Konstans Wells

Top-down control of herbivory by birds and bats in the canopy of temperate broad-leaved oaks (Quercus robur)

The intensive foraging of insectivorous birds and bats is well known to reduce the density of arboreal herbivorous arthropods but quantification of collateral leaf damage remains limited for temperate forest canopies. We conducted exclusion experiments with nets in the crowns of young and mature oaks, Quercus robur, in south and central Germany to investigate the extent to which aerial vertebrates reduce herbivory through predation. We repeatedly estimated leaf damage throughout the vegetation period. Exclusion of birds and bats led to a distinct increase in arthropod herbivory, emphasizing the prominent role of vertebrate predators in controlling arthropods. Leaf damage (e.g., number of holes) differed strongly between sites and was 59% higher in south Germany, where species richness of vertebrate predators and relative oak density were lower compared with our other study site in central Germany. The effects of bird and bat exclusion on herbivory were 19% greater on young than on mature trees in south Germany. Our results support previous studies that have demonstrated clear effects of insectivorous vertebrates on leaf damage through the control of herbivorous arthropods. Moreover, our comparative approach on quantification of leaf damage highlights the importance of local attributes such as tree age, forest composition and species richness of vertebrate predators for control of arthropod herbivory.

Grassland management intensification weakens the associations among the diversities of multiple plant and animal taxa

Land-use intensification is a key driver of biodiversity change. However, little is known about how it alters relationships between the diversities of different taxonomic groups, which are often correlated due to shared environmental drivers and trophic interactions. Using data from 150 grassland sites, we examined how land-use intensification (increased fertilization, higher livestock densities, and increased mowing frequency) altered correlations between the species richness of 15 plant, invertebrate, and vertebrate taxa. We found that 54% of pairwise correlations between taxonomic groups were significant and positive among all grasslands, while only one was negative. Higher land-use intensity substantially weakened these correlations (35% decrease in r and 43% fewer significant pairwise correlations at high intensity), a pattern which may emerge as a result of biodiversity declines and the breakdown of specialized relationships in these conditions. Nevertheless, some groups (Coleoptera, Heteroptera, Hy...

Impact of rain-forest logging on helminth assemblages in small mammals (Muridae, Tupaiidae) from Borneo

Parasites are ubiquitous in wild animals, with host-specific life histories considered as major determinants of prevalence and parasite assemblage patterns. It is predicted that habitat differences in logged rain forests influence population performances of small mammals and consequently may change the infection patterns of local animal populations with regard to endo- and ectoparasites. We investigated patterns of helminth species assemblages (Nematoda, Platyhelminthes) in two rat species ( Leopoldamys sabanus , Niviventer cremoriventer ) and two tree shrew species ( Tupaia tana , T. longipes ) in three logged and three unlogged rain forests in Borneo by examining 337 faecal samples with non-invasive faecal egg count (FEC). Nematode eggs prevailed in 95% of all samples with up to five (mean 1.9 ± 1.1) morphotypes. Whereas members of Strongylida were most prevalent in L. sabanus , T. tana and T. longipes , Spirurida dominated in N. cremoriventer that revealed at the same time the lowest average nematode prevalence and FEC. Cestode eggs were only found in L. sabanus and T. tana . Composition and abundance patterns of the parasitic helminth assemblages were influenced by logging. As hypothesized, species richness of nematode morphotypes and mean number of infections per host of T. longipes were larger in logged than in unlogged forest. In contrast, L. sabanus was more heavily infected with cestodes in unlogged than in logged forest and also revealed larger egg counts for strongylids and spirurids in unlogged forest. Our results suggest that forest degradation and altered environmental conditions influence helminth diversity and infection patterns of small mammals with contrasting trends among host species. The inconsistent logging-induced changes in helminth assemblages from different hosts indicate that specific sets of habitat-host-parasite interactions are uniquely influenced by the effects of logging. Consequently, predictions on changes of parasite diversity and prevalence with regard to habitat disturbance need to be based on the individual life histories of the hosts (and the parasites).

Bartonella and Rickettsia in arthropods from the Lao PDR and from Borneo, Malaysia !

Rickettsioses and bartonelloses are arthropod-borne diseases of mammals with widespread geographical distributions. Yet their occurrence in specific regions, their association with different vectors and hosts and the infection rate of arthropod-vectors with these agents remain poorly studied in South-east Asia. We conducted entomological field surveys in the Lao PDR (Laos) and Borneo, Malaysia by surveying fleas, ticks, and lice from domestic dogs and collected additional samples from domestic cows and pigs in Laos. Rickettsia felis was detected by real-time PCR with similar overall flea infection rate in Laos (76.6%, 69/90) and Borneo (74.4%, 268/360). Both of the encountered flea vectors Ctenocephalides orientis and Ctenocephalides felis felis were infected with R. felis. The degrees of similarity of partial gltA and ompA genes with recognized species indicate the rickettsia detected in two Boophilus spp. ticks collected from a cow in Laos may be a new species. Isolation and further characterization will be necessary to specify it as a new species. Bartonella clarridgeiae was detected in 3/90 (3.3%) and 2/360 (0.6%) of examined fleas from Laos and Borneo, respectively. Two fleas collected in Laos and one flea collected in Borneo were co-infected with both R. felis and B. clarridgeiae. Further investigations are needed in order to isolate these agents and to determine their epidemiology and aetiological role in unknown fever in patients from these areas.

Pitchers of Nepenthes rajah collect faecal droppings from both diurnal and nocturnal small mammals and emit fruity odour

The pitchers of Nepenthes rajah, a montane carnivorous plant species from Borneo, are large enough to capture small vertebrates such as rats or lizards, which occasionally drown therein. The interactions of N. rajah with vertebrates, however, are poorly understood, and the potential mechanisms that lure vertebrates to the pitchers are largely unknown. We observed frequent visits (average: one visit per 4.2 h) of both the diurnal tree shrew Tupaia montana and the nocturnal rat Rattus baluensis to pitchers by infrared sensor camera and video recording. Both mammalian species often licked the inner surface of the pitcher lid, which harbours numerous exudate-producing glands. Analysis of volatiles extracted from the secretions of the pitcher lids by gas chromatography coupled to mass spectrometry (GC/MS) revealed 44 volatile compounds, including hydrocarbons, alcohols, esters, ketones and sulphur-containing compounds, which are commonly present in sweet fruit and flower odours. The faeces of small mammals were repeatedly observed inside the pitcher, whereas we found the body of only one Tupaia montana drowned in the 42, vital and reasonably large, surveyed pitchers. Our findings suggest that the N. rajah pitcher makes use of the perceptual biases of rats and tree shrews by emitting volatiles known from fruits. The profits that the plant obtains from the repeated visits of two small mammals, together with the provision of exudates for the mammals, comprise an exceptional case of plant–vertebrate interaction.

Timing and severity of immunizing diseases in rabbits is controlled by seasonal matching of host and pathogen dynamics

Infectious diseases can exert a strong influence on the dynamics of host populations, but it remains unclear why such disease-mediated control only occurs under particular environmental conditions. We used 16 years of detailed field data on invasive European rabbits (Oryctolagus cuniculus) in Australia, linked to individual-based stochastic models and Bayesian approximations, to test whether (i) mortality associated with rabbit haemorrhagic disease (RHD) is driven primarily by seasonal matches/mismatches between demographic rates and epidemiological dynamics and (ii) delayed infection (arising from insusceptibility and maternal antibodies in juveniles) are important factors in determining disease severity and local population persistence of rabbits. We found that both the timing of reproduction and exposure to viruses drove recurrent seasonal epidemics of RHD. Protection conferred by insusceptibility and maternal antibodies controlled seasonal disease outbreaks by delaying infection; this could have also allowed escape from disease. The persistence of local populations was a stochastic outcome of recovery rates from both RHD and myxomatosis. If susceptibility to RHD is delayed, myxomatosis will have a pronounced effect on population extirpation when the two viruses coexist. This has important implications for wildlife management, because it is likely that such seasonal interplay and disease dynamics has a strong effect on long-term population viability for many species.

Co‐infections and environmental conditions drive the distributions of blood parasites in wild birds

Experimental work increasingly suggests that non-random pathogen associations can affect the spread or severity of disease. Yet due to difficulties distinguishing and interpreting co-infections, evidence for the presence and directionality of pathogen co-occurrences in wildlife is rudimentary. We provide empirical evidence for pathogen co-occurrences by analysing infection matrices for avian malaria (Haemoproteus and Plasmodium spp.) and parasitic filarial nematodes (microfilariae) in wild birds (New Caledonian Zosterops spp.). Using visual and genus-specific molecular parasite screening, we identified high levels of co-infections that would have been missed using PCR alone. Avian malaria lineages were assigned to species level using morphological descriptions. We estimated parasite co-occurrence probabilities, while accounting for environmental predictors, in a hierarchical multivariate logistic regression. Co-infections occurred in 36% of infected birds. We identified both positively and negatively correlated parasite co-occurrence probabilities when accounting for host, habitat and island effects. Two of three pairwise avian malaria co-occurrences were strongly negative, despite each malaria parasite occurring across all islands and habitats. Birds with microfilariae had elevated heterophil to lymphocyte ratios and were all co-infected with avian malaria, consistent with evidence that host immune modulation by parasitic nematodes facilitates malaria co-infections. Importantly, co-occurrence patterns with microfilariae varied in direction among avian malaria species; two malaria parasites correlated positively but a third correlated negatively with microfilariae. We show that wildlife co-infections are frequent, possibly affecting infection rates through competition or facilitation. We argue that combining multiple diagnostic screening methods with multivariate logistic regression offers a platform to disentangle impacts of environmental factors and parasite co-occurrences on wildlife disease.

Shifts from native to invasive small mammals across gradients from tropical forest to urban habitat in Borneo

AbstractUrbanization has paved the way for the spread of commensal rodents at global scale. However, it is largely unknown how these species use tropical anthropogenic landscapes originally covered with forests and inhabited by diverse small mammal assemblages. We surveyed non-flying small mammals in various urban and suburban habitat types and adjacent forest in the tropical town of Kota Kinabalu in Borneo. We used occupancy and polynomial regression models to determine variation in species occurrences along gradients of land-use intensity. Müller’s sundamys (Sundamys muelleri) was the only native small mammal species found in urban and suburban landscapes with a continuous decrease in occurrence probability from forests to urban habitats. The invasive Asian black rat (Rattus rattus species complex) and the invasive Asian house shrew (Suncus murinus) had the highest occurrence probabilities in habitats of intermediate land-use intensity, but Asian black rats are also likely to occasionally invade forested habitats and occupied urban habitats in sympatry with the Norway rat (Rattus norvegicus). In urban and suburban habitats, fallow land possibly favoured the occurrence of S. muelleri and S. murinus. Other native small mammal species (Muridae, Sciuridae, Tupaiidae) were found only in forested areas. Our study shows that native small mammals found in forest are largely replaced by invasive species in urban and suburban habitats. Due to their occurrence in habitats of various land use intensities, S. muelleri and R. rattus comprise central links between forest wildlife and urban species, an association that is important to consider in studies of parasite and disease transmission dynamics.

Seed consumption by small mammals from Borneo

Fruit and seed consumers can both positively and negatively affect plant recruitment through seed dispersion and seed predation, respectively. In turn, fruits influence the abundance and distribution of consumers sustained by local plant assemblages. These interactions are key processes in plant recruitment and the dynamics of tropical forests, where most plants depend on dispersal by frugivorous animals (Corlett 1998). An understanding of these interactions and the functional role of particular seed-dispersing animals is increasingly important nowadays, given that human impact on tropical forest ecosystems may negatively impact seed dispersal and forest regeneration in both natural and human-altered forests (Wright et al. 2000). Small mammals generally comprise species-rich and abundant assemblages in tropical forests that exploit various habitats on the ground and in the different strata above (Bourliere 1989, Malcolm 1991, Voss & Emmons 1996, Wells et al. 2004). Most of the commonly encountered small mammals in South-East Asian forests are considered to have an omnivorous and opportunistic diet, including a wide array of plant and animal materials (Emmons 1991, Langham 1983, Leighton & Leighton 1983, Lim 1970). However, detailed information on diet composition and the types or quantities of seeds being consumed are rarely available, and the presumably important role of small mammals as seed dispersers remains anecdotal and has yet to be quantified. Generally, the majority of studies on small mammals and seeds have been centred on seed dispersal via

Host specificity and niche partitioning in flea–small mammal networks in Bornean rainforests

The diversity of ectoparasites in Southeast Asia and flea–host associations remain largely understudied. We explore specialization and interaction patterns of fleas infesting non‐volant small mammals in Bornean rainforests, using material from a field survey carried out in two montane localities in northwestern Borneo (Sabah, Malaysia) and from a literature database of all available interactions in both lowland and montane forests. A total of 234 flea individuals collected during our field survey resulted in an interaction network of eight flea species on seven live‐captured small mammal species. The interaction network from all compiled studies currently includes 15 flea species and 16 small mammal species. Host specificity and niche partitioning of fleas infesting diurnal treeshrews and squirrels were low, with little difference in specialization among taxa, but host specificity in lowland forests was found to be higher than in montane forests. By contrast, Sigmactenus alticola (Siphonaptera: Leptopsyllidae) exhibited low host specificity by infesting various montane and lowland nocturnal rats. However, this species exhibited low niche partitioning as it was the only commonly recorded flea from rats on Borneo. Overall complementary specialization was of intermediate intensity for both networks and differed significantly from random association; this has important implications for specific interactions that are also relevant to the potential spread of vector‐borne diseases.