The wheat curl mite Aceria tosichella (Acari: Eriophyoidea) is a complex of cryptic lineages with divergent host ranges: evidence from molecular and plant bioassay data

Aceria tosichella (the wheat curl mite, WCM) is a global pest of wheat and other cereals, causing losses by direct damage, as well as the transmission of plant viruses. The mite is considered to have an unusually wide host range for an eriophyoid species. The present study tested the commonly held assumption that WCM is a single, highly polyphagous species by assessing the host range of genetically distinct lineages of WCM occurring in Poland on different host plants. Genotyping was performed by analyzing nucleotide sequence data from fragments of the mitochondrial cytochrome c oxidase subunit I (COI) and the nuclear D2 region of 28S rDNA. Mean between‐lineage distance estimated using COI data was found to be one order of magnitude greater than the within‐clade lineage and, in some cases, comparable to distances between WCM lineages and a congeneric outgroup species. Host acceptance was tested by quantifying population growth for different WCM mitochondrial (mt)DNA lineages when transferred from source host plants to test plants. These experiments revealed significant differences in host colonization ability between mtDNA lineages, ranging from highly polyphagous to more host‐specific. The present study reveals that WCM is composed of several discrete genetic lineages with divergent host‐acceptance and specificity traits. Genetic variation for host acceptance within A. tosichella s.l. may act as a reproductive barrier between these lineages, most of which had narrow host ranges. Two lineages appear to have high pest potential on cereals, whereas several others appear to specialize on wild grass species. We conclude that WCM is not a homogeneous species comprising polyphagous panmictic populations rather it is a complex of genetically distinct lineages with variable host ranges and therefore variable pest potential. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 165–180.     

Phylogenetic analyses reveal extensive cryptic speciation and host specialization in an economically important mite taxon

The wheat curl mite (WCM) is a major pest in cereal crops around the world and the vector of at least four known pathogens capable of reducing yields in crops such as wheat, corn, barley, oats, millet and rye. Current taxonomy recognizes WCM as a single species, Aceria tosichella; however, recent genetic, physiological and ecological studies have shown that WCM is likely to be a species complex. In this study we assessed genetic variation and phylogenetic relationships among WCM from four continents and a wide range of host plants using DNA sequence data from one mitochondrial gene, one nuclear gene and a single nuclear intergenic spacer region. Phylogenetic analyses revealed 11 unique mite lineages associated with specific plant hosts including wheat and barley. Host associations were consistent across continents, often with a single haplotype dominating a host plant regardless of geographic origin. The genetic and ecological differences identified in this study support the notion that WCM is a species complex in need of major taxonomic revision. These findings have implications for control of WCM globally, particularly within the context of identifying plants that form ‘green bridge’ refuges, assessing disease transmission risk, and identifying resistance in cereal genotypes to WCM and associated pathogens.     

Wheat curl mite and dry bulb mite: untangling a taxonomic conundrum through a multidisciplinary approach

Two economically important eriophyoid mites, Aceria tosichella (wheat curl mite; WCM) and Aceria tulipae (dry bulb mite; DBM), were frequently confounded in the world literature until the late 20th Century. Their morphological similarity and ambiguous data from plant‐transfer and virus‐transmission trials contributed to this confusion. Until recently, there was a general lack of knowledge about the existence of species complexes and it was not possible to accurately genotype tested mites. In the present study, two WCM genotypes of divergent host specificity (MT‐1 and MT‐2) and one DBM genotype were tested for the acceptance of Poaceae, Amarylidaceae, and Liliaceae species that were reported or suspected as hosts of WCM or DBM. The MT‐1 lineage colonized all tested plants. Onion‐ and garlic‐associated DBM populations did not colonize tulip and wild garlic, suggesting that host‐acceptance variability exists within A. tulipae s.l. Morphometric analysis did not discriminate closely‐related MT‐1 and MT‐2 genotypes but completely separated both WCM genotypes from DBM based on the larger overall body size of the latter. Three morphological traits combined to discriminate between the DBM and MT‐1 genotypes, both of which can infest Amarylidaceae bulbs. In total, these combined DNA sequence, host‐acceptance, morphometrical results unambiguously separated two WCM and one DBM genotypes. Similar studies on additional lineages of both WCM and DBM should ultimately dispel previous taxonomic confusion between these two species. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 421–436.     

Population genetics of the wheat curl mite (Aceria tosichella Keifer) in Australia: implications for the management of wheat pathogens

The wheat curl mite (WCM), Aceria tosichella Keifer, is a polyphagous eriophyoid mite and the primary vector of wheat streak mosaic virus (WSMV) and five other viral pathogens in cereals. Previous research using molecular markers and a series of laboratory experiments found A. tosichella in Australia to consist of two genetically distinct lineages, which have broad overlapping distributions and differ in their ability to transmit WSMV under controlled conditions. This pattern of transmission also appears to be apparent in the field, whereby a strong association between WSMV detection and a single WCM lineage has been detected. In this study, we conduct a population genetic analysis and provide information on the genetic structure of the Australian viruliferous WCM lineage. We assessed genetic differentiation of 16 WCM populations using nine microsatellite markers. Strong evidence for extensive gene flow and low genetic structuring throughout the Australian wheatbelt was evident, with an exception for Western Australian and far north Queensland populations that appear to be genetically isolated. The data also indicate genetic patterns consistent with an arrhenotokous parthenogenetic mode of reproduction. Implications of these findings are discussed with reference to the management of WCM and associated cereal pathogens in Australia and overseas.     

The distribution of wheat curl mite (Aceria tosichella) lineages in Australia and their potential to transmit wheat streak mosaic virus

The wheat curl mite (WCM), Aceria tosichella , is an eriophyid pest of cereals, and the vector responsible for the transmission of wheat streak mosaic virus (WSMV). In a previous study, the taxonomic status of A. tosichella in Australia was assessed using molecular markers. A. tosichella was shown to consist of two genetically distinct lineages likely to represent different species. Here we show that both lineages occupy similar distributions, occurring throughout the entire Australian wheat belt, and that the lineages are often found in sympatry. CLIMEX analysis suggests that tolerance to heat and desiccation limit the distribution of A. tosichella . In the laboratory, only one WCM lineage transmitted WSMV virus under controlled conditions. These results have implications for the management of WCM and WSMV within Australia.     

Wheat curl mite, Aceria tosichella, and transmitted viruses: an expanding pest complex affecting cereal crops

The wheat curl mite (WCM), Aceria tosichella, and the plant viruses it transmits represent an invasive mite-virus complex that has affected cereal crops worldwide. The main damage caused by WCM comes from its ability to transmit and spread multiple damaging viruses to cereal crops, with Wheat streak mosaic virus (WSMV) and Wheat mosaic virus (WMoV) being the most important. Although WCM and transmitted viruses have been of concern to cereal growers and researchers for at least six decades, they continue to represent a challenge. In older affected areas, for example in North America, this mite-virus complex still has significant economic impact. In Australia and South America, where this problem has only emerged in the last decade, it represents a new threat to winter cereal production. The difficulties encountered in making progress towards managing WCM and its transmitted viruses stem from the complexity of the pathosystem. The most effective methods for minimizing losses from WCM transmitted viruses in cereal crops have previously focused on cultural and plant resistance methods. This paper brings together information on biological and ecological aspects of WCM, including its taxonomic status, occurrence, host plant range, damage symptoms and economic impact. Information about the main viruses transmitted by WCM is also included and the epidemiological relationships involved in this vectored complex of viruses are also addressed. Management strategies that have been directed at this mite-virus complex are presented, including plant resistance, its history, difficulties and advances. Current research perspectives to address this invasive mite-virus complex and minimize cereal crop losses worldwide are also discussed.     

Distribution and influence of grazing on wheat curl mites (Aceria tosichella Keifer) within a wheat field

Wheat streak mosaic virus (WSMV) is a serious disease of wheat and is primarily transmitted from infected to healthy plants by the wheat curl mite, Aceria tosichella Keifer. Although wheat is the primary plant host of A. tosichella, wheat curl mites have been recorded on more than 60 different plant hosts; this broad host range allows mites to survive outside the wheat‐growing season by providing a ‘green bridge’. Despite the fact that A. tosichella can only crawl short distances, the mites can disperse via wind and thus have the capacity to readily infest wheat crops from neighbouring refuges. In this study, we undertook field trials to investigate the temporal movement of A. tosichella, as well as the importance of wind and livestock grazing on mite dispersal late in the cropping season. We demonstrate there is a window in spring when A. tosichella undergo significant movement in south‐eastern Australia, and this is likely related to the development stage of wheat plants, and may also be influenced by wind direction. We found that grazing wheat crops reduced mite numbers, suggesting that any increase in WSMV issues in ‘grain and graze’ crops is likely due to the longer season wheat varieties used in these systems rather than the direct effects of grazing. These results emphasize the importance of crop management strategies in the control of A. tosichella.     

Influence of volunteer wheat plant condition on movement of the wheat curl mite, Aceria tosichella, in winter wheat

The wheat curl mite (WCM), Aceria tosichella Keifer, is the vector of wheat streak mosaic virus and high plains virus which cause significant crop loss in winter wheat throughout the western Great Plains. Volunteer wheat emerging before harvest, as a result of severe hail, is the primary source of mites and virus that infect fall-planted winter wheat. Wind-borne movement of the WCM is of key importance in the spread and infection of the virus complex. Significant movement of WCM from wheat has been thought to be closely tied to the senescence or deterioration of the host. Results from field and greenhouse studies indicated that movement from un-vernalized winter wheat was not closely associated with the deterioration of the wheat host. Greenhouse studies showed no correlation between WCM movement and plant condition, but there was a highly significant relationship between WCM movement and mite population on the host plant. Field studies did not demonstrate increased movement associated with deteriorating un-vernalized winter wheat. However, healthier hosts which were able to support a larger population of mites were associated with increased movement. The main influence on the level of mite movement relates to the size of the source population and not the condition of the host plant, but plant condition appears to be a factor in limiting the increase of the WCM population.     

Potential new sources of wheat curl mite resistance in wheat to prevent the spread of yield-reducing pathogens

The wheat curl mite (WCM), Aceria tosichella Keifer (Trombidiformes: Eriophyidae), is a major pest in cropping regions of the world and is recognised as the primary vector of several yield-reducing pathogens, primarily affecting wheat. Management of WCM is complicated due to several aspects of the mite’s biology and ecology; however, commercially viable mite resistant wheat varieties may offer practical long-term management options. Unfortunately, mite populations have adapted to previously identified sources of resistance, highlighting the need for further sources of resistance and the value of stacking different resistances to give greater degrees and longevity of control. In this study we assessed the susceptibility of 42 wheat-derived genotypes to mite population growth using a new experimental method that overcomes methodological limitations of previous studies. Experimental wheat lines included a variety of wheat genotypes, related Triticeae species, wheat-alien chromosome amphiploids, and chromosome addition or substitution lines. From these we identify new promising sources of WCM resistance associated with Thinopyrum intermedium, Th. ponticum and Hordeum marinum chromosomes. More specifically we identify group 1J and 5J chromosomes of the L3 and L5 wheat-Th. intermedium addition lines as new sources of resistance that could be exploited to transfer resistance onto homoeologous wheat chromosomes. This study offers new methods for reliable in situ estimations of mite abundance on cereal plants, and new sources of WCM resistance that may assist management of WCM and associated viruses in wheat.     

Biogeography and diversification of colletid bees (Hymenoptera: Colletidae): emerging patterns from the southern end of the world

Aim The evolutionary history of bees is presumed to extend back in time to the Early Cretaceous. Among all major clades of bees, Colletidae has been a prime example of an ancient group whose Gondwanan origin probably precedes the complete break‐up of Africa, Antarctica, Australia and South America, because modern lineages of this family occur primarily in southern continents. In this paper, we aim to study the temporal and spatial diversification of colletid bees to better understand the processes that have resulted in the present southern disjunctions. Location Southern continents. Methods We assembled a dataset comprising four nuclear genes of a broad sample of Colletidae. We used Bayesian inference analyses to estimate the phylogenetic tree topology and divergence times. Biogeographical relationships were investigated using event‐based analytical methods: a Bayesian approach to dispersal–vicariance analysis, a likelihood‐based dispersal–extinction–cladogenesis model and a Bayesian model. We also used lineage through time analyses to explore the tempo of radiations of Colletidae and their context in the biogeographical history of these bees. Results Initial diversification of Colletidae took place at the Late Cretaceous (≥ 70 Ma). Several (6–14) lineage exchanges between Australia and South America via Antarctica during the Late Cretaceous and Eocene epochs could explain the disjunctions observed between colletid lineages today. All biogeographical methods consistently indicated that there were multiple lineage exchanges between South America and Australia, and these approaches were valuable in exploring the degree of uncertainty inherent in the ancestral reconstructions. Biogeographical and dating results preclude an explanation of Scrapterinae in Africa as a result of vicariance, so one dispersal event is assumed to explain the disjunction in relation to Euryglossinae. The net diversification rate was found to be highest in the recent history of colletid evolution. Main conclusions The biogeography and macroevolutionary history of colletid bees can be explained by a combination of Cenozoic vicariance and palaeoclimatic changes during the Neogene. The austral connection and posterior break‐up of South America, Antarctica and Australia resulted in a pattern of disjunct sister lineages. Increased biome aridification coupled with floristic diversification in the southern continents during the Neogene may have contributed to the high rates of cladogenesis in these bees in the last 25–30 million years.     

Effects of host plant on life‐history traits in the polyphagous spider mite Tetranychus urticae

Studying antagonistic coevolution between host plants and herbivores is particularly relevant for polyphagous species that can experience a great diversity of host plants with a large range of defenses. Here, we performed experimental evolution with the polyphagous spider mite Tetranychus urticae to detect how mites can exploit host plants. We thus compared on a same host the performance of replicated populations from an ancestral one reared for hundreds of generations on cucumber plants that were shifted to either tomato or cucumber plants. We controlled for maternal effects by rearing females from all replicated populations on either tomato or cucumber leaves, crossing this factor with the host plant in a factorial design. About 24 generations after the host shift and for all individual mites, we measured the following fitness components on tomato leaf fragments: survival at all stages, acceptance of the host plant by juvenile and adult mites, longevity, and female fecundity. The host plant on which mite populations had evolved did not affect the performance of the mites, but only affected their sex ratio. Females that lived on tomato plants for circa 24 generations produced a higher proportion of daughters than did females that lived on cucumber plants. In contrast, maternal effects influenced juvenile survival, acceptance of the host plant by adult mites and female fecundity. Independently of the host plant species on which their population had evolved, females reared on the tomato maternal environment produced offspring that survived better on tomato as juveniles, but accepted less this host plant as adults and had a lower fecundity than did females reared on the cucumber maternal environment. We also found that temporal blocks affected mite dispersal and both female longevity and fecundity. Taken together, our results show that the host plant species can affect critical parameters of population dynamics, and most importantly that maternal and environmental conditions can facilitate colonization and exploitation of a novel host in the polyphagous T. urticae, by affecting dispersal behavior (host acceptance) and female fecundity.     

Molecular probes of phylogeny and biogeography in toads of the widespread genus Bufo

Genetic relationships among 25 species of Central and South American Bufo and among representative North, Central, and South American, Asian, and African Bufo were probed, using the quantitative immunological technique of microcomplement fixation (MC'F) which indicated a clear separation of North, Central, and South American lineages of Bufo. The South American lineage likely diverged from the Central and North American lineages in the Eocene; the latter two lineages diverged later, probably in the mid-Oligocene. Some species groups of South American toads, defined on the basis of traditional morphological studies, are genetically quite similar within groups, whereas others are genetically divergent. The amount of albumin evolution does not appear to parallel the amount of karyotypic, morphological, ecological, or behavioral evolution documented. Comparisons suggest that the African lineages separated from the American and Asian lineages in the late Cretaceous, corresponding to the time of the final separation of Gondwanaland, the southern supercontinent including the modern continents of South America, Africa, Australia, Antarctica, and India. The Asian lineages diverged from the lineage giving rise to all of the American species in the early Paleocene.      

Wheat curl mite resistance: interactions of mite feeding with wheat streak mosaic virus infection

The majority of plant viruses are dependent on arthropod vectors for spread between plants. Wheat streak mosaic virus (family Potyviridae, genus Tritimovirus, WSMV) is transmitted by the wheat curl mite, Aceria tosichella Keifer, and this virus and vector cause extensive yield losses in most major wheat (Triticum aestivum L.)-growing regions of the world. Many cultivars in use are susceptible to this vector-virus complex, and yield losses of 10-99% have been documented. wheat curl mite resistance genes have been identified in goat grass, Aegilops tauschii (Coss) Schmal., and transferred to hexaploid wheat, but very few varieties contain effectively wheat curl mite resistance, due to virulent wheat curl mite populations. However, wheat curl mite resistance remains an effective strategy to reduce losses due to WSMV. The goal of our project was to identify the most effective, reproducible, and rapid method for assessing wheat curl mite resistance. We also wanted to determine whether mite resistance is affected by WSMV infection, because the pathogen and pest commonly occur together. Single and group wheat curl mite infestations produced similar amounts of leaf rolling and folding on wheat curl mite-susceptible wheat varieties that were independent of initial wheat curl mite infestation. This finding will allow accurate, efficient, large-scale screening of wheat germplasm for wheat curl mite resistance by infesting plants with sections of wheat leaf tissue containing mixed stages of wheat curl mite. The wheat curl mite-resistant breeding line 'OK05312' displayed antibiosis (reduced wheat curl mite population development). The effect of WSMV infection on wheat curl mite reproduction was genotype-dependent. Mite populations increased on infected wheat curl mite- and WSMV-susceptible plants compared with uninfected plants, but WSMV infection had no significant effect on wheat curl mite populations on resistant plants. OK05312 is a strong source of wheat curl mite resistance for wheat breeding programs.     

Divergent host acceptance behavior suggests host specialization in populations of the polyphagous mite Abacarus hystrix (Acari: Prostigmata: Eriophyidae)

For phytophagous arthropods, host acceptance behavior is a key character responsible for host plant specialization. The grain rust mite, Abacarus hystrix (Nalepa), is an obligately phytophagous, polyphagous eriophyid mite recorded from at least 70 grass species. In this study, the hypothesis that two host populations of this mite (one collected from quackgrass and the other from ryegrass) are highly host-specific was tested using behavioral data. For this purpose, female behavior when exposed to familiar and novel host plants was observed in no-choice cross experiments. Altogether, 13 variables were used to describe mite behavior. Data were subjected to principal component analysis, and host acceptance behavior was subsequently tested with generalized estimating equations (GEE). Distinct variation in female behavior between familiar and novel hosts was observed. Females from neither population accepted novel hosts. This was recorded as significant differences in the occupation of and overall activity on particular plant parts. On their familiar host, females were not active and showed little tendency to move. On novel hosts females were more active and mobile, spending more time walking, running, and climbing on the whole plant surface and showing a tendency to disperse. Other differences in behavior between studied populations were also observed. Thus, the results suggest that mites of these two studied populations (1) differ in their behaviors during plant exploitation and (2) can quickly distinguish between their familiar host and an unfamiliar host used by a conspecific. These findings support the hypothesis of narrow host specialization of ryegrass and quackgrass populations of this highly polyphagous species.     

Decay of similarity of gamasid mite assemblages parasitic on Palaearctic small mammals: geographic distance, host-species composition or environment

Aim The similarity between parasite assemblages should decrease with increasing geographic distance between them, increasing dissimilarity in environmental conditions, and/or increasing dissimilarity of the local host fauna, depending on the dispersal abilities of the parasites and the intimacy of their associations with the host. We tested for a decay in the similarity of gamasid mite assemblages parasitic on small mammals with increasing geographic, ‘environmental’ and ‘host faunal’ (= ‘host’) distances. Location We used data on assemblages of haematophagous gamasid mites (superfamily Dermanyssoidea) parasitic on small mammals (Insectivora, Lagomorpha and Rodentia) from 26 different regions of the northern Palaearctic. Methods Similarity in mite assemblages was investigated at the compound community level across all regions, and at the component community level, across populations of the same host species for each of 11 common host species. Similarity between pairs of mite communities was estimated using both the Jaccard and the Sorensen indices. Environmental distance was estimated as the dissimilarity between locations in a composite measure of climatic variables, and host faunal distance was simply taken as the reciprocal of indices of similarity between the composition of host faunas in different locations. Generalized Linear Models (GLM) and Akaike's Information Criterion were used to select the best model of decay in similarity as a function of geographic, ‘environmental’ and ‘host faunal’ distances. Results Overall, despite slight differences among host species, the similarity in mite assemblages decreased with both increasing ‘environmental’ distance and increasing ‘host faunal’ distance, but was generally unaffected by geographic distance between regions. The similarity of component communities of gamasid mites among host populations was determined mainly by similarity in the physical environment, whereas that of compound communities varied mainly with host‐species composition. Main conclusions Our results indicate that the general decay in community similarity with increasing geographic distances does not apply to assemblages of gamasid mites; it is possible that they can overcome great distances by means of passive dispersal (either by phoresy or wind‐borne), or more likely they occur wherever their hosts are found as a result of tight cospeciation in the past. Mite assemblages on small mammalian hosts seem to be affected mainly by local environmental conditions, and, to a much lesser extent, by the species composition of local host communities.     

Phylogeographic structuring of Plasmodium lineages across the North American range of the house finch (Carpodacus Mexicanus)

The determinants of the geographic distribution of avian hematozoa are poorly understood. Sampling parasites from one avian host species across a wide geographic range is an accepted approach to separate the potential influence of host species distribution from geographic effects not directly related to host species biology. We used polymerase chain reaction to screen samples for hematozoan infection from 490 house finches (Carpodacus mexicanus) collected at 8 sites spanning continental North America. To explore geographic patterns of parasite lineage distributions, we sequenced a portion of the mitochondrial cytochrome b gene of Plasmodium species infecting 77 house finches. We identified 5 distinct Plasmodium haplotypes representing 3 lineages that likely represent 3 species. One lineage was common at all sites where we detected Plasmodium species. The second lineage contained 3 haplotypes that showed phylogeographic structuring on a continent-wide scale, with 1 haplotype common in eastern North America and 2 common in western North America. The third divergent lineage was recovered from 1 individual host. Considered together, the partial phylogeographic structuring of Plasmodium cytochrome b lineages over the range of the house finch suggests that parasite lineage distribution is not solely dependent on host species distribution, and other factors such as arthropod vector competence and distribution may be important.     

Multiple cryptic species with divergent substrate affinities in the Serpula himantioides species complex

Serpula himantioides is a widespread saprotrophic morphospecies mainly colonising coniferous wood in nature, but it appears frequently in buildings as well. From an earlier study, it is known that at least three divergent lineages occur within the S. himantioides species complex. In this study, a broader sample of S. himantioides isolates has been analysed by multi-locus sequencing, including new isolates from Asia, North and South America. Altogether five phylogenetical species (PS1-5) were detected, all recognised across independent gene phylogenies. A new southern South American phylogenetic species (PS1) was found, representing an early diverging lineage within the S. himantioides species complex. The two closely related PS2 and PS3 lineages included isolates from North America only, and PS4 was also dominated by North American isolates. Most of the investigated isolates (76%) clustered into PS5, a lineage that has been found on most continents, including North America. Overall, little phylogeographical structure was found in PS5, indicating frequent and recent long-distance dispersal events within this widespread lineage. Our analyses indicate that South and North America are the centres of divergence for the S.?himantioides species complex. Some of the lineages seem adapted to various substrates, but PS5 is able to decay a wide array of angiosperms and gymnosperms, which may have facilitated the spread of this lineage throughout the world.     

Divergence and gene flow in the globally distributed blue‐winged ducks

The ability to disperse over long distances can result in a high propensity for colonizing new geographic regions, including uninhabited continents, and lead to lineage diversification via allopatric speciation. However, high vagility can also result in gene flow between otherwise allopatric populations, and in some cases, parapatric or divergence‐with‐gene‐flow models might be more applicable to widely distributed lineages. Here, we use five nuclear introns and the mitochondrial control region along with Bayesian models of isolation with migration to examine divergence, gene flow, and phylogenetic relationships within a cosmopolitan lineage comprising six species, the blue‐winged ducks (genus Anas), which inhabit all continents except Antarctica. We found two primary sub‐lineages, the globally‐distributed shoveler group and the New World blue‐winged/cinnamon teal group. The blue‐winged/cinnamon sub‐lineage is composed of sister taxa from North America and South America, and taxa with parapatric distributions are characterized by low to moderate levels of gene flow. In contrast, our data support strict allopatry for most comparisons within the shovelers. However, we found evidence of gene flow from the migratory, Holarctic northern shoveler A. clypeata and the more sedentary, African Cape shoveler A. smithii into the Australasian shoveler A. rhynchotis, although we could not reject strict allopatry. Given the diverse mechanisms of speciation within this complex, the shovelers and blue‐winged/cinnamon teals can serve as an effective model system for examining how the genome diverges under different evolutionary processes and how genetic variation is partitioned among highly dispersive taxa.     

The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae)

Abstract. To reconstruct the phylogeny of the Sericini and their systematic position among the scarabaeid beetles, cladistic analyses were performed using 107 morphological characters from the adults and larvae of forty‐nine extant scarabaeid genera. Taxa represent most ‘traditional’ subfamilies of coprophagous and phytophagous Scarabaeidae, with emphasis on the Sericini and other melolonthine lineages. Several poorly studied exoskeletal features have been examined, including the elytral base, posterior wing venation, mouth parts, endosternites, coxal articulation, and genitalia. The results of the analysis strongly support the monophyly of the ‘orphnine group’ + ‘melolonthine group’ including phytophagous scarabs such as Dynastinae, Hopliinae, Melolonthinae, Rutelinae, and Cetoniinae. This clade was identified as the sister group to the ‘dung beetle line’ represented by Aphodius + Copris. The ‘melolonthine group’ is comprised in the strict consensus tree by two major clades and two minor lineages, with the included taxa of Euchirinae, Rutelinae, and Dynastinae nested together in one of the major clades (‘melolonthine group I’). Melolonthini, Cetoniinae, and Rutelinae are strongly supported, whereas Melolonthinae and Pachydemini appear to be paraphyletic. Sericini + Ablaberini were identified to be sister taxa nested within the second major melolonthine clade (‘melolonthine group II’). As this clade is distributed primarily in the southern continents, one could assume that Sericini + Ablaberini are derived from a southern lineage. Plausibly, ancestors of Sericini + Ablaberini and Athlia were separated by a vicariance event, such as the separation of the African plate from the rest of Gondwana, whereas Sericini and Ablaberini probably diversified during the early Tertiary, with dispersal of some basal Sericini to South America.