The geometric approach to explore the Bactrocera tau complex (Diptera: Tephritidae) in Thailand

Specimens of the genus Bactrocera were collected from several host plants in northern and western Thailand. They were morphologically recognized as Bactrocera tau and were subdivided into eleven samples according to host plant, geographic origin and time of collection. Twelve landmarks of the right wing were described in a total of 264 males and 276 females. An exploratory analysis using kernel density estimates was performed on the multivariate morphometric space. Non-parametric classification highlighted the existence of two non-overlapping clusters within both males and females. The clusters were not congruent with geography. One cluster (cluster I) contained only one plant, Momordica cochinchinensis, the other one (cluster II) contained five different plants including M. cochinchinensis. Further morphometric analyses on selected samples indicated that the influence of the plants on the shape of the wing could not explain satisfactorily the presence of two clusters. Genetic techniques identified the presence of B. tau cryptic species C in M. cochinchinensis from cluster I, and of B. tau cryptic species A in Coccinia grandis from cluster II. Our working hypothesis is that the two clusters identified by geometric morphometrics were species A and C, respectively.     

Phenotypic evidence for hidden biodiversity in the <Emphasis Type="Italic">Merodon aureus</Emphasis> group (Diptera,Syrphidae) on the Balkan Peninsula: conservation implication

Cryptic species and phenotypic divergent units provided useful information about hidden biodiversity in the Merodon genus (Diptera, Syrphidae) on the Balkan Peninsula. Discrimination three cryptic species within both the M. aureus (M. aureus A, M. aureus B and M. aureus C) and M. cinereus (M. cinereus A, M. cinereus B and M. cinereus C) complexes was done by study of subtle wing variation (wing size and shape) using a geometric morphometric approach. Observed interspecific differentiation is generally in agreement with a previous study using molecular markers (allozyme loci, COI mtDNA). A considerable evolutionary and phenotypic intraspecies diversity of the M. aureus A, M. aureus B, M. aureus C, M. cinereus A, M. cinereus B, M. cinereus C and M. funestus species from the Balkan Peninsula has important implications for diagnosing biodiversity, including endemic and cryptic species. Observed phenotypic divergent units within the species might be considered as an evolutionary potential of the M. aureus group and used for defining conservation priorities. This study has contributed to the recognition of the value of wing traits in order to decipher the hidden diversity and evolutionary diversification.     

Population genetic structure of Zeugodacus tau species complex in Thailand

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Wing shape as a potential discriminator of morphologically similar pest taxa within the Bactrocera dorsalis species complex (Diptera: Tephritidae)

Four morphologically cryptic species of the Bactrocera dorsalis fruit fly complex (B. dorsalis s.s., B. papayae, B. carambolae and B. philippinensis) are serious agricultural pests. As they are difficult to diagnose using traditional taxonomic techniques, we examined the potential for geometric morphometric analysis of wing size and shape to discriminate between them. Fifteen wing landmarks generated size and shape data for 245 specimens for subsequent comparisons among three geographically distinct samples of each species. Intraspecific wing size was significantly different within samples of B. carambolae and B. dorsalis s.s. but not within samples of B. papayae or B. philippinensis. Although B. papayae had the smallest wings (average centroid size=6.002 mm±0.061 SE) and B. dorsalis s.s. the largest (6.349 mm±0.066 SE), interspecific wing size comparisons were generally non-informative and incapable of discriminating species. Contrary to the wing size data, canonical variate analysis based on wing shape data discriminated all species with a relatively high degree of accuracy; individuals were correctly reassigned to their respective species on average 93.27% of the time. A single sample group of B. carambolae from locality 'TN Malaysia' was the only sample to be considerably different from its conspecific groups with regards to both wing size and wing shape. This sample was subsequently deemed to have been originally misidentified and likely represents an undescribed species. We demonstrate that geometric morphometric techniques analysing wing shape represent a promising approach for discriminating between morphologically cryptic taxa of the B. dorsalis species complex.     

ITS2 sequence variations among members of Diachasmimorpha longicaudata complex (Hymenoptera: Braconidae) in Thailand

The parasitoid Diachasmimorpha longicaudata complex in Thailand contains at least 3 cryptic species informally designated as species D. longicaudata A, B and BB. DNA sequence data of nuclear ITS2 (second internal transcribed spacer) were used to characterize members of this D. longicaudata complex. The polymerase chain reaction (PCR) amplicon of ITS2 region of D. longicaudata B (≈ 650 bp) clearly differentiated this species from A and BB (amplicon of ≈ 590 bp). Sequence alignment of individual parasitoids revealed that low intraspecies differences ranged from 0.457 to 3.991%, but interspecies differences ranged from 7.566 to 12.989%. Phylogenetic trees constructed using Neighbor-Joining (NJ) and Maximum Parsimony (MP) methods, taking the parasitoid Psyttalia concolor complex as an outgroup, revealed that D. longicaudata A, B and BB formed a monophyletic group, with species A and BB being more closely related than species B. ITS2 characterization of D. longicaudata complex has revealed an interesting divergence of the three cryptic sibling species in Thailand.     

Competitive Interactions between Immature Stages of Bactrocera cucurbitae (Coquillett) and Bactrocera tau (Walker) (Diptera: Tephritidae) under Laboratory Conditions

The melon fly, Bactrocera cucurbitae (Coquillett), and the pumpkin fly, Bactrocera tau (Walker), are economically important pests that attack mainly cucurbitacean fruits. The two fruit fly species have similar natural distributions, host ranges, and population growth capacities. This study was designed to assess the asymmetrical competitions through resource exploitation between the larvae of B. cucurbitae and B. tau at different density levels and temperatures, and on different hosts by comparing the relative effects of interspecific and intraspecific interactions on four life history parameters: survival rate, puparial mass, puparial duration, and developmental duration. Our results showed that intraspecific and interspecific competitions occurred under some laboratory conditions, and B. cucurbitae took advantage over B. tau at the high-density level and at low and high temperatures on pumpkin, bitter gourd, and bottle gourd when interspecific competition took place. Intraspecific and interspecific competitions mainly affected the puparial mass and the survival rate of the two fruit fly species but had no marked effect on the puparial duration or development duration.     

Temporal dynamics of Bactrocera (Zeugodacus) tau (Diptera: Tephritidae) adults in north Jiangxi,a subtropical area of China revealed by eight years of trapping with cuelure

Bactrocera (Zeugodacus) tau (Walker) (Diptera: Tephritidae), a quarantine pest of international concern, has become widespread and economically important in South and Southeast Asia. It has been introduced into north Jiangxi from Fujian, China and become a threat to local fruits and vegetables. Field trapping with cuelure (CUE) as lure is needed, not only for monitoring population dynamics of B. tau but also for direct control of population. In this study, we collected weekly tephritids from 4 CUE-baited McPhail traps every year from 2008 to 2015, analysing seasonal and inter-year B. tau population fluctuations. The total number of captured tephritids reached 7811, including B. tau (5321), B. scutellata (Hendel) (2375) and B. cucurbitae (Coquillett) (1 1 5). Among them, B. tau was the dominant species. It had 3–4 generations a year in north Jiangxi with high generation overlapping. The average number of trapped B. tau males did not differ significantly in different years, while the average monthly number showed large variation from June to December. No B. tau males were trapped before June in any year. Adult males were present from mid-June to late December, with distinct peaks between late August and mid-October. This fruit fly can withstand below 13℃ as a monthly minimal temperature under field conditions, and adult capture rates were significantly related to monthly minimal temperature and monthly total precipitation. These results can be used to predict the occurrence time and population size of B. tau in different seasons, enabling growers and extension personnel to take efficient management measures before they damage the host crops.     

Addition to the revision of the Pristaulacus comptipennis species-group: Description of two new species from Laos and Thailand (Hymenoptera: Aulacidae)

Two species belonging to the recently revised Pristaulacus comptipennis species-group, P. manuelae Turrisi and Madl, sp. n. from Laos and P. iosephi Turrisi and Madl, sp. n. from Thailand, are described, illustrated and compared with most related species. The former species most resembles P. watanabei Turrisi and Smith, 2011, from Thailand, but is readily recognizable, mainly by the shape of the head, which is elongated, with a well-developed temple. The latter species resembles two other species, P. nobilei Turrisi and Smith, 2011, from south China, and P. emarginaticeps Turner, 1922, from Vietnam, but it is easily recognizable mainly by the shape of the medial occipital groove, which is shallow and V-shaped, as well as the shape of the mesosoma, with anterior lobe of mesoscutum, anterior to notauli, strongly elevated dorsally and anteriorly and distinctly overhanging pronotum.     

Allozyme data reveal genetic diversity and isolation by distance in sympatric Glyphidrilus Horst, 1889 (Oligochaeta: Almidae) of the Lower Mekong River Basin

The genus Glyphidrilus, comprised of semi-aquatic freshwater earthworms that live in an ecotone between terrestrial and freshwater ecosystems, are widely distributed along riverbanks and rice paddy systems. Two Glyphidrilus species (Glyphidrilus vangviengensis and Glyphidrilus mekongensis) are endemic in the Lower Mekong River Basin and are sympatric from Northern Thailand to Southern Laos. However, species delimitation among the Mekong Glyphidrilus remains unclear because the key morphological traits in semi-aquatic earthworms are highly polymorphic. This study assessed the distinction between G. vangviengensis and G. mekongensis using allozyme electrophoresis. A total of 752 individuals collected from 33 localities were screened for 10 putative loci from seven enzymatic systems, revealing that G. vangviengensis and G. mekongensis are two distinct species, according to their different allelic patterns and high genetic distance. A low genetic differentiation within each species was indicated by the pairwise Nei's D and F_ST analyses, and the absence of population structure was detected by AMOVA and Bayesian structure analyses. However, a significant isolation by distance, but not vicariance, was observed, which is probably due to the river current causing translocation downstream and so gene flow between adjacent localities. The genetic diversity of the Mekong Glyphidrilus was relatively high and comparable to other earthworm taxa, and several localities showed deviation from Hardy–Weinberg equilibrium. An additional cryptic species from Ban Hat Khamphi, Loei, Thailand, was inferred.     

Six cryptic species on a single species of host plant: morphometric evidence for possible reproductive character displacement

Abstract 1. Diversification of some highly host‐specific herbivorous insects may occur in allopatry, without shifts in host use. Such allopatric divergence may be accelerated by sexual selection operating on courtship displays. Wing size and shape may affect visual and vibrational courtship displays in tephritid fruit flies. Geometric morphometric methods were used to examine wings of six sympatric cryptic species in the neotropical genus Blepharoneura. All six species feed on flowers of the same species of host (Gurania spinulosa), a neotropical vine in the Cucurbitaceae. Three of the fly species court and mate in close proximity on the host. Thus, courtship behaviours could serve as important reproductive isolating mechanisms. Two sets of hypotheses were tested: (i) species differ in wing shape and wing size; and (ii) species are sexually dimorphic in wing size and wing shape. Wing size differed among a few species, but wing shape differed significantly among all six species. Sexual dimorphism in wing size was found in only one species, but sexual dimorphism in wing shape was found in two of the three species known to court on the same host plant. In the two sexually dimorphic species, wing shape differed among males, but not among females. This suggests that selection for reproductive character displacement might accelerate divergence in wing shape.     

Does character displacement demonstrate density-dependent expression in females? A test on the wing shape of two species of European damselflies

Character displacement (CD) is the evolutionary process which leads to the divergence in trait expression of closely related species in regions where species co-occur, compared to allopatric populations. In Europe CD has been investigated in males of Calopteryx splendens and C. virgo and has been related to species recognition. If species recognition is relevant for males, also females should benefit from CD. The most obvious differences between females of these two species are wing profile and colour. We sampled females from allopatric and from sympatric populations with different relative abundances of these species. Wing shape and pigmentation were evaluated for each damselfly. CD was found in wing profile but not in wing transparency. The relative abundance of species significantly affected CD, but with a different pattern in each species. The prediction that wing shape become more different from the allopatric state when the species was relatively rare, but more similar to the allopatric state when the species was common was evident only for C. splendens. Wing shape changes might increase differences in flying patterns making males more effective to discriminate between heterospecific females. So, CD we observed may be the result of a selection directed to reduce interspecific reproductive interference.     

DNA barcoding reveals the occurrence of cryptic species in host-associated population of Conogethes punctiferalis (Lepidoptera: Crambidae)

Conogethes punctiferalis (Guénee) is a critical pest that commonly infests castor (Ricinus communis Linnaeus) and cardamom (Elettaria cardamomum Maton) in India. The moths of both castor and cardamom appear to be similar in wing pattern and color. However, the results of behavioral studies elicited a doubt that there may be differences in terms of host specialization. In the present study, we conducted morphological studies and DNA barcode analyses using cytochrome oxidase I gene, which unraveled the mystery of C. punctiferalis breeding on castor and cardamom. The differences in male aedeagus and female bursae were prominent, yet, not sufficient enough to say that they are different species. The results showed high haplotype diversity (0.817 ± 0.073) and nucleotide diversity (0.0285 ± 0.002) in C. punctiferalis. In addition, topologies of neighbor-joining trees indicate that Conogethes sp. breeding on castor belongs to C. punctiferalis while those on cardamom are of a separate clade. Further genetic analysis revealed significant genetic differentiations among the two sampled populations, reflecting limited gene flow. Neutrality tests and mismatch distributions showed population expansion in C. punctiferalis, while the results of an analysis of molecular variance (AMOVA) indicated the existence of significant genetic variation among the examined host races. Conclusively, analysis using mitochondrial DNA showed an amount of genetic divergence between the two host-associated populations compatible with cryptic species rather than host races.     

Wing geometry of Phlebotomus stantoni and Sergentomyia hodgsoni from different geographical locations in Thailand

Geographic populations of the two main sandflies genera present in Thailand were studied for species and population identification. Size and shape of Phlebotomus stantoni and Sergentomyia hodgsoni from different island and mainland locations were examined by landmark-based geometric morphometrics. Intraspecific and interspecific wing comparison was carried out based on 12 anatomical landmarks. The wing centroid size of P. stantoni was generally larger than that of S. hodgsoni. Within both species, wings from the continent were significantly larger than those from island populations. Size variation could be significant between geographic locations, but could also overlap between genera. The wing venation geometry showed non-overlapping differences between two species. The within-species variation of geometric shape between different geographical locations was highly significant, but it could not interfere with the interspecies difference. The lack of species overlapping in shape, and the high discrimination between geographic populations, make geometric shape a promising character for future taxonomic and epidemiological studies.     

World Catalogue of the Family Ratardidae (Lepidoptera,Cossoidea) with Description of a New Species from Thailand and Establishment of New Synonymy and a New Status for <Emphasis Type="Italic">Ratarda javanica</Emphasis> Roepke, 1937

An illustrated catalogue is given of the family Ratardidae in the world fauna which includes 10 species from 3 genera. A new species, Callosiope elenae Yakovlev sp. n., is described from Lampang Province, Thailand. Externally, the new species differs well from the type species of the genus, C. banghaasi. The fore wing of C. elenae sp. n. has a pattern of alternating wide black bands against pure white background (in C. banghaasi, the fore wing is strongly darkened, without bands). The hind wing of C. elenae sp. n. shows a pattern of large dropshaped black smears at the wing edge and a more or less expressed spotted pattern in the discal and postdiscal areas (in C. banghaasi, the hind wing is completely black, with no pattern). A new synonymy: Ratarda marmorata Moore, 1879 = Ratarda guttifera Hering, 1925 syn. n. and a new status: Ratarda mora javanica Roepke, 1937 stat. n., are established. Ratarda excellens (Strand, 1917) is recorded in the fauna of Thailand for the first time.     

Taxonomical studies on Oriental Platygastridae (Hymenoptera: Platygastroidea)

14 species new to science are described, viz. Amblyaspis joenssoni, Euxestonotus sabahensis, Leptacis cheyi (all from Malaysia), L. jani (from Laos), L. maliauensis (from Malaysia), L. ongkudoni (from Malaysia), L. pederseni (from Laos), L. reticulaticeps (from Malaysia), L. solodovnikovi, L. vilhelmseni (both from Laos), Sacespalus viklundi (from Malaysia), Synopeas laosianum (from Laos), S. opaciceps and S. waidii (both from Malaysia). The following species described in Leptacis by Ushakumari, R., Narendran, T.C., 2007. A taxonomic revision of Leptacis Foerster (Hymenoptera: Platygasteridae) of India. Rec. Zool. Surv. India 107, 7–32 are transferred to Synopeas: L. aeros, L. alus, L. asiaticus, L. benazeer, L. diversus, L. manii, L. mustus, L. nuperus and L. scaposus. Synopeas saltaense is a nom. nov. for S. intermedius Buhl, 2005 preoccupied by S. intermedius (Ushakumari, R., 2004. Diversity of Platygaster Latreille (Hymenoptera: Platygastridae) of Kerala. In Rajmohana, K., Narendran, T.C., Perspectives on biosystematics and biodiversity: Prof. T. C. Narendran commemoration volume. Systematic Entomology Research Scholars Association, University of Calicut, Kozhikode, India, pp. 573–591). New locality records for 20 already known Oriental platygastrid species are given.     

Niche partitioning in a sympatric cryptic species complex

Competition theory states that multiple species should not be able to occupy the same niche indefinitely. Morphologically, similar species are expected to be ecologically alike and exhibit little niche differentiation, which makes it difficult to explain the co‐occurrence of cryptic species. Here, we investigated interspecific niche differentiation within a complex of cryptic bumblebee species that co‐occur extensively in the United Kingdom. We compared the interspecific variation along different niche dimensions, to determine how they partition a niche to avoid competitive exclusion. We studied the species B. cryptarum, B. lucorum, and B. magnus at a single location in the northwest of Scotland throughout the flight season. Using mitochondrial DNA for species identification, we investigated differences in phenology, response to weather variables and forage use. We also estimated niche region and niche overlap between different castes of the three species. Our results show varying levels of niche partitioning between the bumblebee species along three niche dimensions. The species had contrasting phenologies: The phenology of B. magnus was delayed relative to the other two species, while B. cryptarum had a relatively extended phenology, with workers and males more common than B. lucorum early and late in the season. We found divergent thermal specialisation: In contrast to B. cryptarum and B. magnus, B. lucorum worker activity was skewed toward warmer, sunnier conditions, leading to interspecific temporal variation. Furthermore, the three species differentially exploited the available forage plants: In particular, unlike the other two species, B. magnus fed predominantly on species of heather. The results suggest that ecological divergence in different niche dimensions and spatio‐temporal heterogeneity in the environment may contribute to the persistence of cryptic species in sympatry. Furthermore, our study suggests that cryptic species provide distinct and unique ecosystem services, demonstrating that morphological similarity does not necessarily equate to ecological equivalence.     

缅甸树萝卜和深裂树萝卜，老挝杜鹃花科植物二新记录种

报道了老挝杜鹃花科树萝卜属2新记录种:缅甸树萝卜(Agapetes megacarpa W. W. Smith)和深裂树萝卜(A. lobbii C. B.Clarke),前者分布于老挝琅勃拉邦省(Luang Prabang Province),后者分布于万象省(Vientiane Province),凭证标本保存在俄罗斯科马洛夫植物研究院标本馆(LE)。提供了每种的凭证标本、分布信息和彩色照片,分别利用繁殖器官性状和营养器官性状编制了老挝3种树萝卜属植物的检索表。     

Hidden Biodiversity in an Ecologically Important Freshwater Amphipod: Differences in Genetic Structure between Two Cryptic Species

Cryptic species, i.e. species that are morphologically hard to distinguish, have been detected repeatedly in various taxa and ecosystems. In order to evaluate the importance of this finding, we have to know in how far cryptic species differ in various aspects of their biology. The amphipod Gammarus fossarum is a key invertebrate in freshwater streams and contains several cryptic species. We examined the population genetic structure, genetic diversity and demographic history of two of them (type A and type B) using microsatellite markers and asked whether they show significant differences. We present results of population genetic analyses based on a total of 37 populations from the headwaters of two major European drainages, Rhine and Rhone. We found that, in both species, genetic diversity was geographically structured among and within drainages. For type A in the Rhine and type B in the Rhone, we detected significant patterns of isolation by distance. The increase of genetic differentiation with geographical distance, however, was much higher in type A than in type B. This result indicates substantial interspecific differences in population history and/or the extent of current gene flow between populations. In the Rhine, type B does not show evidence of isolation by distance, and population differentiation is relatively low across hundreds of kilometres. The majority of these populations also show signatures of recent bottlenecks. These patterns are consistent with a recent expansion of type B into the Rhine drainage. In summary, our results suggest considerable and previously unrecognized interspecific differences in the genetic structure of these cryptic keystone species.     

Geometric morphometric study of geographic and host-related variability in Aceria spp. (Acari: Eriophyoidea) inhabiting Cirsium spp. (Asteraceae)

The russet mite, Aceria anthocoptes (Nalepa), is the only eriophyoid that has been recorded on Cirsium arvense (L.) Scop. It has been noted in several European countries and recently in the USA. In this study we explored the geographic and host-related variability of Aceria spp. inhabiting different Cirsium spp. We applied landmark-based geometric morphometric methods to study morphological variability of three body regions (ventral, coxigenital and prodorsal) of 13 Aceria spp. populations inhabiting five Cirsium spp. in Serbia (Europe) and four Cirsium spp. in Colorado (North America). Analyses of size and shape variation revealed statistically significant differences between Aceria spp. living on European native and North American native Cirsium spp., as well as between A. anthocoptes s.s. inhabiting European C. arvense and North American C. arvense. The coxigenital region was the most informative when considering inter-population shape differences. European Aceria spp. dwelling on Cirsium spp., including A. anthocoptes s.s. from C. arvense, are characterized by higher inter-population size and shape variability than their North American counterparts. This finding supports a Eurasian origin of A. anthocoptes, presumed to consist of a complex of cryptic taxa probably coevolved with host plants in the native environment. Morphological similarity among Aceria spp. inhabiting North American native Cirsium spp. may indicate that speciation of A. anthocoptes started relatively soon after the host shift to plants different from C. arvense in the invaded region.     

Discrimination of Eubazus (Hymenoptera, Braconidae) sibling species using geometric morphometrics analysis of wing venation

Abstract.  Complexes of sibling and cryptic species are encountered frequently in parasitic Hymenoptera. Geometric morphometrics is a useful tool to detect minimal morphological variations, which often are undetectable by traditional morphological studies and even by classical morphometric approaches. We applied geometric morphometrics to wing venation to assess a complex case of sibling species in the genus Eubazus (Hymenoptera, Braconidae), parasitoids of conifer bark weevils of the genus Pissodes (Coleoptera, Curculionidae). The results and methods were compared with previous taxonomic studies on the same species, involving classical multivariate morphometrics, isoenzyme analyses, cross-mating experiments and biological observations. Geometric morphometrics confirmed the previous division into four distinct species. However, this approach enabled the four species to be separated simultaneously, with a reliability of 98.6% for well-classified females and 93.1% for males. A similar result in previous studies was obtained only by combining isoenzyme analyses and several canonical variate analyses, including many morphometric characters. Furthermore, measurements of wing venation were less time-consuming, more reliable and required less prior knowledge of braconid taxonomy than the measurements needed for the classical morphometrics methods. Geometric morphometrics was used also to test the effect of host species on wing shape. Several female populations of Eubazus semirugosus originating from three different Pissodes spp. were compared. Significant differences were found in wing shape between conspecific Eubazus from different host species. The results are discussed in relation to reproductive isolation and genetic flow between the four species.