-degrading bacteria in an agricultural soil

In the last 10 years, accelerated mineralization of Atrazine (2-chloro-ethylamino-6-isopropylamino-s-triazine) has been evidenced in agricultural soils repeatedly treated with this herbicide. Here, we report on the interaction between earthworms, considered as soil engineers, and the Atrazine-degrading community. The impact of earthworm macrofauna on Atrazine mineralization was assessed in representative soil microsites of earthworm activities (gut contents, casts, burrow linings). Soil with or without earthworms, namely the anecic species Lumbricus terrestris and the endogenic species Aporrectodea caliginosa, was either inoculated or not inoculated with Pseudomonas sp. ADP, an Atrazine-degrading strain, and was either treated or not treated with Atrazine. The structure of the bacterial community, the Atrazine-degrading activity and the abundance of atzA, B and C sequences in soil microsites were investigated. Atrazine mineralization was found to be reduced in representative soil microsites of earthworm activities. Earthworms significantly affected the structure of soil bacterial communities. They also reduced the size of the inoculated population of Pseudomonas sp. ADP, thereby contributing to the diminution of the Atrazine-degrading genetic potential in representative soil microsites of earthworm activities. This study illustrates the regulation produced by the earthworms on functional bacterial communities involved in the fate of organic pollutants in soils.     

Distribution, prevalence and intensity of earthworm populations in arable land and grassland in Scotland

The earthworms of arable and pasture fields from 100 randomly chosen arable farms were identified and counted and their relationship with soil factors examined. Thirteen species of earthworm were recorded including Lumbricus friendi from Scotland for the first time. The most prevalent and numerous species were Aporrectodea longa, Aporrectodea caliginosa and Lumbricus terrestris. All earthworm species had cosmopolitan distributions apart from Aporrectodea nocturna and Lumbricus festivus which were confined to southern/central Scotland. Four soil characteristics (% moisture, % sand, % organic matter and pH) shared no relationship with species recorded but tillage may have had a detrimental effect on species composition and size of population. It is suggested that the magnitudes of the earthworm populations found, particularly in permanent pasture, were important in maintaining soil structure and fertility, and the spread of the New Zealand flatworm (Artioposthia triangulata), an obligate predator of earthworms, could have a considerable impact on agricultural productivity and wildlife in Scotland.     

Invasion of exotic earthworms into ecosystems inhabited by native earthworms

The most conspicuous biological invasions in terrestrial ecosystems have been by exotic plants, insects and vertebrates. Invasions by exotic earthworms, although not as well studied, may be increasing with global commerce in agriculture, waste management and bioremediation. A number of cases has documented where invasive earthworms have caused significant changes in soil profiles, nutrient and organic matter dynamics, other soil organisms or plant communities. Most of these cases are in areas that have been disturbed (e.g., agricultural systems) or were previously devoid of earthworms (e.g., north of Pleistocene glacial margins). It is not clear that such effects are common in ecosystems inhabited by native earthworms, especially where soils are undisturbed. We explore the idea that indigenous earthworm fauna and/or characteristics of their native habitats may resist invasion by exotic earthworms and thereby reduce the impact of exotic species on soil processes. We review data and case studies from temperate and tropical regions to test this idea. Specifically, we address the following questions: Is disturbance a prerequisite to invasion by exotic earthworms? What are the mechanisms by which exotic earthworms may succeed or fail to invade habitats occupied by native earthworms? Potential mechanisms could include (1) intensity of propagule pressure (how frequently and at what densities have exotic species been introduced and has there been adequate time for proliferation?); (2) degree of habitat matching (once introduced, are exotic species faced with unsuitable habitat conditions, unavailable resources, or unsuited feeding strategies?); and (3) degree of biotic resistance (after introduction into an otherwise suitable habitat, are exotic species exposed to biological barriers such as predation or parasitism, “unfamiliar” microflora, or competition by resident native species?). Once established, do exotic species co-exist with native species, or are the natives eventually excluded? Do exotic species impact soil processes differently in the presence or absence of native species? We conclude that (1) exotic earthworms do invade ecosystems inhabited by indigenous earthworms, even in the absence of obvious disturbance; (2) competitive exclusion of native earthworms by exotic earthworms is not easily demonstrated and, in fact, co-existence of native and exotic species appears to be common, even if transient; and (3) resistance to exotic earthworm invasions, if it occurs, may be more a function of physical and chemical characteristics of a habitat than of biological interactions between native and exotic earthworms.     

蚯蚓在我国南方土壤修复中的应用

蚯蚓作为生物量最大的土壤动物, 对土壤生态系统和环境质量影响深远。本研究介绍了华南地区主要应用的皮质远盲蚓(Amynthas corticis)、毛利远盲蚓(A. morrisi)、壮伟远盲蚓(A. robustus)、参状远盲蚓(A. aspergillum)、南美岸蚓(Pontoscolex corethrurus)和赤子爱胜蚓(Eisenia fetida)的生态特征, 阐述了它们与土壤pH值、酶活性、金属富集和有效性改变、孔道和微团聚体形成之间的紧密关系: (1)蚯蚓生存的土壤酸碱性范围较广(pH为3.8-7.9), 其存活率与土壤类型、有机质含量和成分、土壤污染程度和蚯蚓种类相关; (2)肠道内、蚓粪和蚓触圈的酶活性分别表征了蚯蚓取食喜好、土壤养分循环及微生物种群特征; (3)蚯蚓能够富集不同种类的金属并改变其有效性, 这些变化具有蚓种间、金属种类间和土壤类型之间的差异; (4)蚯蚓活动及其生产的蚓粪能改变土体结构、产生孔道、影响土壤团聚体数量、大小和分布。蚯蚓的上述作用使其在解决中国南方红壤酸化、土壤金属污染、茶园土壤养分不平衡、高速公路建设临时用地土壤损毁等方面具有广阔的应用前景。目前, 由于华南远盲蚓的生理特征差异研究较少, 远盲蚓繁育技术的缺乏一定程度上限制了这些蚯蚓在中型和大型尺度下应用技术的研究和推广。有必要进一步挖掘蚯蚓在土壤修复中的潜力, 进行蚯蚓主导的相关技术研发, 深入探讨其影响机制。     

The population of earthworms (Lumbricidae) in the main types of dark coniferous forests in Pechora-Ilych Nature Reserve

The population of earthworms has been studied in the main types of old-growth dark coniferous forests of Pechora-Ilych Nature Reserve (Komi Republic) that have not been subject to anthropogeniс impact for a long time. Ten species of earthworms have been identified. The greatest diversity (7 species), abundance, and biomass of earthworms has been revealed in the tall-grass fir–spruce forests. P. diplotetratheca had the greatest abundance. E. nordenskioldi nordenskioldi and E. atlavinyteae had the greatest biomass. The lowest species diversity (3 species), abundance, and biomass of earthworms have been found in the largefern, blueberry–green moss, and sphagnum–horsetail fir–spruce forests. The role of deadwood in maintaining the species diversity of Lumbricidae in dark coniferous forests has been demonstrated. The complexes of Lumbricidae have been considered in anthropogenically disturbed territories, where the following species with a habitat range to the south of the northern and middle taiga have been identified: L. rubellus, A. rosea, A. caliginosa caliginosa, and E. fetida.     

Earthworm-mycorrhiza interactions can affect the diversity, structure and functioning of establishing model grassland communities

Both earthworms and arbuscular mycorrhizal fungi (AMF) are important ecosystem engineers co-occurring in temperate grasslands. However, their combined impacts during grassland establishment are poorly understood and have never been studied. We used large mesocosms to study the effects of different functional groups of earthworms (i.e., vertically burrowing anecics vs. horizontally burrowing endogeics) and a mix of four AMF taxa on the establishment, diversity and productivity of plant communities after a simulated seed rain of 18 grassland species comprising grasses, non-leguminous forbs and legumes. Moreover, effects of earthworms and/or AMF on water infiltration and leaching of ammonium, nitrate and phosphate were determined after a simulated extreme rainfall event (40 l m(-2)). AMF colonisation of all three plant functional groups was altered by earthworms. Seedling emergence and diversity was reduced by anecic earthworms, however only when AMF were present. Plant density was decreased in AMF-free mesocosms when both anecic and endogeic earthworms were active; with AMF also anecics reduced plant density. Plant shoot and root biomass was only affected by earthworms in AMF-free mesocosms: shoot biomass increased due to the activity of either anecics or endogeics; root biomass increased only when anecics were active. Water infiltration increased when earthworms were present in the mesocosms but remained unaffected by AMF. Ammonium leaching was increased only when anecics or a mixed earthworm community was active but was unaffected by AMF; nitrate and phosphate leaching was neither affected by earthworms nor AMF. Ammonium leaching decreased with increasing plant density, nitrate leaching decreased with increasing plant diversity and density. In order to understand the underlying processes of these interactions further investigations possibly under field conditions using more diverse belowground communities are required. Nevertheless, this study demonstrates that belowground-aboveground linkages involving earthworms and AMF are important mediators of the diversity, structure and functioning of plant communities.     

Earthworm numbers,biomass and respiratory metabolism in a beech woodland-Wytham Woods,Oxford

Summary The mean annual population density of earthworms was found to be 164.6 m^-2 during a period of detailed study between October 1971 and September 1972. In a year of less detailed study between November 1972 and October 1973 the population density was 117.5 m^-2 (139.8 m^-2 when the type of extraction method was allowed for). Mean biomass densities in the two years of investigation were 41.0 g preserved wet wt m^-2 (1971–1972) and 38.6 (possibly 39.2) g preserved wet wt m^-2 (1972–1973).Comparison of the Brogden's Belt population and biomass densities with those reported from other woodlands indicates that soil type is more important than leaf litter type in determining the numerical abundance of earthworms. Population densities are lower in beechwoods on mor soils, mor soils also support significantly fewer species. As with numbers, mean biomass density in beechwoods on mor soils was significantly lower than that occurring in beechwoods on mull soils; the latter, in turn, being lower than those found in mixed deciduous woods on mull soils. Unlike population density biomass density is influenced by both soil and litter type, this is discussed by reference to mean body weights and food quality as reflected by tannin, nitrogen and carbohydrate content.The annual respiratory metabolism of the Brogden's Belt earthworms was calculated to be between 10.7 and 13.41 O₂ m^-2 a^-1, which is equivalent to between 4.1 and 5.1% of the total soil metabolism. A production/biomass ratio of 0.49–0.58 was estimated, as was a net population efficiency of 22%.