Multiple invasions in urbanized landscapes: interactions between the invasive garden ant <Emphasis Type="Italic">Lasius neglectus</Emphasis> and Japanese knotweeds (<Emphasis Type="Italic">Fallopia</Emphasis> spp.)

Urbanized landscapes are the theater of multiple simultaneous biological invasions likely to affect spread dynamics when co-occurring introduced species interact with each other. Interactions between widespread invaders call for particular attention because they are likely to be common and because non-additive outcomes of such associations might induce negative consequences (e.g., enhanced population growth increasing impacts or resistance to control). We explored the invasions of two widespread invasive taxa: the Japanese knotweed species complex Fallopia spp. and the invasive garden ant Lasius neglectus, in the urban area of Lyon (France). First, we investigated landscape habitat preferences as well as co-occurrence rates of the two species. We showed that Fallopia spp. and L. neglectus had broadly overlapping environmental preferences (measured by seven landscape variables), but their landscape co-occurrence pattern was random, indicating independent spread and non-obligatory association. Second, as Fallopia spp. produce extra-floral nectar, we estimated the amount of nectar L. neglectus used under field conditions without ant competitors. We estimated that L. neglectus collected 150–321 kg of nectar in the month of April (when nectar production is peaking) in a 1162 m² knotweed patch, an amount likely to boost ant population growth. Finally, at six patches of Fallopia spp. surveyed, herbivory levels were low (1–6% loss of leaf surface area) but no relationship between ant abundance (native and invasive) and loss of leaf surface was found. Co-occurrences of Fallopia spp. and L. neglectus are likely to become more common as both taxa colonize landscapes, which could favor the spread and invasion success of the invasive ant.     

<Emphasis Type="Italic">Lasius neglectus</Emphasis> Van Loon et al. (Hymenoptera,Formicidae), an Invasive Ant Species in Crimea

Data on distribution of Lasius neglectus Van Loon et al. in Crimea are reported. The structure of foraging areas of 111 monocalic and polycalic colonies, the daily activity rhythm, and the visiting of 26 species of trees by L. neglectus workers were studied; over a third of the visited tree species were conifers. Most colonies of L. neglectus in Crimea are monocalic. No replacement of the 12 native ant species present in the territories of the monocalic and polycalic colonies of L. neglectus was observed. Invasion of L. neglectus to Crimea probably started in the early 1970s.     

Mass Occurrence and Dominant Behavior of the European Ant Species <Emphasis Type="Italic">Formica fuscocinerea</Emphasis> (Forel)

Recently, masses of the ant Formica (Serviformica) fuscocinerea (Forel) have been occurring at numerous sites in Southern Germany. Although F. fuscocinerea is native to Southern Germany, these mass occurrences resemble ant invasions in density and dominance. This study aimed to investigate the underlying mechanisms that promote sudden mass occurrence of a previously inconspicuous ant species within its native range. To estimate the competitive dominance of F. fuscocinerea, species occurrence and abundance considering biotic and abiotic parameters were studied in a natural habitat where F. fuscocinerea co-occurred with two other common ant species, Myrmica ruginodis (Nylander) and Lasius niger (Linnaeus). To understand the species’ distribution in the field, laboratory experiments on interspecific competition were conducted. Finally, the colony structure of F. fuscocinerea was investigated with intraspecific aggression tests. Formica fuscocinerea dominated an area that, as indicated by strongly frequented foraging trails on the trees, provided important food sources, e.g. trophobionts, to the ants. Other ant species coexisted only at the periphery of the F. fuscocinerea range. Laboratory experiments revealed F. fuscocinerea as highly dominant species. Additionally, F. fuscocinerea showed a complete lack of intraspecific aggression between ants originating from distances up to 58 km, indicating weak or nonexistent behavioral boundaries among ants of physically separated nests. Since extraordinarily high worker densities, strong interspecific dominance and a lack of colony boundaries within supercolonies are considered to be important traits of several invasive ant species we conclude that the same traits also promote the dominance of F. fuscocinerea.     

Collapse of the invasive garden ant, <Emphasis Type="Italic">Lasius neglectus</Emphasis>, populations in four European countries

The invasive garden ant Lasius neglectus (Hymenoptera: Formicidae) has been spreading rapidly in Europe ever since the 1990s. This ant established enormous supercolonies in many European cities and poses a serious threat to the local native faunas. The spread of this species has not slowed down in the last decades, but in the recent years the sizes of the known L. neglectus populations have generally been declining or have stagnated. For 29 supercolonies checked in four countries, in 10 cases L. neglectus individuals have not been found on the former area of their occurrence. On the other hand, only two supercolonies have expanded. In this paper, we summarize these monitoring data collected by the personal independent, diligent monitoring activities of myrmecologists on populations of the invasive garden ant in Bulgaria, Hungary, Poland and Spain. The reasons for this collapse are thought to be: (1) depletion of the local resources, (2) gradation of pathogens and (social)parasites, (3) climatic factors, (4) intra-population mechanisms, (5) confrontation with highly competitive native species, and (6) lack of suitable nesting microhabitats. As similar phenomena were observed in the cases of supercolonies of other invasive ant species, it seems that they decline more generally than has been thought.     

Stratification of the ant species (Hymenoptera,Formicidae) in the urban broadleaf woodlands of the city of Kiev

Investigations were carried out in June–August 2012 and 2013 in broadleaf forests and planted stands of the northern red oak Quercus rubra in Kiev. The ant visitation rates of 3–6 most abundant plant species within each vegetation layer were analyzed. In all, 16 species of ants were found in the broadleaf forest (Quercus robur + Acer spp. + Carpinus betulus), with the dominance of three species: Formica rufa, Lasius fuliginosus, and L. emarginatus. Eight ant species occurred in Q. rubra stands with the undergrowth of Acer campestre and A. platanoides; the dominant species were Lasius fuliginosus and L. emarginatus. The northern red oak stands with conspecific undergrowth had only four ant species with no dominants among them. Plants with the highest ant visitation rates in the herbage layer of the broadleaf forest were the invasive Impatiens parviflora and the native Aegopodium podagraria. Ants were rare or absent on the remaining herbs: Impatiens noli-tangere, Carex sylvatica, Stellaria holostea, and Galium odoratum. The herbage layer was mostly visited by influents, the dominants being represented only by Lasius emarginatus. Within the shrub layer, ants most frequently visited the undergrowth of the maple Acer platanoides, often harboring colonies of the aphid Periphyllus lyropictus; the highest visitation rates were recorded in the dominants L. emarginatus and L. fuliginosus. The arboreal layer had the highest visitation rate and was most often visited by the dominants Lasius fuliginosus, L. emarginatus, and F. rufa. The common oak Q. robur was the most visited tree in broadleaf forests. In addition to providing ants with food (colonies of the aphids Lachnus roboris and Stomaphis quercus, and also phytophagous insects and other invertebrates), these trees were used for nesting by the dominants L. fuliginosus and L. emarginatus. The presence of aphid colonies was also typical of ripe trees and undergrowth of the maple A. platanoides, the second-visited tree in broadleaf forests. The common hornbeam Carpinus betulus was the least frequently visited, though some ant species, usually L. emarginatus, nested in the trunks and branches of old trees. The vertical distribution of ants in broadleaf forests and red oak stands formed two clusters: the influents occupied the lower (herbage and shrub) vegetation layers, while the subdominants and dominants occupied the upper (shrub and arboreal) ones. The smallest number of ant species (1 or 2) in all the vegetation layers was recorded in the areas with F. rufa; areas with L. fuliginosus had twice as many species; the greatest numbers of ant species were found in the areas with L. emarginatus and in those without dominants. This pattern may result from different territoriality of the dominants: the strongest in F. rufa (defending the whole territory) and the weaker in the other two species (defending only a part of the territory). Plants of all the vegetation layers varied in their attractiveness to ants. As a result, the layers had a mosaic structure in their vertical (between-layer) and horizontal (between different plant species within one layer) arrangement. From 60 to 100% of plants of certain species were visited by ants while plants of other species were not visited at all. The main reason for visiting plants by ants was the presence of aphid colonies.     

Interaction between a threatened endemic plant (<Emphasis Type="Italic">Anchusa crispa</Emphasis>) and the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Seed dispersal mutualisms are essential to ensure the survival of diverse plant species and communities worldwide. Here, we investigated whether the invasive Argentine ant can replace native ants by fulfilling their functional role in the seed dispersal of the rare and threatened endemic myrmecochorous plant, Anchusa crispa, in Corsica (France). Our study addressed the potential of Linepithema humile to disperse elaiosome-bearing seeds of A. crispa, examining L. humile’s effects on (1) the composition of communities of ants removing seeds, (2) the number of seed removals, (3) seed preference, (4) the distance of seed dispersion, and (5) seed germination. We caught seven native species at the control site, but only the Argentine ant at invaded sites. L humile removed A. crispa seeds in greater numbers than did native ants, respectively 66 and 23%, probably due to their higher worker density. The invader was similar to native ants with respect to distance of seed transport. Finally, rates of seed germination were not significantly different between seeds previously in contact with either Argentine ants or not. Taken all together, these results suggest that the Argentine ant is unlikely to pose a threat to A. crispa population. These results have important implications for the management of this rare and threatened endemic plant and provide an example of non-negative interactions between invasive and native species.     

Ants (Hymenoptera: Formicidae) as potential egg predators of lepidopteran pests of sweet potato, <Emphasis Type="Italic">Ipomoea batatas</Emphasis> (Solanales: Convolvulaceae), in Japan

The sweet potato Ipomoea batatas (L.) Lam., which has extrafloral nectaries, was visited by nine indigenous ant species in south Kyushu, Japan. These ant species were evaluated for their ability to remove the eggs of two lepidopteran species—Aedia leucomelas (Linnaeus) and Spodoptera litura (Fabricius)—through ant-exclusion experiments. The results showed that Tetramorium bicarinatum (Nylander), one of the dominant ant species, is an efficient predator of lepidopteran eggs. This ant species removed A. leucomelas eggs from ant-visited plants in six trials conducted from mid-July to mid-October, with high removal rates (90–100%) occurring from mid-September onward. On ant-excluded plants, only 0–18% of exposed eggs were removed by ants that accidentally invaded the plants. Tetramorium bicarinatum also removed egg masses and the hatchlings of S. litura. The mean percent area of egg-laden leaves eaten by S. litura larvae was lower on ant-visited plants (0.9%) than on ant-excluded plants (57.3%). In contrast, Formica japonica Motschulsky, another dominant ant species, was not observed removing the eggs or larvae of the lepidopteran species. These results show that certain indigenous ant species are efficient as biocontrol agents. Thus, individual ant species need to be investigated for their ability to control insect pests in various crops.     

Effects of alien invasion by <Emphasis Type="Italic">Bombus terrestris</Emphasis> L. (Apidae) on the visitation patterns of native bumblebees in coastal plants in northern Japan

When alien pollinator species enter a native community of pollinators in which resource partitioning has been established, the pollination network between plants and pollinators may be modified through the interactions between the pollinators over the use of floral resources. We observed the floral-use patterns of native (Bombus hypocrita and B. deuteronymus) and alien (B. terrestris) bumblebee species in a coastal grassland in northern Japan. We analyzed the factors determining resource partitioning patterns. B. hypocrita tended to visit flowers with shallow or wide open corollas, such as Rosa rugosa, whereas B. deuteronymus visited flowers with complex or deeper corollas, such as Lathyrus japonicus. Given the wider floral preference of B. terrestris, floral use by the alien bumblebees consistently overlapped with that of native bumblebees. The visitation of B. terrestris to R. rugosa flowers was positively correlated with that of B. hypocrita. These bumblebee species frequently used similar floral resources, in part because of the large overlap in the seasonality of their foraging activity. The visitation frequency of B. deuteronymus to L. japonicus flowers was independent of the visitation frequency of other bumblebee species. The major visitation periods of the bumblebees to L. japonicus flowers reciprocally differed between B. deuteronymus and B. terrestris, suggesting phenological resource partitioning between these species. Our study suggests that phenological niche partitioning is more common in specialized flowers (L. japonicus) than in generalized flowers (R. rugosa).     

Reduced ant defenses in <Emphasis Type="Italic">Macaranga</Emphasis> myrmecophytes (Euphorbiaceae) infested with a winged phasmid <Emphasis Type="Italic">Orthomeria cuprinus</Emphasis>

Macaranga is a tree genus that includes many species of myrmecophytes, which are plants that harbor ant colonies within hollow structures known as domatia. The symbiotic ants (plant–ants) protect their host plants against herbivores; this defense mechanism is called ‘ant defense’. A Bornean phasmid species Orthomeria cuprinus feeds on two myrmecophytic Macaranga species, Macaranga beccariana and Macaranga hypoleuca, which are obligately associated with Crematogaster ant species. The phasmids elude the ant defense using specialized behavior. However, the mechanisms used by the phasmid to overcome ant defenses have been insufficiently elucidated. We hypothesized that O. cuprinus only feeds on individual plants with weakened ant defenses. To test the hypothesis, we compared the ant defense intensity in phasmid-infested and non-infested M. beccariana trees. The number of plant–ants on the plant surface, the ratio of plant–ant biomass to tree biomass, and the aggressiveness of plant–ants towards experimentally introduced herbivores were significantly lower on the phasmid-infested trees than on the non-infested trees. The phasmid nymphs experimentally introduced into non-infested trees, compared with those experimentally introduced into phasmid-infested trees, were more active on the plant surface, avoiding the plant–ants. These results support the hypothesis and suggest that ant defenses on non-infested trees effectively prevent the phasmids from remaining on the plants. Thus, we suggest that O. cuprinus feeds only on the individual M. beccariana trees having decreased ant defenses, although the factors that reduce the intensity of the ant defenses remain unclear.     

Patterns of floral resource use by two dominant ant species in a biodiversity hotspot

Ecological dominance in ants is often fuelled by carbohydrate intake. Most studies have focused on the importance of invasive ant mutualistic associations with trophobionts whereas few studies have investigated the importance of floral nectar on invasion success. In this study, utilisation of temporarily available floral nectar by the invasive Argentine ant, Linepithema humile, was compared to that of the dominant native ant, Anoplolepis custodiens, within the Cape Floristic Region (CFR), a biodiversity hotspot. The effect of these two focal ant species on species composition and abundance of ground foraging ants as well as floral arthropod visitors in inflorescences of Proteacea species was assessed. Foraging activity, and trophic ecology inferred from the abundance of natural stable isotopes of Carbon (δ¹³C) and Nitrogen (δ¹⁵N), and the ratio of Carbon to Nitrogen (C:N) were compared between the two ant species during three flowering periods. Linepithema humile significantly reduced the abundance and species diversity of both above-ground and floral arthropod species abundance and composition. Linepithema humile increased its foraging activity with increasing nectar availability, switching its diet to a more herbivorous one. Anoplolepis custodiens did not respond as effectively to increasing floral nectar or negatively impact floral arthropod visitors. This study showed that the availability of floral nectar and ability of L. humile to more effectively utilise this temporarily available resource than native ants, can contribute significantly to the further spread and persistence of L. humile in natural environments in the CFR.     

Temperature and photoperiodic control of diapase induction in the ant <Emphasis Type="Italic">Lepisiota semenovi</Emphasis> (Hymenoptera,formicidae) from Turkmenistan

The ants L. semenovi has been found to belong to species with endogenous-heterodynamic seasonal life cycles with the obligate diapause induced predominantly by factors internal for a colony, whereas external ecological factors (photoperiods and temperature) produce merely modifying effects by accelerating or delaying the diapause onset. The photoperiodic and temperature regulation of diapause induction in larvae and queens is shown. Under effect of short days and low temperature the periods of larval pupation and queen oviposition in a colony are shortened markedly, i.e., the diapause of larvae and queens occurs earlier. The daily rhythms of temperature 15/25°C and particularly 20/30°C as compared with constant temperatures 20 and 25°C that correspond to the mean circadian temperatures of the thermorhythm, inhibit manifestations of the short day effects by stimulating the non-diapause development and increasing duration of the seasonal development cycle of ant colonies. The L. semenovi photoperiodic reaction is quantitative, as development and pupation of larvae and egg-laying of queens cease sooner or later under both the short and the long days, but in the latter case significantly later. Thus L. semenovi is one more example among very rare ant species that are revealed to have the photoperiodic regulation of the colony development seasonal cycle.     

Truffle diversity (<Emphasis Type="Italic">Tuber</Emphasis>, Tuberaceae) in British Columbia

To improve baseline data for the developing truffle industry in British Columbia, we compiled existing Tuber species sequences from published and unpublished studies and generated new ITS sequences for truffles belonging to Tuber collected in the province. In doing so, we obtained evidence that 13 species of Tuber occur in the province, including six introduced and seven native species, two of which are putative undescribed species. Of the native species, the Tuber anniae species complex is widely distributed in the province while Tuber beyerlei appears to be much more restricted in distribution. Four of the introduced species have commercial value (Tuber melanosporum, Tuber aestivum, Tuber brumale, and Tuber borchii) as do two of the native species (Tuber gibbosum and Tuber oregonense). Focused sampling on likely tree hosts, both hardwood and Pinaceae species, as well as in currently unexplored parts of the province seems likely to expand our knowledge of the diversity and distribution of Tuber species in British Columbia.     

New and previously unknown Eurytomidae (Hymenoptera) species from Yemen

Ten new species of the genera Eurytoma (8 species) and Tetramesa (2 species) from Yemen are described (Eurytoma lahji Zerova, E. thoraxica Zerova, E. cyrtophorae Zerova, E. longipes Zerova, E. yemeni Zerova, E. mabari Zerova, E. tibiaspinae Zerova, E. longitarsis Zerova, Tetramesa sanai Zerova, and T. rujumi Zerova). The type specimens of the new species are deposited in the collection of the Institute of Zoology, National Academy of Sciences of Ukraine, Kiev.     

Larvae of the exotic predatory ladybird <Emphasis Type="Italic">Platynaspidius maculosus</Emphasis> (Coleoptera: Coccinellidae) on citrus trees: prey aphid species and behavioral interactions with aphid-attending ants in Japan

An aphidophagous ladybird, Platynaspidius maculosus (Weise) (Coleoptera: Coccinellidae), is originally distributed in China, Taiwan, and Vietnam. The ladybird has recently intruded into the southern and central parts of Japan. The present study found that the larvae of this ladybird preyed on three aphid species, Aphis spiraecola, Aphis gossypii, and Toxoptera citricidus (all Hemiptera: Aphididae), feeding on young shoots of various Citrus species in August to early October in Shizuoka Prefecture, central Japan. Laboratory rearing of the sampled larvae confirmed that the larvae completed their development (adult emergence) by consuming each of the three aphid species. The ladybird larvae were observed foraging in aphid colonies attended by one of the four ants, Lasius japonicus, Pristomyrmex punctatus, Formica japonica, and Camponotus japonicus (all Hymenoptera: Formicidae). Field observations revealed that the foraging/feeding larvae were almost completely ignored by honeydew-collecting ants even when they physically contacted each other. Thus, in Japan, the larvae of the exotic ladybird exploit colonies of the three aphid species attended by one of the four ant species on many Citrus species. On the basis of the results, I discuss the possibility of the ladybird’s reproduction on citrus trees in Japan, probable adaptations of the ladybird larvae to aphid-attending ants, and potential impacts of the ladybird on native insect enemies attacking ant-attended aphids on citrus.     

Ants Associated with <Emphasis Type="Italic">Turnera subulata</Emphasis> (Turneraceae): Elaiosome Attraction,Seed Dispersion and Germination

Symbiosis between plants and ants include examples in which the plant provides shelter and/or food for ants that, in turn, act in the defense or in the dispersion of seeds from the host plant. Although traditionally referred as mutualistic, the results of these interactions may vary with the ecological context in which patterns are involved. A range of species have facultative association with Turnera subulata (Turneraceae). Here, using behavioral bioassays, we investigated the effects of the most frequent ant species associated with T. subulata (Brachymyrmex sp.1, Camponotus blandus (Smith), Dorymyrmex sp.1, Crematogaster obscurata Emery, and Solenopsis invicta Buren) in the dispersion of plant host seeds and in the number of seedlings around the associated ant nests. We also evaluated the effects of these ant species in the germination of T. subulata seeds, in the consumption of elaiosome, and in the attractiveness to elaiosome odor. Our results showed that the ant species associated with T. subulata presented variation in the attraction by the odor and in the rate of consumption of the elaiosomes. However, none of the ant species studied contributed significantly to the increase of seed germination and seedling growth. Our results suggest that the consumption of the elaiosome by ant species is not a determinant factor to the success of germination of T. subulata. However, such species could contribute indirectly to seed germination by carrying seeds to sites more fertile to germination. In general, our results help to elucidate the results of ecological interactions involving ants and plants.     

Weak nestmate discrimination behavior in native and invasive populations of a yellowjacket wasp (<Emphasis Type="Italic">Vespula pensylvanica</Emphasis>)

In geographic regions with warm winters, invasive yellowjacket wasp colonies (genus Vespula) often exhibit polygyny (multiple queens) and persist for multiple years, despite these phenomena being rare in the native range. Here, we test the hypothesis that polygyny, caused by foreign queens being accepted into an existing colony, is the result of relaxed nestmate recognition in the invasive range, as has been observed in some supercolonial invasive ants. In bioassays with wild colonies in the field, we found that nestmate discrimination was weak in both invasive (Hawaii) and native (California) populations of Vespula pensylvanica, with significant nestmate discrimination in only ~?30% of trials. We also found that the diversity and variability of cuticular hydrocarbons, chemical compounds that mediate nestmate recognition, were not reduced in introduced populations, unlike several supercolonial invasive ant species. Our findings suggest that ancestral weak nestmate discrimination behavior of V. pensylvanica may make this species pre-adapted to transition to polygyny and extended colony lifespans when introduced into environments with benign winters that facilitate foreign queens joining existing colonies in late season.     

Heritable bacterial endosymbionts in native and invasive populations of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

Harmonia axyridis Pallas (1773) (Coleoptera: Coccinellidae) is the well-studied system of invasive insect species. Native and invasive parts of the area of H. axyridis are isolated geographically. We studied the species composition and the distribution of bacterial symbionts Spiroplasma and Rickettsia in seven localities of the native area and six localities of the invasive area of H. axyridis. Rickettsia was detected in H. axyridis populations for the first time. We found that the proportion of beetles infected with Rickettsia in native and invasive populations of H. axyridis is about 0.03. Spiroplasma was found only in native populations of H. axyridis. The proportion of infected individuals with Spiroplasma in native populations of H. axyridis is about 0.08. All studied native populations of H. axyridis are infected with Spiroplasma, while all invasive populations are not. We discuss the possible influence of Spiroplasma and Rickettsia in the formation of invasive populations of H. axyridis.     

Effect of host species,host nest density and nest size on the occurrence of the shining guest ant <Emphasis Type="Italic">Formicoxenus nitidulus</Emphasis> (Hymenoptera: Formicidae)

Understanding habitat requirements of species is important in conservation. As an obligate ant nest associate, the survival of the globally vulnerable shining guest ant, Formicoxenus nitidulus, is strictly tied to that of its hosts (mound building Formica ants). We investigated how host species, nest density, inter-nest distance and nest mound size relate to the occurrence of F. nitidulus. In total, 166 red wood ant nests were surveyed in SW Finland (120 Formica polyctena, 25 F. rufa, 14 F. aquilonia, 5 F. pratensis, and 2 F. lugubris). Overall, F. nitidulus was found in 60% of the nests. For the actual analysis, only F. polyctena and F. rufa nests were included due to the small number of other nests. F. nitidulus was more likely to be found among F. polyctena than F. rufa. Also, while inter-nest distance was not important, a high nest density, commonly found in polydomous (multi-nest) wood ant colonies, was beneficial for F. nitidulus. The guest ant was also more likely to be found in large host nests than small nests. Thus, our results show that the best habitat for the guest ant is a dense population of host nest mounds with a high proportion of large mounds. Conservation efforts should be directed at keeping the quality of the red wood ant habitats high to preserve their current populations and to increase colonization. This will not only benefit the guest ant, but also a plethora of other species, and help in maintaining the biodiversity of forests.     

<Emphasis Type="Italic">Lioptilodes friasi</Emphasis> (Lepidoptera: Pterophoridae) Niche Breadth in the Chilean Mediterranean Matorral Biome: Trophic and Altitudinal Dimensions

Understanding the factors driving the diet breadth of phytophagous insects remains one of the main questions in ecological research. In this study we explored the diet breadth and plant-insect associations in the plume moth Lioptilodes friasi Vargas & Parra (Lepidoptera: Pterophoridae). This phytophagous insect was originally described in association with a single host species, Haplopappus foliosus (Asteraceae), a native shrub of the Chilean Mediterranean matorral. In order to address the breadth of host plant choice, we surveyed other Haplopappus species growing along the elevation gradient of central Chile from sea level to 2600 m. We were able to obtain L. friasi adults from five additional Haplopappus species: Haplopappus chrysantemifolius and Haplopappus decurrens from the coastal zone, Haplopappus multifolius and Haplopappus schumanii from the mid-elevation zone, and Haplopappus scrobiculatus at high elevation. Our results demonstrate that the genus-specialized endophagous herbivore L. friasi has a wider distribution and climatic tolerance than previously described. Its biogeographical range extends from the lowland coastal habitats up to the Andean subnival level. We propose that shared flower phenotypic traits such as morphology and chemical composition may have allowed the colonization of closely related Haplopappus species in central Chile, the expansion of which is limited by the harsh high elevation conditions.