Observations on the burrows and burrowing behaviour of two mud-dwelling decapod crustaceans,Nephrops norvegicus and Goneplax rhomboides

The burrows of the Norway lobster Nephrops norvegicus (L.) and of the crab Goneplax rhomboides (L.) were studied in Loch Torridon, Scotland. Polyester resin casts of burrows in the sea were made by divers to reveal their subsurface form. Tunnels made by N. norvegicus were usually simple, with two or more openings on the mud surface, and penetrated to a depth of about 30 cm. G. rhomboides burrows did not descend more than about 15 cm beneath the surface, but were usually more complex than the lobster burrows and had several openings. The methods of burrow construction used by the two crustaceans are described from aquarium observations. Neither N. norvegicus nor G. rhomboides show obvious morphological adaptations for burrowing, and it is suggested that the fossorial habit was adopted very early by decapods. The burrows of N. norvegicus do not seem to have assumed any functions in addition to the original one of providing refuge from predators. There is not sufficient known of the biology of the crab to indicate whether the same is true in its case.     

Functional morphology of Lithotrya dorsalis (Cirripedia: Thoracica) in relation to its burrowing habit

Burrowing mechanisms of the scalpellid barnacle Lithotrya dorsalis collected from Indian Key, Florida, USA and Fort Point, Jamaica, West Indies, in the fall and spring of 1983 and 1984 were investigated. Calcite spicules covering the peduncule of this animal mechanically abrade carbonate substratum and are described by light and scanning electron microscopy. These spicules undergo a morphological transformation during the molt cycle. Initial observations on the burrowing habit of L. dorsalis indicate that it also employs a chemical agent during bioerosion. Characteristic etching patterns on both optically pure calcite (Iceland spar) and shell material lend further support to this contention. Possible sources of this chemical agent are discussed.     

Cellulose digestion and phenol oxidation in coastal isopods (Crustacea: Isopoda)

In order to test three hypotheses on digestive constraints that may have affected the colonization of land by isopods, two marine isopods and one semi-terrestrial species were screened for their ability to oxidize phenolic compounds and digest cellulose in natural and artificial diets. Ligia pallasii (Isopoda: Oniscidea) and Gnorimosphaeroma oregonense (Isopoda: Sphaeromatidea) oxidized dietary phenolics, but Idotea wosnesenskii (Isopoda: Valvifera) did not, even though it feeds on seaweeds that are rich in phenolics. All three species were able to digest some cellulose, but this ability was least developed in the marine phytophagous species, Idotea wosnesenskii, and best developed in the semi-terrestrial L. pallasii. After reducing the number of endosymbiotic bacteria in the hepatopancreas (midgut digestive gland) by feeding antibiotics, cellulose digestion in L. pallasii was significantly reduced. Our results are consistent with the hypotheses that (1) the ability to oxidize phenolics is absent in phytophagous marine isopods, but present in saprophagous marine and semi-terrestrial species, (2) the ability to digest cellulose was an important pre-adaptation facilitating a fully terrestrial life-style in isopods, and (3) endosymbiotic bacteria in the hepatopancreas aid digestion in terrestrial isopods, and to a lesser degree in semi-terrestrial species, but not in marine isopods.     

The mouthparts of the members of the Jaera albifrons group of species (Crustacea: Isopoda)

The mouthparts of Jaera albifrons Leach, Jaera ischiosetosa Forsman, Jaera praehirsuta Forsman and Jaera forsmani Bocquet were examined with the light microscope and the Stereoscan electron microscope, and were found to be identical in all species. The structure of the mouthparts indicate that all four species feed by scouring and abrading algal or bacterial films from rocks or Fucus. Food, therefore, does not seem to be an isolating factor involved in the microdistribution of the four species on a shore.     

Is helping a beneficial learning experience for red-cockaded woodpecker (Picoides borealis) helpers?

Several hypotheses have been suggested to explain the evolution and maintenance of helping behavior in cooperatively breeding birds, one of which we investigated in the red-cockaded woodpecker (Picoides borealis). Helping may provide a learning experience that improves reproductive success once the helper becomes a breeder. We used data collected from a population of red-cockaded woodpeckers in the Sandhills of North Carolina to compare the reproductive success of 2-year-old breeders that were helpers at age 1 (helping experience) to those that were floaters or solitary males at age 1 (no helping experience). Reproductive success of the two groups was similar, indicating that helping provides no experience useful in reproduction. We reject the learning experience hypothesis as an explanation for the maintenance of helping behavior in red-cockaded woodpeckers. Received: 29 July 1996 / Accepted after revision: 27 April 1997     

Hepatopancreatic endosymbionts in coastal isopods (Crustacea: Isopoda), and their contribution to digestion

Three isopod species (Crustacea: Isopoda), commonly found in the intertidal and supratidal zones of the North American Pacific coast, were studied with respect to symbiotic microbiota in their midgut glands (hepatopancreas). Ligia pallasii (Oniscidea: Ligiidae) contained high numbers of microbial symbionts in its hepatopancreatic caeca. Numbers of endosymbionts were strongly reduced by ingestion of antibiotics. By contrast, the hepatopancreas of Idotea wosnesenskii (Valvifera: Idoteidae) and Gnorimosphaeroma oregonense (Sphaeromatidea: Sphaeromatidae) did not contain any microbiota. Results of feeding experiments suggest that microbial endosymbionts contribute to digestive processes in L. pallasii, the most terrestrial of the three isopods that we studied. The acquisition of digestion-enhancing endosymbionts may have been an important evolutionary step allowing isopods to colonize terrestrial habitats where relatively indigestible leaf litter is the primary food source. By contrast, the ability to digest phenolic compounds was most developed in one of the more marine species, suggesting that this trait may have evolved independently in isopod species that consume a phenolic-rich diet, whether in marine habitats or on land. Received: 28 August 2000 / Accepted: 8 December 2000     

Population dynamics and reproductive biology of the commensal isopod Colidotea rostrata (Crustacea: Isopoda: Idoteidae)

Colidotea rostrata (Benedict, 1898) is the only known commensal idoteid isopod, living on and mimicking the color of two northeastern Pacific sea urchins of the genus Strongylocentrotus. The population dynamics and reproductive biology of C. rostrata on its host urchins were studied at a low rocky intertidal area in southern California (33°40N; 118°30W) from December 1984 to December 1986. Isopod populations remained relatively stable throughout the 2 yr study, with isopods inhabiting an average of 56.1% of the urchins at 6.1±0.6 (x±2 SE) isopods per urchin. Female isopods reach sexual maturity at a length of 6.8 mm. Fecundity in C. rostrata averaged 11.8±0.9 (x±2 SE) embryos per brood, and increased with female body size. Breeding occurs all year long in C. rostrata, with a main reproductive period between the warmer spring and summer months. Newly released mancas and juvenile isopods were present during all months of the study. C. rostrata differs from the free-living Idoteidae in its smaller maximum size, reduced fecundity, 1:1 sex ratio, and low juvenile mortality. These features may represent adaptations to a commensal life style that reflect a reduced mortality pressure on these isopods.     

Locating food in a spatially heterogeneous environment: Implications for fitness of the macrodecomposer Armadillidium vulgare (Isopoda: Oniscidea)

To assess the fitness consequences of foraging on patchy resources, consumption rates, growth rates and survivorship of Armadillidium vulgare were monitored while feeding in arenas in which the spatial distribution of patches of high quality food (powdered dicotyledonous leaf litter) was varied within a matrix of low quality food (powdered grass leaf litter). Predictions from behavioural experiments that these fitness correlates would be lower when high quality food is more heterogeneously distributed in space were tested but not supported either by laboratory or field experiments. To investigate whether A. vulgare can develop the ability to relocate high quality food patches, changes in foraging behaviour, over a comparable time period to that used in the fitness experiments, were monitored in arenas in which there was a high quality food patch in a low quality matrix. A. vulgare increased its ability to relocate the position of high quality food over time. It reduced time spent in low quality food matrices and increased time spent in high quality food patches with time after the start of the experiment. When the position of a high quality food patch was moved, the time spent in the low quality food matrix increased and less time was spent in high quality food patches, compared to arenas in which the food was not moved. The fitness benefits for saprophages of developing the ability to relocate high quality patches while foraging in spatially heterogeneous environments are discussed.     

Extent and variation of sound enhancement inside burrows of the frog Eupsophus emiliopugini (Leptodactylidae)

Variation of the amplification effect of burrows of the leptodactylid frog Eupsophus emiliopugini on conspecific calls generated externally was investigated. Advertisement calls broadcast through a loudspeaker placed in the vicinity of a burrow were monitored with small microphones positioned inside and outside the cavity. For 150 presentations of calls of 15 individuals in 12 burrows, 134 were amplified and 16 were attenuated (range –6–13 dB). The fundamental resonant frequency of burrows, measured with broadcast noise and pure tones, averaged 814 Hz (range 302–1361 Hz) and covaried with burrow length. The dominant frequency of the calls of burrow occupants (average 1062 Hz, range 636–1459 Hz) was not correlated with the fundamental resonant frequency of these cavities. In burrows with low resonant frequencies, externally broadcast calls with high dominant frequencies were attenuated, or amplified to a lower extent than calls with lower dominant frequencies. The dominant frequencies of the calls experienced shifts towards the burrows’ fundamental resonant frequencies. The amplification of calls inside burrows of E. emiliopugini exhibits manifest variability, with considerable potential for facilitating acoustic interactions in this species. Received: 18 July 1999 / Received in revised form: 19 July 1999 / Accepted: 25 July 1999     

Facultative lecithotrophy during larval development of the burrowing shrimp Callianassa tyrrhena (Decapoda: Callianassidae)

Survival, developmental and consumption rate (Artemia nauplii ingested per day) as well as predation efficiency (ingested per available Artemia nauplii) were studied during the larval development of the shallow-water burrowing thalassinid Callianassa tyrrhena (Petagna, 1792), which exhibits an abbreviated type of development with only two zoeal stages and a megalopa. The larvae, hatched from berried females from S. Euboikos Bay (Aegean Sea, Greece), were reared at 10 temperature–food density combinations (19 and 24 °C; 0, 2, 4, 8 and 16 Artemia nauplii d⁻¹). Enhanced starvation resistance was evident: 92 and 58% of starved zoeas I molted to zoea II, while metamorphosis to megalopa was achieved by 76 and 42% of the hatched zoeas at 19 and 24 °C, respectively. The duration of both zoeal stages was affected by temperature, food density and their interaction. Nevertheless, starvation showed different effects at the two temperatures: compared to the fed shrimp, the starved zoeae exhibited accelerated development at 19 °C (8.4 d) but delayed metamorphosis at 24 °C (5.9 d). On the other hand, both zoeal stages were able to consume food at an increased rate as food density and temperature increased. Predation efficiency also increased with temperature, but never exceeded 0.6. Facultative lecithotrophy, more pronounced during the first zoeal stage of C.tyrrhena, can be regarded as an adaptation of a species whose larvae can respond physiologically to the different temperature–food density combinations encountered in the wide geographical range of their natural habitat. Received: 28 February 1998 / Accepted: 21 October 1998     

Does oxygen deficiency modify the functional response of Saduria entomon (Isopoda) to Bathyporeia pilosa (Amphipoda)?

The functional response of the predatory isopod Saduria entomon to the prey amphipod Bathyporeia pilosa was measured in normoxia (95% O₂ saturation), moderate hypoxia (45% O₂ saturation) and hypoxia (35% O₂ saturation) in aquarium experiments. The prey densities tested ranged from 400 to 8000 ind m⁻². Prey density influenced consumption rates of S. entomon in normoxia and 45% O₂ saturation, but there was no difference between consumption rates at these two oxygen levels. Nevertheless the form of functional response differed. In normoxia S. entomon showed a positively density-dependent functional response to B. pilosa, indicating a potentially stabilizing effect on the prey population. In moderate hypoxia the variance in consumption increased, decreasing the statistical power to distinguish between response models. The functional response of S. entomon in moderate hypoxia was best described with a density-independent response, characterized as destabilizing for the prey population. In hypoxia (35% O₂) predation by S. entomon did not respond to increasing prey density, as almost no amphipods were eaten at this oxygen level. The results are discussed in terms of the usability of theoretical models to examine predator–prey relationships in stressful environments. Received: 26 April 1997 / Accepted: 20 May 1997     

Studies on digestion and the fine structure of digestive caeca in Eurydice pulchra (Crustacea: Isopoda)

In the carnivorous isopod Eurydice pulchra Leach the mid-gut caeca have a pH of 5.6 and produce a strong acid proteinase, a lipase and a carbohydrase. Electron microscope studies of the digestive caeca of starved and fed animals show that secretion is merocrine and may originate from a complex involving the basal cell membrane and associated mitochondria, producing lysosomes. Absorbed materials, in fed animals, appear to take the form of neutral lipid bodies, glycogen, and proteinaceous crystals, the latter forming within the golgi complex. Digestion of both lipid and protein inclusions was observed to occur intra-cellularly by fusion with free lysosomes. Secretion and absorption may occur in the same cell, and the scheme for the sequence of epithelial cell stages is proposed.     

Effects of helpers on breeder survival in the red-cockaded woodpecker (Picoides borealis)

Helpers can gain future indirect fitness benefits by increasing the survival of breeders that produce offspring related to the helper. Helping augments group size through the helper's presence and, in some cases, by increasing fledging success. Breeders may then experience enhanced survivorship because of the benefits of living in large groups. Helping may also reduce the workload of the breeder, which in turn may increase the likelihood that the breeder will survive to breed again. We used Cox's proportional hazards model to examine whether breeders' survival in two populations of the red-cockaded woodpecker (Picoides borealis) was enhanced when group size was increased in the presence of (1) the helper itself, or (2) "extra" fledglings (fledglings produced by the breeder because of helping behavior). We found that in the presence of helpers, the risk of a breeder dying declined by 21–42% for males and 0–14% for females. Our results suggest reduced breeder workload as one mechanism to explain reduced breeder mortality in the presence of helpers: breeders spent less time incubating and provisioning nestlings when assisted by helpers. The risk of a breeder dying declined by 16–42% in males and 26–43% in females in the presence of "extra" fledglings. We speculate on possible mechanisms by which fledglings might affect breeder survival. Our results support the hypothesis that helpers gain future indirect benefits by reducing breeder mortality. Electronic Publication     

Influence of increased cod abundance and temperature on recruitment of northern shrimp (Pandalus borealis)

Despite an increase in northern shrimp (Pandalus borealis) female biomass in the past years, the recruitment of the offshore population north and northeast of Iceland has remained very low. In this study, the influence of abiotic and biotic factors was studied in relation to shrimp recruitment. Two factors, cod (Gadus morhua) abundance and summer sea surface temperature (SST), were found to have a negative effect on offshore shrimp recruitment, explaining 71 % of the observed variation. Both cod abundance and temperature on the offshore shrimp grounds have increased in the past years, while recruitment has decreased and been at historically low levels since 2005. No significant relationship was found between recruitment and spawning biomass, indicating that recruitment variability is mainly driven by other factors. Cod abundance and summer SST are likely to affect different life stages of shrimp, as SST influences shrimp during its planktonic phase while cod abundance influences the demersal stage.     

Associative behavior of the fish Cryptocentrus cryptocentrus (Gobiidae) and the pistol shrimp Alpheus djiboutensis (Alpheidae) in artificial burrows

Observations and experiments on the associative behavior of the fish Cryptocentrus cryptocentrus (Valenoiennes) and the shrimp Alpheus djiboutensis De Man from the Red Sea were carried out in artificial burrows. A signal system, bringing about correlated behavior in fish and shrimp, consisted of antennal contact of the shrimp with the fish, tail undulations by the fish and, finally, emergence of the shrimp. The shrimp emerged from the burrow only in the presence of the fish. A constant antennal contact was maintained by the partners while the shrimp was outside the burrow. Emerging from the burrow, the shrimp pushed the fish towards the entrance. Following visual stimulation, the shrimp always retreated back into the burrow with the fish. However, the further back the shrimp was located inside the burrow, the less intense was its retreat with the fish. In the absence of the fish, visual stimulation of the shrimp had no effect. Cleaning of fish by shrimp was observed within the burrow.     

Biodiversity in Littorina species (Mollusca: Gastropoda): a physiological approach using heat-coma

 The upper critical thermal limits were measured as heat-coma temperatures in Littorina species as a tool for examining physiological diversity. Thermal tolerance was found to be stable within a population, but extensive variation occurred between populations and species. All species examined displayed heat-coma values of ∼30 °C and did not show a positive correlation with shore height. Eulittoral-fringe species tended to have higher coma temperatures than eulittoral species. Coma temperatures varied both seasonally and geographically. Lethal thermal limits were also investigated; these were ∼10 C° higher than coma temperatures. Received: 16 August 1999 / Accepted: 13 June 2000     

Respiratory regulation and the role of the blood in the burrowing shrimp Callianassa californiensis (Decapoda: Thalassinidae)

Questions relating to oxygen extraction and hemolymph oxygen transport have been investigated in the burrowing shrimp Callianassa californiensis Dana. The relationship of oxygen consumption to oxygen tension was determined in intermolt and postmolt shrimp. Soft postmolt individuals had higher rates of oxygen consumption and a somewhat lower critical oxygen tension than intermolt specimens. Both oxygen-carrying capacity and hemocyanin content are somewhat lower in postmolt compared to intermolt shrimp. The measured gradient between postbranchial (pericardial sinus) and prebranchial (abdominal sinus) hemolymph oxygen content is small and highly variable, and is not affected by exposure to anoxic stress. Hemolymph pH and buffering capacity are high. The oxygen consumption-oxygen tension relationships and respiratory properties of the hemolymph are discussed in relation to the hypoxic environment associated with the habitat of these crustaceans.Supported in part by Institutional Sea Grant 04-3-158-4 and Public Health Service Grant HL 12326.Communicated by J.S. Pearse, Santa Cruz     

Latitudinal variation in the reproductive biology of the burrowing ghost shrimp Callianassa filholi (Decapoda: Thalassinidea)

Population structure and reproductive ecology of the burrowing ghost shrimp Callianassa filholi Milne-Edwards, 1878 were studied in populations along a latitudinal gradient throughout New Zealand during the breeding season. Size-frequency distributions revealed unimodal populations, with predominantly sexually mature shrimp. All populations showed an unbiased sex-ratio, and there appeared to be no significant difference in size (carapace length, CL) between sexes. At the same time, CL and size at maturity differed significantly between populations; however, a general increase in sizes from north to south was not consistent throughout the latitudinal range studied. The timing of the breeding cycle differed significantly between populations, and breeding started earlier in southern populations. Number of embryos (fecundity) increased linearly with female CL at each location, but rates differed significantly between populations. Embryo size was not related to number of embryos, and the former increased significantly with latitude. With the exception of embryo size, observed differences in body size/size at maturity and reproductive timing between C. filholi populations are thought to be determined by food availability rather than temperature. Thus, further study is suggested on these aspects of thalassinid reproductive biology. Received: 17 June 1999 / Accepted: 8 January 2000     

Trophic diversity of idoteids (Crustacea, Isopoda) inhabiting the Posidonia oceanica litter

The coexistence of three idoteid species in Posidonia oceanica litter raises the question of trophic diversity and their role in the litter degradation process. Hence, diet composition of Idotea balthica, Idotea hectica and Cleantis prismatica was studied using a combination of gut contents and stable isotopes analysis. Gut content observations indicate that P. oceanica dead leaves are an important part of the ingested food for the three species, although their tissues are constituted of only a small to medium fraction of P. oceanica carbon. Our results also underlined the potential role of these species in the degradation of P. oceanica litter by mechanically fragmenting the litter and by assimilating a small to medium fraction of carbon. Moreover, we showed that there were considerable inter- and intra-specific differences in diet composition. Diet differed between juveniles and adults for I. balthica. Crustaceans are an important food source for adults of I. balthica, while I. hectica indicated a major contribution of algal material. C. prismatica showed an intermediate diet. This trophic diversity is probably one of the factors allowing these species to coexist in the same biotope.