EXPERIMENTAL STUDIES OF PONDEROSA PINE. III. DIFFERENCES IN PHOTOSYNTHESIS,STOMATAL CONDUCTANCE,AND WATER-USE EFFICIENCY BETWEEN TWO GENETIC LINES

As part of an intensive study of heritable differences among the progeny of Pinus ponderosa parents from two contrasting habitats (coastal vs. interior, continental), we examined the potential for differences in photosynthesis rate, stomatal conductance, and photosynthetic water-use efficiency. Plants from a cross between two coastal parents (ponderosa × ponderosa) exhibited lower photosynthetic water-use efficiencies, relative to plants from a coastal × interior cross (ponderosa × scopulorum). The lower water-use efficiencies in the ponderosa × ponderosa plants were evident as a lower ratio of external to intercellular CO₂ concentrations and higher stomatal conductances at any given rate of photosynthesis. The ponderosa × scopulorum plants exhibited lower stomatal conductances over a range of leaf-to-air water vapor concentration differences, which was partially explained by lower stomatal densities. The ponderosa × scopulorum plants also exhibited lower maximum photosynthesis rates and lower needle nitrogen concentrations. Taken together, the results suggest that in adapting to drier habitats, P. ponderosa has acquired improved water-use efficiencies and lower transpiration rates, but at the expense of reduced maximum photosynthesis rates.     

EXPERIMENTAL STUDIES IN PONDEROSA PINE. I. RELATIONSHIP BETWEEN VARIATION IN PROTEINS AND MORPHOLOGY

Ponderosa pine from California (Pinus ponderosa var. ponderosa) were crossed to one another and also to individuals from the Rocky Mountains (P. p. var. scopulorum). All crosses involved a single mother and a single pollen donor. Patterns of inheritance of electrophoretically-detectable loci followed Mendelian expectations with one exception. Shikimate dehydrogenase showed unpredictable banding patterns in intervarietal crosses. Variability at these biochemical loci was compared to variability at 14 morphometric characters. The relationship between these two levels of variation is complex and permits only one generalization: variability at one level is not a good predictor of variability at the other level.     

Genomic mating in outbred species: predicting cross usefulness with additive and total genetic covariance matrices

Diverse crops are both outbred and clonally propagated. Breeders typically use truncation selection of parents and invest significant time, land, and money evaluating the progeny of crosses to find exceptional genotypes. We developed and tested genomic mate selection criteria suitable for organisms of arbitrary homozygosity level where the full-sibling progeny are of direct interest as future parents and/or cultivars. We extended cross variance and covariance variance prediction to include dominance effects and predicted the multivariate selection index genetic variance of crosses based on haplotypes of proposed parents, marker effects, and recombination frequencies. We combined the predicted mean and variance into usefulness criteria for parent and variety development. We present an empirical study of cassava (Manihot esculenta), a staple tropical root crop. We assessed the potential to predict the multivariate genetic distribution (means, variances, and trait covariances) of 462 cassava families in terms of additive and total value using cross-validation. Most variance (89%) and covariance (70%) prediction accuracy estimates were greater than zero. The usefulness of crosses was accurately predicted with good correspondence between the predicted and the actual mean performance of family members breeders selected for advancement as new parents and candidate varieties. We also used a directional dominance model to quantify significant inbreeding depression for most traits. We predicted 47,083 possible crosses of 306 parents and contrasted them to those previously tested to show how mate selection can reveal the new potential within the germplasm. We enable breeders to consider the potential of crosses to produce future parents (progeny with top breeding values) and varieties (progeny with top own performance).     

GENETIC VARIATION IN THE PONDEROSAE OF THE SOUTHWEST

Ninety-five seedling populations of southwestern ponderosa pine (Pinus ponderosa var. scopulorum) along with single populations of Pinus engelmannii and Pinus arizonica were compared in four environmentally disparate common gardens. Differentiation among ponderosa pine populations was detected for a diverse assortment of variables that included patterns of shoot elongation, measures of growth potential, winter and spring freezing damage, and leaf characteristics. Multiple regression models accounted for as much as 85% of the variance among populations and described complex clines that were dominated by elevational and latitudinal effects. Although P. ponderosa, P. arizonica, and P. engelmannii were readily differentiated, the performance of progenies from one population suggested introgression primarily involving P. ponderosa and P. arizonica but also implicating P. engelmannii.     

VARIATION IN SEED CHARACTERS IN NEMOPHILA MENZIESII: EVIDENCE OF A GENETIC BASIS FOR MATERNAL EFFECT

A growing body of evidence indicates that phenotypic selection on juvenile traits of both plants and animals may be considerable. Because juvenile traits are typically subject to maternal effects and often have low heritabilities, adaptive responses to natural selection on these traits may seem unlikely. To determine the potential for evolutionary response to selection on juvenile traits of Nemophila menziesii (Hydrophyllaceae), we conducted two quantitative genetic studies. A reciprocal factorial cross, involving 16 parents and 1960 progeny, demonstrated a significant maternal component of variance in seed mass and additive genetic component of variance in germination time. This experiment also suggested that interaction between parents, though small, provides highly significant contributions to the variance of both traits. Such a parental interaction could arise by diverse mechanisms, including dependence of nuclear gene expression on cytoplasmic genotype, but the design of this experiment could not distinguish this from other possible causes, such as effects on progeny phenotype of interaction between the environmental conditions of both parents. The second experiment, spanning three generations with over 11,000 observations, was designed for investigation of the additive genetic variance in maternal effect, assessment of paternal effects, as well as further partitioning of the parental interaction identified in the reciprocal factorial experiment. It yielded no consistent evidence of paternal effects on seed mass, nor of parental interactions. Our inference of such interaction effects from the first experiment was evidently an artifact of failing to account for the substantial variance among fruits within crosses. The maternal effect was found to have a large additive genetic component, accounting for at least 20% of the variation in individual seed mass. This result suggests that there is appreciable potential for response to selection on seed mass through evolution of the maternal effect. We discuss aspects that may nevertheless limit response to individual selection on seed mass, including trade-offs between the size of individual seeds and germination time and between the number of seeds a maternal plant can mature and their mean size.     

Classification tree and minimum-volume ellipsoid analyses of the distribution of ponderosa pine in the western USA

Aim Ponderosa pine (Pinus ponderosa Douglas ex Lawson & C. Lawson) is an economically and ecologically important conifer that has a wide geographic range in the western USA, but is mostly absent from the geographic centre of its distribution – the Great Basin and adjoining mountain ranges. Much of its modern range was achieved by migration of geographically distinct Sierra Nevada (P. ponderosa var. ponderosa) and Rocky Mountain (P. ponderosa var. scopulorum) varieties in the last 10,000 years. Previous research has confirmed genetic differences between the two varieties, and measurable genetic exchange occurs where their ranges now overlap in western Montana. A variety of approaches in bioclimatic modelling is required to explore the ecological differences between these varieties and their implications for historical biogeography and impending changes in western landscapes. Location Western USA. Methods We used a classification tree analysis and a minimum‐volume ellipsoid as models to explain the broad patterns of distribution of ponderosa pine in modern environments using climatic and edaphic variables. Most biogeographical modelling assumes that the target group represents a single, ecologically uniform taxonomic population. Classification tree analysis does not require this assumption because it allows the creation of pathways that predict multiple positive and negative outcomes. Thus, classification tree analysis can be used to test the ecological uniformity of the species. In addition, a multidimensional ellipsoid was constructed to describe the niche of each variety of ponderosa pine, and distances from the niche were calculated and mapped on a 4‐km grid for each ecological variable. Results The resulting classification tree identified three dominant pathways predicting ponderosa pine presence. Two of these three pathways correspond roughly to the distribution of var. ponderosa, and the third pathway generally corresponds to the distribution of var. scopulorum. The classification tree and minimum‐volume ellipsoid model show that both varieties have very similar temperature limitations, although var. ponderosa is more limited by the temperature extremes of the continental interior. The precipitation limitations of the two varieties are seasonally different, with var. ponderosa requiring significant winter moisture and var. scopulorum requiring significant summer moisture. Great Basin mountain ranges are too cold at higher elevations to support either variety of ponderosa pine, and at lower elevations are too dry in summer for var. scopulorum and too dry in winter for var. ponderosa. Main conclusions The classification tree analysis indicates that var. ponderosa is ecologically as well as genetically distinct from var. scopulorum. Ecological differences may maintain genetic separation in spite of a limited zone of introgression between the two varieties in western Montana. Two hypotheses about past and future movements of ponderosa pine emerge from our analyses. The first hypothesis is that, during the last glacial period, colder and/or drier summers truncated most of the range of var. scopulorum in the central Rockies, but had less dramatic effects on the more maritime and winter‐wet distribution of var. ponderosa. The second hypothesis is that, all other factors held constant, increasing summer temperatures in the future should produce changes in the distribution of var. scopulorum that are likely to involve range expansions in the central Rockies with the warming of mountain ranges currently too cold but sufficiently wet in summer for var. scopulorum. Finally, our results underscore the growing need to focus on genotypes in biogeographical modelling and ecological forecasting.     

The effects of pollen composition on fitness components in a neotropical herb

Summary Experimental pollinations of Costus allenii (Zingiberaceae) were conducted to assess the effects of pollen composition on fitness. Plants were selfed, outcrossed with the first nearest neighbor, and outcrossed with pollen mixtures obtained from the nearest 2, 3, and 5 plants. Cross type had a significant effect on seed production, seed weight and total-plant dry weight. Progeny from crosses with 3, and 5 parents grew significantly larger than selfed progeny, or those from 1-parent crosses. Competition experiments indicated the superiority of progeny from 3-, and 5-parent crosses over progeny from 1-parent crosses, but no differences in competitive ability were observed between progeny from 3-, and 5-parent crosses. Relative fitness, based on 1) seed production, 2) percent germination, and 3) dry weight, varied significantly among crosses, and was greatest for crosses with 3 parents and lowest for selfs. The relative fitness of progeny from 5-parent crosses was lower than that of all other outcrossed classes. We suggest that the significant effect of pollen composition on fitness results from variation in the genetic similarity of seed and pollen parents, which is a function of spatial distribution and population structure.     

Drought responses of conifers in ecotone forests of northern Arizona: tree ring growth and leaf δ<Superscript>13</Superscript>C

We sought to understand differences in tree response to meteorological drought among species and soil types at two ecotone forests in northern Arizona, the pinyon-juniper woodland/ponderosa pine ecotone, and the higher elevation, wetter, ponderosa pine/mixed conifer ecotone. We used two approaches that provide different information about drought response: the ratio of standardized radial growth in wet years to dry years (W:D) for the period between years 1950 and 2000 as a measure of growth response to drought, and ¹³C in leaves formed in non-drought (2001) and drought (2002) years as a measure of change in water use efficiency (WUE) in response to drought. W:D and leaf ¹³C response to drought for Pinus edulis and P. ponderosa did not differ for trees growing on coarse-texture soils derived from cinders compared with finer textured soils derived from flow basalts or sedimentary rocks. P. ponderosa growing near its low elevation range limit at the pinyon-juniper woodland/ponderosa pine ecotone had a greater growth response to drought (higher W:D) and a larger increase in WUE in response to drought than co-occurring P. edulis growing near its high elevation range limit. P. flexilis and Pseudotsuga menziesii growing near their low elevation range limit at the ponderosa pine/mixed conifer ecotone had a larger growth response to drought than co-occurring P. ponderosa growing near its high elevation range limit. Increases in WUE in response to drought were similar for all species at the ponderosa pine/mixed conifer ecotone. Low elevation populations of P. ponderosa had greater growth response to drought than high-elevation populations, whereas populations had a similar increase in WUE in response to drought. Our findings of different responses to drought among co-occurring tree species and between low- and high-elevation populations are interpreted in the context of drought impacts on montane coniferous forests of the southwestern USA.     

Variation and inheritance of resistance to cypress aphid, Cinara cupressi Buckton in Cupressus lusitanica Miller

Cupressus lusitanica seedlings from open-pollinated seeds of 18 families were inoculated with day-old first instar Cinara cupressi. Aphid survival was used to determine the genetic basis and inheritance of resistance to the insect. There was marked variation in aphid survival both between and within families. An individual-tree narrow-sense heritability of 0.76 ± 0.30 shows strong additive genetic control which could allow effective selection and breeding for resistance. Resistant parents produced resistant progeny while susceptible parents produced susceptible progeny. There were notable exceptions as some susceptible parents produced highly resistant progeny, indicating that they had acquired pollen from resistant neighbours. Recovery of aphid damaged trees is evident implying that care should be taken in selecting for resistance. The strong additive variance and potentially high heritability indicate that one cycle of selection may yield a resistant population, while intraspecific crossing may give better results. Implications of the results in a C. lusitanica breeding programme are discussed.     

HISTORICAL SEPARATION AND PRESENT GENE FLOW THROUGH A ZONE OF SECONDARY CONTACT IN PONDEROSA PINE

I examined the effects of historical division and secondary contact between eastern and western varieties of ponderosa pine (Pinus ponderosa Laws Pinaceae) on extant patterns of genetic variation. Fossil and biogeographic evidence both indicate that the current point of contact between these two varieties represents secondary contact following historical separation during the Wisconsin glaciation. Current gene flow was assessed by observing the degree of introgression of paternally inherited cpDNA and maternally inherited mtDNA polymorphisms. Both seeds and pollen are wind dispersed in ponderosa pine. Introgression was primarily from west to east, the direction of the prevailing wind, for both organelles, but introgression of cpDNA far exceeded that of mtDNA. Thus pollen is the main agent of contemporary gene flow between the two varieties. Neither seeds nor pollen showed enough introgression since secondary contact to have homogenized the two gene pools. However, allozyme differentiation was minimal. This calls into question assumptions of selective neutrality for at least some of the markers. Theory predicts that nuclear markers will show a high locus-to-locus variance of F_ST following historical separation. This prediction is confirmed by the allozyme data for ponderosa pine, and may provide a useful means of identifying historical separations from allele frequency data.     

Primers designed to amplify a mitochondrial nad1 intron in ponderosa pine, Pinus ponderosa, limber pine, P. flexilis, and Scots pine, P. sylvestris

The b/c intron of the mitochondrial nad1 gene, was sequenced to characterize the indel region of ponderosa pine, Pinus ponderosa. The sequence in ponderosa pine was aligned with the sequence in Scots pine, Pinus sylvestris, to design seven primers that are useful for sequencing and for revealing size variation in amplified fragments in ponderosa pine, Scots pine, and limber pine, Pinus flexilis. These primers reveal variability in all three species, and the pattern of variability within ponderosa pine is described by a preliminary survey. The indel region of ponderosa pine contains three distinct elements with lengths of 31, 32, and 34 bp. Received: 1 March 2000 / Accepted: 14 April 2000<@head-com-p1a.lf>Communicated by P.M.A. Tigerstedt     

INHERITANCE OF MERISTIC VARIATION AND THE EVOLUTION OF DEVELOPMENTAL STABILITY IN RAINBOW TROUT

We found relatively high heritabilities in the narrow sense for seven of eight meristic characters in a population of rainbow trout using regression of mean progeny values on mid-parent values. In sharp contrast, there is no statistically significant additive genetic variance controlling developmental stability, as measured by fluctuating asymmetry (h² = 0.02). However, there is a significant correlation between the average heterozygosity of each family at isozyme loci and the average number of asymmetric traits per individual. We have previously reported a strong correlation between heterozygosity at protein loci and decreased fluctuating asymmetry in this and other salmonid populations. Thus, there is little or no additive, but substantial dominance, genetic variation affecting fluctuating asymmetry. This suggests that there has been directional selection for increased developmental stability.     

QUANTITATIVE GENETICS OF SEQUENTIAL LIFE-HISTORY AND JUVENILE TRAITS IN THE PARTIALLY SELFING PERENNIAL,AQUILEGIA CAERULEA

We determined the genetic basis of several traits related to overall fitness of Aquilegia caerulea, a perennial herb of the Rocky Mountains in western North America. To obtain measures of heritability relevant to the evolutionary potential of wild populations, we performed full and partial diallel crosses and studied progeny performance in the field. Based on a joint analysis of two designs with a total of 18 parents and 102 crosses, we detected significant maternal variance for seed mass and emergence time, but this component was negligible for later-expressed traits. Low heritability and evidence that maternal effects on seed mass are largely environmental suggest that in this population there is little evolutionary potential for change in seed mass under conditions experienced during the study. Seed mass varied depending on particular combinations of parents and cross direction. Such an interaction can have several different biological interpretations, including that particular maternal parents selectively provision embryos sired by particular pollen genotypes. Width of the first true leaf after 4 wk of growth and leaf size of juvenile plants at years one and two were significantly heritable and positively genetically correlated. Juvenile survival exhibited significant dominance variance, as expected from evidence of inbreeding depression in this trait. In contrast, for other traits that exhibit inbreeding depression in this population (seed mass and third-year leaf size), dominance variance was negligible.     

Prediction of progeny variation in oat from parental genetic relationships

Summary The ability to predict agronomic performance of progeny from a cross would be a great benefit to plant breeders in selecting parents. The predictive value of parental genetic relationships estimating F₁ progeny means and F₄ family variances of nine argronomic traits was tested in 76 oat crosses, using genetic distance measures based on coefficients-of-parentage, quantitatively inherited morphological characters, and discretely inherited biochemical and morphological characters. Coefficients-of-parentage were better predictors of F₁ performance than similarity measures derived from plant morphology or discretely inherited characters. Combined distance measures were better estimators of F₁ specific combining ability (SCA) effects than any single measure. Among cultivars of similar adaptation and quantitative morphology, crosses between parents with high coefficients-of-parentage gave higher SCA effect values than crosses of distantly related parents for grain yield and total biomass. The opposite was found for crosses among cultivars of different adaptation or quantitative morphology. The best predictor of trait variances among F₄ families was coefficients-of-parentage. Crosses between more distantly related parents produced larger variances among families than crosses between closely related parents for plant biomass. For grain yield, test weight, heading date, grain filling period, and maturity date, crosses between more closely related parents produced larger among-family variances than crosses of distantly related parents. Crosses between more distantly related parents involved at least one parent unadapted to central New York, and resulted in most of the progeny being generally unadapted. This, in part, may account for the low genetic variances for heading date, test weight, and grain yield in crosses of distantly related parents.     

Genetic variability of mineral concentrations in Festuca arundinacea Schreb.

Summary Nine randomly chosen clones of tall fescue (Festuca arundinacea Schreb.) were mated in all possible combinations to determine the nature of genetic variation for Mg, Ca, K, and P concentrations in a broad genetic base population. General combining ability mean squares were significant for most variables, whereas specific combining ability mean squares were not significant in most instances indicating that additive genetic variance was more important. Genotype x year interactions were significant for most variables, suggesting that selection should be evaluated over many environments. Broad-sense heritability estimates based on parental and progeny variance components were generally high for P, K, Ca, and Mg but low for the ratio K/(Ca + Mg). Narrow-sense heritabilities for these minerals were close to the broad sense values since the additive genetic variance was the largest component of the total genetic variation. Correlations between mineral concentrations and herbage dry matter yield were low. It was concluded that adequate genetic variation exists to improve mineral concentration without altering herbage dry matter yields.Journal Series no. 5886     

GENETIC STRUCTURE OF GEOGRAPHICALLY MARGINAL POPULATIONS OF PONDEROSA PINE

Eleven populations of ponderosa pine (Pinus ponderosa) located on an east-west transect of the Palmer Divide of Central Colorado were studied. Within each population transects of approximately 20 × 250 m were established. In seven populations the transects represented contrasting ecological conditions such as north and south facing slopes. Within each transect 200–250 trees greater than 10 cm in height were tagged, their locations mapped, their ages determined and their diameters and heights measured. Electrophoretic analyses of mature needle tissue indicated that while most of the allozyme variation resided within transects, significant genetic heterogeneity could be found between populations and transects. Among populations consistent patterns of genetic variation were observed at only two loci. Five of the 13 loci had consistent but very small differences between transects within populations. Considerable genetic heterogeneity occurred among 20 m sections of the transects and among age classes within transects. The genetic structure of these populations can be explained by mosaic patterns of seedling recruitment characteristic of these ponderosa pine stands. Only a few maternal trees contribute to each patch of seedlings, and this localized founder effect produces the genetic structure observed in these populations.     

Genetics of nodulation in Chickpea (Cicer arietinum L.)

Combining ability, components of genetic variance and graphic analysis revealed that nodulation in the cultivars of Chickpea (Cicer arietinum L.) under study, was predominantly under the control of non-additive gene action although substantial additive effect was also present. The crosses giving high specific combining ability effects also manifested highly significant positive heterosis. The parents F-61, Giza and Annegiri possessed mostly dominant alleles while Phule G-5, NEC-249 and N-31 possessed mostly recessive alleles having positive effect on nodule weight.     

Genetic studies on resistance to Valsa canker in apple: genetic variance and breeding values estimated from intra- and inter-specific hybrid progeny populations

Malus sieboldii Rehd. exhibits high levels of resistance to Valsa canker caused by Valsa ceratosperma (Tode ex Fr.) Maire while cultivated apples (Malus domestica Borkh.) are susceptible to the disease. In this study, progenies from 23 full-sib families derived from both inter- and intra-specific hybridization among 16 Malus genotypes as parents were assessed for resistance to V. ceratosperma (Vc) for two seasons using an excised shoot assay to determine the pattern of inheritance of the resistance and to also estimate the variance components, narrow-sense heritability, and breeding values of parental genotypes. Generally, M. sieboldii × M. domestica and its reciprocal crosses had more resistant progenies to Vc than intra-specific crosses of M. domestica. Resistance to Vc expressed as the relative lesion length among progenies showed continuous variation irrespective of cross, suggesting the quantitative nature of the resistance to the three virulent isolates of Vc that were tested. Resistance to Vc using the progeny population was analyzed using a mixed linear model based on restricted maximum likelihood. The parental effect (general combining ability (GCA)) was significant while the interaction effect between parents (specific combining ability (SCA)) was relatively small and non-significant. The ratio of SCA/GCA variance was about 32%, suggesting that additive genetic variance had a major contribution to the total genetic variance for resistance to Vc. There was a positive correlation (r = 0.49, p < 0.01) between mid-parental GCA and SCA predictions among 23 full-sib families for the resistance. Narrow-sense heritability estimated by sib analysis was moderate ( [^(h)]² = 0.29 ) \left( {{{\hat{h}}^2} = 0.29} \right) . The predicted breeding values (BV) of the 16 parents indicated that M. sieboldii “Sanashi 63” and “Hayanarisanashi 1” would be useful for breeding for high levels of resistance to Vc.     

Combining ability analysis over environments in diallel crosses of linseed (Linum usitatissimum L.)

Summary This paper reports on combining ability studies for yield and its component traits in diallel crosses involving ten ecogeographically and genetically diverse linseed (Linum usitatissimum L.) cultivars in the F₂ generation over three locations. The general combining ability (GCA) and specific combining ability (SCA) mean squares were significant at all three locations for all traits. Combined analysis over locations showed the same trend of significance. The ratio of GCA to SCA mean squares was significant for all the traits in individual location analysis as well as in combined analysis. This indicated the predominant role of additive gene effects in the inheritance of these characters. The GCA mean squares were several times larger than SCA mean squares for all the traits, indicating the presence of considerable magnitude of additive genetic variance and the additive x additive components of the epistatic variance. Consequently, effective selection should be possible within these F₂ populations for all characters. Significant genotype-location and GCA-location interactions indicated that more than one test location is required to obtain reliable information. The inexpensive and reliable procedure used for making the choice of parents was the determination of breeding values of the parents on the relative performance of their F₂ progeny bulks.Part of the thesis submitted by the senior author in partial fulfillment of the requirements for the Ph. D. degree of the Marathwada Agricultural University, Parbhani, 431 402, India     

Scope for genetic enhancement of the parasitisation potential of four native strains of Trichogrammatoidea sp. nr. lutea Girault (Hymenoptera: Trichogrammatidae) in Kenya

In response to emerging interest in commercial mass production of Trichogramma for Helicoverpa armigera biocontrol in eastern Africa, laboratory experiments were undertaken to assess the scope for genetic enhancement of the parasitisation potential of native strains of the local common trichogrammatid species, Trichogrammatoidea sp. nr. lutea. Four promising strains (ex-Kilifi – Kilifi District, ex-Kwa Chai – Kibwezi District, ex-Rarieda – Bondo District and ex-Ebuhayi, Kakamega District) were tested for cross-mating in reciprocal combinations with focus on fecundity and progeny female ratio. While all the crosses resulted in F1 progeny of both sexes, significant differences were observed between homogamic and reciprocal heterogamic crosses in fecundity, progeny production, proportion of female progeny and adult longevity. Among all the crosses, the cross between ex-Rarieda strain females and ex-Kilifi strain males resulted in progeny that was significantly superior in fecundity and progeny female ratio. Conversely, Kilifi strain females crossed to males from ex-Rarieda strain gave rise to progeny with relatively low fecundity and female ratio. There were significant differences between homogamic crosses and most reciprocal heterogamic crosses in the major biological attributes. Genotypic and phenotypic variance-covariance matrices generated for six life-history traits showed high positive correlations for most traits in both inbred (P<0.05) and reciprocal heterogamic crosses (P<0.05 and P<0.001). Fecundity and number of female offspring were the most important factors in the heterogamic crosses. The results confirmed the scope for genetic enhancement through inter-strain crossing for improving the field impact potential of T. sp. nr. lutea being targeted for commercial mass production.