Beetle assemblages of indigenous and alien decomposing fruit in subtropical Durban,South Africa

Ecological theory predicts that insect assemblages on indigenous plants will be more diverse than those on alien plants. However, this theory refers primarily to herbivores, and its applicability to decomposers is unclear. Here, we compare beetle assemblages from the fallen fruit of the two most common plants producing large fleshy fruit at our study site in Durban, South Africa: one indigenous African plant (the toad tree, Tabernaemontana ventricosa, Apocynaceae) and one invasive tropical American one (the granadilla, Passiflora edulis, Passifloraceae), at various stages of the decomposition process, thus spanning a continuum between herbivory and detritivory. Beetles found on the two plants included both alien (some categorized as pests of stored fruit) and indigenous species (including some localized endemics). We found that the mean diversity and abundance were not significantly different between the two plant species (with nine beetles from four morphospecies in the average sample) and that the beetle assemblages from the two plant species largely overlap. Statistical analyses suggested that other factors such as stage of fruit decomposition and seasonality could have greater influence on diversity and abundance than the provenance of the plant species. We conclude that insect–plant interactions in the emerging ecosystems that include indigenous and alien species in both groups are complex and that the importance of interactions between species of different provenances may have previously been underestimated.     

Reverse influence of riparian buffer width on herbivorous and predatory Hemiptera

Insects are major contributors to farmland biodiversity, and their economic roles are also diverse. Many herbivorous species are crop pests, while predatory insects have the potential to act as biological controls against pests. Overall insect diversity has declined as a result of intensified agricultural practices. Riparian buffers may support insect populations in intensively cultivated areas, but their actual impact on the balance between harmful pests and beneficial predators is not known. It can be postulated that this impact may vary depending on the characteristics and location of the riparian buffer itself. We investigated the possibility that the biotic and abiotic attributes of agricultural riparian buffers adjoining crop fields and watercourses can explain the species composition of hemipteran assemblages. In particular, we were interested in the abundances of species belonging to the genus Nabis (generalist predators) and recognized and potential pests of cereal crops. Riparian buffer width and the presence or absence of woody plants were not associated with hemipteran species turnover among riparian buffers. In contrast, differences in the degree of dominance by grasses, in plant species turnover, and in which crop plant was cultivated in the adjacent field, explained a significant proportion of the variance in hemipteran species turnover. The abundance of predatory Nabis species increased with increasing riparian buffer width, whereas the abundance of recognized and potential crop pests decreased. The reverse patterns in the predatory and herbivorous Heteroptera suggest that increasing riparian buffer width might enhance biological control by Nabis predators.     

外来植食性广聚萤叶甲对非靶标植物的潜在影响

为评估以入侵豚草为食的外来广聚萤叶甲（Ophraella communa）对非靶标植物的潜在生态风险,采用选择性试验测定了广聚萤叶甲成虫和幼虫对当地代表性植物的取食选择,观察了幼虫在选出的植物上生长发育的适合度表现。结果表明：在5大类52种测试植物中,广聚萤叶甲成虫和幼虫不同程度地取食向日葵、苍耳、天明精、菊芋（仅成虫）、紫茎泽兰（仅幼虫）和石胡荽（仅幼虫）;野外罩笼非选择性测定发现,初孵幼虫在苍耳和向日葵8个品种上可完成发育,直到成虫产卵,并观察到在菊芋上完成幼虫发育,个别到成虫但未产卵;在这些植物上幼虫存活率明显降低,蛹质量明显减轻,成虫产卵明显减少。本文还分析了广聚萤叶甲对少数本土植物以及经济作物向日葵的潜在生态风险。     

Taxonomic homogenization and differentiation across Southern Ocean Islands differ among insects and vascular plants

Aim To investigate taxonomic homogenization and/or differentiation of insect and vascular plant assemblages across the Southern Ocean Islands (SOI), and how they differ with changing spatial extent and taxonomic resolution. Location Twenty‐two islands located across the Southern Ocean, further subdivided into five island biogeographical provinces. These islands are used because comprehensive data on both indigenous and non‐indigenous insect and plant species are available. Methods An existing database was updated, using newly published species records, identifying the indigenous and non‐indigenous insect and vascular plant species recorded for each island. Homogenization and differentiation were measured using Jaccard’s index (JI) of similarity for assemblages across all islands on a pairwise basis, and for island pairs within each of the biogeographical provinces. The effects of taxonomic resolution (species, genus, family) and distance on levels of homogenization or differentiation were examined. To explore further the patterns of similarity among islands for each of the taxa and groupings (indigenous and non‐indigenous), islands were clustered based on JI similarity matrices and using group averaging. Results Across the SOI, insect assemblages have become homogenized (0.7% increase in similarity at species level) while plant assemblages have become differentiated at genus and species levels. Homogenization was recorded only when pairwise distances among islands exceeded 3000 km for insect assemblages, but distances had to exceed 10,000 km for plant assemblages. Widely distributed non‐indigenous plant species tend to have wider distributions across the SOI than do their insect counterparts, and this is also true of the indigenous species. Main conclusions Insect assemblages across the SOI have become homogenized as a consequence of the establishment of non‐indigenous species, while plant assemblages have become more differentiated. The likely reason is that indigenous plant assemblages are more similar across the SOI than are insect assemblages, which show greater regionalization. Thus, although a suite of widespread, typically European, weedy, non‐indigenous plant species has established on many islands, the outcome has largely been differentiation. Because further introductions of insects and vascular plants are probable as climates warm across the region, the patterns documented here are likely to change through time.     

Diversity and abundance patterns of phytophagous insect communities on alien and native host plants in the Brassicaceae

The herbivore load (abundance and species richness of herbivores) on alien plants is supposed to be one of the keys to understand the invasiveness of species. We investigate the phytophagous insect communities on cabbage plants (Brassicaceae) in Europe. We compare the communities of endophagous and ectophagous insects as well as of Coleoptera and Lepidoptera on native and alien cabbage plant species. Contrary to many other reports, we found no differences in the herbivore load between native and alien hosts. The majority of insect species attacked alien as well as native hosts. Across insect species, there was no difference in the patterns of host range on native and on alien hosts. Likewise the similarity of insect communities across pairs of host species was not different between natives and aliens. We conclude that the general similarity in the community patterns between native and alien cabbage plant species are due to the chemical characteristics of this plant family. All cabbage plants share glucosinolates. This may facilitate host switches from natives to aliens. Hence the presence of native congeners may influence invasiveness of alien plants.     

Do alien plants escape from natural enemies of congeneric residents? Yes but not from all

As predicted by the enemy release hypothesis, plants are supposedly less attacked by herbivores in their introduced range than in their native range. However, the nature of the natural enemies, in particular their degree of specificity may also affect the level of enemy escape. It is therefore expected that ectophagous invertebrate species, being generally considered as more generalists than endophagous species, are more prompt to colonise alien plants. In Swiss, Siberian and Russian Far East arboreta, we tested whether alien woody plants are less attacked by native herbivorous insects than native congeneric woody plant species. We also tested the hypothesis that leaf miners and gall makers show stronger preference for native woody plants than external leaf chewers. In all investigated regions, leaf miners and gall makers were more abundant and showed higher species richness on native woody plants than on congeneric alien plants. In contrast, external leaf chewers did not cause more damage to native plants than to alien plants, possibly because leaf chewers are, in general, less species specific than leaf miners and gall makers. These results, obtained over a very large number of plant-enemy systems, generally support the hypothesis that alien plants partly escape from phytophagous invertebrates but also show that different feeding guilds may react differently to the introduction of alien plants.     

Insect diversity and trophic structure differ on native and non‐indigenous congeneric rushes in coastal salt marshes

Displacement of native plant species by non‐indigenous congeners may affect associated faunal assemblages. In endangered salt marshes of south‐east Australia, the non‐indigenous rush Juncus acutus is currently displacing the native rush Juncus kraussii, which is a dominant habitat‐forming species along the upper border of coastal salt marshes. We sampled insect assemblages on multiple plants of these congeneric rushes in coastal salt marshes in Sydney, New South Wales, Australia, and compared the abundance, richness, diversity, composition and trophic structure between: (i) J. acutus and J. kraussii at invaded locations; and (ii) J. kraussii at locations either invaded or not invaded by J. acutus. Although J. acutus supported a diverse suite of insects, species richness and diversity were significantly greater on the native J. kraussii. Moreover, insect assemblages associated with J. kraussii at sites invaded by J. acutus were significantly different from, and more variable than, those on J. kraussii at non‐invaded sites. The trophic structure of the insect assemblages was also different, including the abundance and richness of predators and herbivores, suggesting that J. acutus may be altering consumer interactions, and may be spreading in part because of a reduction in herbivory. This strongly suggests that J. acutus is not playing a functionally similar role to J. kraussii with respect to the plant‐associated insect species assemblages. Consequently, at sites where this non‐indigenous species successfully displaces the native congener, this may have important ecological consequences for community composition and functioning of these endangered coastal salt marshes.     

Relationship of different feeding groups of true bugs (Hemiptera: Heteroptera) with habitat and landscape features in Pannonic salt grasslands

Eastern European grasslands are still inhabited by a rich arthropod fauna, but the drivers and mechanisms influencing their communities have to be understood to ensure their future survival. Heteroptera communities were studied in 20 plot-pairs in Pannonic salt steppe–salt marsh mosaics in Hungary. The effects of vegetation characteristics, landscape diversity and the proportion of surrounding grasslands on the composition, species richness and abundance of different feeding groups of true bugs (carnivores, specialist and generalist herbivores) were examined using ordinations and mixed-effect models. We found distinct herbivorous assemblages corresponding to microtopography-driven differences in water regime and vegetation between steppe and marsh plots, but this pattern was less pronounced in carnivorous assemblages. A higher species richness of true bugs was found in the more diverse steppe vegetation than in the salt marsh vegetation, while the abundance pattern of true bugs was opposite. Landscape diversity had a positive effect on the species richness and abundance of generalist herbivores and carnivores. Our results suggested that generalist herbivores and carnivores appear to drive diversity patterns in the local landscape due to their high dispersal abilities and the broader range of resources they can utilize. Specialist herbivores strongly influence the local insect biomass in relation to the distribution and density of their host plants. The present study highlights the importance of both habitat and landscape diversity for local insect diversity in Pannonic salt grasslands and suggests that the main threats for arthropod diversity are those processes and activities that homogenize these areas.     

山东省外来植物的区系特征及空间分布

外来植物及其生态入侵对当地生物多样性保护和生态系统产生的不利影响,是目前生态学研究的热点之一。本文在文献查阅和野外调查基础上,建立了山东省外来植物数据库,并对其组成、地理区系、空间分布等进行统计分析,结果表明:1)山东省现有外来植物共827种,隶属122科,416属,其中境外外来种348种;2)外来植物比例高,占山东植物区系的39.03%,且优势科和表征科明显;3)外来植物属的地理分布多样,温带成分占优势(52.51%),热带成分丰富(44.06%);4)外来植物来源地和分布地相对集中,反映了环境因子和人类活动在入侵因素中的共同影响;5)温带气候带来源的植物较易入侵山东。     

A preliminary assessment of changes in plant-dwelling insects when threatened plants are translocated

Translocation of threatened species is a tool used increasingly to conserve biodiversity, but the suite of co-dependent species that use the threatened taxa as hosts can be overlooked. We investigate the preliminary impact of translocating three threatened plant species on insect species and the integrity of insect assemblages that depend on these plants as their hosts. We compare the insect assemblages between natural populations of the threatened species, related non-threatened plant species growing wild near the threatened plants, and threatened plants translocated to another site approximately 40?km away. We used host breadth models and a coextinction risk protocol to determine which insect species are potentially host-specific on the threatened plants, and then assessed these insects?? potential presence at the translocation site. We found that insect assemblages on naturally-occurring threatened plants had more individuals, higher species density and higher species richness than assemblages on translocated plants. For one plant species, Leucopogon gnaphalioides, species composition differed significantly between wild and translocated populations (P?<?0.001). Furthermore, four insect species that were host-specific to Banksia brownii and B. montana were not detected on the translocated plants. Instead, translocated plants supported insect assemblages more similar to those of related plant species from the surrounding area. We conclude that threatened plant translocations that involve seed collection and propagation may have limited benefit for individual dependent species or the supported insect assemblage. Additional conservation actions will be required to maintain the diversity of insect assemblages and host-dependent relationships.     

Integration of alien plants into a native flower-pollinator visitation web

Introduced alien species influence many ecosystem services, including pollination of plants by animals. We extend the scope of recent 'single species' studies by analysing how alien plant species integrate themselves into a native flower visitation web. Historical records for a community in central USA show that 456 plant species received visits from 1429 insect and 1 hummingbird species, yielding 15 265 unique interactions. Aliens comprised 12.3% of all plant species, whereas only a few insects were alien. On average, the flowers of alien plants were visited by significantly fewer animal species than those of native plants. Most of these visitors were generalists, visiting many other plant species. The web of interactions between flowers and visitors was less richly connected for alien plants than for natives; nonetheless, aliens were well integrated into the native web. Because most visitors appear to be pollinators, this integration implies possible competitive and facilitative interactions between native and alien plants, mediated through animal visitors to flowers.     

Human impacts, energy availability and invasion across Southern Ocean Islands

Aim Ongoing biological invasions will enhance the impacts of humans on biodiversity. Nonetheless, the effects of exotic species on diversity are idiosyncratic. Increases in diversity might be a consequence of similar responses by species to available energy, or because of positive relationships between human density, energy and propagule pressure. Here we use data from the Southern Ocean island plants and insects to investigate these issues. Location The Southern Ocean Islands ranging from Tristan da Cunha to Heard Island and South Georgia. Methods Generalized linear models are used to explore the relationships between indigenous and exotic species richness for plants and insects on two different islands. Similar models are used to examine interactions between indigenous and exotic species richness, energy availability and propagule pressure at the regional scale. Results Positive relationships were found between indigenous and exotic species richness at local scales, although for plants, the relationship was partially triangular. Across the Southern Ocean Islands, there was strong positive covariation between indigenous and exotic plant species richness and insect species richness, even taking spatial autocorrelation into account. Both exotic and indigenous plant and insect species richness covaried with energy availability, as did human visitor frequency. When two islands with almost identical numbers of human visits were contrasted, it was clear that energy availability, or perhaps differences in climate‐matching, were responsible for differences in the extent of invasion. Conclusion In plants and insects, there are positive relationships between indigenous and exotic diversity at local and regional scales across the Southern Ocean islands. These relationships are apparently a consequence of similar responses by both groups and by human occupants to available energy. When visitor frequency is held constant, energy availability is the major correlate of exotic species richness, though the exact mechanistic cause of this relationship requires clarification.     

Genetic differentiation associated with host plants and geography among six widespread species of South American Blepharoneura fruit flies (Tephritidae)

Tropical herbivorous insects are astonishingly diverse, and many are highly host‐specific. Much evidence suggests that herbivorous insect diversity is a function of host plant diversity; yet, the diversity of some lineages exceeds the diversity of plants. Although most species of herbivorous fruit flies in the Neotropical genus Blepharoneura are strongly host‐specific (they deposit their eggs in a single host plant species and flower sex), some species are collected from multiple hosts or flowers and these may represent examples of lineages that are diversifying via changes in host use. Here, we investigate patterns of diversification within six geographically widespread Blepharoneura species that have been collected and reared from at least two host plant species or host plant parts. We use microsatellites to (1) test for evidence of local genetic differentiation associated with different sympatric hosts (different plant species or flower sexes) and (2) examine geographic patterns of genetic differentiation across multiple South American collection sites. In four of the six fly species, we find evidence of local genetic differences between flies collected from different hosts. All six species show evidence of geographic structure, with consistent differences between flies collected in the Guiana Shield and flies collected in Amazonia. Continent‐wide analyses reveal – in all but one instance – that genetically differentiated flies collected in sympatry from different host species or different sex flowers are not one another's closest relatives, indicating that genetic differences often arise in allopatry before, or at least coincident with, the evolution of novel host use.     

Recovery of indigenous butterfly community following control of invasive alien plants in a tropical island’s wet forests

Invasive alien species pose one of the highest threats to biodiversity, especially in isolated oceanic islands where high rates of both endemism and extinction risk also usually prevail. Few studies have investigated the impact of invasive alien plants on butterflies in insular ecosystems, despite butterflies representing a key indicator group for terrestrial arthropod diversity. Using the Pollard Technique, we quantified butterfly species richness and abundance in eight wet lowland forest areas invaded by alien plants, principally the strawberry guava (Psidium cattleianum Sabine) on the tropical volcanic island of Mauritius, and compared the results with paired adjacent forest plots that had been weeded of alien plants between 2 and 12 years previously. Butterfly assemblages in weed-infested and weeded forests were distinctly different with higher species richness and much higher butterfly abundance in the latter. At least some of these differences seemed attributable to weed removal effects on forest structure, but understanding the precise mechanisms involved will require further study. The results suggest that alien plant invasion may have contributed to the extinction of certain endemic taxa and can increase the likelihood of butterfly species extinction by reducing population sizes through reduced habitat quality. Such a shift in a forest’s butterfly assemblage is likely to have negative effects on both their indigenous predators and the plants they pollinate. It is argued that in order to maintain butterfly and other arthropod diversity and function in these forests, alien plant control must be maintained and extended beyond the current 1% of surviving forest remnants.     

Comparison of invertebrate herbivores on native and non‐native Senecio species: Implications for the enemy release hypothesis

The enemy release hypothesis posits that non‐native plant species may gain a competitive advantage over their native counterparts because they are liberated from co‐evolved natural enemies from their native area. The phylogenetic relationship between a non‐native plant and the native community may be important for understanding the success of some non‐native plants, because host switching by insect herbivores is more likely to occur between closely related species. We tested the enemy release hypothesis by comparing leaf damage and herbivorous insect assemblages on the invasive species Senecio madagascariensis Poir. to that on nine congeneric species, of which five are native to the study area, and four are non‐native but considered non‐invasive. Non‐native species had less leaf damage than natives overall, but we found no significant differences in the abundance, richness and Shannon diversity of herbivores between native and non‐native Senecio L. species. The herbivore assemblage and percentage abundance of herbivore guilds differed among all Senecio species, but patterns were not related to whether the species was native or not. Species‐level differences indicate that S. madagascariensis may have a greater proportion of generalist insect damage (represented by phytophagous leaf chewers) than the other Senecio species. Within a plant genus, escape from natural enemies may not be a sufficient explanation for why some non‐native species become more invasive than others.     

Effects of an invasive alien tree on the diversity and temporal dynamics of an insect assemblage on an oceanic island

Native vegetation is frequently replaced by alien plants on isolated oceanic islands. The effects of such replacements by invasive plants on the diversity and temporal dynamics of island-endemic insects remain unclear. We examined flying insect communities using Malaise traps on the small island of Nishi-jima in the oceanic Ogasawara Archipelago in the northwestern Pacific. On the island, an alien tree, Casuarina equisetifolia, has become dominant, occupying 57.3?% of the vegetation area. The species richness, composition, and abundance of pollinators (bees), predators (wasps), and wood-boring beetles (cerambycids, mordellids, and elaterids) were compared in each summer season of 4?years among three vegetation types: C. equisetifolia forest, natural forest, and grassland. In the traps, 82.3?% of species captured were endemic to the archipelago. The grassland harbored the highest species richness of native bees and wasps, whereas the natural forest had the highest species richness of native wood-boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups. Principal response curves indicated that differences in species composition among the three vegetation types were consistent through time for all insect groups. Most insect species were more abundant in natural forest or grassland than in C. equisetifolia forest. Standard deviations in both the numbers of individuals and species estimated under a Bayesian framework suggested that annual fluctuations of abundance and species density were similar among vegetation types (except for elaterid abundance). Therefore, replacement by C. equisetifolia has likely altered insect species composition but has not necessarily dramatically affected the temporal dynamics of insect assemblages on the island.     

Comparative footprint of alien, agricultural and restored vegetation on surface-active arthropods

Both invasive alien trees and agricultural conversion have major impacts on biodiversity. We studied here the comparative impact of these two types of land transformation on a wide range of surface-active arthropod species using pitfall traps, with evergreen sclerophyllous natural vegetation (fynbos) as the control. The study was in the Cape Floristic Region, a global biodiversity hotspot, where alien trees are of major concern and where vineyards replace natural fynbos vegetation. Surface-active arthropods were selected as they are species rich, relatively immobile, and occur in high abundance. We hypothesized that the impact of the two types of land cover transformation would produce similar qualitative and quantitative effects on the arthropods. We also compared the results in the transformed and natural areas with those in areas cleared of alien trees. Arthropod species richness in cleared areas was higher than in vineyards and more similar to that in natural fynbos, while alien trees had the lowest. Overall abundance scores were highest in cleared areas, closely followed by fynbos, then vineyards and lowest in alien trees. Several species were restricted to each vegetation type, including alien trees. In terms of assemblage composition, all vegetation types were significantly different, although fynbos and vineyards grouped, suggesting that vineyards have less impact on the arthropod community than do alien trees. When rare species were excluded, vineyards and cleared sites grouped, indicating some recovery but only involving those species that were common and habitat tolerant. Our results suggest that vineyards retain a greater complement of indigenous species than alien trees, but that clearing of these aliens soon encourages establishment of indigenous species. Although there were significant differences in soil moisture and litter depth within and between vegetation types, we did not record them as significantly affecting species richness or abundance, even in alien vegetation, an encouraging sign for restoration.     

Effect of alien riparian vegetation and its removal on a highly endemic river macroinvertebrate community

Invasive alien trees along river banks can reduce indigenous biodiversity, while their removal can restore it. We assessed here family- and species-level responses of river benthic macroinvertebrate assemblages to three riparian vegetation types (natural, alien trees, cleared of alien trees) in the Cape Floristic Region biodiversity hotspot. High species beta diversity of this highly endemic fauna meant that between-river, as well as seasonal effects, dominated assemblage patterns. SASS5, a qualitative, rapid bioassessment technique, based on the sensitivity of the families present, was used as a measure of river health and, indirectly, of water quality. SASS indicated a decline in water quality conditions after alien clearing, a likely response to the greater insolation and apparent erosion of cleared banks, resulting in elevated temperatures and suspended solids and lowered oxygen levels. Overall, cleared and natural sites were more similar to each other than to alien sites, suggesting some post-clearing recovery. However, many sensitive, endemic taxa survived in alien-invaded sites, and more than in the natural sites. These endemic species made use of shady, cool, high-oxygen levels under the alien tree canopy. However, endemics declined in overall abundance in sites cleared of aliens, being replaced by more tolerant, widespread taxa. Clearance of the alien trees opened up the rivers to sunny conditions, which had a major impact on community composition. Vegetation types, oxygen levels and river width were important environmental variables affecting these macroinvertebrate responses. Re-establishment of invertebrate biodiversity matched that of indigenous vegetation, with the most sensitive endemic taxa only recovering after establishment of bushy indigenous and shade-producing fynbos. Therefore, for biodiversity conservation objectives to be achieved, it is essential that indigenous plants are maintained and encouraged during and after clearing to ensure the recovery of endemic and sensitive taxa.     

Performance on different host plants of an alien and an indigenous Bemisia tabaci from China

The capacity of the Middle East‐Asia Minor 1 putative species of the whitefly Bemisia tabaci (Gennadius) species complex, commonly referred to as the ‘B biotype’, to invade has often been linked to its presumed wider host range than the indigenous competitors. To determine whether this alien putative species and the indigenous Asia II 1 whitefly putative species, commonly referred to as the ‘ZHJ2 biotype’, differ in their ability to use different host plants, we compared their development, survival and reproduction on eight crop species/cultivars that are commonly cultivated in Zhejiang, China. Of the eight host plants tested, B performed substantially better than ZHJ2 on squash, tomato and tobacco, B and ZHJ2 preformed equally well on cotton and sweet potato, while ZHJ2 performed better than B on kidney bean and pepper. These results indicate that while B generally has a wider host range than many indigenous B. tabaci, an indigenous B. tabaci can perform as well as or better on some host plants. These results combined with the cropping patterns in Zhejiang suggested that the differential capacity to use various host plants between whitefly species is important in mediating the process of invasion by an alien whitefly species.     

Habitat complementarity and butterfly traits are essential considerations when mitigating the effects of exotic plantation forestry

Habitat loss threatens insect diversity globally. However, complementary vegetation types in remaining habitat increases opportunities for species survival. We assess the extent to which indigenous forest patches moderate the impact of exotic commercial afforestation on grassland butterflies. Butterflies were sampled in grassland along uncorrelated gradients of landscape-scale indigenous forest and plantation cover, while controlling for variation in local vegetation composition. We separately assessed responses by butterfly groups differing in habitat preference, larval diet, and mobility. There was no effect of landscape- or local-scale variables on species richness, but there was a strong interactive effect of forest and plantation cover on butterfly assemblage structure. The effect varied according to species traits. When forest cover was high, assemblages did not differ at different levels of plantation cover. However, plantation cover significantly influenced assemblage structure when forest cover was low. Grassland with limited forest cover in the protected area supported unique assemblages with high frequency of less mobile, specialized species with herbaceous larval host plants, whereas grassland with low forest cover near plantations had a prevalence of mobile, generalist species. A positive association between forest cover and butterflies with woody larval host plants suggests that indigenous forest patches improved the suitability of fragmented grassland for a subset of butterflies, emphasising the value of natural heterogeneity in transformed areas. However, certain butterfly traits associated with large, open grassland were under-represented in grassland between plantations, underscoring the importance of open areas in the broader landscape to conserve the full diversity of species.