摘要
唐鱼(Tanichthys albonubes)是一种小型溪流鱼类,属于国家Ⅱ级重点保护水生野生动物,也是广东最有代表性的鱼类之一。1998年版的中国动物红皮书将唐鱼的濒危等级列为野外绝迹。2003年唐鱼自然种群在广州从化重新发现。为弄清唐鱼自然种群的分布区域,了解唐鱼的栖息环境和不同种群的遗传多样性特征,掌握其生理生态特点,更好地开展唐鱼资源的保护和恢复工作,本文开展了一系列的野外调查和实验研究。首先以从化地区为重点丌展唐鱼自然种群和生境调查,分析不同种群的遗传多样性特征,结合唐鱼分布区域历史上的重大地质事件,探讨其自然分布格局的生物地理学过程。在此基础上研究了唐鱼的繁殖行为和胚胎发育,完成了从化市唐鱼自然保护区的总体规划和技术设计。主要结果如下:
1唐鱼的自然种群分布和生境
调查发现目前唐鱼自然种群的分布区域主要有从化市、增城市、清远市、高明市以及陆河市,分别属于流溪河水域、东江、北江、西江和莲花山水系。其中增城市、高明市、陆河市唐鱼自然分布区域以及从化市的7个分布点为历史没记载而本调查首次发现的。从野生唐鱼的体色来看,各个分布区域的唐鱼表现为两种表型。增城、从化、清远以及高明的唐鱼在个体颜色无差异,陆河的唐鱼体色偏狄暗,躯体和尾鳍分布大小不等的黑色斑块,背鳍边缘为红色。
野生唐鱼的生境可以分为沼泽地、泥沙底质溪流和沙石底质溪流三种类型。唐鱼生境主要有以下几个特点:1)溪流或草本沼泽,水流缓慢。唐鱼偶而会在溪流下游的大水面边缘出现;2)水生植物分布广泛,以莎草科和禾本科植物为主。在沙石底质的栖息地水绵分布广泛;3)伴生鱼类少。在沼泽栖息地无其它鱼类分布;在泥沙和沙石底质溪流栖息地基本只有条纹二须鱼巴、拟细鲫等少数鱼类伴生。
位于从化市的唐鱼自然保护区是典型的唐鱼自然分布区域。保护区内存在三种类型的唐鱼生境,水生植物群落和浮游动植物丰富,在此建立唐鱼自然保护区,对唐鱼自然资源的保育具有重要意义。
2唐鱼自然种群和养殖群体的形态学差异分析
作者在从化5个不同分布点分别采集野生唐鱼各30尾,和40尾养殖唐鱼进行传统形态的和多变量形态度量学差异分析,结果表明5个自然种群之间无显著形态差异,而人工养殖群体和从化自然种群之间形态差异显著。
3唐鱼遗传多样性的RAPD分析
作者在5个唐鱼自然种群分布地分别采集6尾野生唐鱼,从养殖场采集6尾养殖唐鱼进行RAPD分析,对100条引物进行了筛选,确定了8条引物进行实验,共获得90个位点。在各组的多态性位点比例和平均杂合度方面,从化种群最高,分别是22.22%和0.091。从化、增城和清远种群遗传相似性最高,三者相似性都在0.9251-0.9551。陆河与其它种群的相似性最低,只有0.6309-0.7459,说明陆河唐鱼种群与其它种群的亲缘关系最远。AMOVA分析显示,在遗传多样性的变异来源中,有74%来源于群体间,26%来源于群体内。通过UPGMA方法构建系统树,从化唐鱼种群和增城唐鱼种群聚类为一个谱系枝,两者关系最为密切;清远唐鱼种群则独立为另一个谱系枝,但和前两者共享一个谱系,三者的关系密切;其次是高明种群和前三者相近;养殖唐鱼群体独立为另一个谱系枝,陆河唐鱼种群也独立成为另一个谱系枝,且与其它种群的关系最为疏远。
4唐鱼遗传多样性的fAFLP分析及自然分布格局的生物地理学过程
作者在5个唐鱼自然种群分布地分别采集6尾野生唐鱼,从养殖场采集6尾养殖唐鱼进行fAFLP分析,对32对引物组合进行了筛选,确定了4对引物组合进行实验,共检出334条清晰扩增片段。6组唐鱼群体的多态位点比例和平均杂合度分别为:从化种群48.50%和0.168,增城种群52.40%和0.190,清远种群42.51%和0.142,养殖群体25.45%和0.100,陆河种群30.84%和0.111,高明种群30.54%和0.113。结果显示唐鱼养殖群体的遗传多样性水平最低,增城、从化自然种群最高。和其它用同样方法研究的野生淡水鱼类相比,例如白杨鱼Gobio gobio、须条鳅Barbatula barbatula、岩原鲤Procypris rabaudi等相比,唐鱼自然种群遗传多样性水平较高。
唐鱼各群体遗传分化系数Hst为0.395,明显高于其它淡水鱼类。AMOVA分析也显示有53%的变异来源于群体间。基因流Nm(0.383)显示各种群之间没有发生基因交流,因此可以认为目前唐鱼自然资源的种群遗传结构属于岛屿模型。
AMOVA显示的各不同群体的遗传多样性变异来源为群体间53%,群体内47%,群体间变异大于群体内变异。而后,作者利用5个自然种群之间的遗传距离与5个地区的实际相隔地理距离,经过Mantle分析得出两者显著相关。结合UPGMA方法构建的系统树以及华南地区地质的演变过程,作者认为唐鱼自然分布格局的形成与在第四纪(晚更新世)以来的海退和海侵有关。第四纪冰期的海退形成广阔陆地,退海后的水网联系为唐鱼的扩散创造了机会。而玉木冰期后的再次海侵,则造成了目前的分布格局。
5唐鱼的繁殖行为、胚胎发育研究以及保护对策
观察了唐鱼的繁殖行为方式和胚胎发育。唐鱼存在几种繁殖行为,即占区行为、张鳍行为、对峙行为、撞击行为、攻击行为、求偶行为以及交配行为。将唐鱼的胚胎发育分为29个时期,得出了唐鱼的最适繁殖温度为24-27℃,据此推测野生唐鱼一年可多次产卵。研究结果表明,唐鱼资源的减少和其栖息地的破坏有很大关系,保护唐鱼栖息地是野生唐鱼资源保护的最有效途径。本研究选择从化市良口镇新村水尾洞横坑作为唐鱼自然保护区的建设地址,规划和设计上充分考虑到唐鱼的栖息条件、繁殖习性、遗传多样性、种群基因交流、种群恢复等需要,符合濒危动物保护原则和唐鱼的生理生态特点。
As one of the most representative fish in Guangdong Province,White Cloud Mountain minnow(Tanichthys albonubes)has been listed in the second class state protected animals in China,and its status is extinct in nature in the China red data book of endangered animals.In 2003,the natural population of T.albonubes were rediscovered in CongHua city,In order to understanding its natural distribution,habitat,the genetic diversity and ecological characteristics, and performing the conservation of its natural resource,a series of investigations in wild and experiment research were performed.Firstly,we investigated the natural population and habitat of T.albonubes,emphasizing on the CongHua city.Secondly,we synthesized its characteristics of genetic diversity,gave the hypothesis that the distribution pattern has taken shape throughout the important geological affairs.Thirdly,we researched its reproductive behaviors and embryonic development,then finished the functional division and technique measures of the CongHua Natural Reserve of T.albonubes.The main results are as follows.
1 Distibution and habitat of its natural population
Investigations indicated that the distributions of its natural population were CongHua city, ZengCheng city,QingYuan city,GaoMing city and LuHe city,Belonging to sub-basin of Liuxi river,Dongjiang,Beijiang,Xijiang and the sub-basin of Lianhua mountain repectively.In this paper,the distributions of ZengCheng city,GaoMing city,LuHe city and seven location of CongHua city were firstly reported.On respect of the body color of T.albonubes.The wild fish displayed two color types,the fish color of LuHe city was different from other distributions,with the red basal margin of basal fin and black spots on its body.
There were three type of habitat of T.albonubes in natural distributions,including marsh, mud-sand-bottom stream and sand-stone-bottom stream.The characteristics of habitat are as follows:1.It mainly lived in stream and marsh.2.There are much aquatic plant in its habitat.3. Other fish are seldom in its habitat.No other fish appear in marsh,only several fish appear in stream,such as Puntius semifasciolatus.
With rich aquatic plants,phytoplankton and zooplankton,CongHua Natural Reserve of T. albonubes was the typical natural distibution including three types of habitat,Building up reserve in this place is scientific and very important to the conservation of T.albonubes.
2 Morphological variations between the natural population and hatchery stock of Tanichtys albonubes
The natural population of T.albonubes in Conghua city were investigated between 2006 and 2007.In this paper,traditional morphological data and truss network data were combined to conduct principal component analysis(PCA) and stepwise discriminant analysis to study the morphological variations between the natural populations and hatchery stock of T.albonubes. The results are showed as follows:1.In traditional morphological data,the morphological variations were significantly different(P＜0.01).2.In truss network data,through PCA with selected 12 morphological parameters,the identification accuracy were 95-100%(PC1) between the five natural population and one hatchery stock,synthetic identification accuracy were 100%.
3 Assessing genetic diversity of T.albonubes using RAPD markers.
Randomly Amplified Polymorphic DNA(RAPD) was used to detect the genetic diversity of five natural populations and one hatchery stock.six individuals from each stock were used.Of the 100 RAPD primers screened,8 primers generated clear and reproducible bands.A total of 90 reproducible loci ranging from 200 to 2000bp were am-plified from all the 36 individuals. Comparing the percentage of polymorphic loci and the average heterozygosity between six stocks,the CongHua population was the highest with 22.22%and 0.091 respectively.The Shannon's diversity indices between CongHua population,ZengCheng population and QingYuan population were about 0.9251-0.9551 with high similarity,.The Shannon's diversity indices between LuHe population and other five stocks is about 0.6309-0.7459,so the LuHe population was low relative of other stocks.Analysis of molecular variance(AMOVA) revealed that the percentage of variation within stock(26%) was lower than among stocks(74%).The six stocks were clustered into four major clades using UPGMA analysis.
4 Assessing genetic diversity of T.albonubes using fAFLP markers.
Genetic diversity of five natural populations and a hatchery stock of T.albonubes was investigated using fluorescence-based AFLP(amplified fragment length polymorphism).6 individuals from each stock were used.A total of 334 reproducible bands amplified with four fAFLP primer combinations were obtained from 36 fish.The percentage of polymorphic loci ranged widely from 25%to 52.4%.The percentage of polymorphic loci and the average heterozygosity of six stocks were as follows:CongHua population(48.5%,0.168),ZengCheng population(52.4%,0.190),QingYuan population(42.51%,0.142),Hatchery stock(25.45%,0.100),LuHe population(30.84%,0.111),GaoMing (30.54%,0.113)respectively.The genetic diversity of hatchery stock was the lowest in six stocks. Comparing other fish with AFLP,such as Gobio gobio,Barbatula barbatula,Procypris rabaudi et al.,the genetic diversity of T.albonubes was higher.
The average Fst value of overall stocks was 0.395,which indicated a high degree of genetic differentiation among the stocks.Also,the analysis of molecular variance(AMOVA) revealed that the percentage of variation within stock(47%) was lower than among stocks(53%).The estimated Nm value was 0.383,indicated that six stocks were geographic,and no gene flow happened.
The six stocks were clustered into four major clades with UPGMA.Through MANTLE analysis, result indicated five natural populations were relative to geologic distance,combining the geological changes,we gave the hypothesis that the natural distribution pattern of T.albonubes has been taken shape throughout the important geological affairs.
5 The reproductive behaviors,embryonic development and conservation of Tanichthys albonubes
Reproductive behaviors of T.albonubes were observed and summaried in this paper,including range behavior,stretching fins,confronting,collision,attacking,courtship and mating.Also,We studied its embryonic development through microscope,divided and illustrated it into 29 developmental stages.Under the water temperature of 27℃(±0.5℃),it took 34 hours and 8 minutes to complete embryonic development.Time for incubation showed a negative correlation to the water temperature.The optimum incubation temperature for T.albonubes embryos was between 24℃and 27℃.
In addition,our research showed that the decrease of habitat was negative to the natural resource of T.albonubes,protecting the habitat of T.albonubes was the most effective method.In the process of the establishment of CongHua Natural Reserve of T.albonubes,we considered the habitat,reproductive,genetic diversity,gene flow and restoration,so our work was conforming to the princilple of endangered conservation and the physiology,ecology of T.albonubes.
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