Nest Sanitation in Social Spider Mites: Interspecific Differences in Defecation Behavior

Nest sanitation behavior is an important adaptation in nest‐building animals. The social spider mite Stigmaeopsis miscanthi (Saito) constructs woven nests on the leaf undersurface of perennial pampas grass (Miscanthus sinensis Anderss) and lives in the nests in large groups. Nest members of this species defecate at only one site inside the nest for nest sanitation, and the waste management is maintained by volatile chemical and tactile cues. Other members of the genus Stigmaeopsis Banks also construct woven nests, live in groups, and manage their fecal wastes, but the species differ in nest and group size, host plant, and social behavior. We investigated the details of waste management in four Stigmaeopsis spp. and found differences in the location of the defecation site and in the way each species recognizes this site. Like S. miscanthi, Stigmaeopsis longus defecates at only one site and uses volatile chemical and tactile cues, whereas Stigmaeopsis takahashii and Stigmaeopsis saharai both defecate at two sites and use only tactile cues. We discuss the reasons for the waste management differences within the genus, and the relationship between the means of waste management, nest and group size.     

‘Harem’ and ‘non-harem’ type mating systems in two species of subsocial spider mites (Acari,Tetranychidae)

The mating system of a subsocial spider mite, Schizotetranychus miscanthi Saitō, which is closely related to Schizotetranychus longus Saitō (the long seta form of Schizotetranychus celarius (Banks) is a synonym of the latter) was studied in comparison with that of S. longus. Comparisons between nesting patterns of the two related species, S. miscanthi and S. longus revealed a difference in distribution of males among nests. Although more than one male sometimes occurred in the large nests of S. miscanthi, most nests were occupied only by a single male. On the other hand, many nests of S. longus included several males. Behavioral experiments revealed that the male and females of S. miscanthi which cohabited in a nest defended their offspring from phytoseiid predators. Observations and a census of the nesting pattern in a wild population indicate that this is the second example of biparental defense and of a subsocial life-pattern in spider mites. Differences in mating systems were experimentally demonstrated in the two species. Only a single male of S. miscanthi survived in a nest, as a result of highly aggressive male-male combat, while two males of S. longus cohabited in a nest. The mating system of the former species is thus considered as harem polygyny, while that of the latter as scramble type polygyny. Furthermore, observations by video recording and scanning electron microscopy showed that the winning male in the male-male combat in S. miscanthi often preyed on the loser, suggesting cannibalism among them.     

Do Kin Selection and Intra-Sexual Selection Operate in Spider Mites?

Between the two subsocial spider mites, Schizotetranychus longus Saito and S. miscanthi Saito, a big difference exists in male reproductive behavior. The males of the former species have an extraordinarily mild relationship with conspecific males, whereas those of the latter species show mortal aggression against conspecific males. Field and experimental observations on the structure of mating populations showed that S. longus is under inbreeding conditions because of a lack of males in early spring, whereas S. miscanthi is under outbreeding conditions. Therefore, I hypothesized that the difference in male antagonism between the two species may reflect the difference in relatedness of males, that is, it has evolved by kin selection. The recent discovery of two clinal trends in male–male aggression in S. miscanthi provided evidence in favour of this hypothesis. Furthermore, a correlation analyses between experimentally evaluated male aggression and distribution patterns of males in the field indicated that the variation in male antagonism is actually reflected in field populations. Based on these studies, I discuss the solidity of the kin-selection hypothesis. Lastly I address how spider mites are fruitful model animals for conducting behavioral, ecological and genetic studies to understand the evolution of haplo-diploidy.     

Function of the web box as an anti-predator barrier in the spider mite,<Emphasis Type="Italic"> Schizotetranychus recki</Emphasis>

Spider mites inhabiting Sasa bamboo show considerable variation in traits believed to be the result of coevolution between predator and prey. In Schizotetranychus recki Ehara inhabiting the hairy leaves of a dwarf bamboo, Sasa senanensis, all quiescent stages, including eggs, appear within web boxes in the leaf hairs of their host plant, and this habit is thought to be a trait involved in predator avoidance. To test this hypothesis, the survival rates of S. recki eggs inside a web box and those freed of a web box were assessed in relation to six predator species that co-occur with the spider mite in the field. The results clearly show that the webbing behavior (web box) of S. recki has a function in avoiding five predator species. However, one predator species, Agistemus summersi Ehara, preyed more on eggs protected by a web box. This suggests that this predator species has a special trait to overcome the nest barriers.     

Egg-depositing behavior as a predator avoidance tactic of Yezonychus sapporensis Ehara (Acari, Tetranychidae)

Spider mites are very small phytophagous organisms living on plant leaves. They possess a diverse array of life types. Such diversity in life type has been supposed to be a result of coevolution between predator and prey, although there is little concrete evidence to support this concept. In Yezonychus sapporensis inhabiting the leaves of a dwarf bamboo, Sasa senanensis, all quiescent stages including eggs appear on the tips of the host leaf hairs during the summer season, and this habit is thought to be a kind of predator avoidance trait. To confirm this, the survival rates of Y. sapporensis eggs deposited on natural sites and experimentally manipulated sites were observed for eight co-occurring predator species. The results clearly showed that the oviposition behavior of Y. sapporensis has a function in avoiding predator attack, at least under experimental conditions. Received: October 26, 2000 / Accepted: December 19, 2000     

Protein differences detected by two-dimensional electrophoresis in the species complex of Schizotetranychus celarius (Banks)

The species complex of Schizotetranychus celarius (Banks) has been thought to involve three morphologically distinct species, S. celarius, Schizotetranychus longus Sait and Schizotetranychus miscanthi Sait. Furthermore, the former species includes at least three morphs which have not yet been examined taxonomically. In the present study, the protein differences of these species and morphs were examined using two-dimensional electrophoresis. In comparison of protein spot patterns between 11 populations of the S. celarius complex and Schizotetranychus recki Ehara, the former group appeared to be very different to S. recki. The phenogram made based on genetic distance suggested that the species complex includes at least two subgroups: S. miscanthi and S. longus. These results are consistent with the previous taxonomic study. On the other hand, the morphs which had been categorized as S. celarius did not form a distinct cluster.This study was supported by a Grant-in-Aid for Scientific Research from the Ministry of Education. Science and Cultures, Japan (No. 02640501) and from the Ministry of Agriculture, Forestry and Fisheries (Bio Cosmos Program, BCP-93-I-B-14), and it is the contribution No. E-163 of the Fruit Tree Research Station.     

Variation in social behavior within a spider mite genus, Stigmaeopsis (Acari: Tetranychidae)

The spider mites belonging to the genus Stigmaeopsis constructextremely dense oval woven roofs (web-nests) over depressionson the lower surface of host leaves and are known to have akind of sociality. The four species that occur on bamboo plantsin Japan show different nest areas. The nest area of Stigmaeopsislongus is the largest, followed by that of S. celarius, S. takahashii,and S. saharai in decreasing order. Smaller nests effectivelyprevent adults of several predator mite species from intruding.We hypothesized that variation in nest size reflects differentanti-intruder adaptations of this mite group in relation totheir sociality. The larger nest makers may adopt an alternativeanti-intruder strategy, namely, counterattack by a large group,so as to compensate for the disadvantage of large nests. S.longus and S. celarius adults effectively defended their largenests against potential predators, and the effects of nest defenseincreased with the number of individuals in a nest. S. takahashiiand S. saharai revealed no counterattack effect. Counterattackabilities that increase with the adult density, and thus, socialitymay compensate for the vulnerability of larger nests.     

Revegetation patterns and seedbank structure on abandoned pastures in northern Japan

To compare revegetation patterns between native and exotic species on abandoned pastures in northern Japan, we surveyed the vegetation structure and the seedbank using a flotation technique in 140 1 1 m plots. Previously introduced grasses such as Poa pratensis, Phleum pratense, and Dactylis glomerata were abundant 20 yr after the pasture abandonment, while Sasa senanensis, native shrub species, regenerated from propagation that had spread from the surrounding forests. S. senanensis shrublands and P. pratensis/P. pratense grasslands established on deep soils while D. glomerata grasslands established on shallow soils. Trees rarely became established on abandoned pastures. The seed density in the seedbank, representing 19 species, ranged from 542 to 2957 seeds/m². The dominant species in the vegetation (P. pratensis, Trifolium repens, and Rumex acetosella) were also common in the seedbank, whereas Cerastium holosteoides var. angustifolium, Chenopodium album var. centrorubrum, and Erigeron canadensis were widespread in the seedbank but did not occur in the extant vegetation. S. senanensis regenerated by vegetative propagation, and P. pratensis and P. pratense developed a seedbank. We concluded that for native species, particularly S. senanensis, vegetative reproduction has an important role on revegetation rather than regeneration from the seedbank, and the dwarf bamboo may be a keystone species in the ecosystems.     

Roles of dominant understory Sasa bamboo in carbon and nitrogen dynamics following canopy tree removal in a cool‐temperate forest in northern Japan

To clarify the role of dense understory vegetation in the stand structure, and in carbon (C) and nitrogen (N) dynamics of forest ecosystems with various conditions of overstory trees, we: (i) quantified the above‐ and below‐ground biomasses of understory dwarf bamboo (Sasa senanensis) at the old canopy‐gap area and the closed‐canopy area and compared the stand‐level biomasses of S. senanensis with that of overstory trees; (ii) determined the N leaching, soil respiration rates, fine‐root dynamics, plant area index (PAI) of S. senanensis, and soil temperature and moisture at the tree‐cut patches (cut) and the intact closed‐canopy patches (control). The biomass of S. senanensis in the canopy‐gap area was twice that at the closed‐canopy area. It equated to 12% of total biomass above ground but 41% below ground in the stand. The concentrations of NO₃^? and NH₄⁺ in the soil solution and soil respiration rates did not significantly change between cut and control plots, indicating that gap creation did not affect the C or N dynamics in the soil. Root‐length density and PAI of S. senanensis were significantly greater at the cut plots, suggesting the promotion of S. senanensis growth following tree cutting. The levels of soil temperature and soil moisture were not changed following tree cutting. These results show that S. senanensis is a key component species in this cool‐temperate forest ecosystem and plays significant roles in mitigating the loss of N and C from the soil following tree cutting by increasing its leaf and root biomass and stabilizing the soil environment.     

Development and characterization of polymorphic microsatellite DNA markers for Sasa senanensis (Poaceae: Bambuseae)

Sasa senanensis is a dwarf bamboo species distributed on the floors of cool temperate forests in Japan and adjacent regions. We isolated eight polymorphic microsatellite loci from this species. The number of alleles ranged from two to eight and the observed heterozygosity per locus from 0.13 to 0.74. Seven of the eight loci were also polymorphic in Sasa nipponica. Most of these markers were successfully amplified in other dwarf bamboo species. These markers will be useful for investigating clonal structure and population genetics in some dwarf bamboo species.     

Effects of Sika Deer (Cervus nippon) and Dwarf Bamboo (Sasa senanensis) on Trillium Populations in Akan National Park,Eastern Hokkaido,Japan

The competition between Trillium spp. and other species could modify the utility of Trillium spp. as indicators of the impact of deer on forest vegetation. To evaluate whether Trillium spp. (T. camschatcense and T. tschonoskii) are appropriate indicators of the impact of sika deer (Cervus nippon) in the presence of dwarf bamboo (Sasa senanensis), which competes with Trillium populations, we examined the size class of large‐growth‐stage Trillium plants, the frequency of the presence of deer pellets and the coverage of S. senanensis in deer‐excluded sites (where deer have been eliminated since 1995) and unfenced control sites in Akan National Park, eastern Hokkaido, Japan. The result of a cumulative link mixed model (CLMM) suggests that size class at deer‐excluded sites is influenced by competitive understory species, even without deer grazing. The CLMM result for unfenced control sites suggests that C. nippon negatively influences the size class of Trillium populations; however, the negative effect weakened with increasing S. senanensis coverage. Thus, S. senanensis can function as an unpalatable neighbor that makes it difficult for C. nippon to detect Trillium plants. We conclude that the size class structure of Trillium spp. can be a useful index of C. nippon usage under sparse or medium S. senanensis understories; however, it is difficult to use these indices under dense S. senanensis understories because of the weakened negative effect of C. nippon grazing.     

Variation in photoinhibition among Sasa senanensis, Quercus mongolica, and Acer mono in the understory of a deciduous broad-leaved forest exposed to canopy gaps caused by typhoons

To elucidate mechanisms for tolerating sudden increases in light intensity following canopy gap formation, we investigated susceptibility to photoinhibition in the evergreen clonal plant bamboo, Sasa senanensis, and two deciduous broadleaf woody plants, Quercus mongolica, and Acer mono. We measured pre-dawn photochemical efficiency of photosystem II (F _v /F _m) in plants exposed to canopy gaps and in shade-grown plants through the month following gap formation. Photoinhibition (indicated by decreased F _v /F _m) was smallest in S. senanensis and largest in A. mono. S. senanensis had the highest area-based net CO₂ assimilation rate (A _area) and electron transport rate (ETR) under high light conditions. This species also had the highest leaf mass per area (LMA) and leaf nitrogen content per area (N _area). Higher values of LMA and N _area under shade conditions probably contribute to circumvent photoinhibition through maintenance of a higher ETR capacity. Q. mongolica, a gap-dependent species, had properties intermediate between S. senanensis and A. mono; it appeared less susceptible to photoinhibition than the shade-tolerant A. mono. None of the species examined had increased photosynthetic capacity 1 month after gap formation, indicating that shade-grown leaves were unable to fully acclimate to increased light.     

Imaginal induction of diapause in several 'adult-female diapausing' spider mites

Abstract. The possibility of imaginal induction of diapause in adult female spider mites is suggested in Stigmaeopsis miscanthi. Here, several spider mites are studied to investigate the generality of this phenomenon. The existence of imaginal induction of reproductive diapause is established in Stigmaeopsis longus, two forms of S. miscanthi and Tetranychus kanzawai adult females. The females that develop under diapause-averting (long days and 18 °C) conditions are induced to diapause under short-daylength conditions at 18 °C after moulting to the adult stage. However, the adult females of Tetranychus urticae never diapause under these conditions.     

Aciculosporium sasicola sp. nov. on witches' broom ofSasa senanensis

A new species ofAciculosporium, A. sasicola (anam.Albomyces sasicola), is described and illustrated. The fungus differs fromAciculosporium take by its small asci and ascospores, though witches' broom and signs probably caused by the former onSasa senanensis are similar to those caused by the latter on bamboos.     

Early post-fire vegetation regeneration in <Emphasis Type="Italic">Larix kaempferi</Emphasis> artificial forests with an undergrowth of <Emphasis Type="Italic">Sasa senanensis</Emphasis>

Post-fire vegetation regeneration was studied for a 6-year period in a 13-year-old-artificial forest consisting of Larix kaempferi with a dense undergrowth of Sasa senanensis. The study site was classified into three fire severity categories according to the degree of Sasa senanensis scorching, that is, a high-severity category, a mid-severity category, and a low-severity category. Study plots were established in areas which fitted the criteria for each category, and in nearby unburned sites. A total of 41 woody species were newly emerged during the 6-year study period in the burned and unburned plots. Only a few seedlings and resprouts emerged in the unburned plots, while many seedlings emerged in the high-severity plots from the first year after fire onward. A high-severity fire that burns the rhizomes of Sasa is necessary for the vegetation recovery by germination of seed. Whereas the establishment of seedlings was restricted to a few years after fire, the regeneration through resprouting continued into the last year of observation. The survival time of resprouts was longer than that of seedlings, and the survival time of shade-tolerant species was longer than that of shade-intolerant species. In contrast, shade-intolerant species grew more rapidly than shade-tolerant species. The plants ability to exceed the maximum height of the Sasa before the bamboo recovers can be critical to the survival of shade-intolerant species. Because resprouts have a stronger resistance to the shade of Sasa than seedlings, the resprouts of shade-tolerant species play a major role in the re-establishment of woody species after fire in sites with considerable Sasa ground-cover.     

Microsite variation in light availability and photosynthesis in a cool-temperate deciduous broadleaf forest in central Japan

An empirical light simulation model was applied to estimate stand scale photosynthesis in a deciduous broadleaved forest in central Japan. Based on diurnal courses of photosynthetically active photon flux density (PPFD), we characterized the components of incoming light within the forest canopy, and found that the instantaneous relative PPFD (PPFD under the canopy relative to that above the canopy) under diffuse light condition was a reliable estimator of the intensity and duration of PPFD. We calculated the daily photosynthesis (A_day) for each PPFD class using photosynthesis–light response curves. Model simulated A_day were corroborated with the estimates obtained from the nearby CO₂ flux tower. The result demonstrated the potential of the light simulation model. The light use efficiency of two dominant species, Betula ermanii as overstory and Sasa senanensis as understory species, were then evaluated. At the forest understory, PPFD under 50 mol m^–2 s^–1 contributed to 77% of the sunshine duration on a completely clear day. Therefore, a higher apparent quantum yield for S. senanensis enhanced the utilization of low PPFD for photosynthesis. On the other hand, at the upper forest canopies, B. ermanii with a higher light-saturated photosynthetic rate used high PPFD efficiently. Consequently, potential of daily net photosynthesis for both B. ermanii and S. senanensis was high under each light condition. Such interspecific difference in the patterns of light utilization was suggested as one of factors allowing coexistence of the two species in the study forest.     

Vegetation succession at the abandoned Ogushi sulfur mine, central Japan

We surveyed plant community development at the abandoned Ogushi sulfur mine. We found seven communities dominated by the following respective species: Deschampsia flexuosa, Miscanthus sinensis, shrub willow, Gaultheria miqueliana–Betula ermanii, Sasa senanensis–Betula ermanii, willow–Betula ermanii, and Sasa kurilensis–Abies veitchii. We examined the succession of these communities, in which younger communities of low height and ground cover contained seedlings of the successive communities that were taller and had higher ground cover. To understand the development of these different communities, we surveyed damage from mining pollution and effects of immature soils formed by landslides. The average pH (H₂O) was 4.12, and aluminum concentrations were not sufficiently high to damage plant growth, except in areas where sulfur had been mined. The organic carbon and nitrogen content in soil samples were very low because of a delay in soil development caused by a large landslide in 1937. Hence, succession was positively correlated with the soil development stage. The delay in soil development after a large landslide influenced the seven successional steps of the plant communities, but mineral poisons at the abandoned Ogushi sulfur mine had no effect on succession.     

Bambusicolous fungi in Japan (4): a new combination, Astrosphaeriella aggregata

Melanopsamma aggregata, described on culms of Phyllostachys bambusoides in Japan, is transferred to the genus Astrosphaeriella, and a new combination, A. aggregata, is proposed. Phyllostachys heterocycla var. pubescens and Sasa kurilensis are newly recorded as host plants of this species. The species produces a Pleurophomopsis-like microconidial state in culture media.     

Genetic basis of woven nest size in subsocial spider mites

The variation in nest size of social spider mites of the genus Stigmaeopsis is assumed to correspond to their anti-predator strategy and to be a key aspect of their social organization and speciation. It is known that the length of the dorsal setae (sc1, 2nd propodosomal setae) correlates with the nest size. We conducted interspecies cross experiments to determine the heredity of sc1 length and nest size using two closely related species that build different sized nests, Stigmaeopsis saharai Saito et Mori and Stigmaeopsis takahashii Saito et Mori. A cross between a S. saharai female and a S. takahashii male produced several viable F₁ females. We measured sc1 length and the nest size of the F₁ females and then compared these values with those of their parent species. The sc1 length of F₁ females and the nest size constructed by these mites were intermediate with respect to the values of the two parent species. Therefore, the length of the sc1 and nest size are heritable. This result sheds light on the importance of considering the genetic basis for the variations in social organization.     

Counterattack success of a social spider mite against two predominant phytoseiid predator species

A survey was conducted of the predator fauna occurring in and around the nests of the two forms (LW: low male aggression and HG: high male aggression) of Stigmaeopsis miscanthi (Saito) that occur in Japan. Two phytoseiid species, Neoseiulus womersleyi (Schicha) and Typhlodromus bambusae Ehara predominated in S. miscanthi nests and their respective occurrence frequencies were the same in LW form nests as in HG form nests. We examined the counterattack success of S. miscanthi LW form males against these two phytoseiid predators. It was shown that while LW form male(s) could kill or effectively drive the larvae of both predator species out of their nests, there were no significant differences in the male counterattack success rate between 1-male and 2-male defended nests, or against the two predator species. On the other hand, there was a significant difference between the two predator species’ behavioral response to male-defended nests.