The diversity, recombination and horizontal transmission of Wolbachia in spiders in China

다국어 초록 (Multilingual Abstract)

Wolbachia are maternally inherited endosymbiotic bacteria. These intracellular bacteria are common in ar thropods and could manipulate host reproduction in diverse ways, such as feminization, parthenogenesis, male killing and cytoplasmic incompatibility. In spiders, infection by Wolbachia has been found in a total of 99 species belonging to 62 genera and 17 families. Furthermore, recent studies analyzed the phylogeny of Wolbachia in Hylyphantes graminicola, 2 cave spiders and Agelenopsis species using multilocus sequence typing (MLST) approach. However, the diversity of Wolbachia strains determined by MLST in spiders from China is still largely unknown.
In this study, we collected 1153 spider individuals from Mangshan in China and screened for Wolbachia in 975 individuals representing 68 spider species belonging to 45 genera of 16 families. We analyzed the phylogenetic relationship between Wolbachia and their host spiders by MLST approach. We found novel infections of Wol bachia in 1 family, 9 genera and 20 species of spiders. We found 13 new Wolbachia strains and suggest that group A is more common than group B in Wolbachia that infect spiders. Our results revealed three recombination events of the concatenated multilocus sequences in Wolbachia that infect spiders. Furthermore, our results demonstrated the phylogenetic incongruence between Wolbachia and spiders, suggesting the horizontal transmission of Wol bachia in spiders.
We suggest that recombination and horizontal transmission may play an important role in the diversity and evolution of Wolbachia in spiders.

참고문헌 (Reference)

1 Yun, Y., "Wolbachia strains typing in different geographic population spider, Hylyphantes Graminicola(Linyphiidae)" 62 : 139-145, 2011

2 Yun, Y., "Wolbachia screening in spiders and assessment of horizontal transmission between predator and prey" 40 : 164-169, 2011

3 Stouthamer, R., "Wolbachia pipientis : Microbial manipulator of arthropod reproduction" 53 (53): 71-102, 1999

4 Kittayapong, P., "Wolbachia infection complexity among insects in the tropical rice-field community" 12 : 1049-1060, 2003

5 Zimmermann, B. L., "Wolbachia in Neotropical terrestrial isopods. FEMS Microbiol" 91-, 2015

6 Baldo, L., "Wolbachia are present in Southern African scorpions and cluster with supergroup F" 55 (55): 367-373, 2007

7 Goodacre, S. L., "Wolbachia and other endosymbiont infections in spiders" 15 : 517-527, 2006

8 Werren, J. H., "Wolbachia : master manipulators of invertebrate biology" 6 : 741-751, 2008

9 Baldo, L., "Widespread recombination throughout Wolbachia genomes" 23 : 437-449, 2006

10 Jiggins, F. M., "The rate of recombination in Wolbachia bacteria" 19 : 1640-1643, 2002

11 Ahmed, M. Z., "The intracellular bacterium Wolbachia uses parasitoid wasps as phoretic vectors for efficient horizontal transmission" 11 (11): e1004672-, 2015

12 Weinert, L. A., "The incidence of bacterial endosymbionts in terrestrial arthropods" 282 : 1-6, 2015

13 Fitton, M. G., "The Hymenoptera associated with spiders in Europe" 90 : 65-93, 1987

14 Malloch, G., "Super-infections of Wolbachia in byturid beetles and evidence for genetic transfer between A and B super-groups of Wolbachia" 14 : 627-637, 2005

15 Rigaud, T., "Success and failure of horizontal transfers of feminizing Wolbachia endosymbionts in woodlice" 8 : 249-255, 1995

16 Dunaj, S. J., "Spider phylosymbiosis:divergence of widow spider species and their tissues’ microbiomes" 20 : 2020

17 Baldo, L., "Revisiting Wolbachia supergroup typing based on WSP : spurious lineages and discordance with MLST" 55 (55): 81-87, 2007

18 Jiggins, F. M., "Recombination confounds interpretations of Wolbachia evolution" 268 : 1423-1427, 2001

19 Martin, D., "RDP : detection of recombination amongst aligned sequences" 16 (16): 562-563, 2000

20 Li, S. J., "Plant mediated horizontal transmission of Wolbachia between whiteflies" 11 : 1019-1028, 2017

21 Gauld, I. D., "Phylogeny of the Polysphincta group of genera(Hymenoptera : Ichneumonidae; Pimplinae) : a taxonomic revision of spider ectoparasitoids" 31 : 529-564, 2006

22 Zhou, W., "Phylogeny and PCR-based classification of Wolbachia strains using wsp gene sequences" 265 : 509-515, 1998

23 Batista, P. D., "Phylogenetic placement and evidence for horizontal transfer of Wolbachia in Plutella xylostella(Lepidoptera : Plutellidae)and its parasitoid, Diadegma insulare(Hymenoptera : Ichneumonidae)" 142 (142): 57-64, 2010

24 Liu, K. -K., "Parasitoidism of the Ichneumon Oxyrrhexis rugosus on the funnel-web spider Allagelena difficilis and developmental characteristics of the parasitoid" 50 : 931-939, 2015

25 Gonzaga, M. O., "Parasitoid-induced mortality of Araneus omnicolor (Araneae, Araneidae) by Hymenoepimecis sp. (Hymenoptera, Ichneumonidae) in southeastern Brazil" 94 (94): 223-227, 2007

26 Swofford, D. L., "PAUP-A computer program for phylogenetic inference using maximum parsimony" 102 : A9-, 1993

27 Cobb, L. M., "Occurrence of parasitoid wasps, Baeus sp. and Gelis sp., in the egg sacs of the wolf spiders Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in southeastern Idaho" 118 : 122-123, 2004

28 Enigl, M., "Negative evidence of Wolbachia in the predaceous mite Phytoseiulus persimilis" 36 (36): 249-262, 2005

29 Baldo, L., "Multilocus sequence typing system for the endosymbiont Wolbachia pipientis" 72 (72): 7098-7110, 2006

30 Baldo, L., "Mosaic nature of the Wolbachia surface protein" 187 (187): 5406-5418, 2005

31 Ilinsky, Y., "Molecular diversity of Wolbachia in lepidoptera : prevalent allelic content and high recombination of MLST genes" 109 : 164-179, 2017

32 Korenko, S., "Modification of Tetragnatha montana(Araneae, Tetragnathidae)web architecture induced by larva of the parasitoid Acrodactyla quadrisculpta(Hymenoptera, Ichneumonidae, Polysphincta genus-group)" 54 : 1-7, 2015

33 Gonzaga, M. O., "Modification of Nephila clavipes (Araneae, Nephilidae) by the parasitoids Hymenoepimecis bicolor and H. robertsae (Hymenoptera, Ichneumonidae)" 22 : 151-165, 2010

34 Posada, D., "Modeltest : testing the model of DNA substitution" 14 : 817-818, 1998

35 Tamura, K., "MEGA4: Molecular evolutionary genetics analysis (MEGA) software version 4.0" 24 : 1596-1599, 2007

36 Baldo, L., "Insight into the routes of Wolbachia invasion : high levels of horizontal transfer in the spider genus Agelenopsis revealed by Wolbachia strain and mitochondrial DNA diversity" 17 : 557-569, 2008

37 Hilgenboecker, K., "How many species are infected with Wolbachia? a statistical analysis of current data" 281 : 215-220, 2008

38 Wasbauer, M. S., "Host records for some north American spider wasps, with notes on prey selection(Hymenoptera : Pompilidae)" 35 : 393-401, 1962

39 Kurczewski, F. E., "Host records for some north American Pompilidae(Hymenoptera)first supplement" 41 : 367-382, 1968

40 Kurczewski, F. E., "Host records for some north American Pompilidae (Hymenoptera). third supplement" 46 : 65-81, 1973

41 Yang, X. -H., "High levels of multiple infections, recombination and horizontal transmission of Wolbachia in the Andricus mukaigawae(Hymenoptera; Cynipidae)communities" 8 : e78970-, 2013

42 Reuter, M., "High levels of multiple Wolbachia infection and recombination in the ant Formica exsecta" 20 : 748-753, 2003

43 Chen, Y. T., "Geography has a greater effect than Wolbachia infection on population genetic structure in the spider mite, Tetranychus pueraricola" 1-10, 2016

44 Ahmed, M. Z., "Evidence for common horizontal transmission of Wolbachia among butterflies and moths" 16 : 1-16, 2016

45 Zug, R., "Epidemiology in evolutionary time : the case of Wolbachia horizontal transmission between arthropod host species" 25 : 2149-2160, 2012

46 Guarisco, H., "Egg parasitism of the jumping spider, Phidippus clarus(Salticidae)by the parasitoid wasp, Idris saitidis(Scelionidae)" 102 : 68-69, 1999

47 Rozas, J., "DnaSP, DNA polymorphism analyses by the coalescent and other methods" 19 (19): 2496-2497, 2003

48 Ros, V. I. D., "Diversity and recombination in Wolbachia and Cardinium from Bryobia spider mites" 12 : S13-, 2012

49 Werren, J. H., "Distribution of Wolbachia among Neotropical Arthropods" 262 : 197-204, 1995

50 Wang, G. -H., "Discovery of a new Wolbachia supergroup in cave spider species and the lateral transfer of phage WO among distant hosts" 41 : 1-7, 2016

51 Salunke, B. K., "Determination of Wolbachia diversity in butterflies from Western Ghats, India, by a multigene approach" 78 (78): 4458-4467, 2012

52 Larkin, M. A., "Clustal W. Clustal X version 2. 0" 23 : 2947-2948, 2007

53 Werren, J. H., "Biology of Wolbachia" 42 : 587-609, 1997

54 Foelix, R., "Biology of Spiders" Oxford University Press 2011

55 Tan, Y. G., "A preliminary report on parasitoid wasps of spiders" 8 : 70-72, 1999