Characterisation of burrow architecture under natural conditions in the sand-dwelling wolf spider Allocosa brasiliensis

Allocosa brasiliensis (Petrunkevitch, 1910) is a wolf spider that constructs silk-lined burrows along the coastal sand dunes of Argentina, Brazil and Uruguay. This species shows a reversal in typical sex roles and sexual size dimorphism expected for spiders. Females are the smaller and mobile sex, which initiates courtship at the male burrow entrance. Mating occurs in the male burrow, and when it ends, the male leaves and the female stays inside. Females prefer to mate with those males showing longest burrows, so burrow dimensions would be under strong sexual selection pressures. Previous studies in the laboratory indicated that male burrows are longer than those of virgin females, which were described as simple silk capsules. Preliminary observations suggested that juvenile burrows presented differences from those of adults; however, detailed observations of A. brasilienisis burrow characteristics at the field were lacking. The aim of this study was to characterise the burrows of adults and juveniles of A. brasiliensis under natural conditions. We recorded the dimensions of burrows inhabited by males, females and juveniles (n = 30 for each category) and created beeswax moulds that reflected burrow shape. Juveniles inhabited tubular burrows with two branches and two openings; on the contrary, adults were found in tubular burrows with a single entry. Males and females inhabited burrows of similar length and width, but those of juveniles were shorter and narrower. We discuss the results and their possible functional explanations according to the selective pressures expected for each developmental stage and sex.     

Juvenile Sphaeroma quadridentatum invading female-offspring groups of Sphaeroma terebrans

Female isopods Sphaeroma terebrans Bate 1866 are known to host their offspring in family burrows in aerial roots of the red mangrove Rhizophora mangle. During a study on the reproductive biology of S. terebrans in the Indian River Lagoon, Florida, USA, juvenile S. quadridentatum were found in family burrows of S. terebrans. Between September 1997 and August 1998, each month at least one female S. terebrans was found with juvenile S. quadridentatum in its burrow. The percentage of S. terebrans family burrows that contained juvenile S. quadridentatum was high during fall 1997, decreased during the winter, and reached high values again in late spring/early summer 1998, corresponding with the percentage of parental female S. terebrans (i.e. hosting their own juveniles). Most juvenile S. quadridentatum were found with parental female S. terebrans, but a few were also found with reproductive females that were not hosting their own offspring. Non-reproductive S. terebrans (single males, subadults, non-reproductive females) were never found with S. quadridentatum in their burrows. The numbers of S. quadridentatum found in burrows of S. terebrans ranged between one and eight individuals per burrow. No significant correlation between the number of juvenile S. quadridentatum and the numbers of juvenile S. terebrans in a family burrow existed. However, burrows with high numbers of juvenile S. quadridentatum often contained relatively few juvenile S. terebrans. The majority of juvenile S. quadridentatum found in family burrows of S. terebrans were smaller than the juvenile S. terebrans that were cared for by their mothers. The results indicate that the presence of S. quadridentatum in S. terebrans family burrows may negatively affect the duration of extended parental care in S. terebrans. It is not known why parental female S. terebrans are not able to discriminate against juvenile S. quadridentatum. Possibly, the fact that the two species are closely related facilitates S. quadridentatum sneaking into S. terebrans family burrows.     

Extended parental care in two endobenthic amphipods

Extended parental care is described for two endobenthic amphipods which inhabit the estuarine soft-bottoms along the Gulf of Maine. The juveniles of both amphipod species, Leptocheirus pinguis and Casco bigelowi, remain in the burrow of their mother after they hatch from the brood pouch. Several consecutive clutches of L. pinguis can inhabit the female's burrow simultaneously, but in C. bigelowi there is always only one clutch of juveniles in the burrow of the female. Larger juveniles of L. pinguis start building their own small tubes at the bottom of the female's burrow. Casco bigelowi females increase the size of their burrows during the time when the juveniles grow. The juveniles of both species leave the female's burrow at about half adult size. The major tasks of female L. pinguis and C. bigelowi during the time of extended parental care is to irrigate and maintain the deep burrow. It is hypothesized that extended parental care in L. pinguis, C. bigelowi and other tube/burrow-living peracarids is a mechanism to protect small offspring from epibenthic predation.     

Egg production of two sympatric species of Hyalella Smith, 1874 (Crustacea,Amphipoda, Dogielinotidae) in aquaculture ponds in southern Brazil

This study tests the hypothesis that sympatric species of Hyalella may show different reproductive strategies that permit their coexistence. Ovigerous females were collected monthly with nets during a 1-year period in two aquaculture trout ponds in Vale das Trutas, São José dos Ausentes County, southern Brazil. In both species there was a decrease in the number of embryos in the brood pouch during embryonic development, and an increase in egg volume. However, the two species differed in the body size of ovigerous females, number of eggs and juveniles per female, brood mortality rate, egg size and hatching size of juveniles. Egg production continued throughout the year, with the highest number of eggs produced during winter and spring by Hyalella pleoacuta, and winter, spring and autumn by Hyalella castroi. These differences in the reproductive traits of species of Hyalella may permit their coexistence in nature.     

Copulatory courtship and morphology of genitalic coupling in seven Phyllophaga species (Coleoptera: Melolonthidae)

Males in six of seven species of Phyllophaga perform apparent courtship behaviour during copulation. As predicted by the hypothesis that such copulatory courtship is under sexual selection by female choice, behavioural details differ between species. The modified male abdominal sternites, which rub against the female's pygidium during copulation and whose morphology also differs between species, and the flattened, setose tarsi of males of one species probably function as contact courtship devices during copulatory courtship. Observations of events inside copulating beetles were made through the semi-transparent abdominal sternites of females and male genitalia in three species. They revealed cryptic genitalic thrusting behaviour of male genitalia within the female which occurred even when there was little or no external movement of the male genitalia. Dissections of pairs frozen in copula demonstrated that males transfer large quantities of material to females during copulation that is not physically related to sperm transfer. The type of material varied between species. In at least two species most of the transfer apparently occurred after sperm transfer. The possibility that some male genitalic structures function as holdfasts and/or stimulatory devices was suggested by their positions relative to female structures, and/or their movements within the female. Portions of male genitalia with strong bristles tended to lie against setose or more heavily sclerotized portions of the female reproductive duct. Some portions of female genitalia which vary between species seem ill-designed to exclude the genitalia of cross-specific males. The female reproductive tract is partially everted during copulation in one species.     

Sexual dimorphism in chelicerae,forelegs and palpal traits in two burrowing wolf spiders (Araneae: Lycosidae) with sex-role reversal

Allocosa alticeps and Allocosa brasiliensis are two sex-role-reversed spiders. Females locate males and initiate courtship. Copulation occurs inside male burrows and after copulation the males deliver their burrows to the females. We explored the occurrence of differences in morphological traits related to sex roles in both Allocosa species and compared the results with two non-burrowing wolf spiders with typical sex roles. We measured the length of the foreleg's tibia-patella and the chelicerae. Scanning electron micrographs of palpal tarsus distal sections were taken. Males showed higher values than females in all the traits considered. Adult and penultimate males lacked true claws compared with juveniles and females, but showed modified spines. The palpal organ was more proximal to the tibia compared with the non-burrowing lycosids considered in this study. Spines and palpal organ location could be associated with more effective digging. We discuss how natural and sexual selection could interact on morphology in this scenario.     

A not so natural history of the tarantula Brachypelma vagans: Interaction with human activity

We describe the structure of a population of Brachypelma vagans (Ausserer, 1875) in relation to the intensity of human activity and report characteristics of the burrows in Campeche, Mexico. During September and October 2003, we established sampling areas in five different classes of vegetation type/land use: mature forest (MF), secondary forest (SF), backyard (BY), and a football field divided into corner area (FC) and goal area (FG). The densities of spiders and the proportion of different age/gender classes of individuals on the sites were compared. Morphological data among adults and juveniles were contrasted, and differences in morphology between juveniles were tested according to land use class. We compared the nearest distances between neighbouring burrows and between burrows and trees. Also, we studied the orientation of the burrows, and compared the diameter of the burrow entrance. Brachypelma vagans was found exclusively in the open areas with densities that ranked from 0.02 to 0.1 individuals per square metre, being among the highest ever reported for Theraphosidae. However, there was a negative relationship between density and intensity of human activity. The population of this tarantula shows segregation in occupation of space. Females occupied exclusively the backyards, whereas juveniles occupied sites according to their stage of development. The youngest juveniles occupied the backyards, while the pre‐adults occupied the football field. The distance between burrows was highly variable at all the sites. However it tended to be shorter in the backyards. The orientation of burrows was in all sites preferentially directed northwards. The diameter of the burrow entrances was a relatively good indicator of the sex and age of its occupier, and related almost directly to the dimension of the body. This study provides better knowledge of the structure of B. vagans populations in a human‐modified environment and gives new information on the natural history of these spiders.     

Natural prey of the crab spider Thomisus onustus (Araneae: Thomisidae), an extremely powerful predator of insects

The natural prey of the crab spider Thomisus onustus inhabiting Eryngium plants was studied in the Absheron Peninsula, Azerbaijan. The total percentage of specimens of T. onustus found while feeding was low (9.1%). However, solitary late instar and adult females exhibited a very high feeding percentage (29.9%). Other sex–age groups of T. onustus, including small juveniles, adult males, and females guarding egg sacs, were found feeding significantly less frequently. Investigation has shown that T. onustus is a polyphagous predator, with representatives of four arthropod orders found in its diet. The primary food of T. onustus was Diptera and Hymenoptera, which collectively accounted for 94.2% of total prey. Worker ants constituted about one‐fifth of the prey, suggesting that T. onustus is a myrmecophagic spider. The length of prey killed by T. onustus ranged between 1.25 and 16.00 mm (mean 6.21 mm) and constituted from 34.4 to 587.9% (mean 194.9%) of length of their captors. The most frequently captured were large arthropods, exceeding the size of the spiders (83.1%). Small and medium‐sized juveniles captured considerably larger prey (in terms of prey–predator length ratio) than late instar and adult females. Over half of their prey constituted arthropods more than two‐fold larger than the spiders.     

Reproductive biology of two species of Hyalella Smith, 1874 (Crustacea: Amphipoda: Hyalellidae) from southern Brazil

Reproduction is a critical event in the life cycle of animals, and a high amount of energy is allocated to it. The aim of this study was to characterize the reproductive biology of two recently described species of Hyalella from the north-east region of state of Rio Grande do Sul in southern Brazil. The specimens were collected during 12 months from a stream source (Hyalella georginae) and from a water reservoir (H. gauchensis), in Palmeira das Missões municipality, state of Rio Grande do Sul. Upon sampling, ovigerous females and pairs in precopulatory behaviour were separated into microtubes. In the laboratory, the cephalothorax length (CL in mm) of all specimens was measured. Male size at onset of sexual maturity was larger than that of the females, in both species. There was a correlation between the size of paired males and females: larger males carried larger females, in both Hyalella species. Precopulatory pairs were found throughout the year; however, H. georginae had a higher frequency in spring, and H. gauchensis in autumn. There was a positive correlation between the CL of ovigerous females and the number of eggs/juveniles. A reduction in the number of eggs throughout the embryonic development was observed in H. georginae. The average fecundity of H. georginae was significantly higher than H. gauchensis. The differences between the reproductive biology of these species may be related to the physical and environmental conditions at which they are exposed in their habitats.     

The reproductive cycle of Mothocya epimerica (Isopoda: Cymothoidae) a parasite of the sand smelt,Atherina boyeri (Osteichthyes: Atherinidae), in the Lesina Lagoon,Italy

The population of sand smelt, Atherina boyeri, living in the Lesina lagoon (Italy, south-western Adriatic Sea) is infested by the cymothoid isopod Mothoyca epimerica. Monthly samples of sand smelt were taken from July 1990 to May 1991. Male hosts were more infested than females. No relationship was found between the prevalence of infestation and the gonadosomatic and hepatosomatic indices of sand smelt. Mothocya epimerica is a protandric hermaphrodite. Reproduction occurs about four times from early summer to early winter, at 2 month intervals. Females produce multiple broods and their life span is at least 6 months during the reproductive season. From February to May reproduction is considerably reduced.     

A new species of Crepidula (Mollusca: Mesogastropoda: Calyptraeidae) from Chile: additional characters for the identification of eastern Pacific planar Crepidula group

A new species of Crepidula is described. It was collected together with Pagurus edwardsii Dana and Pagurus perlatus H. Milne Edwards, inhabiting the inside of empty shells of Tegula euryomphala Jones and other gastropods. The shell is low convex-oval in juveniles, low convex-oblong in males and concave-oblong elongated in females. In the adult phases the apex projects slightly at the posterior border. The sinusoidal septum is ‘V’ shaped at its centre, with a smooth sulcus in the left quarter. The osphradium has 2–13 digitiform leaflets according to phases of the life cycle. The characteristically thin, external-lateral teeth of the taenioglossate radula possess a row of denticles. In addition, evidence is given that: (1) males have a lanceolate penis with a spermatic tube; (2) the intracapsular-embryonic development starting with many embryos, occurs with a decrease at the larvae-protoconch stage, each one with an adhered-vitelline mass, and culminates with a few prehatching juveniles of 1 mm size. In the expression of the consecutive-protandric hermaphroditic sexuality, the male phase is truly separated from the female phase by a transition period at the population level. The shell and radular morphology, intracapsular development and ecological antecedents allow the proposal of Crepidula coquimbensis as a new species, and suggest a future taxonomic revision of the group based, moreover, on osphradium structure and new characters of the reproductive system, such as penis morphology.     

Aspects of the life history and reproductive biology of the introduced terrestrial amphipod Arcitalitrus dorrieni (Hunt) at two sites in Co. Galway,Ireland

The reproductive biology of Arcitalitrus dorrieni (Hunt) was investigated over a period of 25 months at a deciduous and a coniferous forest site at Kylemore, Co. Galway, Ireland. The species has an annual, univoltine reproductive cycle with ovigerous females found throughout the year. Recruitment of juveniles begins in May–June with a peak of reproductive activity in August–September. The populations at each site show a bimodal size-distribution for most of the year, with a unimodal distribution in March, April and May. Mortality occurs exponentially, and the life-span is 20–22 months. Mean fecundity was 5·65 eggs per female and was dependent on female size, egg size and brood stage. Brood mortality was 37·5% at Addergoole and 29·8% at Tullywee Bridge.     

Reproductive strategies of two sympatric species of Hyalella Smith, 1874 (Amphipoda,Dogielinotidae) in laboratory conditions

This study was performed in order to test theoretical predictions that sympatric species might show different reproductive strategies to facilitate their coexistence. The duration of precopulatory mate‐guarding behaviour, body size of ovigerous females, embryonic and postembryonic development time, fecundity, and viability of the sympatric amphipod species Hyalella pleoacuta and H. castroi were investigated under laboratory conditions (19±1°C and 12 h light/dark photoperiod). Specimens were collected with nets in autumn and spring 2004, and summer and winter 2005 in the Vale das Trutas, São José dos Ausentes County, southern Brazil, and transported to the laboratory. Both species of Hyalella showed similar reproductive strategies, especially in respect to the duration of the precopula, duration of the embryonic period, and fecundity. However, these species differed in body size of ovigerous females, duration of the postembryonic period, viability, and body size of juveniles that remained in the marsupium and those released by females. These differences in the reproductive traits of H. pleoacuta and H. castroi may facilitate their coexistence in nature.     

A report on two large collections of the squat lobster Munidopsis platirostris (Decapoda,Anomura, Munidopsidae) from the Caribbean,with notes on their parasites,associates, and reproduction

The reproduction and parasite associates of the squat lobster Munidopsis platirostris (A. Milne Edwards and Bouvier, 1894) were investigated based on collections made in the Bahamas and Curaçao with grassmat and bundled fishing net traps used to collect sipunculan worms and other small invertebrates. Size of ovigerous M. platirostris was significantly correlated with clutch size for females from both localities but females from the Bahamas produced significantly more eggs (on average 10.1 eggs/clutch) than females from Curaçao (on average 6.6 eggs/clutch). Early embryos of M. platirostris from the Bahamas were 0.74–0.82 mm in diameter, similar to some other species of Munidopsidae as well as Chirostylidae. Two species of crustaceans, another squat lobster and a leptostracan, as well as a limpet mollusc, were collected with M. platirostris in the Bahamas, while a sipunculan was an associated species in a Curaçao collection. One specimen of M. platirostris had an unidentified cryptoniscoid epicaridean isopod, possibly representing a new genus and species. Two specimens of M. platirostris each had one rhizocephalan externa of a species belonging to Lernaeodiscus Müller, 1862 but their morphology does not match that of L. schmitti Reinhard, 1950, the only species in the genus known from squat lobsters in the western Atlantic. Additional materials and tools, such as DNA analysis, are needed to describe these potentially new parasites and we suggest that use of these traps may be an effective method to obtain additional samples.     

Some new parasitic copepods (Siphonostomatoida: Nicothoidae) from deep-sea asellote isopods

Summary

The highly transformed siphonostomatoid Choniorhiza enaulis gen. et sp. nov. is described from an asellote isopod, Dendromunna sp., in the deep North Atlantic Ocean to the south west of the Azores. Male and female live inside a cyst on the dorsal side of the branchial chamber of the isopod. The new genus may be distinguished from all other genera of the Nicothoidae, except Rhizorhina, by the presence of oral processes replacing the cephalic appendages. It differs from Rhizorhina in the presence of a distinct urosome in the female. A key to all genera of the Nicothoidae is provided and Pseudonicothoe Avdeev & Avdeev is recognized as a junior synonym of Paranicothoe Carton. The male syntype of Paranicothoe cladocera Carton is designated as the lectotype because the female is an isopod. Sphaeronella bradfordae sp. nov. is described from an isopod of the genus Haploniscus caught on the Chatham Rise off New Zealand.     

Comparative population biology of Uca rapax (Smith, 1870) (Brachyura,Ocypodidae) from two subtropical mangrove habitats on the Brazilian coast

The structure of two populations of the fiddler crab Uca rapax in two subtropical mangrove habitats near Ubatuba, State of São Paulo, Brazil were compared. The size–frequency distribution, sex ratio, and recruitment were evaluated. Sampling was performed monthly from April 2001 to March 2002 in the Itamambuca and Ubatumirim habitats. Crabs were caught manually for 15?min by two collectors during low tide. The carapace width of each crab was measured with a digital caliper, and the sex and ovigerous state were recorded. The median size of the carapace width of males was greater than that of females at both sites (P<0.05). The median size of the crabs from Itamambuca was larger than at Ubatumirim (P<0.05). Only 28 ovigerous females were obtained from both mangroves, which suggested that females might remain in their burrows during the incubation period. The highest recruitment pulse occurred in winter for both populations, probably as a consequence of high reproductive activity during summer. The sex ratio in the size classes showed an anomalous pattern, with a higher frequency of females in the intermediate size classes. This may be related to a greater energy requirement for reproduction in females, thus delaying growth. The variable environmental conditions to which Uca rapax populations are subject appear to act directly or indirectly on the population, causing variations in growth and reproductive processes in the different populations investigated here.     

A new fecampiid of the genus Kronborgia (Platyhelminthes: Turbellaria: Neorhabdocoela) parasitic in the intertidal isopod Exosphaeroma obtusum (Dana) from New Zealand

Kronborgia isopodicola sp. n. (Turbellaria: Neorhabdocoela: Fecampiidae) is described from the South Island of New Zealand. The species has a parasitic phase within the haemocoel of the intertidal isopod Exosphaeroma obtusum (Dana). Sexes are separate: each female worm, emerging when mature from its host, secretes a cocoon around itself in which it is joined by a male. Cocoons are found under rocks, are approximately 10 mm long and reflect the shape of the contained female. The worms spawn and lay numerous egg capsules, each containing two embryos. Although placed in Kronborgia Christensen and Kanneworff, 1964 because it is dioecious, K. isopodicola exhibits some characters typical of the genus Fecampia Giard, 1886. It differs from all other described fecampiids by its possession of eyes throughout life.     

Offspring discrimination by female parents of the matriphagous spider Cheiracanthium japonicum (Araneae: Eutichuridae)

The ability of females of the matriphagous spider Cheiracanthium japonicum to discern their own offspring was examined based on a comparison of her maternal care for her own eggs and spiderlings with other conspecific females as well as those of other congeneric species. Cheiracanthium japonicum females did not discern their own nest from those of conspecific and congeneric individuals. In addition, the female did not distinguish her own egg sac from those of other conspecific or congeneric individuals. On the other hand, the female clearly discriminated between her own spiderlings and those of other congeneric species only if the congeneric spiderlings were introduced into the breeding nest. Consequently, the female will allow conspecific spiderlings to predate her (i.e. cannibalism). However, it is hardly expected that spiderlings of a certain female would eat another female in the field, as the reproductive period is quite short and synchronised in a population. Considering that the female can only recognise spiderlings of other species, the recognition ability of C. japonicum females will be evolved only to work as a protection against intruders of other species into the breeding nest.     

Reproductive ecology and behaviour of a species of Adenomera (Anura,Leptodactylinae) with endotrophic tadpoles: Systematic implications

The species of the Leptodactylus fuscus group and those of the genus Adenomera lay their eggs in underground chambers. According to current systematic arrangement of these genera, this behaviour is convergent. Here we describe reproduction, courtship, tadpole morphology, calls, chamber structure, and populational phenology of an Adenomera species, and compare some of these features with those of other species of the genus and with species of Leptodactylus of the fuscus and pentadactylus groups. We tested the tadpoles of the Adenomera sp. and those of Leptodactylus labyrinthicus (pentadactylus group) for their foam‐making abilities. There was sexual dimorphism in size; males averaged 22.0?mm and females 24.3?mm in snout–vent length. Males called from late September to late February; calling often began about 2–3?h before sunset. Males called exclusively in open (non‐forest) areas. Egg clutches were found around male calling sites. Males excavated spherical chambers which had a direct entrance. During mating, the male led the female towards a previously excavated chamber. Territorial interactions (aggressive calls and fights) occurred when an intruder male approached a calling male. Late‐stage tadpoles and newly metamorphosed juveniles were found within underground chambers. Clutch size averaged 10.6; eggs averaged 3.7?mm in diameter and were cream coloured. Tadpoles had horny beaks, but no labial tooth; spiracle was present. All tested groups of tadpoles generated foam within 10?h. As we report for Adenomera sp., the males of Leptodactylus species of the fuscus group lead the female to a previously excavated chamber. Tadpole foam‐making behaviour was reported in Leptodactylus of the fuscus group and was previously unknown for any species of Adenomera or for Leptodactylus of the pentadactylus group. The way in which the tadpoles of Adenomera sp., Leptodactylus spp. (of the fuscus group) and L. labyrinthicus made foam was quite similar. The foam‐making behaviour of the studied tadpoles may act to avoid compaction of the tadpoles at the bottom of the basin or underground chamber, avoiding overcrowding and increasing respiratory and excretory efficiency. As presently recognized, the phylogenetic position of Adenomera suggests that reproductive major behavioural features are convergent with some Leptodactylus species. Alternatively, our data point to close phylogenetic relationships between Adenomera and Leptodactylus of the fuscus and pentadactylus groups, reinforcing the paraphyletic nature of the genus Leptodactylus.     

The morphology and biology of the entomophilic Thripinema fuscum (Tylenchida: Allantonematidae), and the histopathological effects of parasitism on the host Frankliniella fusca (Thysanoptera: Thripidae)

We used light and electron microscopy to detail the in vivo life cycle of the nematode Thripinema fuscum and to determine the effects of parasitism on tissues of the thrips host Frankliniella fusca. The parasitic T. fuscum female produced eggs within 4–5 days after ingress and the host haemocoele became packed with eggs and developing juvenile nematodes. Mature juveniles migrated to the hindgut and fully developed juveniles first emerged from the posterior end of the male or female F. fusca 9 days after parasitization. Parasitization induced a displacement of the host alimentary tract, an atrophy of the ovaries and fat body, and an alteration of energy and waste reserves in host tissues. Our observations reveal a specialized relationship showing that this parasite is well adapted to exploit the host digestive, reproductive and excretory systems.