Recovery from topkill of shortleaf pine × loblolly pine hybrids compared to their parent populations

Hybrids between shortleaf pine (Pinus echinata Mill.) and loblolly pine (Pinus taeda L.) have increased since the 1950s throughout the southeastern USA. Previously, greater sprouting capacity and the formation of a basal crook that lowers the height of dormant buds may have favored pure shortleaf pine populations on fire prone sites. The objective of this study was to determine how seasonal timing of topkill by both fire and topclipping affect sprouting of shortleaf × loblolly pine F1 hybrids compared to their parent open-pollinated populations during their third growing season. A factorial combination of top-clipping (hand pruners) and girdling by fire (propane torch) was conducted on November 2010, January, March, and April 2011 and sprouting response was measured after the growing season. Survival of topkilled shortleaf pine (94 %) was greatest followed by hybrid (78 %) and loblolly pine (35 %). However, species effects varied with topkill treatment and treatment date because survival was relatively lower for loblolly and hybrid pines in the burn-only as well as the November and April treatment dates while survival of shortleaf pine was consistently high. The number of sprouts was greatest for shortleaf (32.3) intermediate for hybrid (23.8) and lowest for loblolly pine (12.0). Overall, 83 % of shortleaf pine, 35 % of hybrid pine, and 5 % of loblolly pine exhibited a basal crook. The height from ground line to the lowest sprout was shortest for shortleaf (3.5 mm), intermediate for hybrids (7.7 mm), and largest for loblolly pine (21.3 mm). While the hybrid saplings exhibited intermediate performance in survival, sprouting capacity, and crooking, pure shortleaf pine were superior and are probably better suited to recover from fire.     

Does landscape composition alter the spatiotemporal distribution of the pine processionary moth in a pine plantation forest?

Landscape composition and physiognomy affect community structure and species distribution across space and time. The pine processionary moth (PPM) (Thaumetopoea pityocampa Den. & Schiff., Lepidoptera, Notodontidae) is a common pine defoliator throughout southern Europe and Mediterranean countries. We surveyed the spatiotemporal distribution of the PPM in a pine plantation forest in southwestern France and used the density of the winter nests as a proxy for population density. The study spanned 4 years (2005–2008) and showed a high temporal variability in nest density. We found a strong edge effect with nest densities at stand edges more than twice as large as within-stand densities. At the landscape scale, the spatial distribution of the moth exhibited a significant spatial autocorrelation in 3 out of 4 years of our study. The spatial scales of the autocorrelation ranged from ca. 2 km to more than 22 km. We found a positive correlation between spatial distributions corresponding to certain sampling years, but the relationship was not systematic. Landscape configuration appeared to be an important driver of the PPM spatial pattern. Bivariate Moran’s I correlograms showed that patch richness density as well as the percentage of local landscape covered by various land uses were correlated with population density. The study showed that accounting for landscape characteristics may be important in order to understand forest insect pest distribution, even in cases where the host species is abundant and homogeneously distributed throughout the study area, e.g., pure plantation forests.     

Mapping quantitative trait loci controlling early growth in a (longleaf pine × slash pine) × slash pine BC<Subscript>1</Subscript> family

Random amplified polymorphic DNA (RAPD) markers were employed to map the genome and quantitative trait loci controlling the early growth of a pine hybrid F₁ tree (Pinus palustris Mill. 2 P. elliottii Engl.) and a recurrent slash pine tree (P. elliottii Engl.) in a (longleaf pine 2 slash pine) 2 slash pine BC₁ family consisting of 258 progeny. Of the 150 hybrid F₁ parent-specific RAPD markers, 133 were mapped into 17 linkage groups covering a genetic distance of 1,338.2 cM. Of the 116 slash pine parent-specific RAPD markers, 83 were mapped into 19 linkage groups covering a genetic distance of 994.6 cM. A total of 11 different marker intervals were found to be significantly associated with 13 of the 20 traits on height and diameter growth using MAPMAKER/QTL. Nine of the eleven marker intervals were unique to the hybrid parent 488 genome, and two were unique to the recurrent parent 18-27 genome. The amount of phenotypic variance explained by the putative QTLs ranged from 3.6% to 11.0%. Different QTLs were detected at different ages. Two marker intervals from the hybrid parent 488 were found to have QTL by environment interactions.     

Can exotic pine trees assist in restoration?

Invasive exotic woody species, including conifers escaped from plantations, are usually regarded as serious threats to native biodiversity. Becerra & Montenegro (Applied Vegetation Science, 16, 2013, 2) present an interesting example of invasive Pinus radiata facilitating native woody regeneration in semi‐arid central Chile. However, the positive value of invasive conifers for restoration and conservation will not necessarily apply in all situations.     

Effect of pine foliage damage on the incidence of larval diapause in the pine caterpillar Dendrolimus punctatus （Lepidoptera： Lasiocampidae）

The pine caterpillar Dendrolimus punctatus （Walker） with a larval facultative diapause is one of the most destructive insect pests of the pine tree Pinus massoniana in China. The larvae feeding on pine trees with different damage levels were studied to determine the induction of diapause under both laboratory and field conditions. Developmental duration of larvae before the third instar was the longest when fed with 75%-90% damaged needles, followed by 25%-40% damaged needles and intact pine needles, whereas mortalities did not differ among different treatments under the conditions of 25℃ and critical photoperiod 13.5：10.5 L：D. At 25 ℃, no diapause was induced under 15：9 L：D, whereas 100% diapause occurred under 12：12 L：D regardless of the levels of needle damage. Incidences of larvae entering diapause when they were fed with intact, 25%-40% and 75%- 90% damaged pine needles were 51.7%, 70.8% and 81% under 13.5：10.5 L：D, respectively. Similar results were obtained in the field experiment. Incidence of diapause was significantly different among the pine needle damage levels of pine trees when the photoperiod was close to the critical day length, indicating that the effect of host plants on diapause induction was dependent on the range of photoperiod. The content of amino acid and sugar decreased and tannin increased in pine needles after feeding by the pine caterpillars, suggesting that changed levels of nutrients in damaged needles or a particular substance emitted by damaged pine trees was perhaps involved in the diapause induction of the pine caterpillar.     

Effects of fertilization and herbicides on growth of young loblolly pine and infestations of Nantucket pine tip moth（Lepidoptera： Tortricidae）

A 2-year-old pine plantation was selected to receive treatments of fertilizers and herbicides to evaluate effects on Nantucket pine tip moth infestations and the tree growth parameters of height, diameter and volume increment. Nitrogen and phosphorus fertilizers, and hexazinone and sulfometuron methyl herbicides were used in creating six treatments： （i） control; （ii） phosphorus; （iii） nitrogen and phosphorus; （iv） phosphorus and herbicide; （v） nitrogen, phosphorus and herbicide; and （vi） herbicide. Treatments were applied in 1987 and 1988. In 1987, trees treated with nitrogen, phosphorus and herbicide had significantly greater height, diameter and volume growth than trees not receiving fertilizer treatments, but did not have significantly higher tip moth infestations than control trees. Treatments receiving phosphorus only had much lower tip moth infestation rates than other treatments except nitrogen and phosphorus. In 1988, tip moth infestations were uniformly low, with no differences in treatment effects observed.     

Does tree diversity increase wood production in pine forests?

Recent experimental advances on the positive effect of species richness on ecosystem productivity highlight the need to explore this relationship in communities other than grasslands and using non-synthetic experiments. We investigated whether wood production in forests dominated by Aleppo pine (Pinus halepensis) and Pyrenean Scots pine (Pinus sylvestris) differed between monospecific and mixed forests (2-5 species) using the Ecological and Forest Inventory of Catalonia (IEFC) database which contains biotic and environmental characteristics for 10,644 field plots distributed within a 31,944 km(2) area in Catalonia (NE Spain). We found that in Pyrenean Scots pine forests wood production was not significantly different between monospecific and mixed plots. In contrast, in Aleppo pine forests wood production was greater in mixed plots than in monospecific plots. However, when climate, bedrock types, radiation and successional stage per plot were included in the analysis, species richness was no longer a significant factor. Aleppo pine forests had the highest productivity in plots located in humid climates and on marls and sandstone bedrocks. Climate did not influence wood production in Pyrenean Scots pine forests, but it was highest on sandstone and consolidated alluvial materials. For both pine forests wood production was negatively correlated with successional stage. Radiation did not influence wood production. Our analysis emphasizes the influence of macroenvironmental factors and temporal variation on tree productivity at the regional scale. Well-conducted forest surveys are an excellent source of data to test for the association between diversity and productivity driven by large-scale environmental factors.     

Modifying transient β-glucuronidase expression in pine species using introns

The effect of introns on gene expression was evaluated. Several intron-promoter combinations were introduced by microparticle bombardment into two pine species, stone pine (Pinus pinea L.) and salgareño pine (Pinus nigra Arn. ssp. Salzmannii (Dunal) Franco). Gene expression was evaluated by measuring transient GUS expression. Two promoters (CaMV35S and double CaMV35S modified) and two introns (intron 1 from maize genes alcohol dehydrogenase-1 and Shrunken-1) were used in our study. In both pine species tested, the Sh1-int1 increased transient GUS expression from 2 to 6-fold compared to the intron-less construction. On the contrary, the inclusion of the Adh1-int1 associated with the double CaMV35S modified resulted in a dramatic decrease in the expression in both pine species analyzed. Our results suggest that Sh1-int1 may be useful for the acquisition of the required levels of genetic activity of new agronomic traits introduced into pines.     

Do multiple herbivores maintain chemical diversity of Scots pine monoterpenes?

A central issue in our understanding of the evolution of the diversity of plant secondary metabolites (PSMs) is whether or not compounds are functional, conferring an advantage to the plant, or non-functional. We examine the hypothesis that the diversity of monoterpene PSMs within a plant species (Scots pine Pinus sylvestris) may be explained by different compounds acting as defences against high-impact herbivores operating at different life stages. We also hypothesize that pairwise coevolution, with uncorrelated interactions, is more likely to result in greater PSM diversity, than diffuse coevolution. We tested whether up to 13 different monoterpenes in Scots pine were inhibitory to herbivory by slugs (Arion ater), bank voles (Clethrionomys glareolus), red deer (Cervus elaphus) and capercaillie (Tetrao urogallus), each of which attack trees at a different life stage. Plants containing more α-pinene were avoided by both slugs and capercaillie, which may act as reinforcing selective agents for this dominant defensive compound. Herbivory by red deer and capercaillie were, respectively, weakly negatively associated with δ(3)-carene, and strongly negatively correlated with the minor compound β-ocimene. Three of the four herbivores are probably contributory selective agents on some of the terpenes, and thus maintain some, but by no means all, of the phytochemical diversity in the species. The correlated defensive function of α-pinene against slugs and capercaillie is consistent with diffuse coevolutionary processes.     

Is fire a selective force of seed size in pine species?

Fire is selectively shaping most of the traits of plants growing in fire-prone environments. However, seed size and other features related to seed production have not been studied in the light of the evolutionary role of fire. Our research tests the hypothesis that larger seeds have a higher chance of surviving wildfires and produce more vigorous seedlings with a lower death rate. To test this hypothesis the germination and early seedling growth of five Spanish pine species were studied. Weight, length and width of all seeds were measured. The biomass (fresh and dry weight) and length (root and total) of subsequent seedlings were also measured after 30 d from emergence. Seeds were submitted to elevated temperatures for periods in which the chance of survival was 50 % (calculated by means of a logistic model for each pine species). The differences observed among species suggests that fire may be adaptively shaping seed size in pines with larger seeds (Pinus canariensis and P. pinaster), because larger seeds are more likely to survive after heat shocks. Furthermore, in P. canariensis, seedlings after heat treatment are even larger than those submitted to control. In P. halepensis, despite being well adapted to fire, our results indicated no relationships between fire and seed characteristics. Finally, although heat treatment has a general adverse effect on seedling growth in the case of the two subalpine pines, we have detected a positive relationship between seed size and seedling growth but only in the largest seeds. This might also suggest the relevance of fire as a selective force for these pines which is outperformed by the relevance of dispersal and emergence time as adaptive traits in the post-fire scenario.     

Postfire regeneration of Aleppo pine – the temporal pattern of seedling recruitment

The temporal dynamics of Pinus halepensis seedling emergence, density and survival were studied in detail during the first postfire year, in five Aleppo pine forests of Attica (Stamata, Villia, Avlona, Kapandriti and Ag. Stefanos; burned in June 1990, July 1990, September 1991, September 1992 and July 1993, respectively). The timing of emergence and establishment of Aleppo pine seedlings was found to be correlated with the prevailing meteorological conditions - mainly rainfall and to a lesser degree temperature. In most cases, seed germination and seedling emergence took place in a ``single massive wave'' (resembling a right-skewed normal distribution), shortly after the onset of the first postfire rainy season, (Stamata, Villia and Avlona; 80-90% of the total cohort emerged during November-December). However, variations in the timing and spread of this pattern were observed in Agios Stefanos and Kapandriti. In the former, there was a considerable delay of seedling emergence (80% by the end of February); in the latter, a bimodal pattern was found (peaks in December-February and March-April). The dynamics of pine seedling mortality (which amounted to a yearly 23-50% of the total cohort) showed two major, discrete peaks of almost equal importance during the first postfire year: at the initial seedling establishment phase (in the winter) and during the dry summer.     

“Reversed” intraguild predation: red fox cubs killed by pine marten

Camera traps deployed at a badger Meles meles set in mixed pine forest in north-eastern Poland recorded interspecific killing of red fox Vulpes vulpes cubs by pine marten Martes martes. The vixen and her cubs settled in the set at the beginning of May 2013, and it was abandoned by the badgers shortly afterwards. Five fox cubs were recorded playing in front of the den each night. Ten days after the first recording of the foxes, a pine marten was filmed at the set; it arrived in the morning, made a reconnaissance and returned at night when the vixen was away from the set. The pine marten entered the den several times and killed at least two fox cubs. It was active at the set for about 2 h. This observation proves that red foxes are not completely safe from predation by smaller carnivores, even those considered to be subordinate species in interspecific competition.     

Growth–climate response of Jack pine on clay soils in northeastern Canada

We used tree-ring data from a major North American boreal tree species, Jack pine (Pinus banksiana Lamb.), to decipher spatial and temporal tree-growth responses to climate variability within the area of northwestern Quebec and northeastern Ontario. Fifteen sites with clay soils were selected and grouped into North and South sub-regions at approximately 49°N and 50°N, respectively. Tree-ring chronologies were analyzed through a response function for the years 1951–2000 to identify growth-limiting climate factors. Increased precipitation in June in the previous year and a warm month of April this year favored radial growth whereas higher temperature in September and increased precipitation in October, both of the previous year, and current June precipitation were negatively related to growth. There was a clear difference in climatic response between the southern and northern sub-regions: southern sites were more responsive to temperature dynamics while on northern sites Jack pine growth appeared negatively influenced by an excess of precipitation. Soil conditions, with larger areas covered by less water permeable clay deposits in the northern sub-region, explain this result.If recently observed trends towards warmer springs continue, Jack pine may increase its radial growth in the study area. However, increases in fall precipitation, also predicted under the future climate, may offset the positive effect of previous years weather on clay sites.     

Do exotic pine plantations favour the spread of invasive herbivorous mammals in Patagonia?

Changes in land use patterns and vegetation can trigger ecological change in occupancy and community composition. Among the potential ecological consequences of land use change is altered susceptibility to occupancy by invasive species. We investigated the responses of three introduced mammals (red deer, Cervus elaphus; wild boar, Sus scrofa; and European hare, Lepus europaeus) to replacement of native vegetation by exotic pine plantations in the Patagonian forest‐steppe ecotone using camera‐trap surveys (8633 trap‐days). We used logistic regression models to relate species presence with habitat variables at stand and landscape scales. Red deer and wild boar used pine plantations significantly more frequently than native vegetation. In contrast, occurrence of European hares did not differ between pine plantations and native vegetation, although hares were recorded more frequently in firebreaks than in plantations or native vegetation. Presence of red deer and wild boar was positively associated with cover of pine plantations at the landscape scale, and negatively associated with mid‐storey cover and diversity at the stand scale. European hares preferred sites with low arboreal and mid‐storey cover. Our results suggest that pine plantations promote increased abundances of invasive species whose original distributions are associated with woodlands (red deer and wild boar), and could act as source or pathways for invasive species to new areas.     

Does availability of potassium affect cold hardening of Scots pine through polyamine metabolism?

The effect of different potassium availability on the polyamines and frost resistance of Scots pine seedlings ( Pinus sylvestris L.) during cold hardening was studied. Scots pine seedlings were grown applying different rates of potassium by using the relative addition rate technique followed by a 2- or 9-week hardening period with decreased light intensity, day length and temperature. After 2 weeks of treatment the seedlings were not hardened (LT₅₀, =−11°C) and showed no differences in frost resistance, although differences in the polyamine levels between the K levels were observed. After 9 weeks of hardening the seedlings at the low, medium and high K levels showed a mean frost resistance (LTs.₅₀) of −81, −63, and −47°C, respectively. A negative effect of K on the frost resistance of the needles was also found in adult trees in September. The results indicate that at the early stage of cold hardening, potassium or free polyamine levels do not affect the frost resistance of Scots pine needles. However, in hardened seedlings and adult trees potassium displays a negative and putrescine a positive correlation with frost resistance, whereas spermidine and spermine show no correlation.     

Mid-term successional patterns after fire of mixed pine–oak forests in NE Spain

This study analyzes the factors affecting the current variability in density and age and size structure of mixed pine–oak forests of Pinus nigra and Quercus faginea in Central Catalonia (NE Spain), 37 years after a wildfire. The objective is to determine whether different post-disturbance responses may be obtained from the same pre-fire community and which factors can determine these different potential responses. The two factors analyzed were the distance to the unburned forest and site conditions (represented in this case by different aspects). The response of pines and oaks was different to the pattern expected for the Mediterranean Basin. Oaks resprouted immediately from stools already present before the fire and dominated during the first years, independent of both disturbance and site conditions. Pines established later, and their response depended on both factors: pine density decreased sharply from the forest edge to the burned area, and the number of pines was also higher in the more mesic than in the more xeric conditions. The age structure analysis for pines and oaks in the different aspects also revealed site-dependent rates of succession manifested by initial differences in post-fire establishment. In mesic plots, the establishment of pines occurred quite early, while in xeric plots, pine recruitment was delayed several years. These different patterns of post-fire recovery have led to pine dominance in more mesic sites and codominance of pines and oaks in more xeric ones, suggesting that different mid-term post-fire patterns can be identified for the same pre-fire forest type, depending on variations in environmental conditions.