黄河源区植被生长季NDVI时空特征及其对气候变化的响应

利用黄河源区MODIS/NDVI数据、1∶100万植被类型图和气象资料,分析了该区不同植被类型生长季NDVI时空特征以及与气候因子的关系。结果表明,1）2000—2011年,黄河源区植被生长呈改善趋势,生长季NDVI年际变化率每10 a为＋2.75%,高寒草原、高寒草甸、高寒灌丛生长季NDVI年际变化率分别为每10 a＋2.84%、＋2.65%、＋2.77%。2）黄河源区植被改善面积占全区总面积的29.39%,主要分布在卡日曲和玛曲上游、扎曲流域、布青山南麓、扎陵湖北部和鄂陵湖周边地区。植被退化面积仅占全区总面积的0.98%,主要分布在约古宗列曲东南部山地和卡日曲北部山地。受水热条件控制,植被改善表现为：①植被改善面积南坡大于北坡;②植被改善面积随海拔升高先增加后减小;③植被改善面积随坡度增加迅速减小。3）黄河源区植被生长季NDVI与同期气温和降水分别存在显著正相关性,其中高寒草原和高寒草甸生长受降水影响更为明显,而高寒灌丛生长受气温影响更为明显。气候的暖湿化趋势可能是促使黄河源区植被生长改善的主要原因。     

基于SPOT—VGT的黄河流域植被覆盖时空演变

植被是土地覆盖中的最主要部分,是连接土壤、大气和生物等要素的自然“纽带”。植被覆盖动态变化对全球能量循环和物质循环具有重要影响,是全球变化研究的重要内容之一。黄河流域作为我国重要的粮食生产基地,其环境变化直接影响到流域经济的可持续发展。为了快速准确地提取地表植被状况,了解黄河流域生态环境,利用1998—2011年的SPOT-VGT遥感数据,结合地理信息技术,采用均值法和趋势分析法对黄河流域植被NDVI时空分布特征和变化趋势进行了动态监测。结果表明,（1）黄河流域14年NDVI均值的空间分布整体特征是东南部平原、盆地和西部山地植被状况要好于北部地区。其次,黄河流域属于干旱半干旱地区,植被发育主要依赖于水文条件,所以沿黄河干流和支流区域也具有较高的植被NDVI值。（2）黄河流域植被NDVl年均值近14年间整体呈缓慢增长趋势,1998--2000年呈现急剧减少态势,2001--2003年出现了较为快速的增长,2004--2011年又出现了较长时间的连续增长过程。（3）黄河流域植被NDVI基本不变的区域约占研究区总面积的71．13％;植被NDVI轻微改善的区域约占流域总面积的27．30％,且主要分布于流域东南部的盆地、平原和西部的山地、丘陵地区,植被NDVI退化的区域面积约占流域总面积的0．98％。黄河流域自1998年以来,植被NDVI整体在不断提高,生态环境在不断改善。     

新疆博格达山北坡表土花粉散布特征

基于博格达山北坡68个表土样品花粉组合特征,对比植物群落样方调查结果,借助聚类分析、主成分分析方法,探讨了表土花粉组合与现代植被分布的关系。研究表明,(1)博格达山北坡表土花粉可划归5个不同植被带,湿度是影响其分布的主要因素,藜科(Chenopodiaceae)、蒿属(Artemisia)、云杉属(Picea)花粉分布受气流影响显著,忽略它们对其他植被带花粉组合的干扰,表土花粉与现代植被分布对应良好。各植被带均有其特有的花粉组合方式,山地荒漠带藜科-蒿属组合占绝对优势,山地草原带演替为蒿属-藜科-禾本科(Poaceae)-蔷薇科(Rosaceae)组合,山地森林带以云杉属-桦木属(Betula)-蒿属-藜科-禾本科为主,高山草甸带以蒿属-云杉属-藜科-莎草科(Cyperaceae)组合为特征,高山垫状植被带表现为蒿属-藜科-蔷薇科-云杉属组合。(2)草本植物花粉含量(62.7%)优势明显,乔、灌木(37.3%)次之。蒿属(23.1%)、藜科(21.5%)、云杉属(18.1%)、莎草科(9.4%)、禾本科(8.6%)、桦木属(5.7%)、蔷薇科(5.3%)等科(属)含量高、变幅大,为最主要的花粉类型,可作为古气候研究的重要依据,藜科、蒿属产量大、易传播,表现出超代表性,云杉属代表性较好,莎草科则受自身结构及保存条件等多重因素影响呈低代表性。(3)蒿属/藜科(A/C)比值不仅能将山地荒漠带、山地草原带区分开,还能指示研究区域湿度变化,古环境重建时可作为区域有效湿度的代用指标。     

京津冀地区近20年NDVI时空变化特征

健康稳定的自然生态系统是保障城市发展的重要基础.了解京津冀城镇快速发展过程中自然生态系统的变化,有助于该区域城镇绿色协调可持续发展.植被指数NDVI时空变化特征可反映地区自然生态系统状况及其演变规律.基于MOD13Q1和Landsat遥感影像数据,利用一元线性回归趋势分析法分析近20年(2000—2019年)京津冀地区...     

1982-2006年华北植被覆盖变化及其与气候变化的关系

利用1982-2006年GIMMSNDVI数据反映华北地区植被覆盖变化状况,结合1982--2006年该地区85个气象站点的气温和降水数据,分别从年际变化、季节变化和月变化三个时间尺度分析华北地区植被覆盖变化及其与气候变化的关系。结果表明,从年际变化来看,华北植被变化与气温变化关系较与降水关系密切;从季节变化来看,华北地区植被生长在不同季节对水热条件变化的响应不同,春季和秋季植被生长与气温的关系较与降水的关系密切,而夏季植被生长主要受降水的影响;从月变化来看,4月和5月植被变化受气温变化影响较明显,一定程度上说明4月和5月植被生长的NDVI值增加可能是由于气候变暖引起的植被生长季提前产生的;6-9月植被生长与前2个月降水变化关系密切,说明植被生长对降水变化具有一定的滞后性。     

榆林地区植被时空分异特征及其影响因素研究

植被作为反映陆地生态系统和气候的重要指标,对研究全球或区域生态环境变化具有重要作用。以地处黄土高原生态脆弱区的榆林市为研究区,基于地理探测器模型,选取坡向、坡度、气温、降水和土壤类型5类自然因子,土地利用类型、人口密度和GDP 3类人文因子,分析榆林地区植被空间分异特征及其驱动力,并揭示了促进植被生长影响因子的最适宜特征。结果表明,（1）研究区2000—2018年植被覆盖趋向改善,NDVI呈现增加趋势,增速为0.11/10a,2008年以后植被增长较为明显;NDVI在2018年中高等级（0.6—0.8）面积比2000年中高等级面积明显增加;中高等级集中于榆林市东部黄土丘陵区,中低等级（0.2—0.4）集中于榆林市西北部的风沙区,植被覆盖呈现东部高西北低的空间分布特征。（2）人口密度和气温因子较好地解释植被NDVI空间分异性,是影响NDVI空间分异性的主要因子,GDP、土地利用类型和坡度是次级影响因子,其他因子对NDVI空间分异存在间接影响;坡向、降水和土壤类型因子与其他自然、人文因素对植被空间分布影响存在显著性差异。（3）自然、人文因子对榆林市NDVI的影响存在交互作用,因子之间的交互...     

三江源地区基于植被生长季的NDVI对气候因子响应的差异性研究

采用1999—2008年NDVI和三江源地区及周边31个气象站点的温度和降水旬数据,在确定每年5月中旬至9月底作为三江源地区植被生长季（14旬）的基础上,分析NDVI对气候因子变化响应的区域差异。结果显示：以巴颜喀拉山为界,三江源西部生长季内NDVI指数对温度、降水的线性相关性明显比中东部普遍,同时,不同海拔地段响应程度存在着明显的差异,且海拔5300m左右为线性响应的最高限,NDVI对气候因子的线性响应主要表现为对温度正相关和降水负相关。因此,对三江源生长季内植被生长而言,水条件明显好于温度条件,并且温度对植被覆盖的影响通过直接正相关和降水负相关表现出来,温度是三江源地区植被生长的主导因子。     

Knowing fire incidence through fuel phenology: A remotely sensed approach

Fire is a basic ecological factor that contributes to determine vegetation diversity and dynamics in time and space. Fuel characteristics play an essential role in fire ignition and propagation; at the landscape scale fuel availability and flammability are closely related to the vegetation phenology that directly affects wildfire pattern in time and space. In this view, the annual normalized difference vegetation index (NDVI) profiles derived from high temporal resolution satellites, like SPOT Vegetation, represent an effective tool for monitoring the coarse-scale vegetation seasonal timing. The objective of this study thus consists in quantifying the explanatory power of multitemporal NDVI profiles on the fire regime characteristics of the potential natural vegetation (PNV) types of Sardinia (Italy) over a 5-year period (2000-2004). The results obtained show a good association between the NDVI temporal dynamics of the PNV of Sardinia and the corresponding fire regime characteristics, emphasizing the role of the bioclimatic timing of the vegetation in controlling the coarse-scale wildfire spatio-temporal distribution of Sardinia. By providing a sound phytogeographical framework for describing different wildfire regimes, PNV maps can thus be considered helpful cartographic documents for fire management strategies at the landscape scale.     

Persistently highest risk areas for hantavirus pulmonary syndrome: potential sites for refugia.

Interannual variation in the number of cases of human disease caused by hantaviruses in North America has been hypothesized to reflect environmental changes that influence rodent reservoir populations. This hypothesis postulates that when cases are rare reservoir populations are geographically restricted in patches of suitable habitat. Identifying these sites, which is needed to test the hypothesis, has proven to be a challenge. Satellite imagery of the U.S. Southwest has shown associations among the likelihood of human hantaviral disease and increases in the rodent populations, as well as increased prevalence of Sin Nombre virus (SNV) in rodent populations. In this study we characterize local areas that had environmental signatures that persisted as predicted highest risk sites for human disease through much of the 1990s. These areas represent a small percentage (0.3%) of the region. Exploratory analyses indicate that these areas were not randomly distributed, but were associated with certain landscape characteristics. Characteristics of elevation, slope, aspect, and land cover were associated with persistent high risk. Using multivariate Poisson regression to control for confounding effects, sites with deciduous- or mixed-forest land cover on moderate to steep slopes (>5 degrees) above 2130 m elevation were associated with increasing numbers of years at highest risk. These are candidate locations for refugia. Sites associated with cleared ground or shrubland were less often associated with high risk compared to reference conditions. The seasonal patterns of vegetation growth in persistently high-risk areas were compared to matched locations using MODIS (moderate resolution imaging spectroradiometer) NDVI (normalized difference vegetation index) during a time of a severe drought in the region from 2002 to 2004. Despite the drought and regardless of land cover, the NDVI in persistently highest risk areas had an early onset, with significantly higher levels of green vegetation that lasted longer than at comparable sites. These observations identify locations that can be monitored for the abundance of P. maniculatus and presence of SNV. If these sites are refugia, we predict they will be occupied by infected deer mice when other monitored sites are unoccupied.     

Translating effects of inbreeding depression on component vital rates to overall population growth in endangered bighorn sheep

Evidence of inbreeding depression is commonly detected from the fitness traits of animals, yet its effects on population growth rates of endangered species are rarely assessed. We examined whether inbreeding depression was affecting Sierra Nevada bighorn sheep (Ovis canadensis sierrae), a subspecies listed as endangered under the U.S. Endangered Species Act. Our objectives were to characterize genetic variation in this subspecies; test whether inbreeding depression affects bighorn sheep vital rates (adult survival and female fecundity); evaluate whether inbreeding depression may limit subspecies recovery; and examine the potential for genetic management to increase population growth rates. Genetic variation in 4 populations of Sierra Nevada bighorn sheep was among the lowest reported for any wild bighorn sheep population, and our results suggest that inbreeding depression has reduced adult female fecundity. Despite this population sizes and growth rates predicted from matrix-based projection models demonstrated that inbreeding depression would not substantially inhibit the recovery of Sierra Nevada bighorn sheep populations in the next approximately 8 bighorn sheep generations (48 years). Furthermore, simulations of genetic rescue within the subspecies did not suggest that such activities would appreciably increase population sizes or growth rates during the period we modeled (10 bighorn sheep generations, 60 years). Only simulations that augmented the Mono Basin population with genetic variation from other subspecies, which is not currently a management option, predicted significant increases in population size. Although we recommend that recovery activities should minimize future losses of genetic variation, genetic effects within these endangered populations-either negative (inbreeding depression) or positive (within subspecies genetic rescue)-appear unlikely to dramatically compromise or stimulate short-term conservation efforts. The distinction between detecting the effects of inbreeding depression on a component vital rate (e.g., fecundity) and the effects of inbreeding depression on population growth underscores the importance of quantifying inbreeding costs relative to population dynamics to effectively manage endangered populations.     

天津植被指数对气候因子响应的敏感性分析

探索植被覆盖与气候变化相互关系是全球环境变化研究领域的重要内容之一,研究特定区域植被对气候变化响应特征对植被重建和生态恢复具有重要意义。然而,目前植被对气候因子响应的敏感性研究还十分缺乏。利用1982—2003年8km×8km的NASA/GIMMS半月合成的归一化植被指数（NDVI）和同期气候数据,研究了天津地区NDVI对气候因子响应特征及其敏感性。结果表明：植被NDVI与气温及降水均有显著非线性正相关关系（P〈0.0001）;在半月平均气温低于0℃时,植被NDVI与气温没有显著性相关,而从气温高于0℃,一直到高于22℃,NDVI与气温的关系均达到显著性水平（P〈0.05）,但相关性是逐渐降低的;当半月平均气温高于23℃及以上时,NDVI与气温没有显著的相关关系（P〉0.05）;当半月降水量〉0mm时,NDVI与降水存在极显著正相关关系（P〈0.0001）,随着降水量的增加,相关关系减弱,在降水量大于50mm时,NDVI与降水没有显著相关关系（P〉0.05）;研究结果证实,天津地区植被指数对气候的响应存有明显的非线性特征,在低温和低降水量条件下植被的响应更为敏感,23℃和50mm分别是该地区影响植被生长的气温和降水阈值。结合1982—2003年逐半月气候条件分析发现,气温的影响主要是春、秋两季,而降水的影响主要表现在春、秋及夏初。     

Captive Breeding and Reintroduction Evaluation Criteria: a Case Study of Peninsular Bighorn Sheep

Abstract: Captive breeding and reintroduction programs are rarely evaluated, and assessment criteria vary widely. We used the following criteria to evaluate a bighorn sheep ( Ovis canadensis ) augmentation program: (1) survival and recruitment rates in the captive population, (2) survival of released animals, (3) recruitment of released animals, (4) growth rate of the reintroduced or augmented population, and (5) establishment of a viable wild population. Captive bighorn survival and recruitment was high, averaging 0.98 (SD = 0.05) and 71.0% (SD = 19.4), respectively. Annual survival of free-ranging captive-reared bighorn ( n = 73, x = 0.80, SD = 0.11) did not differ (   Z = −0.85, p = 0.40; n = 14) from survival of wild-reared bighorn ( n = 43, x = 0.81, SD = 0.12). Recruitment was unusually low for both captive-reared (  x = 13.7%, SD = 0.24) and wild-reared ewes (  x = 13.7%, SD = 0.20). Although reintroduction did not result in population growth or establishment of a viable population, it helped prevent extirpation of the reinforced deme, preserved metapopulation linkage, and aided habitat preservation. Chronic low recruitment and low adult survivorship precluded achievement of criteria 3–5. Environmental conditions in the release area also appeared to hinder program success. Standard evaluation criteria for ongoing reintroductions allow for informative assessments and facilitate comparisons needed to refine reintroduction science as a recovery tool for threatened or endangered populations.     

Secondary sexual characters signal fighting ability and determine social rank in Alpine ibex (Capra ibex)

Social dominance is a fundamental aspect of male evolutionary ecology in polygynous mammals because it determines access to estrous females. As it is rarely possible to monitor marked individuals of known morphology, little is known about the determinants of male dominance. We studied the social structure of Alpine ibex males in Gran Paradiso National Park, Italy in 2003, 2006, and 2007. Dominance interactions produced a linear social hierarchy. In ibex males, body mass and horn length are key traits in male-male combat, and both increase with age. We explored the links between age, body mass, horn length, and social rank. Ibex males showed much age-independent phenotypic heterogeneity and rapidly growing males reached high rank at a younger age than slow-growing males. Because there is no trade-off between horn growth and longevity, fast-growing males may face weak potential costs of rapid growth and high fitness benefit of achieving high rank. Violent interactions were more likely to occur between males similar in both mass and horn length, independently of age. We suggest that only high-quality individuals can afford a strategy of rapid growth for both secondary sexual characters, and likely reap substantial fitness benefits.     

Intra-sexual variability in feeding behaviour of a mountain ungulate: size matters

Many studies comparing the behaviour of individuals of different genders or species showed that animal body mass and forage quality/quantity are key elements of the foraging ecology of herbivores. Since body mass could also influence the animal’s sensitivity to predation risk, its vigilance behaviour should consequently be affected. Alpine ibex (Capra ibex) is characterised by a strong dimorphism among males of different ages, thus representing an ideal case study for testing the Jarman-Bell principle, avoiding possibly misleading effects resulting from the comparison between different species or genders. We analysed the fine-scale foraging behaviour of male ibex in order to assess the effect of body mass and the effects of vegetation quality/quantity on both foraging and vigilance behaviour. Our results showed that smaller males were more selective than larger ones, on account of their lower capability of digesting plant. Smaller males scanned the environment more frequently than larger ones. Male ibex grazed more selectively in sites with high quality forage and their bite rate increased as forage biomass decreased. Vigilance frequency increased with increasing forage biomass as, under these circumstances, ibex are able to prolong anti-predator vigilance while chewing bites that have already been cropped. Our findings highlight the effects of body mass per se on both foraging and anti-predator behaviours in herbivores, thus supporting the Jarman-Bell principle. Foraging can arguably be considered a very flexible behaviour with high evolutionary relevance as it enables herbivores to optimally adjust their total energy intake under varying conditions of food resources.     

Bite rates in Rocky Mountain bighorn sheep (<Emphasis Type="Italic">Ovis canadensis</Emphasis>): effects of season,age, sex and reproductive status

We investigated the effects of vegetation biomass, crude protein content of consumed forage, age, sex and reproductive status on bite rates in Rocky Mountain bighorn sheep. We expected higher bite rates and vigilance in lactating females with young and higher bite rates in young growing individuals, than in non-reproducing females or rams. Lactating ewes had higher bite rates than yeld ewes and than subadult or adult rams. Subadult rams had higher bite rates than adult rams or yeld ewes. On recently burned grassland in spring, however, rams had a higher bite rate than adult females, while the contrary was true on control plots and on the burned plots in autumn. Bite rates declined for both ewes and rams from April to September and varied from year to year. While rams of different ages had significantly different bite rates, there was no effect of age on bite rates for ewes. There was no correlation between bite rates and available total biomass or biomass of live vegetation, or the numbers of steps taken while foraging for either ewes or rams. Adult rams had a lower vigilance rate than adult ewes, and vigilance decreased with increasing bite rates for all sheep. Bite rates in bighorn sheep vary greatly according to age, season and vegetation structure. An increase in bite rates during the forage growing season may compensate for higher energy demands during lactation and growth. There is a potential trade-off between foraging and vigilance as vigilance decreased with increasing bite rates.     

Growth of larval krill,<Emphasis Type="Italic"> Euphausia superba</Emphasis>, in fall and winter west of the Antarctic Peninsula

A quantitative characterization of growth-related parameters for larval and juvenile Antarctic krill, Euphausia superba (Dana), is essential to understanding the early life history of this key species in the Southern Ocean ecosystem. To this end, instantaneous growth rate experiments were conducted with larval and juvenile krill to determine growth increments and molting frequency in situ. All experiments were carried out during fall and winter months (April through September) on nine separate cruises west of the Antarctic Peninsula between 1987 and 1999. A consistent seasonal pattern across years was observed: growth rates decreased during fall (April/May), were minimal in early winter (June), and increased to maximum rates by late winter (September). Habitat-specific differences (water column vs under-ice) in growth rates of larvae collected on the same cruise were not observed in early winter (June 1993; within-year comparison). However, in a between-year comparison, larvae from the under-ice habitat (June/July 1987) had significantly higher growth increments than larvae from the open-water habitat (July 1989). The difference between these two comparisons may be a function of the degree of contrast in food availability in the water column and the sea ice at different times in the winter. Daylength at the time of collection explained 74% of the variation in larval and juvenile growth rates. This correlation may be an indirect effect of the influence of diel cycles on krill behavior and/or primary production in both the water column and ice.Communicated by J.P. Grassle, New Brunswick     

Spatio-temporal analysis of alpine ecotones: A spatial explicit model targeting altitudinal vegetation shifts

There is general agreement in literature that Alpine vegetation belt ecotones have shown a trend of upward migration in the last few decades. Despite the potential of such shifts as indicators of global change effects in mountain ecosystems, there are relatively few works focused on their assessment in a systematic and spatially explicit way. In this work our aim is to quantify the altitudinal shifts and analyse the spatial pattern dynamics of mountain ecotones. We developed a novel procedure to delineate the current and former state of three characteristic mountain ecotones, which we formalised as forest, tree and tundra lines. Our approach is based on the recognition of altitudinal extreme outposts identified with ecotone locations at a slope scale. The integration of multi-temporal datasets allows the identification and quantification of altitudinal advances and retreats in the outpost locations for a given period. We tested the method in a section of the Italian Alps for the period 1957-2003. Results show a general trend of an increase in altitude for the three ecotones, despite the occurrence of occasional decreases. We estimate decadal altitude increments of 25 m for forest line, 13 m for treeline and 11 m for tundra line. We also identified changes in ecotone spatial morphology between the two dates, with significant implications in connectivity and colonisation dynamics.     

Woody plant richness and NDVI response to drought events in Catalonian (northeastern Spain) forests

The role of species diversity on ecosystem resistance in the face of strong environmental fluctuations has been addressed from both theoretical and experimental viewpoints to reveal a variety of positive and negative relationships. Here we explore empirically the relationship between the richness of forest woody species and canopy resistance to extreme drought episodes. We compare richness data from an extensive forest inventory to a temporal series of satellite imagery that estimated drought impact on forest canopy as NDVI (normalized difference vegetation index) anomalies of the dry summer in 2003 in relation to records of previous years. We considered five different types of forests that are representative of the main climatic and altitudinal gradients of the region, ranging from lowland Mediterranean to mountain boreal-temperate climates. The observed relationship differed among forest types and interacted with the climate, summarised by the Thorntwaite index. In Mediterranean Pinus halepensis forests, NDVI decreased during the drought. This decrease was stronger in forests with lower richness. In Mediterranean evergreen forests of Quercus ilex, drought did not result in an overall NDVI loss, but lower NDVI values were observed in drier localities with lower richness, and in more moist localities with higher number of species. In mountain Pinus sylvestris forests NDVI decreased, mostly due to the drought impact on drier localities, while no relation to species richness was observed. In moist Fagus sylvatica forests, NDVI only decreased in plots with high richness. No effect of drought was observed in the high mountain Pinus uncinata forests. Our results show that a shift on the diversity-stability relationship appears across the regional, climatic gradient. A positive relationship appears in drier localities, supporting a null model where the probability of finding a species able to cope with drier conditions increases with the number of species. However, in more moist localities we hypothesize that the proportion of drought-sensitive species would increase in richer localities, due to a higher likelihood of co-occurrence of species that share moist climatic requirements. The study points to the convenience of considering the causes of disturbance in relation to current environmental gradients and historical environmental constraints on the community.     

广东省NDVI时空变化特征及其对气候因子的响应

广东省植被类型丰富、气候条件复杂,开展气候因子对植被的影响研究,对于全省保护生态环境、应对气候变化具有重要意义.利用MODIS NDVI数据和地面气象观测数据,基于变化趋势分析、空间自相关分析、相关性分析等方法,研究广东省NDVI时空分布特征及其对气温、降水、日照时数等气候因子的响应.结果表明,2000—2018年广东...     

Extra-pair paternity in the monogamous Alpine marmot revealed by nuclear DNA microsatellite analysis

The genetic parentage and pedigrees of 35 litters from 12 family groups of monogamous Alpine marmot (Marmota marmota) in the French Alps were analyzed using six hypervariable sequence repeat nuclear loci (microsatellites). All of the members of these family groups were sampled during a 5-year period. Hairs taken directly from animals served as a source of DNA for amplification of the loci. Our results indicate that the genetic mating system of the Alpine marmot is quite different from a strictly monogamous breeding system. Extra-pair paternity occurred in 11 of the 35 litters (31.4%). Of the 134 juveniles typed, 26 (19.4%) could be attributed to extra-pair copulation (EPC). We examine hypotheses which could explain the evolution of EPC and discuss the different patterns of extra-pair mating. Received: 11 July 1997 / Accepted after revision: 23 May 1998