Seasonal aspects of the life cycles of pseudoscorpions (Arachnida, Pseudoscorpiones)

Analysis of available information on the seasonal features of life cycles in pseudoscorpions (Arachnida, Pseudoscorpiones) shows that in the temperate climate of Europe with distinct seasonality, the development of these peculiar arachnids can be either eurychronous (= homodynamic), with a poorly pronounced seasonal arrangement, or stenochronous (= heterodynamic), with a distinct seasonal arrangement. This is similar to some other arachnids, particularly spiders and harvestmen. Eurychronous pseudoscorpions are characterized by the approximately year-round development (quite often with winter activity in the whole or part of the population), an overlapping of consecutive generations, the presence of alternative development with or without the dormant state (at all the postembryonic life cycle stages), lack of brood chamber production by females, and the start of free-living life at the protonymphal stage. Stenochronous pseudoscorpions, on the contrary, possess clearly arranged (usually univoltine) development with overwintering deutonymphs and tritonymphs (more rarely adults), a clear separation of adjacent generations, the production of brood chambers where the regressive protonymphs develop until their molt into deutonymphs, and the start of free-living life at the deutonymphal stage. These two types of life cycles are exemplified by two pseudoscorpions from South England, namely Neobisium muscorum with eurychronous development, and Chthonius orthodactylus with stenochronous, univoltine development (Goddard, 1976). There is no correlation between the taxonomic position and the seasonal characters of life cycles in representatives of Neobisiidae, Chthoniidae, and Chernetidae. There is, instead, a close connection between the eco-physiological state of these arachnids and the type of their gas exchange (continuous, diffusive and non-cyclic in the active state, as opposed to discontinuous, with cyclic emission of CO₂ and uptake of O₂ in the dormant state). The latter information may be useful for distinguishing different kinds of dormancy (diapause and quiescence) in these arachnids.     

Dormant stages and their participation in adjustment and regulation of life cycles of harvestmen (Arachnida,Opiliones)

Information on seasonal adaptations in the life cycles of harvestmen (order Opiliones) summarized in this review reveals a great diversity of these arachnids with respect to duration, voltinism, and phenology of their life cycles, as well as to the number and ontogenetic position of the dormant stages required for survival during the winter season and also for synchronizing development with seasonal climate rhythms. Most harvestmen have stenochronous development with univoltine life cycles synchronized by arrest of development in hibernating eggs or (rarely) diapausing nymphs and adults. The number and arrangement of dormant stages represent stable species-specific traits, with some rare cases of interpopulation differences (e.g., in Phalangium opilio). Eurychronous harvestmen exhibit the year-round occurrence of main ontogenetic stages which show equal abilities for either active development or quiescence (depending upon the external factors). Two alternative types of development arrest are common in univoltine opilionids: (1) diapause induced at the early stage of embryogenesis and terminated during cooling (in Opilio parietinus and eleven other species) with transformation into postdiapause quiescence; (2) cold quiescence enforced by low temperatures at the last stage of egg development just before hatching (in Phalangium opilio and four other species). In conclusion, the systems of seasonal adaptations in Opiliones are compared with those in other arachnids, insects, and crustaceans. Some promising directions in the study of seasonal adaptations in opilionid life cycles are suggested.     

Population dynamics, distribution patterns and life cycles of Neobisium muscorum and Chthonius orthodactylus (Pseudoscorpiones: Arachnida)

The population dynamics, life cycles and distribution patterns of Neobisium muscorum (Leach) and Chthonius orthodactylus (Leach) were studied over a two year period in a Chiltern beechwood site. The data obtained for the two species are compared, and the importance of pseudoscorpions in the beechwood community is discussed.     

Karyotype analysis and achiasmatic meiosis in pseudoscorpions of the family Chthoniidae (Arachnida: Pseudoscorpiones)

Karyotypes of pseudoscorpions (Arachnida, Pseudoscorpiones) are largely unknown. Here we describe for the first time karyotypes of the suborder Epiocheirata, represented by 9 European species of two genera of Chthoniidae, Chthonius and Mundochthonius. Diploid chromosome numbers of males range from 21 to 37. Karyotypes of both genera differ substantially. Acrocentric chromosomes predominate in karyotypes of the genus Chthonius, whereas M. styriacus exhibits a predominance of metacentric chromosomes. These differences suggest that the two genera belong probably to distant branches of the family Chthoniidae. It is proposed that karyotype evolution of the genus Chthonius was characterised by a reduction of chromosome numbers by tandem and centric fusions as well as gradual conversion of acrocentric chromosomes to biarmed ones, mostly by pericentric inversions. A tendency towards reduced chromosome numbers is evident in the subgenus Ephippiochthonius. All species display X0 sex chromosome system that is probably ancestral in pseudoscorpions. The X chromosome exhibits conservative morphology. It is metacentric in all species examined, and in the majority of them, a subterminal secondary constriction was found at one of its arms. In contrast to chthoniids, secondary constriction was not reported on sex chromosomes of other pseudoscorpions. Analysis of prophase I chromosomes in males revealed an achiasmatic mode of meiosis. Findings of the achiasmatic meiosis in both genera, Chthonius and Mundochthonius, indicate that this mode of meiosis might be characteristic of the family Chthoniidae. Amongst arachnids, achiasmatic meiosis has only been described in some scorpions, acariform mites, and spiders.     

Analysis of the Stridulation in Solifuges (Arachnida: Solifugae)

Stridulation in solifuges has not been investigated yet. We performed a comparative analysis of the stridulatory organs and sounds produced by juveniles of various developmental stages and adults (both sexes) of Galeodes caspius subfuscus Birula. The stridulatory organ is of similar morphology in all developmental stages. The sound that they produced was a broad frequency hissing, composed of one or two chirps with maximum at 2.4 kHz. The intensity of the sound was found to increase with body size. Otherwise, no differences were observed between stridulation in juvenile, male and female individuals. Therefore, we suggest that the stridulation in solifuges has primarily a defensive role. As solifuges are neither venomous nor unpalatable, they might imitate an accoustically aposematic organism that shares the same habitat and has similar circadian activity, e.g. vipers. It may also have an intraspecific function in reduction of cannibalistic tendencies.     

Species richness and biogeography of non-acarine arachnids in Namibia

A total of 821 species, 296 genera, and 69 families of non-acarine arachnids (Araneae, Solifugae, Scorpiones, Pseudoscorpiones, Opiliones and Amblypygi) are presently known from Namibia. Patterns of spider, solifuge and scorpion species richness and endemism are summarized relative to the 14 major Namibian types of vegetation. Spiders are most speciose in the higher rainfall areas, solifuges in the dry areas, and scorpions in rocky areas. Namibia probably has the world's greatest diversity of solifuges. Many arachnids endemic to Namibia are associated with the Namib Desert sand dunes.     

烟蚜茧蜂寄生对烟蚜生长发育和繁殖的影响

【目的】为探究烟蚜茧蜂Aphidius gifuensis对烟蚜Myzus persicae寄生后烟蚜生长发育和繁殖影响及其在烟蚜防控中的作用。【方法】采用室内叶片培养皿法,测定了不同龄期烟蚜(1-4龄若蚜及成蚜)被烟蚜茧蜂寄生后烟蚜当代(F₀)及后代(F₁)的发育历期、寿命、产仔量。【结果】烟蚜1龄若蚜被烟蚜茧蜂寄生后,1龄若蚜历期受到显著影响,较未被寄生对照缩短了0.65 d (33.00%),而2-4龄若蚜期和成蚜历期未受显著影响;2龄若蚜被寄生后,4龄若蚜和成蚜历期均受到显著影响,较未被寄生的对照分别缩短了1.50 d (100.00%)和0.83 d (100.00%),而2和3龄若蚜期未受显著影响;3龄若蚜被寄生后,4龄若蚜和成蚜期均受到显著影响,较未被寄生对照分别缩短了1.50 d (100.00%)和0.83 d (100.00%),而3龄若蚜期未受显著影响。2龄若蚜被寄生后烟蚜的寿命和产仔量均受到显著影响,寿命较未被寄生对照缩短了8.39 d(54.00%),每雌产仔量较未被寄生对照减少了48.67头(100.00%)。此外,2-3龄若蚜被寄生后其后代(F₁)1龄若蚜历期受影响最大,较未被寄生烟蚜后代1龄若蚜期缩短了0.80 d(40.00%),其F₁代平均寿命较未被寄生烟蚜后代平均寿命延长了5.72 d(41.20%),但其产仔量与对照差异不显著。【结论】不同龄期烟蚜被烟蚜茧蜂寄生后,其发育历期和寿命比未被寄生的缩短,产仔量比被未寄生的少,但其F₁代平均寿命比未被寄生后代烟蚜的长且其产仔量未受显著影响。当代2龄若蚜期是控制烟蚜增长的关键时期。这些结果对确定烟蚜茧蜂防治烟蚜适期及认识其机理具有重要参考意义。     

Ontogenetic variation in the sensory structures on the pedipalps of cosmetid harvestmen (Arachnida,Opiliones, Laniatores)

In arachnids, pedipalps are highly variable appendages that may be used in feeding, courtship, defense, and agonistic encounters. In cosmetid harvestmen, adults have pedipalps that feature flattened femora, spoon‐shaped tibiae, and robust tarsal claws. In contrast, the pedipalps of nymphs are elongate with cylindrical podomeres and are adorned with delicate pretarsi. In this study, we used scanning electron microscopy to examine the distribution of cuticular structures (e.g., sensilla chaetica, pores) on the elements of the pedipalps of adults and nymphs of three species of cosmetid harvestmen. Our results indicate that there is considerable ontogenetic variation in the morphology of the trochanter, femur, patella, tibia, and tarsus. The pretarsus of the nymph has a ventral patch of setae that is absent from the adult tarsal claw. We observed this structure on all three cosmetid species as well as on the pedipalps of an additional seven morphospecies of nymphs collected in Belize and Costa Rica. This structure may represent a previously unrecognized autapomorphy for Cosmetidae. Examinations of the pedipalps of antepenultimate nymphs of additional gonyleptoidean harvestmen representing the families Ampycidae, Cranaidae, Manaosbiidae, and Stygnidae revealed the occurrence of unusual, plumose tarsal setae, but no setal patches on the tarsal claw.     

Reinterpretation of Dracochela deprehendor (Arachnida: Pseudoscorpiones) as a stem‐group pseudoscorpion

Abstract: The middle Devonian (Givetian–Eifelian) pseudoscorpion Dracochela deprehendor Schawaller, Shear and Bonamo is redescribed from the type material and an additional palpal fragment. Dracochela differs from extant pseudoscorpions in having numerous spinules on the leg tarsi, the femur at least as long as the patella on the posterior legs, the stem of the arolia thick, most blades of the serrulae only weakly fused and in lacking a spinneret on the chelicera. The blades of the cheliceral rallum are shown to have been arranged in two rows, as in most Heterosphyronida. The cheliceral serrulae are compared with analogous structures in other arachnids (Notostigmata, Opiliones, Palpigradi, Schizomida and Scorpiones), and it is concluded that the panctenal state (all lamellae attached to finger) is plesiomorphic relative to the hemictenal state (apical lamellae raised), which has evolved independently in Heterosphyronida and Neobisioidea. The trichobothriotaxy of the chela of Dracochela is shown to be similar to that of the extant family Pseudotyrannochthoniidae. The growth of the chelal fingers followed the same pattern as that seen in modern pseudoscorpions, with most of the increase in length occurring at the base of the fingers. The family Dracochelidae Schawaller, Shear and Bonamo is treated as a plesion and assigned to the stem‐group of Pseudoscorpiones. The ordinal name Chelonethi Thorell is restricted to crown‐group pseudoscorpions, and the superordinal name Pseudoscorpiones Latreille is adopted for the total‐group (i.e. stem‐group plus Chelonethi).     

Ultrastructure of the Geogarypus nigrimanus Spermatozoon (Arachnida,Pseudoscorpionida)

The ultrastructure of the spermatozoon of Geogarypus nigrimanus (Arachnida, Pseudoscorpionida) is described. The spermatozoon is composed of a small elliptic nucleus, a short flagellum and a very long and complex acrosome. In the male genital ducts, as in other studied species of pseudoscorpions, the sperm components are rolled up to form a globular structure enclosed in a cyst wall. The Geogarypus spermatozoon with a reduced flagellum and a giant acrosome seems to be evolutionary more advanced than spermatozoa from other pseudoscorpions.     

The role of parthenogenesis in the biology of two species of mayfly (Ephemeroptera)

1. From a single founder virgin female of each species, nine and seven successive parthenogenetic generations of Cloeon dipterum and Cloeon simile, respectively, were reared: all offspring were female. Eggs were removed from final instar nymphs, subimagos and virgin imagos of each generation and their development assessed. The life cycles of 487 individual C. dipterum and 315 C. simile were recorded, together with details of growth rate, egg production and adult size of these and other representatives of each generation. In order to distinguish between the effects of parthenogenesis and the effects of culture conditions, fertilized eggs of both species were reared under the same laboratory conditions. 2. Fewer unfertilized than fertilized eggs hatched. There was no consistent trend in the percentage hatch in successive parthenogenetic generations. In both species the time taken for parthenogenetic individuals to complete a generation varied considerably, and no relationship was found between generation length and the number of parthenogenetic generations that had passed, the developmental stage of the donor, or the time taken for the eggs to hatch. After 50 weeks there was, in both species, an overlap of up to six generations. The number of eggs produced by both species was significantly greater in the first parthenogenetic generation than in later generations but the numbers for later generations lie within the recorded range of egg number in imagos captured in the field. 3. To find if parthenogenetic individuals occur in the field, the sex ratios of last instar nymphs and subimagos of C. simile were measured in samples collected over 13 successive years from a set of small isolated pools. There was a female bias both in winter and at the start of the emergence season. It is possible that, when mature nymphs and emerging subimagos are damaged by predators, the eggs which are released from them survive and develop. 4. Mature, egg-containing nymphs of both species were present in the field throughout the year, although their numbers were small at the beginning of winter. 5. Oviposition by imagos captured from mating swarms and by virgin imagos, of both species, was recorded for the first time, showing that C. dipterum is not always ovoviviparous. It is suggested that early instar nymphs occurring in mid-winter come not only from unfertilized eggs released from injured nymphs and subimagos, but also from fertilized eggs laid in autumn.     

Comparative assessment of feeding damage by pod-sucking bugs (Heteroptera: Coreoidea) associated with cowpea, Vigna unguiculata ssp. unguiculata in Nigeria

Feeding trials were conducted on three (young, mid-fill and mature) developmental stages of cowpea Vigna unguiculata ssp. unguiculata pods in the screenhouse using fourth instar nymphs and adults of Anoplocnemis curvipes (Fabricius), Riptortus dentipes (Fabricius), Mirperus jaculus (Thunberg), Clavigralla tomentosicollis St?l and C. shadabi Dolling. Anoplocnemis curvipes was observed to be the most damaging coreoid species causing a yield reduction of 26.4-51.7% followed by R. dentipes (24.4-29.4%), M. jaculus (21.9-26.9%), C. tomentosicollis (17.9-22.4%) and C. shadabi (15.9-20.4%). The fourth instar nymphs of each pod-sucking bug species caused a significantly higher cowpea yield reduction than their respective adults. Similarly, infestation on young pods compared to mid-fill and mature stages resulted in significantly higher yield reduction. The results suggest that infestation levels of two fourth instar nymphs of A. curvipes or three fourth instar nymphs of the other four pod-sucking bug species per young pod should be adequate for screening of cowpea varieties for resistance to the coreoid bugs.     

中国伪蝎名录（蛛形纲：伪蝎目）

本文根据我国伪蝎目已有种类记述,归纳并列出10科34属73种（包括亚种）伪蝎的中文名、拉丁学名、引证及其地理分布,以供今后研究参考。     

Effects of photoperiod,temperature and melatonin on nymphal development,polyphenism and reproduction in Halyomorpha halys (Heteroptera: Pentatomidae)

Influences of photoperiod, temperature and melatonin were examined on development and color patterns of nymphs, and sternum color and reproductive fate of adults in Halyomorpha halys (brevis). Short-day accelerated nymphal development, whereas long-day accelerated reproductive maturation. The two types of photoperiodic responses at different stages may help maintain the univoltinism of this species in the field, assuring the right timing for diapause and reproduction. The pronotum of fifth instar nymphs reared under LD 11:13 shows a brown-marbled color pattern darker, with less creamy-yellowish speckles, than that of nymphs reared under LD 16:8. Short-day-reared fifth instar nymphs, which are destined to diapause in adults, had shorter white stripes on the pronotum, smaller body size, less frequent feeding and more lipid accumulation than the long-day-reared. The longer the exposure to LD 13:11 at 20 degrees C during the nymphal stage, the greater the expression of short-day associated characteristics observed in the fifth nymphal instar and adult stage. Melatonin orally administered (500 microg/ml) to insects under LD 16:8 produced slightly reduced body size, suppressed feeding, extended nymphal period and increased accumulation of lipid compared to the untreated bugs, but did not affect the incidence of diapause in females. In males, melatonin retarded gonadal development under LD16:8. These results may suggest a possible role of melatonin for the control of seasonal polyphenism and development.     

Seasonal patterns of abundance and host relationships of the Australian paralysis tick,Ixodes holocyclus Neumann (Acarina: Ixodidae), in southeastern Queensland

Estimates of seasonal abundance of larvae, nymphs and females of the paralysis tick, Ixodes holocyclus, were obtained by collecting ticks that engorged on small mammals and birds trapped in two localities in southeastern Queensland: Brisbane (wet sclerophyll forest) and Tamborine Mountain (cleared rain forest). The long-tailed short-nosed bandicoot, Isoodon macrourus, was the most common mammal trapped but small numbers of other marsupials, rodents and ground frequenting birds were also captured. Small numbers of five other tick species were also collected. In both habitats there was clearly one dominant generation of I. holocyclus;7er year, although the presence of all stages at most times of the year indicated overlapping of smaller cohorts. Females were most abundant in spring and early summer, larvae in summer-autumn, and nymphs in autumn-winter. I. holocyclus was abundant on I. macrourus and rare on most other mammals and birds captured. At the peak of abundance of each instar, each bandicoot dropped from 500 to 2000 engorged larvae, 100 to 200 engorged nymphs, and four to six engorged females. Life tables were compiled for the tick in both habitats and these indicate that there was relatively high survival from engorged larva to engorged nymph and thence to engorged female and that most mortality occurred between detaching of the engorged female and the detaching of the engorged larva. The tick was more abundant on bandicoots from cleared rain forest and rain forest edge, than on those from sclerophyll woodland. The survival of engorged larvae and nymphs of I. holocyclus and the larval productivity of engorged females were examined in a warm moist climate where the tick was abundant (Tamborine Mountain) and in a hotter dryer climate where the tick was rare or absent (Amberley). In both localities engorged larvae and nymphs survived to the next instar in all seasons of the year. On most occasions engorged females produced eggs which hatched. Mature bandicoots from tick infested areas showed little or no resistance to infestation with larvae or nymphs of I. holocyclus, whereas other small mammals from the same area showed an appreciable degree of resistance to the immature stages of the tick. Feeding larvae and nymphs exposed to normal light-dark cycles in the laboratory detached during the afternoon and early evening. This behaviour and host resistance are discussed in relation to the daily activity cycles of host species, their habitat preferences, and their role as hosts for I. holocyclus.     

Sex pheromone of the larch caterpillar moth, Dendrolimus superans, from northeastern China

A study was set up to determine the sources and rates of mortality of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae) on field‐grown cassava in Uganda. Using a cohort‐based approach, daily direct observations were used to construct partial life tables for 12 generations of egg and nymph populations which were studied over a 1‐year period. Mortality was categorized as dislodgement, predation, parasitism (for nymphs only), unknown death, and inviability (for eggs only). The highest mean rate of marginal mortality across all stages was attributed to parasitism, with dislodgement and predation following, respectively. Across all factors, the highest mean rate of marginal mortality was observed in the fourth instar followed by the eggs, first‐, second‐, and third‐instars, respectively. Key factor analysis revealed that dislodgement was the major mortality factor contributing to generational mortality in eggs while for nymphs, parasitism in the fourth instar was the main driving force behind the observed generational mortality. Highest irreplaceable mortality in both the egg and nymph stages was attributed to dislodgement followed by parasitism and predation, and least was due to unknown death. Across stages, highest irreplaceable mortality rates were observed in the eggs and the fourth‐instar nymphs. The other stages had relatively low rates of irreplaceable mortality. Rain‐protection experiments revealed no significant differences in marginal mortality rates when compared to the open field situation.     

Thermal requirements for the development and reproduction of Nysius huttoni White (Heteroptera: Lygaeidae)

Nysius huttoni White is an economically important pest of wheat and brassica crops in New Zealand. Because of its frequent presence in export fruit packages, it is also considered an important quarantine pest to countries that trade with New Zealand. To provide critical information for the pest risk analysis, forecast and management of N. huttoni, we investigated the effect of five consistent temperatures (10, 15, 20, 25, and 30 degrees C) on its development, survival and reproduction. At 10 degrees C both eggs and nymphs did not develop but the latter grew. Nymphs could survive 10 degrees C for >1.5 mo, with the fifth instar nymphs surviving for up to 145 d. Adults could live for at least 100 d at this temperature. This species could not complete its lifecycle at or below 15 degrees C. Between 15 and 30 degrees C, fifth instar stage was significantly longer than other nymphal stages. Egg hatch rate and total survival rate for all stages were significantly higher at 20 degrees C than at other test temperatures. The developmental rate of different life stages increased linearly with the increase of temperatures from 15 to 30 degrees C. The estimated low temperature threshold for the completion of lifecycle was 11.9 degrees C, and that for mating and oviposition was 12.3 and 16.8 degrees C, respectively. The thermal requirement for completing a life cycle of N. huttoni was 625 DD. The time needed for completing a life cycle was similar for both sexes. Temperature had little effect on adult body weight and sex ratio. Implications of the above findings are discussed.     

九香虫形态及生物学特征研究

通过建立网室近自然实验种群,结合野生个体观察记述了云南玉溪九香虫Coridius chinensis (Dallas)种群的各虫期形态特征、在云南玉溪红塔区的年生活史、寄主范围、取食偏好、成虫及若虫的行为习性,为该虫的保育和规模化养殖提供依据。结果显示,九香虫卵有绿色和粉色两种生态型,若虫有5个龄期;在云南玉溪红塔区1年仅发生1代,仅以成虫态越冬;红塔区九香虫野生个体主要以佛手瓜和南瓜等葫芦科植物为食;取食偏好选择试验表明该虫最喜取食葫芦,冬瓜和南瓜次之;雌雄成虫可多次交配多次产卵,单雌单次平均产卵量为16.3±6.0粒。研究表明,云南玉溪红塔区的九香虫种群为典型的一化性昆虫,休眠期可达8个月;该虫食性较广,可取食多科植物,但以葫芦科瓜类为主;羽化成虫每年9月中旬至10月上旬陆续从佛手瓜、南瓜等寄主植物上迁至枯树皮下、柴堆等隐蔽场所越冬,翌年4月下旬陆续解除休眠,迁至佛手瓜、南瓜等作物上取食繁殖。