Impacts of silviculture on genetic diversity in the native forest species Eucalyptus sieberi

Potential impacts of regeneration practices ongenetic diversity in the Australian nativeforest species Eucalyptus sieberi L.A.S.Johnson. (silvertop ash) were assessed usingDNA markers. Three different silviculturaltreatments were examined: clear-felling withaerial re-sowing, and the seed tree system withsite preparation by either burning ormechanical disturbance. In addition, twounharvested stands were chosen as controls. Atotal sample of 825 trees were genotyped at 35Mendelian markers: 26 single-copy nuclear RFLPsand 9 microsatellites. No significantdifferences were found among the treatments inany of four population genetic statistics:allelic richness, effective number of alleles,expected heterozygosity and the panmictic index(f). Rare alleles were prevalent, and a MonteCarlo simulation showed that the apparent lossof four rare alleles from the saplingregenerants was highly statisticallysignificant. There was no evidence for recentbottlenecks from analyses of either the levelsof expected heterozygosity relative to thatexpected under mutation drift equilibrium, orthe allele frequency profiles. A dendrogram ofthe relationships between the sampledpopulations suggested that the seed tree systemmay result in the promotion of genetic drift(slight expansion of the dendrogram) whileaerial re-sowing of clear falls with the sameseedlot will lead to genetic homogenisation(contraction of the dendrogram). The apparentgenetic robustness of E. sieberi tonative forest regeneration practices isattributed to its local abundance combined withthe favourable properties of its reproductivebiology, as well as to the limitation that onlya single rotation was examined.     

Microsatellite loci for two East African tree species, Leptonychia usambarensis (Sterculiaceae) and Sorindeia madagascariensis (Anacardiaceae)

We isolated 20 trinucleotide microsatellites from two African tree species: Sorindeia madagascariensis (nine microsatellites) and Leptonychia usambarensis (11 microsatellites). Number of alleles ranged from three to seven in Sorindeia and two to 10 in Leptonychia. Observed heterozygosity ranged from 0.025 to 0.829 for Sorindeia and from 0.226 to 0.933 for Leptonychia. Two loci from each species departed from Hardy-Weinberg equilibrium. These microsatellite markers will be used to study how forest fragmentation affects pollination and seed dispersal processes of these tree species.     

基于SSR标记的不同优势等级云南松遗传多样性分析

基于SSR分子标记技术,按树高和胸径（地径）因子将样地内的植株分为优势木、中等木和劣势木3类,对不同生长优势等级云南松林木的遗传多样性变异特征进行了研究。结果表明：在林冠层,优势木的有效等位基因数、Shannon's信息指数、观测杂合度和期望杂合度分别为2.083、0.762、0.290和0.423,略高于中等木和劣势木;在更新层,有效等位基因数、Shannon's信息指数、观测杂合度和期望杂合度分别为2.063、0.774、0.272和0.410,除观测杂合度外,其余3个指标也均表现为优势木略高于中等木和劣势木。从差异显著性检验来看,不同生长优势等级云南松林木间的遗传多样性差异不显著,即林木生长分化对遗传多样性的影响不明显。     

Effects of reforestation methods on genetic diversity of lodgepole pine: an assessment using microsatellite and randomly amplified polymorphic DNA markers

 We examined the effects of different methods of forest regeneration on the genetic diversity of lodgepole pine (Pinus contorta var ‘latifolia’) using two different DNA-based molecular markers [randomly amplified polymorphic DNA (RAPDs) and microsatellites or simple sequence repeats (SSRs)]. Genetic diversity was estimated for 30 individuals in each of four populations for the following three stand types: (1) mature lodgepole pine (>100 years); (2) 20- to 30-year-old harvested stands left for natural regeneration; (3) 20- to 30-year-old planted stands (4 stands of each type); and one group of 30 operationally produced seedlings. There was no significant effect of stand type on expected heterozygosity, although allelic richness and diversity were much higher for SSRs than for RAPDs. Expected heterozygosity ranged from 0.39 to 0.47 based on RAPDs and from 0.67 to 0.77 based on SSRs. The number of alleles per locus for SSRs ranged from 3 to 34 (mean 21.0), and there was a significant relationship between sequence repeat length and the number of alleles at a locus. Both marker types showed that over 94% of the variation was contained within the populations and that the naturally regenerated stands sampled had lower (not significant) expected heterozygosity than the planted or unharvested stands. The group of seedlings (assessed by RAPDs only) had expected heterozygosity and allele frequencies similar to those of the unharvested stands. Genetic distance measures were higher than obtained previously in the species using isozyme markers. There was no correlation between the two marker types for pair-wise genetic distances based on populations analyzed by both methods. Pair-wise genetic distance measures and an ordination of allele frequencies for both marker types showed little effect of geographic location or stand type on genetic similarity. Received: 18 August 1998 / Accepted: 17 September 1998     

Eight microsatellite markers for the neotropical tree Luehea seemannii (Tiliaceae)

We isolated eight polymorphic microsatellites from the neotropical tree Luehea seemannii for gene flow and genetic structure studies. We used a streptavidin subtractive enrichment technique to develop a library of CA/GT repeats. Eight loci were screened for diversity from 96 individuals from Barro Colorado Island (BCI) and neighbouring Gigante peninsula in Panama. Luehea seemannii shows moderate levels of genetic diversity within these two populations. Allelic richness ranged from four to nine alleles and averaged 6.44 alleles per locus. Average expected heterozygosity was 0.65 on BCI and 0.60 on Gigante. Results are compared to microsatellite data from another wind‐dispersed gap colonizing species common in Panama.     

黑龙江省张广才岭北部小飞鼠的遗传多样性与种群历史动态

小飞鼠 (Pteromys volans) 为树栖夜行滑行类啮齿动物,在森林种子传播和维持生态系统平衡等方面发挥着重要的生态学作用。本研究利用mtDNA Cytb、控制区和nDNA微卫星3种分子标记,对黑龙江省张广才岭北部的小飞鼠种群进行遗传多样性与历史动态分析。检测出Cytb全序列 (1 140 bp) 的平均单倍型多样性为0.909,平均核苷酸多样性为0.616%;控制区全序列 (1 066 bp) 的平均单倍型多样性为0.945,平均核苷酸多样性为1.698%;微卫星检测出种群平均等位基因数13.167个,观测杂合度0.727,期望杂合度0.864,近交系数0.159。结果表明,小飞鼠种群遗传多样性丰富,但存在一定程度的杂合度不足和近亲繁殖;未检测到种群近期遗传瓶颈效应,种群内无遗传分化。高比例的稀有单倍型 (≥ 60%) 、低频率等位基因与近亲繁殖,提示未来种群面临遗传多样性下降的风险,建议加大对该物种的关注和保护力度。基于Cyt b基因的系统进化关系结果表明,小飞鼠存在3个明显的遗传谱系：远东、欧亚大陆北部和日本北海道,本研究中张广才岭和大兴安岭的样本单倍型归属为远东谱系。     

Effects of Logging on the Genetic Diversity of Quercus tiaoloshanica Chun et Ko in a Tropical Montane Forest of Hainan Island, Southern China

Quercus tiaoloshanica Chun et Ko, which has a narrow range of distribution, is one of the important endemic species of the tropical montane rain forest on Hainan Island, southern China. Long-term logging and habitat destruction have resulted in population decline and distribution retreat of Q. tiaoloshanica. To determine the impact of logging on the genetic diversity of Q. tiaoloshanica, the authors investigated the genetic structures using amplified fragment length polymorphism markers in four regenerated stands after logging and in one unlogged stand. Compared with the unlogged stand, the effective number of alleles per locus dropped by 1.0% in selective logging stands and by 2.0% in clear logging stands, corresponding to reductions of 3.8% and 5.2%, respectively, in mean Nei＇s gene diversity and 2.9% and 3.5%, respectively, in mean Shannon diversity index. No substantial genetic erosion was detected in any of the regenerated stands owing to the high tree density and high heterogeneity of the Q. tiaoloshanica stands investigated. Meanwhile, there was no natural regeneration of the species observed in a Dacrydium pierrei Hickel plantation 700 m away from the regenerated stands, suggesting the limited ability of seed dispersal of Q. tiaoloshanica. Clear logging should be undertaken cautiously because the total number of plant species dropped by 15.2% in the clear-logged stands compared with the unlogged stand. To conserve the genetic diversity of this species, as well as the plant biodiversity of tropical forests, the habitats of Q. tiaoloshanica should be protected against exploitation in terms of agricultural or other forms of land use, and some mature trees should be preserved as seed sources to maintain an adequate regeneration base for this species in the management of logging.     

森林砍伐对苦槠种群遗传结构的影响

人类活动严重干扰着自然生态系统,其中砍伐是对森林生态系统最常见的干扰之一,它导致森林退化,植物种群变小,甚至灭绝,遗传多样性也随之下降。当被破坏的森林未被转换性利用时,则会逐渐恢复,但由于瓶颈效应,恢复起来的生态系统中植物种群的遗传结构可能会改变。恢复种群遗传组成的改变一方面与干扰的强度、频度和持续时间有关,另一方面,也受植物生活史特点的深刻影响。然而,我国对于砍伐后恢复起来的森林生态系统中生物多样性的改变,尤其是遗传多样性的改变的研究并不多见。研究在浙江省宁波市天童国家森林公园及周边地区选择了5个苦槠种群,采用SSR微卫星标记来分析砍伐对苦槠种群遗传结构的影响。5对多态SSR引物共得到了29个等位基因。种群内维持了较高的遗传多样性,种群间遗传分化程度较低,基因流达8．68。恢复林和成熟林种群的遗传多样性相差不大,以阿育王寺地区恢复种群的最高;表明砍伐对于苦槠种群遗传多样性的影响不大,这与苦槠较强的萌条能力有关。尽管如此,在恢复种群中观察到近期的种群瓶颈,显示出砍伐对种群遗传组成的影响;而在一个成熟林中也观察到种群瓶颈,这是因片断化导致种群变小之故。植被保存最好的天童国家森林公园内苦槠种群的遗传多样性却较低,这可能与成熟林中苦槠优势度较低有关。     

Polymorphic microsatellite loci for population studies of the razor clam, <Emphasis Type="Italic">Sinonovacula constricta</Emphasis>

The razor clam (Sinonovacula constricta) is an important aquacultured bivalve in China. The natural populations of this species are decreasing quickly. To facilitate studies on genetic diversity and population structure of wild populations, microsatellites were isolated from a CA enriched genomic library. Eight microsatellite loci were polymorphic in 30 individuals from Chongming in Shanghai, China. The number of alleles per polymorphic locus varied from 6 to 13 and the values of observed heterozygosity and expected heterozygosity ranged from 0.350 to 1.000 and from 0.602 to 0.902, respectively. These microsatellites are being used in studying population differentiation and genetic diversity for effective conservation and management genetic resources of S. constricta.     

Pollen flow in fragmented landscapes maintains genetic diversity following stand-replacing disturbance in a neotropical pioneer tree,Vochysia ferruginea Mart

In forests with gap disturbance regimes, pioneer tree regeneration is typically abundant following stand-replacing disturbances, whether natural or anthropogenic. Differences in pioneer tree density linked to disturbance regime can influence pollinator behaviour and impact on mating patterns and genetic diversity of pioneer populations. Such mating pattern shifts can manifest as higher selfing rates and lower pollen diversity in old growth forest populations. In secondary forest, where more closely related pollen donors occur, an increase in biparental inbreeding is a potential problem. Here, we investigate the consequences of secondary forest colonisation on the mating patterns and genetic diversity of open-pollinated progeny arrays for the long-lived, self-compatible pioneer tree, Vochysia ferruginea, at two Costa Rican sites. Five microsatellite loci were screened across adult and seed cohorts from old growth forest with lower density, secondary forest with higher density, and isolated individual trees in pasture. Progeny from both old growth and secondary forest contexts were predominantly outcrossed (t_m=1.00) and experienced low levels of biparental inbreeding (t_m−t_s=0.00–0.04). In contrast to predictions, our results indicated that the mating patterns of V. ferruginea are relatively robust to density differences between old growth and secondary forest stands. In addition, we observed that pollen-mediated gene flow possibly maintained the genetic diversity of open-pollinated progeny arrays in stands of secondary forest adults. As part of a natural resource management strategy, we suggest that primary forest remnants should be prioritised for conservation to promote restoration of genetic diversity during forest regeneration.     

栖息条件对中华蜜蜂遗传多样性的影响

为了评估不同栖息条件对中华蜜蜂(简称中蜂)种群遗传多样性的影响,本研究运用微卫星标记的方法,对皖南山区4个不同采样点120个中华蜜蜂群体的2400只工蜂样本进行了分析.结果表明:与天然林野生中蜂群相比,单一茶树种植结构下,皖南中蜂群的等位基因数、平均期望杂合度和多态信息含量的差异均不显著;人工驯养中蜂群的等位基因数、多态信息含量差异显著,平均期望杂合度差异不显著;2006年采集的野生中蜂群与天然林野生中蜂群的等位基因数、平均期望杂合度和多态信息含量均差异显著.各种群间存在极显著的遗传分化,平均分化系数为0.32.说明皖南地区茶树单一化种植不会影响中蜂的遗传结构与资源保护.     

Genetic diversity impacts of forest fires,forest harvesting,and alternative reforestation practices in black spruce (Picea mariana)

Benchmarks were established for genetic diversity inherent in natural mature populations, and genetic diversity impacts of forest fires, clearcut harvesting and alternative natural and artificial silvicultural regeneration practices were determined in black spruce (Picea mariana). Allozymes of 32 loci were used to determine and compare genetic diversity and genetic relationships of adjacent or nearby four stand types: post-fire natural mature (FNM), post-fire natural young (FNR), post-harvest natural young (HNR) and post-harvest plantation (PLT), of black spruce at each of the four study sites located in two ecoregions in Manitoba: Ecoregion 90-Lac Seul Upland (Eastern) and Ecoregion 158 - Mid-Boreal Lowland (Northern). Both allelic- and genotypic-based genetic diversity parameters, as well as latent genetic potential, were determined. Black spruce populations showed typical moderate to high levels of allozyme genetic diversity. The mean genetic diversity parameters over the 16 black spruce populations sampled were as follows: percent loci polymorphic - 67%, mean number of alleles per locus - 2.52, effective number of alleles per locus - 1.70, observed heterozygosity - 0.222, expected heterozygosity - 0.308, mean number of observed genotypes per locus - 3.65, mean number of expected genotypes per locus - 5.03, genotype additivity (observed) - 116.8, genotype additivity (expected) - 161, genotype multiplicity (observed) - 6.16 x 10(15), genotype multiplicity (expected) - 2.06 x 10(19) and latent genetic potential - 26.12. The four stand types (FNM, FNR, HNR and PLT) had comparable and statistically similar genetic diversity levels at each of the four study sites as well as overall. No significant differences in black spruce genetic diversity levels were observed between the two ecoregions in Manitoba, as well as between the post-fire and post-harvest regenerated stands. No particular order of genetic relatedness among the four stand types was observed. Black spruce populations showed some sort of site-related differentiation in their genetic constitution. Allelic heterogeneity and genetic distances among populations within stand types and among four stand types suggest that the genetic diversity was maintained at the landscape level in black spruce. The results of our study demonstrate that forest fires and currently used clearcut harvesting, and alternative natural and artificial silvicultural regeneration practices, do not adversely affect genetic diversity in black spruce, and that the genetic diversity effects of clearcut harvesting are not significantly different from those due to forest fires in black spruce.     

Genetic diversity of Vietnamese domestic chicken populations as decision‐making support for conservation strategies

The aims of this study were to assess the genetic diversity of 17 populations of Vietnamese local chickens (VNN) and one Red Jungle Fowl population, together with six chicken populations of Chinese origin (CNO), and to provide priorities supporting the conservation of genetic resources using 20 microsatellites. Consequently, the VNN populations exhibited a higher diversity than did CNO populations in terms of number of alleles but showed a slightly lower observed heterozygosity. The VNN populations showed in total seven private alleles, whereas no CNO private alleles were found. The expected heterozygosity of 0.576 in the VNN populations was higher than the observed heterozygosity of 0.490, leading to heterozygote deficiency within populations. This issue could be partly explained by the Wahlund effect due to fragmentation of several populations between chicken flocks. Molecular analysis of variance showed that most of genetic variation was found within VNN populations. The Bayesian clustering analysis showed that VNN and CNO chickens were separated into two distinct groups with little evidence for gene flow between them. Among the 24 populations, 13 were successfully assigned to their own cluster, whereas the structuring was not clear for the remaining 11 chicken populations. The contributions of 24 populations to the total genetic diversity were mostly consistent across two approaches, taking into account the within‐ and between‐populations genetic diversity and allelic richness. The black H'mong, Lien Minh, Luong Phuong and Red Jungle Fowl were ranked with the highest priorities for conservation according to Caballero and Toro's and Petit's approaches. In conclusion, a national strategy needs to be set up for Vietnamese chicken populations, with three main components: conservation of high‐priority breeds, within‐breed management with animal exchanges between flocks to avoid Wahlund effect and monitoring of inbreeding rate.     

Impact of selection and breeding on the genetic diversity in Douglas-fir

Genetic changes following domestication of Douglas-fir were studied using isozyme data derived from two generations of seed orchards and their 49 wild progenitor populations. In addition, the breeding, production, and infusion populations used in the seed orchards were compared to their wild counterparts. Several parameters of gene diversity were measured (number of alleles per locus N _a, per cent of polymorphic loci PLP, and expected heterozygosity H, and population divergence D). These measures were similar or higher in the domesticated populations compared to their natural progenitors, indicating that early selection and breeding of a highly polymorphic species does not significantly reduce genetic variation. The two generations of seed orchards also did not differ, indicating that genetic variation may remain stable over future generations of forest plantations. Interestingly, compared to the natural populations, heterozygosity was higher in the seed orchards, probably due to pooling of widely distributed natural populations; however, rare localized or private alleles seemed to be less frequent in the domesticated populations. Differentiation values were not significant between the first generation orchards and the natural populations, but significant differences were observed between the second generation orchards and the wild progenitor populations, probably due to the interbreeding that forms the advanced generation seed orchards.     

Isolation and characterization of microsatellite loci in Santalum austrocaledonicum,Santalaceae

The forest tree Santalum austrocaledonicum is endemic to the archipelagos of New Caledonia and Vanuatu, and is threatened by the reduction of the populations due to exploitation. In order to investigate the genetic diversity and structure of this species, we developed eight pairs of primers for nuclear microsatellites. These loci were polymorphic in all the populations, with a mean of three to 33 alleles per locus.     

PCR primers for trinucleotide and tetranucleotide microsatellites in greater amberjack,Seriola dumerili

Eighteen nuclear‐encoded microsatellites from a genomic DNA library of greater amberjack, Seriola dumerili, were isolated and characterized. The microsatellites include 13 perfect (five tetranucleotide and eight trinucleotide) and five imperfect (three tetranucleotide, one trinucleotide and one combination dinucleotide/trinucleotide) repeat motifs. The number of alleles at the 18 microsatellites among a sample of 29 fish ranged from two to 20; gene diversity (expected heterozygosity) ranged from 0.068 to 0.950, whereas observed heterozygosity ranged from 0.069 to 0.966. Following Bonferroni correction, genotypes at all 18 microsatellites fit expectations of Hardy–Weinberg equilibrium, and all pairwise comparisons of microsatellites did not deviate significantly from genotypic equilibrium. Greater amberjack support commercial and recreational fisheries along both the Atlantic and the Gulf coasts of the USA and represent a species with potential for worldwide aquaculture. The microsatellites developed will be useful for population genetic studies of ‘wild’ populations and breeding studies of domesticated populations.     

Isolation and characterization of microsatellite loci in Santalum insulare,Santalaceae

The forest tree Santalum insulare is endemic to the archipelagos of eastern Polynesia and is threatened by important anthropic pressures. In order to investigate the genetic diversity and structure of this species, we developed eight pairs of primers for nuclear microsatellites. One hundred and sixty‐two individuals distributed in 10 islands were analysed. These loci were all polymorphic and displayed three to 15 alleles per locus. Significant Hardy–Weinberg deviation was detected and interpreted as a result of Wahlund effect. These markers constitute an efficient tool in investigating the genetic patterns of S. insulare populations.     

Genetic evaluation of alternative silvicultural systems in coastal montane forests: western hemlock and amabilis fir

Genetic diversity and mating system were quantified for shelterwood, patch cut and green tree-retention silvicultural systems, and compared to adjacent old-growth. This is a component of a larger study conducted in montane old-growth forests of coastal British Columbia to evaluate the feasibility and ecological consequences of alternative silvicultural systems. The experiment includes replicated treatments representing a range of overstory removal adjacent to old-growth and clearcut areas. Based on 22 electrophoretically assayed loci, the effects of silvicultural systems on genetic parameters of amabilis fir (Abies amabilis and western hemlock (Tsuga heterophylla were assessed by comparing an average number of alleles per locus, the percent polymorphic loci, and observed and expected heterozygosity between parental populations and naturally regenerated progeny as well as among treatments. Genetic variation in natural regeneration was greater than in parental populations, especially for low-frequency alleles. Silvicultural treatments caused no significant differences in amabilis fir genetic-diversity parameters, while the shelterwood system resulted in lower observed and expected heterozygosity in western hemlock. Nei's genetic distance revealed that all parental populations were extremely similar. The two species had contrasting mating system dynamics with amabilis fir producing higher levels of correlated paternity and inbreeding with wider variation among individual tree outcrossing-rate estimates. Western hemlock had significant levels of correlated paternity only for the green tree and shelterwood treatments demonstrating family structuring inversely related to stand density. Inbreeding in western hemlock was significant but lower than that observed for amabilis fir with a J-shaped distribution for individual tree multilocus outcrossing-rate estimates. The pollination and dispersal mechanisms of the two species represent the most-likely factors causing these differences. Artificial regeneration may be utilized to augment the genetic resources of natural ingress.     

Development of microsatellite markers for <Emphasis Type="Italic">Manilkara maxima</Emphasis> T.D. Penn. (Sapotaceae) and their use in conservation genetics

Manilkara maxima is an endemic tree species of the Atlantic Forest in southern Bahia, Brazil. It is considered important for forest conservation due to its mutualistic interactions with endemic and endangered animals. Our aim was to develop microsatellite markers to estimate genetic diversity in order to provide information for effectiveness of future conservation programs. We used next generation sequencing technology to develop the first specific microsatellite markers for M. maxima. Seventeen new microsatellite loci were applied in 72 individuals sampled in three natural populations. On average, the number of alleles per loci was 8.8. The expected heterozygosity varied between 0.72 and 0.77, indicating that the developed set of molecular markers is useful for genetic diversity studies. Additionally, the estimated value for the combined probability of exclusion (Q) was greater than 0.999, which indicates the powerful of these molecular tools for paternity and kinship analysis. Our results demonstrate that the set of microsatellites developed in this work is a powerful tool for population genetics, molecular ecology and conservation biology purposes.     

Small effect of fragmentation on the genetic diversity of Dalbergia monticola, an endangered tree species of the eastern forest of Madagascar, detected by chloroplast and nuclear microsatellites

Background and Aims

The oriental forest ecosystem in Madagascar has been seriously impacted by fragmentation. The pattern of genetic diversity was analysed on a tree species, Dalbergia monticola, which plays an important economic role in Madagascar and is one of the many endangered tree species in the eastern forest.

Methods

Leaves from 546 individuals belonging to 18 small populations affected by different levels of fragmentation were genotyped using eight nuclear (nuc) and three chloroplast (cp) microsatellite markers.

Key Results

For nuclear microsatellites, allelic richness (R) and heterozygosity (H_e,nuc) differed between types of forest: R = 7·36 and R = 9·55, H_e,nuc = 0·64 and H_e,nuc = 0·80 in fragmented and non-fragmented forest, respectively, but the differences were not significant. Only the mean number of alleles (N_a,nuc) and the fixation index F_IS differed significantly: N_a,nuc = 9·41 and N_a,nuc = 13·18, F_IS = 0·06 and F_IS = 0·15 in fragmented and non-fragmented forests, respectively. For chloroplast microsatellites, estimated genetic diversity was higher in non-fragmented forest, but the difference was not significant. No recent bottleneck effect was detected for either population. Overall differentiation was low for nuclear microsatellites (F_ST,nuc = 0·08) and moderate for chloroplast microsatellites (F_ST,cp = 0·49). A clear relationship was observed between genetic and geographic distance (r = 0·42 P < 0·01 and r = 0·42 P = 0·03 for nuclear and chloroplast microsatellites, respectively), suggesting a pattern of isolation by distance. Analysis of population structure using the neighbor-joining method or Bayesian models separated southern populations from central and northern populations with nuclear microsatellites, and grouped the population according to regions with chloroplast microsatellites, but did not separate the fragmented populations.

Conclusions

Residual diversity and genetic structure of populations of D. monticola in Madagascar suggest a limited impact of fragmentation on molecular genetic parameters.