Linking Temporal and Spatial Scales in the Study of an Argentine Ant Invasion

Our long-term study of an invasion of Argentine ants (Linepithema humile) in a 481-ha biological preserve in Northern California links multiple spatial and temporal scales. We have investigated, at local spatial scales of tens of meters, how nests spread and contract seasonally and from year to year. Microsatellite analysis shows population genetic structure on the scale of about 100 m. At the landscape scale, we have surveyed the spread and impact of Argentine ants in the biological preserve since 1993, and have found high variability both seasonally and from year to year. Here, we describe how seasonal patterns in nesting behavior at the local scale help to explain how the location of the invasion edge changes. Thus, the growth and spread of nests on the scale of tens of meters, from season to season, produce the dynamics of the invasion from year to year at the scale of hundreds of meters.     

Seed dispersal of the Korean pine,Pinus koraiensis,by the red squirrel,Sciurus vulgaris

Seed hoarding behavior of the red squirrel,Sciurus vulgaris, was studied in relation to the amount of dispersed seeds of the Korean pine,Pinus koraiensis, and the distribution of its seedlings. After removing a cone from a tree, squirrels sat on the ground and ripped off its cone scales before transporting it. A mean of 3.2 seeds were scatter-hoarded per hole. Of 7.7×10⁴ mature seeds produced in a 0.21 ha planted Korean pine forest, 22% were estimated to be directly eaten by four squirrels, 9% were hoarded by them in the pine forest and 65% were cached outside the forest. Squirrels rediscovered hoarded seeds frequently, until the ground was covered with snow, during the period from snow fall until seed germination the next spring, few hoarded seeds were utilized. Korean pine seedlings were found up to 600 m from their mother trees. Scatter-hoarding by squirrels extensively contributes to seed dispersal to places suitable for the regeneration of the Korean pine. The large size of the cone, the absciss-layer at the cone penduncle, the infrequent dehiscence of cone scales, the large and wingless seeds, and the thick seed-coats have probably all been specialized to facilitate utilization by GenusSciurus.     

Prevalence of coccidia parasites (Protozoa) in red squirrels ( Sciurus vulgaris): effects of host phenotype and environmental factors

We investigated the relative importance of environmental factors versus host phenotype in determining parasite prevalence in Eurasian red squirrels ( Sciurus vulgaris). One hundred and forty-three fecal samples of 116 different squirrels collected in 2000 and 2001 from five study areas in the Italian Alps, were examined for intestinal protozoans. Two species of Eimeria were present with a medium to high prevalence in both years and in all areas, while two other species were rare, occurring only in some areas and not in all years. Cryptosporidium parvum had a high prevalence in the two study areas of the Western Alps, while in the three areas of the Central Alps it was recorded only once. The prevalence of Eimeria sciurorum and C. parvum fluctuated in parallel with squirrel density, suggesting a possible correlation between the presence of these protozoans and host density. A gender effect on E. sciurorum prevalence at low density could be explained by different space use patterns and social organization of males and females. C. parvum occurred more frequently in young squirrels, suggesting an acquired immunity in adults, but age-related susceptibility was not found for eimerian species. The coccidian community was more similar within than between regions, and study area and year were key parameters in predicting coccidia infection. There was no evidence of competition between coccidian species, but one positive interaction between E. sciurorum and E. andrewsi was observed. Our results suggest that the effects of geographic region, area features, and year effects probably related to fluctuations in host population density, were more important than individual phenotypic host characteristics in structuring the coccidian assemblage and determining levels of parasite prevalence in red squirrel populations.     

Patterns of genetic diversity in the red squirrel (Sciurus vulgaris L.): Footprints of biogeographic history and artificial introductions

British S. vulgaris are classified as aseparate subspecies, S. v. leucourus, tomainland Europe. While S. vulgaris is notunder threat across most of its Eurasian range,in Britain, Ireland and Italy populations aredeclining, mainly due to the introduction ofthe American grey squirrel (S.carolinensis). In this study, we conducted anextensive survey of mitochondrial DNA variationin British S. vulgaris populations and apreliminary survey of continental Europeanpopulations. Our main aims were to determinethe extent to which any populations of S.vulgaris in Britain are partially or whollythe product of artificial translocation of redsquirrels from continental Europe, and whethercontinental population variation will provideinformation on post-glacial reafforestationpatterns in Europe. We found that the majorityof extant populations of British S.vulgaris are of continental ancestry, manywith a very recent (last 40 years) Scandinavianancestry. The Scandinavian haplotype hasrapidly become the most dominant innortheastern Britain, despite not appearing innorthern English populations until 1966. Thissuggests that these squirrels may have anadaptive advantage in the non-native sprucedominated conifer plantations of northernEngland. Our preliminary examination ofcontinental populations demonstrated that theyare sufficiently differentiated to allow aphylogeographic study of this species.     

Community level impacts of an ant invader and food mediated coexistence

Ant invasions exert a range of effects on recipient communities, from displacement of particular species to more complex community disruption. While species loss has been recorded for a number of invasion events, a little examined aspect of these invasions is the mechanisms for coexistence with resident ant species.The yellow crazy ant, Anoplolepis gracilipes (Smith), is considered one of the world’s worst ant invaders and has recently undergone rapid population growth in Tokelau. We surveyed the ground-dwelling ant fauna in two plots on each of five invaded and three uninvaded islands across two atolls in Tokelau to examine community characteristics of the ant fauna in areas with and without yellow crazy ants. We also used three types of food bait (tuna, jam and peanut butter) to experimentally test if species are able to coexist by consuming different food resources. Anoplolepis gracilipes was found to coexist with two to six other ant species at any one site, and coexisted with a total of 11 ant species. Four species never co-occurred with A. gracilipes. Non-metric multidimensional scaling showed significant differences in community composition and the relative abundance of species between areas that had, and had not, been invaded by A. gracilipes. The number of other ant species was significantly lower in communities invaded by the yellow crazy ant, but did not decline with increasing A. gracilipes abundance, indicating that impacts were independent of population density. The yellow crazy ant dominated all tuna and jam baits, but had a low attendance on peanut butter, allowing four other ant species to access this resource. Our results demonstrate community level impacts of an ant invader on a tropical oceanic atoll and suggest that differing use of food resources can facilitate coexistence in ant communities. Received 11 September 2006; revised 15 January 2007; accepted 22 February 2007.     

Origin of, and conservation units in, the Irish red squirrel (Sciurus vulgaris) population

Knowledge of genetic relationships among wildlife populations is fundamental to their conservation, particularly where translocations are concerned. This study involved a survey of mitochondrial DNA variation in the Irish red squirrel population. Our main aims were: (1) to determine whether the Irish red squirrel population is distinct from that found in Britain, given known translocations that took place from Britain in the 1800’s; and (2) whether inclusion of Irish data into a reanalysis of European red squirrel data could reveal patterns of postglacial spread in Ireland. We found evidence that the current Irish red squirrel population may be a mixture of native and translocated stock, and relationships between Irish and European haplotypes supported a number of colonisation events of the island. Although only one haplotype was common to both Ireland and Britain, it is probable that the most common haplotypes in Ireland are British introductions that have since become extinct in Britain. There was a significant regional genetic structure in Ireland (P < 0.001), as well as between all Irish and British regions. Although it is likely that the red squirrel will not be fundamental in tracing the colonisation of Ireland by mammals, the data demonstrated that individual regions within Ireland, as well as the Irish population as a whole, are distinct both from the British population and from each other and, therefore, these populations should be treated as separate Management Units (MU) in conservation strategies.     

Food Shortage Disrupts Social Organization: The Case of Red Squirrels in Conifer Forests

A recent conceptual model of spatial organization in vertebrates, based upon changes in home range overlap with habitat quality, ‘the space-use model’, predicts large and strongly overlapping home ranges and absence of territorial behaviour in habitats with poor food availability. We investigated whether the model can be extended to predict intra-population variation in space-use in a habitat with strong temporal variation in resource abundance. We studied space use of Eurasian red squirrels (Sciurus vulgaris) in subalpine conifer forests over 4 years with strong fluctuations in seed production. Norway spruce dominated forests are resource limited habitats for squirrels, and tree seed availability is gradually depleted in the year following a poor seed-crop. Male and female squirrels responded to spruce seed-crop failure in 2000 by strongly increasing their home ranges and core-areas in the following year (mean MCP ± SE in summer and autumn, respectively: 2001, 122.9±14.1 ha and 84.3±9.4 ha, against 43.0±6.4 ha and 20.8±3.8 ha in 2002, and 22.7±1.5 ha and 21.4±2.0 ha in 2003). In 2001, half of the animals, those with the poorest quality pre-dispersal home ranges, emigrated to areas with more larch. Residents had multi-nuclear core-areas. Also, intra-sexual core-area overlap (males by males, females by females) was higher in 2001 and summer 2002, than in autumn 2002 and in 2003 (means ± SE in summer and autumn, respectively: males by males, 2001, 52±8 and 44±7%, 2002, 80±30 and 15±11%, 2003, 37±19 and 26±14%; females by females 2001, 52±10 and 112±32%, 2002, 55±27 and 0±0%, 2003, 12±8 and 0.1±0%). Red squirrels responded to food shortage by moving to patches with other food resources and abandoned the spacing pattern of reduced core-area overlap among males and nearly exclusive core-areas among females, found in less variable habitats. After richer seed-crops in 2001 and 2002, it took squirrels about a year to reduce the size of their home ranges and core-areas and return to a spacing pattern of stable home ranges and intra-sexual territoriality of adult females. These results are consistent with the space-use model and show that spacing behaviour in red squirrels is a plastic, conditional strategy with individuals adapting the size and/or location of their home ranges in relation to local distribution and abundance of food resources.Co-ordinating editor: R. Härdling     

Escape from Gregarine Parasites Affects the Competitive Interactions of an Invasive Mosquito

When a species is introduced into a new location, it may escape, at least temporarily, from its natural enemies. In field surveys, we found that when the exotic, invasive mosquito, Aedes albopictus, invades new sites, it initially experiences reduced infection by its gut parasite, Ascogregarina taiwanensis. To determine the effect of this escape from parasitism on the competitive ability of A. albopictus, we performed a laboratory competition experiment in which infected and uninfected A. albopictus larvae were reared in microcosms alone and in competition with larvae of the native mosquito, Ochlerotatus triseriatus. We analyzed the effect of parasitism by A. taiwanensis on A. albopictus performance when subjected to intra- and interspecific competition across a range of larval densities, as well as the effect of A. albopictus parasitism by A. taiwanensis on the competitive impact of A. albopictus on O. triseriatus. At a density of 30 O. triseriatus larvae, O. triseriatus survivorship was significantly reduced by the addition of 30 unifected A. albopictus, but not by addition of 30 infected A. albopictus, and not by addition of 15 A. albopictus whether infected or uninfected. Although estimated finite rate of population increase (') showed similar trends, and was significantly affected by treatments, no pairwise differences in rate of increase were significant. Infection by A. taiwanensis also significantly prolonged A. albopictus female development time and reduced the intraspecific competitive effect of increased density of A. albopictus, but did not affect A. albopictus survivorship, mass, or estimated finite rate of population increase. Thus, when A. albopictus escapes from this parasite as it colonizes new sites, this escape may give it a small, but significant, added competitive advantage over O. triseriatus, which may facilitate range expansion of A. albopictus and enhance A. albopictus's initial impact on resident species.     

Discovery of a life history shift: precocious flowering in an introduced population of Prosopis

The genus Prosopis contains many valuable, long-lived, multi-purpose legume trees, some of which are also invasive species. The time of first flowering is important for increasing production of sweet, protein-rich pods in plantations, especially under short rotations, and affects the rate of spread as a weed. Trees generally begin flowering at 3–5 years of age. However, seedlings from a small seedlot collected from an introduced population in southern Mauritania were observed to begin flowering at 3–4 months after germination. This is unknown in the native range of any Prosopis species, and appears to represent a major evolutionary event triggered after naturalization. This paper reports a detailed investigation with seed collected from Aleg, Brakna region, Mauritania, in 1998 (Prosopis sp. ‘Aleg’). Two experiments were established, in glasshouses at Cirad, Montpellier, France, and Coventry University, UK. Flowering began at the two sites 104 and 169 days after sowing, respectively, and 97.5% of plants had flowered at Coventry after 306 days. Flowers produced abundant pollen with 50–60% viability indicated by FDA staining and 40–50% germination on an artificial medium. No pods were formed. All plants analysed were diploid. Morphologically, these juvenile plants were similar to American species of section Algarobia, and exhibited leaf characters typical of P. pallida. However, due to the large variation in morphology within this section of the genus, further studies are required to confirm the actual species. This is the first record of precocious flowering in Prosopis, and has been found only in a single, introduced population. Implications for genetic improvement of tropical Prosopis, and particularly the spread of these species as invasive weeds, are discussed. Such a life history shift in the increased reproductive ability of a species following introduction, with potentially significant environmental effects, may not be restricted to Prosopis and merits further detailed investigation.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

From a Howling Wilderness to Howling Safaris: Science,Policy and Red Wolves in the American South

This article details the complex natural and cultural history of red wolf (Canis rufus) restoration in the American South. The decisions and methods utilized in the red wolf’s recovery after 1960 were unprecedented and creative but not geographically limited. The federal red wolf recovery experiment highlights the debate over what constitutes a species in a dynamic world, and the practical challenges and unexpected results in endangered species management in peopled landscapes. This wildlife restoration story illustrates not only the “hands-on” management role humans played, and continue to play, but also reveals cultural assumptions about what constitutes a “wild” wolf and about the necessity of wilderness. The red wolf recovery project provides constructive lessons for future species restoration involving flora and fauna on public and private land, and demonstrates human and animal engagement in the making of nature and culture.     

Spatial Dynamics of Lyme Disease: A Review

Lyme disease (LD), the most frequently reported vector-borne disease in the United States, requires that humans, infected vector ticks, and infected hosts all occur in close spatial proximity. Understanding the spatial dynamics of LD requires an understanding of the spatial determinants of each of these organisms. We review the literature on spatial patterns and environmental correlates of human cases of LD and the vector ticks, Ixodes scapularis in the northeastern and midwestern United States and Ixodes pacificus in the western United States. The results of this review highlight a need for a more standardized and comprehensive approach to studying the spatial dynamics of the LD system. Specifically, we found that the only environmental variable consistently associated with increased LD risk and incidence was the presence of forests. However, the reasons why some forests are associated with higher risk and incidence than others are still poorly understood. We suspect that the discordance among studies is due, in part, to the rapid developments in both conceptual and technological aspects of spatial ecology hastening the obsolescence of earlier approaches. Significant progress in identifying the determinants of spatial variation in LD risk and incidence requires that: (1) existing knowledge of the biology of the individual components of each LD system is utilized in the development of spatial models; (2) spatial data are collected over longer periods of time; (3) data collection and analysis among regions are more standardized; and (4) the effect of the same environmental variables is tested at multiple spatial scales.     

Intraspecific Phylogeography of Red Squirrels (Tamiasciurus hudsonicus) in the Central Rocky Mountain Region of North America

We used variation in a portion of the mitochondrial DNA control region to examine phylogeography of Tamiasciurus hudsonicus, a boreal-adapted small mammal in the central Rocky Mountain region. AMOVA revealed that 65.66% of genetic diversity was attributable to variation within populations, 16.93% to variation among populations on different mountain ranges, and 17.41% to variation among populations within mountain ranges. Nested clade analysis revealed two major clades that likely diverged in allopatry during the Pleistocene: a southern clade from southern Colorado and a northern clade comprising northern Colorado, Wyoming, eastern Utah, and eastern Idaho. Historically restricted gene flow as a result of geographic barriers was indicated between populations on opposite sides of the Green River and Wyoming Basin and among populations in eastern Wyoming. In some instances genetic structure indicated isolation by distance.     

Temporal and Spatial Expression of Hoxa-2 During Murine Palatogenesis

1. Mice homozygous for a targeted mutation of the Hoxa-2 gene are born with a bilateral cleft of the secondary palate associated with multiple head and cranial anomalies and these animals die within 24 hr of birth (Gendron-Maguire et al., 1993; Rijli et al., 1993; Mallo and Gridley, 1996). We have determined the spatial and temporal expression of the Hoxa-2 homeobox protein in the developing mouse palate at embryonic stages E12, E13, E13.5, E14, E14.5, and E15.2. Hoxa-2 is expressed in the mesenchyme and epithelial cells of the palate at E12, but is progressively restricted to the tips of the growing palatal shelves at E13.3. By the E13.5 stage of development, Hoxa-2 protein was found to be expressed throughout the palatal shelf. These observations correlate with palatal shelf orientation and Hoxa-2 protein may play a direct or indirect role in guiding the palatal shelves vertically along side the tongue, starting with the tips of the palatal shelves at E13, followed by the entire palatal shelf at E13.5.4. As development progresses to E14, the stage at which shelf elevation occurs, Hoxa-2 protein is downregulated in the palatal mesenchyme but remains in the medial edge epithelium. Expression of Hoxa-2 continues in the medial edge epithelium until the fusion of opposing palatal shelves.5. By the E15 stage of development, Hoxa-2 is downregulated in the palate and expression is localized in the nasal and oral epithelia.6. In an animal model of phenytoin-induced cleft palate, we report that Hoxa-2 mRNA and protein expression were significantly decreased, implicating a possible functional role of the Hoxa-2 gene in the development of phenytoin-induced cleft palate.7. A recent report by Barrow and Capecchi (1999), has illustrated the importance of tongue posture during palatal shelf closure in Hoxa-2 mutant mice. This along with our new findings of the expression of the Hoxa-2 protein during palatogenesis has shed some light on the putative role of this gene in palate development.     

松鼠几项生态生理指标季节变化的比较

研究了分布于我国大、小兴安岭林区松鼠体重、身体热值、脂肪含量、灰分含量以及水分含量的季节变化。松鼠体重存在明显的季节性变化和性别差异 ,春季 ,雄性松鼠体重明显小于雌性个体 ,而其它季节 ,雄性大于雌性。冬季体重最低(3 3 1 .9～ 3 45 .1 g) ,秋季体重最高 (3 78.1～ 3 89.9g) ,体重降低了 9.41 %～ 1 4.0 7%。大、小兴安岭松鼠身体热值、脂肪含量和水分含量季节变动范围为 6.2 71～ 8.60 9k J/g和 6.891～ 8.41 3 k J/g,8.81 %～ 2 1 .3 7%和 9.47%～ 2 2 .0 1 % ,67.1 2 %～73 .3 8%和 66.3 5 %～ 71 .73 % ;灰分含量不存在季节差异 ,大、小兴安岭年平均值分别为 1 2 .0 4%± 1 .1 7%和 1 1 .2 7%±2 .0 7%。环境温度、食物资源和繁殖状况是影响东北松鼠体重和身体组成季节性差异的主要因素     

Ontogenetic Variation in the Diurnal Food and Habitat Associations of an Endemic and an Exotic Fish in Floodplain Ponds: Consequences for Niche Partitioning

In floodplain ponds with low piscivore abundance, both endemic Midgley's gudgeons, Hypseleotris sp. 5, and exotic mosquitofish, Gambusia holbrooki, showed significant ontogenetic variation in the use of food and space. Small gudgeons were generally associated with surface and benthic habitats, then restricted their distribution to benthic habitats at a size of approximately 24mm (standard length). The ontogenetic variation in mosquitofish habitat use was less discrete, and could be described as a tendency for larger individuals to be associated with the bottom of the littoral macrophyte beds than with the surface of the macrophyte beds or surface of the limnetic zone. Small gudgeons exhibited high spatial overlap with mosquitofish within the surface habitats of the ponds. All size-class/species comparisons showed significant partitioning of food resources, however, the diets of small gudgeons and mosquitofish were very similar. Therefore, juvenile gudgeons may have to pass through a similar spatial and trophic niche to introduced mosquitofish before recruiting to the adult stage. Possible mechanisms driving the ontogenetic variation in gudgeon and mosquitofish habitat use are discussed. This paper demonstrates that ontogenetic niche shifts at fine spatial scales can affect our interpretation of interactions between native and introduced fishes.     

The Distribution of an Introduced Mollusc and its Role in the Long-term Demise of a Native Confamilial Species

The estuarine mud snail, Batillaria attramentaria, was transported to the Pacific coast of North America with the Pacific oyster, Crassostrea gigas (C. gigas), imported from Japan in the early part of this century, and has proliferated in several bays where the oyster was introduced. Since the arrival of Batillaria there have been declines in populations of its native ecological equivalent, Cerithidea californica. This study documents the distribution of the exotic Batillaria throughout its entire introduced range, concentrating on the few bays in northern California where both snails exist sympatrically. Using dates of initial importation of C. gigas and dates of first documentation of Batillaria within a bay, I established the earliest possible date for the introduction of Batillaria in a particular area. In cases where Cerithidea also was, or had been, present within a bay, I calculated a range of time for either the continued coexistence of the two species or the time until local exclusion of the native. Density measurements of Cerithidea within these bays where the species co-occurred allowed comparison of present Cerithidea numbers to historical accounts. Results indicated that Batillaria is replacing Cerithidea in the northern marshes of California. This replacement of the northernmost Cerithidea populations is not only reducing Cerithidea's overall range, but also eliminating a race recently shown to be a genetically distinct from southern Cerithidea populations. Other studies that have demonstrated superior exploitative competitive ability by Batillaria provide a potential mechanism for this displacement pattern. Regardless, the results presented here indicate that the displacement process is slow, taking on average >50 years to complete. This study illustrates a gradual, but predictable process of exotic replacement of native species, and argues strongly against complacency toward invaders that may currently seem innocuous. This revised version was published online in July 2006 with corrections to the Cover Date.     

Spatial and temporal pattern of introduced Bombus terrestris abundance in Hokkaido, Japan, and its potential impact on native bumblebees

A commercial colony of Bombus terrestris (L.) was introduced to Japan in 1992 for crop pollination in greenhouses. Since then wild colonies have developed and spread in some regions. In the present study, we measured the spatial distribution and temporal change in abundance of B. terrestris in the Chitose River Basin, Hokkaido, Japan to elucidate the relation of greenhouses to the bees distribution and to evaluate its potential effects on native bumblebees. Bumblebees were collected with window traps in windbreak forests roughly 1, 2, 4, and 6 km NNW and SSE of a large greenhouse. The peak catch of B. terrestris queens occurred in early June, suggesting that they had successfully hibernated in the field. The distributions of B. terrestris and the native B. ardens were mutually exclusive, while the native B. hypocrita appeared at all sites. Catches of B. terrestris were restricted to within 4 km of the nearest greenhouse, suggesting that the invasion was still in the initial phase in this area. The reduction in abundance of the native bumblebees in the sites of high B. terrestris abundance suggests the presence of interspecies competition between B. terrestris and the native bumblebees during the early part of the colony activity, although such reduction in B. ardens can be explained by habitat suitability.     

Apparent facilitation of an invasive mealybug by an invasive ant

Summary In the southeast United States, the invasive ant Solenopsis invicta is known to derive important carbohydrate (honeydew) resources from mealybugs utilizing grasses. Most important appears to be an invasive mealybug, Antonina graminis. We studied whether this mealybug and a similar native species also benefit from association with S. invicta. We found that mealybug occurrence increases significantly with increasing proximity to S. invicta mounds, suggesting that mealybugs benefit as well. Mutual benefits derived by S. invicta and A. graminis are consistent with a hypothesis proposing that associations among invasive species can be important in their success at introduced locations.