Foraging strategy quick response to temperature of Messor barbarus (Hymenoptera: Formicidae) in Mediterranean environments

Animals principally forage to try to maximize energy intake per unit of feeding time, developing different foraging strategies. Temperature effects on foraging have been observed in diverse ant species; these effects are limited to the duration of foraging or the number of foragers involved. The harvester ant Messor barbarus L. 1767 has a specialized foraging strategy that consists in the formation of worker trails. Because of the high permeability of their body integument, we presume that the length, shape, and type of foraging trails of M. barbarus must be affected by temperature conditions. From mid-June to mid-August 1999, we tested the effect on these trail characteristics in a Mediterranean forest. We found that thermal stress force ants to use a foraging pattern based on the variation of the workers trail structure. Ants exploit earlier well-known sources using long physical trails, but as temperatures increases throughout the morning, foragers reduce the length of the foraging column gradually, looking for alternative food sources in nonphysical trails. This study shows that animal forage can be highly adaptable and versatile in environments with high daily variations.     

Submerged macrophyte seed bank in a Mediterranean temporary marsh: abundance and relationship with established vegetation

The abundance and composition of the submerged macrophyte seed bank in the Doñana marsh (southwestern Spain) was evaluated to assess its relationship with the overlying vegetation. The results obtained demonstrate the existence of a dense seed-bank, both in terms of the number of seeds and their biomass, which represented about 10% (5% for angiosperms and >20% for Charophyta) of the total plant biomass, which ensures the maintenance of the annual submerged macrophyte populations of the seasonally inundated Donñana marsh. Seed bank and established vegetation were coupled, as reflected in the existence of significant correlations between their structure and abundance. This coupling was lacking for Charophyta, whose oospores are widespread and abundant across the marsh, even at locations where they are absent, or rare, in the established vegetation. These differences between the seed bank of annual angiosperms and Charophyta appear to reflect, in part, structural differences between angiosperm seeds and Charophyta oospores, with important ecological consequences. An important aspect of these differences is the allocation of Charophyta reproductive effort to many small propagules lacking embryo storage, compared to those of angiosperms, which ensures their efficient dispersal and numerical abundance in the seed bank.     

Regeneration processes of biological soil crusts,macro-cryptogams and vascular plant species after fine-scale disturbance in a temperate region: Recolonization or successional replacement?

Open base-rich sandy habitats are often characterised by an extensive cover of biological soil crusts (BSCs). We studied the impact of mechanical disturbances on cyanobacteria/algae-dominated BSCs, macro-cryptogams (bryophytes, lichens), phanerogams and their regeneration processes in a 3-year field experiment in a temperate region. We compared controls with two fine-scale disturbance regimes, each based on 50 plots (20 cm×20 cm): 1. BSCs were raked (weak disturbance), 2. BSCs were completely removed (strong disturbance). We used a Sileno conicae-Cerastion semidecandri habitat as a model area for the field study and ask if the crusts are able to regenerate or will be replaced by macro-cryptogams and/or phanerogams.     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

Plant species richness in calcareous grasslands as affected by dispersability in space and time

Abstract. Species richness in calcareous grassland is discussed against the background of historical dispersal processes associated with traditional land-use management such as grazing, but also the artificial establishment by hayseed. Important vectors in the traditionally man-made landscape were sheep and cattle or other livestock such as goats. Calcareous grasslands were not only connected to each other but also to other habitats such as villages, forests, arable fields and heathlands by these vectors which could cover large distances (e.g. transhumance shepherding), which is not the case in the current man-made landscape. Species richness after restoration management of abandoned and afforested calcareous grasslands was predicted by using characters of dispersability in space and time. These were the persistence of the species in the vegetation and the diaspore bank after abandonment or afforestation and the dispersal capacity through wind and sheep. The results reveal that reintroduction of sheep grazing is necessary to reestablish the original species richness. The first validation of the prediction of the succession on clear-cut sites and a comparison with data of species composition in abandoned quarries and the surroundings made it obvious that a species' own dispersal capacity in space is very low except for some well wind-dispersed species. Therefore, it is recommended to include and to simulate dispersal processes in future management to be able to restore the original species richness.     

Role of soil seed bank along a disturbance gradient in an alpine meadow on the Tibet plateau

We examined the role of the soil seed bank along a grazing disturbance gradient and its relationship with the vegetation of alpine meadows on the Tibet plateau, and discussed the implications for restoration. The seed bank had a high potential for restoration of species-rich vegetation; 62 species were identified in the vegetation and 87 in the seed bank, 39 species being common to both. Mean seed density was 3069–6105 viable seeds m⁻². The density of buried seeds increased significantly with increasing disturbance, indicating that restoration of disturbed areas is not seed limited. Seed density and species richness decreased with depth. The proportion of perennial species decreased with decrease in disturbance both in seed bank and in vegetation. A large portion of species with persistent seeds in the disturbed areas indicate that this seed type can be regarded a strategy of adaptation to current disturbances. Detrended correspondence analysis (DCA) showed significant differences of species composition between seed bank and vegetation, except for the seriously disturbed site. Our results suggest that the establishment of new species in severely disturbed areas is more dependent on the seed bank. By contrast, the restoration in less-disturbed and mature meadows does not rely on seed banks, and the establishment of the vegetation in these communities is more likely to rely on seed dispersal from the standing vegetation and on species with vegetative reproduction.     

Seed bank dynamics in tall‐tussock grasslands along an altitudinal gradient

Abstract. We studied the germinable soil seed bank of tall‐tussock grasslands along an altitudinal gradient in the mountains of central Argentina. We selected 10 sampling plots at three altitudinal levels (1200 m, 1600 m and 2200 m). We assessed the composition of the established vegetation and took ten compound soil samples (0 ‐ 5 cm depth) at each plot in autumn and spring. The soil samples were sieved, chilled, and incubated in a glasshouse to assess the composition of the seed bank. The similarity between the composition of the seed bank flora and that of the established vegetation was low throughout the gradient. Most species did not change their seed bank strategy along the gradient. Seed bank richness and density increased with altitude. Most species had a persistent seed bank at all altitudinal levels, and the proportion of such species increased with altitude. These results suggest that a cold climate directly and/or indirectly favours the formation of seed banks and seed persistence in the soil.     

Burial of canopy-stored seeds in the annual psammophyte Agriophyllum squarrosum Moq. (Chenopodiaceae) and its ecological significance

Agriophyllum squarrosum Moq. is a dominant annual on sand dunes in the arid regions of central Asia. A high percentage of seeds is retained on dead plants which become covered by moving sand, but little is known about the ecological significance of burial of canopy-stored seeds. We investigated the size and dynamics of the buried canopy-stored seed bank and effects of burial on seed germination. In March (during the windy season), May (beginning of the germination season), and July (middle of the growing season), the number of seeds per square meter in sample plots in the dunes was 623, 223 and 22, respectively, with 54.6, 30.6 and 12.9% of the total seeds retained on buried plant canopies. In a controlled experiment, more seedlings emerged from released (dispersed) than from canopy-stored seeds when burial depth was the same. No viable ungerminated released seeds were found, but 45–80% of the ungerminated canopy-stored seeds were viable. In general, with an increase in applied water germination of released seeds buried at a depth of 1 or 2 cm and of canopy-stored seeds buried at 1 cm increased, but regardless of watering regime few or no released seeds at 4 cm or canopy-stored seeds at 2 or 4 cm germinated. Significantly more seedlings emerged from plants buried in a horizontal than in a vertical position. Seedlings originating from buried canopy-stored seeds on an active dune accounted for only 5.4% of the total seedlings emerging, and most of them emerged later than those from released seeds. Thus, seed release is more effectively postponed in buried than in exposed canopies, and burial of canopy-stored seeds is a mechanism that helps regulate seed germination and seedling emergence of A. squarrosum on active dunes.     

Seed rain and seed bank of Chinese yew (Taxus chinensis var. mairei) population in Tianmu Mountain☆

Seed rain and seed bank of a Chinese yew (Taxus chinensis var. mairei) population in Tianmu Mountain were researched in 2008 and 2009. The seed rain lasted from 16th–23th of October to 5th–14th of December, and the heaviest seed falling period was from 2nd to18th of November. The intensity of seed rain showed a great inter-annual variation, with a good harvest in 2008. The fallen seeds were composed of 49.9% proportion of immature seed, 33.8% proportion of chewed seed and 16.3% proportion of mature seed. The analysis on the soil seed bank under mother forest showed that the number of intact seeds was 122.75 ± 108.08 grain/m² in October, 279.25 ± 210.73 grain/m² in December 2008, and 166.5 ± 165.34 grain/m² in October, 322.5 ± 275.73 grain/m² in December 2009. The increased number of seed was 156.5 ± 222.723 grain/m² in 2008 and 156 ± 275grain/m² in 2009, which showed a significant variation. Large number of intact seeds added into soil seed bank after seed rain each year. The number of intact seeds in soil seed bank decreased 112.75 ± 47.74 grain/m² from December 2008 to October 2009. Large number of intact seeds lost from seed rot and seed predation by animals. The number of seeds in soil bank under bamboo forest was much lower than that of mother tree forest, and the increased number of seeds was 0.63 ± 1.60 grain/m² in 2008 and 2.88 ± 1.86 grain/m² in 2009. The number of seedling was 0.73 ± 1.10 trees/m² in mother tree forest and 0.09 ± 0.35 trees/m² in bamboo forest. Seedling survival ratio was 0.37% in mother tree forest and 10.23% in bamboo forest. The micro-habitat in bamboo forest was fit for seed germination. Birds transported seeds to bamboo forest, and had an important effect on the regeneration of Chinese yew.     

Seed rain and seed bank of Chinese yew (Taxus chinensis var. mairei) population in Tianmu Mountain

Seed rain and seed bank of a Chinese yew (Taxus chinensis var. mairei) population in Tianmu Mountain were researched in 2008 and 2009. The seed rain lasted from 16th–23th of October to 5th–14th of December, and the heaviest seed falling period was from 2nd to18th of November. The intensity of seed rain showed a great inter-annual variation, with a good harvest in 2008. The fallen seeds were composed of 49.9% proportion of immature seed, 33.8% proportion of chewed seed and 16.3% proportion of mature seed. The analysis on the soil seed bank under mother forest showed that the number of intact seeds was 122.75 ± 108.08 grain/m² in October, 279.25 ± 210.73 grain/m² in December 2008, and 166.5 ± 165.34 grain/m² in October, 322.5 ± 275.73 grain/m² in December 2009. The increased number of seed was 156.5 ± 222.723 grain/m² in 2008 and 156 ± 275grain/m² in 2009, which showed a significant variation. Large number of intact seeds added into soil seed bank after seed rain each year. The number of intact seeds in soil seed bank decreased 112.75 ± 47.74 grain/m² from December 2008 to October 2009. Large number of intact seeds lost from seed rot and seed predation by animals. The number of seeds in soil bank under bamboo forest was much lower than that of mother tree forest, and the increased number of seeds was 0.63 ± 1.60 grain/m² in 2008 and 2.88 ± 1.86 grain/m² in 2009. The number of seedling was 0.73 ± 1.10 trees/m² in mother tree forest and 0.09 ± 0.35 trees/m² in bamboo forest. Seedling survival ratio was 0.37% in mother tree forest and 10.23% in bamboo forest. The micro-habitat in bamboo forest was fit for seed germination. Birds transported seeds to bamboo forest, and had an important effect on the regeneration of Chinese yew.     

Strategies of the beetle Oochrotus unicolor (Tenebrionidae) thriving in the waste dumps of seed-harvesting Messor ants (Formicidae)

1. A diverse group of arthropods have adapted to the niches found inside the nests of social insects. Studies mostly focused on very specialised parasites residing in the brood chambers. However, the biology and strategies of symbionts occupying other niches, such as waste dumps, are underexplored. 2. Using a series of complementary experiments, this study demonstrated that the Mediterranean beetle Oochrotus unicolor has adapted to the waste dump niche found in the nests of Messor harvester ants. 3. Laboratory experiments confirmed field observations that the beetle preferentially resided in the refuse pits. Next, it was shown that the beetles readily consumed seeds and flour, whereas other food sources were poorly accepted and ant brood was never even eaten. The beetles did not elicit a strong aggression response in Messor ants, and they could tolerate very high densities of workers without clear costs. The beetles modestly mimicked the nest recognition cues of their Messor host. This imperfect mimicry could promote the adoption of the beetle in the ant colony, in concert with mechanical defence generated by its tank-like body. Isolation of the beetle from its host did not significantly affect the beetle's chemical cuticular profile nor did it provoke elevated ant aggression, indicating that the beetle does not acquire the chemicals passively from its host. 4. This paper discusses the fact that waste dumps in social insect nests are hotspots for arthropod symbionts. It shows that symbionts in this niche may employ behavioural, trophic and chemical strategies that are different from those found in other niches of social insect nests.     

Seed dispersal in Erythronium grandiflorum (Liliaceae)

Primary and secondary seed dispersal was investigated for the glacier lily Erythronium grandiflorum in the Colorado Rocky Mountains. These heavy seeds have no obvious adaptations for biotic or abiotic dispersal, but can be thrown short distances when the dehiscent fruits are shaken by wind. We used sticky traps to measure primary transport of seeds up to 1 m away from individual plants. A seed cafeteria experiment examined the role of ants and rodents in secondary seed transport. Primary dispersal by wind was positively skewed and median transport distances were influenced by variation in plant height. Secondary dispersal was negligible compared to Viola nuttallii, an elaiosome-bearing species. Thus, seed dispersal was highly restricted in E. grandiflorum, and a 1 m radius encompassed the modal section of the seed dispersal curve. The seed dispersal component of gene flow was quantified and combined with previous measurements of pollen flow to yield a more complete estimate of Wright's neighborhood size, N _e, for E. grandiflorum. The lack of a special seed dispersal mechanism in E. grandiflorum is discussed in terms of a source-sink model for seedling establishment with respect to distance from the parental plants.     

Task-Matching and Short-Term Size Shifts in Foragers of the Harvester Ant, Messor pergandei (Hymenoptera: Formicidae)

Some social insects produce workers of variable size and/or shape. One hypothesis to explain such variation is that it increases colony efficiency in performing diverse tasks, with greater efficiency leading to greater colony-level fitness. Using the granivorous ant Messor pergandei, which produces a unimodal distribution of worker sizes, I explored two corollaries of this ergonomic hypothesis: that there should be detectable task-matching and that the mean size should shift predictably with a change in task. In five experiments conducted in different years and sites in California deserts, colonies were baited with fragments of wheat or millet ground and sieved to different sizes. In six of eight colonies so baited the minimum linear dimension or mass of seeds carried by foragers was significantly related to forager size, with r² values ranging from 3 to 30%. In four of the five experiments, colonies were subsequently baited with either large seed fragments alone or small fragments alone. In three of these four experiments I detected a rapid response, over a few hours or days, in which foragers deployed by colonies shifted to larger or smaller mean size in directions predicted by size of the bait. These results suggest that an ergonomic function is one contributor to worker size polymorphism in M. pergandei. Although task-matching did not explain a substantial amount of the variance in forager size in all experiments, the rapid size shifts provide behavioral evidence that task-matching is important to colonies.     

Restoration of wooded meadows ‐ a comparative analysis along a chronosequence on Öland (Sweden)

Abstract. Wooded meadows on the Baltic Island of Öland result from traditional agricultural management over centuries which has led to a species‐rich vegetation with high species diversity. Today, nearly all of these meadows have been abandoned and became rapidly overgrown by deciduous shrub and tree species forming a closed canopy which resulted in a rapid and strong decrease in species numbers of the herb layer. Recent efforts aim to restore overgrown wooded meadows by cutting single shrubs and trees to open the canopy. However, the effects of abandonment as well as of any restoration management in wooded meadows have rarely been documented until now. Mechanisms driving succession after restoration such as the dispersal potential of the respective species over time and space have not been analysed yet. Therefore, a chronosequence was studied which included a traditionally managed wooded meadow, an overgrown meadow which has been abandoned for more than 100 yr and a meadow which was restored 36 yr ago by cutting and is now grazed. We analysed the soil seed bank of the 3 meadows in comparison with the established vegetation and endozoochorous seed dispersal by cattle and sheep. After abandonment 87% of the typical grassland species vanished from the established vegetation and were replaced by species characteristic of woodland and disturbed grassland communities. The mean number of species decreased from 52 species per plot (4 m²) to 18 species. Mean species number and number of seeds in the seed bank declined significantly from the traditionally managed to the overgrown meadow. Most of the grassland species were assigned to a transient seed bank type while only 1/3 could be classified as having a short‐term persistent seed bank. Thus, restoration of wooded meadows cannot rely on the soil seed bank. Endozoochorous seed dispersal by cattle and sheep was shown for 15% of the species with seed densities per 100 g air dried dung of 737 and 767, respectively. Movement of animals between ancient and restored wooded meadows is recommended since many of the species only occurred in low densities and therefore, will probably not be found in the dung samples.     

Allocation to reproduction following experimental defoliation in Platanthera bifolia (Orchidaceae)

The roles of herbivory and pollination success in plant reproduction have frequently been examined, but interactions between these two factors have gained much less attention. In three field experiments, we examined whether artificial defoliation affects allocation to attractiveness to pollinators, pollen production, female reproductive success and subsequent growth in Platanthera bifolia L. (Rich.). We also recorded the effects of inflorescence size on these variables. We studied the effects of defoliation on reproductive success of individual flowers in three sections of inflorescence. Defoliation and inflorescence size did not have any negative effects on the proportion of opened flowers, spur length, nectar production or the weight of pollinia. However, we found that hand-pollination increased relative seed production and defoliation decreased seed set in most cases. Interactions between hand-pollination and defoliation were non-significant indicating that defoliation did not affect female reproductive success indirectly via decreased pollinator attraction. Plants with a large inflorescence produced relatively more seeds than plants with a small inflorescence only after hand-pollination. The negative effect of defoliation on relative capsule production was most clearly seen in the upper sections of the inflorescence. In addition to within season effects of leaf removal, defoliated P. bifolia plants may also have decreased lifetime fitness as a result of lower seed set within a season and because of a lower number of reproductive events due to decreased plant size (leaf area) following defoliation. Our study thus shows that defoliation by herbivores may crucially affect reproductive success of P. bifolia.     

A first report of water hyacinth (Eichhornia crassipes) soil seed banks in South Africa

We investigated water hyacinth seed banks in several aquatic systems in South Africa. Fifteen sites, mainly in the Eastern and Western Cape provinces, were surveyed from August to October 2009. Soil seed density varied between 0 and 2534 seeds/m² but did not differ significantly between the type of waterbody (impoundment vs. river) or the history of control carried out at a site. Average germination was 54.17% with very fast velocities (Vigour Index = 36.66) and maximum germination around three days. Although we demonstrated the existence of an important reservoir of seeds, results from this study indicated that a combination of factors such as water fluctuation, eutrophication and seed decomposition might have had a great influence on dispersal and persistence of seeds.     

Secondary successional patterns in a sagebrush (Artemisia tridentata) community as they relate to soil disturbance and soil biological activity

The relationship between secondary succession, soil disturbance, and soil biological activity were studied on a sagebrush community (Artemisia tridentata) in the Piceance Basin of northwestern Colorado, U.S.A. Four levels of disturbance were imposed. I: the vegetation was mechanically removed and as much topsoil as possible was left; 2: the vegetation was mechanically removed and the topsoil scarified to a depth of 30 cm; 3: topsoil and subsoil were removed to a depth of 1 m, mixed and replaced; 4: topsoil and subsoil were removed to a depth of 2 m and replaced in a reverse order. Plant species composition, dehydrogenase and phosphatase enzymatic activity, mycorrhizae infection potentials, and percent organic matter were the variables measured.Treatment 4 drastically altered the pattern of vegetation succession. Treatments 2, 3, and 4 started with Salsola iberica as the dominant species but six years later, 3 and to lesser extent 2 changed in the direction of the species composition of 1, dominated by perennial grasses and perennial forbs. Treatment 4 developed a shrub dominated community. The rate of succession was not decreased by the increased levels of disturbance. Both dehydrogenase enzyme activity and mycorrhizae infection potential (MIP) increased with the change from Salsola iberica to a vegetation dominated by either perennial grasses and forbs or shrubs. The intensity of disturbance in 2, 3, and 4 reduced drastically dehydrogenase activity and MIP, but in six years they recovered to levels comparable to 1. Phosphatase enzyme activity and organic matter were unrelated to species composition but related to treatment and time elapsed. In both cases a significant decrease was observed throughout the six-year period.Nomenclature followsThis study was funded by the United States Department of Energy under Contract No. DE-AS02-76EV04018.     

Top‐soil translocation as a technique in the re‐creation of species‐rich meadows

Abstract. In France, most civil engineering and excavation projects are at present accompanied by compensatory measures with the aim of preserving biodiversity. In order to avoid the destruction of a habitat of high conservation interest in NE France, harbouring two legally protected plant species, an experiment of soil translocation was conducted on an area of 1 ha. The donor site was an extensively managed mesophilic meadow and the receiving site was a neighbouring arable land. The vegetation of the translocated meadow was described 8 and 17 months after soil translocation, and compared (1) with vegetation resulting from more classical restoration techniques tested on the arable land (natural regeneration and seed mixture sowing) and (2) with the soil seed bank and vegetation previously present on the donor site. Results showed that the soil translocation technique permitted the development of many meadow species, including two legally protected species, and few ruderal species despite a large area of bare ground. This technique seems effective in terms of number and abundance of meadow species compared to natural regeneration and commercial seed sowing. In the case of the two classical methods, species richness was lower and only widespread species were present. Topsoil translocation provides a good compensatory method to avoid habitat and species destruction. However, the study should be continued, with the aim of assessing the longer term development and stabilization of the vegetation of the translocated meadow.     

Ant foraging on extrafloral nectaries of Qualea grandiflora (Vochysiaceae) in cerrado vegetation: ants as potential antiherbivore agents

Summary Qualea grandiflora is a typical tree of Brazilian cerrados (savanna-like vegetation) that bears paired extrafloral nectaries (EFNs) along its stems. Results show that possession of EFNs increases ant density on Q. grandiflora shrubs over that of neighbouring non-nectariferous plants. Frequency of ant occupancy and mean number of ants per plant were much higher on Qualea than on plants lacking EFNs. These differences resulted in many more live termitebaits being attacked by foraging ants on Qualea than on neighbours without EFNs. Termites were attacked in equal numbers and with equal speeds on different-aged leaves of Qualea. The greatest potential for herbivore deterrence was presented by Camponotus ants (C. crassus, C. rufipes and C. aff. blandus), which together attacked significantly more termites than nine other ant species grouped. EFNs are regarded as important promoters of ant activity on cerado plants.