Assemblages of terrestrial isopods (Isopoda, Oniscidea) in a fragmented forest landscape in Central Europe

Terrestrial isopods were collected in 13 forest fragments differing in area (within the range of 0.1 and 254.5 ha), shape and composition of forest vegetation (thermophilous oak, mesophilous oak-hornbeam, thermophilous oak-hornbeam, acidophilous oak, basiphilous oak, beech oak-hornbeam, moist mixed deciduous forest, plantations of deciduous and coniferous trees), all situated in the Český kras Protected Landscape Area, Czech Republic, Central Europe. Number of sites sampled in each fragment of forest depended on its size and ranged from 1 to 7. Altogether 30 sites were sampled. Soil samples (5 per site collected twice a year) and pitfall trapping (5 traps per site in continuous operation throughout a year) during 2008–2009 yielded a total of 14 species of terrestrial isopods. The highest densities and highest epigeic activities of terrestrial isopods were recorded in the smallest fragments of woodland. Although a wider range of habitats were sampled in the larger fragments of woodland there was not a greater diversity of species there and the population densities and epigeic activities recorded there were lower. Porcellium collicola was most abundant in small fragments of woodland regardless the vegetation there. Armadillidium vulgare and Protracheoniscus politus were statistically more abundant in the larger fragments of woodland. The results indicate that forest fragmentation does not necessarily result in a decrease in the species richness of the isopod assemblages in such habitats.     

Histological studies on the marsupium of two terrestrial isopods (Crustacea,Isopoda, Oniscidea)

The marsupium, a brood pouch in peracarid crustaceans (Crustacea, Malacostraca) has evolved in terrestrial environment for providing nutrition and optimal conditions for embryogenesis. In the present study we give details on the histology and ultrastructure of its constituting elements such as oostegites and cotyledons. Marsupia of two different eco-morphological types of woodlice, namely the non-conglobating species Trachelipus rathkii Brandt, 1833 and the conglobating species Cylisticus convexus De Geer, 1778 were investigated. Light microscopic (LM) studies showed some differences in the main structure of the two species’ brood pouch: in Trachelipus rathkii, a ‘clinger’ type woodlice, the oostegites bend outwards during brood incubation as growing offspring require more space, while in Cylisticus convexus, a ‘roller’ type isopod, the sternites arch into the body cavity to ensure space for developing offspring and still allowing conglobation of the gravid females. The quantitative analysis of the oostegites’ cuticle proved that the outer part is about 2.5 - 3 times thicker compared to the inner part in both species. Electron microscopic (TEM) examinations show only small histological differences in the oostegites and cotyledon structure of the two species. Cellular elements and moderately electron dense fleecy precipitate are found in the hemolymph space between the two cuticles of oostegites. The cells contain PAS positive polysaccharide areas. TEM studies revealed some differences in the cotyledon ultrastructure of the two species. Cotyledons of Trachelipus rathkii consist of cells with cristate mitochondria and granular endoplasmic reticulum with cisterns. Cotyledons of Cylisticus convexus consist of cells with densely cristate mitochondria and ribosomes attached to vesicular membrane structures. In both species cells with electron dense bodies were observed. We conclude that - besides the differences in marsupial shapes - the fine structure of the oostegites and cotyledons is hardly affected by the eco-morphological type, specifically the conglobating or non-conglobating character of the studied species.     

Biogeography of body size in terrestrial isopods (Crustacea: Oniscidea)

This study tries to unveil the contribution of climatic shift in shaping the extreme body size diversity in terrestrial isopods (Oniscidea). Trying to explain size variation at an interspecific level, we test five hypotheses: (1) Bergmann's Rule and the temperature‐size rule postulate large size in cold areas; (2) The metabolic cold adaptation theory postulates small animal sizes in cold environments; (3) The primary productivity hypothesis predicts size increase in resource‐rich areas; (4) The aridity resistance hypothesis predicts large size in arid regions; and (5). The acidosis hypothesis predicts smaller size with decreasing soil pH. Globally, Bergmann's rule and the aridity hypothesis are weakly supported. Among families and genera, results are variable and idiosyncratic. Conglobating species sizes provide weak support for the acidosis hypothesis. Overall, size is strongly affected by familial affiliation. Isopod size evolution seems to be mainly affected by phylogenetically constrained life‐history traits.     

Occurrence and assemblage composition of millipedes (Myriapoda, Diplopoda) and terrestrial isopods (Crustacea, Isopoda, Oniscidea) in urban areas of Switzerland

Terrestrial isopods and millipedes, members of the invertebrate macro-decomposer guild, were collected through pitfall traps in three Swiss cities (Zurich, Lucerne, Lugano). A total of 7,198 individuals of 17 isopod species (7093 ind.), and 10 millipede species (105 ind.) were captured. Besides the Alpine endemic isopod (Trichoniscus alemannicus) and millipede (Cylindroiulus verhoeffi), urban assemblages were mainly composed of widespread, native European and even cosmopolitan species, which are frequent in anthropogenic areas. Overall species richness (isopods and millipedes combined) was similar in Zurich (17 species) and Lucerne (16), while only 13 species were sampled in Lugano. According to the Sørensen index of similarity, species composition of Zurich and Lucerne were more alike, while the one of Lugano was more distinct from the other two cities. This result can be explained by the spatial proximity of Zurich and Lucerne in the north of the Alps compared to Lugano, which is located more distantly and in the south of the Alps. Dominant isopods and millipedes in Zurich and Lucerne were found to be widespread synanthropic species in temperate Europe(Porcellio scaber, Trachelipus rathkii and Ophyiulus pilosus) while the dominant isopod in Lugano (Trachelipus razzautii) is a species with a north-eastern Mediterranean distribution. Our study reveals that the urban millipede and isopod fauna in Swiss cities mainly consists of widespread species, but species of narrower distribution (e.g. Trichoniscus alemannicus, Cylindroiulus verhoeffi) may also find suitable habitats in cities. Despite some signs of biotic homogenization, our study also found compositional differences of millipede and isopod assemblages between northern and southern cities that suggest geographical effects of the regional species pool.     

Description of the postmarsupial manca stages of Armadillidium granulatum (Crustacea,Isopoda, Oniscidea)

The development of the postmarsupial manca stages of Armadillidium granulatum Brandt, 1833 was studied in detail by morphological analysis. Ovigerous females were reared separately under controlled conditions, allowing us to follow the stages of development from release from the marsupium until the appearance of the first juvenile stage, identified by the full development of the seventh pair of pereopods. Each newborn was followed to record the subsequent moults that identify the three postmarsupial manca stages. Manca stage M I had a mean duration of 6 h, manca stage M II 15 days, and manca stage M III 32 days. The cephalothorax width was measured to provide a growth measure for each stage. The mean values of the cephalothorax width were: 0.512 mm (± 0.083 SD) for M I, 0.677 mm (± 0.058 SD) for M II and 1.194 mm (± 0.079 SD) for M III. The morphological modifications in the three postmarsupial manca stages were described, the body parts illustrated, and SEM images taken. The distinguishing characteristics among mancas were discussed, and comparisons made with manca stages of other terrestrial isopod species.     

The postmarsupial development of Porcellio siculoccidentalis, with some data on reproductive biology (Crustacea, Isopoda, Oniscidea)

In the broader context of research on the Sicilian Porcellio imbutus-complex, the postmarsupial development of Porcellio siculoccidentalis Viglianisi, Lombardo & Caruso, 1992 was studied in detail. This research was conducted in the laboratory under controlled conditions, allowing us to follow the stages of development, from the formation of the marsupium in ovigerous females until the larval stages and development of the seventh pair of legs. The timing of developmental stages and the morphological modifications of appendages in the postmarsupial manca stages (M I-M III) are described. The manca stage M I had a duration of about one hour. Ovigerous females were collected and reared separately, and the number of parturial molts in the absence of males was counted. The results showed a maximum of four successive parturial molts. Fecundity and fertility were evaluated as the number of eggs and embryos, respectively, inside the marsupium of the ovigerous females. Both parameters were positively correlated with the size of the females. The maximum numbers of eggs and embryos in the marsupium were 113 and 141, respectively. Data describing the total number of postmarsupial mancas released per month indicated that the highest release occurred in April.     

The occurrence of a rickettsial disease in British woodlice (Crustacea,Isopoda, Oniscidea) populations

During a structural study of the gut and tegumental glands of the woodlouse Porcellio scaber Latreille, numerous darkly staining spindle-shaped bodies were found in certain tissues. These proved to be elementary bodies of Rickettsiella grylli (Rickettsiales, Rickettsiaceae). Details of its structure and distribution within the host tissues are given, including the rosette-type tegumental glands and a type of tissue not seen in woodlice before, the vesicular bodies (also found in healthy specimens). This is the first time that this pathogenic organism has been recorded in P. scaber and the first record for R. grylli from the British Isles. The rickettsial disease may also be present in British populations of the woodlice Oniscus asellus Linnaeus and Armadillidium vulgare (Latrielle).     

Morphology and evolution of respiratory structures in the pleopod exopodites of terrestrial Isopoda (Crustacea, Isopoda, Oniscidea)

The morphology of the respiratory structures located in the pleopod exopodites of terrestrial Isopoda is described for representative species of different genera. Special emphasis is set on the evolution of these lungs in the context of phylogenetic relationships as revealed by other morphological characters. In the most primitive species of the Oniscidea, and still of subordinate taxa of the Crinocheta, respiration takes place in the thin ventral integument of the exopodites. The evolution of dorsal respiratory fields with a weakly wrinkled surface occurred at least six times within the Crinocheta. In five of these cases, a further development can be observed. The evolution of a partly covered respiratory field with strongly wrinkled surface may have taken place six times. It is assumed that completely internalized lungs with spiracles surrounded by a water‐repellent surface microsculpture, evolved at least six times independently within the Oniscidea: in the Tylidae, Actaecia, Aphiloscia, the Eubelidae, the Armadillidae and in a taxon probably comprising Porcellionidae plus Armadillidiidae.     

The diversity of terrestrial isopods in the natural reserve "Saline di Trapani e Paceco" (Crustacea, Isopoda, Oniscidea) in northwestern Sicily

Ecosystems comprising coastal lakes and ponds are important areas for preserving biodiversity. The natural reserve "Saline di Trapani e Paceco" is an interesting natural area in Sicily, formed by the remaining strips of land among salt pans near the coastline. From January 2008 to January 2010, pitfall trapping was conducted in five sampling sites inside the study area. The community of terrestrial isopods was assessed using the main diversity indices. Twenty-four species were collected, only one of them endemic to western Sicily: Porcellio siculoccidentalis Viglianisi, Lombardo & Caruso, 1992. Two species are new to Sicily: Armadilloniscus candidus Budde-Lund, 1885 and Armadilloniscus ellipticus (Harger, 1878). This is high species richness for a single reserve in Sicily. The extended sampling period also allowed us to study species phenology. Most of the species exhibited higher activity in spring than in autumn while some species also exhibited lower activity in the summer. The species richness revealed that the study area is in an acceptable conservation status; Shannon and Pielou indices also confirmed a more or less even distribution of individuals belonging to different species.     

Reproductive phenology of a natural population of Armadillidium pelagicum Arcangeli, 1955 (Isopoda, Oniscidea)

Armadillidium pelagicum, the most abundant species of the genus in Tunisia is endemic to the North of Tunisia and the circumsicilian islands. In the laboratory and under the natural conditions of temperature and photoperiod, virgin females of A. pelagicum exhibit a spontaneous ovarian maturation, followed by a parturial moult. Nevertheless, the onset of reproduction is greatly accelerated by the presence of a male. In fact, mating shortened the lag time (from the beginning of experiment to the parturial moult) by reducing the number of normal moult preceding the first parturial moult and the duration of the preparturial intermoult. In this species, only one mating is enough to ensure the fertility for several egg layings. Otherwise, it has been shown that A. pelagicum has a reproductive period rather than a reproductive activity and that mating females have a longer reproductive period than virgins. Like several species of terrestrial Isopods, the duration of the gestation period is temperature-dependent.     

Morphology and evolution of the respiratory apparatus in the family Eubelidae (Crustacea,Isopoda, Oniscidea)

The morphology of the respiratory apparatus in the pleopodal lungs of the family Eubelidae was investigated. The family is a monophyletic group including more than 240 species in 53 genera (three of which are nomina dubia), mostly distributed in the Afrotropical Region (tropical Africa and Arabian Peninsula). In all the Eubelidae, except for the monospecific genus Parelumoides and two species of the genus Elumoides, the exopods of pleopods have lungs. All the pulmonary morphologies present in the entire suborder Oniscidea are found: 1) uncovered lungs, composed of a pleated respiratory surface, directly exposed to the air (Atracheodillo-type) or partially enclosed within the appendage (Synarmadilloides-type); 2) covered lungs with several spiracles and respiratory trees, housed within the appendages, with spiracles surrounded by a specialized, nonrespiratory, structure (perispiracular area) (Eubelum- and Somaloniscus-types); 3) covered lungs with only one spiracle, with or without perispiracular area, and one respiratory tree (Aethiopopactes- and Periscyphis-types), which in taxa with Periscyphis-type lung crosses the insertion of the appendage and penetrates into the pleon with bundles of respiratory tubules. The evolution of the various types of lungs is discussed. It is concluded that the two main evolutionary lines, i.e., uncovered lungs and covered lungs, originated independently from an ancestral respiratory structure-the semilunar area. A first mechanism of development of the semilunar area by folding of its surface produced the Atracheodillo-type (all folds coplanar with the surface of the exopod) and Synarmadilloides-type (folds partly coplanar and partly intraflexed inside the exopod) uncovered lungs. A second mechanism of development by tubular invagination of the cuticle of the semilunar area produced the polyspiracular Eubelum-type lungs (numerous arborescent invaginations) and the monospiracular Aethiopopactes-type lungs (only one arborescent invagination), probably passing through a common intermediate pattern. From the common pattern, both the poly- and monospiracular types would have inherited the characteristic concave cell arrangement of the perispiracular area. The Somaloniscus-type and Periscyphis-type lungs are forms specialized for arid environments, directly derived from the Eubelum-type and Aethiopopactes-type, respectively.     

The agnarid terrestrial isopods (Isopoda,Oniscidea, Agnaridae) of the province of Qazvin,Iran, with a description of a new species

Six species of terrestrial isopods from the province of Qazvin, central Iran, are recorded. Three species, Hemilepistus klugii (Brandt, 1833), Protracheoniscus ehsani Kashani, 2014 and Mongoloniscus persicus Kashani, 2014, were previously reported from the province. Hemilepistus elongatus Budde-Lund, 1885 and Protracheoniscus major (Dollfus, 1903) are recorded for the first time, and one species, Protracheoniscus sarii sp. n., is described as new. The diagnostic characters of the new species are figured.     

Burmoniscus kitadaitoensis Nunomura, 2009 (Crustacea,Isopoda, Oniscidea) from southern Japan,a junior synonym of B. meeusei (Holthuis, 1947)

Re-examination of the holotype of Burmoniscus kitadaitoensis Nunomura, 2009 from Kitadaitojima Island, southern Japan reveals that this species is a junior synonym of B. meeusei (Holthuis, 1947). Partial regions of mitochondrial COI, 12S and 16S rRNA genes, and nuclear 18S and 28S rRNA genes were detected for species identification in the future.     

Mitochondrial DNA polymorphism and feminizing sex factors dynamics in a natural population of Armadillidium vulgare (Crustacea,Isopoda)

Sex determination in Armadillidium vulgare may be under the control of two parasitic sex factors that reverse genetic males into functional neo-females. The first feminizing factor (F) is a Wolbachia and the other (f) is probably a sequence of the F bacterial DNA unstably integrated into the host genome. Both of these feminizing factors are mainly maternally transmitted. Here we investigate the mitochondrial DNA polymorphism of wild iso-female lineages harbouring either F or f. Among the four haplotypes present in the population, two were the f-harbouring lineages, while two were common to the F- and f-harbouring lineages. This result suggests that there has been an introgression of the f factor into lineages infected by F Wolbachia. Based on previous data, we propose two different ways to account for such introgression. Given the particular dynamics of feminizing factors (f-harbouring lineages increase in populations at the expense of F-harbouring lineages), such an introgression should prevent the replacement of F-linked mitochondrial types by f-linked mitochondrial types in wild populations.     

Population structure and reproductive biology of Atlantoscia floridana (van Name, 1940) (Crustacea, Isopoda, Oniscidea) in southern Brazil

Data were obtained on the population structure and reproduction of Atlantoscia floridana, one of the most common species of terrestrial isopods in the restinga (coastal dune) forests of the state of Rio Grande do Sul, southern Brazil. During a 19-month period, a total of 7833 individuals were sampled: 2792 males, 3400 females and 1691 mancas. There was a significant difference between the size of both males and females collected in 2000 and 2001: the mean size was smaller in the second year when individuals in the larger size classes were lacking. Population density varied with season. The minimum population was 131 ind per m² individuals, the maximum 1040 ind per m² and the mean 450  per m². While the overall sex ratio was clearly female biased, the operational sex ratio favored males, and showed no changes with season. Because both ovigerous and post-ovigerous females were present throughout the year, reproduction is considered continuous; however, reproduction peaked during autumn and spring. Ovigerous females were measured (CW = cephalothorax width) and the number of eggs was counted. Fecundity (F) varied from 5 to 23 eggs ( = 11.18 ± 4) per female, and was expressed by the regression F = –18.48 + 22.59 CW, with the female cephalothorax width varying from 1.04 to 1.68 mm. Marsupial mortality was only 0.9%. Egg production was 588 eggs per m² in spring and 660 eggs per m² in autumn. Recruitment occurred in all months, and eggs, embryos and marsupial mancas were also present year-round. A. floridana is the dominant species of terrestrial isopod in the study area. Its most remarkable characteristic is its high reproductive investment.     

Patterns of neotenic differentiation in a subterranean termite, Reticulitermes speratus (Isoptera: Rhinotermitidae)

Plasticity in the caste developmental pathway is a remarkable characteristic of termite societies. In Reticulitermes, two types of neotenic reproductive, nymphoids and ergatoids, may differentiate from nymphs and workers and take over reproduction in the colony after the death of the original primary reproductive pair. We examined the dynamics of newly differentiated nymphoids and ergatoids in experimentally orphaned laboratory colonies of R. speratus with different caste compositions. The period required for differentiation of nymphoids was shorter than that for differentiation of ergatoids. The sex ratio of neotenics was strongly female‐biased, particularly in ergatoids. The results suggested that the number of differentiated ergatoids was restricted by the existence of nymphs or nymphoids in a colony. Workers were assumed to kill most newly differentiated neotenics. Attack reflecting conflict between colony members is probably an important mechanism to control neotenic emergence.     

Spermiophagic activity in the female genital system of some species of terrestrial isopods (Crustacea,Halophilosciidae)

The females of some species of the family Halophilosciidae receive in the course of mating a quantity of sperm considerably redundant with respect to the number of eggs that can be fertilized; this is possible thanks to the peculiar morpho-functional organization that characterizes their genital system and that allows them to store the sperm not only in the great seminal receptacle but also within the ovary. While most of the sperm stay free in the lumen of the seminal receptacle, a part of those present in the ovary undergoes a process of capture by the follicular cells with consequent internalization within endocellular cavities. This process concerns exclusively the immotile tail, that characterizes the peculiar spermatozoon of the isopods and which is essentially of proteic nature. After their capture the sperm tails undergo a gradual process of digestion, which seems to be apparently realized without the intervention of lysosomes. The possible role of this spermiophagic activity might be to represent a significant trophic paternal investment aimed at improving the fitness of the female and of the offsprings.     

A new species of Oniscus Linnaeus, 1758 (Crustacea: Isopoda: Oniscidea) from northern Spain, with a revised key to members of the genus

Oniscvs ancarenisis from the Sierra de Ancares in northern Spain is described as new. The genus Oniscus now contains four species: ancarensis sp. nov. , asellus Linnaeus, lusitanus Verhoeff and simoni Budde-Lund. The distribution of these species is figured and discussed, and a key is given for their determination.     

A twenty-one-year study of maternal dominance and secondary sex ratio in a colony group of stumptailed macaques (Macaca arctoides)

Trivers and Willard's theory of sex-ratio adjustment, as applied to cercopithecines, predicts that the ratio of male to female offspring will be greater for dominant than for subordinate mothers. A local-resourcexyhcompetition hypothesis predicts the reverse. To date, results from several species of macaque are inconsistent and often not statistically significant. In this 21 year study, a colony group of stumptailed macaques is added to the species previously studied. Seventy-five offspring were born to eight mothers for whom long-term dominance was established. Chi-square analyses of data from these 75 offspring failed to yield a significant relationship between sex-ratio and mother's dominance; consequently, consistent with a growing body of cercopithecine literature, neither of the competing theories was supported. © 1994 Wiley-Liss, Inc.