Influence of temperature and relative humidity on the moulting success of Amblyomma limbatum and Aponomma hydrosauri (Acari: Ixodidae) larvae and nymphs

This study compared the duration of the moulting periods of engorged larvae and nymphs of the ixodid ticks, Amblyomma limbatum and Aponomma hydrosauri, at different temperature/relative humidity regimes, and examined the relationships between the engorged weight of ticks and their weights after moulting. The results showed that for each species, there was a significant relationship between the weights of unfed nymphs and engorged larvae, and the weights of unfed adults and engorged nymphs. The weight of engorged nymphs was also a good indicator of their sex, with female ticks having heavier weights as engorged nymphs. Temperature and relative humidity had a marked effect on the moulting success of engorged ticks of both species. Aponomma hydrosauri larvae and nymphs were able to moult at lower temperatures than Amb. limbatum but most ticks, except Ap. hydrosauri larvae, failed to moult at 13 degrees C. Additionally, there was a marked decrease in the pre-moult times of ticks at higher temperatures, with larvae taking less time to moult than nymphs. At temperatures greater than 21 degrees C, Amb. limbatum took less time to moult than Ap. hydrosauri but this interspecific difference was less marked for nymphs. The interspecific differences in the responses of engorged larvae and nymphs to different temperatures and relative humidities correlated with interspecific differences in off-host behaviour and with the different climates the two species experience throughout most of their distributional range.     

Interspecific differences in the movements of female ticks on reptiles.

Movements of females of two reptile ticks, Amblyomma limbatum and Aponomma hydrosauri were studied to determine whether the behaviour of the species on hosts was similar or different. Experiments were conducted to determine if the movement of females on hosts prior to mating was influenced by the delayed attachment of conspecific males. A field experiment was also undertaken to determine whether data obtained during laboratory experiments reflected the behaviour of females on hosts in the field. Both in laboratory and field experiments, a significantly greater proportion of Ap. hydrosauri females changed their site of attachment prior to mating. Individual Ap. hydrosauri females made significantly more positional changes than Amb. limbatum females. These interspecific differences occurred irrespective of the absence of conspecific males or presence of non-conspecific adults on hosts. These data may have important ecological implications in relation to the success of colonizing females at their parapatric boundary.     

Competition for sites of attachment to hosts in three parapatric species of reptile tick

Summary Competition for sites of attachment to hosts by three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum was investigated as a possible cause for the parapatric distributions found in these species throughout southern Australia. Two localities were chosen for detailed study; a boundary between Ap. hydrosauri and Amb. limbatum near Mt. Mary in the mid-north of South Australia and a boundary between Ap. hydrosauri and Amb. albolimbatum near Arno Bay on the Eyre Peninsula of South Australia. Comparisons of sites of attachment to hosts were made between hosts infested by one species of tick and hosts infested by two species of tick. At Mt. Mary, Ap. hydrosauri and Amb. limbatum adults attach more commonly in the ears and on the midback of their hosts, however, no evidence was found to suggest that competition between the species occurred for these sites. At Arno Bay, Amb. albolimbatum adults had similar sites of attachment to hosts as the other two species, however, significantly more of this species attach in the ears of their hosts. There is no difference in sites of attachment to hosts of Ap. hydrosauri females at Arno Bay and Mt. Mary. Ap. hydrosauri males at Arno Bay, however, show a shift in sites of attachment to hosts away from those sites occupied by Amb. albolimbatum males and females. This shift occurs only in cases where Amb. albolimbatum did not infest the same host. Although the shift in male Ap. hydrosauri sites of attachment to hosts can be explained in terms of past competition, there is no evidence to suggest that such competition in the past, or competition at present, maintains the parapatric boundaries found in these species of thick.     

Reproductive interference between three parapatric species of reptile tick

Summary Reproductive interference between three species of reptile tick, Aponomma hydrosauri, Amblyomma albolimbatum and Amb. limbatum was investigated. Adults of two species attach together on the same lizard hosts in narrow overlap zones at parapatric boundaries between species, providing opportunities for interference. The possibility of reproductive interference was suggested because of similarities in the sexual communication systems.Three forms of interference were found in laboratory experiments. Firstly male movement to search for sexually receptive females was inhibited when females of two species were present on the same host, probably because of signal interference. Secondly, a nonspecific, short range attractant pheromone led males into non-conspecific courtship attempts which reduced the time for conspecific courtships. Thirdly Amb. albolimbatum males physically blocked the genitalia of female Ap. hydrosauri after unsuccessful non-conspecific courtship attempts. Field data showed this behaviour was common in both Amblyomma species after conspecific matings. Reproductive interference would reduce the fitness of each species in sympatry, and may contribute to the maintenance of the narrow parapatric boundaries.     

Niche changes between parasite populations: An example from ticks on reptiles

Summary Two Australian tick species Aponomma hydrosauri and Amblyomma albolimbatum have the same major host species, the lizard Trachydosaurus rugosus. While females of Amb. albolimbatum are most often attached in the ears and on the neck of their hosts, Ap. hydrosauri females prefer to attach further back, under the forearms and on the back. Males show the same interspecific difference but there is also a difference between populations. Ap. hydrosauri males from populations in contact with Amb. albolimbatum attach more often in posterior positions than Ap. hydrosauri males from populations isolated from Amb. albolimbatum. These differences were found in both field populations and laboratory reared ticks. Phylogenetic evidence suggests that the change in male attachment site between Ap. hydrosauri populations followed the colonization of T. rugosus. We propose that the most likely reason for the change of attachment sites has been interspecific interactions with Amb. albolimbatum and that competition has been for space for efficient reception of female signals.     

Ant predation on different life stages of two Australian ticks

Laboratory colonies of Rhytidoponera ants were allowed to prey on the fed and unfed stages of the Australian ticks Aponomma hydrosauri and Amblyomma limbatum. The unfed tick stages had a higher survival than the fed stages. The ants took longer to handle the adult ticks than the nymphs and longer to handle the nymphs than the larvae. The ants also took longer to handle the unfed than fed nymphs, but longer to handle the fed than unfed females. As well as the differences between the tick stages, there was a species effect, with the ants taking longer to handle A. limbatum, and with that tick species having a higher survival than A. hydrosauri after ant predation. These stage and species differences may influence the tick population dynamics. This revised version was published online in November 2006 with corrections to the Cover Date.     

Mating behaviour and parapatry in two species of Australian reptile tick

Summary Few quantitative studies have examined the ecological consequences of similarities and/or differences in mating behaviour of parapatric species. Reproductive interference occurs between several parapatric species of Australian reptile tick, due to similarities in their mating behaviour (Andrews et al. 1982a). Attempts to determine whether reproductive interference serves to maintain parapatry between Amblyomma limbatum and Aponomma hydrosauri have been hindered because of difficulties in providing conditions conducive to conspecific mating in Amb. limbatum. The present study examined whether off-host and/or onhost temperature influenced the subsequent mating behaviour (i.e. the proportion of females that mate and the time when mating occurs) of these two species. Irrespective of the temperature experienced by ticks prior to host attachment, specific on-host temperatures were needed to induce mating in Amb. limbatum (i.e. host cloacal temperatures >32° C prior to the time of peak mating activity). Significantly more Amb. limbatum females were mated and the time taken by females to mate decreased with increasing on-host temperatures. mating in Ap. hydrosauri occurred over a wider range of on-host temperatures and the time when mating occurred did not alter at different on-host temperatures. In addition, significantly more Ap. hydrosauri males moved and each male made more moves on hosts than did Amb. limbatum males. It is suggested that Ap. hydrosauri may in consequence have a competitive mating advantage over Amb. limbatum at a boundary. Similarities in mating behaviour, on the other hand, increase the probability of reproductive interference, hence reduce the reproductive fitness of colonizing females of both species. We propose that similarities and differences in mating behaviour could play a critical role in the maintenance of parapatric boundaries.     

Laboratory studies of ant predation on parapatric reptile ticks

Abstract Two reptile tick species, Aponomma hydrosauri and Amblyomma limbatum, have a parapatric distribution in South Australia. Predation may play a role in maintaining the boundary. Laboratory colonies of Rhytidoponera and Iridomyrmex ants were collected from near Mt Mary, South Australia, close to the tick boundary. They were tested as predators of the two tick species. In the experiments, ticks in leaf litter were more protected from predation than those on bare soil. When comparing leaf litter types from the Mt Mary area, mallee litter was more protective than bluebush litter of equivalent depth. Ticks positioned at the base of the litter layer were more protected from predation than those at the litter surface, and Amb. limbatum ticks were more resistant to predation than Ap. hydrosauri ticks. These results contribute to our understanding of the mechanisms maintaining the abrupt parapatric boundary between the two tick species. Predators may contribute to preventing the more susceptible Ap. hydrosauri from spreading further north, where bluebush litter is more common, and so predation risk is higher. Predators probably have less influence in preventing Amb. limbatum from spreading further south.     

Regional‐scale patterns of soil microbes and nematodes across grasslands on the Mongolian plateau: relationships with climate,soil, and plants

Belowground communities exert major controls over the carbon and nitrogen balances of terrestrial ecosystems by regulating decomposition and nutrient availability for plants. Yet little is known about the patterns of belowground communities and their relationships with environmental factors, particularly at the regional scale where multiple environmental gradients co‐vary. Here, we describe the patterns of belowground communities (microbes and nematodes) and their relationships with environmental factors based on two parallel studies: a field survey with two regional‐scale transects across the Mongolia plateau and a water‐addition experiment in a typical steppe. In the field survey, soils and plants were collected across two large‐scale transects (a 2000‐km east–west transect and a 900‐km south–north transect). At the regional‐scale, the variations in soil microbes (e.g. bacterial PLFA, fungal PLFA, and F/B ratio) were mainly explained by precipitation and soil factors. In contrast, the variation in soil nematodes (e.g. density of trophic groups and the bacterial‐feeding/fungal‐feeding nematode ratio) were primarily explained by precipitation. These variations of microbe or nematode variables explained by environmental factors at regional scale were derived from different vegetation types. Along the gradient from nutrient‐poor to nutrient‐rich vegetation types, the total variation in soil microbes explained by precipitation increased and that explained by plant and soil decreased, while the opposite was true for soil nematodes. Experimental water addition, which increased rainfall by 30% during the growing season, increased biomass or density of belowground communities, with the nematodes being more responsive than the microbes. The different responses of soil microbial and nematode communities to environmental gradients at the regional scale likely reflect their different adaptations to climate, soil nutrients, and plants. Our findings suggest that the soil nematode and microbial communities are strongly controlled by bottom‐up effects of precipitation alone or in combination with soil conditions.     

Rural–urban gradient in seasonal forcing of measles transmission in Niger

Seasonally driven cycles of incidence have been consistently observed for a range of directly transmitted pathogens. Though frequently observed, the mechanism of seasonality for directly transmitted human pathogens is rarely well understood. Despite significant annual variation in magnitude, measles outbreaks in Niger consistently begin in the dry season and decline at the onset of the seasonal rains. We estimate the seasonal fluctuation in measles transmission rates for the 38 districts and urban centres of Niger, from 11 years of weekly incidence reports. We show that transmission rates are consistently in anti-phase to the rainfall patterns across the country. The strength of the seasonal forcing of transmission is not correlated with the latitudinal rainfall gradient, as would be expected if transmission rates were determined purely by environmental conditions. Rather, seasonal forcing is correlated with the population size, with larger seasonal fluctuation in more populous, urban areas. This pattern is consistent with seasonal variation in human density and contact rates due to agricultural cycles. The stronger seasonality in large cities drives deep inter-epidemic troughs and results in frequent local extinction of measles, which contrasts starkly to the conventional observation that large cities, by virtue of their size, act as reservoirs of measles.     

Tropical metacommunities along elevational gradients: effects of forest type and other environmental factors

Elevational gradients provide a natural experiment for assessing the extent to which the structure of animal metacommunities is molded by biotic and abiotic characteristics that change gradually, or is molded by aspects of plant community composition and physiognomy that change in a more discrete fashion. We used a metacommunity framework to integrate species‐specific responses to environmental gradients as an approach to detect emergent patterns at the mesoscale in the Luquillo Mountains of Puerto Rico. Elements of metacommunity structure (coherence, species turnover and range boundary clumping) formed the basis for distinguishing among random, checkerboard, Gleasonian, Clementsian, evenly spaced and nested patterns. Paired elevational transects (300–1000 m a.s.l.) were sampled at 50 m intervals to decouple underlying environmental mechanisms: a mixed forest transect reflected changes in abiotic and biotic conditions, including forest type (i.e. tabonuco, palo colorado and elfin forests), whereas another transect reflected changes in environmental conditions but not forest type, as its constituent plots were located within palm forest. Based on distributional data (presence versus absence of species), the mixed forest transect exhibited Clementsian structure, whereas the palm forest transect exhibited quasi‐Gleasonian structure. In contrast, the distribution of modes in species abundance was random with respect to the latent environmental gradient in the mixed forest transect and clumped with respect to the latent environmental gradient in the palm forest transect. Such contrasts suggest that the environmental factors affecting abundance differed in form or type from those affecting distributional boundaries. Variation among elevational strata with respect to the first axis of correspondence from reciprocal averaging was highly correlated with elevation along each transect, even though axis scores were not correlated between mixed forest and palm forest transects. This suggests that the identity of the environmental characteristics, or the form of response by the fauna to those characteristics, differed between the two elevational transects. Despite the proximity of the transects, the patchy configuration of palm forest, and the pervasive distribution of the dominant palm species, the relative importance of abiotic variables and habitat in structuring gastropod metacommunities differed between transects, which is remarkable and attests to the sensitivity of metacommunity structure to environmental variation.     

Environmental predictions and distributional limits of chromosomal taxa in the Australian grasshopperCaledia captiva (F.)

Summary Correlations between environmental parameters and the distribution of four chromosomal taxa in the acridine grasshopperCaledia captiva (F.) have been analyzed. A bioclimate prediction system (BIOCLIM) was used as a model to determine their bioclimatic envelopes and to predict their probable geographic distributions, based on climatic similarities with actual distributions. New, and formerly unsuspected, distribution areas are predicted. Parapatric boundaries are defined that actually coincide with known limits. The general distribution of the four taxa is correlated with rainfall seasonality and a temperature gradient. Their distributional boundaries are mostly in accordance with extreme values in rainfall in an east-west direction and temperature in a north-south direction. A nonmetric multidimensional scaling analysis of one selected narrow parapatric boundary confirms the nature of the geographic correlation with several climatic extreme values. The climatic and vegetation analyses suggest that sharp parapatric distributions can be maintained by gradual environmental influences and that coincidence with abrupt ecotones is not necessary. At least one of the parapatric boundaries seems to be relatively recent in origin (2000–6000 years) and on the basis of the location of extreme climatic values as well as chromosomal and genetic markers it is expected to slowly move further south and east of its present location apparently following a climatically established boundary.     

Plant compositional change over time increases with rainfall in Serengeti grasslands

Previous work from the Plains of Serengeti National Park (SNP) suggested that plant species composition (i.e. the composite measure of species' identities and abundances) is unchanged on the temporal scale of decades ( Belsky 1985 ), raising the possibility that African grazing ecosystems may be resistant to climate change. In this study, grassland plant composition was measured three times between 2000 and 2007 at eight permanent sites spanning the SNP environmental gradient. Confirming previous work, plots in the Serengeti Plains were relatively stable with respect to plant composition through time. In contrast, sites in all other regions of the ecosystem showed greater change in plant species composition during the study. Three factors, annual rainfall, dry season rainfall, and fire frequency, were linearly correlated (all r>0.73) with the amount of composition change at the sites. A path model, which accounted for the strong correlation between annual and dry season rainfall and for the known effect of annual rainfall on fire frequency, was used to elucidate which factors were associated with composition change and which were spuriously correlated. The path model demonstrated that change in plant composition was positively related to annual rainfall, but not dry season rainfall or fire frequency. However, dry season rainfall was positively associated with changes in plant species identity, one component of compositional change, across sites over time. Because climate models predict both increased wet season and decreased dry season rainfall in East Africa, these findings raise uncertainty about how these grasslands will respond to future changes. This study and others ( Belsky 1985 ) suggests that plant assemblages adapted to low rainfall in Serengeti may exhibit compositionally stability, while the same may not be true in regions of higher average rainfall and larger species pools.     

A comparison of the reproductive parameters of females of two reptile tick species

In comparisons of females of two reptile tick species Aponomma hydrosauri and Amblyomma limbatum, Ap. hydrosauri was initially larger, and after mating on the host engorged faster and remained attached for a shorter time before completing engorgement and detaching. Amb. limbatum had a longer period of engorgement, and achieved a greater engorged weight. Engorged Amb. limbatum females laid significantly more eggs than equivalent sized Ap. hydrosauri. Although the two species are ecologically similar and were collected from the same site for this study, their reproductive differences probably reflect adaptations to different conditions in their largely allopatric ranges.     

Contrasting relationships between precipitation and species richness in space and time

Future changes in precipitation regimes are likely to impact species richness in water-limited plant communities. Regional, spatial relationships between precipitation and richness could offer information about how altered rainfall will impact local communities, assuming that processes driving the regional relationship are also dominant at fine spatial and short temporal scales. To test this assumption, we compared spatial and temporal relationships between precipitation and both species richness and species turnover in central North American grasslands. Across a broad geographic gradient, mean plant species richness in 1-m² plots increased significantly with mean annual precipitation. In contrast, over a 36-yr period at one mixed-grass prairie in the center of the regional gradient, single-year precipitation and richness were poorly correlated, and consecutive wet years had little effect on richness. Instead, richness increased most in wet years that followed dry years. Geographically dispersed sites receiving different levels of mean annual precipitation displayed strong differences in species composition, whereas temporal variation in precipitation at one site was not related to compositional dissimilarity, indicating that species turnover plays a key role in generating the regional relationship. Analyses of individual species' presence-absence suggest that the lagged temporal responses reflect environmental germination cues more than resource competition. These complex cues may dampen the initial impact of altered precipitation on diversity, but over the long term, turnover in species composition should lead to changes in richness, as in the regional, spatial relationship. How quickly this long-term response develops may depend on the colonization rates of species better adapted to the altered rainfall regime.     

Plant Functional Traits and the Distribution of West African Rain Forest Trees along the Rainfall Gradient

Plant morphological and physiological traits affect the way plants tolerate environmental stresses and therefore play an important role in shaping species distribution patterns in relation to environmental gradients. Despite our growing knowledge on the role of functional traits in structuring plant communities, few studies have tested their importance at large scales in the wet tropics. Here, we describe the distribution patterns of the most important West African rain forest timber species along the regional rainfall gradient and relate them to their functional traits. We found that the distribution patterns of 25 out of the 31 studied species (80%) were significantly related to mean annual rainfall. Shade tolerance and drought resistance were identified as the main strategy axes of variation. Wood density and leaf deciduousness emerged as the best predictor traits of species position along the rainfall gradient, explaining respectively 32 and 15 percent of the variation. Species traits tended to show stronger relationships with estimated optimum annual rainfall for each species than to the extreme rainfall conditions where they occur. The significant role of rainfall in shaping timber species distribution and the strong relationships between species traits and rainfall indicate that changes in climate, especially declining rainfall, could have strong effects on species composition and abundance in these tropical forests.     

Intermediary disturbance increases tree diversity in riverine forest of southern Brazil

Floods are frequently associated with disturbance in structuring riverine forests and they lead to environmental heterogeneity over space and time. We evaluated the distribution of tree species, ecological groups, species richness and diversity from the point bar to the slope of a riverside forest in southern Brazil (Lat. 30°01′S, Long. 52°47′W) to analyze the effects of flooding on soil properties and forest structure. A plot of 50 × 200 m divided in five contiguous transects of 10 × 200 m parallel to the river was installed, where we measured all the individual trees with pbh ≥ 15 cm. A detailed topographical and soil survey was carried out across the plot and indicated significant differences in organic matter and most mineral nutrients through the topographical gradient. The 1,229 surveyed individuals belonged to 72 species and 35 families. We used Partial CCA and Species Indicator Analysis to observe the spatial distribution of species. Both analyses showed that species distribution was strongly related to the flooding gradient, soil properties and also by space and pure spatial structuring of species and environmental variables (spatial autocorrelation), although a large part of variation remains unexplained. The ecological groups of forest stratification, plant dispersal and requirements for germination indicated slight differences among frequently, occasional and non-flooded transects. Species richness and diversity were higher at intermediate elevations and were associated to the increased spatial–temporal environmental heterogeneity. Across the plot, the direct influence of flooding on tree species distribution created a vegetation zonation that is determined by predicted ecological traits.     

北京东灵山海拔梯度上辽东栎种群结构和空间分布

种群年龄结构和空间分布格局是种群生态学的核心研究内容.为了阐明辽东栎种群海拔梯度分布特点,在北京东灵山地区辽东栎海拔分布范围(1000～1800m)内调查10条样带,研究种群大小级结构和空间分布的变异.种群的平均胸径在海拔梯度上表现出两段式的分布特征,海拔1480m为两段分布的分界点,在每一段内随海拔增加平均胸径也增加, 这反映了海拔梯度上种群的不同发育历史.种群密度、种群的聚集程度、种群的结构在海拔梯度上的分布特征都与平均胸径分布相似,种群密度和聚集程度与平均胸径为负相关系,其分布趋势与平均胸径相反.总体上,东灵山海拔梯度上辽东栎种群还是比较稳定的.辽东栎种群结构和空间分布在海拔梯度上的分布特征是种群发育历史、物种特性、环境、干扰等因素在海拔梯度上综合作用的结果.     

Unproportionally high concentrations of diadenosine triphosphate (Ap3A) and diadenosine tetraphosphate (Ap4A) in heavy platelets. Consequences for in vitro studies with human platelets

Platelets from whole blood were separated into five density subpopulations using a discontinuous Percoll gradient. The content of diadenosine triphosphate (Ap3A), diadenosine tetraphosphate (Ap4A), ADP and ATP were determined in the subfractions. The dinucleotides were directly measured in neutralized, acid-soluble extracts of human platelets with a bioluminescence method not requiring any chromatographic step. When comparing the nucleotide contents of the density subpopulations it became evident that all nucleotides steadily increased with increasing density. Ap3A, Ap4A, ADP and ATP were present in 10-, 7-, 4- and 2-fold higher amounts in the heaviest platelets, respectively, as compared to the subfraction with the lowest density. This finding is practically relevant since the most dense platelet subpopulations may be lost during conventional centrifugation to obtain platelet-rich plasma. Therefore we compared a platelet population obtained from PRP with the platelet population, which had been prepared from whole blood by means of a continuous Percoll gradient. All the four nucleotides investigated were represented in 1.5- to 2-fold higher amounts in the whole blood platelet population. This indicates that PRP does not contain a representative population but lacks part of the large heavy platelets containing the highest amounts of nucleotides.     

Sampling Techniques Influence Understory Plant Trajectories After Restoration: An Example from Ponderosa Pine Restoration

Abstract Although there is no one correct technique for sampling vegetation, the sampling design chosen may greatly influence the conclusions researchers can draw from restoration treatments. Considerations when designing vegetation sampling protocol include determining what sampling attributes to measure, the size and shape of the sampling plot, the number of replicates and their location within the study area, and the frequency of sampling. We installed 20 point‐intercept transects (50‐m long), 8 belt transects (10 × 50 m), 10 adapted Daubenmire transects (four 0.5 × 2‐m plots), and 4 modified‐Whittaker plots (20 × 50 m with smaller nested plots) in treatment and control units to measure understory herbaceous response in a forest restoration experiment that tested different treatments. Point‐intercept transects on average recorded at least twice as much plant cover as did adapted Daubenmire transects and modified‐Whittaker plots taken at the same location for all control and treatment units. Point‐intercept transects and adapted Daubenmire plots on average captured fewer rare and exotic species in the control and treatment units in comparison with the belt transects and modified‐Whittaker plots. Modified‐Whittaker plots captured the highest species richness in all units. Early successional understory response to restoration treatments was likely masked by the response of the herbaceous community to yearly climatic variation (dry vs. wet years). Species richness and abundance were higher in wet years than dry years for all control and treatment units. Our results illustrate that sampling techniques can greatly influence perceptions of understory plant trajectories and therefore the interpretation of whether restoration goals have been achieved. In addition, our results suggest that restoration monitoring needs to be conducted for a sufficient length of time so that restoration treatment responses can be detected.