Understanding the factors controlling the expression of longevity in trees is still an outstanding challenge for tree biologists and forest ecologists. We gathered tree-ring data and literature for broadleaf deciduous (BD) temperate trees growing in closed-canopy old-growth forests in the Northern Hemisphere to explore the role of geographic patterns, climate variability, and growth rates on longevity. Our pan-continental analysis, covering 32 species from 12 genera, showed that 300-400 years can be considered a baseline threshold for maximum tree lifespan in many temperate deciduous forests. Maximum age varies greatly in relation to environmental features, even within the same species. Tree longevity is generally promoted by reduced growth rates across large genetic differences and environmental gradients. We argue that slower growth rates, and the associated smaller size, provide trees with an advantage against biotic and abiotic disturbance agents, supporting the idea that size, not age, is the main constraint to tree longevity. The oldest trees were living most of their life in subordinate canopy conditions and/or within primary forests in cool temperate environments and outside major storm tracks. Very old trees are thus characterized by slow growth and often live in forests with harsh site conditions and infrequent disturbance events that kill much of the trees. Temperature inversely controls the expression of longevity in mesophilous species (Fagus spp.), but its role in Quercus spp. is more complex and warrants further research in disturbance ecology. Biological, ecological and historical drivers must be considered to understand the constraints imposed to longevity within different forest landscapes.

Old-growth mountain forests represent an ideal setting for studying long-term impacts of climate change. We studied the few remnants of old-growth forests located within the Pollino massif (southern Italy) to evaluate how the growth of conspecific young and old trees responded to climate change. We investigated two conifer species (Abies alba and Pinus leucodermis) and two hardwood species (Fagus sylvatica and Quercus cerris). We sampled one stand per species along an altitudinal gradient, ranging from a drought-limited low-elevation hardwood forest to a cold-limited subalpine pine forest. We used a dendrochronological approach to characterize the long-term growth dynamics of old (age > 120 years) versus young (age < 120 years) trees. Younger trees grew faster than their older conspecifics during their juvenile stage, regardless of species. Linear mixed effect models were used to quantify recent growth trends (1950-2015) and responses to climate for old and young trees. Climate sensitivity, expressed as radial growth responses to climate during the last three decades, partially differed between species because high spring temperatures enhanced conifer growth, whereas F. sylvatica growth was negatively affected by warmer spring conditions. Furthermore, tree growth was negatively impacted by summer drought in all species. Climate sensitivity differed between young and old trees, with younger trees tending to be more sensitive in P. leucodermis and A. alba, whereas older F. sylvatica trees were more sensitive. In low-elevation Q. cerris stands, limitation of growth due to drought was not related to tree age, suggesting symmetric water competition. We found evidence for a fast-growth trend in young individuals compared with that in their older conspecifics. Notably, old trees tended to have relatively stable growth rates, showing remarkable resistance to climate warming. These responses to climate change should be recognized when forecasting the future dynamics of old-growth forests for their sustainable management.

In a network of six old-growth and three managed beech forests, distributed over a wide altitudinal range in the Central Apennines, we used principal component analysis to analyse a set of 15 commonly used structural and chronological indicators of old-growthness, related to both live trees and deadwood. Our goals were to (1) quantify the effectiveness of such indicators for ranking forests according to their degree of old-growthness and (2) disentangle the effects of site-related (e.g. elevation) and human-related (e.g. past management) factors on their variation. We selected eight indicators as best descriptors of forest old-growthness: mean age of the five oldest trees; snag volume; coarse woody debris (CWD) volume; density of medium-size snags [diameter at breast height (DBH) ≥ 47.5 cm]; forest stature; basal area; established regeneration and density of large trees (DBH ≥ 67.5 cm). We analysed their relative importance in informing us about the degree of old-growthness in our forests. Our findings suggest that several old-growthness indicators are to be considered carefully, because they are strongly linked to site conditions and particularly to the biogeoclimatic context. Moreover, we provide evidence for how past management can produce negative effects on some indicators.

In the Mediterranean basin, agriculture and other forms of human land use have shaped the environment since ancient times. Intensive and extensive agricultural systems managed with a few cultured plant populations of improved varieties are a widespread reality in many Mediterranean countries. Despite this, historical cultural landscapes still exist in interior and less intensively managed rural areas. There, ancient fruit tree varieties have survived modern cultivation systems, preserving a unique genetic heritage. In this study, we mapped and characterized 106 living fruit trees of ancient varieties in the Basilicata region of southern Italy. Tree ages were determined through tree ring measurements and radiocarbon analyses. We uncovered some of the oldest scientifically dated fruit trees in the world. The oldest fruit species were olive (max age 680 ± 57 years), mulberry (647 ± 66 years), chestnut (636 ± 66 years), and pear (467 ± 89 years). These patriarchs hold a unique genetic resource; their preservation and genetic maintenance through agamic propagation are now promoted by the Lucan Agency for the Development and Innovation in Agriculture (ALSIA). Each tree also represents a hub for biodiversity conservation in agrarian ecosystems: their large architecture and time persistence guarantee ecological niches and micro-habitats suitable for flora and fauna species of conservation significance.