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.     

Seasonal cycles of noctuid moths of the subfamily Plusiinae (Lepidoptera,Noctuidae) of the Palaearctic: Diversity and environmental control

Analysis of data on seasonal development of noctuid moths of the subfamily Plusiinae shows that the control of their seasonal cycles is poorly understood. At the same time, the available data demonstrate considerable diversity of the seasonal patterns of Plusiinae species from different regions. The homodynamic type of seasonal development has been found in Trichoplusia ni and Ctenoplusia agnata of the tribe Argyrogrammatini and in Autographa gamma of the Plusiini. The seasonal development of these southern noctuids is accompanied by regular interzonal migrations of flying adults. When spreading northwards, they can produce a different number of annual generations, depending on the local climatic conditions, and establish temporary local populations whose longevity is limited by the available thermal resources. Adults of some species may fly back southwards, but it is more likely that individuals from temporary local populations cannot survive long winters and are destined to die. The heterodynamic type of seasonal cycles allows insects to survive in the regions with pronounced seasonality of climate. This type of seasonal development includes univoltine, multivoltine, and semivoltine seasonal cycles. Univoltine seasonal cycles with obligate diapause are known in Autographa buraetica, A. excelsa, and Syngrapha ain (Plusiini). Diapause provides tolerance to both low temperatures and a prolonged period when food is unavailable. In Syngrapha ottolenguii (Plusiini), the same result is achieved by inclusion of two photoperiodically controlled diapauses (winter larval and summer adult ones) into the life cycle. The semivoltine seasonal cycle has been reported in only one species of Plusiinae, namely Syngrapha devergens. Larvae of this moth overwinter twice before pupation. Multivoltinism is common in the tribe Plusiini. Depending on the latitude, different species of this tribe can produce up to four generations per year and overwinter as middle-instar larvae in the state of facultative diapause. However, the characteristics of diapause vary substantially between the species: diapause can be deep and stable (as in Diachrysia chrysitis, Plusiini) or unstable and thus not ensuring successful overwintering and steady population growth (as in Macdunnoughia confusa, Plusiini). The seasonal adaptations known in Plusiinae include migrations, winter and summer diapauses, photoperiodic control of larval growth rates, and seasonal polyphenism of larval body coloration. In general, seasonal adaptations of Plusiinae are determined by local environmental conditions and only loosely associated with the systematic position of particular taxa. Only the tribe Abrostolini stands apart from other taxa of Plusiinae: moths of this tribe differ not only in morphology but also in peculiarities of their seasonal development, because all the species of this tribe overwinter as pupae and their seasonal cycles are therefore different from those of the rest of Plusiinae.     

Harvestman (Opiliones) fauna associated with Maine lowbush blueberry fields in the major production areas of Washington and Hancock counties

Over a period of 19 yr, the harvestman (Opiliones) community associated with the lowbush blueberry agro-ecosystem in Maine was studied. Eight species representing five genera, four subfamilies, and two families of harvestmen belonging to the suborder Eupnoi were collected. The harvestman community was dominated by two introduced, synanthropic species: Phalangium opilio in all but 1 yr (that year dominated by Rilaena triangularis). Rilaena was recorded for the first time from eastern North America. Relative abundance of harvestman adults increases throughout the season and the temporal pattern of trap capture does not refute speculated life cycles of the harvestmen being univoltine with overwintering eggs. Some blueberry management practices were found to affect trap capture. We did find that on average (with opposite results 1 yr) trap captures are greater in pruned fields than in fruit-bearing fields. Organic fields were found to have higher relative abundance of harvestmen than conventionally managed fields. Conventionally managed fields with reduced-risk insecticides showed no difference in harvestmen relative abundance compared with those conventionally managed fields using the older more persistent organophosphate insecticides. Insecticide trials with common insecticides used in blueberry insect pest management showed that the organophosphate insecticide, phosmet, and the pyrethroid insecticide, esfenvalerate, were detrimental to P. opilio adults when exposed to leaf residues, whereas the reduced-risk insecticide, spinosad, showed no negative effects compared with nonsprayed foliage.     

Histologic Structure of the Harvestman Leiobunum japonicum (Arachnida: Opiliones: Sclerosomatidae)

Opiliones are an order of arachnids commonly known as harvestmen. Although they belong to the class of arachnids, harvestmen are not spiders. Harvestmen are well known for their exceptionally long legs compared to body size, and the made up of two main parts, cephalothorax and abdomen. Although over 6,400 species of harvestmen have been discovered worldwide, the research on these arachnids is still an unexplored field comparing to spiders or insects. The Harvestmen is the Arachnids disregarded to Korean researchers. Therefore, in Korea, it doesn't become nearly the investigation of the harvestmen. Histological data about the harvestmen tries to be provided through the research of the internal structure of the harvestmen. The study of structures of the opiliones was made with the histological image and it focused mainly on the eye, leg, and genital organs.     

Seasonal aspects of the life cycle of solifuges (Arachnida, Solifugae) as compared with pseudoscorpions (Arachnida, Pseudoscorpiones)

Solifuges (order Solifugae) and pseudoscorpions (order Pseudoscorpiones) united into the superorder Haplocnemata (Shultz, 2007) are nevertheless characterized by essential differences both in morphological and biological characters. Analysis of available information on the biology and life cycles of these arachnids revealed a clear difference between the daily rhythms of activity: their presence in solifuges and their absence in pseudoscorpions. However, this concerning the seasonal adaptations in the two orders is not simple since they demonstrate not only differences but also a lot of similarities. All the studied solifuges are characterized by the seasonally timed stenochronous (heterodynamic) type of development which is characteristic of species with uni-, bi-, and semi-voltine development (i.e., to life cycles completed within a year, half a year, and several years), as well as to species combining different forms of voltinism. This type of development is not only prevalent in solifuges (as in pseudoscorpions and other arachnids) but appears to be the only one, since no cases of eurychronous (homodynamic) development have been found in solifuges; whereas pseudoscorpions and other arachnids develop both steno- and eurychronously. The initial ontogenetic stages remain in underground shelters (brood burrows in solifuges and brood chambers in pseudoscorpions). The first nymphal stages (I instar nymphs in solifuges, protonymphs in pseudoscorpions) are embryonized; active life outside the brood burrows starts with II instar nymphs in solifuges and with deutonymphs in pseudoscorpions.     

Dynamics of snow crab (<Emphasis Type="Italic">Chionoecetes opilio</Emphasis>) movement and migration along the Newfoundland and Labrador and Eastern Barents Sea continental shelves

This study uses survey and tagging data and cluster analysis to interpret dynamics of ontogenetic movements and seasonal migrations in snow crab (Chionoecetes opilio) along the Newfoundland and Labrador (NL) continental shelves. Most historic literature from Atlantic Canada suggests snow crab undertake small-scale ontogenetic movements while observations of seasonal migrations had been near-exclusive to the inshore. Information on both types of movement in the most spatially expansive offshore region of Atlantic Canada, in NL, was lacking. We find that that both ontogenetic movements and seasonal migrations occur in most areas of the NL offshore, with ontogenetic movements generally down-slope and seasonal migrations generally up-slope. Conservative estimates of average ontogenetic movements range from 54 to 72 km for both males and females in the largest offshore regions while seasonal migrations are slightly smaller, with two independent studies on the Grand Bank producing average estimates of 43–46 km and an adjacent tagging study in a smaller inshore bay producing an average estimate of 25 km. Ontogenetic movements appear associated with a search for warm water while seasonal migrations appear associated with both mating and molting in shallow water. On average, morphometrically mature crab of both sexes move less vertical distance than morphometrically immature crab during seasonal migrations. We investigate plausible explanations for ontogenetic movements and spring migrations and detail how bottom temperature affects crab distribution and life history dynamics. We further document movement patterns from tagging studies on the burgeoning snow crab stock in the Eastern Barents Sea toward establishing consistencies in species behaviour on the global scale. Finally, we discuss explanations for historical disparities in the literature between scales of movement for snow crab in the Eastern Bering Sea of Alaska versus Atlantic Canada and advance perspectives on life history theory for the species.     

Change of carotenoid composition in crabs during embryogenesis

Changes of the qualitative and quantitative compositions of carotenoids are studied at various development stages of the external hard roe, determined based on color differences, for the species C. opilio, P. camtschaticus, and P. platypus. It has been revealed that the major carotenoids of the new egg are astaxanthin and β-carotene. Intermediate products of transformation of β-carotene into astaxanthin are identified: echinenone, canthaxanthine, and phenicoxanthine. The carotenoid content per embryo for the new hard roe of C. opilio (the orange egg) amounted to 22.7 ng, of P. camtschaticus and P. platypus (the violet egg)—to 49.2 and 23.3 ng, respectively. In the hard roe at the later development stage (the brown egg) the carotenoid content was decreased to 13.1 ng in C. opilio and to 20.1 ng in P. camtschaticus. Development of embryos is accompanied by accumulation of esterified carotenoids and a decrease of β-carotene and astaxanthine concentrations in all studied species.     

Nutrient availability reduced in older rural impoundments in coastal Bay of Fundy,Canada

The caddisfly species Micropterna lateralis is an abundant representative of limnephilids in intermittent streams. Yet, its basic life history characteristics and adaptations related to environmental factors, such as stream drying, are comparatively understudied. Here, we investigated larval growth and metabolic energy reserves (glycogen, triglycerides) through development in their natural habitat. We concentrated on the larval development because this period represents the important phase of energy accumulation necessary for growth, metamorphosis and embryogenesis. Besides larval physiology, female adults were studied in terms of ovarian maturation. Our results indicate that adult females lack an imaginal diapause, which is otherwise often observed in intermittent stream-inhabiting Limnephilidae. Further, M. lateralis is univoltine and exhibits a relatively fast larval development with five distinct instars, of which four are characterised here (instars II–V). Accrual of biomass occurs in final instars, where a high amount of glycogen is accumulated. Lipid concentrations, on the other hand, are kept constant in final stages and slightly lower than in preceding instars. This dominance of glycogen in final instars found in M. lateralis is highly unusual in insects and of potential adaptive significance for the species’ ability to exploit intermittent habitats.     

Susceptibility to disease varies with ontogeny and immunocompetence in a threatened amphibian

Ontogenetic changes in disease susceptibility have been demonstrated in many vertebrate taxa, as immature immune systems and limited prior exposure to pathogens can place less developed juveniles at a greater disease risk. By causing the disease chytridiomycosis, Batrachochytrium dendrobatidis (Bd) infection has led to the decline of many amphibian species. Despite increasing knowledge on how Bd varies in its effects among species, little is known on the interaction between susceptibility and development within host species. We compared the ontogenetic susceptibility of post-metamorphic green and golden bell frogs Litoria aurea to chytridiomycosis by simultaneously measuring three host-pathogen responses as indicators of the development of the fungus—infection load, survival rate, and host immunocompetence—following Bd exposure in three life stages (recently metamorphosed juveniles, subadults, adults) over 95 days. Frogs exposed to Bd as recently metamorphosed juveniles acquired higher infection loads and experienced lower immune function and lower survivorship than subadults and adults, indicating an ontogenetic decline in chytridiomycosis susceptibility. By corresponding with an intrinsic developmental maturation in immunocompetence seen in uninfected frogs, we suggest these developmental changes in host susceptibility in L. aurea may be immune mediated. Consequently, the physiological relationship between ontogeny and immunity may affect host population structure and demography through variation in life stage survival, and understanding this can shape management targets for effective amphibian conservation.     

Life cycles of 17 riffle-dwelling mayfly species (Baetidae,Heptageniidae, and Ephemerellidae) in central Japan

We described the life cycles of 17 riffle-dwelling mayfly species in a central Japanese stream. Both species of Baetidae (Alainites yoshinensis Gose and Baetis thermicus Uéno) and two of nine species of Heptageniidae (Ecdyonurus scalaris Kluge and Epeorus latifolium Uéno/l-nigrus Matsumura) in this stream were multivoltine (more than one generation per year). Seven other heptageniid species (two unidentified species of Cinygmula, Epeorus aesculus Imanishi, E. curvatulus Matsumura, E. ikanonis Takahashi, E. napaeus Imanishi, and Rhithrogena japonica Uéno) and all six species of Ephemerellidae [Cincticostella elongatula (McLachlan), C. nigra (Uéno), Drunella basalis (Imanishi), D. ishiyamana Matsumura, D. sachalinensis (Matsumura), and D. trispina (Uéno)] were essentially univoltine.     

Annual development of mat-forming conjugating green algae <Emphasis Type="Italic">Zygnema</Emphasis> spp. in hydro-terrestrial habitats in the Arctic

Conjugating green algae of the genus Zygnema (Zygnematophyceae, Streptophyta) are dominant eukaryotic components of hydro-terrestrial microbial mats in the Arctic. Considering the harsh environmental conditions, the aim of this study was to elucidate mechanisms that enable Zygnema spp. to thrive in this habitat. We hypothesized that changes in morphology, physiological performance, and stress tolerance take place during the annual life cycle of the algae. We thus selected four natural populations of Zygnema spp. on Svalbard and investigated them throughout the vegetation season by means of light microscopy and chlorophyll a fluorescence. Additionally, we also investigated one overwintering population. No formation of specialized resting stages (e.g., dormant zygospores) was observed. Markedly, Zygnema spp. survived harsh periods as modified vegetative cells, i.e., pre-akinetes. Pre-akinetes tolerate both desiccation during summer and freezing in winter. These cells are not dormant and therefore recover their physiological activity immediately after transfer to favorable conditions, undergoing rapid growth in the early spring. Nevertheless, once pre-akinetes begin to grow, these newly produced vegetative cells lose stress tolerance. Such rapid dehardening explains their high mortality due to frequent freeze–thaw cycles in the early spring. Arctic Zygnema spp. thus face a phenological trade-off between missing the early growing season and experiencing frost damage.     

Feeding ecology of the Kızılırmak toothcarp, <Emphasis Type="Italic">Aphanius marassantensis</Emphasis>: ontogenetic shift and seasonal diet variation

We studied the feeding ecology of an endemic cyprinodontid fish, Aphanius marassantensis (K?z?l?rmak Toothcarp), with special emphasis on seasonal and ontogenetic diet shift. The dietary composition revealed an omnivorous diet with great seasonal and ontogenetic variation. Cladocera and Calanoid Copepods (Diaptomus sp.) dominated the diet in Spring and Summer, whereas the food items of plant origin e.g. filamentous algae and diatoms were the most important components in Autumn and Winter. An ontogenetic diet shift was also demonstrated. While the feeding of juveniles was based mostly on planktonic organisms, adult individuals preferred larger prey taxa, such as Gammarus sp. and Gastropods. There was no significant sex-related variation in feeding with the exception for the volume of gut content. The investigation on the feeding strategy of A. marassantensis suggested a generalised feeding pattern with some specialised individuals. This generalist feeding habits may account for the well-established population of this species in a big reservoir under coexistence of two non-native fish species, Pseudorasbora parva and Atherina boyeri.     

A comparative analysis of the chemical nature of defensive secretions of Gonyleptidae (Arachnida: Opiliones: Laniatores)

The arachnids of the order Opiliones (harvestmen) produce substances used in defense. In the present paper, we analyzed 22 species of Gonyleptidae to explore the use of defensive substances in taxonomy and evolutionary biology. Thirty-seven different compounds were detected, 18 of which were preliminarily identified. These compounds were mapped onto a phylogenetic tree showing the relationships within the Gonyleptidae. Data from Cosmetidae were used as an outgroup. Five ketones and six alkyl phenols were reported for the first time in harvestmen.     

Life History and Mating Behavior of <Emphasis Type="Italic">Rhagoletis solanophaga</Emphasis> (Diptera: Tephritidae), a Non-Diapausing Species with Highly Variable Mating Duration

As an initial contribution to understanding the adaptive value of behavioral and life-history strategies, the life cycle and mating behavior of an unstudied species of tephritid fruit fly in the genus Rhagoletis are characterized for the first time. Over a 9-month fruiting period, a small proportion of Solanum appendiculatum Dunal (< 10 %) was found to be infested with a single larva of Rhagoletis solanophaga (Hernández & Frías). The average duration of R. solanophaga lifecycle (c.a. 140 days from egg laying to death of adults) exceeded the three month fruitless period. Additionally, R. solanophaga is capable of exploiting Solanaceous plants in at least two genera. These features could have selected for a non-diapausing species of Rhagoletis, a genus where most species are univoltine. Nevertheless, some individuals in the population became dormant. As other members of the genus, R. solanophaga exhibited a resource defense mating system with forced copulations and multiple mating. Both males and females could be highly promiscuous and individual mating success exhibited a wide range of outcomes. Regardless of mating success, mated females stored similar amounts of sperm in two spherical spermathecae. Long copulations were observed, perhaps functioning as a form of mate guarding with probable disadvantages for females. We outline hypotheses and opportunities for future comparative studies examining sperm competition and mate guarding.     

The ontogeny of the Late Permian terebratulids of the family Beecheriidae Smirnova (Brachiopoda)

Based on the study of the ontogenetic changes of cardinalia and brachidium in the genera Beecheria Hall et Clarke, 1893 and Calycelasma Smirnova, 2007 (family Beecheriidae), a complex development of the brachidium, which corresponds to the angustothyridid type, and the formation of crural plates, which functioned as the outer hinge plates at the early developmental stages, are revealed.     

Muscle satellite cells and regulation of recovery potential of muscle

Major aspects of the biology of muscle satellite cells are reviewed: the identification, origin in early development, mechanisms of self-renewal mediated by asymmetric divisions, content in different muscle types and in different ontogenetic stages, role of control genes of the Pax family (in particular, Pax7) and their products in proliferation control, and involvement of growth factors (HGF, FGF, IDF, and TGF-β) in the activation of these cells after muscle damage. The characteristics of the early stages of myogenic differentiation of activated satellite cells along the pathway similar to muscle formation in embryonic development are discussed.     

Limited effective gene flow between two interfertile red oak species

Key message

Highly elevated differentiation in different life stages between two interfertile oak species at a CONSTANS -like gene suggests a role of this gene in pre-zygotic isolation and adaptive divergence between species.

Abstract

Genome-wide differentiation patterns among oak species suggest that divergent selection can maintain species-specific adaptations and morphological integrity by reducing effective interspecific gene flow. While there is evidence for both pre- and post-zygotic isolation mechanisms in oaks (e.g., differences in flowering time, selection against hybrids), these mechanisms are rarely studied at each life stage from acorns to adult trees within the same forest. To assess the reproductive isolation mechanisms between two ecologically divergent species, we (1) quantified the number of hybrids in different life stages in Quercus rubra and Quercus ellipsoidalis, two interfertile red oaks with different adaptations to drought, and (2) assessed the timing of bud burst in both natural populations and in a seedling common garden trial. The low number of hybrids in all life stages suggested pre-zygotic isolation between species or selection in very early life stages that have not been sampled (e.g., early seed abortion). Significant differences in bud burst were found in two consecutive years between species in a common garden seedling trial but not in natural populations of the same provenance. In addition, we found evidence for divergent selection on several gene loci between species in each life stage. In particular, an SSR repeat located within the coding sequence of a CONSTANS-like gene, a locus involved in the photoperiodic regulation of flowering time and development, showed very high interspecific differentiation between species in all life stages (mean F _ST = 0.83), compared to the average neutral differentiation of 3.7 %.

     

The bitter crab syndrome in commercial crabs in the Western part of the Bering and Chukchi Seas

We studied the distribution of the “bitter-crab” syndrome, a disease caused by the parasitic dinoflagellate Hematodinium sp., in eight commercial species of crabs in the west of the Bering and Chukchi seas. The crabs (25 388 individuals) were sampled during bottom trawl surveys of July?September 2010 and October?November 2012. The disease was first identified visually by a color change of the exoskeleton and the hemolymph of the animals and then using microscope analysis of hemolymph samples. Infestation was detected in crabs of three species, Chionoecetes opilio, C. bairdi, and Paralithodes platypus. The prevalence of the disease (the percent of infected individuals relative to all of those examined) in C. bairdi and P. platypus was very low, 0.1 and 0.3%, respectively. Infestation was widespread among C. opilio, its peak in the Bering Sea was in the fall. The average prevalence of the crab disease in different areas of the Bering Sea ranged from 0.8 to 10.8%. A high rate of crab infection was recorded in the Korfa Bay. In the Chukchi Sea, the average prevalence was 2%. Infestation by Hematodinium sp. was not revealed in the deep-sea snow crabs Chionoecetes tanneri and C. angulatus, and in three species of lithodid crabs, Paralithodes camtschaticus, P. brevipes, and Lithodes couesi. This can be explained by the small sample volume and/or ecology of these species, since the disease was registered in other areas in four of them.