Long‐term colonization ecology of forest‐dwelling species in a fragmented rural landscape – dispersal versus establishment

Species colonization in a new habitat patch is an efficiency indicator of biodiversity conservation. Colonization is a two‐step process of dispersal and establishment, characterized by the compatibility of plant traits with landscape structure and habitat conditions. Therefore, ecological trait profiling of specialist species is initially required to estimate the relative importance of colonization filters. Old planted parks best satisfy the criteria of a newly created and structurally matured habitat for forest‐dwelling plant species. We sampled species in 230 ancient deciduous forests (source habitat), 74 closed‐canopy manor parks (target habitats), 151 linear wooded habitats (landscape corridors), and 97 open habitats (isolating matrix) in Estonia. We defined two species groups of interest: forest (107 species) and corridor specialists (53 species). An extra group of open habitat specialists was extracted for trait scaling. Differing from expectations, forest specialists have high plasticity in reproduction mechanisms: smaller seeds, larger dispersules, complementary selfing ability, and diversity of dispersal vectors. Forest specialists are shorter, less nutrient‐demanding and mycorrhizal‐dependent, stress‐tolerant disturbance‐sensitive competitors, while corridor specialists are large‐seeded disturbance‐tolerant competitors. About 40% of species from local species pools have immigrated into parks. The historic forest area, establishment‐related traits, and stand quality enhance the colonization of forest specialists. The openness of landscape and mowing in the park facilitate corridor specialists. Species traits in parks vary between a forest and corridor specialist, except for earlier flowering and larger propagules. Forest species are not dispersal limited, but they continue to be limited by habitat properties even in the long term. Therefore, the shady parts of historic parks should be appreciated as important forest biodiversity‐enhancing landscape structures. The habitat quality of secondary stands can be improved by nurturing a heterogeneous shrub and tree layer, and modest herb layer management.     

Higher β‐ and γ‐diversity at species and genetic levels in headwaters than in mid‐order streams in Hydropsyche (Trichoptera)

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Fine‐scale spatiotemporal patterns of genetic variation reflect budding dispersal coupled with strong natal philopatry in a cooperatively breeding mammal

The relatedness structure of animal populations is thought to be a critically important factor underlying the evolution of mating systems and social behaviours. While previous work has shown that population structure is shaped by many biological processes, few studies have investigated how these factors vary over time. Consequently, we explored the fine‐scale spatiotemporal genetic structure of an intensively studied population of cooperatively breeding banded mongooses (Mungos mungo) over a 10‐year period. Overall population structure was strong (average F_ST = 0.129) but groups with spatially overlapping territories were not more genetically similar to one another than noncontiguous groups. Instead, genetic differentiation was associated with historical group‐fission (budding) events, with new groups diverging from their parent groups over time. Within groups, relatedness was high within but not between the sexes, although the latter increased over time since group formation due to group founders being replaced by philopatric young. This trend was not mirrored by a decrease in average offspring heterozygosity over time, suggesting that close inbreeding may often be avoided, even when immigration into established groups is virtually absent and opportunities for extra‐group matings are rare. Fine‐scale spatiotemporal population structure could have important implications in social species, where relatedness between interacting individuals is a vital component in the evolution of patterns of inbreeding avoidance, reproductive skew and kin‐selected helping and harming.     

Between‐patch natal dispersal declines with increasing natal patch size and distance to other patches in the endangered Southern Dunlin Calidris alpina schinzii

Natal dispersal has profound consequences for populations through the movement of individuals and genes. Habitat fragmentation reduces structural connectivity by decreasing patch size and increasing isolation, but understanding of how this impacts dispersal and the functional connectivity of landscapes is limited because many studies are constrained by the size of the study areas or sample sizes to accurately capture natal dispersal. We quantified natal dispersal probability and natal dispersal distances in a small migratory shorebird, the Southern Dunlin Calidris alpina schinzii, with data from two extensively monitored endangered metapopulations breeding in Sweden and Finland. In both metapopulations philopatry was strong, with individuals returning to or close to their natal patches more often than expected by chance, consistent with the patchy distribution of their breeding habitat. Dispersal probabilities were lower and dispersal distances were shorter in Sweden. These results provide a plausible explanation for the observed inbreeding and population decline of the Swedish population. The differences between Sweden and Finland were explained by patch‐specific differences. Between‐patch dispersal decreased with increasing natal patch size and distance to other patches. Our results suggest that reduced connectivity reduces movements of the philopatric Dunlin, making it vulnerable to the effects of inbreeding. Increasing connectivity between patches should thus be one of the main goals when planning future management. This may be facilitated by creating a network of suitably sized patches (20–100 ha), no more than 20 km apart from each other, from existing active patches that may work as stepping stones for movement, and by increasing nest success and pre‐fledging survival in small patches.     

Mother-offspring interactions do not affect natal dispersal in a small rodent

According to kin selection and inbreeding avoidance hypotheses,natal dispersal should be facultatively adjusted to balancingthe costs and benefits of mother–offspring interactions.In polygynous mammals, it is hypothesized that female offspringshould seek to avoid local resource competition with their mother,whereas male dispersal should be determined by inbreeding avoidance.We tested these hypotheses with a field experiment investigatingthe relationship between territory acquisition and mother'spresence in the root vole Microtus oeconomus. This species hasa flexible social system in which sisters' and mother's homeranges overlap substantially, whereas sons disperse to a greaterextent. Immature sibling voles aged 20 days were released for20 days together with an unrelated adult male in a 2-patch systemeither in the presence of their mother or in the presence ofan unrelated adult female. Offspring movements were not influencedby mother's presence, but offspring, especially females, avoidedthe patch occupied by the adult female irrespective of kinship.Offspring remaining in contact with their mother were reproductivelysuppressed at the middle, but not by the end, of the experimentalperiod. These results indicate that juvenile root voles adoptedan opportunistic settlement strategy where they avoided theadult female irrespective of kinship and inbreeding risks.     

Limited gene dispersal and spatial genetic structure as stabilizing factors in an ant‐plant mutualism

Comparative studies of the population genetics of closely associated species are necessary to properly understand the evolution of these relationships because gene flow between populations affects the partners' evolutionary potential at the local scale. As a consequence (at least for antagonistic interactions), asymmetries in the strength of the genetic structures of the partner populations can result in one partner having a co‐evolutionary advantage. Here, we assess the population genetic structure of partners engaged in a species‐specific and obligatory mutualism: the Neotropical ant‐plant, Hirtella physophora, and its ant associate, Allomerus decemarticulatus. Although the ant cannot complete its life cycle elsewhere than on H. physophora and the plant cannot live for long without the protection provided by A. decemarticulatus, these species also have antagonistic interactions: the ants have been shown to benefit from castrating their host plant and the plant is able to retaliate against too virulent ant colonies. We found similar short dispersal distances for both partners, resulting in the local transmission of the association and, thus, inbred populations in which too virulent castrating ants face the risk of local extinction due to the absence of H. physophora offspring. On the other hand, we show that the plant populations probably experienced greater gene flow than did the ant populations, thus enhancing the evolutionary potential of the plants. We conclude that such levels of spatial structure in the partners' populations can increase the stability of the mutualistic relationship. Indeed, the local transmission of the association enables partial alignments of the partners' interests, and population connectivity allows the plant retaliation mechanisms to be locally adapted to the castration behaviour of their symbionts.     

Within‐season changes in habitat use of forest‐dwelling boreal bats

Bats utilize forests as roosting sites and feeding areas. However, it has not been documented how bats utilize these habitats in the boreal zone with methods afforded by recent technological advances. Forest structure and management practices can create a variety of three‐dimensional habitats for organisms capable of flight, such as bats. Here, we study the presence of boreal bats in a forest forming a mosaic of different age classes, dominant tree species, canopy cover, soil fertility, and other environmental variables, throughout their active season in the summer using passive ultrasound detectors. Our results indicate a preference for mature forest by Eptesicus nilssonii and a pooled set of Myotis bats. Both groups of bats also showed temporal changes in their habitat use regarding forest age. In June and July, both groups occurred more often in mature than young forests, but from August onwards, the difference in occurrence became less evident in Myotis and disappeared completely in E. nilssonii. In addition, E. nilssonii was more often present in forests with low canopy cover, and its occurrence shifted from coniferous forests to deciduous forests during the season. The results reflect the within‐season dynamics of bat communities and their ability to utilize different types of forest as environmental conditions change. Yet, the results most importantly emphasize the importance of mature forests to bat diversity and the need to conserve such environments in the boreal zone.     

Assessing the effect of complex ground types on ground‐dwelling arthropod movements with video monitoring: Dealing with concealed movements under a layer of plant residues

Understanding the effect of ground types on foraging movements of ground‐dwelling arthropods is a key step to managing their spatial distribution as required for successful conservation biological control. Indeed, fine movements at the centimeter scale can strongly influence the foraging ability of pest predators. However, because radio frequency identification or harmonic tracking techniques are not yet suitable for small species and video tracking focuses on uniform and light backgrounds, foraging movements have rarely been studied in relation to ground types. We present a method to track a ground‐dwelling arthropod (the earwig Euborellia caraibea) at night, walking on two contrasted ground types: bare soil and soil partly covered with a stratum of banana plant residues allowing individuals to hide periodically. The tracking of individuals within these ground types was achieved by infrared light, tagging individuals, video treatments, and semi‐automatic cleaning of trajectories. We tested different procedures to obtain segments with identical durations to quantify speeds and sinuosities. These procedures were characterized by the junction time gap between trajectory fragments, the rediscretization time of trajectories, and whether or not to use interpolation to fill in missing points in the trajectories. Earwigs exhibited significantly slower and more sinuous movements on soil with banana plant residues than on bare soil. Long time gaps for trajectory junction, extended rediscretization times, and interpolation were complementary means to integrate concealed movements in the trajectories. The highest slowdown in plant residues was detected when the procedure could account for longer periods under the residues. These results suggest that earwigs spent a significant amount of time concealed by the residues. Additionally, the residues strongly decreased the earwigs'' movement. Since the technical solutions presented in this study are inexpensive, easy to set up, and replicate, they represent valuable contributions to the emerging field of video monitoring.     

Destination‐based seed dispersal homogenizes genetic structure of a tropical palm

As the dominant seed dispersal agents in many ecosystems, frugivorous animals profoundly impact gene movement and fine‐scale genetic structure of plants. Most frugivores engage in some form of destination‐based dispersal, in that they move seeds towards specific destinations, resulting in clumped distributions of seeds away from the source tree. Molecular analyses of dispersed seeds and seedlings suggest that destination‐based dispersal may often yield clusters of maternal genotypes and lead to pronounced local genetic structure. The long‐wattled umbrellabird Cephalopterus penduliger is a frugivorous bird whose lek mating system creates a species‐specific pattern of seed dispersal that can potentially be distinguished from background dispersal processes. We used this system to test how destination‐based dispersal by umbrellabirds into the lek affects gene movement and genetic structure of one of their preferred food sources Oenocarpus bataua, a canopy palm tree. Relative to background dispersal processes, umbrellabird mating behaviour yielded more diverse seed pools in leks that included on average five times more seed sources and a higher incidence of long‐distance dispersal events. This resulted in markedly lower fine‐scale spatial genetic structure among established seedlings in leks than background areas. These species‐specific impacts of destination‐based dispersal illustrate how detailed knowledge of disperser behaviour can elucidate the mechanistic link driving observed patterns of seed movement and genetic structure.     

Effects of individual condition and habitat quality on natal dispersal behaviour in a small rodent

1. Individuals should benefit from settling in high-quality habitats, but dispersers born under favourable conditions have a better physical condition and should therefore be more successful at settling in high-quality habitats. 2. We tested these predictions with root voles (Microtus oeconomus) by a manipulation of individual condition through litter-size enlargement and reduction during lactation combined with a manipulation of habitat quality through degradation of the vegetation cover. We accurately monitored movements of 149 juveniles during a settlement and breeding period of 3 months. 3. The litter size treatment had long-lasting effects on body size, life-history traits and home range size, but did not influence dispersal behaviour. 4. Different stages of dispersal were influenced by habitat quality. In low-quality patches, females dispersed earlier, spent more time prospecting their environment before settling, and settlers had a smaller adult body size than in high-quality patches. Preference and competition for high-quality patches is likely adaptive as it increased fitness both in terms of survival and reproduction. 5. We found no interactive effect of individual condition and habitat quality on natal dispersal and habitat selection. 6. These findings suggest that immediate conditions are more important determinants of dispersal decisions than conditions experienced early in life.     

Multiple environmental stressors increase the realised niche breadth of a forest‐dwelling fish

Understanding the determinants of species' niche breadth is important due to the negative relationship between niche breadth and extinction probability. Species tolerant to extremely harsh abiotic conditions (e.g. low pH or hypoxia) often have relatively small realised niches due to a trade‐off between abiotic and biotic stress tolerance, and are therefore particularly vulnerable to extinction. We hypothesised that tolerance to multiple extreme conditions (e.g. low pH, hypoxia and drought) in brown mudfish Neochanna apoda, would increase their realised niche breadth because each extreme would provide additive refuge against a dominant species, the banded kokopu Galaxias fasciatus. Fish distributions were surveyed in 65 peat‐swamp‐forest streams and pools in New Zealand, which varied in dissolved oxygen, acidity and hydroperiod. Mudfish distribution was extremely patchy, and almost completely allopatric with kokopu. Allopatry was driven mainly by pool hydroperiod; mudfish occupied 88 percent of temporary pools due to their tolerance of habitat drying, whereas kokopu were absent from temporary pools. Within permanent pools, mudfish occurrence was negatively related to pool oxygen and pH, whereas kokopu occurrence was positively related to these conditions. Pool conditions were independently distributed in the landscape, suggesting that each abiotic stressor offered additive refuge for mudfish from kokopu predation/competition. Consequently, the mudfish realised niche breadth depended on the number of abiotic factors driving their allopatry with kokopu. Given the widespread negative relationships between niche breadth and species extinction probabilities, our results indicate that tolerance to multiple stressors may play an important role in insuring species persistence against the multiple drivers of global environmental change.     

Animal movements in fire‐prone landscapes

Movement is a trait of fundamental importance in ecosystems subject to frequent disturbances, such as fire‐prone ecosystems. Despite this, the role of movement in facilitating responses to fire has received little attention. Herein, we consider how animal movement interacts with fire history to shape species distributions. We consider how fire affects movement between habitat patches of differing fire histories that occur across a range of spatial and temporal scales, from daily foraging bouts to infrequent dispersal events, and annual migrations. We review animal movements in response to the immediate and abrupt impacts of fire, and the longer‐term successional changes that fires set in train. We discuss how the novel threats of altered fire regimes, landscape fragmentation, and invasive species result in suboptimal movements that drive populations downwards. We then outline the types of data needed to study animal movements in relation to fire and novel threats, to hasten the integration of movement ecology and fire ecology. We conclude by outlining a research agenda for the integration of movement ecology and fire ecology by identifying key research questions that emerge from our synthesis of animal movements in fire‐prone ecosystems.     

Does nonnutritive tree gouging in a rainforest‐dwelling lemur convey resource ownership as does loud calling in a dry forest‐dwelling lemur?

Nonhuman primates may defend crucial resources using acoustic or chemical signals. When essential resources are limited, ownership display for a resource may be enhanced. Defending resources may depend on population density and habitat characteristics. Using the Milne Edwards' sportive lemurs (Lepilemur edwardsi) and weasel sportive lemurs (L. mustelinus) as models, we tested whether two cryptic nocturnal lemur species differing in population density and habitat show differences in their vocal and chemical communication for signaling ownership of resources. L. edwardsi inhabits a western dry deciduous forest in a high‐density population, whereas L. mustelinus is found in an eastern rainforest in low density. We followed ten L. edwardsi (six males and four females) and nine L. mustelinus (four males and five females) for 215 hr during the early evening (06:00–10:00 p.m.) and the early morning (02:00–05:00 a.m.) and recorded their behavior using focal animal sampling. We found that both species differed in their vocal and chemical communication. L. edwardsi was highly vocal and displayed loud calling in the mornings and evenings while feeding or in the vicinity of resting places. In contrast, L. mustelinus never vocalized during observations, but displayed tree‐gouging behavior that was never observed in L. edwardsi. Tree gouging occurred more often during early evening sessions than early morning sessions. Subjects gouged trees after leaving their sleeping hole and before moving around. We suggest that, in weasel sportive lemurs, non‐nutritive tree gouging is used as a scent‐marking behavior in order to display ownership of sleeping sites. Altogether, our findings provide first empirical evidence on the evolution of different communication systems in two cryptic nocturnal primate species contrasting in habitat quality and population density. Further investigations are needed to provide more insight into the underlying mechanisms. Am. J. Primatol. 72:1062–1072, 2010. © 2010 Wiley‐Liss, Inc.     

Female-biased natal and breeding dispersal in an alpine lizard, Niveoscincus microlepidotus

We measured two aspects of dispersal in the alpine Australian scincid lizard, Niveoscincus micolepidotus : (1) natal dispersal, i.e. shift in home range over the lizard's first year of life, and (2) breeding dispersal, i.e. shifts of home ranges between breeding attempts as adults. On average, displacements were surprisingly small. Female neonates dispersed about twice as far as did males in the same cohort (means of 12 m vs. 6 m). A female's natal dispersal distance was not correlated with her body size or our estimate of physiological performance (sprint speed). However, larger, faster-running male neonates dispersed further than did smaller, slower males. As was the case for neonates, adult females moved significantly further between breeding seasons than did adult males (14.2 m vs. 9.6 m). Because of a female's long gestation period (more than 1 year), two groups of females occur simultaneously in the population, non-ovulated (i.e. with yolking folicles) and pregnant females (i.e. approaching parturition). Females that were not yet ovulated showed a markedly stronger dispersal in response to high reproductive effort (i.e. clutch size in relation to body condition) than did pregnant females. In adult males, body size was negatively correlated with dispersal distance, suggesting that although males have overlapping territories, they exhibit an increasing level of site tenacity with age and/or size. Thus, selection for the relatively more pronounced site tenacity in adult males may have resulted in the more marked philopatric behaviour compared to females also as neonates.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 277–283.     

Limited male dispersal in a social spider with extreme inbreeding

Cooperatively breeding animals commonly avoid incestuous mating through pre-mating dispersal. However, a few group-living organisms, including the social spiders, have low pre-mating dispersal, intra-colony mating, and inbreeding. This results in limited gene flow among colonies and sub-structured populations. The social spiders also exhibit female-biased sex ratios because survival benefits to large colonies favour high group productivity, which selects against 1 : 1 sex ratios. Although propagule dispersal of mated females may occasionally bring about limited gene flow, little is known about the role of male dispersal. We assessed the extent of male movement between colonies in natural populations both experimentally and by studying colony sex ratios over the mating season. We show that males frequently move to neighbouring colonies, whereas only 4% of incipient nests were visited by dispersing males. Neighbouring colonies are genetically similar and movement within colony clusters does not contribute to gene flow. Post-mating sex ratio bias was high early in the mating season due to protandry, and also in colonies at the end of the season, suggesting that males remain in the colony when mated females have dispersed. Thus, male dispersal is unlikely to facilitate gene flow between different matrilineages. This is consistent with models of non-Fisherian group-level selection for the maintenance of female biased sex ratios, which predict the elimination of male dispersal.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society 2009, 97 , 227–234.     

Behavioral signature of intraspecific competition and density dependence in colony‐breeding marine predators

In populations of colony‐breeding marine animals, foraging around colonies can lead to intraspecific competition. This competition affects individual foraging behavior and can cause density‐dependent population growth. Where behavioral data are available, it may be possible to infer the mechanism of intraspecific competition. If these mechanics are understood, they can be used to predict the population‐level functional response resulting from the competition. Using satellite relocation and dive data, we studied the use of space and foraging behavior of juvenile and adult gray seals (Halichoerus grypus) from a large (over 200,000) and growing population breeding at Sable Island, Nova Scotia (44.0 ^oN 60.0 ^oW). These data were first analyzed using a behaviorally switching state‐space model to infer foraging areas followed by randomization analysis of foraging region overlap of competing age classes. Patterns of habitat use and behavioral time budgets indicate that young‐of‐year juveniles (YOY) were likely displaced from foraging areas near (<10 km) the breeding colony by adult females. This displacement was most pronounced in the summer. Additionally, our data suggest that YOY are less capable divers than adults and this limits the habitat available to them. However, other segregating mechanisms cannot be ruled out, and we discuss several alternate hypotheses. Mark–resight data indicate juveniles born between 1998 and 2002 have much reduced survivorship compared with cohorts born in the late 1980s, while adult survivorship has remained steady. Combined with behavioral observations, our data suggest YOY are losing an intraspecific competition between adults and juveniles, resulting in the currently observed decelerating logistic population growth. Competition theory predicts that intraspecific competition resulting in a clear losing competitor should cause compensatory population regulation. This functional response produces a smooth logistic growth curve as carrying capacity is approached, and is consistent with census data collected from this population over the past 50 years. The competitive mechanism causing compensatory regulation likely stems from the capital‐breeding life‐history strategy employed by gray seals. This strategy decouples reproductive success from resources available around breeding colonies and prevents females from competing with each other while young are dependent.