The composition and density of fauna utilizing burrow microhabitats created by a non-native burrowing crustacean (<Emphasis Type="Italic">Sphaeroma quoianum</Emphasis>)

The non-native isopod, Sphaeroma quoianum, has invaded many estuaries of the Pacific coast of North America. It creates extensive burrow microhabitats in intertidal and subtidal substrata that provide habitat for estuarine organisms. We sampled burrows to determine the effects of substratum type on the community of inquilines (burrow inhabitants). The density of inquilines was higher in wood and sandstone than marsh banks. Inquilines, representing 58 species from seven phyla, were present in 86% of samples. Inquilines equaled or outnumbered S. quoianum in 49% of the samples. Non-native fauna comprised 29% of the species and 35% of the abundance of inquilines, which is higher than other estuarine habitats in Coos Bay. Sessile non-native species were found living within burrows at tidal heights higher than their typical range. Thus, the novel habitat provided by burrows of S. quoianum may alter the densities and intertidal distribution of both native and non-native estuarine fauna.     

Distribution,density, and habitat use among native and introduced populations of the Australasian burrowing isopod <Emphasis Type="Italic">Sphaeroma quoianum</Emphasis>

The Australasian burrowing isopod (Sphaeroma quoianum) has been introduced to numerous embayments along the Pacific coast of North America. In some bays, populations of S. quoianum can exceed tens of thousands of individuals m⁻³ and bioturbation by the isopods can exacerbate shoreline erosion. Within their native range, however, studies recognize S. quoianum primarily as a woodborer. We measured the distribution, prevalence, habitat use, density, and associated fauna of S. quoianum in two bays within the native range [Tamar estuary (Tamar), Tasmania and Port Phillip Bay (PPB), VIC, Australia] and in one bay where the isopods had been introduced (Coos Bay, OR, USA). Distribution, prevalence, and habitat use were determined from shoreline surveys. Densities and the associated fauna of S. quoianum were measured in three intertidal substrata (marsh bank, wood, and friable rock). In all embayments, S. quoianum occurred primarily between 5 and 30 salinity and 55−68% of sites harbored isopods. Habitat use varied between embayments. Distributional patterns suggest salinity is the primary factor that limits the establishment and spread of S. quoianum. Isopod densities in all substrata were greater in Coos Bay than in the Tamar or PPB, although only densities within marsh banks varied significantly. Similarities in the amount of habitat and food, and the burrow dwelling lifestyle of S. quoianum suggest habitat availability/quality, food levels, predation, and competition are not responsible for the large differences in density. Lack of parasites or disease in populations of S. quoianum introduced to Coos Bay could be responsible for the prolific densities observed.     

Activity rhythms and cyclical changes of microhabitat preferences in the intertidal pipefish Nerophis lumbriciformis (Pisces: Syngnathidae)

As an inhabitant of the intertidal zone, Nerophis lumbriciformis predictably should show a clear rhythmic activity, with its maximum closely correlated with the high tide period, as observed in several other rocky intertidal fish species. We investigated the rhythmic activity patterns of N. lumbriciformis and analysed to what extent specific substratum preferences may be linked to water level and, ultimately, to the particular activity rhythm patterns of the fish. To determine N. lumbriciformis substratum preferences, two different experiments were conducted. The first evaluated substratum preferences from among a group of the four most common substrates found in the sample area. The second experiment evaluated the fish's maintenance of that substratum preference, after alteration of the water level in a simulated ebb tide. From the available substrata, N. lumbriciformis displayed a clear preference for algae until the water level began to decrease. Then, the preference radically shifted from algae to boulders. N. lumbriciformis showed a clear rhythmic activity, affected by both tidal and circadian periodicity, producing asymmetrical activity peaks. Unlike the rhythms of other rocky intertidal fish species, the maximum activity peaks were not centred at the expected high tide period. The adaptive explanation for the apparently peculiar activity peaks appears to be related to the particular substratum preferences of N. lumbriciformis. The detected circatidal period seems, consequently, closely related to small migrations between substrata, whilst the observed increase in diurnal activity, independent of the tidal cycle, may be related to predatory activity, since N. lumbriciformis is a visual feeder. Electronic Publication     

Effect of the substratum in the recruitment and survival of the introduced barnacle Balanus glandula (Darwin 1854) in Patagonia, Argentina

The barnacle Balanus glandula was introduced in Argentina in the 1970s, and today it dominates the high intertidal level in most Argentinean rocky shores. The aim of this work was to evaluate the effect of the type of substrata and intertidal height on a population of Balanus glandula by conducting field surveys and one-year field experiments in which we combined different substrata (hardness: hard and soft, and texture: smooth and rough) at two intertidal heights (mid and high). In natural populations, the highest density of adults and recruits occurred on soft-rough substratum and in the high intertidal. The different textures were important only on the soft substrata and high intertidal, and the density of barnacles of the soft-rough substrata was higher than soft and smooth ones. The most suitable experimental substratum was the soft-rough of the high intertidal, which had the highest recruitment, survival and final density of barnacles at the end of the experiment. In contrast, the hard and smooth of the high and middle intertidal were the least suitable in all cases. Although the recruitment of B. glandula occurred throughout the year, it was higher in the high intertidal, and it showed a recruitment peak in the winter and a second in the summer. While most studies on this barnacle investigated the effects of granite or other volcanic hard substrata, our study also focused on soft substrata. The effects of soft substrata are particularly important because soft sedimentary rocks characterise the southern Atlantic coast of South America and the presence of soft rocks appears to optimize the success of Balanus glandula.     

The effect of particle size frequency distribution of the substratum on the burrowing ability of Chiridota rigida (Semper) (Echinodermata: Holothuroidea)

Chiridota rigida (Semper) burrows easily into well‐sorted substrata, moving between particles in coarse substrata and pushing aside particles in fine substrata. Chiridota rigida does not burrow easily into poorly sorted substrata because spaces which can be penetrated are not available and because the particles cannot be moved as a result of increased stability of the substratum. In poorly sorted substrata, burrowing ability increases with increases in the proportion of fine particles. It would appear that the distribution and abundance of Chiridota rigida and other chiridotids would be affected by the effect of the substratum on their ability to burrow.     

The use of open burrows to estimate abundances of intertidal estuarine crabs

Estimation of absolute (or true) abundances of intertidal burrowing crabs is a difficult problem in some estuarine habitats because the nature of the substratum and behaviour of crabs can restrict researchers to the use of sampling methods which at best estimate only apparent, or relative, abundances. One method that has been used is to count open burrows to estimate population densities. This paper discusses the results of a test examining the validity of using burrows to estimate apparent abundances of a temperate ocypodid crab, Heloecius cordiformis. inhabiting mangrove forests of south-eastern Australia. Under appropriate circumstances, this method may provide a quick and reliable estimate of abundance of crabs.     

Studies on the behaviour and substratum preference in juveniles of the marine prawn,Metapenaeus monoceros (Fabricius)

Circadian rhythmicity in the behaviour of the marine prawn, Metapenaeus monoceros (Fab) and its substratum preference has been observed under laboratory conditions. In the morning at 9. a.m. 80% prawns remain buried and in the afternoon at 3 p.m. 60% burrowing was observed. A contrasting situation was observed in the night i.e. 17% burrowing at 9 p.m. and 30% burrowing at 3 a.m. was registered. In the second set of experiments juveniles were subjected to four types of substrata viz. mud, black fine sand, white coarse sand and stones. It was noted that M. monoceros showed more affinity towards the mud and less preference to other substrata. The obtained results are discussed to provide clues for prawn fishing.This work was supported by I.C.A.R., New Delhi through a project on Reprod. Endocr. Edible prawns.     

Selective crab predation on native and introduced bivalves in British Columbia

Experiments were conducted to determine whether locally abundant crab species prefer co-occurring littleneck clams, Protothaca staminea (Conrad, 1837) and Tapes philippinarum (A. Adams and Reeve, 1850), relative to a recently introduced species, the varnish clam, Nuttallia obscurata, (Reeve, 1857). Prey preference, handling time, pick-up success, profitability and consumption rates were investigated for two crab species, Dungeness crab, Cancer magister (Dana, 1852) and red rock crab, Cancer productus (Randall, 1839) crabs. Both crab species preferred varnish clams over the native species. This may be attributable to the lower handling time, higher pick-up success and increased profitability of consuming varnish clams. Handling time appeared to be a factor not only in species preference, but also in the degree of preference, with shorter handling times corresponding to stronger preference values. Both native and introduced bivalves burrow into the substratum, with the varnish clam burrowing deepest. When feeding on clams in limited substratum both crab species preferred the varnish clam. In the unlimited substratum trials Dungeness crabs preferred varnish clams (although to a lesser degree) while red rock crabs preferred littleneck clams. This was likely due to the significantly deeper burial of the varnish clam, making it less accessible. Although the morphology (i.e. thin shell, compressed shape) of the invader increases its vulnerability to predation, burial depth provides a predation refuge. These results demonstrate how interactions between native predators and the physical characteristics and behaviour of the invader can be instrumental in influencing the success of an invasive species.     

Distribution,substratum preference and burrowing behaviour of Lovenia elongata (Gray) (Echinoidea: Spatangoida) in the Gulf of Elat ('Aqaba), Red Sea

Lovenia elongata (Gray), the Indo-West Pacific spatangoid, was studied in the Gulf of Elat ('Aqaba) at the northern end of the Red Sea. The occurrence and local distribution of this macrobenthic burrowing species was determined and related to the particle size distribution of the substrata in which it was found. Its habitat preference is for clean, grassless, sandy bottoms in intertidal areas (beaches and sandbars), and on sublittoral level bottoms of protected bays and lagoons sheltered from heavy wave action. The preferred sediments were found to range from medium to fine sand. The population density, size frequency distribution, growth, reproduction, recruitment, and mortality of a sublittoral population of L. elongata (at Wadi Taba, Sinai) were studied.The mode of life of this burrowing heart urchin was observed in situ and in aquaria. Its burrowing and emergence behaviour are described. Experiments were carried out on the effect of particle size on burrowing behaviour. There are significant differences in the ability of L. elongata to manipulate substrata of varying grain size. The burrowing process was found to be most rapid in natural sand of medium-fine composition, corresponding to the normal sediment of the urchin's habitat. Such sediments were also found to be suitable for successful larval settlement; juveniles did not survive in coarser sand.Differences in population density and spatial distribution between various size groups are attributed to differences in their ability to manipulate the substratum. L. elongata is found to be morphologically suited to sand, particularly medium-fine sand; however, it is restricted by its morphology to shallow burrowing, since it lacks specialized mechanisms for deeper burrowing and funnel building.     

Evidences of habitat displacement between two common soft-bottom SW Atlantic intertidal crabs

The intertidal burrowing crab Chasmagnathus granulatus Dana is the dominant species in soft sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 41°S) where it produces dense and extensive burrowing beds. The mud crab Cyrtograpsus angulatus Dana coexists with Ch. granulatus in this area, but it also inhabits areas to the south (northern and central Argentinean Patagonia). A survey covering both areas showed that C. angulatus rarely live in burrows when coexisting with Ch. granulatus, but form large burrowing beds when not coexisting with Ch. granulatus. When both species coexisted, burrowing beds of C. angulatus are restricted to sandy-muddy areas. Only rarely are burrows of C. angulatus found within Ch. granulatus beds. However, when Ch. granulatus were experimentally excluded within their burrowing beds, new settlers of C. angulatus made burrows and maintained them until they reached large size. Paired (inside and outside Ch. granulatus burrowing bed) sampling during high tide using beach nets showed that C. angulatus rarely venture inside the Ch. granulatus crab beds. Other field experiments showed that adults Ch. granulatus always displace C. angulatus from burrows. Furthermore, in several sites located south of the limit of distribution of Ch. granulatus at the Patagonian coast, soft bare intertidals are dominated by burrowing beds of C. angulatus mixed with the congener C. altimanus Dana. Together, these evidences suggest that the mud crab C. angulatus is displaced from soft bottom areas by the burrowing crab Ch. granulatus. It is an example of competitive exclusion through aggressive interference in soft-bottom habitats when the shared resource is the access to sediment surface, a two-dimensional well-defined resource.     

External Microflora of a Marine Wood-Boring Isopod

Bacteria associated with the marine wood-boring isopod Limnoria lignorum were enumerated by acridine orange epifluorescence microscopy and by plate counts on several media; the plate-viable bacteria were isolated and identified. Similar procedures were followed to enumerate and identify bacteria associated with the wood substrate from which the isopods were collected and with the surrounding water from the isopod habitat. Approximately 1.4 × 10⁷ bacterial cells were associated with each individual L. lignorum. Aeromonas hydrophila, Pseudomonas, and Vibrio were the most common genera in the isopod microflora. Wood from L. lignorum burrows had an associated bacterial flora of 1.6 × 10⁷ cells per mg (damp weight). A. hydrophila also predominated in the wood microflora. The water from which the isopod population was collected contained 2.3 × 10⁶ bacteria per ml. Pseudomonas and Vibrio species were very common in the water microflora, but A. hydrophila was not detected. Interactions between the isopod, its associated microorganisms, and the microorganisms within the wood substrate are discussed in the light of the known absence of a resident digestive tract microflora in these animals.     

Live coral repels a common reef fish ectoparasite

Coral reefs are undergoing rapid changes as living corals give way to dead coral on which other benthic organisms grow. This decline in live coral could influence habitat availability for fish parasites with benthic life stages. Gnathiid isopod larvae live in the substratum and are common blood-feeding parasites of reef fishes. We examined substrate associations and preferences of a common Caribbean gnathiid, Gnathia marleyi. Emergence traps set over predominantly live coral substrata captured significantly fewer gnathiids than traps set over dead coral substrata. In laboratory experiments, gnathiids preferred dead coral and sponge and tended to avoid contact with live coral. When live gnathiids were added to containers with dead or live coral, significantly fewer were recovered from the latter after 24 h. Our data therefore suggest that live coral is not suitable microhabitat for parasitic gnathiid isopods and that a decrease in live coral cover increases available habitat for gnathiids.     

Burrow plasticity in the deep-sea isopod Bathynomus doederleini (Crustacea: Isopoda: Cirolanidae)

We investigated whether the deep-sea isopod Bathynomus doederleini has the capacity to change burrow length in response to changes in environmental conditions. We observed burrowing behavior in individuals that were placed on substrates with either simple (ST) or complex (CT) surface topographies. Individuals in the ST group (N = 10) constructed seven burrows. The mean ratio of the burrow length to body length was 1.8. The individuals in the CT group (N = 10) constructed eight burrows with a mean ratio of burrow length to body length of 2.5. Thus the burrows were significantly longer in the CT group. In addition, the isopods in the CT group often incorporated a chamber in the mid-section of the burrow. Our results may be used to infer the determinants of burrow morphology and speculate about the lifestyle of this species in the deep sea.     

The effects of burrowing of Helice tridens (De Haan) on the soil of a salt-marsh habitat

The physical and chemical effects of the burrowing activity of the mud crab Helice tridens (De Haan) on the soil of a salt-marsh habitat were investigated. Soil-turnover rate caused by burrowing activity was found to be ≈ 3% of the soil from the surface to a depth of 40 cm every day during the summer. The vertical distributions of leaf and stem fragments of the salt-marsh plant Phragmites australis (Trin.) and the vertical distribution of ammonium N concentration in the soil were also investigated. At locations in the marsh where there were many large burrows, numerous leaf and stem fragments were recognized in the soil, while in areas in the marsh containing only a few small burrows these fragments were scanty. The soil depths at which leaf and stem fragments were abundant, corresponded to the depths of the burrows. These results show that mud crabs bury fallen plant fragments in the soil by their burrowing activity. Ammonium N in the soil was also abundant at locations in the marsh where there were many burrows, indicating that organic matter, such as fallen leaves and stems, may be decomposed to inorganic nutrients which are useful to the salt-marsh plants.     

Activity patterns, feeding and burrowing behaviour of the crab Ucides cordatus (Ucididae) in a high intertidal mangrove forest in North Brazil

Activity patterns, feeding and burrowing behaviour of the economically important semi-terrestrial mangrove crab Ucides cordatus (Ucididae, L. 1763) was studied in a high intertidal Rhizophora mangle forest stand in Bragança, North Brazil. Video observations in the rainy and dry season were conducted over 24 h cycles at different lunar phases to investigate the behaviour of these litter-feeding crabs outside their burrows. During the rainy season, crabs stayed inside their burrows for 79% and 92% of the time during day and night, respectively. Time spent for feeding, burrowing and other activities outside their burrows was significantly longer during the day with 9.9% (night: 1.7%) and at waning and waxing moon with 9% (full and new moon: 0.9%). At neap tides (no tidal inundation) foraging and feeding activities outside burrows were clearly light-dependent, increasing at dawn and decreasing at dusk. Highest activities during daytime relate to the visual localisation of food. During the dry season, crabs spent less time inside burrows at neap tides than during the rainy season (80% and 91%, respectively). However, time spent for feeding activities was similar during both seasons. During almost all observation periods crabs collected leaf litter, but rarely fed on it outside burrows. At neap tides nearly all available litter was collected, suggesting that the U. cordatus population is litter-limited during these times. At spring tides (regular tidal inundation) the surface activity of U. cordatus was tide-dependent. Crabs closed their burrow entrances 2-3 h before flooding and re-emerged as soon as the tide retreated. During the day, burrow maintenance was the second most frequent behaviour after feeding. Agonistic interactions were regularly observed and were mainly related to burrow defence. The mean foraging radius of the crabs was only 19 cm (max: 1 m) underneath high Rhizophora mangle trees where crab densities were high. The results point to a high competition for burrows and show that U. cordatus is territorial. It is concluded that several exogenous factors, in particular light, leaf litter availability, flooding of burrows and the presence of conspecifics are important in controlling the crabs' activity patterns.     

长江河口湿地不同植被中无齿螳臂相手蟹的分布及其洞穴利用

无齿螳臂相手蟹(Chiromantes dehaani)是长江河口中、高潮滩的优势底栖动物之一,具有重要的生态功能.由于其经济价值较低,所受关注很少.本文调查了相似高程芦苇(Phragmites australis)带、芦苇斑块、菰(Zizania aquatica)植被以及裸地4种生境中无齿螳臂相手蟹的数量分布及其洞穴的利用情况.有植被分布的3种生境间植株密度、植株高度和植物干重生物量(地上部分)差异显著(P＜0.001);4种生境问,陷阱桶采样获取的无齿螳臂相手蟹数量分布(P＜0.001)和性比(P=0.001)差异显著;裸地区域没有无齿螳臂相手蟹可用的洞穴分布,芦苇带、芦苇斑块和菰植被间无齿螳臂相手蟹可用的洞穴密度(P＜0.019)、单位洞穴蟹个体数(P＜0.001)差异显著,挖掘洞穴采样获取的无齿螳臂相手蟹密度差异接近显著水平(P=0.067),洞穴占用率(P=0.667)、不同洞穴占用方式频次组成(P＞0.05)差异不显著;无齿螳臂相手蟹可利用洞穴密度与植株密度、植物干重生物量显著负相关(P＜0.05),而挖掘洞穴采样获取的无齿螳臂相手蟹密度与植株高度(P=0.077)相关性接近显著水平且与洞穴密度(P＜0.001)显著正相关.这些结果表明盐沼植被对无齿螳臂相手蟹的分布和洞穴利用存在影响.而进一步探讨盐沼不同植被对无齿螳臂相手蟹分布的影响,需要先了解无齿螳臂相手蟹在不同植被中分布格局在时间上的变化,并结合无齿螳臂相手蟹在不同植被间的迁移行为以及繁殖周期等进行探讨.     

ISOPOD TRACKWAYS FROM THE CRAYSSAC LAGERSTÄTTE, UPPER JURASSIC, FRANCE

Abstract:  Well-preserved arthropod trackways are described from the laminated limestones of the Crayssac Lagerstätte (south-west France, Lower Tithonian). They occur in sediments deposited in the temporary coastal mudflats of intertidal to supratidal zones. The trackways are referred to Pterichnus isopodicus isp. nov., and are interpreted as the locomotion traces of isopods. Different trackway morphotypes are recognized and clearly resulted from variations in the original consistency of the sediment. Sinuous trackways may correspond to vagrant activity on wet mud whereas numerous straight ones indicate a more rapid crawling on a soft-to-firm substratum (e.g. tidal flat during emersion). The preferred orientation of trackways indicates that isopods were crawling in a direction perpendicular to shoreline as a result of possible taxis induced by sediment wetness and/or by a migratory behaviour controlled by tidal rhythm. Unusually long emergence of the sediments may have favoured the preservation of dense networks of trackways. An isopod identity is supported by the general morphology of the tracks and the association of trackways with isopod body fossils. Archaeoniscus , which occurs abundantly in Late Jurassic deposits of England and France, was probably the trace-maker.     

The food and feeding habits of two co-occurring marine catfish Galeichthys feliceps and G. ater (Osteichthyes: Ariidae) along the south-east coast of South Africa

The feeding-associated morphological structures of the two Galeichthys species were found to be similar in all respects. Interspecific competition for food was avoided through habitat separation. Habitat preference, as established by dietary analysis and fishing trials, was based on substratum type. Galeichthys feliceps fed over sandy and muddy substrata in marine and estuarine environments, while G. ater fed exclusively over marine reefs. The two species have different caudal fin structures which are probably evolutionary responses to habitat-associated behavioural requirements in the two different environments.
The diets of the two species were investigated and compared using calorific values of prey items. While both species fed predominantly on benthic crustaceans, polychaetes and molluscs, little dietary overlap occurred at the species level. In the estuarine environment, G. feliceps fed mainly on the anomurans Upogebia africana and Calianassa kraussi , crabs Hymenosoma orbiculare and Cleislosoma edwardsii , and several isopod species. In the marine environment G. feliceps fed mainly on two species of crabs, Thaumastoplax spiralis and Goneplax angulata , the echiurid Ochaetostoma capense and the sedentary polychaete Sternaspsis scutata. A high incidence of teleost scales in the diet of G. feliceps juveniles was found to be a consequence of scavenging rather than a lepidophagous habit. Galeichthys ater fed widely on several reef-associated crabs, isopods, polychaetes and cephalopods.     

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.