Mass mortality of native anuran tadpoles in tropical Australia due to the invasive cane toad (Bufo marinus)

Specific pathways of the ecological impact of invasive species remain poorly known. Although the spread of toxic cane toads (Bufo marinus) through tropical Australia is widely believed to have caused extensive mortality of native reptiles and mammals, effects of toad ingestion on native anurans have been virtually ignored. Our studies on the Adelaide River floodplain show that the most numerous vertebrate victims of toad invasion are native tadpoles that die when they attempt to consume toad eggs. We documented 11 episodes of mass mortality, totalling >1300 tadpoles of 10 species, in five waterbodies within a single wet-season shortly after the toads invaded. A causal link between toad breeding and tadpole mortality is supported by observations that: (1) in at least 9 of the 11 waterbodies involved, toads bred immediately prior to mortality events; (2) water quality was indistinguishable from that of control ponds, and tadpoles placed in that water remained healthy; (3) dead tadpoles showed no sign of disease; and (4) laboratory trials showed rapid, 100% mortality in native tadpoles exposed to freshly-laid toad eggs. Despite these high mortality rates, toad invasion does not appear to threaten the viability of anuran populations because frogs often breed in ponds not used by toads, and because density-dependent growth and survival within tadpole communities mean that additional mortality may not reduce the total effective recruitment of metamorph frogs from a waterbody.     

Frogs under friendly fire: How accurately can the general public recognize invasive species?

Members of the general public interact with wildlife in many ways, and an inability to distinguish between species can have significant implications for conservation. For example, attempts by environmentally-concerned private citizens to control invasive species may cause collateral damage unless people can reliably distinguish native fauna from the invader. We tested the Australian public’s ability to distinguish invasive cane toads (Bufo marinus) from native frogs at egg, tadpole, subadult and adult life-history stages. Errors were common, especially for eggs and tadpoles (27-31% error rates) and less so for subadult and adult toads and frogs (5-43% error rates). Accuracy of identification was higher in people living in areas where toads occur (compared to other parts of Australia or overseas), and similar in men and women (but with a decrease in older men). Ability to identify anurans was increased by toad-identification awareness programs and membership of “toad-busting” community groups, but direct killing of cane toads by the general public may inflict substantial “friendly fire” on native frogs. In the absence of any clear evidence for ecological benefits from toad-killing, we suggest that such collecting activities should be conducted only in areas where toads are known to occur, and under the supervision of trained personnel (to identify any anuran before it is killed), rather than as an ad hoc activity pursued independently by local residents. More generally, conservation activities that involve public participation should carefully evaluate the potential rates and consequences of species misidentification by well-intentioned but untrained people.     

Effects of nonnative fish and habitat characteristics on lentic herpetofauna in Yosemite National Park, USA

Nonnative fishes have been introduced into naturally fishless mountain lakes worldwide, often with negative consequences for native fauna. In this study, I used data collected during a census of native herpetofauna, nonnative trout, and habitat characteristics at all lentic water bodies in Yosemite National Park (n=2655) to quantify the effect of trout introductions and habitat on the distribution of four amphibian species and two reptile species. Impacts of trout on amphibians and reptiles were characterized using generalized additive models that included as predictor variables fish presence/absence, amphibian presence/absence (only in models for the two reptile species), elevation, water depth, littoral zone and shoreline substrate composition, relative survey date, and location. All species showed significant associations with habitat characteristics, and elevation and water depth appeared particularly important in influencing distributions. In addition, distributions of the mountain yellow-legged frog (Rana muscosa) and Pacific treefrog (Hyla regilla) were strongly negatively associated with the presence of nonnative trout while those of the Yosemite toad (Bufo canorus) and Sierra newt (Taricha torosa sierrae) were unrelated to trout presence. The distribution of the mountain garter snake (Thamnophis elegans elegans) was strongly negatively associated with the presence of nonnative trout and positively associated with the presence of native amphibians. Regression results for the Sierra garter snake (Thamnophis couchi couchi) were similar except that the direct effect of nonnative trout was considerably weaker. Together, these results indicate that trout introductions have resulted in considerable alteration of Yosemite's herpetofauna. Long-term studies will be necessary to determine whether removal of nonnative trout populations, where possible, would allow these impacts to be partially reversed.     

Habitat and flower resource partitioning by an exotic and three native bumble bees in central Hokkaido, Japan

Studies of the interspecific relationships between alien and native pollinator species can help forecast the success of alien species as well as assess the extent of disturbance to native plant-pollinator interactions. We examined the habitat and flower resource occupancy by the invasive introduced bumble bee Bombus terrestris and three dominant native bumble bees in central Hokkaido, northern Japan, in relation to a landscape factor (forest cover ratio) and flower morphology with respect to the proboscis length of bees. Three years of monitoring the invasive boundary of B. terrestris indicated that this species, which dominates open agricultural areas, probably will not invade the forests in which the native species dominate. This habitat partitioning likely followed the displacement of the natives by the invader in open agricultural lands. In forested areas, the native species partitioned flower resources on the basis of the relationship between proboscis length and the lengths of the corolla tubes of flowers. However, in open agricultural areas, both the long- and short-tubed flowers were primarily visited by the exotic short-tongued B. terrestris, which foraged illegitimately (by nectar robbing) on long-tubed flowers (Trifolium pratense L.) and legitimately on short-tubed flowers (T. repens L. and Lavandula angustifolia Mill.). The invasion of B. terrestris into open natural vegetation, in addition to open agricultural areas, has recently been reported in Hokkaido. Even though the exotic and dominant native bumble bees partition their habitat according to landscape factors, the invasive bee has the potential to alter the overall interactions within plant-pollinator systems in the regions of open vegetation on Hokkaido Island.     

Impacts of sediment loads, tadpole density, and food type on the growth and development of tadpoles of the spotted tree frog Litoria spenceri: an in-stream experiment

The impact of increased-deposited sediment loads on growth and development of tadpoles of the spotted tree frog, Litoria spenceri, in south-eastern Australia was assessed in conjunction with food type and tadpole density. This investigation was conducted in stream enclosures subjected to varying treatments of tadpole density, substrata and deposited-sediment load. Tadpole growth and development were not significantly affected by food type, but there was a trend toward increased performance on periphyton substrata. Tadpole growth and development were significantly reduced by increases in tadpole density and sediment load. These effects were independent, suggesting that increased sediment load reduced food quality rather than quantity. Activities in catchments that increase sediment loads in streams may adversely affect growth and development of tadpoles of L. spenceri. Increased sediment input into streams may therefore reduce tadpole fitness and recruitment to the terrestrial stage. Disturbance processes that increase stream sediment loads, such as forestry activities and associated road construction, may potentially have contributed to the observed declines of L. spenceri and other lotic anurans in south-eastern Australia.     

Pollination of the invasive exotic shrub Lupinus arboreus (Fabaceae) by introduced bees in Tasmania

Exotic plant invasions threaten ecological communities world-wide. Some species are limited by a lack of suitable pollinators, but the introduction of exotic pollinators can facilitate rapid spread. In Tasmania, where many non-native plants are naturalised, exotic honeybees (Apis mellifera) and bumblebees (Bombus terrestris) have become established. We determined how these species affect the pollination of Lupinus arboreus, an invasive, nitrogen-fixing shrub, which is rarely visited by native pollinators. The proportion of flowers setting seed and the number of ovules fertilised per flower were positively related to the visitation rates of both exotic bee species. There was no effect of bee visitation rates on the proportion of seeds aborted prior to maturity, possibly due to post-fertilisation environmental constraints. We conclude that the spread of B. terrestris may not alter the fecundity of L. arboreus because of the pollination service provided by A. mellifera, and discuss potential interactions between these two bee species.     

Terrestrial vertebrate communities at black-tailed prairie dog (Cynomys ludovicianus) towns

We capitalized on a regional-scale, anthropogenic experiment—the reduction of black-tailed prairie dog (Cynomys ludovicianus) towns across the Great Plains of North America—to test the hypothesis that decline of this species has led to declines in diversity of native grassland vertebrates of this region. We compared species richness and species composition of non-volant mammals, reptiles and amphibians at 36 prairie dog towns and 36 paired sites in the Panhandle Region of Oklahoma during the summers and falls of 1997, 1998 and 1999. We detected 30 species of mammals, 18 species of reptiles and seven species of amphibians. Comparisons between communities at prairie dog towns and paired sites in the adjacent landscape indicated that while richness per se was not necessarily higher in towns, they did harbor significantly more rare and imperiled species. Species that were positively associated with prairie dog towns during one or both seasons (summer and fall) included badgers (Taxidea taxus), eastern cottontails (Sylvilagus floridanus), coyotes (Canis latrans), grasshopper mice (Onychomys leucogaster), swift fox (Vulpes velox), pronghorn antelopes (Antilocapra americana), striped skunks (Mephitis mephitis), white-tailed deer (Odocoileus virginianus), cattle, thirteen-lined ground squirrels (Spermophilus tridecemlineatus), black-tailed jackrabbits (Lepus califonicus), barred tiger salamanders (Ambystoma tigrinum), plains spadefoot toads (Scaphiopus bombifrons), Great Plains toad (Bufo cognatus), Woodhouse's toad (Bufo woodhousii), prairie rattlesnakes (Crotalis viridis), western plains garter snakes (Thamnophis radix), Texas horned lizards (Phrynosoma cornutum), and ornate box turtles (Terrapene ornata).     

Recent, substantial, and unexplained declines of the common toad Bufo bufo in lowland England

A decline in numbers of a large common toad (Bufo bufo) population in south-east England during the 1990s, together with anecdotal reports of similar trends in other toad populations, prompted a nation-wide survey of this species. The survey also included the common frog (Rana temporaria) as a control for which there was no comparable evidence of recent decline. A questionnaire requesting information on the fate of toad and frog populations in the last 15 years of the 20th century was distributed to professional and amateur herpetologists during autumn 2001. Ninety-five respondents provided data on a total of 277 sites, including 232 frog and 202 toad populations in England, Scotland and Wales. More than 80% of the reported sites were rural for both species. Rural frog populations were generally doing well in most parts of Britain, with almost as many increasing as decreasing populations and a high proportion of stable populations. Rural toads also showed no overall trend across the country as a whole. However, regional analysis indicated that although toads were faring at least as well as frogs in the north and west of Britain, they were declining seriously in central and eastern/south-eastern areas. In these regions 50% or more of toad populations have experienced recent declines, whereas frogs have fared as well as they have elsewhere. Toads breeding alone have fared significantly worse than toads breeding at sites where frogs were also present. Reasons for apparently toad-specific declines in lowland England remain unknown.     

Invasive cane toads (Bufo marinus) cause mass mortality of freshwater crocodiles (Crocodylus johnstoni) in tropical Australia

Invasive species are frequently blamed for faunal declines, but there is little direct evidence about the pathways, magnitude and size-selectivity of mortality induced by invaders. Top predators are of particular interest in this context, because their removal can generate substantial cascades of secondary effects on community composition. Cane toads (Bufo marinus) are large South American anurans currently spreading rapidly through tropical Australia. Native predators that attempt to consume these highly toxic toads may die as a result. During surveys of the Victoria River in the semi-arid tropical region of the Northern Territory, we documented massive mortality of freshwater crocodiles (Crocodylus johnstoni) at the toad invasion front. Dead crocodiles spanned a wide size range (0.6-2.1 m long) but with significant biases; intermediate-sized animals (0.6-1.5 m long) were more likely to be found dead. Population densities of crocodiles plummeted by as much as 77% following toad invasion, and population size-structures changed. The negative impacts of toads on crocodiles appear to be greater in these hot semi-arid landscapes than in cooler, higher rainfall areas where crocodiles have access to a wider prey base, and the toads are less prone to desiccation and can rehydrate in small, scattered water bodies rather than in the main river. Hence, the impact of cane toad invasion on this top predator may increase with increasing aridity.     

Monitoring distributions using call surveys: estimating site occupancy, detection probabilities and inferring absence

Monitoring programs serve to track changes in the distribution and abundance of species. A major problem with most monitoring programs is that species detection is imperfect and some populations are inevitably missed. Therefore, in most monitoring programs the true distribution of a species will be underestimated. Here, we report a field test of the reliability and performance of a commonly used method to monitor the distribution of amphibians (anuran call surveys). We surveyed the distribution of four anuran species in western Switzerland, and estimated detection probabilities to account for imperfect species detection and used these estimates to adjust our estimate of site occupancy (i.e., distribution). Next, we assessed how detection probabilities were affected by weather and how site occupancy was affected by site specific covariates. For one species (Hyla arborea), call surveys proved efficient in determining the regional distribution with only few site visits because detection probabilities were relatively high. The call surveys apparently missed many populations of another common species (Bufo calamita) because detection probabilities were lower. Two other species (Bombina variegata and Alytes obstetricans) were uncommon and strong inference from the analysis is not possible. Thus, multispecies surveys may be inefficient for rare species. Estimates of detection probabilities were used to calculate how many site visits are necessary to infer the absence of a species with some predetermined statistical certainty. The implications of “false absences” are important in ecology as they are known to bias usual habitat suitability models and overestimate extinction/colonization events in metapopulations. Large-scale monitoring programs would benefit from the application of an estimation-based approach to monitoring the distribution of species.     

The effects of earthworms on the demography of annual plant assemblages in a long-term mesocosm experiment

Earthworms have been shown to influence plant growth, survival and fecundity. They can therefore affect plant demography in plant communities changing their composition. A long term mesocosm experiment was set-up to test the effects of an endogeic (Aporrectodea caliginosa) and an anecic (Lumbricus terrestris) earthworm species on assemblages of four species of annuals: one grass (Poa annua), two forbs (Veronica persica and Cerastium glomeratum) and one legume (Trifolium dubium). The number of individuals and the biomass of each species were investigated. A. caliginosa and L. terrestris affected the density of T. dubium at each of the three monitored census dates. The other plant species responded to A. caliginosa and L. terrestris at the second and third generations. The presences of A. caliginosa and L. terrestris reduced the total number of plant individuals from the second to the third generation. At harvest (3rd generation), T. dubium and V. persica had more and larger individuals in the presence of A. caliginosa. When both earthworm species were present, T. dubium had few but larger individuals. Our study confirms that earthworms affect plant demography and plant community structure. Our results also show that accurate prediction of long-term effects of earthworms on plant communities cannot be achieved using results on their short-term effects on plant growth. This is due to the poor understanding of the effects of earthworms on plant resource allocation and demography, and also the possibility that earthworms may exert the opposite effect on the short and long-term availability of nutrients.     

Earthworm-mediated maternal effects on seed germination and seedling growth in three annual plants

Many ecological studies have pointed out maternal effects in plants and shown that plant maternal environment influences germination of their seed and subsequent seedling growth. However, few have tested for maternal effects induced by soil macroorganisms. We tested whether two earthworm species (Aporrectodea caliginosa and Lumbricus terrestris) trigger such maternal effects on seed germination and seedling growth of three plant species (Veronica persica, Poa annua and Cerastium glomeratum). Our results show that, through maternal effects, A. caliginosa enhanced seed germination (V. persica and P. annua) and seedling growth (C. glomeratum and P. annua) while L. terrestris reduced seed germination only in V. persica. In some cases, the increase in germination rates of seeds produced in the presence of earthworms was associated with a reduction of nitrogen content in seeds. These results show that earthworms induce maternal effects in plants and that the size and direction of these effects depend on the combination of plant and earthworm species.     

大蟾蜍蝌蚪生长发育的高密度效应

将大蟾蜍(B uf o g arg arizans)蝌蚪分别饲养在40、80、120、160、200和240只/L 6个不同的密度条件下,通过定时定量喂食及测量其全长、体重和死亡率的变化,分析密度对蝌蚪生长发育的影响。结果表明:(1)饲养密度和生长发育时间及两者的交互作用对大蟾蜍蝌蚪的生长发育影响都极显著(P<0.01);(2)在40只/L的密度下,大蟾蜍蝌蚪的全长、体重的增长速率最快,在240只/L的密度下,大蟾蜍蝌蚪的全长、体重的增长速率最慢;(3)在相同时间内,40只/L的密度条件下大蟾蜍蝌蚪的存活率最高,在240只/L的密度条件下大蟾蜍蝌蚪的存活率最低;(4)大蟾蜍蝌蚪的生长发育速度随饲养密度的升高而减慢。     

Impacts of earthworms and arbuscular mycorrhizal fungi (Glomus intraradices) on plant performance are not interrelated

Earthworms and arbuscular mycorrhizal fungi (AMF) might interactively impact plant productivity; however, previous studies reported inconsistent results. We set up a three-factorial greenhouse experiment to study the effects of earthworms (Aporrectodea caliginosa Savigny and Lumbricus terrestris L.) and AMF (Glomus intraradices N.C. Schenck & G.S. Sm.) on the performance (productivity and shoot nutrient content) of plant species (Lolium perenne L., Trifolium pratense L. and Plantago lanceolata L.) belonging to the three functional groups grasses, legumes and herbs, respectively. Further, we investigated earthworm performance and plant root mycorrhization as affected by the treatments. Our results accentuate the importance of root derived resources for earthworm performance since earthworm weight (A. caliginosa and L. terrestris) and survival (L. terrestris) were significantly lower in microcosms containing P. lanceolata than in those containing T. pratense. However, earthworm performance was not affected by AMF, and plant root mycorrhization was not modified by earthworms. Although AMF effectively competed with T. pratense for soil N (as indicated by δ¹⁵N analysis), AMF enhanced the productivity of T. pratense considerably by improving P availability. Remarkably, we found no evidence for interactive effects of earthworms and AMF on the performance of the plant species studied. This suggests that interactions between earthworms and AMF likely are of minor importance.     

Climatic effects on calling and spawning of the natterjack toad Bufo calamita: Discriminant analyses and applications for conservation monitoring

Unlike the other anurans native to Britain, the rare natterjack toad Bufo calamita has a protracted breeding season. Surviving populations are regularly monitored for assessment of population trends and conservation requirements. We have examined the factors influencing calling and spawning behaviour at two British sites and used discriminant analysis to identify climatic conditions associated with reproductive activity. These conditions include rainfall over a short period preceding spawning, though not necessarily on the day itself, and preferably following a spell of dry weather; maximum temperatures on the day before spawning of at least 10°C, and a series of preceding minimum temperatures of at least 5°C (on average) over 2–4 nights. We anticipate that this kind of climatic information, which is accessible without actually visiting toad sites, should maximise the efficiency of monitoring operations by allowing judicious use of limited resources.     

The effect of feeding behavior on Hg accumulation in the ecophysiologically different earthworms Lumbricus terrestris and Octolaseon cyaneum: A microcosm experiment

 A soil microcosm experiment was performed to assess the uptake of Hg from various Hg-spiked food sources (soil, leaf litter and root litter of Trifolium alexandrinum) by two earthworm species, Lumbricus terrestris (anecic) and Octolaseon cyaneum (endogeic). Treatments were applied in which one of the three food sources was Hg spiked and the other two were not. Additional treatments in which all or none of the food sources were Hg spiked were used as controls. Uptake of Hg from soil into tissues of both earthworm species was significantly higher than uptake of Hg from leaf litter or root litter, indicating that soil may be the most important pool for the uptake of Hg into earthworms. In addition, the anecic L. terrestris significantly accumulated Hg from all Hg-spiked food sources (leaf litter, root litter and soil), whereas the endogeic O. cyaneum took up Hg mainly from soil particles. Interestingly, there was no further increase in Hg in L. terrestris when all food sources were Hg spiked compared to the single Hg-spiked sources. This may be attributed to the relatively high Hg content in the soil, which may have influenced the feeding behavior of the earthworms, although their biomass did not significantly decline. We suggest that, in addition to the physiological differences, feeding behavior may also play a role in the contrasting uptake of Hg by the two earthworm species.     

Are differences in fitness traits related to genetic clusters? An empirical test on the European tree frog

Microsatellites are one of the preferred marker types to estimate the level of genetic diversity and subdivision in natural populations of endangered species. Many studies thereby attempted to identify genetically defined management units for conservation by applying a variety of genetic clustering methods. It is not clear, however, whether genetic clusters inferred from neutral molecular markers reflect differences in fitness or adaptation. In this study, I conducted a common garden experiment on the endangered European tree frog (Hyla arborea) to clarify whether fitness-related traits of larval development differed between three genetic groups defined by Bayesian clustering analysis. I reared larvae under semi-natural conditions and measured growth and developmental rates as well as survival at five larval stages from eclosion to completion of metamorphosis. Nested general and generalized linear models showed significant cluster differences for several variables in terms of smaller growth rates and body sizes at early larval stages. These differences were, however, probably not linked to adaptive divergence among clusters but rather to neutral genetic processes in the breeding sites of one cluster, which were spatially isolated and subject of recent bottlenecks. Hence, a genetic load effect (e.g. inbreeding depression) may have acted on the populations of this particular cluster. I advise studies aiming to define management units for conservation to not only use genetic clustering methods but to complement their findings with experimental approaches on fitness-related traits.     

Worms from the cold: Lumbricid life stages in boreal clay during frost

The vertical distribution and activity of earthworm life stages were studied in an arable field during 0.5 m deep frost. The anecic Lumbricus terrestris L. were below the frost at the bottom of their home burrows (max. depth 1.0 m) and remained there apparently active. Their burrows were open, free of ice and water. The endogeic Aporrectodea caliginosa Sav., mainly small juveniles, were aestivating in the frost layer, which confirms freeze-tolerance in this species. Large A. caliginosa individuals were actively burrowing below the frost down to 1 m depth at soil temperatures close to +1 °C, frost evidently triggering much deeper burrowing than summer droughts. Demonstrating cold-hardiness, viable cocoons of both A. caliginosa and L. terrestris were obtained within a 0-0.25 m layer, frozen for ca. one month prior to sampling. These two common earthworms of boreal soils seem to over-winter in all life stages and remain active below the frost, potentially contributing to the maintenance of subsoil processes during the winter months.     

Homing ability widens the sphere of influence of the earthworm Lumbricus terrestris L.

We report that the deep-burrowing earthworm Lumbricus terrestris L. possesses short-range homing capability. For 2 months we followed the soil surface activity of 18 field-collected L. terrestris in an experimental laboratory population. We recorded 26 cases where an individual after completely leaving its burrow returned to the burrow by backing along its own outward trail. Maximally, homing occurred from a distance of 0.70 m and after more than 3 h on the soil surface. The evolution of homing ability in L. terrestris is most likely related to the importance of the burrow for survival and reproduction. This adaptation also guarantees individuals an expanded foraging area, which we believe can partly explain the remarkable efficiency of L. terrestris in the burial of surface litter.