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1.

Question

Coastal environments have often been described as azonal. While this characteristic is clear for the foredune system, it seems less evident for more inland fixed dunes, which host habitats of major conservation concern, whose features seem to be more related to local climatic conditions. We hypothesized that, unlike other coastal habitats, dune perennial grasslands differ floristically and structurally across their European range and that patterns of variation are linked to the corresponding climate.

Location

European coasts (Atlantic Ocean, Baltic, Mediterranean, Black Sea).

Methods

We used a large data set of phytosociological relevés, representative of coastal grasslands throughout their European range. The role of climatic variables (temperature, precipitation and continentality) in determining the variability in species composition and vegetation structure (by means of life forms) was investigated through CCA, DCA and GLM. The degree of concentration of species occurrences within groups was calculated through the Phi coefficient.

Results

Through multivariate analyses we identified seven major types of coastal grassland, corresponding to different geographic areas. The groups significantly differed in their climatic envelope, as well as in their species composition and community structure.

Conclusion

Our results confirm the hypothesis that coastal dune perennial grasslands are subjected to local climate, which exerts significant effects on both floristic composition and community structure. As a consequence, coastal grasslands are particularly prone to the effect of possible climate change, which may alter species composition and distribution, and lead to shifts in the distribution of native plant communities.
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2.

Premise

Plant lineages differ markedly in species richness globally, regionally, and locally. Differences in whole-genome characteristics (WGCs) such as monoploid chromosome number, genome size, and ploidy level may explain differences in global species richness through speciation or global extinction. However, it is unknown whether WGCs drive species richness within lineages also in a recent, postglacial regional flora or in local plant communities through local extinction or colonization and regional species turnover.

Methods

We tested for relationships between WGCs and richness of angiosperm families across the Netherlands/Germany/Czechia as a region, and within 193,449 local vegetation plots.

Results

Families that are species-rich across the region have lower ploidy levels and small monoploid chromosomes numbers or both (interaction terms), but the relationships disappear after accounting for continental and local richness of families. Families that are species-rich within occupied localities have small numbers of polyploidy and monoploid chromosome numbers or both, independent of their own regional richness and the local richness of all other locally co-occurring species in the plots. Relationships between WGCs and family species-richness persisted after accounting for niche characteristics and life histories.

Conclusions

Families that have few chromosomes, either monoploid or holoploid, succeed in maintaining many species in local communities and across a continent and, as indirect consequence of both, across a region. We suggest evolutionary mechanisms to explain how small chromosome numbers and ploidy levels might decrease rates of local extinction and increase rates of colonization. The genome of a macroevolutionary lineage may ultimately control whether its species can ecologically coexist.
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3.

Premise

Habitat fragmentation negatively affects population size and mating patterns that directly affect progeny fitness and genetic diversity; however, little is known about the effects of habitat fragmentation on dioecious, wind pollinated trees. We assessed the effects of habitat fragmentation on population sex ratios, genetic diversity, gene flow, mating patterns, and early progeny vigor in the tropical dioecious tree, Brosimum alicastrum.

Methods

We conducted our study in three continuous and three fragmented forest sites in a Mexican tropical dry forest. We used eight microsatellite loci to characterize the genetic diversity, gene flow via pollen distances, and mean relatedness of progeny. We compared early progeny vigor parameters of seedlings growing under greenhouse conditions.

Results

Sex ratios did not deviate from 1:1 between habitat conditions except for one population in a fragmented habitat, which was female biased. The genetic diversity of adult trees and their offspring was similar in both habitat conditions. Pollen gene flow distances were similar across habitat types; however, paternity correlations were greater in fragmented than in continuous habitats. Germination rates did not differ between habitat conditions; however, progeny from fragmented habitats produced fewer leaves and had a lower foliar area, total height, and total dry biomass than progeny from continuous habitats.

Conclusions

Changes in mating patterns because of habitat fragmentation have negative effects on early progeny vigor. We conclude that negative habitat fragmentation effects on mating patterns and early progeny vigor may be a serious threat to the long-term persistence of tropical dioecious trees.
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4.

Aim

We present the first continental‐scale study of factors controlling the species richness of groundwater‐fed fens, comparing land snails, vascular plants and bryophytes. We separately analyse two ecologically distinct groups differing in conservation value and colonization/extinction dynamics, that is habitat specialists, and matrix‐derived species. Considering the island‐like nature of fen habitats, we hypothesize larger differences in the species richness–environment relationships between habitat specialists and matrix‐derived species than among the taxonomic entities.

Location

Seven European regions

Methods

Richness was counted at 373 well‐preserved fens with undisturbed hydrology using the same protocols. Relationships between the species richness and water pH, waterlogging, climate and geography were explored by GLMs.

Results

Land snail richness responded mainly to water pH, regardless of habitat specialization. Richness of vascular plant and bryophyte specialists was strongly driven by geographical location of the sites, while that of matrix‐derived species was driven by waterlogging and water pH. The richness of matrix‐derived species of all taxa significantly increased with the decreasing waterlogging. Residual richness of specialists of all taxa decreased towards southern Europe.

Main conclusions

In island‐like terrestrial habitats, differences between specialists and matrix‐derived species may outweigh differences among taxa, unless there is one strong physiological determinant of species richness such as pH in land snails. The richness of specialists seems to be strongly related to difficult‐to‐measure regional factors such as historical frequency and connectivity of fen habitats. The richness of matrix‐derived species depends mainly on local conditions, such as pH and waterlogging, determining the degree of habitat contrast against the surrounding matrix. Sufficient waterlogging maintains a high representation of habitat specialists in fen communities, and disturbance of water regime may cause the increase in the number of matrix‐derived species and potentially trigger successional shifts towards non‐fen communities.
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5.

Questions

Which major syntaxa of dry grasslands supported by carbonate bedrock occur in the central and southern Balkans? What is their position along major ecological gradients and in the context of phytogeographic patterns of the region?

Location

Central and southern Balkans, including western Bulgaria, northern Greece, Kosovo, Macedonia (FYROM) and Serbia.

Methods

We compiled a matrix of 660 relevés of dry grasslands over lime‐rich bedrock, previously classified in the Festuco‐Brometea. We applied clustering techniques to classify separately synoptic and relevé data, and applied NMDS with passive projection of indicator values, climatic data and biogeographic geo‐elements onto ordination diagrams to assist interpretation of the syntaxonomic patterns. We constructed elevation distribution profiles for alliances and classes of grasslands of several grassland classes from a broader study area to elucidate the relationship of the elevational sorting of the syntaxa in relation to latitude.

Results

The analysis revealed six major vegetation types, classified into four orders: (1) Stipo pulcherrimae‐Festucetalia pallentis, incl. (sub)montane rocky steppic grasslands of the Saturejion montanae of central Balkans, and the Koelerio‐Festucion dalmaticae – submontane rocky grasslands of southern Serbia and Kosovo; (2) Astragalo onobrychidis‐Potentilletalia represented by the Saturejo‐Thymion (low‐elevation steppic grasslands of southern Balkans); (3) Festucetalia valesiacae represented by grasslands on deep soil and low elevation of northern Greece, and finally (4) high‐elevation rocky grasslands of southern Balkans, classified as a new alliance – Diantho haematocalycis‐Festucion hirtovaginatae, that might belong to a new, yet undescribed, syntaxonomic order. Ordination suggests that the major differentiation of the high‐rank syntaxa follows north–south geographic and low–high elevation gradients.

Conclusions

Because of the transitional biogeographic position of the studied region, as well as considerable large elevation span across latitudes, the diversity of vegetation types is high. The indication a putative new dry grassland order, the mid‐high altitudes of the southern Balkans points to a need to re‐assess the Balkan vegetation occupying the community niche between the low‐elevation dry grasslands (Festuco‐Brometea) and those typical of high elevations (Elyno‐Seslerietea and Daphno‐Festucetea), seeking parallels to patterns described from the western Alps, Pyrenees, and Apennines. This syntaxonomic unit is poised to expand the concept of the Festuco hystricis‐Ononidetea striatae to the Balkans.
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6.
7.

Premise

Reproductive fitness in plants is often determined by the quantity and quality of pollen transferred by pollinators. However, many fitness studies measure only female fitness or rely on proxies for male fitness. Here we assessed how five bee taxon groups affect male fitness in a prairie plant by quantifying pollen removal, visitation, and siring success using paternity assignments and a unique pollinator visitation experiment.

Methods

In Echinacea angustifolia, we measured per-visit pollen removal for each pollinator taxon and estimated the number of pollen grains needed for successful ovule fertilization. Additionally, we directly measured pollinator influence on siring by allowing only one bee taxon to visit each pollen-donor plant, while open-pollinated plants acted as unrestricted pollen recipients. We genotyped the resulting offspring, assigned paternity, and used aster statistical models to quantify siring success.

Results

Siring success of pollen-donor plants differed among the five pollinator groups. Nongrooming male bees were associated with increased siring success. Bees from all taxa removed most of the flowering head's pollen in one visit. However, coneflower-specialist bee Andrena helianthiformis removed the most pollen per visit. Female fitness and proxy measures of male fitness, such as pollinator visitation and pollen removal, did not align with our direct quantifications of male fitness.

Conclusions

Our results illustrate the need for more studies to directly quantify male fitness, and we caution against using male fitness proxy measures. In addition, conservation efforts that preserve a diverse pollinator community can benefit plants in fragmented landscapes.
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8.

Premise

Understanding establishment and spread of non-native plants is important in the face of a homogenizing global flora. While many studies focus on successful, invasive species, fewer have studied failed plant introductions. Until the early 1900s, large quantities of ship ballast, often containing foreign plant propagules, were deposited in New Jersey (USA). The resulting ballast flora is documented in extensive herbarium records, providing us a unique opportunity to analyze successes and failures of novel plant species introductions.

Methods

We used digitized specimens from 75 herbaria to study 264 non-native species introduced into New Jersey through 19th century ballast deposition. We used spatial (density-based clustering; HDBSCAN) and temporal analyses of species retention and geographic spread to quantify disappearance rate, survival, and dispersion through time and define trajectory groups.

Results

Four distinct trajectory groups were identified: waif (only present during import; 32% of species), short-term (disappeared quickly; 20%), established–limited spread (survives locally, 30%), and established–widespread (widespread, 18%). Species disappearance rate was highest during ballast deposition and decreased soon after deposition stopped around 1900. Spatial patterns showed a strong association with 19th century railroads for inland dispersal from ports. The disappearance rate and spatial analyses are robust to herbarium collection bias.

Conclusions

This study using New Jersey as a model is one of the few documenting multispecies successes and failures in inadvertent plant introductions. Results reveal distinct trends in species establishment and geographic spread and highlight the utility of herbarium specimens in answering questions that span large time scales.
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9.

Premise

While angiosperms respond uniformly to abscisic acid (ABA) by stomatal closure, the response of ferns to ABA is ambiguous. We evaluated the effect of endogenous ABA, hydrogen peroxide (H2O2), nitric oxide (NO), and Ca2+, low and high light intensities, and blue light (BL) on stomatal opening of Pleopeltis polypodioides.

Methods

Endogenous ABA was quantified using gas chromatography-mass spectrometry; microscopy results and stomatal responses to light and chemical treatments were analyzed with Image J.

Results

The ABA content increases during initial dehydration, peaks at 15 h and then decreases to one fourth of the ABA content of hydrated fronds. Following rehydration, ABA content increases within 24 h to the level of hydrated tissue. The stomatal aperture opens under BL and remains open even in the presence of ABA. Closure was strongly affected by BL, NO, and Ca2+, regardless of ABA, H2O2 effect was weak.

Conclusions

The decrease in the ABA content during extended dehydration and insensitivity of the stomata to ABA suggests that the drought tolerance mechanism of Pleopeltis polypodioides is independent of ABA.
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10.

Question

What are the composition and spatial patterns of native woody plant communities in the southern Great Chaco and Espinal?

Location

Córdoba Province, central Argentina, an area of ca. 161,000 km2.

Methods

We collected 351 geo‐referenced relevés representative of the geographic, topographic and ecological variation of the Chaco and Espinal woody vegetation in central Argentina. The relevés were classified into vegetation types using the hierarchical ISOPAM method. Forest and shrubland types were described on the basis of diagnostic species occurrences and their distribution in relation to environmental factors. A map of the actual vegetation derived from remote‐sensed images (Landsat) and field data was used to describe the current distribution and abundance of the different vegetation types.

Results

The classification of the 351 plots × 837 species matrix revealed two major clusters comprising seven woody vegetation types corresponding to Chaco lowland and mountain forests and shrublands, Espinal forests and edaphic vegetation. The most important gradients in woody vegetation types are related to elevation, temperature and rainfall variables.

Conclusions

Subtropical seasonally dry woody plant communities from the southern extreme of the Great Chaco and Espinal forests were described for the first time based on complete floristic data. Our results show that lowland Chaco native forests, as well as replacement communities, are still present in its southern distribution range and are well distinguishable from other vegetation types such as the Espinal and mountain forests. Overall, extensive Espinal forests have almost disappeared while Chaco vegetation is highly fragmented and degraded.
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11.

Objectives

Food sharing is a costly form of cooperation that was likely critical to human evolutionary success, including the emergence of human's life history strategy. Food sharing in human communities may be maintained through a number of pathways, including direct dyadic reciprocity, reputation-based processes, and kin-biased exchange. Differences in reproductive demands, labor, and cultural norms may also result in gendered differences in cooperative networks. Here, we examine cooperative networks in egalitarian BaYaka foragers from the Congo Basin.

Materials and Methods

We collected social network data from 112 adults in 41 households in this subsistence community. We implement a Bayesian latent network model to assess individual-, dyadic-, and block-level predictors of food sharing partners.

Results

Conditioning on covariates, we found limited evidence for direct dyadic reciprocity in food sharing. Despite local norms regarding prestige avoidance, we found status-based homophily. High-status individuals—council members and local healers—were more likely to share with one another. Importantly, our results highlight gender differences in patterns of food sharing, interacting with genetic relatedness. Women were more likely to share with one another, especially with kin as genetic relatedness increased.

Discussion

Our results align with evolutionary framing emphasizing kin selection in costly cooperation. The results showing that women cooperate with other women, particularly kin, also complement sex-based patterns in some other mammalian species, potentially reflecting the social support necessary to manage reproductive costs and childcare. BaYaka women's subsistence productivity and local cultural dynamics for autonomy and egalitarianism may likewise help facilitate women's preferential cooperation with one another.
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12.

Premise

The Amazonian hyperdominant genus Eperua (Fabaceae) currently holds 20 described species and has two strongly different inflorescence and flower types, with corresponding different pollination syndrome. The evolution of these vastly different inflorescence types within this genus was unknown and the main topic in this study.

Methods

We constructed a molecular phylogeny, based on the full nuclear ribosomal DNA and partial plastome, using Bayesian inference and maximum likelihood methods, to test whether the genus is monophyletic, whether all species are monophyletic and if the shift from bat to bee pollination (or vice versa) occurred once in this genus.

Results

All but two species are well supported by the nuclear ribosomal phylogeny. The plastome phylogeny, however, shows a strong geographic signal suggesting strong local hybridization or chloroplast capture, rendering chloroplast barcodes meaningless in this genus.

Conclusions

With our data, we cannot fully resolve the backbone of the tree to clarify sister genera relationships and confirm monophyly of the genus Eperua. Within the genus, the shift from bat to bee and bee to bat pollination has occurred several times but, with the bee to bat not always leading to a pendant inflorescence.
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13.

Premise

Bees provision most of the pollen removed from anthers to their larvae and transport only a small proportion to stigmas, which can negatively affect plant fitness. Though most bee species collect pollen from multiple plant species, we know little about how the efficiency of bees' pollen transport varies among host plant species or how it relates to other aspects of generalist bee foraging behavior that benefit plant fitness, such as specialization on individual foraging bouts.

Methods

We compared the pollen collected and transported by three bee species for 46 co-occurring plant species. Specifically, we compared the relative abundance of pollen taxa in the individual bees' scopae, structures where bees store pollen to provision larvae, with the relative abundance of pollen taxa on the rest of bees' bodies, which is more likely to be transferred to stigmas.

Results

Bees carried five times more pollen grains in their scopae than elsewhere on their bodies. Within foraging bouts, bees were relatively specialized in their pollen collection, but transported proportionally less pollen for the host plants on which they specialized. Across foraging bouts, two bee species transported proportionally less pollen for some of their host plants than for others, though differences didn't consistently follow the same trend as at the foraging bout scale.

Conclusions

Our results suggest that foraging-bout specialization, which is known to reduce heterospecific pollen transfer, also results in less-efficient pollen transport. Thus, bee foragers that visit predominantly one plant species may have contrasting effects on that plant's fitness.
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14.

Premise

Floral scent, usually consisting of multiple compounds, is a complex trait, and its role in pollinator attraction has received increasing attention. However, disentangling the effect of individual floral scent compounds is difficult due to the complexity of isolating the effect of single compounds by traditional methods.

Methods

Using available quasi-isogenic lines (qILs) that were generated as part of the original mapping of the floral scent volatile-related loci CNL1 (benzaldehyde) and TPS2 (β-ocimene) in Capsella, we generated four genotypes that should only differ in these two compounds. Plants of the four genotypes were introduced into a common garden outside the natural range of C. rubella or C. grandiflora, with individuals of a self-compatible C. grandiflora line as pollen donors, whose different genetic background facilitates the detection of outcrossing events. Visitors to flowers of all five genotypes were compared, and the seeds set during the common-garden period were collected for high-throughput amplicon-based sequencing to estimate their outcrossing rates.

Results

Benzaldehyde and β-ocimene emissions were detected in the floral scent of corresponding genotypes. While some pollinator groups showed specific visitation preferences depending on scent compounds, the outcrossing rates in seeds did not vary among the four scent-manipulated genotypes.

Conclusions

The scent-manipulated Capsella materials constructed using qILs provide a powerful system to study the ecological effects of individual floral scent compounds under largely natural environments. In Capsella, individual benzaldehyde and β-ocimene emission may act as attractants for different types of pollinators.
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15.

Objectives

The present study investigated the association of skeletal indicator of stress presence with mean age-at-death as a means of understanding whether commonly studied indicators are indeed indicative of increased frailty.

Materials and Methods

Using a medieval Gaelic population from Ballyhanna (Co. Donegal), the present study assessed the association between skeletal indicators of stress and mean age-at-death using the Kaplan–Meier survival function with log rank test to determine whether these indicators were associated with younger age-at-death, and therefore increased frailty, in sub-adults only (0 to 18 years, N = 139) and through comparison to an all-ages cohort (N = 318).

Results

Only linear enamel hypoplasia was found to be associated with significantly decreased survivorship across the all-ages cohort but, conversely, was associated with increased survivorship when analysis was restricted to sub-adults. All other indicators assessed were associated with increased age-at-death for both all-age cohorts and sub-adult cohorts (cribra orbitalia), increased age-at-death when assessing all ages only (porotic hyperostosis and healed periosteal lesions); or were sufficiently rare in adults to prevent comparative analysis (stunting and micronutrient deficiency). Increased survivorship in individuals with higher numbers of co-morbid skeletal indicators was observed for both sub-adults alone and all age cohort.

Discussion

These findings suggest that these commonly recorded skeletal indicators may be more accurately viewed simply as records of stressor exposure and subsequent survival only, rather than providing evidence that these sub-adults are frailer than their similarly aged-at-death peers. Thus, the demographic and sociocultural context is essential to the interpretation of observed skeletal indicators of stress.
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16.

Premise

Many flowering plants depend on insects for pollination and thus attract pollinators by offering rewards, mostly nectar and pollen. Bee pollinators rely on pollen as their main nutrient source. Pollen provides all essential micro- and macronutrients including substances that cannot be synthesized by bees themselves, such as sterols, which bees need for processes such as hormone production. Variations in sterol concentrations may consequently affect bee health and reproductive fitness. We therefore hypothesized that (1) these variations in pollen sterols affect longevity and reproduction in bumble bees and (2) can thus be perceived via the bees' antennae before consumption.

Methods

We studied the effect of sterols on longevity and reproduction of Bombus terrestris workers in feeding experiments and investigated sterol perception using chemotactile proboscis extension response (PER) conditioning.

Results

Workers could perceive several sterols (cholesterol, cholestenone, desmosterol, stigmasterol, β-sitosterol) via their antennae but not differentiate between them. However, when sterols were presented in pollen, and not as a single compound, the bees were unable to differentiate between pollen differing in sterol content. Additionally, different sterol concentrations in pollen neither affected pollen consumption nor brood development or worker longevity.

Conclusions

Since we used both natural concentrations and concentrations higher than those found in pollen, our results indicate that bumble bees may not need to pay specific attention to pollen sterol content beyond a specific threshold. Naturally encountered concentrations might fully support their sterol requirements and higher concentrations do not seem to have negative effects.
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17.

Premise

Intersexual mating facilitation in flowering plants has been largely underexplored. Duodichogamy is a rare flowering system in which individual plants flower in the sequence male-female-male. We studied the adaptive advantages of this flowering system using chestnuts (Castanea spp., Fagaceae) as models. These insect-pollinated trees produce many unisexual male catkins responsible for a first staminate phase and a few bisexual catkins responsible for a second staminate phase. We hypothesized that duodichogamy increases female mating success by facilitating pollen deposition on stigmas of the rewardless female flowers through their proximity with attractive male flowers responsible for the minor staminate phase.

Methods

We monitored insect visits to 11 chestnut trees during the entire flowering period and explored reproductive traits of all known duodichogamous species using published evidence.

Results

In chestnuts, insects visited trees more frequently during the first staminate phase but visited female flowers more frequently during the second staminate phase. All 21 animal-pollinated duodichogamous species identified are mass-flowering woody plants at high risk of self-pollination. In 20 of 21 cases, gynoecia (female flower parts) are located close to androecia (male flower parts), typically those responsible for the second minor staminate phase, whereas androecia are often distant from gynoecia.

Conclusions

Our results suggest that duodichogamy increases female mating success by facilitating pollen deposition on stigmas by means of the attractiveness of the associated male flowers while effectively limiting self-pollination.
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18.

Premise

Plant ecological strategies are often defined by the integration of underlying traits related to resource acquisition, allocation, and growth. Correlations between key traits across diverse plants suggest that variation in plant ecological strategies is largely driven by a fast–slow continuum of plant economics. However, trait correlations may not be constant through the life of a leaf, and it is still poorly understood how trait function varies over time in long-lived leaves.

Methods

Here, we compared trait correlations related to resource acquisition and allocation across three different mature frond age cohorts in a tropical fern species, Saccoloma inaequale.

Results

Fronds exhibited high initial investments of nitrogen and carbon, but with declining return in photosynthetic capacity after the first year. In the youngest fronds, we found water-use efficiency to be significantly lower than in the oldest mature fronds due to increased transpiration rates. Our data suggest that middle-aged fronds are more efficient relative to younger, less water-use efficient fronds and that older fronds exhibit greater nitrogen investments without higher photosynthetic return. In addition, several trait correlations expected under the leaf economics spectrum (LES) do not hold within this species, and some trait correlations only appear in fronds of a specific developmental age.

Conclusions

These findings contextualize the relationship between traits and leaf developmental age with those predicted to underlie plant ecological strategy and the LES and are among the first pieces of evidence for when relative physiological trait efficiency is maximized in a tropical fern species.
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19.

Objectives

The diploic venous system has been hypothesized to be related to human brain evolution, though its evolutionary trajectory and physiological functions remain largely unclear. This study examines the characteristics of the diploic venous channels (DCs) in a selection of well-preserved Homo neanderthalensis and Upper Paleolithic Homo sapiens crania, searching for the differences between the two taxa and exploring the associations between brain anatomy and DCs.

Materials and Methods

Five H. neanderthalensis and four H. sapiens fossil specimens from Western Europe were analyzed. Based on Micro-CT scanning and 3D reconstruction, the distribution pattern and draining orifices of the DCs were inspected qualitatively. The size of the DCs was quantified by volume calculation, and the degree of complexity was quantified by fractal analyses.

Results

High-resolution data show the details of the DC structures not documented in previous studies. H. neanderthalensis and H. sapiens specimens share substantial similarities in the DCs. The noticeable differences between the two samples manifest in the connecting points surrounding the frontal sinuses, parietal foramina, and asterional area.

Discussion

This study provides a better understanding of the anatomy of the DCs in H. neanderthalensis and H. sapiens. The connection patterns of the DCs have potential utility in distinguishing between the two taxa and in the phylogenetic and taxonomic discussion of the Neandertal-like specimens with controversial taxonomic status.
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20.

Question

How does the plant species composition of Pontic–Pannonian salt‐rich habitats vary on a large geographical scale? Do the floristic differences between Pannonia and the Balkans correspond to the current phytosociological classification?

Location

Pannonia (Hungary, Slovakia, Austria, Czech Republic, Croatia, Serbia, Romania) and the Balkans (Bulgaria, Macedonia, Greece).

Methods

Two thousand four hundred and thirty‐seven relevés from halophytic and sub‐halophytic habitats were classified using a modified TWINSPAN. The crispness of classification was checked. DCA and CCA with climate data as explanatory variables were applied.

Results

The classification was best interpreted at the level of 15 clusters. The vegetation changed along the salinity gradient from sub‐halophytic grasslands (including Trifolion resupinati alliance of the Molinio‐Arrhenatheretalia class and Beckmannion eruciformis and Festucion pseudovinae p. p. alliances of the Festuco‐Puccinellietea class) and reed beds (Bolboschoenion maritimi p. p. alliance; the Phragmito‐Magnocaricetea class), through steppe and wet inland halophytic vegetation (Festucion pseudovinae p. p., Puccinellion limosae, Pucinellion convolutae, Bolboschoenion maritimi p. p. and Juncion gerardii of the Festuco‐Puccinellietea class) towards the extreme halophytic vegetation of the Thero‐Salicornietea, Crypsietea and Juncetea maritimi classes. This gradient was longer in the Balkan region, where it spanned from the sub‐mediterranean salt‐rich grasslands to the extremely halophytic vegetation at the Black Sea coast. The second most important gradient coincided with the water regime. Some vegetation types appeared to be confined to either the Pannonian or the Balkan region (especially within dry sub‐halophytic and steppe halophytic grasslands), while others were distributed across the entire study area. The above‐mentioned pattern did not always correspond with current classification systems.

Conclusions

Variation in salt‐rich vegetation predominantly follows the salinity and water regime gradients. Geographical variation, generally coinciding with climatic and historical effects, is also important, especially in drier salt‐rich habitats. Our large‐scale analysis of the floristic variation of salt‐rich habitats might be useful for the unification of classification systems that differ substantially between the countries involved. In addition, the analysis may be useful for adjustment of a classification system in the poorly explored Balkan region, where particular vegetation types were identified with, or delimited from, Central European vegetation types without detailed comparative analysis until now.
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