Grégory Mahy

Toward an old‐growth concept for grasslands, savannas, and woodlands

We expand the concept of “old growth” to encompass the distinct ecologies and conservation values of the worlds ancient grass-dominated biomes. Biologically rich grasslands, savannas, and open-canopy woodlands suffer from an image problem among scientists, policy makers, land managers, and the general public, that fosters alarming rates of ecosystem destruction and degradation. These biomes have for too long been misrepresented as the result of deforestation followed by arrested succession. We now know that grassy biomes originated millions of years ago, long before humans began deforesting. We present a consensus view from diverse geographic regions on the ecological characteristics needed to identify old-growth grasslands and to distinguish them from recently formed anthropogenic vegetation. If widely adopted, the old-growth grassland concept has the potential to improve scientific understanding, conservation policies, and ecosystem management.

Mucilage and polysaccharides in the halophyte plant species Kosteletzkya virginica: Localization and composition in relation to salt stress

Mucilage is thought to play a role in salinity tolerance in certain halophytic species by regulating water ascent and ion transport. The localization and composition of mucilage in the halophyte Kosteletzkya virginica was therefore investigated. Plants were grown in a hydroponic system in the presence or absence of 100mM NaCl and regularly harvested for growth parameter assessment and mucilage analysis with the gas liquid chromatography method. NaCl treatment stimulated shoot growth and biomass accumulation, had little effect on shoot and root water content, and reduced leaf water potential (Psi(w)), osmotic potential (Psi(s)) as well as stomatal conductance (g(s)). Mucilage increased in shoot, stems and roots in response to salt stress. Furthermore, changes were also observed in neutral monosaccharide components. Levels of rhamnose and uronic acid increased with salinity. Staining with a 0.5% alcian blue solution revealed the presence of mucopolyssacharides in xylem vessels and salt-induced mucilaginous precipitates on the leaf abaxial surface. Determination of ion concentrations showed that a significant increase of Na(+) and a decrease of K(+) and Ca(2+) simultaneously occurred in tissues and in mucilage under salt stress. Considering the high proportion of rhamnose and uronic acid in stem mucilage, we suggest that the pectic polysaccharide could be involved in Na(+) fixation, though only a minor fraction of accumulated sodium appeared to be firmly bound to mucilage.

Hybridization and morphogenetic variation in the invasive alien Fallopia (Polygonaceae) complex in Belgium

The invasive alien knotweeds, Fallopia spp. (Polygonaceae), are some of the most troublesome invasive species in Europe and North America. Invasive success in Fallopia may be enhanced by multiple hybridization events. We examined the pattern of hybridization and its evolutionary consequences in Belgium with a concerted analysis of ploidy levels (chromosome counts and flow cytometry), morphological variation, and genetic variation (RAPDs). At least four taxa with different ploidy levels were part of the pattern of invasion in Belgium. Hybrid F. ×bohemica with various chromosome numbers restored the genotypic diversity that was lacking in the parental species. Hybrid genotypes were mainly assigned to a specific genetic pool and not to a mixture between the genetic pools of the putative parental species as would be expected for hybrids. Parental species and hexaploid hybrids differed significantly for a set of well-defined morphological characters, enabling future researchers to distinguish these taxa. On the basis of our results, the importance of hybridization has probably been underestimated in large parts of the adventive range of alien Fallopia species, pointing to the need for concerted molecular and morphological analyses in the study of the evolutionary consequences of hybridization.

Allozyme evidence for genetic autopolyploidy and high genetic diversity in tetraploid cranberry, Vaccinium oxycoccos (Ericaceae).

Polyploidy has been important in the evolution of angiosperms and may significantly affect population genetic diversity and structure. Nineteen isoenzyme loci were studied in diploid and tetraploid populations of Vaccinium oxycoccos (Ericaceae), and the results are compared with data previously reported for the related V. macrocarpon. Diploid V. oxycoccos and V. macrocarpon were readily discriminated based on their allozymic variation. No evidence for fixed heterozygosity was found in tetraploid V. oxycoccos. In contrast, all polymorphic loci exhibited both balanced and unbalanced heterozygotes, with some individuals exhibiting a pattern consistent with the presence of three alleles. These results support an autopolyploid origin for tetraploid V. oxycoccos. However, tetraploid V. oxycoccos possessed a suite of alleles not found in diploid V. oxycoccos; half of these alleles were shared with V. macrocarpon. This suggests that autotetraploid V. oxycoccos may have undergone hybridization with V. macrocarpon or that the autotetraploid retained the genetic variation present in an ancestral diploid species. Following theoretical expectations, proportion of polymorphic loci, mean number of alleles, and observed heterozygosity were significantly higher for the autotetraploid than for the diploid. Mean inbreeding (F(IS)) was similar for diploid and tetraploid V. oxycoccos. The latter exhibited population differentiation (F(ST)) exceeding both diploid species.

Forest refugia revisited: nSSRs and cpDNA sequences support historical isolation in a wide-spread African tree with high colonization capacity, Milicia excelsa (Moraceae).

The impact of the Pleistocene climate oscillations on the structure of biodiversity in tropical regions remains poorly understood. In this study, the forest refuge theory is examined at the molecular level in Milicia excelsa, a dioecious tree with a continuous range throughout tropical Africa. Eight nuclear microsatellites (nSSRs) and two sequences and one microsatellite from chloroplast DNA (cpDNA) showed a deep divide between samples from Benin and those from Lower Guinea. This suggests that these populations were isolated in separate geographical regions, probably for several glacial cycles of the Pleistocene, and that the nuclear gene pools were not homogenized despite M. excelsa’s wind‐pollination syndrome. The divide could also be related to seed dispersal patterns, which should be largely determined by the migration behaviour of M. excelsa’s main seed disperser, the frugivorous bat Eidolon helvum. Within Lower Guinea, a north–south divide, observed with both marker types despite weak genetic structure (nSSRs: FST = 0.035, cpDNA: GST = 0.506), suggested the existence of separate Pleistocene refugia in Cameroon and the Gabon/Congo region. We inferred a pollen‐to‐seed dispersal distance ratio of c. 1.8, consistent with wide‐ranging gene dispersal by both wind and bats. Simulations in an Approximate Bayesian Computation framework suggested low nSSR and cpDNA mutation rates, but imprecise estimates of other demographic parameters, probably due to a substantial gene flow between the Lower Guinean gene pools. The decline of genetic diversity detected in some Gabonese populations could be a consequence of the relatively recent establishment of a closed canopy forest, which could negatively affect M. excelsa’s reproductive system.

Patterns of allozymic variation within Calluna vulgaris populations at seed bank and adult stages

We investigated the spatial genetic structure within and between two plots of Calluna vulgaris and the extent to which the soil seed bank differed genetically from adults at seven allozyme loci. Averaged over the two plots, the seed bank and adult populations contained very similar levels of genetic diversity. Moreover, seeds contained in a single soil core (100 cm3) exhibited similar mean allozyme diversity to the surrounding adult population, indicating that the seed bank preserves genetic diversity at a very local scale. Few differences in allelic frequencies were found between the seed bank and its surrounding adult population in each plot. Mean GST indicated a lack of differentiation between the two plots at adult (GST = 0.008) and seed bank (GST = 0.002) stages. Low interplot differentiation is consistent with the outcrossing mating system of the population (tm = 0.91 in one plot) and its history of human disturbance. In contrast, spatial autocorrelation analysis of adults indicated a genetic structure at a very local scale, with positive autocorrelation for all alleles below 2 m in one plot and with a pattern of positive autocorrelation below 8 m in the two plots. Current limitation to seed dispersal rather than spatial extension of clones is thought to be responsible for local genetic structure.

Tyranny of trees in grassy biomes.

![Figure][1] Highland grassland in Brazil is considered a forest landscape restoration opportunity PHOTO: G. W. FERNANDES Tree planting, fire suppression, and exclusion of megafaunal herbivores (native or domestic) are ecologically reasonable restoration strategies in deforested landscapes

Soil influence on Cu and Co uptake and plant size in the cuprophytes Crepidorhopalon perennis and C. tenuis (Scrophulariaceae) in SC Africa

Cuprophytes are plants that mostly occur on Cu-rich soil in SC Africa. Crepidorhopalon perennis is endemic of a single site. C. tenuis has a broader niche, from normal to Cu-rich soil. Both have been considered as Cu-Co accumulators. We examined soil factors controlling heavy metal accumulation and plant fitness in natural populations. Plant mass and element concentrations in plants and soil were determined in 153 samples from five populations of C. tenuis on copper soil (CTC), two on normal soil (CTN) and the single population of C. perennis (CP). Soil in Cu-sites had higher concentrations of Ca, Mg, P, Mn, Zn, Cu, Co. Plants from Cu-sites were larger and had higher Cu and Co content, and lower Mg, Mn and Ca. Cu in shoots was influenced positively by Cu and Mn and negatively by Ca in the soil. Co in shoots was influenced positively by Co and negatively by Mn and Fe in the soil. Shoot mass was influenced positively by Cu and Mn (CT) or by Cu and Co (both species pooled) in the soil. The results suggest that C. tenuis and C. perennis are genuinely cuprophilous species. Large variation in metal accumulation in shoots can be accounted for by synergistic and antagonistic interactions among several heavy metals, yielding specific accumulation patterns in different populations.

Spatial genetic structure in Milicia excelsa (Moraceae) indicates extensive gene dispersal in a low-density wind-pollinated tropical tree

In this study, we analysed spatial genetic structure (SGS) patterns and estimated dispersal distances in Milicia excelsa (Welw.) C.C. Berg (Moraceae), a threatened wind‐pollinated dioecious African tree, with typically low density (∼10 adults/km2). Eight microsatellite markers were used to type 287 individuals in four Cameroonian populations characterized by different habitats and tree densities. Differentiation among populations was very low. Two populations in more open habitat did not display any correlation between genetic relatedness and spatial distance between individuals, whereas significant SGS was detected in two populations situated under continuous forest cover. SGS was weak with a maximum Sp‐statistic of 0.006, a value in the lower quartile of SGS estimates for trees in the literature. Using a stepwise approach with Bayesian clustering methods, we demonstrated that SGS resulted from isolation by distance and not colonization by different gene pools. Indirect estimates of gene dispersal distances ranged from σg = 1 to 7.1 km, one order of magnitude higher than most estimates found in the literature for tropical tree species. This result can largely be explained by life‐history traits of the species. Milicia excelsa exhibits a potentially wide‐ranging wind‐mediated pollen dispersal mechanism as well as very efficient seed dispersal mediated by large frugivorous bats. Estimations of gene flow suggested no major risk of inbreeding because of reduction in population density by exploitation. Different strategy of seed collection may be required for reforestation programmes among populations with different extent of SGS.

Testing coexistence of extinction debt and colonization credit in fragmented calcareous grasslands with complex historical dynamics

Calcareous grasslands are among the most species-rich ecosystems in temperate countries. However, these ecosystems have suffered from fragmentation and destruction during the last century. We studied the response of calcareous grassland plant diversity to landscape changes in Belgium. Results indicated that high area loss (since 1965) old habitat patches exhibited an extinction debt inverse to low area loss old habitat patches, little depending on the area loss threshold (60%, 70%, 80% or 90%) considered for the distinction between the high and low area loss patches. However, human activities also created new habitat patches in the landscape and therefore provided opportunities for calcareous grassland plant species to colonize new habitats. This also provided opportunities to study species colonization abilities in the context of habitat restoration. We analyzed species richness in new patches compared to old patches in order to detect colonization credit. We detected the presence of a colonization credit in new patches when using high loss old patches (area loss >80%, exhibiting an extinction debt) or all old patches as a reference. However, when the reference was low loss old patches alone (area loss <80%, less likely to exhibit an extinction debt), no colonization credit was detected. In addition, species composition was similar between new patches and old patches. These results are encouraging for restoration programs. However, the results indicated that the presence of an extinction debt in reference habitats could lead to inaccurate conclusions in restoration monitoring. Therefore, extinction debt should be considered when choosing reference habitats to evaluate restoration success.