Testing of selected South African <Emphasis Type="Italic">Pinus</Emphasis> hybrids and families for tolerance to the pitch canker pathogen, <Emphasis Type="Italic">Fusarium circinatum</Emphasis>

Plantations of Pinus spp. constitute approximately 50% of the South African forestry industry. The first aim of this study was to develop a reliable inoculation technique to screen Pinus spp., for tolerance to infection by F. circinatum, which threatens pine forestry in South Africa. Inoculation of branches was compared with stem inoculations and we considered the number of branches or trees required to obtain statistically significant results. Furthermore, variation in the susceptibility of some Pinus families, clones and hybrids was considered. Results showed that branch inoculations were closely correlated with those from stem inoculations, and that it is important to consider branch and stem diameters when assessing susceptibility of trees. Subsequent trials using branch inoculations showed significant differences in F. circinatum tolerance amongst a range of pine species and hybrids of potential interest to forestry in South Africa. Significant differences in susceptibility were also found among clones of two P. radiata families. The most tolerant trees were P. elliottii × caribaea and P. patula × oocarpa hybrids, while the most susceptible species were P. patula, P. greggii and hybrids of these two. This is the first trial considering the susceptibility of Pinus hybrids, Pinus clones and some P. patula provenances, and the results indicate excellent potential for breeding for tolerance to pitch canker in South Africa. Application The accurate selection of disease tolerant planting stock for the South African forestry industry is crucially important for the continued sustainability of this important industry. The work described here provides valuable information on an artificial inoculation technique that will assist the industry in screening trees for tolerance to the pitch canker fungus, F. circinatum. It also provides some indication of the relative susceptibility of a number of Pinus spp., hybrids and families currently being evaluated in the country.     

Variation in pitch canker resistance among provenances of <Emphasis Type="Italic">Pinus patula</Emphasis> and <Emphasis Type="Italic">Pinus tecunumanii</Emphasis> from Mexico and Central America

Pinus patula and high-elevation (HE) sources of P. tecunumanii exhibit intermediate levels of resistance to pitch canker (Fusarium circinatum), compared to extremely resistant species such as P. oocarpa, and extremely susceptible species such as P. radiata. Seedlings from 20 P. patula provenances and 15 HE P. tecunumanii provenances were artificially inoculated with the pitch canker fungus at 21 and 12 weeks of age, respectively, and assessed for resistance 12–20 weeks later. There was important provenance variation in pitch canker resistance for both species. The 20-week LiveStem percentage ranged from 70.3% to 43.6% among the P. patula provenances and 59.6% to 11.7% among HE P. tecunumanii provenances. There was a geographic pattern to the provenance variation, and in both species, low altitude sources demonstrated more resistance than those from high elevation. Provenance variation in pitch canker resistance could be useful when making selection and breeding decisions with these species.     

High genetic diversity of Fusarium circinatum associated with the first outbreak of pitch canker on Pinus patula in South Africa

The disease known as pitch canker results from infection of Pinus species by the fungus Fusarium circinatum. This fungus also causes a serious root disease of Pinus seedlings and cuttings in forestry nurseries. Pinus radiata and P. patula are especially susceptible to the pathogen, but there are no records of pitch canker on P. patula in established plantations. To date, only planting material of this tree species in nurseries or in plantations at the time of establishment have been infected by F. circinatum. Symptoms of pitch canker have recently emerged in an established P. patula plantation in South Africa and this study sought to determine whether the symptoms were caused by F. circinatum. Isolates from cankers were identified as F. circinatum using morphology and DNA-based diagnostic markers. Microsatellite markers were then used to determine the genetic diversity of a collection of 52 isolates. The entire population included 17 genotypes representing 30 alleles, with a greater number of genotypes collected from younger (three- to six-year-old) than older (12- to 19-year-old) trees. Both mating types of F. circinatum were present, but no evidence of sexual recombination was inferred from population genetic analyses. This is the first record globally of pitch canker on P. patula trees in managed plantations. It is of significant concern to South Africa, where P. patula is the most important Pinus species utilised for plantation forestry.     

The taxonomic status of closely related closed cone pines in Mexico and Central America

A taxonomic study on 14 cone and needle traits of 281 trees from 33 provenances of Pinus oocarpa, P. oocarpa var. ochoterenae, P. patula var. longipedunculata, P. patula var. patula and P. tecunumanii in Central America and Mexico was conducted to quantify the affinity between closely related species. A second objective was to determine the geographic range of P. tecunumanii in Mexico. Cluster analysis placed 52% of all provenances in taxa different from those assigned by field foresters. Trees from 15 provenances of what was locally known as Pinus oocarpa var. ochoterenae from Chiapas were statistically indistinguishable from high elevation Central American sources of P. tecuncumanii when assessed for cone and needle characteristics. However, trees from two provenances known locally as P. oocarpa var. ochoterenae from southwestern Oaxaca, Juquila and Tlacuache, were found to be distinct from P. tecunumanii and may be a variant of P. patula. Canonical discriminant analysis was used to determine the taxonomic affinity among newly formed clusters. The spatial (Mahalanobis) distance between P. tecunumanii and P. patula var. patula, P. patula var. longipedunculata, and P. oocarpa was 42.0, 44.4, and 109.4 respectively, and highly significant. The geographic range of Pinus tecunumanii in Mexico appears to be confined to the state of Chiapas.     

Growth and dynamic modulus of elasticity of Pinus patula × Pinus tecunumanii hybrids in Mpumalanga,South Africa

Field establishment of South Africa’s most important commercial pine species, Pinus patula, is severely hampered by the pitch canker fungus, Fusarium circinatum. Importantly, hybrids between P. patula and other pine species tolerant to the pitch canker fungus, such as P. tecunumanii and P. oocarpa, have been identified as an alternative planting stock. In this study, variation in tree volume and dynamic modulus of elasticity (MOE_dym) of the P. patula × P. tecunumanii (low- and high-elevation [LE and HE] ecotypes) hybrid was compared with the P. elliottii × P. caribaea hybrid, and the pure species P. tecunumanii (LE) and P. patula. The MOE_dym was assessed using the Fakkop TreeSonic microsecond instrument across three sites. The results of the study showed that P. patula × P. tecunumanii LE performed significantly better than P. patula × P. tecunumanii HE for volume and MOE_dym, which in turn was significantly better than P. patula. The MOE_dym and tree growth decreased with an increase in elevation. There was significant taxon × site interaction for volume and MOE_dym. The results of these trials suggested that P. patula × P. tecunumanii LE is a suitable alternative to P. patula in the Sabie region of Mpumalanga in South Africa on frost-free sites, in terms of the traits that were assessed.     

The use of field and artificial freezing studies to assess frost tolerance in natural populations of Pinus oocarpa

The susceptibility of Pinus oocarpa to freezing temperatures limits the commercial deployment of the highly productive Pinus patula × P. oocarpa hybrid in South Africa. Identifying P. oocarpa germplasm with increased frost tolerance is important. Twenty-three P. oocarpa provenances, originating from Mexico, Honduras, Guatemala and Nicaragua, were therefore assessed for their tolerance to freezing conditions by analysing field survival after frost events, subjecting needles to freezing temperatures and assessing damage using the electrolyte leakage test, and exposing young plants to freezing temperatures in a semi-controlled environment and scoring tissue damage based on a visual assessment. The performance of many of the provenances represented in the field and artificial freezing studies were similar and there was a strong correlation between provenance ranking in the whole plant freezing and electrolyte leakage test. We therefore support the use of these techniques as a means to assess cold tolerance in P. oocarpa at the provenance level. Provenances from north-western Mexico demonstrated more frost tolerance than those from southern Mexico. Provenances representing Honduras and Guatemala appear to be highly susceptible to frost.     

Selection of Pinus spp. in South Africa for tolerance to infection by the pitch canker fungus

The increasing threats from pests and diseases demand that the South African forest industry explores options to deploy alternative pine species in plantation development. This is especially true for species, such as Pinus patula Schiede and Deppe ex Schltdl. and Cham., which are highly susceptible to the pitch canker fungus Fusarium circinatum. Losses due to F. circinatum have been confined mostly to nurseries and at field establishment resulting in a significant cost to the industry. Although, the fungus has not as yet resulted in stem and branch infections on established P. patula in South Africa, it has caused pitch canker on other, more susceptible species such as P. radiata D. Don., and P. greggii Engelm. ex Parl. As alternatives to P. patula, on the warmer and cooler sites in South Africa, families of P. elliottii Engelm var. elliottii, P. tecunumanii (Schw.) Eguiluz and Perry, P. maximinoi H. E. Moore and P. pseudostrobus Lindl. were screened for tolerance to infection by F. circinatum in greenhouse studies. Seedlings were wounded and inoculated with spores of F. circinatum. Lesion development following inoculation was used to differentiate the levels of tolerance between families. The results showed that P. maximinoi, P. pseudostrobus, and the low elevation variety of P. tecunumanii are highly tolerant to infection with very little family variation. The narrow sense heritability estimates for these species were less than 0.06. In contrast, P. elliottii showed good tolerance with some family variation and a heritability of 0.22, while the high elevation source of P. tecunumanii showed a high degree of family variation and a heritability of 0.59. These results provide the industry with valuable information on pine species tolerant to F. circinatum that could be used as alternatives to P. patula in South Africa.     

Alternative pine hybrids and species to Pinus patula and P. radiata in South Africa and Swaziland

Through the collaborative efforts of companies affiliated with the International Program for Tree Improvement and Conservation (Camcore), a number of pine hybrids have been produced over the last decade. Many of these have been planted in trials across southern Africa that broadly represent winter and summer rainfall areas, with the latter ranging from warm to cold temperate sites. The five-year survival and growth of the hybrids and other pines in 12 of these trials were compared with Pinus radiata in the winter rainfall, and P. patula in the summer rainfall, regions where these species have been planted extensively. Except for the highest altitude site, where freezing conditions are common, the survival of most hybrids and tropical pines was better than P. patula or P. radiata. This was, in part, attributed to their improved tolerance to the pitch canker fungus, Fusarium circinatum, which was present in the nursery at the time of planting. In the winter rainfall area, the P. elliottii × P. caribaea hybrid, P. maximinoi and, surprisingly, the P. patula hybrids performed well. In the summer rainfall regions, hybrids with tropical parents such as P. caribaea, P. oocarpa and P. tecunumanii were more productive in the subtropical/warm temperate zone and, with increasing elevation, those hybrids crossed with P. patula performed relatively better. The P. patula × P. tecunumanii hybrid, particularly when crossed with low-elevation P. tecunumanii, performed exceptionally across most sites.     

Susceptibility of provenances and families of Pinus maximinoi and Pinus tecunumanii to frost in South Africa

The future of South Africa’s most important pine species, Pinus patula, is threatened by the pitch canker fungus, Fusarium circinatum. Pinus maximinoi and P. tecunumanii represent two subtropical species that provide an alternative to planting P. patula on the warmer sites of South Africa. Extending the planting range of P. tecunumanii and P. maximinoi to include higher and colder altitude sites will reduce the area planted to P. patula and the risk of F. circinatum. During 2007 progeny trials of P. tecunumanii and P. maximinoi were planted on a sub-tropical and sub-temperate site. Shortly after the establishment of these trials, unusually cold weather conditions were experienced across South Africa (?3°C at the sub-temperate site) resulting in severe mortality. This provided the opportunity to assess the variation in survival as a measure of frost tolerance within these two species to determine whether it could be improved upon through selection. Results indicated that the variation in survival was under genetic control in P. tecunumanii (h _(0,1) ²  = 0.16, h _L ²  = 0.27) and P. maximinoi (h _(0,1) ²  = 0.11, h _L ²  = 0.23) at the sub-temperate site. Correlations in provenance ranking for survival across sites were high for both species. Moderate correlations in family survival for P. tecunumanii (r = 0.52) were found at the two sites. Improvements in cold tolerance can thus be made in both species extending their planting range to include greater areas planted to P. patula thereby limiting the risk of F. circinatum.     

Genetic characterisation of a Eucalyptus nitens base breeding population in South Africa

The measurement and statistical analysis of data from eight Eucalyptus nitens trials, established in the summer rainfall forestry region of South Africa during the 1980s and 1990s, have enabled the characterisation of the Institute for Commercial Forestry Research's breeding population. Provenance testing showed that the more northerly New South Wales (Australia) Eucalyptus nitens provenances of Barren Mountain and Barrington Tops are distinctly better suited to the summer rainfall areas of South Africa than the southern New South Wales provenances and the Victorian provenance, Penny Saddle. Generally, the species was not badly affected by Coniothyrium canker. High type B genetic correlations for all site pairs, except one comparison, ranged from 0.75 to 0.99 for diameter at breast height (dbh), indicating very little or no genotype × environment interaction for dbh for the genotypes tested in this study. Narrow-sense heritability coefficients ranged from 0.01 to 0.34, indicating that the species generally exhibited sufficient breeding opportunity for improvement of diameter growth. High genetic correlations of greater than 0.90 between diameter measurements at 52 to 62 months after establishment and diameter measurements at 94 or 113 months were found, indicating that selections can be reliably made at five or six years. Predicted genetic gains were highest in the trials at Goedehoop and Arthur's Seat, with increases in dbh of 3.07 cm (17.1%) and 3.17 cm (20.7%), respectively, at full rotation.     

Tolerance of Pinus patula full-sib families to Fusarium circinatum in a greenhouse study

The pitch canker fungus, Fusarium circinatum, has caused large-scale mortality of young Pinus patula Schiede and Deppe ex Schltdl. and Cham. seedlings in nurseries in South Africa since 1990. Diseased seedlings have been inadvertently carried to the field, which in turn have died and has reduced stocking below an acceptable level. Tree breeders have suggested that the only long-term solution to limit infection by this pathogen is to identify and deploy tolerant P. patula families. A commonly used technique to identify tolerant clones is to artificially inoculate open-pollinated progeny from orchard clones with F. circinatum under greenhouse conditions. In these trials, large variation in tolerance to the pathogen among seedlings within open-pollinated families was observed and this could be influenced by the pollen parent. Therefore, identifying individual full-sib families, where both parents are known, should improve the identification of tolerant families, which can then be repeated. In this study, cuttings from control-pollinated P. patula seedling hedges were inoculated with F. circinatum in a greenhouse. The results showed large family variation where some of the full-sib families were similar in tolerance to P. elliottii Engelm. var. elliottii seedlings. Therefore, it is recommended that breeders focus on identifying specific family combinations that are more tolerant to F. circinatum.     

The tolerance of Pinus patula × Pinus tecunumanii, and other pine hybrids, to Fusarium circinatum in greenhouse trials

The field survival of Pinus patula seedlings in South Africa is frequently below acceptable standards. From numerous studies it has been determined that this is largely due to the pitch canker fungus, Fusarium circinatum. Other commercial pines, such as P. elliottii and P. taeda, show good tolerance to this pathogen and better survival, but have inferior wood properties and do not grow as well as P. patula on many sites in the summer rainfall regions of South Africa. There is, thus, an urgent need to improve the tolerance of P. patula to F. circinatum. Operational experience indicates that when P. patula is hybridized with tolerant species, such as P. tecunumanii and P. oocarpa, survival is greatly improved on the warmer sites of South Africa. Field studies on young trees suggest that this is due to the improved tolerance of these hybrids to F. circinatum. In order to test the tolerance of a number of pine hybrids, the pure species representing the hybrid parents, as well as individual families of P. patula × P. tecunumanii, a series of greenhouse screening trials were conducted during 2008 and 2009. The results indicated that species range in tolerance and hybrids, between P. patula and these species, are intermediate in tolerance to F. circinatum. Within P. patula × P. tecunumanii, large family variation exists when pollen from the high elevation source of P. tecunumanii is used. The results of these studies illustrate the importance of developing pine hybrid breeding programs to overcome the susceptibility of our pure species to pathogens such as F. circinatum.     

Provenance variation in growth characters of four subtropical pine species planted in Mexico

In 1995 a species/provenance test was established in the state of Puebla, Mexico to assess and compare the growth of Pinus greggii var. australis, P. maximinoi, P. patula and P. tecunumanii, including 10, 5, 11 and 4 provenances of each species, respectively. Each provenance test lot consisted of seed from 7 to 10 individual trees collected in natural stands. Range-wide samples were included in the test for P. greggii var. australis and P. patula. Material of P. maximinoi and P. tecunumanii from Guatemala, Honduras and Nicaragua, as well as Mexico was established in the test. The test site is in a mountainous area at 1440m elevation, near 20°N latitude. At 5 years of age significant differences were seen at both the species and provenance levels in survival, total height, diameter and volume. The overall test survival was 93%. Variance among species was three time greater than among provenances, comprising 25–33% of the total variation. P. maximinoi had the greatest height, diameter and volume (62dm³) followed by P. tecunumanii (53dm³). P. greggii var. australis had an average volume of 43dm³ and P. patula produced 30dm³. Although P. maximinoi had the lowest survival rate, the San Jeronimo provenance was the most productive of all 30 provenances tested; 98% survival and 73dm³ volume. This was equivalent to 2.8cm annual diameter increment and 2m in height growth per year. Significant growth differences among provenances within species indicate that potential gain exists using provenance selection.     

Disease incidence in Eucalyptus microtheca provenances

A provenance trial of Eucalyptus microtheca F. Muell. was studied in order to select resistant provenances against Cytospara eucalypti twig blight, Pestalotiopsis versicolor stem canker and Botryodiplodia theobromae root collar canker. Out of 21 provenances tried, 4 provenances were screened as resistant against diseases and showed best performance.     

Future outlook for Pinus patula in South Africa in the presence of the pitch canker fungus (Fusarium circinatum)

Approximately 50% of the area planted to softwood trees in South Africa has been established with Pinus patula, making it the most important pine species in the country. More effort has gone into developing this species for improved growth, tree form and wood properties than with any other species. This substantial investment has been threatened in the last 10 years by the pitch canker fungus, Fusarium circinatum. The fungus infects and contaminates nursery plants and, once transferred to the field, causes severe mortality of young trees in the first year after establishment. Although nurserymen have some control of the disease, it is recognised that the best long-term solution to mitigate damage because of F. circinatum infection is to identify tolerant species, clones and hybrids for deployment in plantations in the future. Research has shown that alternative species such as P. tecunumanii, P. maximinoi and P. elliottii are suitable for warm sites. Pine hybrids, particularly between P. patula and the high-elevation sources of P. tecunumanii, appear to be a suitable replacement on subtemperate and temperate sites. Although these alternative species and hybrids are more sensitive to subfreezing temperatures than P. patula, their planting range can be increased by including cold tolerance as a selection criterion. Future breeding efforts will most certainly focus on improving the tolerance of pure P. patula to F. circinatum, which can be achieved by identifying specific family crosses and tolerant clones. The commercial deployment of disease-tolerant control-pollinated P. patula and hybrid families will most likely be established as rooted cuttings, which requires more advanced propagation technology. In the long term, new seed orchards comprised of P. patula clones tolerant to F. circinatum could be used to produce seed for seedling production.     

Provenance and progeny results for the tropical white pine, Pinus chiapensis, at five and eight years of age

Eleven provenances and 132 open-pollinated families of Pinus chiapensis were established at six locations in Brazil, Colombia, and South Africa and assessed for height, volume, and forking percent at five and eight years of age. Provenance × site and family × site interactions were examined in a subset of three trials with 38 open-pollinated families in common. Average volume production of P. chiapensis in Colombia and South Africa was 25 m³/ha/yr and 12 m³/ha/yr, respectively, at eight years of age. Provenance and family differences for height and volume were significant at the individual test sites at eight years of age. The La Trinidad (Chiapas) source was the most productive. Forking percent ranged from 3% in South Africa to 38% in San Jose, Colombia at eight years. Provenance × site interactions were significant for forking but not for height and volume. Family × site interactions were significant for height and volume but not for forking. Forty-six percent of the interaction variance between height and location was contributed by only six families. Height/diameter ratios for P. chiapensis were 0.29 and 0.39 in Colombia and South Africa, respectively, and significantly different.     

Diplodia scrobiculata found in the southern hemisphere

Diplodia scrobiculata, a latent pathogen of Pinus spp. and other conifers with a limited distribution in the United States, Mexico and southern Europe, has not been reported previously in the southern hemisphere. This is unlike its close relative Diplodia pinea that is found in most parts of the world where pines are native or have been introduced. During an intensive D. pinea survey conducted in Pinus patula plantations in eastern parts of South Africa, a small number of isolates atypical of D. pinea were found. Morphological studies and DNA sequence comparisons showed that these isolates represent D. scrobiculata. Microsatellite analyses suggest that the South African isolates of D. scrobiculata might have originated from California. Pathogenicity tests showed that some of the D. scrobiculata isolates were as pathogenic as those of D. pinea on Pinus radiata and Pinus elliottii.     

Endothia gyrosa associated with severe stem cankers on plantation grown Eucalyptus nitens in Tasmania,Australia

Severe stem cankers in Eucalyptus nitens, from a 14-year-old mixed provenance plantation, were associated with infection by Endothia gyrosa, present in its teleomorph state. Surveys of incidence among canker severity classes were carried out in a thinned and pruned stand and an adjacent unthinned and unpruned stand within the affected plantation. No differences in incidence among the canker severity classes were found between the thinned/pruned and unthinned/unpruned stands or between different crown dominance classes within the unthinned/unpruned stand. However, the incidence among canker severity classes was strongly associated with bark roughness with 97% of rough-barked trees developing either annual cankers or cankers causing cambial damage. Stem cankers were found on only 11% of trees with smooth bark. Bark roughness in E. nitens was shown to differ significantly between provenances. Deployment of provenances prone to rough bark in routine plantation establishment may pose a risk of damaging stem canker outbreaks.     

Genetic and site effects on stem breakage in Pinus tecunumanii

Fifteen provenance/progeny tests of Pinus tecunumanii (Schw.) Eguiluz et Perry were assessed for upper-crown stem breakage at 5 to 8 years of age in Brazil, Colombia and South Africa. Five year results indicated that the mean percent stem breakage was moderate in Colombia (14% to 21%), low in South Africa (2% to 6%) and very low in Brazil (<1%). In the Colombian tests, trees from provenances of Mexican origin were more susceptible to wind damage than those from Central America, 20% versus 14%. Provenance order of percent stem breakage at 5 years of age in Colombia was Montebello (27%), La Soledad (25%), San Jose (20%), Chempil (20%), Jitotol (18%), Las Piedrecitas (18%), Celaque (17%), Km 47 (16%), San Vicente (13%), San Jeronimo (13%), San Lorenzo (12%) and Las Trancas (7%). Two half-sib progeny tests assessed at 8 years of age in Colombia with trees from the Mountain Pine Ridge source of P. tecunumanii exhibited 13% stem breakage. Family variation in stem breakage ranged from 0 to 44% in Colombia and 0 to 33% in South Africa at 5 years of age and were highly significant. The Spearman rank correlations in stem breakage of 36 half-sib families common to tests in Colombia and South Africa were r=0.13 and not significant. Family heritabilities for stem breakage at 5 and 8 years of age ranged from 0.17 to 0.63. Results indicate that stem breakage was more common on trees with branches of large diameter and was not strongly related to growth rate.     

Bark thickness and bark volume of Pinus patula in South Africa

In South Africa, tree volume tables and volume functions are usually based on the under bark stem volume. Many forest management studies, however, give the estimated over bark volume, in which case equations are required that estimate bark volume and possibly bark thickness as a function of DBH and tree height. This study deals with the evaluation of geographical regions and tree characteristics on bark thickness and bark volume of Pinus patula in South Africa.