Causal mechanisms underlying host specificity in bat ectoparasites

In parasites, host specificity may result either from restricted dispersal capacity or from fixed coevolutionary host-parasite adaptations. Knowledge of those proximal mechanisms leading to particular host specificity is fundamental to understand host-parasite interactions and potential coevolution of parasites and hosts. The relative importance of these two mechanisms was quantified through infection and cross-infection experiments using mites and bats as a model. Monospecific pools of parasitic mites (Spinturnix myoti and S. andegavinus) were subjected either to individual bats belonging to their traditional, native bat host species, or to another substitute host species within the same bat genus (Myotis). The two parasite species reacted differently to these treatments. S. myoti exhibited a clear preference for, and had a higher fitness on, its native host, Myotis myotis. In contrast, S. andegavinus showed no host choice, although its fitness was higher on its native host M. daubentoni. The causal mechanisms mediating host specificity can apparently differ within closely related host-parasite systems.     

Host specificity among Maculinea butterflies in Myrmica ant nests

Summary Ecological studies have been made of all 5 European species of Maculinea. These confirm that M. nausithous and M. rebeli live underground in Myrmica ant nests for 10 months of the year, as has long been known for the other 3 species. The main discovery was that each Maculinea species depends on a single, and different, host species of Myrmica. This specificity contradicts previous papers and scientific reviews of the relationship between Maculinea and ants. Therefore, early records are re-examined and 3 reasons are given to explain why most are misleading when applied to wild populations. Dependence on a single, rather than any, species of Myrmica explains why Maculinea populations exist in only a small minority of biotopes where their foodplants and Myrmica ants abound. It also explains the puzzling disappearance of Maculinea populations from apparently suitable sites. The discovery that M. alcon and M. rebeli depend on separate species of Myrmica that are not even closely related strengthens the argument that these butterflies are good species.     

Host specificity dynamics: observations on gyrodactylid monogeneans

The directly transmitted viviparous gyrodactylids have high species richness but low morphological and biological diversity, and many species are recorded from only a single host. They therefore constitute a guild of species ideal for studies of the evolutionary significance of host specificity. The group has the widest host range of any monogenean family, being found on 19 orders of bony fish. However, individual species range from narrowly specific (71% of 402 described species recorded from a single host) to extremely catholic (Gyrodactylus alviga recorded from 16 hosts). Gyrodactylid-host interactions extend from 60 mya (G. lotae, G. lucii) down to 150 years (G. derjavini on Oncorhynchus mykiss). Co-evolution with the host is comparatively rare within the gyrodactylids, but host switching or ecological transfer is common, and has been facilitated by the mixing of fish strains that followed glaciation. In this review, we consider the factors responsible for gyrodactylid specificity patterns, using examples from our work on salmonid gyrodactylids including G. salaris, responsible for major epidemics on wild Atlantic salmon (Salmo salar) in Norway since 1975, and G. thymalli from grayling and G. derjavini from trout.G. salaris has a wide host range with highest population growth rates on Norwegian salmon strains. However, growth rates are variable on both host strains and species, because of the multitude of micro- and macro-environmental factors influencing parasite mortality and fecundity. A better predictor of performance is the proportion of fishes of a strain which are innately resistant to the parasite, a measure which is negatively correlated with the time to peak infection in a host strain. Population growth rate is also negatively correlated with age of infection; the initial rate, therefore, predicts best the suitability of a fish as host for G. salaris. The host response to gyrodactylids appears to be the same mechanism in all salmonids with innate resistance as one end of a spectrum, but influenced by stress and probably under polygenic control. Hybrid experiments show that performance of G. salaris on a host is heritable, and usually intermediate between that of the parents. This host response mechanism, coupled with the initial parasite population growth on a fish, determines the host specificity, i.e. whether the fish will be susceptible, a responder or innately resistant. The use of population growth rate parameters allows comparison of different hosts as a resource for a gyrodactylid. In the case of G. salaris, East Atlantic and Baltic strains of Atlantic salmon are core hosts, but other salmonids can physiologically sustain infections for considerable periods, and may be important in parasite dispersal and transmission. A further group of non-salmonid fishes are unable to sustain G. salaris reproduction, but can act as transport hosts.Population growth parameters are very labile to stressors and environmental factors, particularly temperature and salinity, and also other aspects of host ecology and water quality. These factors may also influence the spectrum of hosts that can be infected under particular conditions, and probably favoured ecological transfer of gyrodactylids between host species in periglacial conditions. G. salaris may still be undergoing post-glacial range expansion (aided by anthropogenic spread) as shown by the increase in the species range over the last 25 years. The origin of G. salaris, G. teuchis and G. thymalli is discussed in relation to glacial refugiums during the last ice age.     

An experimental evaluation of host specificity: the role of encounter and compatibility filters for a rhizocephalan parasite of crabs

The encounter/compatibility paradigm of host specificity provides three qualitative pathways to the success or failure of a potential host-parasite interaction. It is usually impossible to distinguish between two of these (encounter and compatibility filters closed versus encounter filter open and compatibility filter closed) because unsuccessful infection attempts are difficult to observe in nature. We were able to open the encounter filter under experimental laboratory conditions. Our analytical system used the rhizocephalan barnacle, Sacculina carcini, a parasitic castrator of the European green crab, Carcinus maenas, and Pachygrapsus marmoratus, a native European crab that occurs with C. maenas but is not parasitized by S. carcini in nature. Penetration followed by unsuccessful infection of P. marmoratus crabs by parasitic barnacle larvae leaves a uniquely permanent record in the thoracic ganglion of the crabs. This provided us with a novel tool to quantify the encounter filter in a host-parasite system in nature. We demonstrated, in the laboratory, that the compatibility filter was closed and that, in nature, even where barnacle larvae were present, the encounter filter was also effectively closed. The closure of both filters in nature explains the failure of this potential host-parasite interaction, an outcome favored by selection in both host and parasite.     

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.     

Host specificity and molecular phylogeny of larval Digenea isolated from New Zealand and Australian topshells (Gastropoda: Trochidae)

The maintenance of strict host specificity by parasites when several closely related host species live in sympatry is poorly understood. Species of intertidal trochid snails in the genera Diloma, Melagraphia and Austrocochlea often occur together and are parasitised by a single digenean morphotype (Platyhelminthes, Trematoda), tentatively placed in Opecoelidae. Of the 10 trochid species (6 from New Zealand, 4 from Australia) we examined, six were found to be infected, and the prevalence of infection was as high as 17.5%. We obtained molecular data (mitochondrial 16S and nuclear rDNA internal transcriber spacer 2 sequences, representing 774 bp), to infer phylogenetic relationships amongst these Digenea. Our phylogeny separated the single morphotype into three clearly defined clades (which are almost certainly separate biological species): (i) those infecting two species of Austrocochlea from Tasmania, (ii) those infecting Diloma subrostrata in Otago and Southland, New Zealand, and (iii) those infecting all the parasitised New Zealand topshells (Melagraphia aethiops, D. subrostrata, Diloma nigerrima and Diloma arida) throughout the country. This last group comprised two subclades, one infecting only D. subrostrata and one infecting the other three species. Two D. subrostrata populations were each found to be infected by genetically distinct parasites, yet sympatric populations of the other snails were not necessarily infected. This study is thus the first to reveal cryptic species of digeneans in a single population of a molluscan first intermediate host. We point out also that the degree of host specificity would have been grossly underestimated if, in the absence of our genetic analysis, we had only considered digenean morphology. Our results shed light on the conditions that may favour switching among intermediate hosts in digeneans, and on the presence/absence of host specificity in these parasites.     

Infestation by Ergasilus coatiarus (Copepoda: Ergasilidae) in two Amazonian cichlids with new host record from Peru: An ectoparasites natural control approach

The occurrence of copepods ergasilid was investigated in two species of cichlids of economic importance for aquaculture in the Amazon region: Cichla monoculus and Chaetobranchus semifasciatus. The fish were collected from a semi-intensive fish farm, near the city of Nauta, Loreto State, Peru. Copepods were found in the gill filaments of 44 of 85 specimens (51.7%) of C. monoculus and in eight of 30 (26.6%) specimens of C. semifasciatus. The parasite was identified as Ergasilus coatiarus based on its morphological features. The occurrence did not vary significantly with host size (P ≥ 0.05) in both species. This is the first report of E. coatiarus parasitizing C. semifasciatus in the Amazon basin and the first report in C. monoculus from Peru. The high occurrence of these copepods in the present study points out the need of improving the strategies of parasitic prevention and control in order to better prevent future disease outbreaks.     

Host specificity ofCostesia yakutatensis [Hym.: Braconidae] on lepidoptera in peppermint and alfalfa

Seven species of Lepidoptera, common to alfalfa and peppermint in western Oregon, were tested as potential hosts forCotesia yakutatensis (Ashmead). Laboratory and field studies demonstrated that onlyAutographa californica (Speyer) andTrichoplusia ni (Hubner) were suitable. Percent parasitism, duration of oviposition, and number of parasitoids per host were highest whenC. yakutatensis was associated withA. californica.     

Influence of feeding habits in the endocrine pancreas of insectivore bat Pteronotus personatus and nectarivore bat Anoura geoffroyi: A comparative stereological and immunohistochemical study

Pteronotus personatus as an insectivore bat and has a diet that consists of a high protein diet, whereas the diet of Anoura geoffroyi, a predominantly nectarivore bat, is rich in simple sugars like sucrose, glucose and fructose. Considering that diet influences the activation of different pathways, which may influence morphological adaptations in the gastrointestinal system, the aim of this study was to compare the morphology of the endocrine pancreas in P. personatus and A. geoffroyi. For this, histological, stereological and immunohistochemical methods were used. In P. personatus, the average diameter of the pancreatic islet was 40.47 μm ± 13.94, while in A. geoffroyi was 88.16 μm ± 36.40. The total number of pancreatic islets in P. personatus was 26150 ± 2346 and in A. geoffroyi was 15970 ± 1666. In P. personatus, the volume density of the pancreatic islets was 3.4%± 2.6, whereas in A. geoffroyi the volume density was 6.1% ± 3.7. In addition, the immunodensity of the α, β and δ cells, in P. personatus was 25.8% ± 11.9, 35.5% ± 13.5, 3.9% ± 0.7, respectively, and in A. geoffroyi was 33.10% ± 12.7, 55.08% ± 7.4, 6.2% ± 4.6, respectively. In conclusion, the results of this study indicate differences in the pancreatic weight/body, weight ratio, diameter and volume density of pancreatic islets and in immunodensity of the β and α cells between both species, which have different dietary habits.     

Role of non-timber forest products in the economy of peripheral communities of knuckles national wilderness area of Sri Lanka: A farming systems approach

The Knuckles range of forests has been identified as a unique biological resource in Sri Lanka with much biodiversity. It is also important as a watershed which feeds the Mahaweli reservoir system. Having considered the present hazardous land-use practices such as cardamum production and shifting cultivation, the Knuckles range has been declared a national wilderness area. The Forest Department is in the process of preparing the management plan for the area. There are about 48 villages in the vicinity of the forest. This study attempts to estimate the composition of income in the peripheral communities, ticularly the extraction of non-timber forest products (NTFP) from this range of forests. Data were collected using structured questionnaires from 60 households in three villages in the vicinity of the forest. The rural economy is described using a farming systems approach and the net income contributed by each activity in the farming system was estimated. Results show that nine components of the farming system use resources from the village and the forest, with at least 47 plant species used by the villagers for different purposes. Cardamum production and shifting cultivation contribute significantly to the total income of the household, but the majority of households in the study depend upon forest resources to supply some portion of their income. Nevertheless, the monetary value of the annual income generated by NTFP is much lower than income from either cardamum production or shifting cultivation. If cardamum production and shifting cultivation are to be restricted under the conservation program, there could be additional pressure put on NTFP. Therefore, we suggest a study of the natural regen-eration capacity of selected NTFP; such information should be used in finalizing the management plan in the Knuckles range of forests.     

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.     

Involvement of host recognition by oncomiracidia and post-larval survivability in the host specificity of Heterobothrium okamotoi (Monogenea: Diclidophoridae)

Heterobothrium okamotoi, a monogenean gill parasite, shows high host specificity for tiger puffer, Takifugu rubripes. In the present study, in vivo and in vitro experimental infections were conducted using various fish species, including T. rubripes, to understand the mechanisms of specificity. In in vivo experiments, T. rubripes, grass puffer, Takifugu niphobles, olive flounder, Paralichthys olivaceus, and red sea bream, Pagrus major, were exposed to oncomiracidia of H. okamotoi labelled with a fluorescent dye, 5- (and -6) carboxyfluorescein diacetate succinimidyl ester, and the numbers of parasites on the gills and skin were recorded at intervals. Oncomiracidia were attached to gills and skin of all the experimental fish species immediately after exposure, and the infection intensity on T. rubripes was higher than that on T. niphobles and much higher than those on the other two species. After 2 days, the attached parasites remained on the gills of T. rubripes, but disappeared from the other hosts. During in vitro experiments, gill filaments excised from seven different fish species (four fish species used in the in vivo experiments and panther puffer, Takifugu pardalis, southern flounder, Paralichthys lethostigma and spotted halibut, Verasper variegates) were exposed to oncomiracidia and the attachment to each fish species and subsequent larval behaviour was observed. The percentage of post-larvae that attached to T. rubripes was slightly higher than those which attached to congeneric fish species and much higher than those of non-tetraodontid fish species. In vivo and in vitro experiments demonstrated that oncomiracidia of H. okamotoi have an affinity for their natural host, T. rubripes, and congeneric fish species. The disappearance of attached post-larvae from 'alien' hosts within 2 days during in vivo experiments suggested that host recognition by oncomiracidia and subsequent post-larval survivability are involved in the host specificity of H. okamotoi.     

Host specificity of the egg pupal parasitoid Fopius arisanus (Hymenoptera: Braconidae) in La Reunion

Fopius arisanus is a polyphagous parasitoid of Tephritidae, which has been recently introduced to La Réunion Island as part of a classical biological control programme. We carried out laboratory experiments to assess the host specificity of this parasitoid, initially reared on Bactrocera zonata, and then offered for parasitization the eight local tephritid pest species. Naive or experienced parasitoid females were given tephritid eggs in no choice tests. Fopius arisanus females parasitize all fly species but parasitism varies with host species. No adult wasps emerge from Bactrocera cucurbitae and the survival of this species is only slightly affected by parasitism. Dissections show that the late instars of this fly may eliminate the parasitoid by encapsulation. When developing on Ceratitis capitata, Ceratitis rosa, Dacus ciliatus, Dacus demmerezi, and Neoceratitis cyanescens, parasitoid survival rate ranges from 10 to 25%. Bactrocera zonata and Ceratitis catoirii are the best hosts, yielding parasitoid survival rates of more than 70% with no premature mortality. The egg-larval mortality of C. capitata, C. rosa, D. ciliatus, and N. cyanescens, and the pupal mortality of D. demmerezi, are significantly increased by parasitism. The size of emerging adults is affected by host species and is correlated to pupal weight. Bactrocera zonata would be a favorable host to support routine colonization of F. arisanus for mass production of this parasitoid.     

Host specificity of a generalist parasite: genetic evidence of sympatric host races in the seabird tick Ixodes uriae

Due to the close association between parasites and their hosts, many ‘generalist’ parasites have a high potential to become specialized on different host species. We investigated this hypothesis for a common ectoparasite of seabirds, the tick Ixodes uriae that is often found in mixed host sites. We examined patterns of neutral genetic variation between ticks collected from Black‐legged kittiwakes (Rissa tridactyla) and Atlantic puffins (Fratercula arctica) in sympatry. To control for a potential distance effect, values were compared to differences among ticks from the same host in nearby monospecific sites. As predicted, there was higher genetic differentiation between ticks from different sympatric host species than between ticks from nearby allopatric populations of the same host species. Patterns suggesting isolation by distance were found among tick populations of each host group, but no such patterns existed between tick populations of different hosts. Overall, results suggest that host‐related selection pressures have led to the specialization of I. uriae and that host race formation may be an important diversifying mechanism in parasites.     

Host specificity testing,release and establishment of Urophora solstitialis (L.) (Diptera: Tephritidae), a potential biological control agent for Carduus nutans L., in Australia

The seed‐fly Urophora solstitialis was imported into Australia from southern France. Previous workers had shown that this seed‐fly limited its attack to selected members of the genus Carduus. Host specificity studies revealed that no native Australian plant tested from the family Asteraceae was a suitable host plant. Permission to release the fly was granted, and it was released in three regions of the New South Wales tablelands. The fly underwent a second generation at two of the release sites, and reasons for its failure to do so at the third are suggested. It has successfully established itself in all three regions, having completed a full seasonal cycle in the field.     

Host specificity of microsporidia pathogenic to the gypsy moth, Lymantria dispar (L.): Field studies in Slovakia

Several species of microsporidia are important chronic pathogens of Lymantria dispar in Europe but have never been recovered from North American gypsy moth populations. The major issue for their introduction into North American L. dispar populations is concern about their safety to native non-target insects. In this study, we evaluated the susceptibility of sympatric non-target Lepidoptera to two species of microsporidia, Nosema lymantriae and Vairimorpha disparis, isolated from European populations of L. dispar and applied in field plots in Slovakia. Application of ultra low volume sprays of the microsporidia maximized coverage of infective spores in a complex natural environment and, thus, exposure of non-target species to the pathogens. Of 653 non-target larvae collected from plots treated with V. disparis in 2002, 18 individual larvae representing nine species in four families were infected. These plots were monitored for two subsequent seasons and V. disparis was not recovered from non-target species. Of 2571 non-target larvae collected in N. lymantriae-treated sites, one larva was found to be infected. Both species of microsporidia, particularly N. lymantriae, appear to have a very narrow host range in the field, even when an inundative technique is used for their introduction. V. disparis infections in L. dispar exceeded 40% of recovered larvae in the treated study sites; infection rates were lower in sites sprayed with N. lymantriae. Several naturally-occurring pathogens were recorded from the non-target species. The most common pathogen, isolated from 21 species in eight families, was a microsporidium in the genus Cystosporogenes.     

Host range testing and life history of the defoliator Hymenomima nr. memor: an unsuitable biological control agent for Schinus terebinthifolia in the U.S.A.

Host range of larvae of Hymenomima nr. memor (Lepidoptera: Geometridae) was examined in quarantine to evaluate its suitability as a biological control of Brazilian peppertree, Schinus terebinthifolia. Brazilian peppertree, S. terebinthifolia is an environmental and agricultural weed from South America that had invaded many subtropical and tropical areas of the world including Florida and Hawaii, USA. Laboratory life history and quarantine host range studies of H. memor were conducted with no-choice feeding tests. These tests included eight species of the Anacardiaceae and one species of Sapindaceae. Larvae of H. memor had five to six instars with each head capsule width increasing by 1.68-X. Development time from neonate to adult was 46.7?±?2.2 days. In host range tests, neonates completed development to the adult stage on all non-target species, except Toxicodendron radicans. Moreover, developmental times were delayed and pupal weights were reduced for larvae fed Spondias purpurea leaves. Due to the broad host range exhibited by H. memor larvae, this species will not be considered as a biological control agent of S. terebinthifolia in the continental U.S.A.     

Host specificity and aspects of the biology ofCalligrapha pantherina (Col.: Chrysomelidae), a biological control agent ofSida acuta [Malvaceae] andS. rhombifolia in Australia

The host specificity of a chrysomelid beetle,Calligrapha pantherina Stål, from Mexico, was verified in the CSIRO quarantine facilities in Brisbane, Australia on 80 species of plants in 28 families, either closely related to Malvaceae or in families containing economically important species.C. pantherina oviposited and developed successfully on its preferred host,S. acuta, and less successfully onS. rhombifolia andS. spinosa, all of which are weeds in Australia. Aspects of oviposition, larval development, and adult feeding and longevity ofC. pantherina are described.C. pantherina was released in Australia during 1989 and readily established onS. acuta but established at only a few sites onS. rhombifolia.     

DNA barcoding of endoparasitoid wasps in the genus Anicetus reveals high levels of host specificity (Hymenoptera: Encyrtidae)

The genus Anicetus includes economically important biocontrol agents that are introduced for control of soft and wax scale insect agricultural pests (Ceroplastes spp.). Understanding of host–parasitoid associations is critical to the successful outcome of their utilization in biological control projects. However, identification of these parasitoids is often difficult because of their small size and generally similar morphological features, and hence, studies on the host–parasitoid associations. Here, nucleotide sequence data were generated from the mitochondrial COI gene and the D2 region of 28S rRNA to assess genetic variation within and between species of Anicetus occurring in China. The results of this study support the use of the COI and the D2 region of 28S rRNA gene as useful markers in separating species of Anicetus, even in cases where morphological differences are subtle. On the other hand, the COI gene is also useful in recognizing species with much variation in morphology. DNA barcoding reveals high levels of host specificity of endoparasitoids wasps in the genus Anicetus. Our results indicate that each Anicetus species is adapted to a limited set of host species, or even are monospecific in their host choice.     

Host specificity of <Emphasis Type="Italic">Tectococcus ovatus</Emphasis> (Hemiptera: Eriococcidae), a potential biological control agent of the invasive strawberry guava, <Emphasis Type="Italic">Psidium cattleianum</Emphasis> (Myrtales: Myrtaceae), in Florida

Strawberry guava, Psidium cattleianum Sabine, is a woody tree or shrub native to coastal southeastern Brazil. Strawberry guava was introduced into Florida in the late 1800s as an ornamental species. The plant escaped cultivation and is invading natural areas throughout the southern half of the state. In addition to negative effects on Florida’s native ecosystems, strawberry guava also is a preferred host of the Caribbean fruit fly, Anastrepha suspensa Loew (Diptera: Tephritidae). In total, 57 plant species representing 21 families were included in the host range tests. First instar nymphs of Tectococcus ovatus Hempel fed on two closely related guava species, Brazilian guava (Psidium friedrichsthalianum O. Berg), and Costa Rican guava (Psidium guineense Sw.). However, none of the nymphs completed their development on these two non-target species. The results of the host specificity tests suggest that T. ovatus is a suitable candidate for classical biological control of strawberry guava in Florida.