The phylogeny of a mutualism: evolution and coadaptation between Trollius and its seed-parasitic pollinators

Interactions between seed-parasitic pollinators and their hosts provide useful model systems for the analysis of evolution of mutualism and potential coevolution between plants and insects. Here I present the systematics, pollination ecology and evolution of one of these interactions. I have documented and analysed the phylogenetic and geographic associations between Trollius (Ranunculaceae: 18 spp.) and Chiastocheta (Diptera: Anthomyiidae; 17 spp.), a host-specific genus of seed-parasitic flies that pollinate their host plants to varying extent. Their interactions are usually facultative mutualisms, but in the specialized T. europaeus three fly species are obligate mutualists and a fourth species is an antagonist. The distribution patterns of fly species among Trollius species suggest that the flies evolved in associations with five highly derived Trollius species, and secondarily colonized four more primitive taxa in the parts of their ranges that overlapped with primary hosts. In general, host specificity is maintained primarily through allopatry, with colonization occurring in regions of overlap between parapatric taxa. Fly speciation has occurred in allopatry, both within and among host taxa. Cospeciation is not evident, but convergent evolution in Trollius flowers of several traits, viz. orange sepals, elongated staminodia and increased carpel number per flower, may be the result of mutualism with Chiastocheta.     

Recent findings of Phlebotomus neglectus Tonnoir, 1921 in Italy and its western limit of distribution

During the sand fly season of 2001, sand flies were collected in two different regions of Italy to investigate the northern distribution of Phlebotomus neglectus. The study areas were two provinces of northern Italy, Ivrea and Verona, in Piedmont and Veneto regions respectively. Sticky traps and CDC miniature light traps were used both inside and outside domestic and peri-domestic environments. Sand flies were also collected in domestic resting sites with mouth and electric aspirators. Of 1,295 sand fly specimens collected, 855 were from the Veneto region and 440 were from the Piedmont region. Five species of the genera Phlebotomus and Sergentomyia were identified. S. minuta (66.3%) was the most abundant sand fly followed by P. perniciosus (22.2%), P. neglectus (10.3%) and P. papatasi (1.2%). Only one specimen of P. mascittii was caught in Piedmont region. P. neglectus was found in 9 out of 15 (60%) of the sites where sand flies were found with a prevalence of 1.5% to 45.8%; the highest value was recorded in Piedmont region. P. neglectus was mostly associated with peridomestic and domestic sites. The results confirm previous observations on the presence of P. neglectus in northern Italy and suggest that this species is probably widespread in the area. From a review of all previous published and unpublished records of P. neglectus, it appears that this species is present in only two areas of the country, namely in the regions of Friuli Venezia Giulia, Veneto and Piedmont in northern Italy and the regions of Calabria, Apulia and Sicily in southern Italy. The apparent absence of P. neglectus in Central Italy is discussed and the resulting settlement through two probable migration routes from the East is analysed.     

Host range and genetic strains of leafminer flies (Diptera: Agromyzidae) in eastern Brazil reveal a new divergent clade of Liriomyza sativae

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Nuisance flies on Australian cattle feedlots: immature populations

Species composition, seasonality and distribution of immature fly populations on a southern Queensland feedlot during 2001-2003 were determined. Similar data were collected on feedlots in central New South Wales and central Queensland. The fly species recovered in the highest numbers were Musca domestica L. (Diptera: Muscidae), Stomoxys calcitrans L. (Diptera: Muscidae) and Physiphora clausa Macquart (Diptera: Ulidiidae). Houseflies were the dominant species at all feedlots. Houseflies preferred the warmer months from October to June, but stable flies preferred the cooler months and peaked in spring (September-November) and autumn (March-May). Larval abundance ratings recorded in the feedlot and numbers of larvae extracted in the laboratory from corresponding samples followed similar trends. Larvae of M. domestica were most abundant in the hospital and induction area and least abundant in horse stables and yards. Pupae of M. domestica were abundant in the hospital and induction area and drains, but least abundant in horse stables and yards. Larvae of S. calcitrans were most abundant in drains and least abundant in horse stables and yards. Pupae of S. calcitrans were most numerous in drains and least numerous in old cattle pens. Feedlot design and management had little effect on fly reduction.     

Barcoding Queensland Fruit Flies (Bactrocera tryoni): impediments and improvements

Identification of adult fruit flies primarily involves microscopic examination of diagnostic morphological characters, while immature stages, such as larvae, can be more problematic. One of the Australia’s most serious horticultural pests, the Queensland Fruit Fly (Bactrocera tryoni: Tephritidae), is of particular biosecurity/quarantine concern as the immature life stages occur within food produce and can be difficult to identify using morphological characteristics. DNA barcoding of the mitochondrial Cytochrome Oxidase I (COI) gene could be employed to increase the accuracy of fruit fly species identifications. In our study, we tested the utility of standard DNA barcoding techniques and found them to be problematic for Queensland Fruit Flies, which (i) possess a nuclear copy (a numt pseudogene) of the barcoding region of COI that can be co‐amplified; and (ii) as in previous COI phylogenetic analyses closely related B. tryoni complex species appear polyphyletic. We found that the presence of a large deletion in the numt copy of COI allowed an alternative primer to be designed to only amplify the mitochondrial COI locus in tephritid fruit flies. Comparisons of alternative commonly utilized mitochondrial genes, Cytochrome Oxidase II and Cytochrome b, revealed a similar level of variation to COI; however, COI is the most informative for DNA barcoding, given the large number of sequences from other tephritid fruit fly species available for comparison. Adopting DNA barcoding for the identification of problematic fly specimens provides a powerful tool to distinguish serious quarantine fruit fly pests (Tephritidae) from endemic fly species of lesser concern.     

Nested distributions of bat flies (Diptera: Streblidae) on Neotropical bats: artifact and specificity in host-parasite studies

We examined the structure of ectoparasitic bat fly infestations on 31 well‐sampled bat species, representing 4 Neotropical families. Sample sizes varied from 22 to 1057 bats per species, and bat species were infested by 4 to 27 bat fly species. Individual bats supported smaller infracommunities (the set of parasites co‐occurring on an individual host), ranging from 1 to 5 fly species in size, and no bat species had more than 6 bat fly species characteristically associated with it (its primary fly species). Nestedness analyses used system temperature (BINMATNEST algorithm) because it is particularly well‐suited for analysis of interaction networks, where parasite records may be nested among hosts and host individuals simultaneously nested among parasites. Most species exhibited very low system temperatures (mean 3.14°; range 0.14–12.28°). Simulations showed that nested structure for all 31 species was significantly stronger than simulated values under 2 of the 3 null hypotheses, and about half the species were also nested under the more stringent conditions of the third null hypothesis. Yet this structure disappears when analyses are restricted to “primary” associations of fly species (flies on their customary host species), which exclude records thought to be atypical, transient, or potential contaminants. Despite comprising a small fraction of total parasite records, such anomalies represent a considerable part of the statistical state‐space, offering the illusion of significant ecological structure. Only well understood and well documented systems can make distinctions between primary and other occurrence records. Generally, nestedness appears best developed in host‐parasite systems where infestations are long‐term and accumulate over time. Dynamic, short‐term infestations by highly mobile parasites like bat flies may appear to be nested, but such structure is better understood in terms of host specificity and accidental occurrences than in terms of prevalence, persistence, or hierarchical niche relations of the flies.     

Ecological studies of Eastern Australian fruit flies (Diptera: Tephritidae) in their endemic habitat

Fortnightly fruit fly captures for a 2-year period at Cooloola (south-east Queensland) contained 11 species. Three species, viz., D. tryoni, D. neohumeralis and D. endiandrae predominated. The peak trap catches of 7 species corresponded with the peak fruiting times of their major hosts. There was no direct relationship between temperature and rainfall and the variations in population numbers. The host plants of some species do not grow in the Cooloola area and there is evidence that large numbers of flies migrate into the region from other breeding areas up to 100 km away. Pockets of tropical rainforest such as Cooloola could be important adult fruit fly feeding areas even in the absence of larval host plants.     

贵州兴义实蝇类昆虫多样性特征

【目的】掌握贵州兴义实蝇的种类组成和不同生境中的发生规律及多样性变化。【方法】2010-2011年,在贵州兴义 5个不同生境的监测点,采用性诱剂、食物引诱剂及被害瓜果培养法对实蝇类昆虫种类和数量进行了系统调查和监测,并对监测数据进行分析。【结果】贵州兴义实蝇优势种包括桔小实蝇 Bactrocera dorsalis (Hendel)、瓜实蝇 B. cucurbitae (Coquillett)、南瓜实蝇 B. tau (Walker)及具条实蝇 B. scutellata (Hendel),且不同年份各监测点实蝇优势种及相似性系数(0.33~0.89)存在差异;但其年种群动态相似。聚类分析结果显示,两个农村生境果园中实蝇种群关系最近,城市混合果园与其他生境果园中实蝇种群关系最远;各监测点的多样性指数在不同年份也不同,监测的2年间 Simpson 指数、Shannon 指数和均匀度指数分别在0.11~0.68、0.28~1.25和0.18~0.62之间。【结论】贵州兴义实蝇种类丰富,种群发生规律与气候、寄主、生态位竞争等因素有关;相近生境混合果园诱捕到的实蝇种数和数量大于单一果园;离城区越近,实蝇发生期越早,数量越大。研究结果为了解该地区实蝇种类分布以及预防实蝇危害提供了科学依据。     

北京地区7种常见嗜尸性蝇类的COI基因序列分析及DNA条形码的建立

嗜尸性蝇类在命案死亡时间和现场推断方面有着十分重要的应用, 而DNA 条形编码技术能摆脱对虫卵和幼虫的饲养以及后续物种鉴定方面专业知识的依赖, 有助于实现现场采集蝇类样本的快速鉴定。本研究采集了北京地区7个嗜尸性蝇类优势种共77个个体的样本, 测定了所有个体线粒体DNA 上细胞色素C氧化酶亚基Ⅰ（COI）基因1 120 bp的序列。基于序列的系统发生分析显示, 同一物种不同个体的序列均以高达99%的支持值聚集在一起。序列间的分歧统计表明这些蝇类在物种内的个体分歧不超过1%, 而不同物种间的净分歧均超过7.74%, 最高可达14.85%。滑动窗口分析表明, 在整个序列区段种间差异位点存在较平均的分布。通过测定COI基因的序列, 建立了北京地区7个嗜尸性蝇类优势种的DNA条形码, 据此实现了对这些物种准确、快速、简单的区分和鉴定, 同时也为后续应用于物种鉴定的种属特异性位点之筛选提供了基础数据。     

Sand fly vectors (Diptera, Psychodidae) of American visceral leishmaniasis areas in the Atlantic Forest, State of Espírito Santo, southeastern Brazil

The aim of this study was to evaluate the sand fly fauna of American visceral leishmaniasis (AVL) endemic areas within the Central Atlantic Forest Biodiversity Corridor, State of Espírito Santo, southeastern Brazil. The sand fly captures were performed between January, 1989 and December, 2003 in localities where autochthonous cases of AVL were recorded, as well as in their boundary areas. Sand flies were collected from surrounding houses and domestic animal shelters using two to five CDC automatic light traps, and manual captures were also performed using mouth aspirators in one illuminated Shannon trap during the first four hours of the night. We used cladistic analysis to determine the geographic relationships among the collected sand fly species as well as the index species for the occurrence of other sand flies. A total of 62,469 sand flies belonging to 17 species and eight genera was collected in 164 localities from nine municipalities with AVL records. The richness (S=17) and diversity (H=0.971) of sand flies were lower than in conservation areas and similar to modified environments in the Atlantic Forest of Espírito Santo. Lutzomyia longipalpis was identified in 79 localities. The cladistic analysis identified Evandromyia lenti as the index species for Lutzomyia longipalpis. The latter seems to be the main vector of AVL in the Central Atlantic Forest Biodiversity Corridor due to its high abundance and distribution matching the disease occurrence. Therefore, Evandromyia lenti may be used as an index species for the occurrence of Lutzomyia longipalpis.     

Spatial distribution of sand fly species (Psychodidae: Phlebtominae), ecological niche,and climatic regionalization in zoonotic foci of cutaneous leishmaniasis,southwest of Iran

Cutaneous leishmaniasis (CL) is a complex vector‐borne disease caused by Leishmania parasites that are transmitted by the bite of several species of infected female phlebotomine sand flies. Monthly factor analysis of climatic variables indicated fundamental variables. Principal component‐based regionalization was used for recognition of climatic zones using a clustering integrated method that identified five climatic zones based on factor analysis. To investigate spatial distribution of the sand fly species, the kriging method was used as an advanced geostatistical procedure in the ArcGIS modeling system that is beneficial to design measurement plans and to predict the transmission cycle in various regions of Khuzestan province, southwest of Iran. However, more than an 80% probability of P. papatasi was observed in rainy and temperate bio‐climatic zones with a high potential of CL transmission. Finding P. sergenti revealed the probability of transmission and distribution patterns of a non‐native vector of CL in related zones. These findings could be used as models indicating climatic zones and environmental variables connected to sand fly presence and vector distribution. Furthermore, this information is appropriate for future research efforts into the ecology of Phlebotomine sand flies and for the prevention of CL vector transmission as a public health priority.     

Population structure and colonization history of the olive fly, Bactrocera oleae (Diptera, Tephritidae)

The olive fly, Bactrocera oleae, is the major pest of olives in most commercial olive-growing regions worldwide. The species is abundant in the Mediterranean basin and has been introduced recently into California and Mexico, creating problems for quarantine protection and international trade. Here, we use nuclear microsatellite markers and mitochondrial sequences to examine the history of olive fly range expansion and colonization. Sampled populations span the current distribution of the olive fly worldwide, including South and Central Africa, Pakistan, Mediterranean Europe and Middle East, California, and Mexico. The Pakistani populations appear to be genetically well differentiated from the remaining populations, though rooting the origins of the species is problematic. Genetic similarity and assignment tests cluster the remaining populations into two genetic groups--Africa and a group including the Mediterranean basin and the American region. That Africa, and not the Mediterranean, is the origin of flies infesting cultivated olive is supported by the significantly greater genetic diversity at microsatellite loci in Africa relative to the Mediterranean area. The results also indicate that the recent invasion of olive flies in the American region most likely originated from the Mediterranean area.     

Species richness and rarity of crane flies (Diptera, Tipuloidea) in a boreal mire

Species richness and abundance are central in biodiversity inventories and in measuring the structure of communities. Neglecting the assessment of sampling efficiency may lead to spurious estimates of species richness and conservation value. Our aim was to examine species richness, sampling effectiveness, species-abundance distribution (SAD) and rarity of a boreal, mire-dwelling crane fly (Diptera, Tipuloidea) assemblage in western Finland. 12 Malaise traps dispersed in 4 subplots and standardized sweep net samples were used to collect adult flies from the mire. A total of 23 species and 1,569 specimens were identified. In general all species richness estimators were highly correlated and indicated rather good sampling effort. Sample completeness, expressed as percentage of observed richness divided by estimated richness, was higher for mire-dwellers (mean 75 %) than for all species (mean 63 %). Crane fly assemblages of subplots and combined data fitted best with log-series SAD. Species spatial distribution was positively correlated with average abundance. In other words, the most abundant species occurred in the most of Malaise traps. Seven mire-dwelling species greatly outnumbered (94 % of the collected specimens) all other members in the assemblage, and only one observed species was rare by several definitions (local abundance, extent of occurrence in Finland and area of occupancy). Although the studied assemblage was characterized by commonness, five of the species have threatened status in Europe south of Finland. Separate species richness estimation of all species (vagrants and occasional species included) and focal species (here mire-dwellers) is supported if ecological information is available on the taxonomic group being studied.     

Bat flies on phyllostomid hosts in the Cerrado region: component community, prevalence and intensity of parasitism

Streblidae flies are specialised parasites of bat hosts, mainly phyllostomids. There is a high richness of streblids in the savannah-like Cerrado region; however, there is little quantitative data available in parasitological indices. Here, we describe the component community, prevalence and intensity of a streblid infestation on a phyllostomid bat assemblage in Serra da Bodoquena, a Cerrado region in Southwest Brazil. We conducted surveys by capturing and inspecting bat hosts during the seven-month period between October 2004-December 2005. All the ectoparasites found on the bats were collected in the field and then counted and identified in the laboratory. We captured 327 bats belonging to 13 species, of which eight species were parasitized by 17 species of streblids. Carollia perspicillata and Glossophaga soricina were infested with seven streblid species, whereas the other bat species were infested with four or fewer streblid species. Megistopoda proxima and Aspidoptera falcata flies were found on Sturnira lilium, and Trichobius joblingi was the most prevalent fly on C. perspicillata. Megistopoda aranea and Aspidoptera phyllostomatis were highly prevalent and had a high intensity of infestation on Artibeus planirostris. Overall comparisons of the available data suggest that the component communities of streblids vary more between the Cerrado and Atlantic Forest phytogeographical regions than between localities within the same phytogeographical region.     

Desolation comes from the sky: Invasive Hymenoptera species as prey of Chilean giant robber flies (Diptera: Asilidae) through field observations and citizen science

Robber flies (Asilidae) are the main predatory fly family feeding on beetles, butterflies, other flies for true flies, and even spiders; however, Hymenoptera is the most common prey. Invasive Hymenoptera species are common in central and southern Chile; however, few predators of these are known. The hunting behavior and prey of Chilean robber fly species are also poorly known. The aim of this study is to provide the first hunting behavior records of five Chilean giant robber fly species on invasive Hymenoptera. In addition, an updated distribution of these species is provided. Records of hunting behavior were based on fieldwork collections and citizen science observations. The historical distribution was compared with citizen science observations using chi-square analyzes. Twelve predation events were recorded. Obelophorus terebratus was the most common predator. Bombus terrestris was the invasive Hymenoptera most preyed upon. Both the extension of occurrence of Lycomya germainii as Obelophorus species showed changes in his distribution. Only O. landbecki shown changes in area of occupancy. Citizen science is playing a key role in the knowledge of biological interactions and distribution of endemic and native Chilean robber fly species.     

云南主要有害实蝇种类及区划

采用诱剂诱捕辅以受害瓜果调查的方法,系统梳理了云南有害实蝇类群,确定主要有害种类25种。基于云南自然地理特征,采用聚类分析法深入探讨了云南实蝇区系,提出云南实蝇地理区划可分为德宏、版纳盆谷区,南部边缘中低山宽谷区,滇西南山原峡谷区,中东部高原、岩溶山原区以及北部高原山地区等5大区域。研究发现,受热量条件制约,云南主要有害实蝇类群呈南北向梯度递减;受纵向岭谷区"通道-阻隔"综合作用影响,云南西南部河谷地区是实蝇危害较为严重的区域。首次系统揭示了云南主要有害实蝇的地理分布规律。     

The prevalence of Stephanofilaria sp. in buffalo fly, Haematobia irritans exigua, in Central Queensland

Abstract  Buffalo fly ( Haematobia irritans exigua ) infestations of cattle are associated with characteristic lesions, the initial cause of which has been attributed to a filarial nematode of the genus Stephanofilaria , for which the fly acts as a vector. Survey work in the 1980s estimated the prevalence of microfilaria in female buffalo fly in Queensland at 2.91%. Since then no information has been published and the current prevalence of microfilarial infection in buffalo fly is not known. Buffalo fly were collected from four geographically distinct sites in Central Queensland in mid-summer 2004 and were dissected to estimate Stephanofilaria sp. infection rates . Larval stages of the nematodes were recovered from female flies from all four sites and the percentage of female flies from which nematodes were recovered ranged from 29% to 57%. The average number of larvae recovered from infected female flies ranged from 1.25 to 1.75. Whereas no infected male flies were found from Sites 2–4, larvae were recovered from 43% of male flies collected at Site 1. This high prevalence of filarial infection in buffalo flies implies a correspondingly high level of transmission to cattle in Central Queensland.     

DNA microsatellite analysis of naturally occurring colour intermediates between Bactrocera tryoni (Froggatt) and Bactrocera neohumeralis (Hardy) (Diptera: Tephritidae)

Abstract  The sympatric tephritid fruit flies Bactrocera tryoni (Froggatt) (Queensland fruit fly) and B. neohumeralis (Hardy) differ in time of mating and for the colour of the humeral callus ('shoulder pad'), which is typically entirely yellow in B. tryoni and typically entirely brown in B. neohumeralis . Field collections in sympatric regions usually include at least 1% of individuals whose humeral calli show mixed patches of yellow and brown ('intermediates'). Over 40 years, a number of studies have debated the possibility that these intermediates are interspecific hybrids. In the present study, we have used microsatellites to show that few if any of these intermediates are hybrids. Instead, most variation humeral callus appears to be confined to one species, B. tryoni . We discuss these results in the context of geneflow between the two species and suggest directions for future research.     

远东地区影响针叶树天然更新的球果种实害虫———球果花蝇 (双翅目 :花蝇科)( 英文)

球果花蝇是远东地区的重要球果种实害虫。尤其是在我国东北地区 ,由于球果花蝇的危害已严重影响了落叶松种子产量 ,进而影响到落叶松的造林与更新。本文系统地总结了近些年危害落叶松、云杉、冷杉种实的球果花蝇的研究进展及存在的问题。文中还涵括了作者的一些原始研究数据和尚未发表的新结果。     

Comparative epidemiology of Dipofilaria roemeri infection in two regions of Queensland

The occurrence of Dirofilaria roemeri in definitive and intermediate hosts in two separate areas in southeast Queensland, is compared. The presence of D. roemeri in a macropod species in a defined region is determined by both the geographic distribution and the biting behaviour of those species of tabanid fly which are capable of acting as intermediate host. The seasonal succession and seasonal abundance of 4 tabanid species at Durakai and 8 tabanid species at Allan are responsible for a seasonal pattern of transmission of this filarioid. Moderate to low levels of transmission occur from November through March. Maximum transmission occurs in April associated with the peak in population of Dasybasis hebes, the most important tabanid involved in transmission of D. roemeri in both areas. Findings indicate that the wallaroo is the source of infection for tabanid flies and acts as the reservoir of infection for grey kangaroos and red-necked wallabies, in southeast Queensland. Transmission of D. roemeri is discussed in the light of behavioural interactions between definitive and intermediate hosts.