The seasonal dynamic s of cambial activity, wood formation , occurrence of false rings (FR), and tangential bands of resin canals (RC) were investigated in Pinus halepensis from three Mediterranean dry and semi-arid ecosystems in Spain. We collected intact tissues of phloem, cambium, and outer xylem at monthly intervals throughout 2004 from each time six trees at the three sites. Cell divisions in the cambium in all trees started before our first sampling in mid-March and ceased between November and December. Cambial activity was characterized by two maxima; one in spring and another in autumn. Trees still grew in summer but at a very low rate. The first solitary RC were formed in May or June and tangential bands ofRC in June or July. In general , tangential bands ofRC were observed in wider growth rings . The formation of tangential bands of RC seems to be induced by drought in the second half of the growing season . FR were formed as a result of cambial reactivation in autumn and were observed in growth rings that contained more than 50 cells in a radial row.It appears that summer drought and early autumn precipitation play an important role in false-ring formation.
Recent studies on the seasonal dynamics of secondary tissue formation in Mediterranean trees have shown that xylogenesis depends on species and site conditions, but many questions still remain open. On the other side of the cambium, even less information is available about phloem structure and timing of its formation. We analysed intra-annual phloem variation in width and cell traits in the conducting, non-collapsed phloem (CPH) of Pinus pinea and Pinus halepensis at Mediterranean sites in southern Italy and Spain. In all investigated trees, it was possible to differentiate among the non-conducting, collapsed phloem (NCPH), and the CPH. CPH showed no evident annual growth layers; no differences in radial dimensions of early- and late phloem sieve cells, and no cyclic patterns of axial parenchyma distribution. Since it was not possible to study the seasonality of the phloem growth, we analysed the entire CPH. CPH width showed seasonal fluctuations and was generally the widest during the maximum cambial activity and narrowest during summer and winter. The radial size of newly formed sieve cells varied in relation to seasonal dynamics of cambial activity and fluctuations in local weather conditions. The number of axial parenchyma cells in CPH increased during the summer. The observed intra-annual variations in CPH width and structure seemed to be correlated with seasonal weather conditions in order to ensure a sufficient amount of conducting phloem tissue for translocation of photosynthates and signalling molecules to the actively growing tissues along the stem of a tree growing in the harsh Mediterranean conditions.
Annual periodicity of cambium production of xylem and phloem cells has rarely been compared in trees from different environments. We compared the structure of cambium and the youngest xylem and phloem increments in four tree species, Fagus sylvatica, Picea abies, Pinus sylvestris and Pinus halepensis, from nine temperate and Mediterranean sites in Slovenia and Spain. In Picea abies, Pinus sylvestris and Fagus sylvatica from temperate locations in Slovenia, xylem and phloem growth ring boundaries could be identified. In Fagus sylvatica growing at two elevations on Moncayo mountain, Spain, phloem increment consisted of only early phloem. In Pinus sylvestris from the same two sites, growth ring boundaries were not as clear as in temperate Slovenian sites. In some cases we could identify phloem growth ring boundaries but in others it was very doubtful, which could be explained by collapse of the outermost early phloem sieve cells. In Pinus halepensis from all sites, we could only distinguish between collapsed and non-collapsed phloem, while phloem rings could not be identified. Widths of the youngest phloem and xylem annual increments could only be compared when phloem increments could be clearly defined, as with Picea abies, Fagus sylvatica and Pinus sylvestris from temperate sites. The visibility of the growth ring boundary in phloem was not related to the width of annual radial growth. The correlation between xylem and phloem ring widths was high, but moderate between the number of dormant cambial cells and xylem ring and phloem ring widths. Based on the structure of the youngest phloem increments, we concluded that there is no typical annual periodicity in cambial production of phloem cells in trees from certain Mediterranean sites. This may be due to continuous yearlong cell production and the absence of true cambium dormancy, at least on the phloem side, under mild winter conditions.
Intra-annual density fluctuations (IADFs) in tree rings of Aleppo pine (Pinus halepensis) are considered to be among the most promising wood anatomical features in dendrochronological studies. They provide environmental information in addition to those obtained from tree-ring widths. We used a network of 35 sites in Spain, ranging from nearly desert to temperate climate. We analysed tree-ring series of 529 trees to study IADF frequencies, and their dependence on climatic factors and cambial age. The results showed that IADF frequency is age dependent, with its maximum at the cambial age of 27 years (evaluated at breast height). The frequencies varied across the network and at different sites we recorded that 0.3% to 33% of the analysed tree rings contained IADFs. They were more frequent where and when the temperatures were higher, summer drought was intense and autumn was the main precipitation season. IADF formation was particularly related to high minimum temperatures and wet conditions in late summer and autumn. These results suggest that IADF formation is not related to stressful conditions during summer but to favourable conditions during autumn. These conditions promote cambial reactivation and consequently formation of wider tree rings.
Recent re-excavation of Mumba Rockshelter unearthed an unbiased lithic sample from Bed V. Technological analysis has permitted a reinterpretation of the so-called Mumba Industry, a transitional industry between Middle and Later Stone Ages originally defined by Mehlman (1989). Our data confirm Mehlman’s observation that the “evolutionary” markers in Mumba Bed V are basically typological. However, our study differs from his in that we classify all of Bed V as LSA based on the combined analyses of typology and technology in our excavated assemblage. From a technological perspective, no changes have been observed throughout the sequence, and continuity is the main technological characteristic of the series. The only transitional marker from Lower through Upper Bed V is the appearance of the geometric crescent in the latter, taking into account that microliths exist throughout the sequence. This evidence casts some doubts on previous interpretations and underscores the need to recover a larger sample using modern excavation techniques. It also stresses the need to define the MSA/LSA transition in better terms, combining techno-typological criteria.
The responses of the vascular cambium and tracheid differentiation to extreme drought in Aleppo pine (Pinus halepensis Mill.) were investigated. The research focused on the drought year of 2005, in the primary study area at Maigmo (MAI) in southeastern Spain, with comparisons in Jarafuel (JAL) and Guardamar (GUA). The climate in this region is typically warm and dry with hot summers. Wood formation throughout the 2005 growing season was studied in transverse microtome sections and integrated with a retrospective dendrochronological analysis of crossdated increment cores collected in 2009. For most anatomical sections collected throughout the growing season at MAI, the vascular cambium appeared to be dormant as indicated by the low number of cells per radial file. Occasionally, immature xylem derivatives were observed during the growing season but without production of an annual ring. In increment cores collected at MAI, the 2005 position in the annual ring series contained either a narrow ring of both earlywood and latewood (47% of samples), a narrow ring of apparent latewood with no earlywood (13%), or a missing ring (50%). We introduce the term “dark ring” to refer to those annual rings of apparent latewood with no earlywood. For trees at JAL, the 2005 ring had below-average width and contained both earlywood and latewood. At GUA, the trees produced the widest 2005 ring of all three sites and mainly contained an intra-annual density fluctuation (IADF). The IADF was formed after cambial reactivation in the autumn. Although dark rings, IADFs, and especially missing rings complicate dendrochronological analysis, these anatomical features may provide an additional proxy record from which to infer climate variability and change in the past.