Descriptive biogeography of Tomicus (Coleoptera: Scolytidae) species in Spain

Aim, location Tomicus (Coleoptera, Scolytidae) species are some of the principal pests of Eurasian forest and are represented by three coexisting species in Spain, Tomicus piniperda (Linnaeus, 1758), Tomicus destruens (Wollaston, 1865) and Tomicus minor (Harting, 1834). The distribution of two taxa are unknown as they have until recently been considered separate species. Therefore, we model the potential distribution centres and establish the potential distribution limits of Tomicus species in Iberia. We also assess the effectiveness of different models by comparing predicted results with observed data. These results will have application in forest pest management. Methods Molecular and morphological techniques were used to identify species from 254 specimens of 81 plots. For each plot, a Geographical Information System was used to extract a set of 14 environmental (one topographic, six climatic) and biotic variables (seven host tree distributions). General Additive Models and Ecological Niche Factor Analysis models are applied for modelling and predicting the potential distribution of the three especies of Tomicus. Results The results of both modelling methodologies are in agreement. Tomicus destruens is the predominant species in Spain, living in low and hot areas. Tomicus piniperda occurs in lower frequency and prefers wet and cold areas of north‐central Spain. We detected sympatric populations of T. destruens and T. piniperda in Northern coast of Spain, infesting mainly P. pinaster. Tomicus minor is the rarest species, and it occupies a fragmented distribution located in high and wet areas. The remarkable biotic variable is the distribution of P. sylvestris, incorporated into the models of T. destruens and T. piniperda. Main conclusions These results indicate that in wet areas of north‐central Spain where T. piniperda occurs (and possibly the high altitudes of the southern mountains), T. destruens has a climatic distribution limit. In the northern border of this area, both species overlap their distributions and some co‐occurrences were detected. Tomicus minor potentially occurs in high and wet fragmented areas.     

Distribution and population genetic structure of the Mediterranean pine shoot beetle Tomicus destruens in the Iberian Peninsula and Southern France

1 The Mediterranean pine shoot beetle Tomicus destruens has long been indistinguishable from its congeneric Tomicus piniperda. Both species attack pines, and can be found in sympatry. The geographical distribution of T. destruens is still unclear in most of the Mediterranean Basin. 2 We aimed to describe the geographical distribution and zones of sympatry of both species in the Iberian Peninsula and France, and to study the molecular phylogeographical pattern of T. destruens. 3 Tomicus spp. adults were sampled in Portugal, Spain and France, and a portion of the mitochondrial genes COI and COII was sequenced for 84 individuals. Sequences were aligned to a data set previously obtained from French localities. 4 Tomicus destruens was found in all populations, except for one locality in Portugal and in the Landes (France). It was in sympatry with T. piniperda in two locations on Pinus pinaster and one location on Pinus radiata. 5 Within‐population genetic diversity was high, but we found a significant pattern of spatial distribution of genetic variation, as well as a significant effect of the host tree. 6 The data suggest the existence of two glacial refugia, from which T. destruens recolonized its current range. One refugium was located in Portugal where the beetle probably evolved on P. pinaster. The corresponding haplotypes show a West–East frequency gradient. The other refugium was probably in the eastern range, where the beetles evolved on Pinus halepensis and P. pinea. The corresponding haplotypes show an East–West frequency gradient.     

Predicting the distribution of the two bark beetles Tomicus destruens and Tomicus piniperda in Europe and the Mediterranean region

• 1 Various factors such as climate and resource availability influence the geographical distributions of organisms. Species sensitive to small temperature variations are known to experience rapid distribution shifts as a result of current global warming, sometimes leading to new threats to agriculture and forests. Tomicus piniperda and Tomicus destruens (Coleoptera, Curculionidae, Scolytinae) cause economic damage to pines in Europe and around the Mediterranean Basin. However, their respective potential distributions have not yet been studied at a large scale. The present study aimed to investigate the influence of climatic and host factors on the geographical distributions of both Tomicus species in Europe and around the Mediterranean Sea, and to establish maps of suitable areas.
• 2 Using 114 published localities where the presence or absence of both species was unambiguously recorded, we gathered WorldClim meteorological records to correlate the occurrence of insects with bioclimatic variables and to build potential distribution maps.
• 3 The two studied Tomicus species presented parapatric distributions and opposite climate demands, with T. destruens occurring in locations with warmer temperatures, whereas T. piniperda occurs under a colder climate. Amongst the investigated climate variables, temperature appeared to be most correlated with both species distributions.
• 4 The potential ranges of both species were further restricted by the availability of pine hosts. It appeared that setting new pine plantations in regions where T. destruens or T. piniperda are still absent could favour a rapid expansion of their distributions. Our data will be useful when aiming to apply management strategies adapted to each species, and to forecast their potential range expansions/contractions as a result of climate warming.

     

Impacts of Tomicus minor on distribution and reproduction of Tomicus piniperda (Col., Scolytidae) on the trunk of the living Pinus yunnanensis trees

This study investigated the impacts of Tomicus minor on Tomicus piniperda when the two Tomicus species coexist in the trunks of living Yunnan pine (Pinus yunnanensis L.) trees growing in the Kunming region, in south-western China. Tomicus piniperda mostly locates in the mid- and upper trunks of Yunnan pine tree; whereas T. minor mainly attacks the mid- and lower trunks. In the mid-trunk area from 1.0 to 5.0 m above ground, there are overlapping attack zones for the two Tomicus species, which accounts for an average of 80% of the entire zone occupied by T. piniperda and an average of 70% of the zone occupied by T. minor. In correspondence with their attack distributions, the average attack densities of the two species varied with height along the trunk, with 165.3 egg galleries per m² at a height of 6 m for T. piniperda, and 138.2 egg galleries per m² at the 1 m height for T. minor. It is suggested that T. minor adjusts its attack pattern with respect to T. piniperda, and thereby minimizes interspecific competition. No remarkable difference of average T. piniperda egg gallery length was found between the zone in which only T. piniperda occurred and the zone in which T. piniperda and T. minor coexisted; this is suggested to be due to low host quality in the upper trunk region where only T. piniperda was present. The number of T. piniperda larval galleries was highest when only T. piniperda was present, and decreased as T. piniperda and T. minor coexisted, particularly in the case when the density of T. piniperda was less than that of T. minor. Average larval density was 1649 larval galleries per m² where only T. piniperda occurred. However, when T. piniperda coexisted with T. minor, T. piniperda larval density averaged 1010 per m² when T. piniperda density was higher than T. minor, and averaged 442 per m² when T. piniperda density was less than T. minor, which led to the conclusion that T. minor makes a negative impact on T. piniperda reproduction when the two Tomicus species jointly colonize the same trunk of Yunnan pine tree.     

Shoot feeding ecology of Tomicus piniperda and T. minor (Col., Scolytidae) in southern China

Abstract: The pine shoot beetle, Tomicus piniperda, is causing substantial tree mortality to Pinus yunnanensis in south‐western China, whereas the same species in Europe seldom kills Pinus sylvestris. In order to understand this difference in aggressiveness, we studied the shoot feeding ecology of the pine shoot beetles in Yunnan, and compared it with European results. We found many similarities in the shoot feeding behaviour of T. piniperda, and also that of Tomicus minor, which was common locally. In contrast to Europe, however, the pine shoot beetles in Yunnan seem to be able to predispose the host trees for stem attack by intensive shoot‐feeding. It has also been observed that beetles aggregate in certain trees during shoot feeding, but we could not verify that in our experiment.     

纵坑切梢小蠹对云南松蛀害研究

在昆明地区,纵坑切梢小蠹Tomicus piniperda L.表现出枝梢聚集、树干蛀害等重要的行为学特征,形成三种基本蛀害模式。横坑切梢小蠹、蓝色伴生真菌参与了纵坑切梢小蠹危害过程,并在其中发挥积极作用。上述因素的综合影响,加强了纵坑切梢小蠹对云南松Pinus yunnanensis寄主树木的危害能力。     

Colonization of disturbed Scots pine trees by bark‐ and wood‐boring beetles

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Breeding performance and longevity of Tomicus destruens on Mediterranean and continental pine species

The pine shoot beetle Tomicus destruens (Wollaston) (Coleoptera: Curculionidae: Scolytinae) is one of the main pests of Mediterranean forests, where it is oligophagous on Mediterranean pines. However, possible global warming may make the insect move to higher latitudes and altitudes, allowing it to attack new pine species. In this respect, the aim of the present article was to assess both the acceptance and performance of T. destruens offered host and non‐host pine species. A no‐choice breeding experiment was set up under laboratory conditions, using logs of three Mediterranean (Pinus pinea L., Pinus pinaster Miller, and Pinus halepensis Aiton) and two continental (Pinus nigra Arnold and Pinus sylvestris L.) pine species. Log debarking at the end of adult emergence assessed parent fecundity, egg, and larval mortality. The quality of callow adults emerging from each tested pine was evaluated on the basis of their longevity on a semiartificial diet. Tomicus destruens colonised all tested pine species, but did not reproduce in Scots pine, taking about 79 days to complete development with no differences among pines tested. The best breeding performance, evaluated as female fecundity and adult production, was observed in P. halepensis, and the lowest in P. pinaster. On average, adults emerging from P. pinea survived longer (83 days) than from other pines, and adult longevity was the lowest in males emerging from P. nigra. Austrian pine, which under natural conditions is usually not a host of T. destruens, allowed insect development and adult production similar to P. pinea and P. pinaster.     

Attack dynamics of the pine shoot beetle, Tomicus piniperda (Col.; Scolytinae) in Scots pine stands defoliated by Bupalus piniaria (Lep.; Geometridae)

Abstract 1 After a 1‐year, extensive pine looper (Bupalus piniaria) outbreak, plots were laid out to study tree susceptibility to attack, and performance of Tomicus piniperda in pine trees suffering from varying levels of defoliation. 2 Tomicus piniperda was the dominating stem‐attacking species among the primary stem colonizers, and 82% of all trees that died had been colonized by T. piniperda. 3 Beetle attacks primarily struck severely defoliated trees, i.e. trees that suffered from 90% to 100% defoliation. 4 Beetle attacks peaked in the second year after cessation of the outbreak, and suppressed trees were both more frequently attacked and more susceptible to beetle attack than intermediate and dominant trees. 5 Trees surviving beetle attacks carried more foliage than trees that did not survive the attacks. 6 A single year of severe defoliation is enough to render pine trees susceptible to secondary pests, such as T. piniperda.     

Hierarchical structure of mitochondrial lineages of Tomicus destruens (Coleoptera,Scolytidae) related to environmental variables

We investigated here the relation of environmental variables with the distribution of mitochondrial lineages using a bark beetle species of Mediterranean distribution as a model. We analysed a total number of 460 DNA sequences of Tomicus destruens provided by intensive and extensive collection and GenBank entries. We combined phylogeography and regression models to study the role of five environmental predictors at fine scale in the distribution of a local genealogy. The analysis revealed a high genetic diversity, with 52 haplotypes present in Sierra Espuña forest (SE Spain) and 21 haplotypes in the other 14 Spanish populations, all included in the western clade of the Mediterranean phylogeography of the species. We found a micro‐distribution of the species related to altitude and putative niche segregation between lineages associated with the micro‐environmental conditions of their host pine trees. We compared the phylogeographic hypothesis obtained here with the phylogeography obtained integrating our data with all data published elsewhere. Here, we demonstrate a relation between the environmental heterogeneity and the haplotypic diversity at Mediterranean Basin scale. This analysis allows us to support the evolutionary scenario where the phylogeography and current molecular diversity of T. destruens is a consequence of the recolonization from two principal refugia at both extremes of the Mediterranean Basin and, according to our data, we propose that the heterogeneity of habitats allows fixation of the mitochondrial lineages.     

Phylogeography and conservation genetics of the hellbender salamander (Cryptobranchus alleganiensis)

We investigated hellbender phylogeography through phylogenetic analyses of individuals sampled from 16 locations throughout their range in the eastern United States. Analyses were conducted on concatenated cytochrome-oxidase I (COI), cytochrome-b (Cytb) and NADH dehydrogenase subunit 4 (ND4) mtDNA sequence, totaling 2160 nucleotides. Hellbender haplotypes differed by 0.1 to 5.8% maximum likelihood (ML) corrected sequence divergence. Phylogenetic analyses reveal that hellbenders are separated into 8 reciprocally monophyletic populations or clades differentiated by a minimum of 0.7 to 5.4% sequence divergence, each of which constitutes a separate Management Unit (MU). High among population divergence and reciprocal monophyly suggest that female-mediated gene flow is severely restricted or non-existent among each MU. Hellbenders are currently divided into two subspecies, Cryptobranchus alleganiensis alleganiensis and C. a. bishopi based on morphological characters. The phylogenetic analyses presented here strongly indicate that these subspecies are paraphyletic. Management priorities for the hellbender should be reconsidered in light of these new molecular data. Results from Bayesian rooting indicate the root of the hellbender mtDNA tree lies on the branch leading to hellbender haplotypes from the Current, Eleven Point and New Rivers. The rooted tree suggests that a common ancestor in the southern Ozarks and/or southern Appalachians gave rise to northern hellbender populations, consistent with a Pleistocene refuge hypothesis.     

Sequence data reflect the introduction pathways of the Sirex woodwasp parasitoid,Ibalia leucospoides (Ibaliidae,Hymenoptera)

1. The parasitoid wasp Ibalia leucospoides is native to the northern hemisphere and has been introduced to the southern hemisphere as a biological control agent for the invasive woodwasp Sirex noctilio. Two subspecies of the parasitoid, Ibalia leucospoides leucospoides (Palearctic distribution) and Ibalia leucospoides ensiger (Nearctic distribution), were introduced and are reported to have hybridized.
2. Despite extensive records of the numbers and origins of the wasps imported into the southern hemisphere, nothing is known regarding their current population diversity. We investigated the genetic variation of I. leucospoides in its native and introduced ranges using mitochondrial (COI) and nuclear (ITS) markers.
3. Mitochondrial DNA diversity in the introduced range was limited, with only five haplotypes, although sequence divergence between these haplotypes was high. Similarly, the ITS rDNA sequences revealed multiple clades present in the introduced range.
4. These results reflect introductions from a wide geographical range but where genetic bottlenecks have possibly reduced the genetic diversity. The data further reflect the origin of the I. leucospoides populations in South America and South Africa from New Zealand or Australia. We found no evidence of hybridization between the two subspecies of the parasitoid in its introduced range, and no evidence that I. leucospoides ensiger has established outside its native range.

     

Phylogeography of the giant harlequin beetle (Acrocinus longimanus)

Aim The extent to which cryptic species contribute to neotropical diversity remains inadequately investigated. Based on its highly distinctive morphology, the giant harlequin beetle, Acrocinus longimanus, is currently described as a single species, ranging from southern Mexico to northern Argentina. However, the discovery of cryptic species in Cordylochernes scorpioides, a pseudoscorpion with obligate dependence on the harlequin beetle for dispersal, strongly suggests the existence of barriers to gene flow in A. longimanus. The aim of this study was therefore to determine whether levels of DNA divergence between geographical populations provided evidence of genetically distinct lineages in the harlequin beetle. Location Trinidad and Panamá. Methods Sequencing of 1245 bp of the mitochondrial cytochrome oxidase I (COI) gene of A. longimanus from seven locations in Trinidad and Panamá. Results Mitochondrial haplotype diversity in the harlequin beetle shows limited evidence of geographical structuring, with a maximum sequence divergence between populations of only 1.29%. This is an order of magnitude less than the level of COI divergence between harlequin beetle riding pseudoscorpions from the same geographical locations. Main conclusions The molecular data on populations from northern South America and Panamá are consistent with the current, morphologically based classification of A. longimanus as a single, pan‐neotropical species. In addition, the relatively low level of population divergence detected in this study indicates that speciation in the hitchhiking pseudoscorpion has occurred in the absence of significant barriers to gene flow in its beetle host. It is proposed that, in the harlequin beetle, the phylogenetic signal of colonization and vicariance associated with the formation of the Isthmus of Panamá has been obscured, although not fully erased, by historical and contemporary gene flow.     

Population structuring in the monogonont rotifer Synchaeta pectinata: high genetic divergence on a small geographical scale

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Morphological similarity and molecular divergence of Trochulus striolatus and T. montanus,and their relationship to sympatric congeners (Gastropoda: Pulmonata: Hygromiidae)

Shell morphology has played an important role in the differentiation of mollusc species. However, extensive morphological variation and the lack of readily available diagnostic characters often lead to confusion and controversy in taxonomy of closely related species, such as the genus Trochulus. Two nominal species Trochulus striolatus and T. montanus show only subtle differences in their reproductive systems and are indistinguishable on the basis of sole conchological traits. Therefore, we performed molecular and morphological analyses to establish robust species limits among the taxa. The canonical discriminant analysis (CDA) of shell characters confirmed strong similarity between the species. It also revealed high intraspecific variability of the shell morphology, which allowed the distinguishing of some populations as two extremely distinct forms, while simultaneously making it difficult to discriminate between the two species. Analysis of the genital morphology indicated that differences were continuous and practically negligible among the species. In contrast, phylogenetic analyses based on newly obtained 114 Trochulus sequences of the mitochondrial cytochrome c oxidase subunit I (COI) gene and 55 homologues retrieved from the GenBank database showed clear genetic divergence between T. striolatus and T. montanus. Moreover, they revealed a close relationship between some T. montanus, some T. caelatus and T. clandestinus samples, which formed a monophyletic group. Some of their haplotypes were even identical. It suggested that their recent common origin or recurrent gene flow occurred between these populations. However, T. sericeus sequences were split into independent diverged lineages, which were separated by many unidentified Trochulus species. It suggested that this taxon may represent a paraphyletic species complex.     

Diversity in the genus Rhabdias (Nematoda,Rhabdiasidae): Evidence for cryptic speciation

Lungworms from the genus Rhabdias are common parasites of amphibians and reptiles distributed worldwide. To assess the diversity of Rhabdias spp., we performed molecular analyses of 35 specimens sampled in different regions of Brazil. Molecular analyses were based on the internal transcribed spacer (ITS), large subunit (28S) ribosomal and the cytochrome oxidase I (COI) mitochondrial genes. DNA sequence divergence was compared among ribosomal and mitochondrial genes, analyses using the general mixed Yule‐coalescent (GMYC) method based on the COI gene were used to identify possible cryptic diversity, and phylogenetic analyses using concatenated ITS and 28S ribosomal genes were used to test the monophyly of Rhabdiasidae. We revealed five morphospecies: R. cf. stenocephala, R. breviensis, R. pseudosphaerocephala and two new species, Rhabdias sp.4 and Rhabdias sp.5. DNA sequence levels of divergence among genes ITS, 28S and COI were compared, and the efficiency of the molecular markers to identify species (ITS and COI) and lineages (COI) was tested. GMYC was assigned to 17 well‐supported clades (i.e., 17 species), and cryptic diversity was detected in the Neotropical region as evidenced by the multiple lineages in R. breviensis and R. pseudosphaerocephala. In addition, our results suggest evidence for host–parasite cophylogeny in the R. pseudosphaerocephala complex and dispersal events among their populations. Phylogenetic analyses supported the monophyly of Rhabdiasidae and improved the resolution of main clades. Rhabdias breviensis is closely related to Rhabdias cf. africanus, Rhabdias cf. stenocephala, R. pseudosphaerocephala, Rhabdias sp.4 and Rhabdias sp.5 grouping together in a main clade with Neotropical‐related species. The large geographical distribution appeared to be a phylogenetic pattern among the species of Rhabdias from the neotropics.     

Genetic structure of two Plusiinae species suggests recent expansion of Chrysodeixis includens in the American continent

1. The Plusiinae subfamily has many polyphagous species, many of which occur in South America. Chrysodeixis includens and Rachiplusia nu are two representatives that mainly occurs in soybeans, cotton, common beans, sunflower and alfalfa.
2. A population genetic study of C. includens and R. nu collected in the Southern Cone of America was performed using a partial COI gene sequencing data and compared with specimens from other American countries.
3. Six haplotypes were identified in C. includens populations of Brazil, Argentina, Paraguay and Uruguay, organized within a star-like haplotype network, with the most common haplotype identified as Chin_MC.
4. R. nu populations are more diverse and stable in comparison to C. includens. Populations from Argentina and Uruguay had the highest haplotype diversity, sharing five haplotypes and putatively indicating haplotype exchange.
5. Demographic change analysis suggested a recent population expansion of C. includens over the American continent.
6. Some C. includens haplotypes were country-specific, suggesting population expansion in the countries where specimens were collected.

     

Long-distance colonization of the Western Mediterranean by Cistus ladanifer (Cistaceae) despite the absence of special dispersal mechanisms

Aim There are few biogeographical and evolutionary studies that address plant colonization and lineage origins in the Mediterranean. Cistus serves as an excellent model with which to study diaspore dispersal and distribution patterns of plants exhibiting no special long‐distance dispersal mechanisms. Here we analyse the pattern of genetic variation and divergence times to infer whether the African–European disjunction of C. ladanifer L. is the result of long‐distance dispersal or of vicariance events. Location Principally the Western Mediterranean region, with a focus on the Strait of Gibraltar. Methods We used DNA sequence phylogenetic approaches, based on plastid (rbcL/trnK‐matK) and nuclear (ITS) sequence data sets, and the penalized likelihood method, to date the diversification of the 21 species of Cistus. Phylogenetic relationships and phylogeographical patterns in 47 populations of C. ladanifer were also analysed using two plastid DNA regions (trnS‐trnG, trnK‐matK). These sequence data were analysed using maximum parsimony, Bayesian inference and statistical parsimony. Results Dating estimates indicated divergence dates of the C. ladanifer lineage in the Pleistocene. Eight nucleotide‐substitution haplotypes distributed on the European (four haplotypes) and African (five haplotypes) sides of the Strait of Gibraltar were revealed from C. ladanifer sequences. Both the haplotype network and the phylogenetic analyses depicted two main Cistus lineages distributed in both Europe and North Africa. An Iberian haplotype forms part of the North African lineage, and another haplotype distributed on both continents is related to the European lineage. Haplotype relationships with respect to outgroup sequences supported the hypothesis that the centre of genetic diversity is in northern Africa. Main conclusions Based on lineage divergence‐time estimates and disassociation between geographical and lineage haplotype distributions, we inferred at least two intercontinental colonization events of C. ladanifer post‐dating the opening of the Strait of Gibraltar (c. 5 Ma). This result supports a hypothesis of long‐distance dispersal rather than a hypothesis of vicariance. We argue that, despite limited dispersal abilities, preference for disturbed habitats was integral to historical colonization after the advent of the Mediterranean climate (c. 3.2 Ma), when Cistus species diverged and became established as a dominant element in the Mediterranean scrub.     

Evaluation of DNA barcoding and identification of new haplomorphs in Canadian deerflies and horseflies

This paper reports the first tests of the suitability of the standardized mitochondrial cytochrome c oxidase subunit I (COI) barcoding system for the identification of Canadian deerflies and horseflies. Two additional mitochondrial molecular markers were used to determine whether unambiguous species recognition in tabanids can be achieved. Our 332 Canadian tabanid samples yielded 650 sequences from five genera and 42 species. Standard COI barcodes demonstrated a strong A + T bias (mean 68.1%), especially at third codon positions (mean 93.0%). Our preliminary test of this system showed that the standard COI barcode worked well for Canadian Tabanidae: the target DNA can be easily recovered from small amounts of insect tissue and aligned for all tabanid taxa. Each tabanid species possessed distinctive sets of COI haplotypes which discriminated well among species. Average conspecific Kimura two‐parameter (K2P) divergence (0.49%) was 12 times lower than the average divergence within species. Both the neighbour‐joining and the Bayesian methods produced trees with identical monophyletic species groups. Two species, Chrysops dawsoni Philip and Chrysops montanus Osten Sacken (Diptera: Tabanidae), showed relatively deep intraspecific sequence divergences (～10 times the average) for all three mitochondrial gene regions analysed. We suggest provisional differentiation of Ch. montanus into two haplotypes, namely, Ch. montanus haplomorph 1 and Ch. montanus haplomorph 2, both defined by their molecular sequences and by newly discovered differences in structural features near their ocelli.