Evolution of Cryptocephalus leaf beetles related to C. sericeus (Coleoptera: Chrysomelidae) and the role of hybridization in generating species mtDNA paraphyly

Gómez‐Zurita, J., Sassi, D., Cardoso, A. & Balke, M. (2011). Evolution of Cryptocephalus leaf beetles related to C. sericeus (Coleoptera: Chrysomelidae) and the role of hybridization in generating species mtDNA paraphyly. —Zoologica Scripta, 41, 47–67. Autochthonous European insect diversity is the result of the very complex geological, geographic and climatic history of the Mediterranean area. The leaf beetle genus Cryptocephalus has over 250 species in this area. Among them stands out a group nearly endemic from this region consisting of conspicuous metallic green or blue beetles which can be found visiting yellow Asteraceae flowers in most mid‐ to high‐altitude European grasslands: the Linnaean species C. hypochaeridis, C. sericeus, and all their relatives. In all, these are 32 species forming several taxonomically complex groups across Europe. We sampled all morphological diversity in this lineage and characterized it for two mitochondrial DNA genes. The mtDNA phylogeny of this assemblage was inferred, as well as the timing of its diversification using standard mtDNA substitution rates and a hypothetical Messinian vicariant split. The origin of the group can be traced back to western continental Eurasia in the Lower Miocene. Its subsequent taxonomic splits can be linked to specific periods in the formation of Europe, with a marked trend to east–west phylogenetic divides throughout time and space, and a nearly constant rate of diversification. Only during the Pleistocene, a significant increase in diversification rate can be associated with species formation in the C. hypochaeridis and C. sericeus species complexes. Within these latter groups, most taxa show some degree of mtDNA paraphyly as a result of their recent separation and remarkably by episodes of gene flow in areas of secondary contact among recently diverged species, possibly driven by climatic change.     

Choice,performance and heritability of performance of specialist and generalist insect herbivores towards cacalol and seneciphylline,two allelochemicals of Adenostyles alpina (Asteraceae)

We compared the effects of a sesquiterpene (ST, cacalol) and a pyrrolizidine alkaloid (PA, seneciphylline), both occurring in Adenostyles alliariae, on food choice and performance of specialist and generalist insect herbivores which are all known to feed or live on A. alliariae. In choice experiments we investigated whether the compounds were preferred, deterrent or had no effect. All specialist species Aglaostigma discolor (Hymenoptera, Tenthredinidae), Oreina cacaliae (Coleoptera, Chrysomelidae) and O. speciosissima avoided feeding when confronted with the combination of compounds. Only larvae of A. discolor avoided the single ST treatment as well. Larvae of the generalist species Callimorpha dominula (Lepidoptera, Arctiidae), Cylindrotoma distinctissima (Diptera, Tipulidae) and Miramella alpina (Caelifera, Acrididae) generally avoided feeding from PA, ST and PAST treatments. The only exception were caterpillars of C. dominula which were indiscriminate towards PA when naive, and preferred to feed on the PA treatment when they had experienced the compound before. Performance, measured as the growth of larvae on the different treatments in a no choice situation over a period of 10–17 days, was not different between treatments in the specialist leaf beetles O. cacaliae and O. speciosissima. Their avoidance of the combination treatment in the choice experiments had no obvious effect on growth when forced to feed from the treatment. In the generalist C. dominula only the high concentration combination treatment (PAST) reduced growth of the larvae due to decreased consumption. In C. distinctissima we found reduced growth in all treatments except one (PA3%). Poor growth performance in C. distinctissima was due to postingestive physiological effects of all treatments and additionally to consumption reduction in high‐dose ST treatments. Genetic variability (broad sense heritability) of growth performance metabolism varied in accordance with the specialization degree of the species. O. cacaliae, the most specialized species, had no significant heritability; O. speciosissima, the less specialized specialist, had a heritability of 0.46; C. dominula, the PA adapted generalist species, had a heritability of 0.64; C. distinctissima, the generalist with no apparent adaptations, had a heritability of 0.84.     

Differences in morphology and reproductive traits of Galerucella nymphaeae from four host plant species

The water lily beetle Galerucella nymphaeae L. (Coleoptera: Chrysomelidae) exploits different hosts, including Nuphar lutea Sm. and Nymphaea alba L. (both Nymphaeaceae), as well as Polygonum amphibium L. and Rumex hydrolapathum Hudson (both Polygonaceae). The present study investigates whether within-species differences in morphological and reproductive traits are associated with differences in host species exploitation. A total of 1103 adult beetles were collected from 11 localities in The Netherlands, one of which contained all four hosts and three other localities contained hosts from both families (sympatric localities). Adults originating from Nuphar and Nymphaea were on average darker in colour and larger in size and had disproportionally bigger mandibles than beetles originating from Polygonum and Rumex across the 11 localities. Head capsules of first instar larvae from Nymphaeaceae hosts were between 17% and 28% larger than those of larvae from Polygonaceae hosts. Furthermore, beetles from Nuphar and Nymphaea laid larger sized eggs, but fewer eggs per clutch than beetles originating from Polygonum and Rumex. Although host related variation was less pronounced at the sympatric localities than in the allopatric localities, differences in larval and adult size were still highly significant at the sympatric localities. It is not clear whether the observed differences are genetically based, as opposed to host induced. However, leaf toughness varied among species in a way suggesting that leaf toughness may be partly responsible for host related differences in G. nymphaeae.     

Contrasting patterns of population structure in the montane caddisflies Hydropsyche tenuis and Drusus discolor in the Central European highlands

1. In this study, we compared mitochondrial sequence data (cytochrome oxidase I) to infer the population structure of the two montane caddisflies Hydropsyche tenuis and Drusus discolor. The two species are contrasting examples of montane aquatic insects with insular distributions: D. discolor is restricted to altitudes above 600 m, H. tenuis is limited to the same mountain ranges in Central Europe but inhabits lower altitudes. 2. In particular, we ask whether these two species with similar regional distributions show similar patterns of population structure and haplotype diversity, and whether any differences can be attributed to population history and/or autecology. 3. To determine the population structure of both species, we applied conventional population genetics analyses to mitochondrial sequence data. We collected and sampled 121 specimens of H. tenuis from 29 sites in 10 different regions of the Central European highlands and 138 individuals of D. discolor from 40 sites in 11 different regions. 4. Nine unique haplotypes were identified for H. tenuis and 34 for D. discolor. There were eight variable positions in H. tenuis and 41 in D. discolor. The maximum difference between haplotypes was 0.8% (4 bp) for H. tenuis and 4.2% (21 bp) for D. discolor. We observed haplotype overlap between geographic regions for both species. Analysis of molecular variance showed that two‐thirds of the total variance in H. tenuis was found among regions while in D. discolor, a larger portion of variance was found within regions and populations due to a higher number of haplotypes observed within regions. Mantel test showed a significant relationship between genetic and geographic distance in D. discolor, but no significant relationship in H. tenuis. 5. Our analyses show that, despite their very similar overall distribution pattern in Europe, the two species exhibit distinct population structures, which may reflect differences in phylogeographic history, dispersal capabilities, habitat specifity or within‐region geographic occurrence.     

Genetic relationships between Korean Calanthe species and some naturally occurring mutants based on multiple DNA markers

RAPD, ISSR and AFLP were used to assess the genetic relationships among Korean Calanthe (Orchidaceae) species. Sixteen accessions belonging to five native Calanthe species and some spontaneous mutants were studied. The mutants clustered with C. sieboldii, indicating that they are genetically closer to C. sieboldii than to the rest of the species. Calanthe bicolor clustered with C. discolor, suggesting that its genetic composition is close to that of C. discolor. Though it is suggested to have originated as a result of natural hybridization between C. sieboldii and C. discolor, introgression is likely to have occurred in the direction of C. discolor. Calanthe reflexa and C. aristulifera were the most genetically differentiated of the species studied. We identified 117, 42, 50 putative makers via RAPD, ISSR and AFLP analysis, respectively.     

Biology of the Eucalyptus leaf beetle Paropsisterna selmani (Chrysomelidae: Paropsini): a new pest of Eucalyptus species (Myrtaceae) in Ireland

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Antifungal activity of the dill (Anethum graveolens L.) seed essential oil against strawberry anthracnose under in vitro and in vivo conditions

Strawberry anthracnose, caused by Colletotrichum nymphaeae, is mainly controlled by the application of synthetic chemical fungicides. The present study assessed the antimicrobial activity of essential oils (EOs) from dill (Anethum graveolens L.) seed against C. nymphaeae. The antifungal effects of dill seed EO on C. nymphaeae was initially evaluated in vitro and further extended as in vivo condition. The results indicate that in the contact and volatile assays, dill seed EO significantly inhibited mycelial growth of C. nymphaeae at all concentrations tested. Conidia germination was also significantly inhibited at concentrations of 250 – 1000 ppm. Disease incidence and severity of anthracnose on strawberry fruits were significantly reduced compared with infected control, from concentrations of 500 and 50 ppm, respectively. The results confirm the efficacy of dill seed EO against C. nymphaeae, which may represent an alternative to synthetic chemical fungicides to control strawberry anthracnose pre- and post-harvest.     

Performance of the invasive weevil Polydrusus sericeus is influenced by atmospheric CO2 and host species

• 1 Natural forest systems constitute a major portion of the world's land area, and are subject to the potentially negative effects of both global climate change and invasion by exotic insects. A suite of invasive weevils has become established in the northern hardwood forests of North America. How these insects will respond to increasing CO₂ or O₃ is unknown.
• 2 The present study examined the effects of elevated atmospheric CO₂ and O₃ on the invasive weevil Polydrusus sericeus Schaller at the Aspen Free Air CO₂ Enrichment (FACE) site near Rhinelander, Wisconsin. A performance assay was conducted in the laboratory during the summer of 2007 using mated pairs of P. sericeus fed a combination of aspen, birch and maple foliage. We recorded leaf area consumption, oviposition and adult longevity. We also conducted visual abundance surveys in the field from 2004 to 2007 on aspen and birch at Aspen FACE.
• 3 Elevated CO₂, but not O₃, significantly affected P. sericeus performance. Female, but not male, longevity was reduced under elevated CO₂. Polydrusus sericeus also produced fewer eggs under elevated CO₂ conditions compared with ambient conditions. Adult P. sericeus strongly preferred birch over both aspen and maple, regardless of fumigation treatment.
• 4 The effects of elevated CO₂ on P. sericeus populations at Aspen FACE were minimal, and varied among years and host tree species. Polydrusus sericeus abundance was significantly greater on birch than aspen. Over the long term, elevated CO₂ may reduce adult female longevity and fecundity of P. sericeus. Further studies are needed to evaluate how this information may scale to ecosystem impacts.

     

Galerucella nymphaeae (Col., Chrysomelidae) grazing increases Nuphar leaf production and affects carbon and nitrogen dynamics in ponds

Summary The grazing effects of the waterlily beetle Galerucella nymphaeae on Nuphar lutea stands were studied in three ponds in Central Finland. Production of floating leaves of N. lutea and growth in the G. nymphaeae population were investigated in the ponds and bioenergetics of the beetle larvae in the laboratory. Combination of field and laboratory data enabled estimation of the effect of the beetle on the production of floating leaves of N. lutea and the consequences of grazing for the input of detritus from Nuphar into the ponds. Adults and larvae of G. nymphaeae consumed 3.0–6.1% of the net annual floating leaf production during the growing period. In addition to consumption losses, feeding accelerated the degradation rate of the leaves. This was associated with an increased flow of detrital material of Nuphar origin, and also with increased production of floating leaves in the ponds. These increments were estimated to be up to 3 times greater in the presence of grazing than without it. Grazing by G. nymphaeae releases substantial amounts of carbon and nitrogen bound in Nuphar, particularly in ponds with a dense Nuphar vegetation. It is hypothesized that feeding by this beetle may markedly affect the structure and functioning of such small aquatic systems.     

Feeding behavior of leaf beetles (Coleoptera,Chrysomelidae)

The feeding behavior of adult leaf beetles (41 species from 18 genera and 8 subfamilies) was studied for the first time. Beetles of the genera Chrysolina, Chrysomela, Cryptocephalus, Galeruca, Gastrophysa, Labidostomis, Leptinotarsa, Timarcha, and Cassida stigmatica gnaw a leaf from the edge, whereas the representatives of Donacia, Galerucella, Lema, Lilioceris, Oulema, Phyllobrotica, Plagiodera, Zeugophora, Hypocassida, and most species of Cassida gnaw the leaf plane. In addition, adults of Lilioceris merdigera and Donacia clavipes feed on young leaves rolled into a tube. New host plants are reported for the first time: Hyoscyamus niger for the larvae of the Colorado potato beetle and Naumburgia thyrsiflora for Galerucella grisescens.     

Plasticity of mycangia in Xylosandrus ambrosia beetles

Insects that depend on microbial mutualists evolved a variety of organs to transport the microsymbionts while dispersing. The ontogeny and variability of such organs is rarely studied, and the microsymbiont*s effects on the animal tissue development remain unknown in most cases. Ambrosia beetles (Coleoptera: Curculionidae: Scolytinae or Platypodinae) and their mutualistic fungi are an ideal system to study the animalfungus interactions. While the interspecific diversity of their fungus transport organ一 mycangia—is well-known, their developmental plasticity has been poorly described. To determine the ontogeny of the mycangium and the influence of the symbiotic fungus on the tissue development, we dissected by hand or scanned with micro-CT the mycangia in various developmental stages in five Xylosandrus ambrosia beetle species that possess a large, mesonotal mycangium: Xylosandrus amputatus. Xylosandrus compactus, Xylosandrus crassiusculus, Xylosandrus discolor, and Xylosandrus germanus. We processed 181 beetle samples from the United States and China. All five species displayed three stages of the mycangium development:(1) young teneral adults had an empty, deflated and cryptic mycangium without fungal mass;(2) in fully mature adults during dispersal, the promesonotal membrane was inflated, and most individuals developed a mycangium mostly filled with the symbiont, though size and symmetry varied;and (3) after successful establishment of their new galleries, most females discharged the bulk of the fun gal inoculum and deflated the mycangium. Experimental aposymbiotic individuals demonstrated that the pronotal membrane invaginated independently of the presence of the fungus, but the fungus was required for inflation. Mycangia are more dynamic than previously thought, and their morphological changes correspond to the phases of the symbiosis. Importantly, studies of the fungal symbionts or plant pathogen transmission in ambrosia beetles need to consider which developmental stage to sample. We provide illustrations of the different stages, including microphotography of dissections and micro-CT scans.     

Host-plant effects on larval survival of a salicin-using leaf beetle Chrysomela aeneicollis Schaeffer (Coleoptera: Chrysomelidae)

Several species of willow leaf beetles use hostplant salicin to produce a defensive secretion that consists of salicylaldehyde. Generalist arthropod predators such as ants, ladybird beetles, and spiders are repelled by this secretion. The beetle larvae produce very little secretion when they feed on willows that lack salicylates, and salicin-using beetles prefer salicylate-rich willows over salicylate-poor ones. This preference may exist because the larvae are better defended against natural enemies on salicylate-rich willows. If this is true, the larvae should survive longer on those willows in nature. However, this prediction has not been tested. I determined the larval growth and survival of Chrysomela aeneicollis (Coleoptera: Chrysomelidae) on five willow species (Salix boothi, S. drummondiana, S. geyeriana, S. lutea, and S. orestera). These species differed in their salicylate chemistries and in leaf toughness but not in water content. The water content varied among the individual plants. Larval growth of C. aeneicollis did not differ among the five species in the laboratory, but it varied among the individual plants and it was related to the water content. In the field, C. aeneicollis larvae developed equally rapidly on the salicylate-poor S. lutea and on the salicylate-rich S. orestera. Larval survival was greater on S. orestera than on S. lutea in one year (1986), but there was no difference between them during three succeeding years. In another survivorship experiment, larval survival was low on the medium-salicylate S. geyeriana, but high on the salicylate-poor S. boothi and on S. orestera. Larval survival in the field was related to the larval growth and water content that had been previously measured in the laboratory. These results showed that the predicted relationship between the host plant chemistry and larval survival did not usually exist for C. aeneicollis. One possible reason for this was that the most important natural enemies were specialist predators that were unaffected by the host-derived defensive secretion. One specialist predator, Symmorphus cristatus (Hymenoptera: Eumenidae), probably caused much of the mortality observed in this study. I discuss the importance of other specialist predators to salicin-using leaf beetles.     

A trophic cascade in a macrophyte-based food web at the land–water ecotone

Trophic cascades may purportedly be more common in aquatic than terrestrial food webs, but herbivory on freshwater vascular plants has historically been considered low. Water lilies are an exception, suffering severe grazing damage by leaf beetles. To test whether a central prediction of cascade models—that predator effects propagate downwards to plants—operates in a macrophyte-based food web, we experimentally manipulated predation pressure on a key herbivore of water lilies in the littoral zone of a lake in Michigan, USA. Field experiments comprised combinations of caging treatments to alter the number of predators (larvae of the ladybird beetle Coleomegilla maculata) that hunt the grazers of the macrophytes (larvae of the leaf beetles Galerucella nymphaeae) on the leaves of the water lily Nuphar advena. Predatory larvae of the ladybird beetles significantly reduced grazing damage to water-lily leaves by 35–43%. The predators reduced plant damage chiefly via density-mediated effects, when lower densities of grazers translated to significant declines in plant damage. Plant damage caused by the surviving herbivores was less than predicted from individual grazing rates under predator-free conditions. This suggests that trait-mediated effects may possibly also operate in this cascade. The observed strong effect of predators on a non-adjacent trophic level concurs with an essential component of the trophic cascade model, and the cascade occurred at the ecotone between aquatic and terrestrial habitats: Nuphar is an aquatic macrophyte with emergent and floating leaves, whereas both beetle species are semi-terrestrial and use the dry, emergent and floating leaves of the water lily as habitat. Also, the cascade is underpinned by freshwater macrophytes—a group for which trophic processes have often been underappreciated in the past.     

Nickel localization on tissues of hyperaccumulator species of Phyllanthus L. (euphorbiaceae) from Ultramafic Areas of Cuba

Two species of perennial Phyllanthus (Euphorbiaceae) (Phyllanthus orbicularis and Phyllanthus discolor, both endemic to ultramafic areas of Cuba, and their natural hybrid, Phyllanthus xpallidus) were selected for metal localization microanalysis. Different plant tissues were analyzed by X-ray fluorescence, inductively coupled plasma—atomic emission spectroscopy, and scanning electron microscopy coupled with an energy-dispersive X-ray probe. All of the studied taxa are nickel (Ni) hyperaccumulators and significant concentrations of this element were found in different leaf and stem tissues. The highest Ni content was found in the laticifer tubes, whereas leaf epidermis Ni content resulted to be much more relevant in terms of total metal storage. Calcium and magnesium were found more evenly distributed in leaf and stem tissues.     

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.     

The Origin of Southeastern Asian Triploid Edible <Emphasis Type="Italic">Canna</Emphasis> (<Emphasis Type="Italic">Canna discolor</Emphasis> Lindl.) Revealed by Molecular Cytogenetical Study<Superscript>1</Superscript>

The Origin of Southeastern Asian Triploid Edible Canna ( Canna discolor Lindl.) Revealed by Molecular Cytogenetical Study. Canna discolor Lindl. (Cannaceae), commonly known as edible canna, is often cultivated in Southeastern Asia for its starchy rhizomes. Based on morphological and karyological features, it is thought to be an allotriploid plant originated from hybridization between the closely allied C. coccinea Mill., C. patens Roscoe, C. plurituberosa T. Koyama & Nb. Tanaka, C. speciosa Roscoe, or C. indica L. In this study, to clarify the origin of triploid edible canna, physical mapping of 5S and 18S rDNA probes in C. discolor and its closely related five putative parental species was conducted. Fluorescence in situ hybridization (FISH) technique provided a useful chromosomal marker for discriminating among the diploid putative parental Canna species, and supported the hybrid origin of C. discolor between C. indica var. indica and C. plurituberosa.     

Ecophysiological responses of six coastal dune species along the eastern seaboard of South Africa

Ecophysiological responses of six co‐occurring dune species were investigated in a field study to determine whether they exhibit similar functional traits in response to environmental stressors. The species included Brachylaena discolor DC, Chrysanthemoides monilifera (L.), Scaevola plumieri (L.) Vahl, Canavalia maritima (Aubl.) Thouars, Gazania rigens (L.) Gaertn. and Cyperus esculentis L. Carbon dioxide exchange was saturated at 1,800 μmol m^?2 s^?1 in S. plumieri and at 1,000–1,300 μmol m^?2 s^?1 in the others. Maximal CO₂ exchange occurred during mid‐morning. Midday stomatal closure occurred in S. plumieri, C. monilifera and B. discolor, while stomatal regulation was achieved by maintaining low conductance. Photoinhibition was minimized by efficient mechanisms for light dissipation. Species such as S. plumieri, C. maritima and B. discolor exhibited sclerophylly, a trait for survival in saline and nutrient‐ and water‐deficient environments. Concentrations of Na⁺ and Cl^?1 were high in C. monilifera, G. rigens, B. discolor and S. plumieri, while the high C:N ratio in S. plumieri and C. esculentis suggested nitrogen deficiency. The responses of the six species to environmental stressors in terms of light use, gas exchange, ion and water relations and degree of sclerophylly suggest similar strategies for survival in the dune environment.     

Control efficiency of leaf beetle,Ophraella communa,on the invasive common ragweed,Ambrosia artemisiifolia,at different growing stages

Ophraella communa LeSage (Coleoptera: Chrysomelidae), an effective biological control agent of A. artemisiifolia, was unintentionally introduced into China. To understand the biological control potential of O. communa on A. artemisiifolia, plant height, the number of branches and leaf control index of A. artemisiifolia injured by the beetle were determined. The results showed that plant height and number of branches decreased and leaf control index increased with the increasing initial release density of O. communa adults, when they were released at 90–100-cm plant height stage in 2008 and 60–70-cm stage in 2009. Even when O. communa was released at a low average density of 1.07 adults per plant at early growth stage of A. artemisiifolia in 2009, it could significantly suppress plant height and number of branches and led to a higher leaf control index compared with the herbicide treatment with Roundup. Prior to the fructicative period, the leaf control index of A. artemisiifolia by O. communa was up to 1.0 on the 47th day after the average initial release of 12 beetles per plant in 2008 or on the 85th day after the average release of 1.07 beetles per plant in 2009, which was significantly higher than that in the herbicide treatment. We suggest that the initial release average density of O. communa adults should be ≥1.07 beetles per plant at the early growth stage, or ≥12 beetles per plant at the late growth stage for an effective control of A. artemisiifolia in the field.     

The rove beetle Drusilla sparsa (Coleoptera: Staphylinidae) is a myrmecophilous species associated with a myrmicine ant,Crematogaster osakensis (Hymenoptera: Formicidae)

The rove beetle genus Drusilla includes some myrmecophilous species. The Japanese species Drusilla sparsa (Sharp, 1874) has been regarded as a non‐myrmecophilous beetle. In Kagawa Prefecture, Shikoku Island, western Japan, however, we often observed that D. sparsa adults were walking in the vicinity of foraging workers of the myrmicine ant Crematogaster osakensis Forel, 1990. The body color of the beetle is similar to C. osakensis as in other myrmecophilous beetles found near the trails of the host ants. To examine whether D. sparsa is myrmecophilous, we investigated the distribution of D. sparsa and C. osakensis in the field, as well as their behavior including prey preference of the beetle in the laboratory. Drusilla sparsa beetles were collected only in sites where C. osakensis ants occurred. When the beetles encountered the ant workers, they bent the abdominal tip toward the ants. The ants licked the abdominal tip, and then the beetles usually walked away. Such behavioral reaction of the ants was not observed when the beetles encountered workers of the formicine ant Nylanderia flavipes (Smith, 1874) that continuously attacked the beetles. Drusilla sparsa preferred to feed on dead workers of C. osakensis even when other ants were available as food, indicating that D. sparsa is a myrmecophilous species associated with C. osakensis. Crematogaster osakensis was frequently found in the stomach in the ant predator, the Japanese treefrog Hyla japonica Günther, 1859. Thus, the significance of body color similarity between the host ants and beetles is not a case of Batesian mimicry.     

Mortality pattern and age structure in two carabid populations with different seasonal life cycles

1. Adult populations of two carabid species, a spring breeder, Carabus yaconinus, and an autumn breeder, Leptocarabus kumagaii, were studied in a lowland habitat (area: 16,500 m²) by pitfall sampling and mark-recapture method. The seasonal change in population number, age structure and mortality pattern were clarified and related to their seasonal life cycles.
2. The survival rate of new adults from emergence to the first reproductive season was on the average 48% in C. yaconinus and 55% in L. kumagaii. In C. yaconinus, beetles which emerged later in the season survived more to the first reproductive season than those emerged earlier. C. yaconinus which had survived the pre-reproductive period mostly died out during the first reproductive season of 5 months, and about 8% survived until the second season. In L. kumagaii which had a short reproductive period in autumn, about 20% survived to the second reproductive season, and a small proportion even to the third reproductive season. Accordingly, the proportion of old beetles in the reproductive population was higher in L. kumagaii than in C. yaconinus.
3. The reproductive population of C. yaconinus contained on the average 1600 beetles and produced 3300 new adults. the L. kumagaii population contained on the average about 530 reproductives in autumn, and about 820 beetles emerged in the following year. The recruitment rate of new adults of C. yaconinus was higher than that of L. kumagaii, and this resulted in its higher population density. In L. kumagaii, however, the high adult survivorship and iteroparous reproduction were important for its population growth.
4. Relationship between seasonal adaptation and demographic strategies in the carabid populations were discussed.