The effect of the taxon and geographic range size of host eucalypt species on the species richness of gall-forming insects

Abstract This field study was designed to test whether the taxonomic group and geographic range size of a host plant species, usually found to influence insect species richness in other parts of the world, affected the number of gall species on Australian eucalypts. We assessed the local and regional species richness of gall-forming insects on five pairs of closely related eucalypt species. One pair belonged to the subgenus Corymbia, one to Monocalyptus, and three to different sections of Symphyomyrtus. Each eucalypt pair comprised a large and a small geographic range species. Species pairs were from coastal or inland regions of eastern Australia. The total number of gall species on eucalypt species with large geographic ranges was greater than on eucalypt species with small ranges, but only after the strong effect of eucalypt taxonomic grouping was taken into account. There was no relationship between the geographic range size of eucalypt species and the size of local assemblages of gall species, but the variation in insect species composition between local sites was higher on eucalypt species with large ranges than on those with small ranges. Thus the effect of host plant range size on insect species richness was due to greater differentiation between more widespread locations, rather than to greater local species richness. This study confirms the role of the geographic range size of a host plant in the determination of insect species richness and provides evidence for the importance of the taxon of a host plant.     

Transmission as a predictor of ecological host specificity with a focus on vertical transmission of microsporidia

Consideration of vertical transmission is particularly important for understanding the life cycles of entomopathogens that are naturally occurring in invertebrate populations, are a problem in beneficial insect colonies, or are under consideration as classical biological control agents. Empirical studies generally corroborate the evolutionary hypothesis that virulence should be relatively low for pathogen species that utilize vertical transmission as one mechanism for maintenance in the host population. Nevertheless, many entomopathogens with significant effects on host populations are vertically as well as horizontally transmitted. In addition to gaining a better understanding of pathogen-host interactions and population dynamics, studies of the host range and specificity of putative biological control agents can benefit by using transmission studies to better predict ecological host specificity from physiological data. Horizontal transmission requires a tightly organized host-pathogen relationship to succeed, but still involves, albeit restricted by host behavior and pathogen dosage, the physiological susceptibility of the nontarget host. Vertical transmission studies can provide increased stringency for determining the ecological host specificity of a species and may be one very accurate predictor of the ability of a pathogen to successfully host-switch when introduced into a na?ve population.     

Use of species richness estimators improves evaluation of understory plant response to logging: a study of redwood forests

The recovery of native understory plant communities after timber harvest has received a great amount of attention worldwide. However, most of these studies have ignored the potentially significant effects that differences in habitat across forest development may have on the species–area relationship, even though sampling efficacy among forest developmental stages may differ markedly as a result of equal sampling within stages. We examined vascular plant community structure in coastal redwood forests of northern California (USA) in forest patches in each of four forest stages that develop with forest age: Initiation, Closure, Mature, and Old-growth. We also used a set of traditional and novel approaches to assess sampling efficacy and guide our sampling design. We initially sampled 75 circular plots (0.04 ha) in stands within each forest stage across a 1,347 km² area. The species–area relationship suggested we had adequately sampled all stages after the first season of data collection, but non-parametric richness estimators (Incidence-based Coverage Estimator or ICE) and Michaelis–Menten means (MMMeans) rarefaction curves, indicated inadequate sampling of the older stages. Thus, we added four more samples within the Old-growth forest stage, establishing a more equitable sample of all stages. Our full data set demonstrated significant differences between plant communities in previously harvested forests (‘managed’ = Initiation, Closure, and Mature) and ‘unmanaged’ forests (Old-growth). Though several understory species were present in all stages, a suite of Old-growth indicator and restricted species suggest that recovery of some species may take long periods in managed forests. Interestingly, a separate suite of common understory species were lacking in Old-growth, and species richness in this stage was lower than in the mature managed forests. We discuss how this departure from the prediction of peak richness in Old-growth may relate to fire suppression in Old-growth forests. Lastly, analysis of our initial, smaller sampling of plots suggested that Old-growth plots were the least species rich. However, additional sampling of the Old-growth stage demonstrated that Old-growth plots were as rich as Closure plots. This has important implications for all studies concerned with measuring recovery after forest disturbances; stage-dependent habitat-mediated differences that affect sampling efficacy may not be captured by traditional sampling methods.     

Scientific advances in the analysis of direct risks of weed biological control agents to nontarget plants

Research on host specificity testing protocols over the last 10 years has been considerable. Traditional experimental designs have been refined and interpretation of the results is benefiting from an improved understanding of agent behavior. The strengths, weaknesses, and best practice for the different test types are now quite clearly understood. Understanding the concept of fundamental host range (the genetically determined limits to preference and performance) and using this to maximize reliability in predicting field host specificity following release (behavioral expression of the fundamental host range under particular conditions) are still inconsistently understood or adopted despite having been identified as the critical steps in analyzing the threats posed by biological control agents to the agriculture and biodiversity of novel environments. This needs to be consistently understood and applied so the process of testing can follow a recognized process of risk analysis from hazard identification (identifying life stages of the agent that pose a threat and defining their fundamental host range) to uncertainty analysis based on the magnitude (predicted field host specificity following release) and likelihood of threats (predicted actual damage and impact) to nontargets. Modern molecular techniques are answering questions associated with subspecific variation in biological control agents with respect to host use and the chance of host shifts of agents following release. Guidelines for assessment of nontarget impacts need to recognize and adopt such recent developments and emphasize a general increased understanding of the evolution of host choice and the phylogenetic constraints to shifts in host use. This review covers all these recent advances for the first time in one document, highlighting how inconsistent interpretation by biological control practitioners can be avoided.     

Biology and host range of Heterapoderopsis bicallosicollis: a potential biological control agent for Chinese tallow Triadica sebifera

Chinese tallow, Triadica sebifera, is an invasive weed that infests natural and agricultural areas of the south-eastern USA. A candidate insect for biological control of Chinese tallow has been studied under quarantine conditions. The biology and host range of a primitive leaf-feeding beetle, Heterapoderopsis bicallosicollis, were examined between July 2008 and February 2010. H. bicallosicollis adults fed on 16 of 29 non-target species and survived >10 days on all 29 species, more than the number of days that adults survived when deprived of food. Additionally, adult feeding, oviposition and partial larval development occurred on a Florida endangered species, Heterosavia bahamensis. These data indicated that H. bicallosicollis adult feeding and oviposition may not be limited to the target weed and included several valued North American natives. Therefore, as its release could pose unacceptable risks, testing of this species was discontinued and the quarantine colony destroyed.     

The use of incidence-based species richness estimators,species accumulation curves and similarity measures to appraise ethnobotanical inventories from South Africa

The incorporation of suitable quantitative methods into ethnobotanical studies enhances the value of the research and the interpretation of the results. Prediction of sample species richness and the use of species accumulation functions have been addressed little in applied ethnobotany. In this paper, incidence-based species richness estimators, species accumulation curves and similarity measures are used to compare and predict species richness, evaluate sampling effort and compare the similarity of species inventories for ethnobotanical data sets derived from the trade in traditional medicine in Johannesburg and Mpumalanga, South Africa. EstimateS was used to compute estimators of species richness (e.g. Jackknife), rarefaction curves, species accumulation curves and complimentarity. Results showed that while the Michaelis–Menten Means estimator appeared to be the best estimator because the curve approached a horizontal asymptote, it was not able to accurately predict species richness for one of the data sets when two of its subsamples were individually tested. Instead, the first-order Jackknife estimator best approximated the known richness.     

Demersal fish parasite fauna around the South Shetland Islands: high species richness and low host specificity in deep Antarctic waters

A total of nine Antarctic fish species belonging to five families were examined for their endohelminth parasite fauna. The fishes Parachaenichthys charcoti (Bathydraconidae), Chaenocephalus aceratus (Channichtyidae), Paradiplospinus gracilis (Gempylidae), Muraenolepis microps (Muraenolepididae), Gobionotothen gibberifrons, Lepidonotothen larseni, L. nudifrons, L. squamifrons, and Trematomus eulepidopus (Nototheniidae) were caught between 80 and 608 m trawling depth off the Antarctic Peninsula (Elephant Island, King George Island) in 1996. Nineteen different parasites species comprising five Digenea, two Cestoda, four Nematoda, and eight Acanthocephala were found. Pseudophyllidean cestodes, the nematodes Contracaecum radiatum and C. osculatum as well as the acanthocephalan Corynosoma bullosum were the most common, infesting eight of the fish species studied with prevalences reaching 100%. Pseudoterranova decipiens s.l. was the only parasite that was isolated from all studied fish species; however, at a lower intensity. The observed parasite host specificity was low, and the species richness in a single fish ranged from one to eleven in a C. aceratus. This icefish and the moray cod M. microps were the most heavily infested fish, harbouring many adult and larval parasitic stages. The benthodemersal P. gracilis had only two larval parasite species, while the nototheniids had very similar parasite communities, harbouring a total of 8–14 species. Larval mammalian parasites were found to utilize fish, especially the nototheniids and channichthyids, as a common transmission route into their final hosts. The fish parasites parallel explored different benthic host systems to reach the most suitable host. In contrast to the coast and continental shelf, the meso/bathypelagiac zone appears to be species poor and is inhabited by few larval forms. The fish parasite fauna off the South Shetland Islands can be characterized by generalistic parasites that distribute within Antarctic waters according to the feeding ecology and depth range of their teleost hosts, not only horizontally but also extending vertically into the deep sea.     

Assessing the accuracy of species distribution models to predict amphibian species richness patterns

1. Evaluating the distribution of species richness where biodiversity is high but has been insufficiently sampled is not an easy task. Species distribution modelling has become a useful approach for predicting their ranges, based on the relationships between species records and environmental variables. Overlapping predictions of individual distributions could be a useful strategy for obtaining estimates of species richness and composition in a region, but these estimates should be evaluated using a proper validation process, which compares the predicted richness values and composition with accurate data from independent sources. 2. In this study, we propose a simple approach to estimate model performance for several distributional predictions generated simultaneously. This approach is particularly suitable when species distribution modelling techniques that require only presence data are used. 3. The individual distributions for the 370 known amphibian species of Mexico were predicted using maxent to model data on their known presence (66,113 presence-only records). Distributions were subsequently overlapped to obtain a prediction of species richness. Accuracy was assessed by comparing the overall species richness values predicted for the region with observed and predicted values from 118 well-surveyed sites, each with an area of c. 100 km(2), which were identified using species accumulation curves and nonparametric estimators. 4. The derived models revealed a remarkable heterogeneity of species richness across the country, provided information about species composition per site and allowed us to obtain a measure of the spatial distribution of prediction errors. Examining the magnitude and location of model inaccuracies, as well as separately assessing errors of both commission and omission, highlights the inaccuracy of the predictions of species distribution models and the need to provide measures of uncertainty along with the model results. 5. The combination of a species distribution modelling method like maxent and species richness estimators offers a useful tool for identifying when the overall pattern provided by all model predictions might be representing the geographical patterns of species richness and composition, regardless of the particular quality or accuracy of the predictions for each individual species.     

Egg size of skippers (Lepidoptera: Hesperiidae) in relation to their host specificity and to leaf toughness of host plants

The adaptive significance of egg size of skippers (Lepidoptera; Hesperiidae) in Japan was evaluated in relation to the leaf toughness of their major host grasses. The hesperids that fed on tougher grasses laid larger eggs. Hesperids that laid larger eggs were larger in body size, but lower in fecundity. They also had a wider host range. Thus, despite the lower fecundity, hesperids may benefit from large eggs by having a wider host range of larvae. Grass feeders had wider range of host plants than broadleaf feeders.     

Evaluating the performance of species richness estimators: sensitivity to sample grain size

1. Fifteen species richness estimators (three asymptotic based on species accumulation curves, 11 nonparametric, and one based in the species-area relationship) were compared by examining their performance in estimating the total species richness of epigean arthropods in the Azorean Laurisilva forests. Data obtained with standardized sampling of 78 transects in natural forest remnants of five islands were aggregated in seven different grains (i.e. ways of defining a single sample): islands, natural areas, transects, pairs of traps, traps, database records and individuals to assess the effect of using different sampling units on species richness estimations. 2. Estimated species richness scores depended both on the estimator considered and on the grain size used to aggregate data. However, several estimators (ACE, Chao 1, Jackknifel and 2 and Bootstrap) were precise in spite of grain variations. Weibull and several recent estimators [proposed by Rosenzweig et al. (Conservation Biology, 2003, 17, 864-874), and Ugland et al. (Journal of Animal Ecology, 2003, 72, 888-897)] performed poorly. 3. Estimations developed using the smaller grain sizes (pair of traps, traps, records and individuals) presented similar scores in a number of estimators (the above-mentioned plus ICE, Chao2, Michaelis-Menten, Negative Exponential and Clench). The estimations from those four sample sizes were also highly correlated. 4. Contrary to other studies, we conclude that most species richness estimators may be useful in biodiversity studies. Owing to their inherent formulas, several nonparametric and asymptotic estimators present insensitivity to differences in the way the samples are aggregated. Thus, they could be used to compare species richness scores obtained from different sampling strategies. Our results also point out that species richness estimations coming from small grain sizes can be directly compared and other estimators could give more precise results in those cases. We propose a decision framework based on our results and on the literature to assess which estimator should be used to compare species richness scores of different sites, depending on the grain size of the original data, and of the kind of data available (species occurrence or abundance data).     

Phelipanche ramosa (L.) Pomel populations differ in life-history and infection response to hosts

Phelipanche ramosa (broomrape) is a root-holoparasitic angiosperm that attacks a wide number of annual crops. According to the host, the duration of its life cycle can range from 14 weeks (on tomato/tobacco) to 40 weeks (on oilseed rape). We conducted a cross-infection experiment to evaluate the intensity and kinetics of infection of broomrape populations. Two parasitic populations, P-long collected on oilseed rape and P-short collected on tobacco, were cultivated on two crops (their natural and outsider hosts). After, 4, 8, 12 and 16 weeks, the intensity of infection and the distribution of developmental stages (fixation to fructification) of broomrape were determined. The two broomrape populations showed distinct patterns of intensity and kinetics of infectivity. P-long showed a higher infection success in the early stages than P-short. Both parasitic populations showed a higher aggressiveness on their natural hosts than on outsider hosts. Only the P-short population completed its life cycle on the two hosts, and with similar development rates on both hosts, while the P-long population was unable to complete its life-cycle during the 16 weeks of the experiment. It is suggested that the shift in host specificity that allowed P. ramosa to infect oilseed rape in recent times has led to a divergence of pathovars having different life-cycle durations.     

Morphological variation and host range of two Ganoderma species from Papua New Guinea

Two species of Ganoderma belonging to different subgenera which cause disease on oil palms in PNG are identified by basidiome morphology and the morphology of their basidiospores. The names G. boninense and G. tornatum have been applied. Significant pleiomorphy was observed in basidiome characters amongst the specimens examined. This variation in most instances did not correlate well with host or host status. Spore morphology appeared uniform within a species and spore indices varied only slightly. G. tornatum was found to have a broad host range whereas G. boninense appears to be restricted to palms in Papua New Guinea.     

Application of species richness estimators for the assessment of fungal diversity

Species richness and distribution patterns of wood-inhabiting fungi and mycetozoans (slime moulds) were investigated in the canopy of a Central European temperate mixed deciduous forest. Species richness was described with diversity indices and species-accumulation curves. Nonmetrical multidimensional scaling was used to assess fungal species composition on different tree species. Different species richness estimators were used to extrapolate species richness beyond our own data. The reliability of the abundance-based coverage estimator, Chao, Jackknife and other estimators of species richness was evaluated for mycological surveys. While the species-accumulation curve of mycetozoans came close to saturation, that of wood-inhabiting fungi was continuously rising. The Chao 2 richness estimator was considered most appropriate to predict the number of species at the investigation site if sampling were continued. Gray's predictor of species richness should be used if statements of the number of species in larger areas are required. Multivariate analysis revealed the importance of different tree species for the conservation and maintenance of fungal diversity within forests, because each tree species possessed a characteristic fungal community. The described mathematical approaches of estimating species richness possess great potential to address fungal diversity on a regional, national, and global scale.     

Multiple factors promoting narrow host range in the sea hare,Aplysia californica

Summary Juvenile sea hares, Aplysia californica, utilize only the red algae Plocamium cartilagineum and Laurencia pacifica as host plants at Santa Catalina Island, CA. I tested three hypotheses which might account for this pattern of host choice: 1) A. californica specialize on the algae on which they grow best, 2) A. california specialize on algae from which they acquire secondary compounds that protect them from predators, and 3) A. californica specialize on certain algae in order to lower their encounter rates with predators. The results suggested that host range in the Aplysia californica system is affected by more than one factor. The first hypothesis was supported. A. californica of three size classes grew well on Plocamium, but could not grow at all on most other species of algae. Larger A. californica were able to grow on species of algae that smaller ones could not. The second hypothesis was also supported. Small A. californica grown on Plocamium were rich in terpenes. Small A. californica grown on Ulva sp. were terpene-free. Rock wrasses, Halichoeres semicinctus, were more likely to eat Ulva-fed A. californica than Plocamium-fed A. californica. Other fish and lobster, Panulirus interruptus, did not discriminate between the two groups. Kelp bass, Paralabrax clathratus, which were force-fed Ulva-fed A. californica regurgitated them less often, and after digesting them more completely, than did Paralabrax force-fed Plocamium-fed A. californica. The third hypothesis was rejected. A. californica located on Plocamium were not more cryptic to the opisthobranch Aglaja inermis (navanax), or to the pomacentrid fish, Hypsypops rubicundus, than were A. californica located on other algae. In addition, navanax, a specialist predator of opisthobranchs, was significantly more abundant on Plocamium than on other algae.     

Factors determining the host plant range of the phytophagous mite, Tetranychus urticae (Acari: Tetranychidae): a method for quantifying host plant acceptance

The host plant acceptance of the phytophagous mite Tetranychus urticae was experimentally quantified. Host plant acceptance is described as the proportion of adult females settling on the test plant on which they have been placed. On the other hand, the host plant suitability of T. urticae on different plant species is expressed as the mean number of eggs produced by the females within 5 days (hereafter 'fecundity'). An inbred T. urticae line was tested with regard to host plant acceptance and fecundity on 11 potential host plants. These two variables were positively correlated across host plants; host plant species on which the fecundity was low were also those on which females settled less readily compared to host plants with high fecundity. The characteristics of host plant acceptance of the T. urticae are discussed in light of their potential food resource under natural conditions.     

Intermediate host switches drive diversification among the largest trematode family: evidence from the Polypipapiliotrematinae n. subf. (Opecoelidae), parasites transmitted to butterflyfishes via predation of coral polyps

Podocotyloides stenometra Pritchard, 1966 (Digenea: Opecoelidae) is the only trematode known to infect anthozoan corals. It causes disease in coral polyps of the genus Porites Link (Scleractinia: Poritidae) and its life-cycle depends on ingestion of these polyps by butterflyfishes (Perciformes: Chaetodontidae). This species has been reported throughout the Indo-Pacific, from the Seychelles to the Galápagos, but no study has investigated whether multiple species are involved. Here, we recollect P. stenometra from its type-host and type-locality, in Hawaiian waters, and describe four new species from examination of 768 butterflyfishes from French Polynesia. On the basis of morphology, phylogeny and life-history, we propose Polypipapiliotrema Martin, Cutmore & Cribb n. gen. and the Polypipapiliotrematinae Martin, Cutmore & Cribb n. subf., for P. stenometra (Pritchard) n. comb., P. citerovarium Martin, Cutmore & Cribb n. sp., P. hadrometra Martin, Cutmore & Cribb n. sp., P. heniochi Martin, Cutmore & Cribb n. sp., and P. ovatheculum Martin, Cutmore & Cribb n. sp. Given the diversity uncovered here and the ubiquity, abundance and diversity of butterflyfishes on coral reefs, we predict that Polypipapiliotrema will prove to comprise a rich complex of species causing disease in corals across the Indo-Pacific. The unique life-cycle of these taxa is consistent with phylogenetic distinction of the group and provides evidence for a broader basis of diversification among the family. We argue that life-cycle specialisation, in terms of adoption of disparate second intermediate host groups, has been a key driver of the diversification and richness of the Opecoelidae, the largest of all trematode families and the group most frequently encountered in coral reef fishes.     

Intermediate host availability masks the strength of experimentally-derived colonisation patterns in echinostome trematodes

A fundamental goal of parasite evolutionary ecology is to elucidate patterns of host use and determine the underlying mechanisms of parasite colonisation. In order to distinguish the relative contributions of host encounter rates and host compatibility to infection outcomes, we compared host use in both field and experimental laboratory settings. Two years of bi-weekly snail sampling at a freshwater pond demonstrated fluctuating availability among three potential second intermediate snail host species and suggested that two trematode species (Echinostoma revolutum and Echinoparyphium sp.) did not colonise the three potential snail host species, Lymnaea elodes, Physa gyrina and Helisoma trivolvis, differentially. However, a series of experimental infections demonstrated that both parasites colonised H. trivolvis more so than the other two host species. Thus, more echinostome parasites utilised snail hosts that cannot serve as their first intermediate host. In experimental infections, host size and vagility were not strong determinants of infection. By utilising field and laboratory approaches, we were able to compare the strength of host compatibility under controlled conditions with patterns of infection in nature. Based on the results from these studies, it appears that host encounter is the primary mechanism dictating infection outcomes in the field.     

Restricted host range of the weevil Aubeonymus mariaefranciscae, a new sugar-beet pest

The extent of feeding, fecundity and fertility of a new sugar beet pest, Aubeonymus mariaefranciscae Roudier (Coleoptera: Curculionidae), on a range of plants from different families was determined. We have found that the adult weevils were only able to feed and oviposit on two closely related cultivated plants, beet and sugar beet, both subspecies of Beta vulgaris L., suggesting a restricted host range for this curculionid. None of 16 secondary metabolites (widely distributed among plants, and representative of different chemical groups), nor any of 19 non-host extracts from 13 different families, deterred the adults of this curculionid from feeding on sugar beet. We found a phagostimulatory effect when beet and sugar beet homogenates from various plant tissues were supplied to adult weevils on weeds that they do not feed on. Conversely, no effect was observed when sucrose was provided. These findings suggest that host selection by A. mariaefranciscae adults is due to the presence of an undetermined phagostimulant compound(s) in beet and sugar beet, instead of the recognition of deterrent substances among the non-host plants.