摘要
耳石形状具有高度的物种特异性和显著的群系特异性,在耳石形态的鱼种识别及鱼类系统学研究方面有重要的研究价值,国际上许多学者近年来已就此开展大量基础研究。本文以近年来笔者在国家海洋局908专项、国家重点基础研究发展计划(973计划)研究过程中搜集的近万枚海洋鱼类耳石为标本,利用专业生物图像分析软件,采用数学傅里叶分析等手段,就鱼类耳石形态学研究涉及到的各个主要方向：鱼类耳石形态特征与量度、我国海洋鱼类耳石形态多样性、鱼类耳石形态学在鱼种及群系识别中的应用等进行了较系统的研究。主要研究结果如下：
(1)根据矢耳石在内耳中的位置形态,为便于描述,将耳石靠在鱼体头部的方向称为前部,靠鱼体尾部的方向称为后部,靠鱼体背部的方向称为背部,靠鱼体腹部的方向称为腹部,朝向鱼体外侧的面称为外侧面,朝向鱼体内侧的面称为内侧面。
对中国近海16目83科255种习见鱼类耳石形态进行了测量与记述。以鲈形目种数最多,此外包括鲱形目、仙女鱼目、鳗鲡目、鲇形目、颌针鱼目、鼬鳚目、鳕形目、金眼鲷目、鲻形目、鮋形目、鲽形目、鮟鱇目和海鲢目等部分鱼类。通过对各种鱼类的耳石形态量度特征：耳石长、耳石宽、周长、面积、矩形趋近率、充实度、密度指标等进行了测量分析,对耳石的外部轮廓以傅里叶谐值加以描述,初步建立数据库,然后进行统计分析,对以鱼类耳石形态来识别鱼种进行了初步研究。
对255个鱼种的聚类分析和判别分析结果均表明,不同科之间耳石形态重叠严重,判别成功率仅有44.71%,不适合作为分类识别的单位,而对属和种的判别成功率分别为81.96%和86.44%,有一定的应用研究前景,据此推测,经过大量细致的研究,耳石形态对鱼种的识别有可能成为分类学的有效的辅助手段。
(2)利用2007、2008年采自黄海五个海区斑尾复虾虎鱼群体标本,探讨了耳石形态的群体间差异,及环境对耳石形态的影响。使用耳石长、耳石宽、周长、面积、充实度、矩形趋近率及20个傅里叶分析得到的谐值描述耳石形态,为了去除体长对耳石形态的影响,文中采用标准化后的变量进行分析。结果显示,雌雄、左右及年际间耳石形态无显著差异,但是一年中春季和秋季的耳石形态间存在显著差异；耳石形态的地域间差异大于年间差异。群体间判别分析结果表明黄海五个区域间斑尾复虾虎鱼的耳石形态无显著差异,判别成功率在29.7%-77.4%之间,黄海北部的威海群体的判别成功率最高,且该群体与黄海中南部4个群体间差异较大,海区之间环境差异可能是形成这个结果的内在原因。
(3)利用2009年7月采自南海北部湾的161尾斑鱚(Sillago maculata)和164尾多鳞鱚(Sillago sihama)标本,以形态学常规测量和傅里叶分析来描述耳石的外部形态特征,并采用判别分析法区分两个鱼种。统计分析时以传统的耳石形态一维、二维量度指标(耳石长、耳石宽、周长、面积、矩形趋近率及充实度)和傅里叶谐值相结合,经协方差分析表明部分变量(耳石长、耳石宽、周长、面积、密度、谐值3、4、5、13、15、16)跟体长显著相关。为了消除体长对耳石形态的影响,在统计时只选取了体长范围在90-140mm之间的标本,并对跟体长显著相关的变量进行转换,以残差来代替原始变量,纳入统计分析中。结果表明：两种鱼判别分析的准确率分别为97.8%和100%,如此高的判别准确率,说明耳石形态学分析手段是识别这两种鱼种的一种有效方法。
(4)分析了中国近海的8种天竺鲷属鱼类的矢耳石形态特征。结果显示,8种鱼类矢耳石形态近似,前部较尖,后部略圆钝,背部呈折线状,腹部圆弧形,背部中央有缺刻,听沟宽阔。8种鱼耳石长宽比1.35～1.63,矩形趋近率0.67～0.77,充实度14.77～24.99。根据耳石听沟形态及外部轮廓特征,编制了上述8种天竺鲷属鱼类的检索表。结果表明,听沟是鱼类耳石形态分类的重要特征之一,耳石形态在鱼类系统分类上有良好的应用研究前景。
(5)分析了中国近海的12种石斑鱼属鱼类的矢耳石形态特征。结果显示,12种鱼类矢耳石形态近似,耳石细长,呈三角形或梭形,由听沟开口处形成的前部凹刻“V”形或浅宽,形成显著或不显著的侧叶。大多种类在耳石外部轮廓、听沟形态方面区别甚微,充分体现出属内的近缘关系特点。耳石外部轮廓、边缘光滑度、听沟前部形态、凹刻类型等一些细节特征成为本属鉴定的主要参数依据,基于傅里叶系数的聚类分析也证明了这一点。结果表明,听沟是鱼类耳石形态分类的重要特征之一,耳石形态在鱼类系统分类上有良好的应用研究前景。
In addition, otoliths are widely used as indicators of fish age and growth at annually, daily and seasonal levels. Recent developments in the tools used for determining otolith's microconstituents have opened up possibilities of studying migrations and stock identity as well as using isotopes to reconstruct past life history of fishes, some good progress has made.
The otolith form is species and group specific with the important value for fish taxonomy and systematics research, and some researchers have carried out the massive basic research in the field in recent years. Based on ten thousand otoliths collected from the recent research programms such as "908" special project of State Oceanic Administration of People's Republic of China, National Basic Research Program of China (973 program) (No.2005CB422306), etc., by bio-image analysis software and the mathematical means of Fourier analysis, some fields of fish otolith morphology:Otolith morphological characteristics and measurement methods, otolith morphological diversity, species and stocks identification, morphological changes of otolith growth were carried out. The main findings were as follows:
(1)Based on otolith position and posture in the inner ear, in order to facilitate description of the otolith, we call the direction of the head as anterior, the direction of the tail as posterior, the direction of the back as dorsal, the direction of the belly as ventral. In the meanwhile, we give the distinction of the lateral and mesial surface of otolith. Rostrum and antirostrum are created by a nick in the anterior otolith region. Sulcus acusticus exists at mesial surface of otolith.
Otoliths collected from 255 Species, belonging to 83 families and 13orders of fishes from China Sea were descripted. Most species were from Perciformes, others were from Clupeiformes, Anguilliformes, Myctophiformes, Siluriformes, Gadiformes, Beloniformes, Mugiliformes, Scorpaeniformes, Elopiformes, Beloniformes, and Pleuronectiformes, etc. Otolith shape, size, smoothness degree, sulcus acusticus, rostrum and antirostrum were descirpted in detail; and some measurement characteristic including otolith length, otolith width, perimeter, area, circularity, rectangularity and compactness were analyzed, the outline of the otolith was descriped by Fourier analysis. Used these shape variables into statistical analysis to identity fish.
Cluster analysis and classify analysis showed that, the relatives relationship among fishes based on otolith morphology was not entirely overlap with the traditional classification system of fishes. The classification accuracy of family was 44.71%, and we suspected that family was not suitable for identification; the classification accuracy of genus and species were 81.96% and 86.44%, which showed application perspectives.
(2) Otolith shape is species specific and is an ideal marker of fish population affiliation. Otolith shape of spottedtail goby Synechogobius ommaturus was used to identify stocks in different spawning locations in the Yellow Sea. o explore the potential existence of local stocks of spottedtail goby in the Yellow Sea by analysis of otolith shape, and to investigate the ambient impacts on otolith shape. Spottedtail goby was sampled in five locations in the Yellow Sea in 2007 and 2008. Otolith was described using variables correlated to size (otolith area, perimeter, length, width, and weight) and shape (rectangularity, circularity, and 20 Fourier harmonics). Only standardized otolith variables were used so that the effect of otolith size on the shape variables could be eliminated. There was no significant difference among variables of sex, year and side (left and right). However, the otolith shapes of the spring stocks and the autumn stocks significantly differed. Otolith shape differences were greater among locations than between years. Correct classification rate of spottedtail goby with the otolith shape at different sampling locations ranged 29.7%-77.4%.
(3) 161 samples of Trumpeter sillago, Sillago maculata and 164 of Silver sillago, Sillago maculata were collected from Beibu gulf in july 2009. Using morphological variables and Fourier harmonics to describe otolith characteristic and use discriminant analyses to separate two species. Otoliths were measured by traditional one and two dimensional measures (otoliht length, width, diameters, area, perimeter, circularity and rectangularity), as well as by Fourier analysis to capture the finer regions of otolith. Analysis of covariance (ANCOVA) showed that there was significant correlation between morphological variables (diameter, perimeter, otolith length, otolith width, otolith area, density, harmonics3, harmonics4, harmonics5, harmonics13, harmonics15, and harmonicsl6) and body length. To minimize size effects on the shape variables between species only fish of a medium to all length (90-140 mm) were included in the data analysis and the variables which has significant relation with body length were transformed using residual. The result shows that:the discriminant analysis accuracy of Trumpeter sillago and Silver sillago were 97.8% and 100%, the high accuracy of discriminant analyses show that otolith shape was described accurately by morphological variables and Fourier harmonics, discriminant analyses is an effective way to identify and separate species.
(4)Morphological characters of the sagittal otoliths were analyzed for the eight species of genus Apogon inhabiting China Sea.The results showed that the sagittae of the eight species was similar in shape, all had narrow anterior part, wide posterior part, zigzag dorsal side and arcuate ventral side with a notch in the middle of the dorsal side. The sulcus was wide. Otoliths had aspect ratio of 1.35～1.63, rectangularity of 0.67 to 0.77 and circularity of 14.77～24.99 for eight fishes. According to the sulcus acusticus shape and otolith form, a systemic taxonomy was designed for eight fishes. The results indicted that the sulcus acusticus shape is one of the important characters of otoliths morphological classification, otoliths morphology have a good prospect of application in fish systematic.
(5) Morphological characters of the sagittal otoliths were analyzed for the twelve species of genus Epinephelus inhabiting China Sea.The results showed that the sagittae of the twelve species was similar in shape, all had narrow anterior part, wide posterior part, zigzag dorsal side and arcuate ventral side with a notch in the middle of the dorsal side. The sulcus was wide. Otoliths had aspect ratio of 1.35～1.63, rectangularity of 0.67 to 0.77 and circularity of 14.77～24.99 for eight fishes. According to the sulcus acusticus shape and otolith form, a systemic taxonomy was designed for eight fishes. The results indicted that the sulcus acusticus shape is one of the important characters of otoliths morphological classification, otoliths morphology have a good prospect of application in fish systematic.
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