Local variation in the thistle species Cirsium grayanum affects its utilization by the herbivorous ladybird beetle Henosepilachna niponica

Host plants are the most critical environmental factor for phytophagous arthropods. Adaptation to a novel host will alter the distributional range of an herbivore to include the area the novel host covers, and might promote divergence between populations utilizing the old and new hosts. On the Oshima Peninsula, Hokkaido Island, northern Japan, the ladybird beetle Henosepilachna niponica (Lewis) (Coleoptera: Coccinellidae) usually occurs exclusively on the thistle Cirsium alpicola Nakai (Asteraceae), which is distributed in the southernmost part of the island, although some other Cirsium species often grow in sympatry. At least at one site (Assabu), however, H. niponica depends on another thistle, Cirsium grayanum (Maxim.) Nakai, the most abundant and widely distributed thistle species on the Oshima Peninsula. We examined adult feeding acceptance and preference, and larval performance, in two populations of H. niponica that utilize different species of thistles (the Assabu population occurs on C. grayanum; the Shiriuchi population, on C. alpicola) by testing three kinds of thistles (C. grayanum from Assabu, C. grayanum from Shiriuchi, and C. alpicola from Shiriuchi). Results strongly suggested that among the three plants tested, C. grayanum from Assabu was the most suitable host for H. niponica. We concluded that local differences in C. grayanum mainly explain why it is not used at Shiriuchi, but that there are also local differences in host use among the beetle populations, suggesting some degree of local adaptation. This suggests that different selective regimes associated with different host plants might promote divergence among beetle populations. We conclude that H. niponica is at present unlikely to expand its range to the whole range of C. grayanum.     

Distribution and host specificity of the thistle‐feeding tortoise beetle Cassida vibex (Coleoptera: Chrysomelidae) in southwestern Hokkaido,northern Japan

The thistle‐feeding tortoise beetle Cassida vibex (Coleoptera: Chrysomelidae) is widespread in the Palearctic region. In Japan, this species has been recorded only from the largest island, Honshu. Here we report the occurrence of this species on Hokkaido, the most northern main island of Japan, along with detailed distributional records from the southwestern part of the island (southern Oshima Peninsula) and information on host plants. We also present the results of laboratory experiments on adult feeding preference and larval developmental performance to determine the specificity of C. vibex for four thistle species (Cirsium grayanum, Cir. alpicola, Cir. yezoense and Cir. aomorense; Asteraceae) common in southwestern Hokkaido. Cassida vibex was detected at only four sites among 97 sites investigated on the Oshima Peninsula, in strong contrast to the common occurrence of the congeneric thistle feeder C. rubiginosa (74 of 97 sites). Cassida vibex was found mainly on Cir. alpicola in the wild, but adult beetles showed no feeding preference for this or three other thistle species when given a choice in the laboratory. Larval performance of C. vibex was similar on Cir. alpicola, Cir. yezoense and Cir. aomorense but significantly lower on Cir. grayanum. Since Cir. alpicola often forms large clumps of individual plants, whereas Cir. yezoense and Cir. aomorense are scattered in their habitats, and C. vibex was collected only at sites where there were dense patches of Cir. alpicola, we postulate that the persistence of C. vibex is possible only where a substantial amount of suitable host plants is available.     

Individual variation in two host plants of the ladybird beetle,Epilachna pustulosa (Coleoptera: Coccinellidae)

Individual variation in two species of host plants (thistle,Cirsium kamtschaticum, and blue cohosh,Caulophyllum robustum) of the herbivorous ladybird beetleEpilachna pustulosa was examined under laboratory conditions for their acceptability to adult beetles as a food resource, for adult preference and for larval performance. When clones of these plants were subjected to non-choice tests using posthibernating female beetles, there was found to be significant intraspecific variation among clones in terms of their acceptability, but interspecific variation was not detected. Significant intraspecific as well as interspecific variation were frequently detected in the two host plants when clones of these plants were subjected to choice tests using posthibernating female beetles; the magnitude of interspecific plant variation for beetle preference is not necessarily larger than that of intraspecific plant variation. Individual variation across plant species with respect to beetle larval performance was also significant. A positive correlation between adult preference and larval performance is suggested across the two taxonomically remote host plant species, thistle and blue cohosh, although this needs further investigation.     

Host Specificity of Altica carduorum Guer. Chrysomelidae: Coleoptera , a Defoliator of Cirsium arvense L. Scop. Asteraceae from North-western China

Traditionally, insects are not approved for release in North America as agents for the biological control of weeds if the larvae complete development on desirable plants in no-choice tests. This criterion excludes the use of the leaf-feeding beetle Altica carduroum Guer. since it can complete development on all Cirsium spp., including rare North American species, although in nature it is only known from the target weed Cirsium arvense. Host suitability is the result of a sequence of selection factors, and this study shows that most discrimination between thistle species is the result of host finding and egg production, and not larval developmental ability. In no-choice tests, the larvae completed development on 18 Cirsium species in five sections of the genus that were tested and on Silybum marianum, but not on other genera in the tribe Cardueae or more distantly related Asteraceae. The adult food consumption on C. arvense was two to eight times greater than on other Cirsium species, many of which, such as C. pitcheri and C. drummondii, received no feeding by over 50 of the beetles in the first 24 h. The best North American thistle for egg production was C. scariosum, but this was only 28 of that on C. arvense. The preference for C. arvense in multiple-choice tests increased with the size 2 of the arena. In the largest, an area of 6 m , 80 of the beetles and eggs were found on C. arvense and none were found on plants outside the subtribe Carduineae. Thus, as theory predicts, host discrimination between thistle species takes place at earlier stages than suitability for larval development, and this should be considered when determining the safety of candidate agents.     

Preference-performance linkage in a herbivorous lady beetle: consequences of variability of natural enemies

We investigated the relationship between oviposition preference and offspring performance in a herbivorous lady beetle Epilachna pustulosa on two co-occurring plant species, thistle Cirsium kamtschaticum and blue cohosh Caulophyllum robustum, in 1994 and 1995. The relative importance of bottom-up effects by host plants and top-down effects by natural enemies on offspring performance were determined using field and laboratory experiments. In both years, egg density on blue cohosh was significantly higher than on thistle. A laboratory experiment demonstrated that larval survival from hatching to adult emergence was significantly higher, and developmental period shorter when larvae were reared on blue cohosh compared to thistle. The positive preference-performance linkage varied between years in the field. Top-down effects had a different impact on larval survival on the two host plant species. Arthropod predators, a lady beetle Harmonia axyridis and an earwig Forficula mikado, considerably depressed immature survival on thistle, while they were negligible on blue cohosh. Although the lack of effective predation increased larval survival on blue cohosh, it led to defoliation due to increased larval feeding late in the season. Because of severe intraspecific competition, old larvae had significantly lower survival on blue cohosh than on thistle. In 1994, as larval survival decreased due to defoliation on blue cohosh, the overall survival rate was significantly higher on thistle than on blue cohosh. This survival pattern was opposite to that found in the laboratory experiment. In contrast, in 1995, the increase in predatory lady beetles on thistle caused greater larval mortality. Thus, the overall survival was significantly lower on thistle than on blue cohosh, although severe intraspecific competition occurred on blue cohosh as it had in 1994. Consequently, the offspring performance on the two host plants is largely determined by the relative importance of arthropod predation determining larval survival on thistle and host plant defoliation reducing late larval survival on blue cohosh. These results indicate the important role of spatial and temporal variability of natural enemies on the preference-performance linkage of herbivorous insects. Received: 19 August 1998 / Accepted: 11 January 1999     

Adaptive change and conservatism in host specificity in two local populations of the thistle-feeding ladybird beetle Epilachna niponica

We investigated geographic differences in the host specificity of Epilachna niponica Lewis (Coleoptera: Coccinellidae). The Yuwaku population feeds mainly on Cirsium matsumurae Nakai (Asteraceae) and secondarily on Cirsrium kagamontanum Nakai. The Asiu population, located 150 km away from the Yuwaku, feeds exclusively on C. ashiuense Yokoyama et T. Shimizu. Under laboratory conditions, we examined the differences between the two populations in adult feeding acceptance, adult feeding preference, and larval performance, using several closely related thistle species and varieties, including their native hosts. In the Asiu population, adult beetles clearly avoided the host of the Yuwaku population, C. kagamontanum, and no larvae were able to complete their development, whereas in the Yuwaku population, adults accepted and even preferred it to some other thistle species, and about 10% of first instar larvae became adults. This indicates that the Yuwaku population evolved its feeding preference and physiological adaptation to C. kagamontanum through a utilization of this low‐ranked host under natural conditions. Apart from C. kagamontanum, the two populations showed a similar host susceptibility pattern, indicating that this ladybird beetle has a conserved hierarchy in feeding preference and growth performance. We also observed adult leaf choice behavior when given different thistle species, and found that difference in biting rate after palpation determined the leaf areas consumed, implying that factors on the leaf surface played an important role in the choice.     

Leaf traits of congeneric host plants explain differences in performance of a specialist herbivore

1. Within the host range of herbivorous insects, performance hierarchies are often correlated with relatedness to a primary host plant, as plant traits are phylogenetically conserved. Therefore, it was hypothesised that differences in herbivore performance on closely related plant species are due to resistance traits that vary in magnitude, rather than in the nature of the traits. 2. This hypothesis was tested by manipulating putative resistance traits of three congeneric thistle species (Cirsium arvense, Cirsium palustre, and Cirsium vulgare) and assessing the performance of the oligophagous, leaf‐feeding beetle, Cassida rubiginosa. Measurements were done of survival, weight gain, and development time of the beetle on its primary host, C. arvense, and two alternative hosts under low and high nutrient availability, and on shaved and unshaved leaves. 3. Survival of C. rubiginosa was strongly dependent on plant species with final mean survival rates of 47%, 16%, and 8% on C. arvense, C. palustre, and C. vulgare, respectively. Survival was primarily explained by leaf trichome densities, and to a lesser extent by specific leaf area. Leaf flavonoid concentrations did not explain differences in beetle survival, and there were no differences in beetle weight gain or development time of individuals that survived to adulthood. 4. No beetles survived on unshaved (hairy) C. vulgare plants, but manipulating leaf trichome densities of the thistle species by shaving the leaves moderated the plant‐specific resistance, and equalised the survival rates. Survival of C. rubiginosa on alternative congeneric hosts was explained by a common physical resistance trait that varied in magnitude.     

Oviposition site selection by herbivorous beetles: a comparison of two thistle feeders, Cassida rubiginosa and Henosepilachna niponica

In insects that feed on plants in both adult and larval stages, it is often difficult to distinguish oviposition preference from adult feeding preference, because oviposition can occur at or in proximity to feeding sites. In the present study, characteristics of oviposition site selection of two beetle species, Cassida rubiginosa Müller (Coleoptera: Chrysomelidae) and Henosepilachna niponica (Lewis) (Coleoptera: Coccinellidae), were investigated in the field and laboratory, with particular attention to relationships with adult feeding sites. In the field, distances between adult feeding scars and egg masses differed for C . rubiginosa and H . niponica , with the former being very small and the latter averaging 24.6 cm. The same tendencies for the distances between adult feeding scars and egg masses of the two beetle species were confirmed in cages in which only female beetles were released. Cassida rubiginosa restricted egg laying to host plants in the field and to leaves in laboratory assays. On the other hand, H . niponica placed 8% of egg masses on plants adjacent to host plants in the field and often placed eggs on artificial substrates rather than leaf discs in laboratory assays. These results suggest that oviposition and female feeding sites are virtually inseparable in the case of C . rubiginosa , while H . niponica females do not necessarily keep to host plant leaves as oviposition substrates and they tend to oviposit at some distance from their feeding sites. Results are discussed in relation to proximate and ultimate causes of host selection behavior.     

Variable correspondence of female host preference and larval performance in a phytophagous ladybird beetle Epilachna pustulosa (Coleoptera: Coccinellidae)

Genetic and environmental factors causing intraspecific variation of the thistle Cirsium kamtschaticum Ledeb. as a host plant of the phytophagous ladybird Epilachna pustulosa Kôno were investigated through simple food-choice tests and rearing of larvae. Two thistle clones (T1U₁ and T4H₂) were used, originally growing approximately 12 km apart. A previous study showed that adult female ladybirds preferred T1U₁ to T4H₂, and that larval performance was better on T1U₁, when leaves from the clones in situ were examined. The two clones retained their characteristics with respect to beetle preference after transplantation into a common garden. However, the difference between T1U₁ and T4H₂ with respect to larval performance was reduced after the transplantation. When leaves from shoots of T1U₁ exposed to different sunlight intensities were offered, adult female ladybirds did not show obvious preferences. Larval eclosion rates increased significantly with the increase in leaf sunlight intensity exposure. These results suggest strongly that both genetic and environmental factors are involved in interclonal variation of thistle quality in beetle preference and/or performance. It is suggested that the quality of thistle leaves for larval performance is largely affected by environmental factors, while leaf quality for beetle preference may be determined strictly by genetic factors. Under certain conditions, E. pustulosa females may behave maladaptively, preferring plants not appropriate for larval growth, or not choosing plants appropriate for the larval growth.     

Conspecific thistle plant selection by a herbivorous ladybird beetle, Epilachna pustulosa

Host selection by Epilachna pustulosa Kôno (Coleoptera: Coccinellidae) was surveyed in an area of about 130 ares, focusing on the role of the spatial distribution pattern of the host plant, thistle Cirsium kamtschaticum Ledeb. (Asteraceae) and the environmental conditions of habitats where thistle plants were growing. A total of 198 thistle clones were found in the area studied, and approximately 40% showed some degree of E. pustulosa infestation by July. Eggs were only oviposited on thistle clones that were fed on by adults. Adult beetles and egg masses of E. pustulosa showed an aggregated distribution in the earlier season (June) among thistle clones. The distribution of adults became more random (but still aggregated) by the later season (July), along with an increase in the number of infested clones. Multiple logistic regression analyses revealed that clone size and soil moisture were consistently important for the beetle's choice of clones to feed on. The other logistic regression analyses indicated that thistle‐clone size and sunlight condition influenced egg distribution. Thistle clone selection by E. pustulosa changed with season from a rather strict selection in June to a more obscure one in July, expanding the range of thistle clones used as feeding and oviposition substrate.     

Life history traits of pest and non-pest populations in the phytophagous ladybird beetle,Epilachna niponica (coleoptera, coccinellidae)

Life history traits of the phytophagous ladybird beetle,Epilachna niponica were compared between a non-pest population feeding on wild thistle and a pest population feeding on cultivated solanaceous crops, mainly potato. The pest population had larger females, a higher population growth rate, a more continuous oviposition schedule, and a shorter developmental period in the immature stages, compared with the non-pest population. The two populations showed no clear differences in phenology from spring to autumn, egg mass size, hatchability, and larval survival rate. Significant differences were found in number of eggs laid per female during the first and second periods, and in the developmental period of the immature stages. These life history traits were influenced primarily by food plant. A higher fecundity and shorter immature period appear to be readily altered by the host shift from thistle to potato.     

Differential acceptance of and survivorship on a non‐host plant between two geographically separate populations of the phytophagous ladybird beetle Henosepilachna yasutomii

Phytophagous specialists among insects occasionally accept plants they do not or only rarely utilize under natural conditions. This non‐host acceptance could represent an initial stage in host‐range evolution. Here, we examined adult acceptance of and larval survivorship on a non‐host plant – wild thistle, Cirsium kamtschaticum Ledeb. (Asteraceae) – for two geographically separate populations of the phytophagous ladybird beetle Henosepilachna yasutomii Katakura (Coleoptera: Coccinellidae), which normally utilizes blue cohosh, Caulophyllum robustum Maxim. (Berberidaceae), in the wild. The two beetle populations showed lower acceptance of and survivorship on the non‐host plant (thistle) than on the normal host plant (blue cohosh). Furthermore, they differed significantly in their responses to thistle but not to blue cohosh. Even for the beetle population that more readily accepted thistle as adults, acceptance was moderate and the eclosion rate was low, indicating reduced performance on the non‐host plant. Although studies have infrequently focused on non‐host acceptance, this is likely common in phytophagous specialists and thus is a potentially important evolutionary factor, as it may determine the future direction of food‐range evolution.     

Use of Risk Analysis for Screening Weed Biocontrol Agents: Altica carduorum Guer. (Coleoptera: Chrysomelidae) from China as a Biocontrol Agent of Cirsium arvense (L.) Scop. in North America

Candidate weed biocontrol agents must be screened to exclude those that could threaten desirable plants . Traditionally , this has been done by rejecting species that develop on economically important plants in laboratory no - choice tests . However , because congeneric plants often support development in these tests , even when they are not utilized in nature , the tests do not meet legislated requirements for rare plant species or the increasing public concern for native plants . Plant suitability for larval development is a poor predictor of host range because insects use a sequence of steps in which the early steps , such as host finding and acceptance for oviposition , tend to be stronger than the later ones , such as suitability for development . This study is a trial of a new approach to screening insects as weed biocontrol agents that uses risk analysis to quantify the suitability of a plant as a host on the basis of inset performance at various stages in its life cycle . The insects used for the study was a NW Chinese biotype of the leaf beetle , Altica carduorum, which in terms of climate adaptation and damage it inflicts on the weedy thistle Cirsium arvense, is a promising biocontrol agent for Canada . However , its ability to develop on all North American Cirsium spp . in laboratory no - choice tests currently excludes its release . We show by risk and factor analyses , with five sequential host - selection parameters , that the suitability of these thistles to A. carduorum is so low that the beetle would not jeopardize the continued existence of rare native thistles , so its release should not be a problem .     

Effect of Herbivore Density, Timing of Attack and Plant Community on Performance of Creeping Thistle Cirsium arvense (L.) Scop. (Asteraceae)

Creeping thistle, Cirsium arvense (L.) Scop. (Asteraceae), is one of the most serious weeds in ecological compensation areas (within-field or border refugia) in Europe. Since conventional weed control measures are restricted in compensation areas, augmenting indigenous agents for biological control of the weed may be a feasible alternative. In this paper, we studied the effect of density of the shield beetle, Cassida rubiginosa Muller (Coleoptera, Chrysomelidae), and two vegetation types typical for ecological compensation areas, on the performance of creeping thistle plants in an open field experiment. Early-season larval feeding had no measurable effect on creeping thistle growth, while late-season feeding significantly reduced shoot growth. These findings were attributed to higher feeding rates of the herbivores at higher ambient temperatures late in the season. Defoliation had a strong effect on the above-ground performance of C. arvense plants, but not on the below-ground performance. In contrast, the plant community affected all below-ground performance parameters measured, but only some of the above-ground performance parameters of creeping thistle. A combination of high levels of plant competition and herbivory by C. rubiginosa larvae led to 50% mortality in C. arvense plants during the growing season. We conclude that augmentation of indigenous herbivores of C. arvense in combination with breaking up the root system by tillage and the establishment of a highly competitive plant community of beneficial herbs may be a feasible way to control this problematic weed in ecological compensation areas.     

Spatial prediction of habitat overlap of introduced and native thistles to identify potential areas of nontarget activity of biological control agents

Nontarget feeding of Rhinocyllus conicus Fr?elich and Trichosirocalus horridus (Panzer) on native North American thistles in the genus Cirsium has been documented. Some species of these native thistles have shown greater infestation levels of R. conicus in populations that are in close proximity to the target plant species, Carduus nutans L. In 2005 a study was initiated to identify areas of potential nontarget feeding by R. conicus and T. horridus on thistle species by predicting habitats of two known introduced hosts [C. nutans and Cirsium vulgare (Savi) Tenore] and two native species [Cirsium carolinianum (Walter) Fernald and Schubert and C. discolor (Muhlenberg ex Willdenow) Sprengel] using Mahalanobis distance (D(2)). Cumulative frequency graphs showed that the D(2) models for all four plant species effectively identified site conditions that contribute to the presence of the respective species. Poisson regression showed an association between D(2) values and plant counts at field-test sites for C. nutans and C. carolinianum. However, negative binomial regression detected no association between D(2) values and plant counts for C. discolor or C. vulgare. Chi-square analysis indicated associations between both weevil species and sites where C. vulgare and Carduus nutans were found, but not between the weevil and native thistle species. Habitats of C. nutans and Cirsium carolinianum overlapped in ≈12% of the study area. Data-based habitat models may provide a powerful tool for land managers and scientists to monitor native plant populations for nontarget feeding by introduced biological control agents.     

How to find a needle in a haystack – host plant finding of the weevil Ceratapion onopordi

The weevil Ceratapion onopordi Kirby (Coleoptera: Apionidae) shows a mutualistic interaction with the rust fungus Puccinia punctiformis (Str.) Röhl (Pucciniaceae). One of the weevil’s host plants, the thistle Cirsium arvense (L.) Scop. (Asteraceae), is also the host of the rust fungus. It has been argued that weevils prefer rust‐infected thistle shoots for egg deposition and consequently should be able to detect such shoots. Olfactory, visual, and gustatory orientation was tested using a four‐chamber olfactometer, a visual testing arena, and feeding choice tests. Whereas the weevils used olfactory cues to find their host plants, visual orientation does not seem to be important. Rust‐infected thistle shoots were not preferred over healthy shoots in any of the tests. We conclude that host plants infected with rust fungi, which are rather rare in the field, appear to be found more or less by chance.     

Genetic architecture for normal and novel host-plant use in two local populations of the herbivorous ladybird beetle, Epilachna pustulosa

Trade-offs in host-plant use are thought to promote the evolution of host specificity. However, usually either positive or no genetic correlations have been found. Whereas factors enhancing variation in overall viability have been claimed to mask negative genetic correlations, alternative hypotheses emphasize the sequential changes in genetic correlation in the course of host-range evolution. In this study, the genetic architectures of performances on different hosts were compared in two populations of the herbivorous ladybird beetle, Epilachna pustulosa, using three host plants, one being normal for both, one novel for only one population, and the other novel for both populations. The genetic correlations between larval periods on normal hosts were significantly positive whereas those between normal and novel hosts were not different from zero. There was no evidence for reduced genetic variation on the normal host-plants. These results suggest that the host-range is not restricted by the antagonistic genetic associations among exploitation abilities on different plant species, but rather that selection of different host-plants may improve the coordination between genes responsible for the use of different plants.     

Cryptic local adaptation of the herbivorous ladybird beetle Henosepilachna pustulosa in the ability to use the thistle Cirsium boreale

Local adaptation to different host plants is important in the diversification of phytophagous insects. Thus far, much evidence of the local adaptation of insects with respect to host use at the physiological level has been gathered from systems involving less mobile insects and/or divergent hosts such as plants belonging to different families or genera. On the other hand, the prevalence of such local adaptation of insects with moderate or high dispersal ability to the intraspecific variation of herbaceous hosts is largely unknown. In the present study, we examined the occurrence and degree of local adaptation of the herbivorous ladybird beetle Henosepilachna pustulosa (Kôno) (Coleoptera: Coccinellidae) to its primary host, the thistle Cirsium boreale Kitam. (Asteraceae), through reciprocal laboratory experiments using beetles and thistles from three locations with a range of approximately 200 km. Concerning the larval developmental ability, obvious patterns of local adaptation to the thistles from respective natal locations were detected, at least in some combinations of beetle populations. Similar tendencies were detected concerning adult feeding acceptance, although the statistical support was somewhat obscure. Overall, our results indicate that the degree of local adaptation of insect species with moderate dispersal ability to conspecific herbaceous hosts is occasionally as strong as that involving less mobile insects and/or heterospecific hosts, indicating the potential of such cryptic local adaptation to promote ecological/genetic differentiation of phytophagous insect populations.     

Temporal and spatial relationships between an herbivorous lady beetleEpilachna niponica and its predator,the earwigAnechura harmandi

The relationships between egg predation of an herbivorous lady beetle Epilachna niponica (Lewis ) and its predator, the earwig Anechura harmandi (Burr ), were examined in both time and space. In spite of little annual, changes in egg densities, egg mortality due to predation varied considerably. There was no, clear relationship between the earwig density and the proportionate predation over the five years. The seasonal occurrence of earwig nymphs on thistle plants, however, was closely synchronized with that of egg predation. Predator attacks on the beetle occurred in a time-restricted manner. Thus, later cohorts mostly escape from heavy predatory pressure. No spatially density-depent egg predation was detected at the level of either thistle plants or thistle patches. Furthermore, there was no indication of aggregative behaviour of the earwig in response to local egg density. The earwig density was more likely to be associated with particular localities with sandy deposits available for its nest site.     

The Natural Host Range of Beetle Species Feeding on Broom, Cytisus scoparius (L.) Link (Fabaceae), in Southwest Europe

The natural host range of beetles feeding on broom ( Cytisus scoparius ) and 14 other species (including six other Cytisus species) in the tribe Genisteae was investigated at 39 sites in Spain , Portugal and France in 1989 and 1992 as part of a biological control programme for broom . Data on host - plant associations were analyzed for 36 phytophagous beetle species from 18 sites , and host records were listed for an additional 58 species . Nine species were apparently restricted to the genus Cytisus : Cryptocephalus octoguttatus, Gonioctena olivacea, Bruchidius lividimanus, B. villosus, Exapion elongatissimum, E. fuscirostre, Lepidapion sp . 1 , Polydrusus confluens and Tychius parallelus. These field records suggest a narrower host - plant range for some beetle species than laboratory host - range tests , and may assist in interpreting host - plant associations reported in the literature . Beetle species with a restricted host - plant range were rarely found on related non - host plants .