椒江口海域春秋季虾类资源分布与环境的关系

利用2010年春季(4月)和秋季(10月)椒江口海域渔业资源调查资料,分析了椒江口虾类资源密度的分布,优势种和水团、水深等环境因子对虾类数量和分布的影响.结果表明:椒江口海域春季和秋季虾类平均重量密度分别为7.89和86.20 kg·km-2,尾数密度均值分别为11.04×103和34.41×103 ind·km-2;虾类密度平面分布特征,在春季,调查海域虾类密度西部高于东部,高密度区出现在椒江河口沿岸区域,尤其是受椒江径流影响明显的北部沿海海域;在秋季,东部明显高于西部,尤其是大陈岛东南部受台湾暖流和浙江上升流影响的海域.逐步回归分析表明:椒江口表层温度与虾类重量密度和尾数密度均呈显著的相关关系(Pw =0.004,PN=0.01),但水深与资源密度相关关系不显著;春季,中国毛虾(Acetes chinensis)对虾类尾数密度贡献率最大,对重量密度贡献率也较大,为春季最重要的优势种;秋季,中华管鞭虾(Solenocera crassicornis)对尾数和重量密度贡献率均最大,为秋季最重要的优势种.     

东海大陆架虾类资源量评估

根据2008年春(5月)、夏(8月)、秋(11月)、冬(2009年2月)四季东海区桁杆拖虾网调查资料, 应用资源密度法首次估算了调查海域主要经济虾类和总体的现存资源量和最大持续产量(MSY)。结果表明: 调查海域虾类季度平均资源密度为203.5 kg/km2。根据季节分布, 以夏季最高, 冬季最低; 根据区域分布, 南部海域最高, 中部最低。调查海域虾类的现存资源量约为10.6×104 t, MSY约为22.2×104 t, 与2008年调查海域虾类渔获量比较, 表明该海域虾类资源总体利用略显不足, 尚有一定潜力。在主要经济虾类的现存资源量与MSY中, 以假长缝拟对虾Parapenaeus fissuroides最高; 与历史资料相比, 某些经济价值较高的种类, 如哈氏仿对虾Parapenaeopsis hardwickii、日本对虾Penaeus japonicus等的资源量明显减少, 而长角赤虾Metapenaeopsis longirostris等经济虾类较低的种类资源量有所上升, 这说明东海大陆架海域虾类资源组成情况已经发生了很大改变。       

北京地区植被景观中斑块大小的分布特征

利用GIS软件ARC/INFO将北京地区的1∶20万植被图数字化，并提取各斑块的面积信息。该图包含72个基本的斑块类型，它们又分属于森林、灌丛、草地、果园、农田和水体6大类型。这些大类又分别包含20、28、4、7、11和2个基本类型。选用斑块个数、总面积、平均斑块面积、标准差、变异系数、中值、最大斑块面积、最小斑块面积、极差和偏态系数等几个描述统计量，以及Γ-分布、对数正态分布、Weibull分布、指数分布和正态分布等5个概率分布来刻画斑块大小的分布特征。结果表明除具有很少斑块的少数基本类型以外，其它基本类型以及所有6个大类的斑块大小的分布都不是对称的，而是右偏的。因此，普通的正态分布不能对它们加以刻画；其他4种概率分布也只能刻画部分类型，并且服从对数正态分布的类型最多，服从负指数分布的斑块类型最少。     

采用渔业声学方法和GIS模型对楠溪江鱼类资源量及空间分布的评估

采用快捷、直接、经济的渔业声学调查方法,结合GIS模型对温州楠溪江4个主要渔业作业区域的鱼类空间分布及资源量进行了评估。结果表明,4个区域的鱼类平均目标强度为(-55.03±5.44)dB,平均体长约为7cm,范围3—10cm,各区域的鱼类大小没有显著的统计学差异(P0.05)。同时4个区域的鱼类平均密度为8.87ind./1000m3,95%置信区间为1.66—16.09ind./1000m3。其中区域Ⅲ的鱼类密度最高,平均密度为(20.06±9.34)ind./1000m3。通过鱼类分布的GIS图形得知,鱼类的空间分布呈现斑块状的分布形式。按照ArcGIS对鱼类密度进行的栅格化数据,并结合各个栅格所代表的水体体积,进行鱼类资源量的估算,结果为4个探测区域的鱼类总尾数为2.5万尾,各个区域的鱼类资源量分别为:3971ind.、11478ind.、6587ind.、2978ind.。     

浙江南部近海小黄鱼资源分布及其与环境因子的关系

根据2015—2016年浙江南部近海4个航次的底拖网资源调查数据,利用广义可加模型分析了调查期内小黄鱼资源的分布特征及其与环境因子的关系.结果表明:浙江南部近海的小黄鱼资源主要集中在鱼山渔场,夏季为小黄鱼资源的高产期,站点平均资源密度达到500.74 kg·h^-1·km^-2.不同季节影响小黄鱼资源密度及其分布的环境因子各不相同.其中,环境因子对秋季小黄鱼资源密度的影响效果并不显著.春季,小黄鱼主要分布于水深较浅的高盐水域;夏季,水温和盐度均与小黄鱼资源密度呈负相关关系,小黄鱼主要分布于中温高盐的鱼山海域;冬季,水温与资源密度呈正相关,小黄鱼栖息于水温适宜的外侧站点水域.总体上,小黄鱼资源的分布特征符合其洄游习性,但个别环境因子与资源密度的关系难以解释,仍需进一步研究.研究结果有助于了解浙江南部近海小黄鱼群体的生活习性,以及对小黄鱼资源的养护和管理.     

北京地区植被景观中斑块大小的分布特征

利用GIS软件ARC／INFO将北京地区的1：20万植被图数字化,并提取各斑块的面积信息。该图包含72个基本的斑块类型,它们又分属于森林、灌丛、草地、果园、农田和水体6大类型。这些大类又分别包含20、28、4、7、11和2个基本类型。选用斑块个数、总面积、平均斑块面积、标准差、变异系数、中值、最大斑块面积、最小斑块面积、极差和偏态系数等几个描述统计量,以及厂一分布、对数正态分布、Weibull分布、指数分布和正态分布等5个概率分布来刻画斑块大小的分布特征。结果表明：除具有很少斑块的少数基本类型以外,其它基本类型以及所有6个大类的斑块大小的分布都不是对称的,而是右偏的。因此,普通的正态分布不能对它们加以刻画;其他4种概率分布也只能刻画部分类型,并且服从对数正态分布的类型最多,服从负指数分布的斑块类型最少。     

苏北浅滩沙脊潮沟地形和潮流对虾类分布的影响

采用2010年11和201 1年5月苏北浅滩海域2个航次的渔业资源调查资料,研究苏北浅滩虾类资源密度、种类组成和优势种,结合该海域沙脊潮沟系统,探讨了地形特征和潮流对虾类数量分布的影响.结果表明:两次拖网调查共鉴定虾类15种;依据相对重要性指数IRI值,得出该海域春季优势种为细螯虾(Leptochela gracilis),秋季为中国毛虾(Acetes chinensis);春季虾类生物量和尾数密度均高于秋季,春季和秋季虾类生物量分别为35.64和9.24 kg·km-2,尾数密度分别为35.42× 103和25.92×103ind·km-2;根据贡献率β值,虾类总密度变化与主要优势种虾类密度变化密切相关,两者平面分布规律也一致;因苏北浅滩海域潮沟和沙脊相间的地形地貌特征,加之受东海前进波和黄海旋转波作用,形成潮流从北、东、南3个方向向心辐聚作用;潮沟的流速大于沙脊的流速,致使无论在春季还是秋季,形成虾类生物量和尾数密度在潮沟内远大于沙脊上、在沙脊与潮间带浅滩之间的深槽处略高于沙脊处的分布特征.     

舟山渔场金塘岛海域春夏季的鱼类群落

为了揭示金塘岛附近海域渔场鱼类资源特征,利用2009年5月和9月在舟山渔场金塘岛水域获得的渔业资源调查资料,研究金塘岛水域鱼类密度的时空分布,分析种类组成、优势种、种类数和多样性等群落结构等指标的变化特征,分析这些特征所反映这一水域的渔场属性。结果表明:5月和9月鱼类重量和尾数密度分别为47.51kg.km-2、3.44×103ind.km-2和92.50kg.km-2、20.51×103ind.km-2;鱼类重量密度与尾数密度分布趋势基本一致,呈现出近舟山群岛一侧的东部水域鱼类密度高于西部水域的趋势;5月和9月龙头鱼(Harpodon nehereus)是本海域最重要的优势种,几乎为幼体组成,而凤鲚(Coilia mystus)是次要优势种。总体上,两季平均鱼类幼体比例高达86%,显示出这一水域是鱼类的产卵场和育幼场;鱼类种类数夏季高于春季;种类数与多样性的分布格局相似,也是舟山群岛一侧的东部水域鱼类密度高于西部水域。可以认为,近金塘岛一侧海域是经济幼鱼的索饵场。     

中西太平洋大眼金枪鱼中心渔场时空分布与温跃层的关系

采用Argo数据和中西太平洋渔业委员会的大眼金枪鱼延绳钓数据,绘制了温跃层和月平均单位捕捞努力量渔获量(CPUE)的空间叠加图,分析中西太平洋大眼金枪鱼渔场时空分布对温跃层的响应关系.结果表明: 全年中心渔场纬向主要分布在南北纬10°之间的低纬度区域.在赤道以南有季节性中心渔场的出现和消失,与温跃层上界温度、深度和温跃层厚度的积极性变化有关.中心渔场主要分布在温跃层上界较深(70～100 m)和厚度较大(>60 m)的海域,厚度小于40 m的区域难以形成中心渔场;适宜分布的上界温度区间在26～29 ℃,在此区间外CPUE多小于中心渔场阈值(Q3).中心渔场的空间分布随上界深度和跃层厚度的季节性变动而变动,当赤道以南海域的上界深度变浅以及厚度变薄时,中心渔场消失.温跃层下界温度、深度和温跃层强度季节性变化不显著,但与中心渔场出现有显著关系.中心渔场主要分布在温跃层下界深度两条高值带之间的区域,温度低于13 ℃以及强度大的区域;在温跃层下界深度超过300 m和小于150 m区域,下界温度超过17 ℃的区域,或者强度小的区域难以形成中心渔场.利用频次分析和经验累积分布函数计算其适宜温跃层特征参数分布,结果表明研究区域大眼金枪鱼适宜分布的上界温度、深度和下界温度、深度分别是26～29 ℃、70～110 m、11～13 ℃和200～280 m;适宜分布的温跃层厚度和强度分别是50～90 m和0.1～0.16 ℃·m^-1.本研究初步得出研究区域大眼金枪鱼CPUE空间分布和温跃层的关系,为我国远洋金枪鱼捕捞作业和资源管理提供理论参考.     

三门湾海域冬夏季口足目和十足目虾类的种类组成、时空分布及多样性分析

根据2007年12月和2008年6月三门湾海域的2个航次的渔业资源调查资料,研究三门湾海域口足目与十足目中虾类在冬季和夏季的密度、优势种及多样性的时空分布,并结合该调查海区的地形地貌和水文等因素对虾类的分布进行分析.结果表明,三门湾海域虾类以广温、广盐种为主,其夏季种类数高于冬季,且夏季虾类质量和尾数密度均值亦高于冬季...     

Characterization and Simulation of Uncertain Frequency Distributions: Effects of Distribution Choice,Variability, Uncertainty,and Parameter Dependence

We use bootstrap simulation to characterize uncertainty in parametric distributions, including Normal, Lognormal, Gamma, Weibull, and Beta, commonly used to represent variability in probabilistic assessments. Bootstrap simulation enables one to estimate sampling distributions for sample statistics, such as distribution parameters, even when analytical solutions are not available. Using a two-dimensional framework for both uncertainty and variability, uncertainties in cumulative distribution functions were simulated. The mathematical properties of uncertain frequency distributions were evaluated in a series of case studies during which the parameters of each type of distribution were varied for sample sizes of 5, 10, and 20. For positively skewed distributions such as Lognormal, Weibull, and Gamma, the range of uncertainty is widest at the upper tail of the distribution. For symmetric unbounded distributions, such as Normal, the uncertainties are widest at both tails of the distribution. For bounded distributions, such as Beta, the uncertainties are typically widest in the central portions of the distribution. Bootstrap simulation enables complex dependencies between sampling distributions to be captured. The effects of uncertainty, variability, and parameter dependencies were studied for several generic functional forms of models, including models in which two-dimensional random variables are added, multiplied, and divided, to show the sensitivity of model results to different assumptions regarding model input distributions, ranges of variability, and ranges of uncertainty and to show the types of errors that may be obtained from mis-specification of parameter dependence. A total of 1,098 case studies were simulated. In some cases, counter-intuitive results were obtained. For example, the point value of the 95th percentile of uncertainty for the 95th percentile of variability of the product of four Gamma or Weibull distributions decreases as the coefficient of variation of each model input increases and, therefore, may not provide a conservative estimate. Failure to properly characterize parameter uncertainties and their dependencies can lead to orders-of-magnitude mis-estimates of both variability and uncertainty. In many cases, the numerical stability of two-dimensional simulation results was found to decrease as the coefficient of variation of the inputs increases. We discuss the strengths and limitations of bootstrap simulation as a method for quantifying uncertainty due to random sampling error.     

武夷山景区主要景观类型斑块大小分布规律及其等级尺度效应分析

以武夷山风景名胜区为研究对象,运用景观分类生态原则和群落生态学原则,将武夷山风景名胜区划分为10类景观类型,并提取各类型斑块的面积、周长、数量等信息。在此基础上,运用正态分布、对数正态分布、Weibull分布、г-分布、Bata-分布及负指数分布等6个常见的概率分布,分别在3种等级划分条件下,对各景观类型斑块大小分布规律进行研究．结果表明,多数斑块大小服从对数正态分布。少数服从r．分布和Weibull分布,而所有类型的斑块大小均不服从正态分布;对于同一景观类型而言,不同划分等级对其斑块大小的分布规律产生一定的影响,即斑块大小分布存在着等级效应,其中以b等级划分相对比较理想;对不同的景观类型而言,斑块大小分布规律依类型的不同而存在差异,而且就整个景区所有斑块大小分布来说,尚未有一种概率分布能理想地加以刻画;由具有相同分布规律的不同景观类型的斑块构成的整体,其斑块大小分布规律基本保持不变,而由不同分布规律的不同景观类型的斑块构成的整体,其斑块大小分布规律发生变化。     

复合种群管理的风险评估——以日本鲐为例

单一种群是目前渔业资源评估的基本假设,但渔业资源常由多个地方种群或产卵种群组成,并且种群间存在交流,构成复合种群。根据复合种群概念,以东、黄海日本鲐为例,对其12种种群动态情况进行了模拟。利用模拟所得的数据及剩余产量模型,分别分析了在复合种群、两独立种群及单一种群假设下所设置的10种评估管理方案,结果表明:(1)基于复合种群假设的评估管理方案与模拟的种群动态一致,在单位捕捞努力量渔获量(CPUE)观测误差较小情况下,该方案为最佳方案,可获得最大可持续产量,但随CPUE观测误差增大,该方案种群灭绝率增大,管理效果随之退化。(2)基于两独立种群假设的评估管理方案均使资源过度开发,不利于资源可持续利用。(3)在单一种群假设下,选择不同CPUE作为资源指数和采用不同捕捞量分配方法的评估管理方案存在过度捕捞和开发不足两种状况,其管理效果受种群本身参数及空间交换率等因素的影响而不同;若采用的CPUE反映部分种群动态信息,则其评估管理方案至少在一种模拟情况下出现种群100%灭绝;若CPUE能反映整个种群资源量的动态变化,且捕捞量能按种群的空间结构进行分配,则管理效果与(1)类似,但不能获得最大可持续产量,若忽略种群的空间结构影响而均匀分配捕捞量,则至少在一种模拟情况下出现种群100%灭绝。据此,对于复合种群的管理,建议:(A)如果种群数据收集及数据精度能得到保证,该资源的评估与管理应基于复合种群假设;(B)如果目前收集种群数据存在较大困难,且CPUE数据存在较大误差,则可采用单一种群假设,但必须设定更保守的捕捞量和采用基于种群空间结构的总许可渔获量(TAC)管理方案;(C)在制定渔业管理政策时,应结合种群生态、数据、模型假设及参数估计方法等方面的不确定性对管理控制规则进行系统的管理策略评价以避免风险。     

应用年龄结构产量模型评估印度洋黄鳍金枪鱼资源

利用年龄结构产量模型(Age structured production model,ASPM)评估了印度洋黄鳍金枪鱼资源状况,同时结合亲体量-补充量曲线陡度系数和年龄组自然死亡系数的敏感性分析,描述了黄鳍金枪鱼资源的发展趋势、判断了开发状况。研究认为,陡度系数设在0.6-0.8才可能使亲体量产生出最大可持续产量(Maximum sustainable yield,MSY)的水平。采用美洲热带金枪鱼委员会推荐的自然死亡系数值时,评估结果最接近渔业现状。研究发现,随着捕捞努力量的增加,总资源量和亲体量呈逐年下降趋势,但总资源量自1990年后趋向稳定,维持在195.9-263.2万t,平均为221万t;亲体量在1994年后下降到100万t以下,1997年以后处在维持MSY所需亲体量的水平之下,目前仍呈下降趋势。补充量在渔业初期呈现大幅度波动,1978年后趋于稳定,并维持在3258.36-6583.35×106尾,平均为4687.66×106尾。未成熟鱼的数量总体较为稳定,但成熟鱼的数量出现剧减,从渔业初期的246.51×106尾减少到2005年的19.02×106尾。模型估计的总捕捞死亡系数从渔业初期开始逐渐上升,1991年后出现大幅度上升,处于0.334-0.456间,2003年时超过FMSY,捕捞产量也于2003年超过MSY。分析认为,2003年以来印度洋黄鳍金枪鱼的持续高产量被认为是不可持续,根据ASPM估算,2003-2006年均产量46.4万t,超过了MSY(36.4万t);S/SMSY为0.76;Fall/FMSY为1.39,由此判断现阶段印度洋黄鳍金枪鱼正处于过度捕捞状态。     

A general production model with dependence between data from multiple surveys

A fishery‐independent survey for stock assessments is made sometimes more than once per year to detect a difference in relative sizes of fish populations (e.g., catch‐per‐unit‐effort [CPUE]) in response to a seasonal change in fish spatial distributions. Many managers tended to treat such data independently instead of systematically synthesizing them. A primary objective of this study was to synthesize all survey data via a simple hierarchical structure. I used the general (Pella‐Tomlinson) surplus production model for the illustration, because the purpose of this study was not a stock assessment, and the model was simpler than an age‐structured model. The surplus production model has about an eight decade history (since Graham's paper in 1935) and has been prominent in fish population dynamics. The logistic (Graham‐Schaefer) version was useful in the sense of simplifying the dynamics of a fish population in relation to its intrinsic growth, natural mortality, recruitment, density‐dependence, and fishery catch, but it was criticized because of its unrealistic limitations. Subsequently, the general version was suggested to accommodate flexibility and be realistic. In this study, I inferred parameters in the general surplus production model, simultaneously synthesizing all available data even from different temporal ranges. I used Georges Bank yellowtail flounder (Limanda ferruginea) data for demonstration.     

Some effects of demersal trawling in Lake Malawi (Lake Nyasa) from 1968 to 1974

Exploratory trawl data were used to define demersal fish stocks of Lake Malawi in terms of abundance by depth and area. The mean weight of the catch generally declined with increasing depth. A comparison of initial standing stock estimates and the yield from a commercial fishery over a 7 year period suggests a potential yield of less than half of the initial stock. Over 160 species of fish have been identified in the trawl catch of which over 80% by weight were cichlids. Some of the commonest species of cichlids are listed. The composition of the catch in the heavily exploited southern end of the lake has changed from one dominated by large species of Haplochromis sp. and Lethrinops sp. to one dominated by smaller species.     

Fishing diseased abalone to promote yield and conservation

Past theoretical models suggest fishing disease-impacted stocks can reduce parasite transmission, but this is a good management strategy only when the exploitation required to reduce transmission does not overfish the stock. We applied this concept to a red abalone fishery so impacted by an infectious disease (withering syndrome) that stock densities plummeted and managers closed the fishery. In addition to the non-selective fishing strategy considered by past disease-fishing models, we modelled targeting (culling) infected individuals, which is plausible in red abalone because modern diagnostic tools can determine infection without harming landed abalone and the diagnostic cost is minor relative to the catch value. The non-selective abalone fishing required to eradicate parasites exceeded thresholds for abalone sustainability, but targeting infected abalone allowed the fishery to generate yield and reduce parasite prevalence while maintaining stock densities at or above the densities attainable if the population was closed to fishing. The effect was strong enough that stock and yield increased even when the catch was one-third uninfected abalone. These results could apply to other fisheries as the diagnostic costs decline relative to catch value.     

基于地统计分析西印度洋黄鳍金枪鱼围网渔获量的空间异质性

西印度洋公海海域是世界上围网黄鳍金枪鱼渔业的主要作业海域之一,根据印度洋金枪鱼委员会1999—2004年的1°×1°的各月黄鳍金枪鱼围网渔获量统计数据,采用地统计方法探索该海域黄鳍金枪鱼渔获量的空间异质性特征及其相关生态动力过程。进行了如下分析:(1)利用GIS制图观察渔获量的时空分布特征,发现其空间格局的变异受到的季节变化和年际变化共同影响,且前者明显强于后者。(2)采用地统计方法计算各月渔获量的空间异质性参数,并按照年际和季节情况分别进行了统计分析,发现渔获量的地统计参数值和变异函数模型有明显的季节和年际差异;渔获量的空间相关距离(变程)平均在1000nm左右,冬季要小于夏季;渔获量的空间变异函数模型主要为相关距离较大且空间依赖性较弱的指数模型;渔获量的空间结构方差比例(平均为65.82%)远大于随机性方差比例(平均为34.18%);渔获量在1°×1°尺度下具有明显的空间自相关性。(3)对地统计参数值和渔获量的相关关系研究,并探讨季节变化下渔获量的空间异质性特征与相关生态动力过程关系,发现各月渔获量随着空间总变异(基台值)增加而增加,两者存在强相关性;各月渔获量和南北和西北-东南向分维数值有一定相关性,意味着海洋动力过程在南北和西北—东南向过程越强,渔获量越低。西印度洋黄鳍金枪鱼围网渔获量的空间变异原因在于季风气候和ENSO循环过程引起的海洋流场、营养盐和温跃层等变化外在因素,以及围网捕捞方式和鱼类的行为方式的内在因素共同导致的。     

An unlikely partnership: fishers’ participation in a small-scale fishery data collection program in the Timor Sea

Traditional fisheries stock assessment methods and fishery independent surveys are costly and time consuming exercises. However fishers trained in data collection and utilising other skills can reduce costs and improve fishery assessments and management. A data collection program was conducted by Australian and Indonesian scientists with small-scale Indonesian sea cucumber fishers to evaluate the approach and then capture its benefits. The data fishers recorded allowed for the first stock assessment of this trans-boundary fishery during its centuries-long existence at Scott Reef in north-western Australia. The program also included interviews with fishers capturing the social, economic, and demographic aspects of the fishery. Economic inputs to fishing were complemented by fishery revenue data voluntarily submitted when fishers returned to port and sold their catch. Catch data recorded by fishers demonstrated much higher abundances than estimates obtained using standard visual transect methods and accurately reflected the true catch composition. However, they also showed extreme rates of exploitation. Interviews revealed social and economic factors that would be important considerations if management interventions were made. The program’s approach and the time scientists spent on the fishers’ vessels were key ingredients to fishers’ participation and the utility of the results. Despite the program’s achievements the information generated has not led to improved management or had any direct benefits for the participants. Sustaining the program in the longer term requires that its value is better captured.     

DENSITY DEPENDENT GROWTH IN NORTH PACIFIC SPERM WHALES

Data from the North Pacific sperm whale (Physeter catodon Linnaeus, 1758) fishery were examined for a possible density dependent change in growth during 40 yr harvesting after World War II. Early in this period males from the eastern stock were 16.8 m or less in length. By the early 1970s the largest males in the catch exceeded 16.8 m in length and reached 18.9 m in the late 1970s. The proportion of males measuring over 16.8 m, among sexually mature males (≥14.0 m), increased from 0 to >20% during the 1970s. Increases in the maximum size of males were possibly preceded by a change in the frequency distribution of body lengths in the middle 1960s when only 10% of the postwar catch had been taken. Testis weights suggested an increase in body length at sexual maturity. Two of the three putative North Pacific stocks showed similar growth changes. Adult males taken in the Bering Sea did not show such changes during the exploitation which ended in 1972 because of overfishing. Females showed no detectable change in body size. It is concluded that: (1) density dependent effects on male growth are greater before sexual maturity than after it, (2) males may show density dependent changes even at a population level above 90% of the carrying capacity, (3) polygynous males acquire more mates and realize higher reproductive success because of increased body size, and (4) females appear to maximize production by maturing earlier and shortening calving intervals in response to density change.