Patterns of species assemblages and geographical distributions associated with mandible size differences in coastal tiger beetles in Japan

Nine species of tiger beetle (Cicindelidae) occur in coastal habitats in Japan, with two to four species co-occurring at each locality. To examine the patterns of coexistence and geographical distribution, the mandible size of co-occurring species at 17 localities in Japan was examined, based on the assumption that competition for food is an important factor in determining these patterns. The interspecific overlap of mandible length was absent or very low in localities with two or three species, whereas it was more or less evident in localities with four species. For four large coastal species, the geographical distributions of two species with similar mandible lengths are either allopatric or parapatric, whereas those of two species with different mandible lengths largely overlap. These results strongly suggest that size-assortment in mandible length is important in determining species assemblage and distribution in coastal tiger beetles in Japan.An erratum to this article can be found at     

Tourism impairs tiger beetle (Cicindeldae) populations – a case studyin a Mediterranean beach habitat

Adult activity and larval development of Lophyridia concolor (Dejean) was examined in three sections of a marine beach: (1) an undisturbed section; (2) a section occasionally used by tourists; and (3) a heavily disturbed beach section. Although the activity of adult tiger beetles was similar at all three sites in spring, it diverged during the tourist season. The adults reached a high activity with two maxima on the natural beach, but it was 40% lower in the occasionally used section. The adult activity decreased abruptly without visible maxima in the tourist area. Larval activities showed similar trends. First and second instar larvae were practically absent from the heavily disturbed section. The reduced tiger beetle activity was most likely caused by disturbance and modification of habitat by the tourists. Mechanisms of this impairment are discussed. The results might be of importance for several beach inhabiting invertebrates in the Mediterranean area. Persistently high levels of tourist traffic on beaches might lead to local extirpation, as the decline of Lophyridia a. aphrodisia (Baudi), another Turkish tiger beetle species, shows.     

Geographical variation in mandible morphologies specialised for collembolan predation depend on prey size in the ant Strumigenys lewisi

1. Ants in the genus Strumigenys are predator ants that feed on tiny soil arthropods. The mandibles are modified into high‐speed traps to capture swift collembolan prey. The peculiar mandible morphologies of these ants have evolved depending on characteristics of the prey. Specifically, the evolution of mandible size and shape may be directly driven by prey size. 2. In the present study, the intraspecific variation of the morphological traits of Strumigenys lewisi populations were observed in central Japan. The relationships between the morphological variations and the prey body size were analysed. 3. In workers and queens, three morphological traits, head width, mandible length, and mandible width were significantly different among the multiple sites. Specifically, the mandible length was shorter in southwestern Japan than in other sampling locations. The ancova model revealed that the allometry of the mandible length to the head width was different among the sites. 4. As predicted, the mandible length was positively correlated with the average body size of collembolans in the Entomobryidae family. Furthermore, multiple regression analysis showed that the variation of the mandible length was affected by environmental factors represented as location information. However, the effect of collembolan body size was more effective at predicting mandible length. The study suggests that the geographical variation of mandible morphologies in S. lewisi has been selected by predator–prey interactions with collembolans.     

Heritability of male mandible length in the stag beetle Cyclommatus metallifer

Numerous coleopteran species express male‐specific “weapon traits” that often show size variations among males, even within a single population. Many empirical studies have demonstrated that environmental conditions during development affect absolute weapon size. However, relatively few studies in horned beetles support the hypothesis that the relationship between weapon size and body size, also referred to as a “scaling relationship” or “static allometry”, is largely determined by genetic factors. In this study, the heritability of absolute mandible length and static allometry between mandible length and body size were estimated in the stag beetle Cyclommatus metallifer. While no significant heritable variation was observed in absolute mandible length, high heritability (h² = 0.57 ± 0.25) was detected in the static allometry between mandible length and body size. This is the first report on the genetic effect on male mandible size in Lucanidae, suggesting that absolute mandible size is largely determined by environmental conditions while the static allometry between weapon size and body size is primarily determined by genetic factors.     

Phylogeography of the leaf beetle Chrysolina virgata in wetlands of Japan inferred from the distribution of mitochondrial haplotypes

The genetic differentiation among populations of the leaf beetle Chrysolina virgata living in wetlands of Japan was studied based on the sequence data of the mitochondrial cytochrome oxidase subunit I gene region (750 bp). Two distinct lineages of mitochondrial haplotypes were found: one (clade A) consisted of 26 haplotypes distributed over the distribution range of C. virgata between north‐east Honshu and Kyushu, whereas the other (clade B) was monotypic and confined to a small region in north‐east Honshu where it coexisted with clade A. Nested clade analysis for these haplotypes suggested that range expansion and following differentiation due to isolation by distance might have resulted in the present distribution pattern of the haplotypes in clade A. We discuss the evolutionary process leading to the occurrence of two distinct haplotype clades in Japan in terms of repeated colonization from the continent and range expansion and contraction during climatic changes.     

Variation in the species of freshwater harpacticoid copepods in Japan. II. Attheyella nakaii (Brehm)

Morphological variation in Attheyella nakaii, a dominant species of the genus in the mountain waters of Japan, is presented. The greatest proportion of male type furcal rami in the females of local populations is seen at the northern border of the range, where this species shows a sympatric distribution with A. yesoensis. Diminution of the inner lateral setae of endopodites on legs 2–4 occur in Kyushu and the Ryukyu Islands. The body size and relative length of the leg 5 exopodite show a latitudinal cline. The population of the type locality of this species exhibits a truncated form of furcal rami.     

Rapid evolution of photoperiodic response in a recently introduced insect Ophraella communa along geographic gradients

The introduced beetle Ophraella communa was first found in 1996 in Japan and has rapidly expanded its distribution to include regions that encompass a wide range of latitude and altitude and are dominated by different host‐plants. In this study, we investigated geographic variation in its photoperiodic response for the induction of reproductive diapause, with which the beetle adjusts its life cycle to local climate and host‐plant phenology. The beetle lines were collected from 18 sites in Japan. The diapause incidence under a photoperiodic condition of 13 h light : 11 h dark (LD 13:11) and the critical day length differed among the beetle lines. Analysis with the generalized linear model showed that latitude, altitude and host‐plant species (Ambrosia artemisiifolia vs. Ambrosia trifida) had significant effects on diapause incidence under LD 13:11. These results suggest that the O. communa populations have rapidly adapted to local environmental conditions after their colonization. However, the photoperiodic response of the O. communa population in Tomakomai, the northernmost part of its distribution range in Japan, deviated significantly from the general trend. We suggest that this deviation is attributed to either: (i) that this beetle has colonized Tomakomai more recently compared to the other sites; or (ii) that the Tomakomai population has adapted to local environments in a different way from other populations.     

Genetic differences and phenotypic plasticity in body size between high- and low-altitude populations of the ground beetle Carabus tosanus

The body size of a univoltine carabid beetle Carabus tosanus on Shikoku Island, Japan, was clearly smaller in higher‐altitude populations (subspecies), which possibly represents incipient speciation. To explore the determinants of altitudinal differences in body size in this species, we studied the degree of phenotypic plasticity by conducting rearing experiments at two constant temperatures and examined genetic differences through interpopulation crosses. At 15 °C, C. tosanus had a longer developmental period and a shorter adult body than at 20 °C. Nevertheless, variation in body size due to temperature effects (phenotypic plasticity) was small compared to the interpopulation differences, which suggests substantial genetic differences between populations (subspecies) at different altitudes. In F₁ offspring from crosses between a low‐altitude (subspecies tosanus) and a high‐altitude population (subspecies ishizuchianus), adult body length was affected by the genotypes of both parents, with an interaction effect of parental genotype and offspring sex. Further analyses revealed that adult body length was affected by sex‐linked factors in addition to autosomal factors. These genetic differences in body size may have resulted from adaptations to different altitudes and may be important for the process of incipient speciation because body size differences could contribute to premating reproductive isolation.     

Size variation and mating success in the stag beetle, Lucanus cervus

Abstract.  The stag beetle, Lucanus cervus , is Britain's largest beetle, with a patchy distribution in southern England. The literature suggests that it displays exceptional size variation, particularly in the males, but no analysis of size inequality has ever been conducted. In the present study, stag beetle adults are measured and allometric relationships derived between various parameters and total body length. Most of the specimens found each year are fragments and head width can be used as a good predictor of total body length in each sex. Body size differs between years and between localities and male beetles show a greater degree of size inequality than females. However, L. cervus does not show greater inequality in size than many other beetle species and populations of males are composed of a relatively large number of small individuals. These males are not at a disadvantage in mating because it is the ratio of male : female size that determines mating success. Very large males are less successful in mating and it is suggested that lack of mating success may act as an additional constraint on mandible size in this species. Size variation is most likely caused by variation in larval food resources, coupled with variation in local climatic conditions.     

Habitat segregation in four species of adult tiger beetles (Coleoptera: Cicindelidae)

ABSTRACT.

• 1 Habitat segregation in four species of tiger beetles, Cicindela cancellata Dejean, C.cardoni Fleutians, C.minuta Olivier and C.sumatrensis Herbst in a river bank ecosystem was studied in dry and wet seasons.
• 2 The four species segregated distinctly along the river beds into separate habitats, with occasional overlapping in both the seasons.
• 3 Among the habitat characteristics considered, vegetation, soil moisture and available prey-size were found to be important in species segregation.
• 4 D ²-analysis showed that the habitat preferences of C.cardoni and C.cancellata were closer to that of C.sumatrensis. The habitat of C.minuta was distinctly separated.
• 5 There was a positive correlation between the mandible length of each tiger beetle species and the length of prey captured.
• 6 Variance in mandible length within species was related to prey-length distribution pattern and to variance in habitats.

     

Genetic variation of two species with different life‐history traits in the endangered renosterveld of South Africa – a comparative analysis of Eriocephalus africanus and Hemimeris racemosa

We tested the effects of life‐history traits on genetic variation and conducted a comparative analysis of two plant species with differing life‐history traits co‐occurring in the highly endangered renosterveld of South Africa. We selected eighteen renosterveld remnants with varying degrees of size and isolation where populations of the herbaceous, annual and insect‐pollinated Hemimeris racemosa and the shrubby perennial and both wind‐ and insect‐pollinated Eriocephalus africanus occurred. We postulated a lower genetic variation within populations and increased genetic variation between populations in the annual than in the perennial species. Genetic variation was lower within populations of H. racemosa than within E. africanus, as is typical for annual compared to perennial species. Variation within populations was, however, not correlated with fragment size or distance in either of the two species and genetic variation between populations of the two species was comparable (Φ_ST = 0.10, 0.09).     

Temperature‐size responses match latitudinal‐size clines in arthropods,revealing critical differences between aquatic and terrestrial species

Two major intraspecific patterns of adult size variation are plastic temperature‐size (T‐S) responses and latitude‐size (L‐S) clines. Yet, the degree to which these co‐vary and share explanatory mechanisms has not been systematically evaluated. We present the largest quantitative comparison of these gradients to date, and find that their direction and magnitude co‐vary among 12 arthropod orders (r² = 0.72). Body size in aquatic species generally reduces with both warming and decreasing latitude, whereas terrestrial species have much reduced and even opposite gradients. These patterns support the prediction that oxygen limitation is a major controlling factor in water, but not in air. Furthermore, voltinism explains much of the variation in T‐S and L‐S patterns in terrestrial but not aquatic species. While body size decreases with warming and with decreasing latitude in multivoltine terrestrial arthropods, size increases on average in univoltine species, consistent with predictions from size vs. season‐length trade‐offs.     

Genital lock-and-key system and premating isolation by mate preference in carabid beetles (Carabus subgenus Ohomopterus)

The shapes and lengths of copulatory pieces and vaginal appendices of the carabid beetle subgenus Ohomopterus (genus Carabus) vary among species. In Japan, the species in the group with a medium body size (C. yaconinus, C. iwawakianus, C. maiyasanus, C. uenoi, C. arrowianus, C. esakii, and C. insulicola) are usually allopatric or parapatric, except at Mt Kongosan, where C. uenoi, C. iwawakianus, and C. yaconinus are sympatrically distributed. The degree of premating isolation by mate preference was high between sympatric populations, irrespective of the genetic distance between them. However, premating isolation was absent between parapatric populations. The degree of premating isolation for allopatric populations spanned a wide range of isolation values. Thus, mate discrimination by males seems to have evolved mostly between sympatric pairs. These results suggest two hypotheses. First, premating isolation has evolved through reinforcement or through reproductive character displacement after sympatric contact. Second, premating isolation has evolved in allopatry, and as a result of premating isolation, the species can coexist in sympatry. We also examined the degree of mechanical isolation between C. uenoi and C. iwawakianus (a sympatric pair), which have a very large difference in the length of the copulatory piece. The insertion success was low and only one female produced viable offspring among 15 crosses; however, death in females due to copulation was rare. For sympatric matings between C. uenoi and C. iwawakianus, a large difference in the genital size might reduce the gene flow with small mating costs. Gene flow that was significantly reduced by genital difference might cause either the evolution of premating isolation through reinforcement/reproductive character displacement or through the maintenance of a high degree of premating isolation following sympatric contact. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society, 2006, 87 , 145–154.     

Size and shape changes of Ips typographus L. (Coleoptera: Scolytinae) in relation to population level

1 High intraspecific competition is known to occur during Ips typographus outbreaks, and is thought to be the main factor regulating epidemic populations by affecting beetle population productivity. However, little is known about the consequences of intraspecific competition on population quality during outbreaks, although it could have consequences on beetle population dynamics. 2 Ips typographus morphological variations among localities, years and beetle population levels were investigated in 10 Norway spruce (Picea abies) stands having various beetle damage intensities. Beetle size and shape estimators based on wing length, and using isometric size and log‐shape ratios, were employed. Field‐caught beetles were compared with beetles emerging from controlled breeding at different densities, performed in the laboratory. Beetles from this colony were also used to check the influence of breeding densities on the size estimator. 3 Size variations occurred among localities and years and were consistent with the epidemic or latent status of the beetle populations. Controlled breeding confirmed the negative effect of beetle densities encountered in the field on offspring size. Two hypotheses are formulated to explain this increase of intraspecific competition during an outbreak, but our data support the effect of host quality change between latent and epidemic populations. 4 Shape variations also occurred among localities but were unrelated to beetle population levels. No groups consistent with a geographical structure were found, suggesting low genetic variation for I. typographus populations in France.     

Geographic variation of host use in the leaf beetle Agelasa nigriceps suggests host range expansion

Host range expansion is an important event in the evolution of host use in phytophagous insects. Herein, we report geographic variation of host use in the chrysomelid leaf beetle, Agelasa nigriceps Motschulsky (Coleoptera: Chrysomelidae), and suggest that this beetle is expanding its host range. This beetle has been recently recorded on Pterostyrax hispidus Sieb. et Zucc. (Styracaceae) in addition to its common host plant Actinidia arguta (Sieb. et Zucc.) Planch. ex Miq. (Actinidiaceae). The A. arguta‐associated populations were widely found in Japan, whereas the P. hispidus‐associated populations were found only in central and southwestern Japan. In the present study, we examined adult feeding behavior and larval performance of 12 A. nigriceps populations collected from eight localities, four localities where beetles occurred only on A. arguta (allopatric localities) and four localities where A. arguta‐ and P. hispidus‐associated populations occurred sympatrically (sympatric localities). Beetles of all populations, irrespective of their host plants and localities, showed high acceptance of and high larval performance on A. arguta leaves. In contrast, we found considerable variation in the beetle response to P. hispidus leaves. The A. arguta‐associated populations of allopatric localities scarcely accepted P. hispidus leaves, whereas those of sympatric localities, particularly those of P. hispidus‐associated populations, accepted and grew on P. hispidus leaves, although the degree of acceptance and larval performance varied among localities. These results strongly suggest that A. arguta is the ancestral host for A. nigriceps, and host range expansion to the P. hispidus has occurred in this beetle.     

Character displacement drives floral variation in Pelargonium (Geraniaceae) communities

Interactions between plant community members are an underexplored driver of angiosperm floral variation. We investigate character displacement as a potential contributor to floral variation in Pelargonium communities. Pelargoniums all place pollen on the ventral sides of their pollinators, potentially leading to interspecific pollen transfer (IPT) in sympatry. We show that the positions of pollen placement and receipt are determined by anther and style exsertion lengths. Using field experiments, we demonstrate that heterospecific species experience higher IPT if they have similar style lengths than when they have greater style length differences. Using crosses, we show that IPT has negative consequences on seed set. In combination, these results suggest that character displacement in style length is likely to reduce IPT and increase female fitness in sympatry. Patterns of style length variation across 29 different Pelargonium communities suggest that character displacement has occurred in multiple communities. Furthermore, analyses using a wide-ranging species pair show that style lengths are more different between sympatric populations than they are between allopatric populations. In addition to pollinators as agents of floral divergence, this study suggests that variation in Pelargonium community structure has driven style length variation through character displacement.     

Mandibular shape correlates of tooth fracture in extant Carnivora: implications to inferring feeding behaviour of Pleistocene predators

Percentages of tooth fracture and mandible shape are robust predictors of feeding habits in Carnivora. If these parameters co‐vary above the species level, more robust palaeobiological inferences could be made on fossil species. A test of association is presented between mandible shape and tooth fracture in a subset of extant carnivorans together with large Pleistocene fossil predators from Rancho La Brea (Canis dirus, Panthera atrox, and Smilodon fatalis). Partial least square (PLS) and comparative methods are employed to validate co‐variation of these two parameters in extant carnivorans. Association between mandible shape and percentage of tooth fracture is strongly supported, even if both blocks of data exhibit a phylogenetic signal to a different degree. Dietary adaptations drive shape/fracture co‐variation in extant species, although no significant differences occur in the PLS scores between carnivores and bone/hard food consumers. The fossil species project into PLS morphospace as outliers. Their position suggests a unique feeding behaviour. The increase in the size of prey, together with consumption of skin and hair from carcasses in a cold environment, might have generated unusual tooth breakage patterns in large predators from Rancho La Brea. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 70–80.