中国云南部分小兽体表吸虱昆虫群落相似性及群落分类研究(英文)

应用系统聚类分析方法对云南省境内24种主要小型哺乳动物(小兽)体表吸虱昆虫群落相似性及群落分类进行了研究。研究中将每一种小兽体表的所有吸虱昆虫定义为一个相应的吸虱群落单位。结果表明,小兽体表吸虱群落结构简单,物种多样性很低。多数小兽有固定的吸虱种类寄生,其吸虱的宿主特异性高。在动物分类上隶属同一个属的小兽,其体表吸虱群落相似程度高,在系统聚类分析中大多被归为一类。在动物分类上近缘的小兽,其体表吸虱群落相似,优势虱种相同或相似,此情形尤其表现在鼠属、白腹鼠属、姬鼠属和绒鼠属。吸虱群落相似性大小与相应小兽宿主在动物分类地位上的近缘性高低呈现高度一致。从生态学角度来看,吸虱昆虫与其所寄生的小兽宿主动物之间存在密切的协同进化关系。     

SIMILARITY COMPARISON AND CLASSIFICATION OF SUCKING LOUSE COMMUNITIES ON SOME SMALL MAMMALS IN YUNNAN, CHINA

Abstract  The similarity and classification of sucking louse communities on 24 species of small mammals were studied in Yunnan Province, China, through a hierarchical cluster analysis. All the louse species on the body surface of a certain species of small mammals are regarded as a louse community unit. The results reveal that the community structure of sucking lice on small mammals is simple with low species diversity. Most small mammals usually have certain louse species on their body surface; there exists a high degree of host specificity. Most louse communities on the same genus of small mammals show a high similarity and are classified into the same group based on hierarchical cluster analysis. When the hosts have a close affinity in taxonomy, the louse communities on their body surface would tend to be similar with the same or similar dominant louse species (as observed in genus Rattus, Niviventer, Apodemus and Eothenomys ). The similarity of sucking louse communities is highly consistent with the affinity of small mammal hosts in taxonomy. The results suggest a close relationship of co-evolution between sucking lice and their hosts.     

云南省吸虱昆虫区系及物种多样性

吸虱是寄生于真兽类哺乳动物体表的专性吸血寄生虫,广布于世界各地。云南省已知吸虱昆虫9科13属44种,分别占中国已知吸虱科、属、种的81.82%,59.09%,45.83%。文章参考大量相关文献,从分类阶元、特有物种、动物地理区划和宿主动物4个方面分析云南省吸虱的物种多样性。云南省吸虱特有种有13种,占云南省已知吸虱种类的29.55%,27种为东洋种,15种为古北和东洋两界兼有种,广布种9种。吸虱在5个地理小区的分布,以横断山中部和横断山南部2个地理小区的吸虱物种多样性较高,其它3个区的物种多样性较低。相对于全国而言,云南省吸虱物种多样性较高,吸虱的宿主动物种类丰富。但蚤、恙螨和革螨等其它体表寄生虫相比较,兽类宿主动物体表吸虱的物种多样性明显低于其它体表寄生虫,1科(属)阶元的吸虱其宿主多为相对一致的1个科(属)动物阶元,反映了吸虱宿主特异性较高的事实,吸虱昆虫与其对应的宿主动物已经形成了比较稳定的"一对一"的寄生关系,这是吸虱昆虫与其宿主动物协同进化的生态学表现。     

Rodent louse diversity,phylogeny, and cospeciation in the Manu Biosphere Reserve,Peru

We investigated the diversity, cophylogenetic relationships, and biogeography of hoplopleurid sucking lice (Phthiraptera: Anoplura) parasitizing rodents (Muridae: Sigmodontinae) in the Manu National Park and Biosphere Reserve. Our morphological and molecular studies reveal that 15 distinct louse species parasitize 19 rodent species. Three of these louse species are new to science, and all but two of the host associations were previously unknown. We find that hoplopleurid lice in South America parasitize multiple host species across a large geographic area, and that Peru represents a new geographic locality for almost all the louse species collected in the present study. Phylogenetic analyses of mitochondrial and nuclear data reveal that the louse family Hoplopleuridae and the genera Hoplopleura and Pterophthirus are not monophyletic, and lice do not appear to group by host tribe, collecting locality, or collection elevation. The lack of monophyly for these apparently natural groups (taxonomic, locality, and elevation) indicates that host switching with or without parasite speciation may be prevalent among hoplopleurid lice. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 598–610.     

Host specificity and genealogy of the louse Polyplax serrata on field mice, Apodemus species: a case of parasite duplication or colonisation?

The genealogy, population structure and population dynamics of the sucking louse Polyplax serrata were analysed across four host species of the genus Apodemus. An analysis of 126 sequences of cytochrome c oxidase subunit I using phylogenetic approaches and haplotype networking revealed a clear structure of European samples, forming three distinct and genetically distant clades with different host specificities. Although a clear connection was detected between the host and parasite genealogies/phylogenies, a uniform pattern of co-speciation was not found. For example, a dramatic shift in the degree of host specificity was demonstrated for two related louse lineages living in sympatry and sharing one of their host species. While one of the louse lineages frequently parasitised two different host taxa (Apodemus sylvaticus and Apodemus flavicollis), the other louse lineage was strictly specific to A. flavicollis. The estimate of divergence time between the two louse lineages indicates that they may have arisen due to parasite duplication on A. flavicollis.     

Evolutionary history of mammalian sucking lice (Phthiraptera: Anoplura)

Background  

Sucking lice (Phthiraptera: Anoplura) are obligate, permanent ectoparasites of eutherian mammals, parasitizing members of 12 of the 29 recognized mammalian orders and approximately 20% of all mammalian species. These host specific, blood-sucking insects are morphologically adapted for life on mammals: they are wingless, dorso-ventrally flattened, possess tibio-tarsal claws for clinging to host hair, and have piercing mouthparts for feeding. Although there are more than 540 described species of Anoplura and despite the potential economical and medical implications of sucking louse infestations, this study represents the first attempt to examine higher-level anopluran relationships using molecular data. In this study, we use molecular data to reconstruct the evolutionary history of 65 sucking louse taxa with phylogenetic analyses and compare the results to findings based on morphological data. We also estimate divergence times among anopluran taxa and compare our results to host (mammal) relationships.     

吸虱昆虫群落数量分类及其与小兽宿主的协同进化关系

以云南省16种主要小兽宿主体表吸虱群落为分类单元,以基于共54个群落特征变量的各群落优势虱种的相对优势度D_r、染虱率R_L、虱指数I_L、总染虱率R_LT和总虱指数I_LT及各群落的结构参数（平均丰富度 、平均均匀度 、平均多样性指数 及平均优势度指数 ）来探讨云南省吸虱昆虫与其宿主协同进化的关系。运用SPSS12.0统计软件中的系统聚类分析,对16种主要小兽宿主体表吸虱群落进行数量分析。结果将16种小兽宿主体表吸虱群落分为8个类群。隶属同一个属的小兽,其体表吸虱群落相似程度高,在系统聚类分析中聚为一类; 大多数吸虱群落相似性大小与相应小兽宿主在动物分类上的近缘性高低呈高度一致。结果提示： 吸虱昆虫与其所寄生的小兽宿主存在同步进化的关系,特定的小兽宿主,其体表吸虱群落的构成比较稳定,宿主动物的近缘关系越近,其体表吸虱群落的相似性越高,这是吸虱昆虫与小兽宿主协同进化的一个重要生态学证据。     

Enderleinellus tamiasis Fahrenholz, 1916 (Anoplura: Enderleinellidae), an introduced species, and a new sucking louse for the French fauna

A new sucking louse is recorded for the French Anopluran fauna, Enderleinellus tomiasis found on the introduced Sciurid Tamias sibiricus. This observation highlights the maintenance of parasites when introduced with their hosts and when their hosts settle into a novel environments. It suggests a common origin for two out of four populations of Siberian chipmunks examined. The authors describe the morphological criteria that allow the distinction between the two species of Enderleinellus and each infecting a sciurid host found in our country.     

Lice and ticks of the eastern rufous mouse lemur, Microcebus rufus, with descriptions of the male and third instar nymph of Lemurpediculus verruculosus (Phthiraptera: Anoplura)

Sucking lice and ticks were collected from live-trapped eastern rufous mouse lemurs, Microcebus rufus Geoffroy, in and around the periphery of Ranomafana National Park, southeastern Madagascar, from 2007 to 2009. Samples of 53 sucking lice (Insecta: Phthiraptera: Anoplura) and 28 hard ticks (Acari: Ixodidae) were collected from 36 lemur captures representing 26 different host individuals. All of the lice were Lemurpediculus verruculosus (Ward) (6 males, 46 females, 1 third instar nymph). Only the holotype female was known previously for this louse and the host was stated to be a "mouse lemur." Therefore, we describe the male and third instar nymph of L. verruculosus and confirm M. rufus as a host (possibly the only host) of this louse. All of the ticks were nymphs and consisted of 16 Haemaphysalis lemuris Hoogstraal, 11 Haemaphysalis sp., and 1 Ixodes sp. The last 2 ticks listed did not morphologically match any of the Madagascar Haemaphysalis or Ixodes ticks for which nymphal stages have been described.     

The mitochondrial genome of the chimpanzee louse,Pediculus schaeffi: insights into the process of mitochondrial genome fragmentation in the blood-sucking lice of great apes

Background

Blood-sucking lice in the genera Pediculus and Pthirus are obligate ectoparasites of great apes. Unlike most bilateral animals, which have 37 mitochondrial (mt) genes on a single circular chromosome, the sucking lice of humans have extensively fragmented mt genomes. The head louse, Pediculus capitis, and the body louse, Pe. humanus, have their 37 mt genes on 20 minichromosomes. The pubic louse, Pthirus pubis, has its 34 mt genes known on 14 minichromosomes. To understand the process of mt genome fragmentation in the sucking lice of great apes, we sequenced the mt genome of the chimpanzee louse, Pe. schaeffi, and compared it with the three human lice.

Results

We identified all of the 37 mt genes typical of bilateral animals in the chimpanzee louse; these genes are on 18 types of minichromosomes. Seventeen of the 18 minichromosomes of the chimpanzee louse have the same gene content and gene arrangement as their counterparts in the human head louse and the human body louse. However, five genes, cob, trnS₁, trnN, trnE and trnM, which are on three minichromosomes in the human head louse and the human body louse, are together on one minichromosome in the chimpanzee louse.

Conclusions

Using the human pubic louse, Pt. pubis, as an outgroup for comparison, we infer that a single minichromosome has fragmented into three in the lineage leading to the human head louse and the human body louse since this lineage diverged from the chimpanzee louse ~6 million years ago. Our results provide insights into the process of mt genome fragmentation in the sucking lice in a relatively fine evolutionary scale.

Electronic supplementary material

The online version of this article (doi:10.1186/s12864-015-1843-3) contains supplementary material, which is available to authorized users.     

Phocid seals, seal lice and heartworms: a terrestrial host-parasite system conveyed to the marine environment

Adaptation of pinnipeds to the marine habitat imposed parallel evolutions in their parasites. Ancestral pinnipeds must have harboured sucking lice, which were ancestors of the seal louse Echinophthirius horridus. The seal louse is one of the few insects that successfully adjusted to the marine environment. Adaptations such as keeping an air reservoir and the ability to hold on to and move on the host were necessary, as well as an adjustment of their life cycle to fit the diving habits of their host. E. horridus are confined to the Northern Hemisphere and have been reported from 9 species of northern phocids belonging to 4 genera, including land-locked seal species. The transmission from seal to seal is only possible when animals are hauled-out on land or ice. Lice are rarely found on healthy adult seals, but frequently on weak and young animals. The seal louse is suggested to play an important role as an intermediate host transmitting the heartworm Acanthocheilonema spirocauda among seals. However, the evidence is restricted to a single study where the first 3 larval stages of the heartworm were shown to develop in the louse. The fourth-stage larvae develop in the blood system of seals and eventually transform into the adult stage that matures in the heart. Since all other studies failed to confirm the presence of heartworm larvae in seal lice, other unknown intermediate hosts could be involved in the transmission of the heartworm. Transplacental transmission of microfilariae in seals has been suggested as an additional possibility, but is not likely to be important since the occurrence of heartworms in adult seals is very rare compared with juveniles. Furthermore, there are no findings of the first 3 larval stages in seals. This review shows that the heartworm infects nearly the same species of seals as the seal louse, except for the grey seal Halichoerus grypus, where the heartworm is absent. Prevalence and intensity of infection differ among regions in the Northern Hemisphere. As for seal lice, heartworms mainly infect immature seals, and after infection the prevalence seems to decrease with increasing age of the host.     

Hoplopleura janzeni n. sp. (Phthiraptera: Anoplura), a new sucking louse from a Central American swimming mouse.

Both sexes of a new species of sucking louse Hoplopleura janzeni (Phthiraptera: Hoplopleuridae) are described and illustrated from the Central American ichthyomyine swimming mouse Rheomys raptor (Rodentia: Muridae) collected in Costa Rica. The morphology of the new species is compared with that of Hoplopleura exima Johnson, the only other species of sucking louse known to parasitize an ichthyomyine rodent. Hoplopleura janzeni is unique in having posteriorly directed spurs on the first antennal segment, the fore- and midcoxae, and the hind femora of both sexes.     

Sucking louse (Hoplopleura erratica: Insecta, Anoplura) exchange between individuals of a wild population of Eastern chipmunks, Tamias striatus, in central Tennessee, U.S.A.

Adult Hoplopleura erratica sucking lice were labelled using a setal clipping technique. Host exchange for lice marked in this manner was monitored through a small population of dye-marked Eastern chipmunks, Tamias striatus, the normal host of this ectoparasite, in a deciduous woodland in central Tennessee, U.S.A. This was accomplished by frequent host live-trapping, anaesthetization and pelage examinations. Louse transfer was most prevalent during the host summer mating period, presumably because of increased chipmunk contacts. At this time, 66·7% of marked adult lice within the host population transferred. Exchanges between opposite sex hosts were more common and adult males donated most lice; no juvenile chipmunks were recorded as louse donors. A greater proportion of male than female lice transferred. Using the exclusive boundary strip method, all exchanging chipmunks had overlapping ranges or shared at least one boundary line.     

中国云南洱海周边小兽体表吸虱多样性

在2003—2004年对云南大理洱海周边小兽体表寄生吸虱进行了调查,诱捕小兽3303只,隶属4目7科、15属、21种,共收集14635只吸虱,共计5科、6属、21种。调查点位于洱海周边3个不同方位,恰好处于无量山（洱海东部）、哀牢山（洱海南部）和苍山（洱海西部）3座山的连接地段,由于洱海的天然隔离使这3个方位形成了同地域（经度、纬度、海拔和动物地理区划相同）异生境的地理景观。此项研究的目的是调查同地域异生境里小兽体表吸虱在物种多样性、丰富度、群落结构、相似性和分布。结果表明吸虱的物种多样性低,群落结构简单。吸虱的丰富度、分布和物种多样性随着宿主和生境的不同而存在着显著的差异,这可能暗示着生态环境影响着吸虱和它相对应的小兽宿主的构成和分布。某种宿主经常有某种固定的吸虱,在动物分类上近缘关系较近的小兽宿主,其体表昆虫群落相似性较高。洱海周边不同方位同种小兽宿主上的吸虱物种相同,这个结果强烈暗示着吸虱与其对应宿主之间有明显的协同进化关系。     

Louse infestations of tree shrews (Tupaia glis)

Nine adult tree shrews, Tupaia glis, recently imported from West Malaysia were visually examined for ectoparasites while under general anesthesia. Three shrews were infested by the sucking louse, Sathrax durus , and six shrews had louse ova belonging to this species; two shrews had neither lice nor ova. A total of 20 adult female, 10 adult male, and three third instar nymphal lice was collected. Lice were located on the head, flanks, and dorsal body of shrews while ova were recorded mainly from the anterior flanks but also from some adjacent host sites. The tree shrews appeared to tolerate the lice well although louse vector capacity was not assessed. The last date that lice were recorded from shrews was 22 days after colony set-up, and the last date on which seemingly viable ova were recorded was 64 days after set-up showing that the infestations were ultimately lost.     

Host behaviour drives parasite genetics at multiple geographic scales: population genetics of the chewing louse,Thomomydoecus minor

Pocket gophers and their symbiotic chewing lice form a host–parasite assemblage known for a high degree of cophylogeny, thought to be driven by life history parameters of both host and parasite that make host switching difficult. However, little work to date has focused on determining whether these life histories actually impact louse populations at the very fine scale of louse infrapopulations (individuals on a single host) at the same or at nearby host localities. We used microsatellite and mtDNA sequence data to make comparisons of chewing‐louse (Thomomydoecus minor) population subdivision over time and over geographic space where there are different potential amounts of host interaction surrounding a zone of contact between two hybridizing pocket‐gopher subspecies. We found that chewing lice had high levels of population isolation consistent with a paucity of horizontal transmission even at the very fine geographic scale of a single alfalfa field. We also found marked genetic discontinuity in louse populations corresponding with host subspecies and little, if any, admixture in the louse genetic groups even though the lice are closely related. The correlation of louse infrapopulation differentiation with host interaction at multiple scales, including across a discontinuity in pocket‐gopher habitat, suggests that host behaviour is the primary driver of parasite genetics. This observation makes sense in light of the life histories of both chewing lice and pocket gophers and provides a powerful explanation for the well‐documented pattern of parallel cladogenesis in pocket gophers and chewing lice.     

Arthropod symbiotes of Laonastes aenigmamus (Rodentia:Diatomyidae)

Arthropod symbiotes of the Laotian rock-rat, Laonastes aenigmamus (Rodentia:Diatomyidae), from Laos are examined. This host is a member of Diatomyidae previously thought to have gone extinct >10 million yr ago. Permanent symbiotes are represented by 2 species, a new species of sucking louse, Polyplax sp., near rhizomydis (Phthiraptera:Polyplacidae), and a new species of fur mite, Afrolistrophorus sp., near maculatus (Acariformes:Listrophoridae). The temporary parasites are represented by 18 species, i.e., 1 mesostigmatan species, i.e., a new species of Androlaelaps near casalis (Parasitiformes:Laelapidae); immature stages of 2 tick species, Ixodes granulatus and Haemaphysalis sp. (Parasitiformes:Ixodidae); and a rich fauna of chiggers (Acariformes:Trombiculidae) comprising 8 genera and 15 species. It is hypothesized that this host completely lost its initial fauna of ectosymbiotes and that ancestors of the recorded symbiotes switched to this host from rodents of the superfamily Muroidea.     

Abrocomaphthirus hoplai, a new genus and species of sucking louse from Chile and its relevance to zoogeography

Both sexes of Abrocomaphthirus hoplai, new genus and new species (Anoplura: Polyplacidae), are described and illustrated. The endemic Chilean chinchilla rat Abrocoma bennetti Waterhouse (Rodentia: Abrocomidae) is the type host. The definition of the family Polyplacidae is amended to accommodate the new genus. Polyplax longa (Werneck), also referred to in the literature as Neohaematopinus longus Werneck, is reassigned to Abrocomaphthirus. The host of A. longus comb.n., is Abrocoma cinerea Thomas, another chinchilla rat, which inhabits parts of Argentina, Bolivia, Chile and Peru. The erection of Abrocomaphthirus as a distinct genus has important zoogeographical and evolutionary implications. The tenuous anomaly of P. longa being the sole native representative of the genus Polyplax in South America, possibly with African affinities, is now refuted. Instead, partial colonization of the neotropics by native species of both Polyplax and Neohaematopinus appears to have been relatively recent and from North America. The phylogenetic affinities of Abrocomaphthirus are unknown, but it appears to be closely related to other, more ancient, native South American polyplacid louse genera, such as Cuyana, Eulinognathus, Galeophthirus, and Lagidiophthirus. Arguments are presented in support of an ancestral zoogeographical link to Africa for these louse genera.     

Molecular phylogeny and novel host associations of avian chewing lice (Insecta: Phthiraptera) from South Africa

Compared with Europe and the Americas, the ectoparasites of African birds are poorly understood, despite the avian fauna being relatively well known. Notably, previous studies documenting the host associations and genetic diversity of parasitic chewing lice of southern African birds have been limited in geographic and taxonomic scope. Recent field expeditions exploring the avian diversity in South Africa facilitated an opportunity to obtain louse specimens from a taxonomically diverse host assemblage. This study is the first to investigate avian louse host associations and diversity across a large portion of South Africa encompassing several distinct habitat types, while incorporating molecular genetic data (from portions of the mitochondrial COI and nuclear EF‐1α genes) for ectoparasite phylogenetic analyses. From 1105 South African bird individuals and 170 species examined for lice, a total of 105 new louse–host associations were observed. Morphological and genetic examination of lice with these new host associations reveals a maximum of 66 louse species new to science. Results of this study support the observation that examining museum specimens is a useful way to investigate louse diversity and host associations.     

Epifaunistic arthropod parasites of the four-striped mouse, Rhabdomys pumilio, in the Western Cape Province, South Africa

Flea, lice, mite, and tick species associated with 510 Rhabdomys pumilio were collected at 9 localities in the Western Cape Province, South Africa. The aims of the study were first to quantify the species richness, prevalence, and relative mean intensity of infestation of epifaunistic arthropod species associated with R. pumilio, and second to determine temporal variations in the mean abundance of the parasitic arthropods. Each mouse was examined under a stereoscopic microscope and its parasites were removed, identified, and quantified. The epifaunal population was made up of more than 25,000 individuals and included 8 flea, 1 sucking louse, 11 mite, and 13 ixodid tick species. Female-biased sex ratios were noted for 9 (30%) of the ectoparasite species. Three undescribed mite and 1 undescribed tick species were recovered, and new locality records for 2 flea, the louse, and 2 mite species were documented. A phoretic host association between a nonparasitic mite species, Psylloglyphus uilenbergi kivuensis, and 3 flea species, Chiastopsylla rossi, Hypsophthalmus temporis, and Listropsylla agrippinae, was recorded. The mean abundance of the parasitic mite and insect species were higher during the cold wet season, whereas ticks were more numerous during the warm dry months. The large number of ectoparasite species on R. pumilio, a locally abundant and regionally widespread species, is of medical and veterinary importance particularly in relation to the transmission of pathogens such as Anaplasma marginale, Babesia caballi, and Babesia canis to domestic animals; Rickettsia conori; Yersinia pestis; and the viral disease Crimean-Congo hemorrhagic fever to humans.