Pollination partners of Mucuna macrocarpa (Fabaceae) at the northern limit of its range

Mucuna macrocarpa is a plant found in tropical and subtropical regions that requires an “explosive opening.” Explosive opening is the process that exposes the stamen and pistil from the opening of the carina. This process is needed for cross pollination; however, the plant cannot open itself and opening by an animal is needed. The most common opener of Mucuna flowers is several nectar‐eating bats (e.g., Syconycteris), but the flying fox, Pteropus dasymallus, is the only opener of M. macrocarpa on the subtropical island of Okinawajima. Here, we present the explosive openers and possible pollinators in the northernmost and temperate Kamae region, Kyushu, Japan, where nectar‐eating bats are absent. The Japanese macaque, Macaca fuscata, and the Japanese marten, Martes melampus, were the explosive openers observed during our survey in Kamae. Martens opened flowers using their snout in a manner similar to that of the flying fox, whereas macaques opened flowers using their hands. This is the first time that an animal has been observed opening these flowers with its hands rather than snout. In total, 97% (n = 283) of explosively opened flowers were opened by macaques, and the macaque largely contributed to the overall flower opening. Because many pollen grains become attached to the explosive openers, they are considered to be primary pollinators. Furthermore, two bee species, Apis cerana japonica and Bombus ardens ardens, also visited opened flowers and collected pollen, and they were possibly secondary pollinators.     

Movements of lumpsucker females in a northern Norwegian fjord during the spawning season

The lumpsucker Cyclopterus lumpus is distributed throughout the North Atlantic Ocean and migrates considerable distances between offshore feeding areas and shallow inshore spawning grounds. The number of the lumpsucker has declined since the mid 1980s, probably as a result of overexploitation. The lumpsucker is the preferred host of the sea louse Caligus elongates, which is a problem for marine aquaculture. However, little is known about the biology of the lumpsucker. The aims of the study were to 1) examine the movements of female lumpsucker during the spawning migration, and 2) assess the potential for lumpsucker to act as a vector for transmission of parasites and diseases between aquaculture farms and wild fish. Twenty female lumpsuckers tagged with acoustic transmitters were released during the spawning season in the inner part of Øksfjord, northern Norway and their distribution was recorded by 22 automatic acoustic receivers. The average time until departure from the fjord was 3 days, and within 1 week all fish had left the fjord. Timing of departure from the fjord was unrelated with either tidal current patterns or the time of the day. A high proportion of the fish (75%) were recorded within 200 m of fish farms, but they did not stay for extended periods at these farms. Our results suggest that mature female lumpsucker exhibit a movement pattern characterized by rapid fjord-scale migrations during the spawning season, and that they are not attracted to salmon farms in the same way as a range of other fish species.     

Dynamics of a northern fulmar (Fulmarus glacialis) population at the southern limit of its range in Europe

In the context of global changes, defining the source–sink dynamics of populations of emblematic species, such as seabirds, within the limits of their distribution range is often crucial to optimize the priorities of surveys and conservation management, especially in protected areas. However, ringing is often not possible and only simple survey methods, such as the ‘apparently occupied sites’ method, can be utilized by managers of protected areas and threatened species. Using data collected between 1997 and 2005, we investigated the population dynamics of the northern fulmar (Fulmarus glacialis) at the southern limit of its range on the western French coast, which hosts increasing populations. Using a robust design spatial occupancy model, we estimated the proportion of occupied nests, the rates of nest colonization, nest extinction and the population growth rates of four colonies of the largest population (Ouessant Island). The estimated annual growth rate was high (average 1.049). A deterministic population dynamics model indicated that the observed rapid increase of Ouessant populations cannot be explained by their intrinsic dynamics, which suggested an important role for immigration. Different demographic scenarios provide several lines of evidence that the large northern fulmar population in this Man and Biosphere Reserve is a sink population. The strong increase in a population located at the limits of the species distributional range implies that it functions as a population sink. Inexpensive effective survey methods could allow investigation of the demographic status of seabird populations and provide relevant information for the hierarchization of conservation priorities.     

A preliminary study on habitat selection of the Japanese serow (Capricornis crispus) at two temporal scales,season and time of day,in a montane forest

Journal of Ethology - Habitat selection has a significant influence on animals’ fitness, and has been well studied in various ungulates. A trade-off between the access to forage and to...     

Recognition of a third callionymid species, Callionymus reticulatus Valenciennes 1837 (reticulated dragonet), in the south-eastern North Sea

During routine coastal Demersal Young Fish and Brown Shrimp Surveys (DYFS) in 2002, the reticulated dragonet Callionymus reticulatus was found in addition to C. lyra in the Wadden Sea of Schleswig Holstein, Germany, using a 3‐m experimental brown shrimp beam trawl. The survey area extended from 7 to 9°E and from 53°30′N to 54°55′N. It is the first proven occurrence of this species east of Helgoland in the Wadden Sea National Park area and adjacent waters. Data from two other beam trawl surveys (in May 2002 on sole and in January 2003 on shrimp) were additionally used to cover a wider area of the German Bight and during different seasons.     

Spreading to a limit: the time required for a neophyte to reach its maximum range

The average range size of alien plants in Spain reaches a maximum at 143 years. This is consistent with estimates of such a maximum in Ireland, Britain, Germany and the Czech Republic. A round figure of about 150 years on average for neophytes to reach their maximum range in European countries is indicated.     

Frontier mutualism: coevolutionary patterns at the northern range limit of the leaf-cutter ant-fungus symbiosis

Tropical leaf-cutter ants cultivate the fungus Attamyces bromatificus in a many-to-one, diffuse coevolutionary relationship where ant and fungal partners re-associate frequently over time. To evaluate whether ant-Attamyces coevolution is more specific (tighter) in peripheral populations, we characterized the host-specificities of Attamyces genotypes at their northern, subtropical range limits (southern USA, Mexico and Cuba). Population-genetic patterns of northern Attamyces reveal features that have so far not been observed in the diffusely coevolving, tropical ant-Attamyces associations. These unique features include (i) cases of one-to-one ant-Attamyces specialization that tighten coevolution at the northern frontier; (ii) distributions of genetically identical Attamyces clones over large areas (up to 81 000 km(2), approx. the area of Ireland, Austria or Panama); (iii) admixture rates between Attamyces lineages that appear lower in northern than in tropical populations; and (iv) long-distance gene flow of Attamyces across a dispersal barrier for leaf-cutter ants (ocean between mainland North America and Cuba). The latter suggests that Attamyces fungi may occasionally disperse independently of the ants, contrary to the traditional assumption that Attamyces fungi depend entirely on leaf-cutter queens for dispersal. Peripheral populations in Argentina or at mid-elevation sites in the Andes may reveal additional regional variants in ant-Attamyces coevolution. Studies of such populations are most likely to inform models of coextinctions of obligate mutualistic partners that are doubly stressed by habitat marginality and by environmental change.     

Atlantic bluefin tuna Thunnus thynnus feeding ecology in the northern Gulf of Mexico: a preliminary description of diet from the western Atlantic spawning grounds

A combination of stomach contents, nitrogen stable‐isotope and tissue C:N values are presented to demonstrate feeding activity of Atlantic bluefin tuna Thunnus thynnus on the Gulf of Mexico (GOMEX) spawning grounds. Diets include teleosts, cephalopods, crustaceans and a pelagic tunicate (Pyrosoma atlanticum). Results reveal the need to classify the GOMEX as a T. thynnus feeding ground.     

Living on the edge: Fig tree phenology at the northern range limit of monoecious Ficus in China

Fig trees (Ficus) are a species-rich group of mainly tropical and subtropical plants that are of ecological importance because of the large numbers of vertebrates that utilise their figs for food. Factors limiting their distributions to warmer regions are still poorly understood, but are likely to include factors linked to their specialised pollination biology, because each Ficus species is dependent on one or a small number of host-specific fig wasps (Agaonidae) for pollination. Adult fig wasps are short-lived, but some species are capable of dispersing extremely long distances to pollinate their hosts. Close to its northern range limit we investigated the phenology of Ficus virens, the monoecious fig tree that reaches furthest north in China. Relatively few trees produced any figs, and very few retained figs throughout the winter. Despite this, new crops produced in spring were pollinated, with seasonally migrant pollinators from plants growing further south the most likely pollen vectors. An inability to initiate new crops at low temperatures may limit the distribution of monoecious fig trees to warmer areas.     

Population at the edge: increased divergence but not inbreeding towards northern range limit in Acer campestre

Ecological conditions shape natural distribution of plants. Populations are denser in optimal habitats but become more fragmented in the areas of suboptimal environmental conditions. Usually, fragmentation increases towards the limits of species distribution. Fragmented populations are often characterised by decreased genetic variation, and this effect is frequent in peripheral populations, mostly due to the reduced effective population size. Interestingly, the genetic consequences of fragmentation seem to be relatively weak in forest trees. Using microsatellite markers, we assessed the impact of population fragmentation on the genetic structure of a European tree species Acer campestre. Within the study area, this medium-size wind-dispersed and insect-pollinated tree reveals a gradual decrease in population density towards the northern range limit. Over the distance of 150 km, we detected the significant decrease in allelic richness, heterozygosity as well as an increase in the rate of population divergence along with latitude. On the other hand, we failed to show that the observed patterns of genetic structure result from the variation in population densities. Moreover, inbreeding levels revealed no association with both density and geographic location, suggesting that pollen limitation does not occur, even at the range margin. As we showed that there is no difference in a dispersal scale between low- and high-density populations in the study species, we argue that the genetic structure is a result of postglacial recolonization. However, unlike many other forest trees, A. campestre showed the sharp latitudinal genetic pattern at a very restricted spatial scale. Limited dispersal and high fragmentation are likely the reasons.     

Breeding behaviour, home range and habitat selection in Rock Firefinches Lagonosticta sanguinodorsalis in the wet and dry season in central Nigeria

Knowledge of a species’ movement behaviour and habitat choice is a prerequisite for assessing its ecological requirements to plan successful conservation strategies. Little is known about these factors in the Rock Firefinch, a recently described species which is probably endemic to the Jos Plateau in central Nigeria. We investigated home range size, habitat use and breeding behaviour of the Rock Firefinch in Amurum forest reserve in central Nigeria during the wet (August–October) and the dry season (November–December) using radiotracking. Birds showed high site fidelity. They mainly moved alone or in pairs but did not have exclusive home ranges. Home range size tended to be larger during the dry season due to long movements to water sources. Birds generally preferred inselberg habitat and avoided farmland. During the dry season they additionally utilized gallery forests where water was readily available, and as a result of having to cross scrub savannah to get to water, scrub savannah was also more heavily used during the dry season. Birds bred between the late rainy (September–October) and the early dry season (November). Nest‐sites were associated with rocky boulders. Both sexes contributed to incubation. Daily egg survival rate calculated using the Mayfield method was 0.89 (0.83–0.95 95% confidence interval); no nests failed during the chick stage, but sample size was only four nests. Of all 14 nests found, 50% were depredated and only 29% of breeding attempts succeeded in producing Rock Firefinch chicks. A second breeding attempt was recorded when the first one failed. Chick production just about compensated adult mortality (measured in a separate study at the site) such that the population is probably stable at present. However, given the large uncertainty in our underlying assumptions, more data are needed to confirm this. We suggest that the presence of inselberg habitat in close proximity to water sources is the essential and limiting resource for this species.     

The genetic structure of the steppe plant Iris aphylla L. at the northern limit of its geographical range

Genetic diversity of Iris aphylla was examined in seven Polish populations located at the northern limit of the geographical range of the species. Using RAPD analysis, 491 DNA fragments were amplified, all of which were polymorphic. The peripheral populations were characterized by a low number of unique bands (one to three per population) and moderate values of genetic diversity parameters (percent of polymorphic loci = 30.6%, Shannon's index = 0.149 and Nei's gene diversity = 0.097, on average), which can be explained as being due to the recent expansion of the European biota and the extremely low frequency or absence of seed set and of juveniles, as well as limited pollen and seed dispersal. There was also no significant correlation between ecological properties (population size; number of flowering ramets and fruits) and genetic diversity parameters. Despite this fact, more fixed loci were noted in the smallest populations than were in the largest ones. All populations were significantly differentiated and geographically structured. Analysis of molecular variance between regions showed a low and nonsignificant value, but relatively high molecular variation was partitioned among and within populations. Only one group of south-eastern populations was structured according to isolation by distance pattern and neighbour-joining cluster analysis. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 245–255.     

Fig trees at the northern limit of their range: the distributions of cryptic pollinators indicate multiple glacial refugia

Climatic oscillations during the last few million years had well‐documented effects on the distributions and genomes of temperate plants and animals, but much less is known of their impacts on tropical and subtropical species. In contrast to Europe and North America, ice‐sheets did not cover most of China during glacial periods, and the effects of glacial cycles were less dramatic. Fig trees are a predominantly tropical group pollinated by host‐specific fig wasps. We employed partial mitochondrial COI (918 bp) and nuclear ITS2 (462 bp) gene sequences to investigate the genetic structure and demographic histories of the wasps that pollinate the subtropical Ficus pumila var. pumila in Southeastern China. Deep genetic divergence in both mitochondrial (7.2–11.6%) and nuclear genes (1.6–2.9%) indicates that three pollinator species are present and that they diverged about 4.72 and 6.00 Myr bp . This predates the Quaternary ice ages, but corresponds with the formation of the Taiwan Strait and uplifting of the Wuyi–Xianxia Mountains. The three pollinators have largely allopatric distribution patterns in China and display different postglacial demographic histories. Wiebesia spp. 1 and 2 occupy, respectively, the northern and southern regions of the mainland host range. Their populations both underwent significant postglacial spatial expansions, but at different times and at different rates. Wiebesia sp. 3 is largely restricted to northern islands and shows less evidence of recent population expansion. Their mainly allopatric distributions and different demographic histories are consistent with host plant postglacial expansion from three distinct refugia and suggest one mechanism whereby fig trees gain multiple pollinators.     

Assessing the climatic sensitivity of Douglas-fir at its northern range margins in British Columbia, Canada

Northern hemisphere tree species growing at their northern range margins may be particularly responsive to climate change and could provide important information regarding future broad-scale responses. We analyzed and compared tree-ring data between five Douglas-fir [Pseudotsuga menziesii var. glauca (Mirb.) Franco] populations growing at the species’ northernmost distribution in British Columbia, Canada, and five populations located 150 km to the southeast. We quantified climate–growth relationships using uni- and multivariate techniques at different temporal scales. Our data suggest that (1) even at its northernmost distributions, precipitation limits long-term mature Douglas-fir radial growth more than temperatures, (2) northernmost Douglas-fir populations are distinct from populations located further within the species’ range in terms of certain key short-term growth responses, and (3) northernmost Douglas-fir growth sensitivities to climate may be increasing over time. In the future, mature Douglas-fir productivity in the northern portion of its range may be primarily limited by precipitation, and responses may be strongest at the species’ range margins.     

Modelling the influence of change in fire regime on the local distribution of a Mediterranean pyrophytic plant species (Cistus salviifolius) at its northern range limit

Aims To assess the potential distribution of an obligate seeder and active pyrophyte, Cistus salviifolius, a vulnerable species in the Swiss Red List; to derive scenarios by changing the fire return interval; and to discuss the results from a conservation perspective. A more general aim is to assess the impact of fire as a natural factor influencing the vegetation of the southern slopes of the Alps. Location Alps, southern Switzerland. Methods Presence–absence data to fit the model were obtained from the most recent field mapping of C. salviifolius. The quantitative environmental predictors used in this study include topographic, climatic and disturbance (fire) predictors. Models were fitted by logistic regression and evaluated by jackknife and bootstrap approaches. Changes in fire regime were simulated by increasing the time‐return interval of fire (simulating longer periods without fire). Two scenarios were considered: no fire in the past 15 years; or in the past 35 years. Results Rock cover, slope, topographic position, potential evapotranspiration and time elapsed since the last fire were selected in the final model. The Nagelkerke R² of the model for C. salviifolius was 0.57 and the Jackknife area under the curve evaluation was 0.89. The bootstrap evaluation revealed model robustness. By increasing the return interval of fire by either up to 15 years, or 35 years, the modelled C. salviifolius population declined by 30–40%, respectively. Main conclusions Although fire plays a significant role, topography and rock cover appear to be the most important predictors, suggesting that the distribution of C. salviifolius in the southern Swiss Alps is closely related to the availability of supposedly competition‐free sites, such as emerging bedrock, ridge locations or steep slopes. Fire is more likely to play a secondary role in allowing C. salviifolius to extend its occurrence temporarily, by increasing germination rates and reducing the competition from surrounding vegetation. To maintain a viable dormant seed bank for C. salviifolius, conservation managers should consider carrying out vegetation clearing and managing wild fire propagation to reduce competition and ensure sufficient recruitment for this species.     

The northern geographic range limit of the intertidal limpet Collisella scabra: a test of performance, recruitment, and temperature hypotheses

A decline in abundance towards a species' range boundary is often interpreted as evidence of a decline in individual success, and is usually assumed to reflect a decline in suitable environmental conditions. Gradual declines towards high latitude range boundaries are frequently attributed to limitations on organismal tolerance of cold temperature. Rarely have these two assumptions been empirically tested. The intertidal gastropod Collisella scabra declines monotonically in abundance from 435 to <1 m^?2 over the northern 300 km of its geographic distribution. I examined temperature, adult performance (survival, growth, reproduction), and recruitment at five locations in this region of decline. Mortality ranged from 4.9 to 11.2% per month, but was highest at the lowest latitude study site. Growth rates ranged from 0 to 5.2 mm yr^?1, but were generally lower at lower latitude sites. Gonad development was high in the three populations examined, but the possibility of infrequent spawning at high latitude sites could not be excluded. Finally, a comparison of performance differences among populations with temperature revealed clear effects of temperature on both growth and mortality; however, the patterns were not consistent with a hypothesis of cold stress at the range boundary. Overall there was little evidence for either reduced performance or increasing cold stress in low density high latitude populations. Over the same 300 km, recruitment declined monotonically from a mean of six recruits per 625 cm² to less than one; suggesting that limitations on recruitment, rather that adult performance, are responsible for low abundance in marginal populations. Several hypotheses for the decline in recruitment are discussed in the paper and the most likely explanation appears to be an increase in the distance between populations at the range margin, reducing the chances that dispersing larvae find suitable habitat for settlement.     

Disappearance of a spring cohort in a population of the dragonet,Repomucenus valenciennei, with spring and autumn spawning peaks in Tokyo Bay, Japan

The occurrence of eggs, pelagic larvae and juveniles, and settled juveniles of the dragonetRepomucenus valenciennei in Tokyo Bay, Japan, were investigated by plankton net and bottom trawl surveys between September 1990 and September 1991. Eggs, and pelagic larvae and juveniles appeared from April to November (spring to autumn), peaking in both spring and autumn. From the temporal pattern of egg and pelagic fish occurrence, and pelagic duration reported elsewhere (ca. one month), settlement could be predicted as occurring from late spring to autumn. However, settled juveniles appeared from August to December, with an abrupt peak in November. Aging from daily increments in the otoliths of settled recruits in 1990 indicated that the latter comprised individuals which had hatched between mid-September and early November (i.e. autumn cohort), implying that individuals which had hatched in spring to summer (April to August) were not recruited. Benthic hypoxia occurs widely in Tokyo Bay, from June to October each year, particularly in the northern part, which is the main nursery area ofR. valenciennei. The timing of dissolved oxygen recovery, and appearance of settled fish coincided closely (i.e. November), indicating that summer hypoxic conditions prevented the settlement of the spring cohort and hence recruitment to the population.     

Habitat selection as a hierarchical spatial process: the green woodpecker at the northern edge of its distribution range

Habitat selection can be envisaged as a hierarchical spatial process, from choice of home range to choice of dietary item. The green woodpecker (Picus viridis) is described as being closely bound to cultivated land and deciduous forests, mainly due to its summer diet composed of ants (Hymenoptera: Formicidae) found on meadows and pastures. To explore possible responses of this woodpecker to recent changes in land use practice, we studied home ranges, feeding habitats and food selection of a marginal population (four radio-marked males and five females) in a 30,000-ha conifer-dominated landscape at the northern edge of its distribution range in south-central Scandinavia. We asked: (1) Is the green woodpecker confined to areas with cultivated land and deciduous forest? (2) If so, are important food items (ants) particularly abundant or exclusively found there? (3) Can clearcuts and young plantations substitute for cultivated land as feeding habitat? Home ranges (mean=100 ha) were invariably confined to the parts of the landscape that contained cultivated land (<1% of the total area). In summer, birds preferred to feed in cultivated land, presumably due to a higher overall biomass of ants compared to forest habitats. They avoided clearcuts, but preyed extensively upon soil-dwelling ants in young conifer stands (16–30 years old). We failed to find preferences for particular ant groups (Lasius niger and L. flavus) associated with cultivated land. The principal summer food was Serviformica, an ant group that was equally abundant in cultivated land and forest habitat. A positive correlation between ant body mass and a preference index suggests that the birds selected the larger ant species independent of habitat type. In winter, birds fed exclusively on mound-building Formica rufa-ants in closed-canopy, older forest stands. Our results indicate that the green woodpecker successfully utilizes young conifer plantations as feeding habitat. At a larger scale, we hypothesize that green woodpecker populations fail to establish in managed forest tracts, not because of food shortage, but because the landscapes lack cultivated land serving as a key stimulus encouraging individuals to settle.