The Behavioural Responses by Larvae of the Water Mite Unionicola Foili (Acari: Unionicolidae) to Gravity, Light and Host Chemicals: Adaptations for Transmission Success?

The life cycle of the water mite Unionicola foili includes larvae that parasitize chironomid dipterans, and nymphs and adults that occur in association with the freshwater mussel Utterbackia imbecillis. The georesponses and photoresponses of larval U. foili in the presence of chemical substances from chironomids and mussels both prior to (= pre-chironomid larvae) and after their parasitism with chironomids (= post-chironomid larvae) were examined in the laboratory. The responses of larval U. foili to light and gravity changed during larval ontogeny and varied depending on the chemistry of the medium in which they were examined. Pre-chironomid larvae preferentially responded to water modified by chironomids given that negative phototaxis and positive georesponses were elicited only in the presence of water that had been modified by these hosts. Post-chironomid larvae exhibited host-induced negative phototaxis only in the presence of mussel-modified water. Post-chironomid larvae were consistently geopositive when exposed to water modified by mussels or chironomids. The pattern of responses exhibited by larval U. foili to light and gravity are interpreted as adaptations for locating potential hosts in the field.     

Random amplified polymorphic DNA analysis of kinship within host-associated populations of the symbiotic water mite Unionicola foili (Acari: Unionicolidae)

Kinship relations within populations of unionicolid water mites are not well known, owing to their complex life cycles and the fact that interactions between active and resting stages for some species are transitory. A number of species of unionicolid water mites are, however, obligate symbionts of freshwater mussels and spend most of their life cycle in association with these hosts. Among these species of mites, parents and offspring are more likely to co-occur and thus provide opportunities to address questions related to the structure of the mating system. The present study employs random amplified polymorphic DNA (RAPD) analysis to address kinship within populations of Unionicola foili living in symbiotic association with the host mussel Utterbackia imbecillis. DNA was amplified from adult mites and a representative number of eggs or larvae (n = 20-30) that were removed from mussels collected on three separate occasions (July, November, and March) over a 12-month period. Parsimony analyses of the molecular data for adults and progeny collected from mussels during July, November, and March revealed distinct groupings, that for the most part, corresponded to mites collected from each of the sampling periods. Many of the genetic markers obtained for male and female U. foili were not evident among the larvae or eggs, suggesting that adults obtained from a host mussel at the time of collection were not the parents of a majority of the progeny. However, female mites and eggs collected from mussels during March and November shared more markers than did females and progeny examined during July. Furthermore, many offspring in the July sampling period were found to have one or more parents absent from the sampled population. Overall, RAPD profiling appears to have limited usage in determining kinship within populations of U. foili, due to its recruitment patterns, and the relatively large number of adults and progeny per mussel. It may, however, prove to be a useful method for assessing genetic relatedness among unionicolid mussel-mites that have substantially lower population densities.     

The symbiotic water mite Unionicola formosa (Acari: Unionicolidae) ingests mucus and tissue of its molluscan host

Unionicola formosa is a symbiotic water mite that passes most of its life cycle in the mantle cavity of freshwater mussels. Although mites of this genus are often referred to as parasitic, little is known about their nutritional biology. A few species reportedly pierce the gill of a host mussel and ingest tissue or hemolymph. The present study was undertaken to identify possible sources of nutrition for U. formosa. To determine if mites ingested particulate matter in the mucous strand produced by a mussel during feeding, mussels with resident mites were exposed to a suspension of fluorescent microspheres. There was no evidence that U. formosa ingested the beads. Histochemical staining did, however, indicate a mucous material present in the midgut of the mites. Sodium dodecyl sulfate-polyacrylamide gel electrophoretic assays revealed a high molecular weight component, consistent with a mucopolysaccharide, present both in the mussel gill and the mites. Results from western blots and an immunoaffinity binding assay with antibodies against mussel gill tissue and hemolymph also indicated that mites ingested host tissue. Whereas U. formosa probably does not ingest particulate material acquired by its host's suspension feeding, it is apparent that this mite utilizes host mucus, gill tissue, or hemolymph for at least part of its nutrition.     

中国蚌螨属支序分类分析(蜱螨亚纲:蚌螨科)

选择35个特征对我国蚌螨属15种的支序分析结果表明:蚌螨P-Ⅳ(胫节)端腹部的指状突是将它们聚集成两群的主要特征.今村蚌螨P-Ⅳ端腹部仅具有短矩,与外群最近,认为是我国的蚌螨属水螨中最原始的种类;雌螨前殖吸盘板的形态影响戊蚌螨亚属与多盘蚌螨亚属的种类聚类,由此推测可以将蚌螨P-Ⅳ端腹部的指状突与雌螨前殖吸盘板的形态也作为蚌螨亚属的区分特征.由于殖吸盘结构简单是祖征态之一,因此认为具有5对殖吸盘的邻近蚌螨比6对殖吸盘的厚蚌螨更原始.雌螨具有1对殖吸盘板的寄蚌螨亚属种类与沃蚌螨亚属种类在支序系统树中形成姊妹群,结果支持Vidrine 将沃蚌螨亚属从寄蚌螨亚属中分离的修订.     

Host sex preferences and transmission success by the water mite Unionicola foili (Acari: Unionicolidae) parasitic on the midge Chironomus tentans (Diptera: Chironomidae)

This study examined whether ecoparasitic larval Unionicola foili exhibited a sex bias when infecting laboratory populations of the host insect Chironomus tentans and whether an association with male or female midges increased the likelihood of larval mites returning to the aquatic habitat. When laboratory populations of C. tentans were exposed to larval U. foili, there was a higher prevalence of mites among female hosts at emergence (17 of 30 males vs. 25 of 30 females infected by mites). However, there was no significant difference in the distribution or abundance of larvae among infected male (mean = 2.3 larvae per host) and female (mean = 2.6 larvae per host) midges. Larval mites parasitizing both male and female chironomids were more likely to return to water than could be expected by chance. Mite larvae infesting female C. tentans were more likely to return to water when female hosts deposited egg masses in water, suggesting that oviposition plays an important role in cueing larvae parasitizing female midges to detach. The mechanism responsible for increasing the likelihood that mites parasitizing male hosts return to water remains unclear. Future studies will address the possibility of parasite-mediated changes in host behavior.     

Guild structure in water mites (Unionicola spp.) inhabiting freshwater mussels: choice,competitive exclusion and sex

Summary Unionicolid water mites inhabit freshwater unionid mussels during the nymphal and adult stages of their life-cycle. Regular sampling of mussels from two sites in St. Mark's River, Fl. established that each of four species of water mite (Unionicola abnormipes, U. fossulata, U. serrata and U. formosa) occurred mainly in one or two of the mussel species available at each site.The role of preference for particular mussel species during host location was assessed for the first three mite species by choice experiments, in which mites were offered different mussel species simultaneously. In five out of six experiments, mites entered normally unused mussels as often as they did normally used ones. Additionally, a sexual difference in choice was found for U. fossulata, with males preferring one mussel species and females showing no preference. One mussel species, (Anodonta imbecilis), normally unused but chosen by mite species during the lab. experiments, is inhabited exclusively by the fourth mite species, U. formosa, in the field. An experiment showed that U. formosa excludes other mite species aggressively from Anodonta imbecilis.The results illustrate the sometimes misleading nature of simple sampling data as an indication of host specificity or host preference in parasites. They suggest also that the population dynamics of some parasites might be more fruitfully compared to unrelated, free-living species than to other parasites.     

Behavioral aspects of host recognition by the symbiotic water mite Unionicola formosa (Acarina,Unionicolidae)

Summary The occurrence and specificity of host recognition behavior of adult and nymphal Unionicola formosa and the capability of adult mites to recolonize various mussel species were examined. Adult U. formosa aggregated on excised tissue from their host mussel, Anodonta imbecilis, in preference to that of two other species of mussels. Nymphs also exhibited an aggregation response to host tissue. A radioisotope (⁵¹Cr) technique was used to monitor the recolonization behavior of U. formosa. Adult female mites preferentially re-entered A. imbecilis rather than the sympatric mussel A. cataracta. The specificity of this behavior parallels the distribution of this water mite among potential bivalve hosts in the southeastern U.S. Host recognition by U. formosa may contribute to re-establishing contact with a host after accidental separation and probably helps to maintain mite-mussel symbioses. Whether or not larval U. formosa employ similar host recognition behavior while selecting a potential host has not as yet been determined.     

Random amplified polymorphic DNA analysis of kinship within host-associated populations of the symbiotic water mite Unionicola foili (Acari: Unionicolidae)

Kinship relations within populations of unionicolid water mites are not well known, owing to their complex life cycles and the fact that interactions between active and resting stages for some species are transitory. A number of species of unionicolid water mites are, however, obligate symbionts of freshwater mussels and spend most of their life cycle in association with these hosts. Among these species of mites, parents and offspring are more likely to co-occur and thus provide opportunities to address questions related to the structure of the mating system. The present study employs random amplified polymorphic DNA (RAPD) analysis to address kinship within populations of Unionicola foili living in symbiotic association with the host mussel Utterbackia imbecillis. DNA was amplified from adult mites and a representative number of eggs or larvae (n = 20-30) that were removed from mussels collected on three separate occasions (July, November, and March) over a 12-month period. Parsimony analyses of the molecular data for adults and progeny collected from mussels during July, November, and March revealed distinct groupings, that for the most part, corresponded to mites collected from each of the sampling periods. Many of the genetic markers obtained for male and female U. foili were not evident among the larvae or eggs, suggesting that adults obtained from a host mussel at the time of collection were not the parents of a majority of the progeny. However, female mites and eggs collected from mussels during March and November shared more markers than did females and progeny examined during July. Furthermore, many offspring in the July sampling period were found to have one or more parents absent from the sampled population. Overall, RAPD profiling appears to have limited usage in determining kinship within populations of U. foili, due to its recruitment patterns, and the relatively large number of adults and progeny per mussel. It may, however, prove to be a useful method for assessing genetic relatedness among unionicolid mussel-mites that have substantially lower population densities.     

中国蚌螨属水螨1新种记述

记述了从我国浙江省淡水蚌中采到的蚌螨科Unionicolidae水螨1新种,壮肢蚌螨Unionicola（Anodontinatax）ischyropalpus sp．nov．,模式标本保存于南昌大学生物科学工程系。     

Competitive Exclusion,Harem Behaviour and Host Specificity of the Water Mite Unionicola ypsilophora (Hydrachnellae,Acari) Inhabiting Anodonta cygnea (Unionidae)

The co-occurring freshwater mussels Anodonta cygnea and A. anatina serve as hosts for the water mites Unionicola ypsilophora and U. intermedia, respectively. Male U. ypsilophora display a territorial behaviour. They fight with other males, and as a result, there is usually only one male per host. As a consequence, this intrasexual aggression results in female-defence polygyny, or a harem mating system. In contrast, U. intermedia shows no antagonistic behaviour between males. A. cygnea can serve as a host for U. intermedia, but this mite species apparently is excluded from the mussel by U. ypsilophora. In this way, U. intermedia is restricted by competitive exclusion to the mussel A. anatina.     

Host specificity among Unionicola spp. (Acari: Unionicolidae) parasitizing freshwater mussels

Water mites of Unionicola spp. are common parasites of freshwater mussels as adults, living on the gills, or mantle and foot of their hosts and using these tissues as sites of oviposition. The present study addresses specialization among North American Unionicola mussel-mites using 2 measures of host specificity: (1) the number of host species used by a species of mite; and (2) a measure that considers the taxonomic distinctness of the hosts utilized by mites, weighted for their prevalence in the different hosts. Results of this study indicate the Unionicola spp. mussel-mites are highly host specific, with most species occurring in association with 1 or 2 species of hosts. If 2 or more host species are utilized, they are typically members of the same genus. These data are consistent with studies examining the dispersal abilities and host recognition behavior for members of the group. When the average values of host specificity for Unionicola subgenera were mapped on a phylogenetic tree for these taxa, a clade comprised of gill mites appeared to be more host specific than a clade consisting of mantle mites. There were, however, no apparent patterns of host specificity within each of the clades. Differences in specificity between the 2 lineages may reflect either a long evolutionary history that gill mites have had with host mussels or the intense competition among gill mites for oviposition sites within unionid mussels, leading to increased host specialization.     

Genetic differences among host-associated populations of water mites (Acari: Unionicolidae: Unionicola): allozyme variation supports morphological differentiation

Unionicola poundsi and U. lasallei are recognized as closely related, morphologically distinct species of water mites living in symbiotic association with the mussels Villosa villosa and Uniomerus declivus, respectively. However, results of a transplant experiment suggested that the morphological characters used to separate these species are plastic and are influenced by the host species in which these mites metamorphose. These results indicate that U. poundsi and U. lasallei are variants of the same species. To test the validity of these contrasting notions, the genetic structure of mite populations from Uniomerus declivus and V. villosa was compared. An examination of allozyme variation at 9 enzyme loci revealed a high degree of genetic differentiation between these host-associated populations, with mites from U. declivus and V. villosa being fixed for different alleles at 3 loci and exhibiting significant allele heterogeneity at 71% of their polymorphic loci. Coefficients of genetic similarity and genetic distance for mites from U. declivus and V. villosa were 0.36 and 0.95, respectively. The results of this study suggest that mite populations from U. declivus and V. villosa are genetically distinct and complement morphological data recognizing them as valid species.     

分子鉴定背瘤丽蚌中弯弓蚌螨的隐藏种

Unionicola sp.与弯弓蚌螨U.arcuata在形态结构上非常相似,为了鉴别弯弓蚌螨U.arcuata 和Unionicola sp.,本研究扩增出Unionicola sp.12S rRNA的部分基因片段,与基因数据库中5种寄生蚌螨序列进行比较分析.结果显示,Unionicola sp.与弯弓蚌螨U.arcuata遗传差异为0.145,而分子系统树表明,Unionicola sp.与弯弓蚌螨U.arcuata聚为一支,具较近的亲缘关系.我们推断Unionicola sp.是弯弓蚌螨U.arcuata的1个隐藏种.这可能是由于不同寄主蚌造成了宿主隔离,从而引起了蚌螨遗传基因的分化.     

中国蚌螨属水螨一新种记述(蜱螨亚纲,蚌螨科)

记述了从我国江西省淡水蚌中采到的蚌螨科Unioncolidae水螨1新种,簇刺蚌螨U.(Anodontinatax)penicillatus sp.nov.,模式标本保存于南昌大学生物科学工程系.簇刺蚌螨,新种Unionicola(Anodontinatax)penicillatus sp.nov.(图1～11)正模♂,副模1♀,江西鄱阳湖,1998-11-11,文春根采自背角无齿蚌A.woodiana woodiana(Lea).鉴别特征新种近似于中间蚌螨U.(Anodontinatax)intermedia(Koenike,1882),但通过以下特征区别后者:雄螨具有背小板,雄螨Ⅳ-L-4-5的密刺毛数量为8～10和22～24(后者为18～20和17～19),射精复合体的形状和须肢的分叉腹小爪.词源:拉丁词"penicillatus"意思为"一簇毛的",指新种因雄螨Ⅳ-L-4-5有一簇刺毛而命名.     

The relations between mites of the genus Unionicola and the mussels Anodonta and Unio

The locations in mussels of four species of watermites, Unionicola aculeata, U. bonzi, U. intermedia and U. ypsilophora and their degrees of parasitism have been compared. Unionicola aculeata and U. bonzi do not live parasitic and need Anodonta anatina and Unio pictorum respectively as a shelter for their transformation stages. Unionicola intermedia and U. ypsilophora live parasitic on the gills of Anodonta anatina and A. cygnea respectively. Every female of Unionicola ypsilophora has a territory in the mussel, as contrasted with U. intermedia, many females of which are found together. The consequences for the population density are discussed.Zoological Laboratory, University of Amsterdam     

蚌螨属1新种记述（蜱螨亚纲：蚌螨科）

报道蚌螨属Unionicola一新种。     

Prey detection by the water mite Unionicola crassipes (Acari: Unionicolidae)

SUMMARY. 1. The importance of different senses in prey detection by the common water mite Unionicola carssipes was investigated.
2. Mechanoreception is likely to be the most important mode of detection; mites preferentially attacked vibrating over non-vibrating glass probes. The net-stance, a characteristic posture in which the mite raises its first two sets of legs from the substrate and orients toward vibrations in the water column, is assumed by U. crassipes when hunting.
3. Chemoreception may have a role in area-restricted search: mites exposed to prey-conditioned water were more sedentary than those exposed to unconditioned water. For a sit-and-wait predator like U. crassipes , remaining still in an area that contains prey will result in a greater predator/prey encounter rate. It is possible that contact chemoreception is used to determine the palatability of an already captured prey item, but mites did not attempt to capture prey they had touched unless the prey moved.
4. Vision is not necessary for prey capture; however, mites captured more prey in the light than in the dark. It is not clear whether this is because vision plays a part in prey detection or because prey behave differently in the two situations.     

Ultrastructural evidence for nutritional exchange between brooding unionid mussels and their glochidia larvae

Abstract. The life history of unionid bivalve molluscs includes retention of developing embryos within the gills of parental mussels. This brooding behavior may facilitate nutrient transfer to the glochidia larvae, i.e., matrotrophy. To address this possibility, morphological relationships between brood chambers and developing larvae of Pyganodon cataracta and Utterbackia imbecillis were examined with TEM, and larval shells were observed with SEM, for features that could be associated with the uptake of dissolved materials. Early in brooding, glochidia are enclosed in a vitelline membrane that physically contacts numerous cilia and microvilli of the epithelial cells lining the brood chamber (marsupium). The vitelline membrane subsequently disappears. Lamellar tissues of parental mussels initially have large deposits of glycogen that diminish during the course of brooding. Septa separating brood chambers from adjacent secondary water tubes have numerous mitochondria and microvilli, suggesting the potential for active transport of materials into or out of the marsupia. Since punctae (pores) in the larval shells become filled with an organic matrix early in brooding, they are unlikely to be involved in nutrient exchange. Ultrastructure of the brood chamber and physical contact between the parental mussel and larvae are consistent with a nutritive role for retention of glochidia in the marsupia.     

Unionicola (Pentatax) billieaehonore n. sp. a sponge-associated Hydracarina (Acari: Unionicolidae) from New Zealand

Abstract

Records of sponge-associated Hydracarina from the Southern Hemisphere are uncommon. Unionicola (Pentatax) billieaehonore n. sp. has been found in association with the freshwater sponge Ephydatia kakahuensis in Lake Rotoiti (North Island) and Lake Taupo in New Zealand. Differences in palpal characteristics and in setal patterns of the epimera distinguish this species from the holotype of Unionicola longiseta Walter 1915 which was reported from New Zealand by Schröder (1935).     

Population dynamics of the water mite Unionicola arcuata (Unionicolidae) in the freshwater bivalve Cristaria plicata (Unionidae) in Poyang Lake, eastern China

Population dynamics of the water mite Unionicola arcuata were investigated in the freshwater bivalve Cristaria plicata during the period from January to December 2002 in Poyang Lake, East China. A pattern of seasonal variation was observed, with prevalence and abundance peaking in early spring and autumn. The number of mites in individual hosts was significantly correlated with the size, but not with the sex, of bivalves. The change in infection level of mites on different infection sites in C. plicata was significant, with >58% of the mites found on the outer and inner gills, indicating that U. arcuata shows site preference.