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Phoretic Mites (Acari: Chaetodactylidae) Associated With Phoretic mites (Acari: Chaetodactylidae) associated with the solitary bee Tetrapedia diversipes (Apidae: Tetrapediini) Guaraci Cordeiro, Mariana Taniguchi, Carlos Flechtmann, Isabel Alves-Dos-Santos To cite this version: Guaraci Cordeiro, Mariana Taniguchi, Carlos Flechtmann, Isabel Alves-Dos-Santos. Phoretic mites (Acari: Chaetodactylidae) associated with the solitary bee Tetrapedia diversipes (Apidae: Tetrapedi- ini). Apidologie, Springer Verlag, 2011, 42 (2), pp.128-139. 10.1051/apido/2010044. hal-01003578 HAL Id: hal-01003578 https://hal.archives-ouvertes.fr/hal-01003578 Submitted on 1 Jan 2011 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Apidologie (2011) 42:1 –128 39 Original article c INRA/DIB-AGIB/EDP Sciences, 2010 DOI: 10.1051/apido/2010044 Phoretic mites (Acari: Chaetodactylidae) associated with the solitary bee Tetrapedia diversipes (Apidae: Tetrapediini)* Guaraci Duran Cordeiro1,MarianaTaniguchi1, Carlos Holger Wenzel Flechtmann2,IsabelAlves-dos-Santos3 1 Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo (USP), Av. Bandeirantes, 3900, Bloco 2, 14040-901, Ribeirão Preto, SP, Brasil 2 Departamento de Entomologia, Fitopatologia e Zoologia Agrícola, Escola Superior de Agricultura Luiz de Queiroz (ESALQ), Universidade de São Paulo (USP), Caixa postal 9, 13418-900, Piracicaba, SP, Brasil 3 Laboratório de Abelhas, Departamento de Ecologia, Instituto de Biociências, Universidade de São Paulo (USP), Rua do Matão 321, trav 14. Cidade Universitária, 05508-900 São Paulo, SP, Brasil Received 7 October 2009 – Revised 25 March 2010 – Accepted 26 March 2010 Abstract – Solitary bees of the genus Tetrapedia have a specific association with mites of the genus Roubikia (Chaetodactylidae). These mites are frequently found attached to active Tetrapedia bees. We quantified the number of mites on individuals of Tetrapedia diversipes Klug and examined the interac- tion between these species. Nests of T. diversipes were obtained from trap-nests placed in four localities in São Paulo, Brazil. The study lasted from March 2007 to February 2009. Out of a total of 650 nests with emergences, 118 were infested with mites (Roubikia sp.). From these nests, 176 individuals of T. diversipes emerged with mites on their bodies. Additionally, six individuals of Coelioxoides waltheriae, the specific kleptoparasitic bee to T. diversipes, emerged. Mites were attached mainly to the mesosoma. All nests in- fected with mites did not presented mortality of the immature. The mortality rate of nests was inversely related to the level of mite infestation, suggesting a mutualistic interaction in which mites may remove fungi from the nests, while the bees would provide the mites with transport, dispersal, and shelter. solitary bee / phoresy / trap-nest / Atlantic Forest / symbiosis 1. INTRODUCTION (Lithurgini) and Apidae (Xylocopini) sug- gest that chaetodactylids may have established Mites and bees have co-existed since the themselves in the Lower Cretaceous, when Cretaceous, and there is evidence of a close re- these two bee families became separated lationship between some taxa, probably result- (Engel, 2001). Another hypothesis is that ing from a co-evolutionary processes (Klimov the mites arose in a later period, during et al., 2007a). Most mite lineages associated the Eocene, when many exchanges among with bees use species of Apidae and Megachil- hosts would have occurred simultaneously idae as hosts (Eickwort, 1994; Klimov et al., with the dispersal of bees through the conti- 2007a). These mites belong to the family nents (Klimov and O’Connor, 2007). Chaetodactylidae (Acari: Astigmata). Associ- Chaetodactylidae comprises over 200 de- ations with basal lineages of Megachilidae scribed species within five genera and dis- tributed worldwide, except for the poles Corresponding author: I. Alves-dos-Santos, (Klimov and O’Connor, 2008). The active ini- [email protected] tial phases of the mites (larva and nymph) * Manuscript editor: Stan Schneider have been described for 85 species. During Phoretic mites (Acari: Chaetodactylidae) associated with the solitary bee... 129 these phases, mites live within the bee nests the sand and soil particles may attach to certain and feed on materials found in the brood bees that collect this material (Roubik, 1987; cells, and may be mutualists (consuming nest Vicidomini, 1996). There are frequent changes wastes), parasitoids (killing the egg or the between relatively close taxa within the same larva), commensals or parasites (consuming genus (Klimov et al., 2007b). Approximately the larval food and so hindering the develop- 170 Sennertia species are associated with 469 ment of the larva) (Krombein, 1962; Roubik, species of Xylocopa (Michener, 2000; Klimov 1987;Quetal.,2002). Most Chaetodactylidae and O’Connor, 2007). species are found in association with bees in the phases of deutonymph or phoretic hypo- Among the solitary bees, the genus Tetra- pus, frequently positioned on the acarinarium pedia (Apidae, Tetrapediini), a group of oil- (Fain, 1981). In this dispersal stage, the mite collecting bees, was reported to be in specific stops feeding and becomes inactive; it is trans- association with mites of the genus Roubikia ported by the host to the next site where a new (Roubik, 1987; Klimov and O’Connor, 2007; cycle begins (Baker et al., 1987). The hypopi Klimov et al., 2007a, b). The mites also lodge have a morphology fully adapted to this life on Coelioxoides (Apidae: Tetrapediini), their phase: they have no mouthparts or digestive specific kleptoparasitic species (Klimov et al., tract, a highly sclerotized carapace, which pre- 2007a). It was known that the interaction be- vents dehydration, robust pretarsal claws, and tween Roubikia and Tetrapedia is commensal- caudoventral suckers for attaching to the host istic: the mites feed on larval food and on nest (Houck and O’Connor, 1991). In most cases, a materials without causing any harm to the im- certain synchronism between the life cycles of mature (Klimov et al., 2007b). The association both associates occurs (Klimov and O’Connor, is restricted to the Neotropical region, which is 2007). an area of occurrence for both groups. Some bee species exhibit specialized re- Alves-dos-Santos et al. (2002) and Camillo gions on the body called acarinaria where (2005) studied the nesting biology and the mites are frequently lodged (O’Connor and nest architecture of Tetrapedia diversipes.This Klompen, 1999). Acarinaria are small cavities, species has a bivoltine cycle: the first gen- sometimes covered with setae, which facili- eration is active in the spring, whereas the tate the attachment of mites to the bee’s body second is active in the summer. The females and provide them protection during trans- nest on wood substrate and frequently use pre- port. These structures occur in many fam- existing cavities in tree bark made by beetles; ilies of aculeate hymenopterans (Eickwort, in such cases, they may be captured in trap- 1994; O’Connor and Klompen, 1999)andare nests (Garófalo et al., 2004). Old nests are fre- evidence for the strong reciprocity that those quently re-used. The oil collected by the fe- interactions have been present through evolu- males is used for nest construction, compacted tionary time. However, not all bees with asso- with sand particles, and mixed together with ciated mites exhibit developed acarinaria, and pollen for larval food. The larvae of Tetrape- the mites can, therefore, be distributed ran- dia develop in cells and do not spin cocoons. domly on the host body (Klimov et al., 2007b). Coelioxoides waltheriae Ducke is the specific Among bees of the family Apidae, acari- kleptoparasitic species of T. diversipes. naria are present in species of Xylocopa, Cer- atina,andTetrapedia (Klimov and O’Connor, We suppose that if the relationship of 2007). However, these features not always Roubikia mites with T. diversipes is mutualis- result from mutualistic associations (Klimov tic, than a lower mortality rate would be found et al., 2007b). when the mites are present. Therefore, the ob- Despite the specificities within associa- jectives of the present study were to verify tions, host changes may occur. In some cases, the infestation by mites of the genus Roubikia they occur by mistake. For instance, a mite both in nests and on emerging individuals of may remain on flowers (ex. Sennertia)andat- the solitary bee Tetrapedia diversipes and to tach to different bees, or a mite that blends in study the interaction between them. 130 G.D. Cordeiro et al. 2. MATERIAL AND METHODS MZ16, software IM50). For further details, images were made with a scanning electron microscope We obtained nests of Tetrapedia diversipes by (SEM) in the laboratory NAP/MEPA at ESALQ – setting up trap-nests. This method is widespread USP Campus Piracicaba. We glued dried specimens for the capture of solitary bees and consists of of- in stubs with a double-sided tape at the desired po- fering artificial cavities through cardboard tubes sition and subjected them
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