Fire controls population structure in four dominant tree species in a tropical savanna

The persistence of mesic savannas has been theorised as being dependent on disturbances that restrict the number of juveniles growing through the sapling size class to become fire-tolerant trees. We analysed the population structures of four dominant tropical savanna tree species from 30 locations in Kakadu National Park (KNP), northern Australia. We found that across KNP as a whole, the population size structures of these species do not exhibit recruitment bottlenecks. However, individual stands had multimodal size-class distributions and mixtures of tree species consistent with episodic and individualistic recruitment of co-occurring tree species. Using information theory and multimodel inference, we examined the relative importance of fire frequency, stand basal area and elevation difference between a site and permanent water in explaining variations in the proportion of sapling to adult stems in four dominant tree species. This showed that the proportion of the tree population made up of saplings was negatively related to both fire frequencies and stand basal area. Overall, fire frequency has density-dependent effects in the regulation of the transition of saplings to trees in this Australian savanna, due to interactions with stem size, regeneration strategies, growth rates and tree–tree competition. Although stable at the regional scale, the spatiotemporal variability of fire can result in structural and floristic diversity of savanna tree populations.     

Impacts of savanna trees on forage quality for a large African herbivore

Recently, cover of large trees in African savannas has rapidly declined due to elephant pressure, frequent fires and charcoal production. The reduction in large trees could have consequences for large herbivores through a change in forage quality. In Tarangire National Park, in Northern Tanzania, we studied the impact of large savanna trees on forage quality for wildebeest by collecting samples of dominant grass species in open grassland and under and around large Acacia tortilis trees. Grasses growing under trees had a much higher forage quality than grasses from the open field indicated by a more favourable leaf/stem ratio and higher protein and lower fibre concentrations. Analysing the grass leaf data with a linear programming model indicated that large savanna trees could be essential for the survival of wildebeest, the dominant herbivore in Tarangire. Due to the high fibre content and low nutrient and protein concentrations of grasses from the open field, maximum fibre intake is reached before nutrient requirements are satisfied. All requirements can only be satisfied by combining forage from open grassland with either forage from under or around tree canopies. Forage quality was also higher around dead trees than in the open field. So forage quality does not reduce immediately after trees die which explains why negative effects of reduced tree numbers probably go initially unnoticed. In conclusion our results suggest that continued destruction of large trees could affect future numbers of large herbivores in African savannas and better protection of large trees is probably necessary to sustain high animal densities in these ecosystems.     

Positive and negative effects of grass,cattle, and wild herbivores on <Emphasis Type="Italic">Acacia</Emphasis> saplings in an East African savanna

Plant–plant interactions can be a complex mixture of positive and negative interactions, with the net outcome depending on abiotic and community contexts. In savanna systems, the effects of large herbivores on tree–grass interactions have rarely been studied experimentally, though these herbivores are major players in these systems. In African savannas, trees often become more abundant under heavy cattle grazing but less abundant in wildlife preserves. Woody encroachment where cattle have replaced wild herbivores may be caused by a shift in the competitive balance between trees and grasses. Here we report the results of an experiment designed to quantify the positive, negative, and net effects of grasses, wild herbivores, and cattle on Acacia saplings in a Kenyan savanna. Acacia drepanolobium saplings under four long-term herbivore regimes (wild herbivores, cattle, cattle + wild herbivores, and no large herbivores) were cleared of surrounding grass or left with the surrounding grass intact. After two years, grass-removal saplings exhibited 86% more browse damage than control saplings, suggesting that grass benefited saplings by protecting them from herbivory. However, the negative effect of grass on saplings was far greater; grass-removal trees accrued more than twice the total stem length of control trees. Where wild herbivores were present, saplings were browsed more and produced more new stem growth. Thus, the net effect of wild herbivores was positive, possibly due to the indirect effects of lower competitor tree density in areas accessible to elephants. Additionally, colonization of saplings by symbiotic ants tracked growth patterns, and colonized saplings experienced lower rates of browse damage. These results suggest that savanna tree growth and woody encroachment cannot be predicted by grass cover or herbivore type alone. Rather, tree growth appears to depend on a variety of factors that may be acting together or antagonistically at different stages of the tree’s life cycle.     

Host-preference and density of woodrose-forming mistletoes (Loranthaceae) on savanna vegetation,South Africa

In the Bushbuckridge region of South Africa host preference and density of two woodrose-forming mistletoes, Erianthemum dregei (Eckl. & Zeyh.) V. Tieghem and Pedistylis galpinii (Schinz ex Sprague) was quantified in relation to land-use (harvested or unharvested), rainfall (high > 660 or low < 660 mm year^–1) and catenal position (top or lower slope). These two mistletoes are generalist hemi-parasites of savanna trees and shrubs occurring on 25 and 17 hosts respectively, seven of which are shared. Thirty-six percent of woody plant species recorded were found to be hosts. Although Sclerocarya birrea (A. Rich.) Hochst. comprised only 4% of woody plant density in the environment, it was the principal host for both mistletoes, accounting for 71% of total E. dregei and 42% of P. galpinii infection. Mistletoe infection relative to density of Ficus stuhlmanii, Trichilia emetica and Cassine transvaalensis indicated that these were preferential hosts to S. birrea. Mistletoe host preference was negatively correlated with host wood density. Mistletoe number per tree had a weak relationship to canopy size. Mistletoes of all size classes were denser at high rainfall relative to low rainfall sites. Interestingly, the overall mistletoe size class distribution was similar between harvested and unharvested sites. The ratio of living to dead mistletoe was 2 to 1 for E. dregei and 1.5 to 1 for P. galpinii. There are sufficient dead mistletoes in unharvested and harvested areas to satisfy present market demand. Living E. dregei predominated in harvested rather than unharvested areas suggesting that current-harvesting levels had little or no negative effect on the population. In contrast, P. galpinii was denser in unharvested areas possibly owing to its higher market value and thus higher harvesting levels.     

Biological control of an alien tree, Acacia cyclops, in South Africa: impact and dispersal of a seed-feeding weevil, Melanterius servulus

Several Australian Acacia species have become highly invasive in a number of ecologically sensitive areas of South Africa. Many have useful attributes that, to an extent, have hampered biocontrol efforts by restricting potential agents to those reducing seed production whilst not affecting vegetative growth. The outcome of the introduction of a seed-feeding weevil, Melanterius servulus, against Acacia cyclops in 1994, was assessed by investigating the plant’s reproductive phenology, as well as levels of damage caused by the beetles. The study provided essential information on host/agent interactions and, on the build up and spread of populations of the agent. Seed damage is commonly as high as 90% (exceptionally it reaches 100%), and dispersal rates of the beetles on average is almost 2 km per year. The consistently high levels of seed damage achieved hold promise for the future of the program, however, its ultimate success will be further enhanced through the integration of biological control with other manual clearing operations that are currently employed on a large scale in South Africa.     

Juggling carbon: allocation patterns of a dominant tree in a fire-prone savanna

In frequently burnt mesic savannas, trees can get trapped into a cycle of surviving fire-induced stem death (i.e. topkill) by resprouting, only to be topkilled again a year or two later. The ability of savanna saplings to resprout repeatedly after fire is a key component of recent models of tree–grass coexistence in savannas. This study investigated the carbon allocation and biomass partitioning patterns that enable a dominant savanna tree, Acacia karroo, to survive frequent and repeated topkill. Root starch depletion and replenishment, foliage recovery and photosynthesis of burnt and unburnt plants were compared over the first year after a burn. The concentration of starch in the roots of the burnt plants (0.08 ± 0.01 g g⁻¹) was half that of the unburnt plant (0.16 ± 0.01 g g⁻¹) at the end of the first growing season after topkill. However, root starch reserves of the burnt plants were replenished over the dry season and matched that of unburnt plants within 1 year after topkill. The leaf area of resprouting plants recovered to match that of unburnt plants within 4–5 months after topkill. Shoot growth of resprouting plants was restricted to the first few months of the wet season, whereas photosynthetic rates remained high into the dry season, allowing replenishment of root starch reserves. ¹⁴C labeling showed that reserves were initially utilized for shoot growth after topkill. The rapid foliage recovery and the replenishment of reserves within a single year after topkill implies that A. karroo is well adapted to survive recurrent topkill and is poised to take advantage of unusually long fire-free intervals to grow into adults. This paper provides some of the first empirical evidence to explain how savanna trees in frequently burnt savannas are able to withstand frequent burning as juveniles and survive to become adults. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Assessing the Effects of Invasive Alien Species on Rural Livelihoods: Case Examples and a Framework from South Africa

The detrimental impacts of invasive alien species (IAS) on ecosystem goods and services and local and regional economies are well documented. However, the use of IAS by rural communities is little understood, and rarely factored into IAS control programmes. Understanding the use of IAS by rural communities and factoring these into cost-benefit models is complex, depending upon a range of local-level attributes such as the time since invasion, abundance, and local-level costs and benefits. This paper reports on two case studies examining the role of IAS in rural livelihoods in the Eastern Cape, South Africa. In both cases, rural communities made widespread consumptive use of the IAS and generally would prefer higher densities, except in certain key localities. Several households traded in IAS products to generate supplementary income. We present a conceptual framework to guide interpretation of these and future case studies, considering attributes such as time since invasion, the competitiveness of the species, and the relative costs and benefits.     

Lifecycle of the bruchid beetle Bruchidius uberatus and its predation of Acacia nilotica seeds in a tree savanna in Botswana

The life cycle of the bruchid beetle Bruchidius uberatus in seeds of Acacia nilotica has been investigated under field and laboratory condition. In contrast to the multivoltine populations in the Sudan, in Botswana most beetles of B. uberatus are univoltine. They emerge from the seed and pod in early spring (October to November). Only a small part of the population, which emerge in late summer (February to March) is multivoltine, as confirmed by rearing experiments. The reproductive activity of females is not stimulated by pollen, as found in feeding experiments. Minimum life-span of adult beetles varies between 4 and 40 days, but it did not differ between univoltine and multivoltine beetles. Other data on life-history are 22 days for hatching of the fist-instar larva at a temperature regime of 20/15 °C (day/night) and 15 days at a temperature regime of 32/15 °C and 3 to 11 months for development from larvae to beetle.During dry storage of seeds B. uberatus can destroy the total amount of stored seeds within a few years. In stored seed pool, host-specificity of larva is low.The life cycles of the Botswana population have been discussed in relation to that of the species in the Sudan.

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Zusammenfassung Der Lebenszyklus des Samenkäfers Bruchidius uberatus in Samen von Acacia nilotica wurde unter Feld- und Laborbedingungen untersucht. Im Gegensatz zu multivoltinen Populationen im Sudan (Peake, 1982) sind die Populationen von B. uberatus in Botswana vor allem univoltin. Adulte Käfer verlassen die Samen von A. nilotica während des Süd-Frühlings (Oktober/November). Im Spätsommer (Februar/März) geschlüpfte Käfer entstammen multivoltinen Genotypen, wie Züchtungsexperimente gezeigt haben. Die reproduktive Aktivität der Weibchen wird nicht durch die Zufuhr von Pollen oder Zucker stimuliert.Die Entwicklungsdauer vom Ei bis zum 1. Larvenstadium beträgt 15 Tage bis 52 und 3 bis 11 Monate bis zum adulten Käfer, deren Lebenserwartung unter Laboratoriumbedingungen 4 bis 40 Tage beträgt. Hierbei besteht kein Unterschied zwischen uni- und multivoltinen Käfern. Der lange Lebenszyklus in Botswana steht im Gegensatz zu dem 40-tägigen Lebenszyklus von B. uberatus im Sudan, der offensichtlich durch die Jahrhunderte andauernde Sammlung von Früchten verursacht ist.Samenvorräte, die zu Forstzwecken längere Zeit aufbewahrt werden, können innerhalb von 5 bis 8 Jahren völlig zerstört werden. Dabei ist die Wirtsspezifität gering. Die Infektion betrifft auch andere Acacia-Arten, z.B. A. burkei, A. erioloba, A. hebeclada, A. mellifera, A. robusta und A. tortilis. Durch die rasche Zerstörung der Embryoachse haben die Käfer keinen positiven Einfluss auf die Keimung dieser hartschaligen Acacia-Samen.

     

Effects of controlled livestock grazing and annual prescribed fire on epigeal termite mounds in a savannah woodland in Burkina Faso

The diversity and abundance of epigeal termite mounds were investigated in response to controlled livestock grazing and annual prescribed fire in a Sudanian savannah-woodland in Tiogo State Forest. Sampling of termite mounds was carried out in 4×4 subplots of 0.25 ha in a split-plot experimental design during the rainy season in 2002. There were two main plots of which one was fenced to exclude livestock grazing and the second exposed to grazing. Each of the main plots included 4 subplots with annual prescribed fire since 1992 and 4 subplots without fire. Data were collected on the number and characteristics of termite mounds. A mean density of 698 mounds ha⁻¹ was recorded. Mounds built by Trinervitermes spp. were the most abundant followed by Cubitermes spp., Macrotermes subhyalinus and M. bellicosus. The large mound-builders Macrotermes spp. dominated the community interms of basal area (96% of the total) and above-ground volume (99%). The diversity of mound types was notaffected by livestock grazing and annual early prescribed fire (P>0.05). There was no statistical effect of livestock grazing on mound density, whereas a strong depressive effect of annual fire was observed for Trinervitermes spp. mound density (P=0.012). In this ecosystem, annual prescribed fire appeared to be the major determinant for termite mound abundance. Received 2 February 2007; revised 23 October 2007 and 21 January 2008; accepted 22 Feburary 2008.     

The distribution,abundance, and the effects of fire on mound building termites (Trinervitermes and Cubitermes spp., Isoptera: Termitidae) in northern guinea savanna West Africa

Summary Termite mound densities in typical guinea savanna, Detarium, and grassland (boval) habitats in northern guinea savanna were determined by random quadratting of 2–3 sites in each habitat (100, 10x10 m quadrats per habitat). Dominant species in guinea savanna were T. geminatus (46 mounds ha^-1) and T. oeconomus (21 mounds ha^-1), in Detarium T. geminatus (59 mounds ha^-1) and C. curtatus (45 mounds ha^-1) and in boval C. curtatus (72 mounds ha^-1) and T. geminatus (22 mounds ha^-1). Only C. curtatus densities and total densities differed significantly between sites within habitats, but all species differed significantly in abundance between habitats. The composition of each community was related to general environment but no particular environmental variable was shown to be a major determinant of termite distribution. Evidence for the limitation of termite populations was obtained from indirect evidence of competition between colonies in Detarium, and by experimental manipulation of fire regimes in the typical guinea savanna habitat. Harvester termites increased four-five fold over two years in fire-protected plots as a result of increased food supplies. Total termite densities in the fire-protected community equilibrated to the new population density (100 mounds ha^-1) after only two-three years.     

Variation in seed and seedling traits in Pithecellobium pedicellare,a tropical rain forest tree

Summary Pithecellobium pedicellare, a mimosoid legume, is a large canopy tree in the tropical rain forests of Costa Rica. We examined the pattern of variation in seed weight, germination date, hypocotyl length (stem), and rachis length (the first leaf) of the seedlings in this species. Seeds collected from widely dispersed individual trees at the La Selva Biological Station, Costa Rica, were randomly planted in blocks, and grown under controlled, indoor conditions for about 2 weeks. There-fore, we were able to quantify the effects of maternal family on mean seed weight and the effects of maternal family and microenvironment on the remaining traits examined. A significant effect of maternal family was detected for all traits. In particular, the maternal effects on germination date and seedling size traits which were consistently significant even after controlling the initial seed weight may indicate that the maternal effects reflect, at least to some extent, maternal genetic control over these traits. Despite overall strong maternal effects, the performance of maternal siblings, such as the rachis length, differed among blocks. The sensitivity of maternal siblings to the local environments may contribute to the maintenance of genetic variability in this highly outcrossing tropical species.     

The Combined Effects of Pre- and Post-Insemination Sexual Selection on Extreme Variation in Male Body Size

Orb-weaving spiders of the genus Nephila are notable for their sexual size dimorphism, with dwarf males and giant females. However, less well known is the extreme size polymorphism of males that is characteristic of some species. For example, adult male body size in N. edulis varies by an order of magnitude. Previous experiments reveal that male mating behaviour covaries with body size, suggesting the size variation is maintained by opposing pre- and post-insemination sexual selection pressures. Here, we test this idea by allowing males of different sizes to compete directly and simultaneously for access to females. Using the sterile-male technique for paternity assessment, we show that two competing males drawn from the extremes of size variations, split paternity the same way as two males of the same size drawn from the intermediate sizes. The paternity of a large male dropped from 50% to 30% on average if he competed against two instead of one small male. The large male increased his mating frequency when there were more rivals but required a much lower total duration of copulation to achieve the same paternity share. These data are consistent with the idea that opposing pre- and post-insemination selection pressures at least partly explain the variation in male body size. Co-ordinating editor: L.D. Hurst     

Effects of the estuarine dinoflagellate Pfiesteria shumwayae (Dinophyceae) on survival and grazing activity of several shellfish species

A series of experiments was conducted to examine effects of four strains of the estuarine dinoflagellate, Pfiesteria shumwayae, on the behavior and survival of larval and adult shellfish (bay scallop, Argopecten irradians; eastern oyster, Crassostrea virginica; northern quahogs, Mercenaria mercenaria; green mussels, Perna viridis [adults only]). In separate trials with larvae of A. irradians, C. virginica, and M. mercenaria, an aggressive predatory response of three strains of algal- and fish-fed P. shumwayae was observed (exception, algal-fed strain 1024C). Larval mortality resulted primarily from damage inflicted by physical attack of the flagellated cells, and secondarily from Pfiesteria toxin, as demonstrated in larval C. virginica exposed to P. shumwayae with versus without direct physical contact. Survival of adult shellfish and grazing activity depended upon the species and the cell density, strain, and nutritional history of P. shumwayae. No mortality of the four shellfish species was noted after 24 h of exposure to algal- or fish-fed P. shumwayae (strains 1024C, 1048C, and CCMP2089) in separate trials at ≤5 × 10³ cells ml⁻¹, whereas higher densities of fish-fed, but not algal-fed, populations (>7–8 × 10³ cells ml⁻¹) induced low (≤15%) but significant mortality. Adults of all four shellfish species sustained >90% mortality when exposed to fish-fed strain 270A1 (8 × 10³ cells ml⁻¹). In contrast, adult M. mercenaria and P. viridis exposed to a similar density of fish-fed strain 2172C sustained <15% mortality, and there was no mortality of A. irradians and C. virginica exposed to that strain. In mouse bioassays with tissue homogenates (adductor muscle, mantle, and whole animals) of A. irradians and M. mercenaria that had been exposed to P. shumwayae (three strains, separate trials), mice experienced several minutes of disorientation followed by recovery. Mice injected with tissue extracts from control animals fed cryptomonads showed no response. Grazing rates of adult shellfish on P. shumwayae (mean cell length ±1 standard error [S.E.], 9 ± 1 μm) generally were significantly lower when fed fish-fed (toxic) populations than when fed populations that previously had been maintained on algal prey, and grazing rates were highest with the nontoxic cryptomonad, Storeatula major (cell length 7 ± 1 μm). Abundant cysts of P. shumwayae were found in fecal strands of all shellfish species tested, and ≤45% of the feces produced viable flagellated cells when placed into favorable culture conditions. These findings were supported by a field study wherein fecal strands collected from field-collected adult shellfish (C. virginica, M. mercenaria, and ribbed mussels, Geukensia demissa) were confirmed to contain cysts of P. shumwayae, and these cysts produced fish-killing flagellated populations in standardized fish bioassays. Thus, predatory feeding by flagellated cells of P. shumwayae can adversely affect survival of larval bivalve molluscs, and grazing can be depressed when adult shellfish are fed P. shumwayae. The data suggest that P. shumwayae could affect recruitment of larval shellfish in estuaries and aquaculture facilities; shellfish can be adversely affected via reduced filtration rates; and adult shellfish may be vectors of toxic P. shumwayae when shellfish are transported from one geographic location to another.     

Effects of Exogenous Electromagnetic Fields on a Simplified Ion Channel Model

In this paper, we calculate the effect of an exogenous perturbation (an electromagnetic field [EMF] oscillating in the range of microwave frequencies in the range of 1 GHz) on the flux of two ion species through a cylindrical ion channel, implementing a continuous model, the Poisson–Smoluchowski system of equations, to study the dynamics of charged particle density inside the channel. The method was validated through comparison with Brownian dynamics simulations, supposed to be more accurate but computationally more demanding, obtaining a very good agreement. No EMF effects were observed for low field intensities below the level for thermal effects, as the highly viscous regime and the simplicity of the channel do not exhibit resonance phenomena. For high intensities of the external field (>10⁵ V/m), we observed slightly different behavior of ion concentration oscillations and ion currents as a function of EMF orientation with respect to the channel axis.     

Long-term impacts of goat browsing on bush-clump dynamics in a semi-arid subtropical savanna

The effects of 16 years of continuous browsing by goats in a South African savanna at stocking rates intended for bush control were compared with plots unbrowsed for the same period of time. Differences in bush-clump density, structure and species composition were recorded. Bush-clump density did not differ between browsed and unbrowsed plots. Within individual bush-clumps, browsing was shown to impact more on structure than species composition, with smaller, shorter bush-clumps, containing fewer species but much greater stem-densities. Although species presence/absence was little affected by browsing, many species showed differences in abundance, growth and location within browsed and unbrowsed bush-clumps. Species reduced in abundance in browsed plots included Cussonia spicata, Ehretia rigida, Grewia occidentalis, Jasminum angulare and Senecio linifolius. Several species showed reduced growth in browsed plots, particularly those located at bush-clump edges. The relatively unpreferred Aloe ferox was a notable exception. Although browsing had little effect on the composition of the main clump founding species, emergents or late arrivals, there were twice as many single plants in browsed plots and emergence of several species was restricted to the middle of bush-clumps. Comparison of our findings with aerial photographic evidence and other literature suggest that browsing alone is unlikely to significantly reduce scrub cover, although it can clearly control further expansion. Combinations of fire and browsing, rather than one factor alone, are considered likely to act fastest and most effectively to significantly reduce or remove scrub cover altogether.     

Carbon assimilation, nitrogen, and photochemical efficiency of different Himalayan tree species along an altitudinal gradient

In the area of Jumla region in Western Nepal, measurements of saturated leaf net photosynthetic rate (P_sat), nitrogen content, leaf fluorescence, carbon isotopic composition, and water status were performed on woody coniferous (Pinus wallichiana, Picea smithiana, Abies spectabilis, Juniperus wallichiana, Taxus baccata), evergreen (Quercus semecarpifolia, Rhododendron campanulatum), and deciduous broadleaved species (Betula utilis, Populus ciliata, Sorbus cuspidata) spreading from 2 400 m up to the treeline at 4 200 m a.s.l. With the exception of J. wallichiana, P_sat values were lower in coniferous than broadleaved species. Q. semecarpifolia, that in this area grows above the coniferous belt between 3 000 and 4 000 m, showed the highest P_sat at saturating irradiance and the highest leaf N content. This N content was higher and P_sat lower than those of evergreen oak species of tempe forests at middle and low altitudes. For all species, P_sat and N content were linearly correlated, but instantaneous nitrogen use efficiency was lower than values measured in lowland and temperate plant communities. The values of carbon isotopic composition, estimated by ₁₃C, showed the same range reported for temperate tree species. The ranking of ₁₃C values for the different tree types was conifers < evergreen broadleaved13C were found along the altitudinal gradient. Quantum yield of photochemistry at saturating irradiance, measured by leaf fluorescence (F/F_m), was highest in J. wallichiana and lowest in T. baccata. Overall, photochemical efficiency was more strongly related to species than to altitude. Interestingly, changes of .F/F_m along the altitudinal gradient correlated well with the reported altitudinal distribution of the species.This revised version was published online in March 2005 with corrections to the page numbers.     

Effects of competition and season on survival and maturation of young bank vole females

In territorial microtines intra-specific density dependent processes can limit the maturation of individuals during the summer of their birth. This may have demographic consequences by affecting the number and the age distribution of breeding individuals in the population. Little is known about this process on a community level, though populations of many northern microtine species fluctuate in synchrony and are known to interfere socially with each other. We experimentally studied the influence of the field vole Microtus agrestis on maturation, breeding, space use and survival of weanling bank voles, Clethrionomys glareolus. Two additive competition experiments on bank vole populations were conducted in large outdoor enclosures, half of them additionally housing a field vole population. In a mid-summer experiment low population density and absence of older breeding females minimised intra-specific competition. Survival was not affected by the presence of field voles. Season had a significant effect on both the probability of maturation and breeding of the weanlings. Competition with field voles significantly delayed breeding, and coupled with seasonal effects decreased the probability of breeding. In a late-summer experiment breeding and survival of bank vole weanlings were studied for three weeks as part of a high density breeding bank vole population. Weanlings did not mature at all nor were their space use and survival affected by the presence of field voles. Our results show that competition with other species can also have an impact on breeding of immatures. In an extreme seasonal environment, even a short delay of breeding may decrease survival chances of offspring. Seasonal and competition effects together may thus limit the contribution of year born females to reproductive output of the population. Other studies have shown that adult breeding bank voles suffer lower survival in the presence of field voles, but this study showed no survival effects on the weanlings. Thus it might be beneficial for weanlings to stay immature especially in the end of the breeding season and postpone reproduction to the next breeding season if densities of competing species are high.