Evolution of larval food plant associations in Delias Hübner butterflies (Lepidoptera: Pieridae)

A review of larval food plants of the genus Delias is presented. Larvae specialize primarily on aerial‐stem and root hemiparasites (“mistletoes”) in the order Santalales. Although butterfly food plant associations have been recorded for only a small proportion of the genus (28 species or ～11%, representing 12/24 species‐groups), available data suggest that the family Loranthaceae is used most frequently (77%), followed by the Santalaceae sensu stricto (14%) and Viscaceae (8%). With the possible exception of Euphorbiaceae (1%), almost all non‐mistletoe records are considered to be erroneous and, in most cases, probably represent the mistletoe host tree on which the larvae sometimes pupate. Of the eight major clades recognized in Delias, food plants have been recorded for six of these, although the majority of records (89%) are for three clades (hyparete, belladonna, nigrina). Optimization of the larval food plant data in the context of recent phylogenetic hypotheses for both butterflies and plants revealed little evidence of cospeciation at the higher systematic levels. The most parsimonious reconstruction was an origin of larval feeding on Loranthaceae, followed by at least six independent colonizations to Santalaceae + Viscaceae. In contrast to related pierids in the Aporiina associated with mistletoes in which further shifts from aerial‐stem mistletoes to distantly related plants (e.g. host trees parasitized by mistletoes) have facilitated differentiation at the generic level, there is no firm evidence to indicate that such secondary, monomorphic shifts have evolved in Delias. However, larvae of D. henningia (pasithoe group of belladonna clade) from Palawan and Luzon, the Philippines, appear to be polymorphic, feeding on both Loranthaceae and Euphorbiaceae.     

Fire frequency effects in a grassy woodland: Trees and grasses

Our study investigated the effects of fire frequency on the tree and grass components of Cumberland Plain Woodland, an endangered grassy eucalypt woodland in eastern Australia. We located three sites within each of three fire frequency classes (high, moderate, low) based on fire history and a similar time since last fire. For trees, we asked whether fire frequency affected density, eucalypt population structure, basal area and spatial patterning. For grasses, we tested for fire frequency effects on total abundance and the abundance of two common species. Density of trees was not significantly affected by fire frequency for juveniles, saplings or adult trees and neither was basal area per hectare. Features of a recruitment bottleneck model were present for eucalypts. There was a large pool of suppressed juveniles making up more than half the population at each fire frequency. Saplings were the smallest group and were susceptible to fire‐induced stem mortality, with particularly low numbers relative to juveniles and small trees where fire was frequent or intense. Despite this difference in sapling mortality, numbers of small trees did not differ with fire frequency. Saplings could transition to small trees in equal numbers if they did so more quickly at high than at lower fire frequencies, and if recruitment of saplings into the tree layer was controlled independently of fire frequency. The size hierarchy of small to medium eucalypt trees was homogeneous over eight of the nine sites, spatial patterning of adult trees was random tending to regular, and mean tree size decreased with density at all sites. These features of eucalypt population structure are indicative of possible resource competition which could regulate tree recruitment. Total grass cover index was high across all fire frequencies, with Themeda triandra dominating at high and moderate fire frequencies and Microlaena stipoides at low fire frequency.     

Competition as an amplifier of short-term vegetation responses to climate: an experimental test

1. Responses to a spring warming treatment were measured on five common herbaceous species grown in outdoor microcosms in Northern England. Although elevated temperature had a beneficial effect on canopy height and plant cover in monospecific cultures of all species, strongly divergent responses to warming occurred in mixtures.
2. We show that the effect of interspecific competition was to modify and amplify the vegetation response to the warming treatment through effects on phenology and morphological development.
3. The observed responses between the species to warming are consistent with predicted differential responses linked to genome size.
4. We conclude that the mechanism of competitive interaction proposed by Boysen-Jensen (1929 ) coupled with the insights related to genome size provide a basis for explaining and predicting the role of interannual variation in temperature in determining year to year fluctuations in the relative abundance of species in productive perennial herbaceous vegetation.     

Size and structure of populations of Oedura reticulata (Reptilia: Gekkonidae) in woodland remnants: Implications for the future regional distribution of a currently common species

Abstract The gecko Oedura reticulata is restricted in distribution to the southwest of Western Australia. Within this region O. reticulata occurs mainly in smooth-barked eucalypt woodland habitat, much of which has become fragmented by clearing. In this study, demographic characteristics of nine populations of O. reticulata that currently persist in eucalypt woodland remnants are assessed and compared with those of populations in three nature reserves. The remnants cover a range of sizes but are similar in age, tree species composition and vegetation structure. Population sizes vary considerably among remnants and are poorly correlated with remnant size or the number of smooth-barked eucalypts. Population size does not appear to be limited by habitat availability in most remnants. The number of adults of breeding age is small in most populations suggesting that they may be susceptible to stochastic extinction pressures. The poor dispersal ability of this species between remnants means that the possibility of recolonization of a remnant following an extinction event is unlikely. As a result, the occupancy rate of O. reticulata in remnant woodland is likely to decline.     

Phylogenetic relatedness as a tool in restoration ecology: a meta-analysis

Biotic interactions assembling plant communities can be positive (facilitation) or negative (competition) and operate simultaneously. Facilitative interactions and posterior competition are among the mechanisms triggering succession, thus representing a good scenario for ecological restoration. As distantly related species tend to have different phenotypes, and therefore different ecological requirements, they can coexist, maximizing facilitation and minimizing competition. We suggest including phylogenetic relatedness together with phenotypic information as a predictor for the net effects of the balance between facilitation and competition in nurse-based restoration experiments. We quantify, by means of a Bayesian meta-analysis of nurse-based restoration experiments performed worldwide, the importance of phylogenetic relatedness and life-form disparity in the survival, growth and density of facilitated plants. We find that the more similar the life forms of neighbouring plants are the greater the positive effect of phylogenetic distance is on survival and density. This result suggests that other characteristics beyond life form are also contained in the phylogeny, and the larger the phylogenetic distance, the less is the niche overlap, and therefore the less is the competition. As a general rule, we can maximize the success of the nurse-based practices by increasing life-form disparity and phylogenetic distances between the neighbour and the facilitated plant.     

Plant community responses to bison reintroduction on the Northern Great Plains,United States: a test of the keystone species concept

Keystone species restoration, or the restoration of species whose effect on an ecosystem is much greater than their abundance would suggest, is a central justification for many wildlife reintroduction projects globally. Following restoration, plains bison (Bison bison L.) have been identified as a keystone species in the tallgrass prairie ecoregion, but we know of no research to document similar effects in the mixed‐grass prairie where restoration efforts are ongoing. This study addresses whether Northern Great Plains (NGP) mixed‐grass prairie plant communities exhibit traits consistent with four central keystone effects documented for bison in the tallgrass prairie. We collected species composition, diversity, abundance, bare ground cover, and plant height data in three treatments: where livestock (Bos taurus L.) continuously grazed, livestock were removed for 10 years, and bison have been introduced and resident for 10 years. We observed mixed support for bison acting as keystone species in this system. Supporting the keystone role of bison, we observed higher species richness and compositional heterogeneity (β‐diversity) in the bison treatment than either the livestock retention or livestock removal treatments. However, we observed comparable forb, bare ground, and plant height heterogeneity between bison‐restored sites and sites where livestock were retained, contradicting reported keystone effects in other systems. Our results suggest that after 10 years of being restored, bison partially fulfill their role as a keystone species in the mixed‐grass prairie, and we encourage continued long‐term data collection to evaluate their influence in the NGP.     

Experiments on tree and shrub establishment in temperate grassy woodlands: Seedling survival

Abstract Experimental studies of the emergence of shrubs and trees in grassy woodlands on the New England Tablelands, New South Wales, Australia, showed that emergence of seedlings was determined by seed supply, seed predators and seed burial. The survival of these seedlings was then observed in an experiment to test the effects of previous land use, grazing by stock and grazing by other vertebrates. The fate of four eucalypts and six shrub species was followed over 5 years. Across all species more than 50% mortality occurred in the first 6 months prior to the imposition of grazing treatments. These deaths were attributed to the combined effects of insect defoliation, cold, and low soil moisture. Average mortality over all treatments showed two distinct trends: eucalypts and one unpalatable shrub (Leptospermum) had greater than 1% survival over 5 years, whereas Acacia, Cassinia, Indigophera, Lomatia and Xanthorrhoea either had very low or no survival after 5 years. The effect of livestock grazing on seedling numbers was rarely detected because of patchy emergence and mortality due to other causes. However, proportional hazard regression models showed that there was often an increased hazard associated with grazing or grazed landscapes. Overall, those species with high hazard coefficients associated with stock are rare in the landscape, whereas those with lesser risk are more common. Recruitment is likely to be an extremely rare event because the highest proportion of germinable seed sown that survived to a juvenile stage was 0.42% and the mean across all species was 0.12%. No natural recruitment of shrub species was observed over 5 years of observation, suggesting that recruitment is episodic and disturbance driven. Enhancing natural ‘regeneration’ of woody plants under these circumstances may be more challenging than simply fencing off remnants.     

Deuterium stable isotope ratios as tracers of water resource use: an experimental test with rock doves

Naturally-occurring deuterium stable isotope ratios can potentially be used to trace water resource use by animals, but estimating the contribution of isotopically distinct water sources requires the accurate prediction of isotopic discrimination factors between water inputs and an animals body water pool. We examined the feasibility of using estimates of water fluxes between a bird and its environment with a mass-balance model for the deuterium stable isotope ratio of avian body water (D_body) to predict isotopic discrimination factors. Apparent fractionation and thus discrimination factors were predicted to vary with the proportion of an animals total water losses than could be attributed to evaporative processes. To test our ability to predict isotopic discrimination, we manipulated water intake and evaporative water loss in rock doves (Columba livia) by providing them with fresh water or 0.15 M NaCl solution in thermoneutral or hot environments. After we switched the birds from drinking water with D=–95 VSMOW (Vienna Standard Mean Ocean Water) to enriched drinking water with D=+52 VSMOW, steady-state D_body was approached asymptotically. The equilibrium D_body was enriched by 10–50 relative to water inputs. After isotopic equilibrium was reached, the degree of enrichment was positively related (r²=0.34) to the fraction of total water loss that occurred by evaporation supporting the major prediction of the model. The variation we observed in discrimination factors suggests that the apparent fractionation of deuterium will be difficult to predict accurately under natural conditions. Our results show that accurate estimates of the contribution of different water sources to a birds body water pool require large deuterium isotopic differences between the sources.     

Reduced resource defence in an uncertain world: an experimental test using captive nutmeg mannikins

Recent models of economic defence in a group-foraging context predict that the frequency of aggressive interactions should decline as resource density increases, but empirical studies have provided only equivocal support for this prediction. We suggest that whether or not foragers have information concerning the location of patches will influence both the intensity of aggressive encounters and the effect that changes in food density will have on aggression. The intensity of aggression should be greatest when patch locations are known to all, making resources spatially predictable and the availability of alternatives more certain. When food is hidden, increasing the density of patches should have little effect on aggression levels, mostly as a result of the greater uncertainty about the availability of replacement food patches. To test these predictions, we investigated the effect of patch density on the use of aggressive behaviour in nutmeg mannikins, Lonchura punctulata, when food patches were either visible (signalled patch location) or hidden (unsignalled patch location) to all foragers. As predicted, we found that the intensity of aggressive encounters was higher when patch location was signalled than when it was not. Moreover, the effect of patch density on aggression depended on whether patch location was signalled or not. When patch location was unknown, the number of aggressive encounters was unaffected by changes in patch density, but when food location was signalled, increasing patch density resulted in the expected decline in the frequency of aggression.     

Shifts from competition to facilitation between a foundation tree and a pioneer shrub across spatial and temporal scales in a semiarid woodland

* Theoretical and empirical research has supported the hypothesis that plant-plant interactions change from competition to facilitation with increasing abiotic stress. However, the consistency of such changes has been questioned in arid and semiarid ecosystems. * During a drought in the semiarid south-western USA, we used observations and a field experiment to examine the interactions between juveniles of a foundation tree (Pinyon pine, Pinus edulis) and a common shrub (Apache plume, Fallugia paradoxa) in replicated areas of high and low stress. * The presence of F. paradoxa reduced P. edulis performance at low-stress sites, but had the opposite effect at high-stress sites. However, the intensity of the interactions depended on temporal variation in climate and age of P. edulis. Both above- and below-ground factors contributed to competition, while only above-ground factors contributed to facilitation. * These results support the hypothesis that interactions can change from competition to facilitation as abiotic stress increases in semiarid environments. A shift from competition to facilitation may be important for the recovery of P. edulis and other foundation species that have experienced large-scale mortality during recent droughts.     

Ecosystem wicks: Woodland trees enhance water infiltration in a fragmented agricultural landscape in eastern Australia

Abstract Since European settlement, Eucalyptus box woodlands have been substantially modified by agricultural practices, and in many areas in southern Australia are now restricted to scattered or clumped trees. We report here on a study to examine the impact of trees on water flow (infiltration) in an agricultural landscape with substantial areas of extant native vegetation. We examined infiltration through coarse‐ and fine‐textured soils within four landscape strata, the zones below Eucalyptus melliodora and Callitris glaucophylla canopies, the intertree zone dominated by perennial grasses and a landscape homogenized by cultivation and dominated by annual crops. We measured sorptivity, the early phase of water flow, and steady‐state infiltration with disc permeameters at two supply potentials. These different potentials enabled us to separate infiltration into (i) flow through large (biopores) and small pores and (ii) flow through small pores only where biopores are prevented from conducting water. On the fine‐textured soils, both sorptivity and steady‐state infiltration were significantly greater (approximately fivefold) under the timbered strata compared with the grassy slopes or cultivation. Differences were attributable to the greater proportion of macropores below the tree canopies compared with the nontimbered strata. The lack of a significant difference on the coarse‐textured soils, despite their macropore status, was attributed to differences in surface litter and plant cover, which would maintain continuous macropores at the surface and thus conduct large amounts of water. The tendency of slopes covered by cryptogamic crusts and grasses to shed run‐off and for the trees to absorb substantial quantities of water reinforced the important ecological service provided by trees, which moderates large run‐off events and captures small amounts of water leaking from the grassy patches. In the absence of these ‘ecosystem wicks’, run‐off would find its way into regional groundwater and contribute to rising salinity.     

Interactions between epiphytes during canopy soil formation: an experiment in a lower montane cloud forest of southeast Mexico

• In several montane forests around the world, epiphytes coexist in mats, sharing the rhizosphere and forming histosol-type soils rich in nutrients. The role of these epiphytes in the formation of canopy soil and the fitness costs that epiphytes face when cohabiting in these mats are unknown.
• In a lower montane cloud forest in central Veracruz, Mexico, a 2-year factorial experiment was carried out with the presence/absence of ramets of Phlebodium areolatum (Polypodiaceae), Tillandsia kirchhoffiana, T. multicaulis and T. punctulata (Bromeliaceae). We examined (i) which epiphyte species contribute to the formation of canopy soil, (ii) the role of epiphyte composition in the soil nutrient composition, and (iii) the fitness costs faced by epiphytes when cohabiting.
• Canopy soil formation highest when P. areolatum is present. Soil nutrient content does not change with epiphyte composition, is influenced by the microbiota, and P content decreases with the presence of epiphytes. The fitness costs show that the species compete, decreasing their survival and growth, but the competitive capacity differs between the species.
• We conclude that P. areolatum is an ecosystem engineer that promotes the creation of canopy soil but is a poor competitor. The results coincide with the model of succession by facilitation. Canopy soil is a slow-created component whose nutrient content does not depend on the epiphytic flora. In epiphyte mats, the dominant interactions are competitive, but there is also facilitation.

     

MAOA genotype,social exclusion and aggression: an experimental test of a gene–environment interaction

In 2002, Caspi and colleagues provided the first epidemiological evidence that genotype may moderate individuals' responses to environmental determinants. However, in a correlational study great care must be taken to ensure the proper estimation of the causal relationship. Here, a randomized experiment was performed to test the hypothesis that the MAOA gene promoter polymorphism (MAOA‐LPR) interacts with environmental adversity in determining aggressive behavior using laboratory analogs of real‐life conditions. A sample of 57 Caucasian male students of Catalan and Spanish origin was recruited at the University of Barcelona. Ostracism, or social exclusion, was induced as environmental adversity using the Cyberball software. Laboratory aggression was assessed with the Point Subtraction Aggression Paradigm (PSAP), which was used as an analog of antisocial behavior. We also measured aggressiveness by means of the reduced version of the Aggression Questionnaire. The MAOA‐LPR polymorphism showed a significant effect on the number of aggressive responses in the PSAP (F_1,53 = 4.63, P = 0.03, partial η² = 0.08), as well as social exclusion (F_1,53 = 8.03, P = 0.01, partial η² = 0.13). Most notably, however, we found that the MAOA‐LPR polymorphism interacts significantly with social exclusion in order to provoke aggressive behavior (F_1,53 = 4.42, P = 0.04, partial η² = 0.08), remarkably, the low‐activity allele of the MAOA‐LPR polymorphism carriers in the ostracized group show significantly higher aggression scores than the rest. Our results support the notion that gene–environment interactions can be successfully reproduced within a laboratory using analogs and an appropriate design. We provide guidelines to test gene–environment interactions hypotheses under controlled, experimental settings.     

Nutrient status,disturbance and competition: an experimental test of relationships in a wet meadow

Abstract. In many species-rich hay meadows in Central Europe, the traditional extensive (low input, low output) management is no longer economical and meadows are either fertilized or abandoned. Both these practices lead to changes in species composition and usually to a loss in species diversity. The response of a species-rich meadow plant community to fertilization, mowing and removal of dominant species was studied in a manipulative factorial experiment over four years. Both species richness and seedling recruitment were positively influenced by mowing and to a lesser extent by removal of the dominant species, Molinia caerulea, and were negatively influenced by fertilization. Fertilization caused an immediate increase in community biomass. Response to removal of the dominant species was delayed by one season, continued over the whole period, and by the fourth year the biomass reached a similar value as in the plots with Molinia present. Changes in species composition were evaluated by RDA for repeated measurements. The best and only significant predictor of species response to fertilization was plant height. This shows that with increased nutrient availability, nutrient limitation weakens and competition for light becomes the decisive factor. Competition for light appears to be more asymmetric than competition for nutrients, and consequently, it is more likely to drive inferior species to extinction. This is, together with seedling recruitment limitations, the most important cause of a decrease in species richness under high nutrient levels.     

Impacts of a native parasitic plant on an introduced and a native host species: implications for the control of an invasive weed

Background and Aims: While invasive species may escape from natural enemies in thenew range, the establishment of novel biotic interactions withspecies native to the invaded range can determine their success.Biological control of plant populations can be achieved by manipulationof a species' enemies in the invaded range. Interactions weretherefore investigated between a native parasitic plant andan invasive legume in Mediterranean-type woodlands of SouthAustralia. Methods: The effects of the native stem parasite, Cassytha pubescens,on the introduced host, Cytisus scoparius, and a co-occurringnative host, Leptospermum myrsinoides, were compared. The hypothesisthat the parasitic plant would have a greater impact on theintroduced host than the native host was tested. In a fieldstudy, photosynthesis, growth and survival of hosts and parasitewere examined. Key Results: As predicted, Cassytha had greater impacts on the introducedhost than the native host. Dead Cytisus were associated withdense Cassytha infections but mortality of Leptospermum wasnot correlated with parasite infection. Cassytha infection reducedthe photosynthetic rates of both hosts. Infected Cytisus showedslower recovery of photosystem II efficiency, lower transpirationrates and reduced photosynthetic biomass in comparison withuninfected plants. Parasite photosynthetic rates and growthrates were higher when growing on the introduced host Cytisus,than on Leptospermum. Conclusions: Infection by a native parasitic plant had strong negative effectson the physiology and above-ground biomass allocation of anintroduced species and was correlated with increased plant mortality.The greater impact of the parasite on the introduced host maybe due to either the greater resources that this host providesor increased resistance to infection by the native host. Thisdisparity of effects between introduced host and native hostindicates the potential for Cassytha to be exploited as a controltool.     

Competitive resource sharing: An experimental test of a learning rule for ESSs

Six sticklebacks (Gasterosteus aculeatus) distributed themselves between two simulated drift food patches in the ratio of patch profitabilities (Milinski 1979). The present study investigated the rule with which each fish decides to stay in one or the other patch, and whether the fish have more difficulties in reaching a decision when the prey is supplied irregularly as is the case under natural conditions. With frame by frame analysis of filmed experiments the hunting success and sampling effort of each individual fish could be determined continuously. The results support the hypothesis that the fish used the relative payoff sum (RPS) learning rule (Harley 1981; Regelmann 1984). There was a close correlation between all decisions of individual fish during a trial, whether to stay or leave a patch, and decisions predicted by the RPS learning rule using the actual experience of the fish. Differences in competitive ability between the fish influenced the dynamics of the distribution in the predicted way, the more successful competitors deciding earlier where to stay than the less successful ones. This had the effect that the good competitors as well as the poor ones were distributed between the patches in the ratio of patch profitabilities. It is also compatible with the RPS learning rule that the fish reached their decision as quickly whether they were supplied with prey regularly or irregularly.     

Ejaculate economics: an experimental test in a moth

Increasing evidence shows that spermatogenesis is costly. As a consequence, males should optimize the use of their sperm to maximize their reproductive outputs in their lifetime. However, experimental evidence on this prediction is largely lacking. Here, we examine how a male moth Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) responds to the presence of rivals or additional mates and how such response influences his lifetime reproductive fitness. We show that when rival males are present around a copulating pair, the male ejaculates more sperm to win a sperm competition battle but in such an environment he inseminates fewer females, sires fewer offspring and lives shorter. The opposite is the case when additional females are present during copulation. These findings reveal that elevated reproductive expenditure owing to sperm competition intensity is made at the expense of longevity and future reproduction.