湛江高桥红树林和盐沼湿地的大型底栖动物次级生产力

为了比较湛江高桥潮间带不同植物生境的大型底栖动物次级生产力,根据2010年4个季度湛江高桥潮间带生境的大型底栖动物数据,运用Brey经验公式计算不同植物生境的大型底栖动物次级生产力.结果表明:湛江高桥红树林和盐沼湿地不同生境大型底栖动物平均次级生产力为11.77 g AFDM·m-2·a-1.其中,无瓣海桑生境次级生产力最高,为18.16 g AFDM·m-2·a-1,其次是桐花树、盐地鼠尾粟和木榄生境,分别为17.67、8.34和2.92 g AFDM·m-2·a-1.在4种生境中,木榄生境的年生产力/年均生物量(P/B)最高,为2.38,其次是无瓣海桑、盐地鼠尾粟和桐花树生境,分别为1.23、0.99和0.48.湛江高桥潮间带不同植物生境大型底栖动物次级生产力和P/B值的差异主要与总有机碳含量、食物类型和动物个体大小有关.     

不同红树植物群落中大型底栖动物群落的比较

2005—2006年对广东湛江红树林国家级自然保护区湿地三种红树植物群落(白骨壤 桐化树群落、桐花树群落、木榄 桐花树群落)的大型底栖动物群落特征进行了分析研究。白骨壤 桐化树群落大型底栖动物群落的物种数、栖息密度、生物量、丰富度指数和多样性指数均最高,优势度指数居中,均匀度指数略低于桐花树群落;桐花树群落大型底栖动物物种数急剧减少,尤其是底内型、底上附着型和穴居型种类减少明显,生物量和栖息密度下降到最低,由于个体数种间分配较为均匀而导致优势度指数下降而均匀度指数增高,虽丰富度指数略低于白骨壤 桐化树群落,但多样性指数接近于白骨壤 桐化群落;木榄 桐花树群落,大型底栖动物群落的物种数,尤其是穴居型和底内型种数继续减少,但生物量和栖息密度有所上升,个体数种间分配不均匀而使优势度指数增高而均匀度下降,加上丰富度指数最低,故多样性指数最小。白骨壤 桐化树群落优势种的生活型是底内型和穴居型;桐花树和木榄 桐花树群落优势种的生活型均是穴居型。三种红树植物群落中的大型底栖动物群落的GS/GSB分别为0.48、0.63、0.80。相同红树植物群落大型底栖动物群落结构都较为相似,木榄 桐花树群落的相似性最高,而不同红树植物群落大型底栖动物群落特征的差异明显,反映了不同红树群落对底栖动物群落作用的差别,同时也展示了各种大型底栖动物对不同红树群落生境的适应情况。     

几种红树植物模拟湿地系统对污水中重金属的净化效应

在温室中建立无瓣海桑(Sonneratia apetala)、桐花树(Aegiceras corniculatum)、木榄(Bruguiera gymnorrhiza)、秋茄(Kandelia candel)、无瓣海桑+木榄(S.apetala+ B.gymnorrhiza)、无瓣海桑+秋茄(S.apetala+ K.candel)、无瓣海桑+桐花树(S.apetala+ A.corniculatum)等7 种模拟湿地系统, 用正常浓度污水(SW)、5 倍浓度污水(FW)和10 倍浓度污水(TW)3 种人工配制的生活污水分别对7 种模拟湿地系统进行污灌6 个月。结果发现: 7 种人工模拟湿地生态系统对污水中重金属的净化率与红树植物种类以及污水中重金属浓度有关。当灌正常浓度污水时, 7 种人工模拟湿地生态系统对Pb 和Mn 的总净化率以无瓣海桑+秋茄最大, 对Cu 的总净化率以桐花树最大, 对Zn 和 Cd 的总净化率以无瓣海桑+桐花树最大。当灌5 倍浓度污水时, 7 种人工模拟湿地生态系统对Pb 总净化率以无瓣海桑+桐花树最大, 对Cu 和Cd总净化率以无瓣海桑+秋茄最大, 对Zn 和Mn 总净化率以无瓣海桑+木榄最大。当灌10 倍浓度污水时, 7 种人工模拟湿地生态系统对Pb、Mn 的总净化率以无瓣海桑+秋茄最大, 对Cu 的总净化率以秋茄最大, 对Zn 的总净化率以无瓣海桑+木榄最大, 对Cd 总净化率以无瓣海桑最大。重金属有95%以上是积累在土壤中, 表明土壤子系统是人工模拟湿地系统净化的主体。     

无瓣海桑与乡土红树植物混交对林地大型底栖动物的影响

选择广东省雷州市附城镇和珠海市淇澳岛沿海1年生无瓣海桑人工林,分别在林下混种乡土红树植物红海榄或木榄幼苗,对混交林和无瓣海桑纯林林地大型底栖动物群落进行比较,探讨无瓣海桑与乡土红树植物混交对林地大型底栖动物的影响。结果显示,混交林和无瓣海桑纯林之间大型底栖动物群落的优势种存在差异;相似性分析检验(One-Way ANOSIM)、等级聚类和非参数多变量标序结果均表明1年无瓣海桑+红海榄混交林、1年无瓣海桑+木榄混交林和1年无瓣海桑纯林之间大型底栖动物群落结构差异显著。研究还发现在无瓣海桑人工林林下混交红海榄或木榄这两种乡土红树植物,可提高林地底栖动物的生物量和物种多样性。BIOENV分析说明大型底栖动物分布与红海榄或木榄这两种混交的乡土树种的凋落物量密切相关,这进一步证实了混交乡土红树植物对林地底栖动物多样性和分布的影响。两个研究地实验结果均显示,在无瓣海桑林下种植木榄的效果要优于红海榄,表现在木榄的平均苗高、凋落物量、凋落物量占群落凋落物总量百分比和幼苗成活率均高于红海榄,其对提高林地大型底栖动物生物量和物种多样性的效果也明显优于红海榄。     

不同红树植物群落中大型底栖动物群落的比较

2005—2006年对广东湛江红树林国家级自然保护区湿地三种红树植物群落（白骨壤+桐化树群落、桐花树群落、木榄+桐花树群落）的大型底栖动物群落特征进行了分析研究。白骨壤+桐化树群落大型底栖动物群落的物种数、栖息密度、生物量、丰富度指数和多样性指数均最高，优势度指数居中，均匀度指数略低于桐花树群落；桐花树群落大型底栖动物物种数急剧减少，尤其是底内型、底上附着型和穴居型种类减少明显，生物量和栖息密度下降到最低，由于个体数种间分配较为均匀而导致优势度指数下降而均匀度指数增高，虽丰富度指数略低于白骨壤+桐化树群落，但多样性指数接近于白骨壤+桐化群落；木榄+桐花树群落，大型底栖动物群落的物种数，尤其是穴居型和底内型种数继续减少，但生物量和栖息密度有所上升，个体数种间分配不均匀而使优势度指数增高而均匀度下降，加上丰富度指数最低，故多样性指数最小。白骨壤+桐化树群落优势种的生活是底内型和穴居型；桐花树和木榄+桐花树群落优势种的生活型均是穴居型。三种红树植物群落中的大型底栖动物群落的GS/GSB分别为0.48、0.63、0.80。相同红树植物群落大型底栖动物群落结构都较为相似，木榄+桐花树群落的相似性最高，而不同红树植物群落大型底栖动物群落特征的差异明显，反映了不同红树群落对底栖动物群落作用的差别，同时也展示了各种大型底栖动物对不同红树群落生境的适应情况。     

不同生态恢复阶段无瓣海桑人工林湿地中大型底栖动物群落的演替

对广东省珠海市淇澳岛红树林自然保护区恢复背景相同的6个月林龄、3a林龄、6a林龄无瓣海桑人工林和光滩湿地中的大型底栖动物群落进行了比较研究。结果显示:处于不同生态恢复阶段的无瓣海桑人工林林地中大型底栖动物优势种(Y>0.02)存在差异;相似性分析检验(One-Way ANOSIM)表明4种生境间大型底栖动物群落结构差异显著。等级聚类和非参数多变量标序也显示4种生境间的大型底栖动物群落结构差异显著。相似性百分比分析(SIMPER)显示,各生境动物群落主要特征及群落间的差异主要是受各生境大型底栖动物优势种所决定。研究结果还表明:光滩、6个月林龄、3a林龄无瓣海桑人工林湿地中大型底栖动物群落的Shannon-Wiener多样性指数、Pielou均匀度指数和Margalef丰富度指数呈上升趋势,这说明无瓣海桑种植的前3a,随着植被的迅速恢复,大型底栖动物多样性增加明显。BIOENV分析结果表明:3a林龄之前的无瓣海桑人工林湿地中,大型底栖动物物种多样性主要受植被特征的影响,快速恢复的植被提供了荫蔽,既减轻了高温和水分蒸发对大型底栖动物的胁迫,又给动物提供了一个躲藏和逃避敌害的场所。而相对于植被的快速恢复,无瓣海桑人工林林地土壤理化性质的改变要缓慢和滞后得多。但随着恢复时间的延长,6a林龄无瓣海桑人工林湿地中的大型底栖动物群落的上述3个物种多样性指数反而明显下降。这是由于6a林龄无瓣海桑人工林林地土壤已具有红树林酸性硫酸盐土的基本特征,林地土壤理化性质的显著变化影响了大型底栖动物群落,还可能与无瓣海桑提供了和乡土红树植物不同营养水平(如C/N比)和单宁含量的"凋零物"食物源有关,具体机制还需进一步研究。     

东、黄海典型站位底栖动物粒径谱研究

对东、黄海典型站位底栖动物粒径谱进行了首次研究 ,得出 Sheldon粒径谱呈典型的双峰模式 ,两个峰形分别对应着大型底栖动物和小型底栖动物的生物量分布。小型底栖动物峰值出现在粒级 6～ 10 ,大型底栖动物峰值在 13～ 14。两峰值之间的波谷位置在粒级 10～ 11,是大型、小型底栖动物的过渡带 ,生物量主要由大型动物的幼龄个体构成。E3站大型动物幼龄个体大量出现 ,使该站的过渡粒级上出现峰值 ,粒径谱呈单峰形。同时构建了正态化粒径谱 ,在此基础上对不同生态系统的比较进行了初步尝试。     

珠海市淇澳岛红树林群落组成初步研究

本文研究了珠海市淇澳岛红树林的组成、群落类型、群落结构外貌、生态序列及红树林内的主要动物种类.初步调查结果显示:该地区有红树植物7科9种,半红树5科6种,红树林伴生植物4科8种,脊椎动物中鸟类10目20科46种.淇澳岛红树林群落类型主要有:秋茄-桐花树-老鼠筋群落、桐花树-芦苇群落、老鼠筋群落、无瓣海桑群落、木榄-银叶树群落.本文还针对珠海淇澳岛红树林湿地资源现状提出可持续发展建议.     

淇澳岛两种红树林区大型底栖动物群落比较

对珠海淇澳岛天然秋茄(Kandelia candel)与人工恢复无瓣海桑(Sonneratia apetala)两种红树林林地间大型底栖动物群落结构进行了比较研究.结果显示,天然秋茄林与人工无瓣海桑林林地大型底栖动物种类组成及数量分布都存在差异,秋茄林大型底栖动物平均生物量、栖息密度分别为668.64g·m^-2和8370个·m^-2,无瓣海桑林分别为511.64g·m^-2和4748个·m^-2.两个红树林剖面不同潮位带大型底栖动物结构存在差异,秋茄林生物量和栖息密度的最高值都在中潮带,主要是由节肢动物和软体动物所主导:无瓣海桑林栖息密度的最高值也是在中潮带,生物量变化比较不规则.两个红树林剖面大型底栖动物群落特征的季节变化较明显,生物量都是秋季处于最高水平:栖息密度都是冬季处于最高水平,秋季最低:生物多样性都是夏季处于最低,但最高值秋茄林剖面是在春季,而无瓣海桑林剖面则是秋季.研究结果表明,林地底质和盐度等生境条件以及红树林群落的高度、郁闭度和林下通风透光等生态特征的差异是造成红树林底栖动物群落差异的重要原因.     

两种相手蟹对不同红树植物叶片取食的偏好性

大型底栖动物是红树林生态系统中重要的消费者,在红树林的营养物质转化和能量传递过程中具有重要作用。通过设置室内模拟饲喂实验,研究了华南沿海红树林大型底栖动物的常见种--无齿相手蟹(Sesarma dehaani)和双齿相手蟹(S.bidens)对外来红树植物无瓣海桑(Sonneratia apetala)和乡土红树植物秋茄(Kandelia obovata)、桐花树(Aegiceras corniculatum)凋落叶片的取食特性。结果表明,随着淋溶作用的进行,各树种叶片中单宁含量和C/N均逐渐降低,两个数值的变化幅度以秋茄叶片最大,分别达(45.85±7.74)g·kg-1和11.24%,无瓣海桑为(45.15±11.14)g·kg-1和2.43%,桐花树则为(18.75±15.15)g·kg-1和-1.36%。随着叶片中单宁含量和C/N的降低,两种相手蟹的取食量相应地呈逐渐上升的趋势,双齿相手蟹从3.097g到10.079g和无齿相手蟹则从2.480g到8.723g。两种相手蟹种间的取食特性差异不显著(P0.05),但对不同树种的取食上却存在显著差异,其中对桐花树的取食偏好性显著差于无瓣海桑和秋茄(P0.01),这主要与桐花树叶片的高单宁含量和低营养价值有关。研究结论说明大面积引种外来物种无瓣海桑将可能对华南沿海红树林生态系统的食物链结构产生潜在影响,并为红树林的生态恢复和科学造林提供了参考。     

Competitive Control of an Exotic Mangrove Species: Restoration of Native Mangrove Forests by Altering Light Availability

Recently, many studies have focused on the possibility of restoring mangrove ecosystems by introducing fast‐growing mangroves. However, methods for managing an exotic fast‐growing species to restore mangrove ecosystems and at the same time preventing invasion by introduced species remains unclear. Sonneratia apetala Buch‐Ham is one example of an exotic mangrove with both high ecological value and potential risk for invasion after introduction. To investigate the possibility of reducing the potential for invasion by altering light availability, we simulated different irradiances of S. apetala understory in the greenhouse. For each irradiance treatment, three levels of competition between S. apetala and native mangroves Aegiceras corniculatum (L.) were used: no competition, intraspecific competition and interspecific competition. Compared with A. corniculatum, S. apetala showed a significantly higher growth rate for both height and biomass accumulation under full irradiation. Compared to the full irradiation treatment, the shading treatment significantly reduced the height, total biomass and biomass allocation to leaves of S. apetala by 61.31, 71.0, and 76.2%, respectively, whereas the growth of A. corniculatum was not affected. The results suggested that lowering light availability could inhibit the growth of S. apetala and increase the competitiveness of A. corniculatum. Planting introduced fast‐growing mangroves at a density of approximately 2,000 plants/hm² is an effective strategy for preventing potential invasion and restoring wetland habitats. By taking advantage of the differences in shade tolerance between fast‐growing exotic mangroves and native mangroves, introduction of fast‐growing mangroves in coastal areas could have huge potential for reforesting mangrove ecosystems.     

不同生长光强下8种红树植物幼苗叶片的光响应特征

为探讨红树植物光适应的生理生态策略,对6种真红树植物[无瓣海桑(Sonneratia apetala)、秋茄(Kandelia candel)、木榄(Bruguiera gymnorrhiza)、桐花树(Aegiceras corniculatum)、老鼠簕(Acanthus ilicifolius)、卤蕨(Acrostichum aureum)]和2种半红树植物[银叶树(Heritiera littoralis)、黄槿(Hibiscus tiliaceus)]的1 a生幼苗在不同生长光强(自然光强的100%、45%、30%、10%)下的光合光响应特征进行了研究。结果表明,不同生长光强对红树植物光响应特征的影响因物种而异,遮荫显著提高了秋茄和木榄的最大净光合速率(Pmax),而对其他红树植物的Pmax没有显著影响;秋茄在45%光强下具有较高的Pmax,木榄的Pmax则在45%和30%光强下显著高于其他2个处理。随着生长光强的下降,秋茄幼苗叶片的光饱和点显著上升,木榄、老鼠簕和卤蕨的光补偿点呈下降趋势,木榄和卤蕨的表观量子效率升高的同时暗呼吸速率下降。木榄、老鼠簕和卤蕨具有较强的耐荫性,...     

Wetland changes and mangrove restoration planning in Shenzhen Bay, Southern China

Mangrove forests and associated gei wai (excavated ponds used for shrimp and fish farming) provide important ecosystem services in Shenzhen Bay. Much of the mangrove and gei wai wetlands, however, have been lost because of intensified human activities in the past 30 years. Using five-phase remote-sensing images, we describe the recent history of the spatial–temporal dynamics for the wetlands in the bay. From 1986 to 2007, mangrove area increased from 1.8 to 4.8 km², while the area of gei wai decreased from 36.6 to 17.2 km². Reclamation of gei wai mainly occurred in western and northern Shenzhen Bay, and changed the tidal water environment. The bay has five typical mangrove communities: Avicennia marina + Kandelia candel + Aegiceras corniculatum, Kandelia candel + Aegiceras corniculatum + Acanthus ilicifolius, Bruguiera gymnorrhiza + Excoecaria agallocha, Aegiceras corniculatum, and Sonneratia apetala + Sonneratia caseolaris. The distribution of these communities and their dominant species in the bay exhibit a spatial pattern and temporal (successional) sequence. We describe a mangrove restoration program based on the mangrove successional sequence and the interaction of mangrove and gei wai in the bay. We have planned six mangrove protection and restoration projects in closed areas, semiclosed areas, and open areas to reconstruct the ecological integrity of the entire Shenzhen Bay.     

Allometric growth and carbon storage in the mangrove Sonneratia apetala

Allometric growth reflects different allocation patterns and relationships of different components or traits of a plant and is closely related to ecosystem carbon storage. As an introduced species, the growth and carbon storage of Sonneratia apetala are still unclear. To derive allometric relationships of the mangrove S. apetala and to estimate carbon storage in mangrove ecosystems, we harvested 12 individual Sonneratia apetala trees from four different diameter classes in the Futian National Nature Reserve, Guangdong, China. Allometric growth models were fitted. The results showed that diameter at breast height (DBH) and wood density were better variables for predicting plant biomass (including above- and below-ground biomass) than plant height. There were significant power function relationships between biomass and DBH, with a mean allometric exponent of 2.22, and stem biomass accounted for 97% of the variation in S. apetala total biomass. Nearly isometric scaling relationships were developed between stem biomass and other biomass components. To better understand the carbon stocks of the S. apetala ecosystem, we categorized all trees into five age classes and quantified vegetation carbon storage. The S. apetala vegetation carbon storage ranged from 96.48 to 215.35 Mg C ha^?1, and the carbon storage significantly increased with stand age. The allometric equations developed in this study are useful to estimate biomass and carbon storage of S. apetala ecosystems.

     

中国红树林生态系统保护空缺分析

红树林作为海岸带重要的生态系统类型,具有维持海岸生物多样性、防风固岸、促淤造陆等重要的生态功能,在气候变化和快速城市化背景下认识红树林受保护状况具有重要意义。以基于遥感解译的红树林分类数据为基础,通过空缺分析,分析了我国红树林的就地保护状况,结果显示我国分布的红树林总面积为264 km~2(尚不含我国港、澳、台的统计数据),其中61.4%在自然保护区内受到保护。从红树林分布的主要省份来看,在海南省分布的红树林面积较少但保护比例高,广西和广东省分布的红树林面积大但受保护面积比例相对较低。在3种红树林类型中,红树-木果楝林和红海榄-木榄林分布面积较小,但受保护的面积都在90%以上,秋茄-桐花树-白骨壤林分布的面积最大,但受保护的面积为52.6%。研究提出自然保护区外红树林分布的关键区域,并建议通过划定生态保护红线等方式来进行保护。     

The distribution of benthic invertebrates along a natural turbidity gradient in Lake Temiskaming,Ontario-Quebec

Lake Temiskaming, a rift valley lake on the Ontario-Quebec border, exhibits a permanent gradient of turbidity due to tributary streams which cut through clay deposits to the north of the lake. Concentrations of total phosphorus (TP) also decreased from north to south, with values suggesting mesotrophic conditions. Concentrations of chlorophyll a were characteristic of oligotrophic lakes and showed little relationship to either turbidity or TP. Large numbers of Tubificidae were found at our northernmost sampling station at a depth of 50 m, probably reflecting the localized impact of allochthonous organic matter introduced by a tributary stream. Numerical abundance of the benthic fauna was much lower and did not vary significantly among the six more southerly 50 m stations, but biomass declined from north to south as Heterotrissocladius oliveri relaced Pontoporeia hoyi. Numerical abundance did not differ significantly among stations at depths of 10 m, but biomass decreased from north to south reflecting the distributions of the largest species, Hexagenia sp. and P. hoyi. Intensive sampling on two transects showed that maximum numbers of invertebrates occurred in the profundal zone. While these results are consistent with the correlation between TP and zoobenthic biomass reported by other investigators, size selective predation by fish may also be important in controlling the distribution of benthic invertebrates in Lake Temiskaming.     

Growth and physiological responses of Kandelia candel and Bruguiera gymnorrhiza to livestock wastewater

Growth and physiological responses of two mangrove species (Kandelia candel and Bruguiera gymnorrhiza) to livestock wastewater under two salinity conditions (seawater with salinity of 30þ and freshwater) were examined in greenhouse pot-cultivation systems for 144 days. Wastewater treatment significantly enhanced growth of Kandelia candel and Bruguiera gymnorrhiza in terms of stem height, stem basal diameter, leaf production, maximum unit leaf area and relative growth rate. Wastewater discharges and salinity levels did not significantly change biomass partitioning of Kandelia candel, however, more biomass of Bruguiera gymnorrhiza was allocated to leaf due to wastewater discharges. In Bruguiera gymnorrhiza, contents of chlorophyll a and chlorophyll b increased with wastewater discharges but such increase was not observed in Kandelia candel. On the other hand, livestock wastewater increased leaf electric conductance in Kandelia candel but not in Bruguiera gymnorrhiza. The peroxidase activity in stem and root of Kandelia candel under both salinity conditions increased due to wastewater discharges, while the activity in root of the treated Bruguiera gymnorrhiza seedlings decreased under freshwater condition but increased at seawater salinity. The superoxide dismutase activity in treated Bruguiera gymnorrhiza decreased but did not show any significant change in Kandelia candel receiving livestock wastewater.     

Spatio‐temporal pattern of specific gravity of mangrove diaspore: implications for upstream dispersal

Mangrove plants, which develop highly productive forests on tropical–subtropical coastlines and river estuaries, rely mainly on river and sea water currents for their diaspore dispersal. Mangrove diaspores are basically dispersed in brackish to sea waters; thus whether they sink or float during the dispersal period could be changed dynamically, both spatially and temporally, depending on the salinity to which they are exposed, as well as on their specific gravity (SG). We found that the SG of diaspores of two mangrove species, Rhizophora stylosa and Bruguiera gymnorrhiza, which are dominant species and form a typical zonation (B. gymnorrhiza in inland areas and R. stylosa in more seaward areas) in the study area, changed spatially and temporally. Bruguiera gymnorrhiza gradually lost its SG in accordance with elapsed time, while R. stylosa lost its SG in the first 6 h and gained gradually according to elapsed time. Hydrodynamic simulation of dispersal patterns, in combination with observed specific gravity changes of both diaspores and river water, demonstrated that the spatio‐temporal pattern of specific gravity change was closely related to the difference in dispersal behaviors between the species. The diaspore of B. gymnorrhiza generally disperses upstream, and covers wider ranges than that of R. stylosa, at least in the early phase of dispersal. These dispersal behaviors agreed well with the distribution pattern of the species across estuaries in the study site. To date, hydrochory has been considered to be a mostly passive process governed by the dynamics of water current and subsequent sediment movement, with fixed diaspore characteristics such as shape and buoyancy. The present study shed new light on an active and dynamic process in hydrochory and found that the buoyancy of diaspores may be controlled by changes in their spatial and temporal SG.     

Restoration of Aegiceras corniculatum mangroves in Jiulongjiang Estuary changed macro-benthic faunal community

Macro-benthic faunal communities were compared between non-vegetation mudflat and Aegiceras corniculatum mangroves with different ages in Jiulongjiang Estuary, China. Faunal species number was highest in the mature mangrove and was higher in mangroves than in the mudflat, as snails and some crustaceans species were only collected in mangroves. The 5-year-old mangrove had the highest infaunal abundance and crustacean biomass. Snails had more abundance in the young mangroves. Uca arcuata was the dominant crab species in the non-vegetation mudflat and 5-year-old mangrove. Mangrove vegetation and sediment characteristics analyses indicated different habitats due to A. corniculatum mangrove restoration. However, overall poor correlations between faunal assemblage and sediment properties indicated that sediment properties were not the major factors influencing faunal distribution.