Disentangling the relative importance of climate,size and competition on tree growth in Iberian forests: implications for forest management under global change

Most large‐scale multispecies studies of tree growth have been conducted in tropical and cool temperate forests, whereas Mediterranean water‐limited ecosystems have received much less attention. This limits our understanding of how growth of coexisting tree species varies along environmental gradients in these forests, and the implications for species interactions and community assembly under current and future climatic conditions. Here, we quantify the absolute effect and relative importance of climate, tree size and competition as determinants of tree growth patterns in Iberian forests, and explore interspecific differences in the two components of competitive ability (competitive response and effect) along climatic and size gradients. Spatially explicit neighborhood models were developed to predict tree growth for the 15 most abundant Iberian tree species using permanent‐plot data from the Spanish Second and Third National Forest Inventory (IFN). Our neighborhood analyses showed a climatic and size effect on tree growth, but also revealed that competition from neighbors has a comparatively much larger impact on growth in Iberian forests. Moreover, the sensitivity to competition (i.e. competitive response) of target trees varied markedly along climatic gradients causing significant rank reversals in species performance, particularly under xeric conditions. We also found compelling evidence for strong species‐specific competitive effects in these forests. Altogether, these results constitute critical new information which not only furthers our understanding of important theoretical questions about the assembly of Mediterranean forests, but will also be of help in developing new guidelines for adapting forests in this climatic boundary to global change. If we consider the climatic gradients of this study as a surrogate for future climatic conditions, then we should expect absolute growth rates to decrease and sensitivity to competition to increase in most forests of the Iberian Peninsula (in all but the northern Atlantic forests), making these management considerations even more important in the future.     

Spatiotemporal mating pattern variation in a wind‐pollinated Mediterranean shrub

Spatiotemporal variation in mating patterns is poorly known in wind‐pollinated plant species. Here, we analysed mating patterns of the wind‐pollinated dioecious shrub Pistacia lentiscus by genotyping 904 seeds from 30 mother plants with eight microsatellite markers in a high‐density population in two consecutive flowering seasons. We found significant differences in some mating system estimates between years, particularly in the levels of correlated paternity. Overall, within‐mothers correlated paternity was higher in 2007 than in 2006 (r_pWM = 0.085 and 0.030), which translated into an effective number of fathers (N_ep) of 11.8 and 33.6 respectively. Using a smoothing interpolation technique, we show that the effective pollen cloud was spatially structured in patches of high‐ and low‐genetic diversity, which do not remain constant from year to year. In 2006, the among‐mothers correlated paternity (r_pAM) showed no trend with distance, suggesting no restriction of pollen dispersal. However, in 2007, r_pAM was greater than zero at short distances, revealing the existence of small‐scale patterns of pollen dispersal. The fact that the studied seasons were climatically homogeneous during the flowering time suggested that the observed differences might be ascribed to between‐year phenological variation of individuals in the studied population or other (unknown) factors. Numerical simulations, based on the real data set, indicated that the clumping of males and decreasing plant density, which is related to different types of pollen limitation, greatly increase correlated mating in this wind‐pollinated species, which is of relevance under the frame of the continuous anthropogenic habitat disturbance suffered by Mediterranean ecosystems.     

Mechanisms of colony selection by first‐year Lesser Kestrels Falco naumanni

In colonial species, first‐time breeders may use the number of settled conspecifics in colony selection, but such a relationship is confused by the correlation between colony size and nest‐site availability. To distinguish conspecific attraction from neutral colony selection, we experimentally increased nest availability for first‐year Lesser Kestrels Falco naumanni, allowing us to dissociate the number of vacant nest‐sites from colony size at the arrival time of first‐year birds. Under natural conditions, the number of first‐year birds settling was positively correlated with both the number of philopatric and the total number of breeding pairs (colony size) already settled. However, the probability of occupation of experimentally manipulated nests by first‐year birds was independent of colony size. In experimental colonies, the number of first‐year birds settling was positively correlated with the number of manipulated nest‐sites but not with the number of conspecifics. Overall, these results support a neutral colony selection by first‐year Lesser Kestrels based on nest‐site availability.     

Fitness components of the hybrid Phlomis x margaritae Aparicio & Silvestre (Lamiaceae)

Studies of microsporogenesis, pollen germination, pollen stainability, fruit and seed set and ovary predation have been undertaken to evaluate the fitness of the hybrid genotype of Phlomis x margaritae relative to its parents (P. purpurea and P. x composita) and allied taxa (P. x composita nothom. composita, P. x composita nothom. trulknqueiz , and P. lychnitis). Phlomis x margaritae shows irregular meiosis with very low pollen germination rate and very high pollen sterility, no seed set, and very high ovary predation. It thus has very low reproductive success and fitness. Phlomis x composita nothom. trullenquei also shows low pollen germination and seed set, but these follow regular microsporogenesis. The existence of cryptic structural sterility is suggested for this taxon. The rest of the studied taxa (P. purpurea, P. lychnitis and P. x composita nothom. composita) were fertile and can support the specific or subspecific taxonomic status of P. x composita. A fixed ovule abortion pattern in the taxa with yellow flowers was found, since 'brood size' consisted just of one seed per fruit. On the contrary, P. purpurea was found to produce from one to four seeds per fruit. Some conservation implications are suggested.     

Genetic variation of early height growth traits at the xeric limits of Austrocedrus chilensis (Cupressaceae)

The study of the genetic variation of early height growth traits in seedlings helps to predict the possible outcomes of tree populations in the face of climate change. Second‐year height growth of 10 geographically marginal populations of Patagonian cypress (Austrocedrus chilensis (D. Don) Pic. Ser. et Bizzarri) (Cupressaceae) was characterized under greenhouse conditions. Variation among and within an average of 15 open‐pollinated families (comprising 21 seedlings per family) for each population was analysed for six size and timing traits obtained from fitted Boltzmann growth curves. The among‐family and among‐population variances were 4.03% and 2.74% of the total phenotypic variation, while the residual variance was 84.57% on average. Genetic differentiation among populations was low, except for the maximum growth rate (Q_ST = 0.35) and for growth initiation (Q_ST = 1). For most traits, genetic variation and heritability were variable across populations, except for growth initiation, which showed in general null intra‐population levels of genetic variance. Although no direct associations were found between the additive genetic variation and latitude or altitude, the north range of the distribution was more variable for the pool of the analysed traits. Although most extreme‐marginal populations of A. chilensis would be very limited in their ability to evolve if climate in north‐west Patagonia turns drier and warmer, their long‐term persistence could largely rely on a phenotypic diversification strategy.     

Temporal variation of heterozygosity‐based assortative mating and related benefits in a lesser kestrel population

Heterozygosity as a target of mate choice has received much attention in recent years and there is growing evidence supporting its role in the evolution of mate preferences. In this study we analyse mating patterns in relation to heterozygosity in a lesser kestrel (Falco naumanni) population intensively monitored over six study years (2002–2007). The magnitude of heterozygosity‐based assortative mating varied over time, being particularly patent in the last study years (2006, 2007). We have found evidence that this mating pattern entails both direct and indirect‐genetic benefits. Clutch size increased with female heterozygosity and more heterozygous males raised a higher number of fledglings particularly in those years when the strength of the heterozygosity‐based assortative mating was markedly higher. In the last study year, parent–offspring correlation of heterozygosity was stronger and higher than the expected if individuals would have randomly mated with respect to heterozygosity. Overall, our results offer empirical support to the heterozygous mate hypothesis of sexual selection but suggest that genetic diversity may act as a temporally variable target for mate choice.     

Isozyme Evidence for Natural Hybridization in Phlomis (Lamiaceae): Hybrid Origin of the RareP.xmargaritae

Phlomisxmargaritae is a rare and sterile hybrid taxon foundin a single endangered population in southern Spain. It waspreviously described as a morphologically intermediate hybrid,putatively between Phlomis purpurea and P. composita. The presentstudy used allozymes as molecular markers to assess the hybrididentity ofP.xmargaritae . Ten putative loci from seven enzymesystems were resolved: five were monomorphic and fixed acrossall taxa studied and the rest (Aat-1, Aat-2, 6-Pgdh-2, Pgi andPgm) were polymorphic in at least one taxon. The two parentaltaxa are fixed for different alleles at 6-Pgdh-2 and show distinctallelic frequency differences for four other loci. Phlomisxmargaritaedisplays fixed heterozygous phenotypes for four of the fivepolymorphic loci, these being composed of combinations of thealleles found in the parental taxa. No unique alleles were detectedin P.xmargaritae. We conclude that this taxon is of hybrid originand confirm the identity of the parental taxa involved. It isfurther suggested that this population is composed of individualsthat are recent F₁hybrids that have not undergone backcrossingor introgression. Global conservation measures are necessaryfor the whole hybrid system in this location since further continuousassessment could reveal the evolutionary input of hybridizationin Phlomis.Copyright 2000 Annals of Botany Company Endangered species, genetic diversity, isozyme variation, hybrid species, Phlomisxmargaritae, plant conservation, population genetics, Lamiaceae, Spain.     

Variation patterns in the Phlomis × composita (Lamiaceae) hybrid complex in the Iberian Peninsula

Hybridization seems to be common in the genus Phlomis (Lamiaceae) in the Iberian Peninsula, especially in the P.  ×  composita complex. In order to detect patterns of morphological variation linked to eco- and geographical variation, ecological (cluster and canonical correspondence) and morphometric (principal component and discriminant function) analyses were performed. Character count procedure was applied to discern between divergence and reticulate events for the origin of the morphologically intermediate plants. Following these analyses clear patterns were detected suggesting the existence of four independent morphological groups also supported by the ecogeographical data. These are P. crinita ssp. crinita (Levante, eastern Spain), P. crinita ssp. malacitana (Andalusia, southern Spain) and P. lychnitis (widespread) being the extremes of the morphological variation, and P.  ×  composita including all the morphologically intermediate individual plants. Furthermore, at the population level significant differences in hybrid plant frequencies between areas were also found.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 97–108.