The distribution and abundance of Helice tridens (De Haan) burrows and substratum conditions in a northeastern Japan salt marsh (Crustacea: Brachyura)

The effects of the physical condition of the substratum on the distribution and abundance of mud-crab burrows were investigated. In the salt-marsh habitats studied, where burrows were abundant, there was a high proportion of silt-clay. In such conditions, the diameters of openings and the depths of the burrows were greater than those when the relative proportion of silt-clay was low.In the laboratory, it was found that with a low proportion of clay, mud-crabs did not dig burrows when the water content was high and thus equivalent to that in the marsh. When the proportion of silt-clay was low, the larger mud-crabs did not make burrows at the depth of the water table in the marsh, because the layer above the water table was too thin to allow construction of the burrow. The field observations could thus be explained by the results of experiments showing that the depth of the burrows was controlled by the depth of the water table and the relative proportion of silt-clay in the substratum.     

The occupation of artificial burrows by Helice tridens (De Haan) and the length of its stay

The mud-crab Helice tridens (De Haan) influences the cycling of matter in salt-marsh ecosystems through its burrowing activity. If the crabs occupy and stay in their burrows for a short time, their burrowing activity will be great, since they will continuously construct new burrows. Therefore, investigation of the relation between the crabs and their burrows is considered to be important. In the present study, the relation between pipes as a form of artificial burrow and their occupation by the crab was analyzed. A close relationship was recognized between the diameter of the pipe opening and the carapace width of the crab which occupied the pipe, while pipes with a length shorter than the depth of the crab burrows were hardly occupied. These results indicate that the diameter and length of an artificial burrow affects the likelihood of its occupation by this species of crab. The length of the crab's stay in this type of artificial burrow was generally 1 day. This result may be related to the field observation that newly made burrows frequently collapse due to water current occurring through tidal action after the crabs have left.     

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.     

Root Foraging Influences Plant Growth Responses to Earthworm Foraging

Interactions among the foraging behaviours of co-occurring animal species can impact population and community dynamics; the consequences of interactions between plant and animal foraging behaviours have received less attention. In North American forests, invasions by European earthworms have led to substantial changes in plant community composition. Changes in leaf litter have been identified as a critical indirect mechanism driving earthworm impacts on plants. However, there has been limited examination of the direct effects of earthworm burrowing on plant growth. Here we show a novel second pathway exists, whereby earthworms (Lumbricus terrestris L.) impact plant root foraging. In a mini-rhizotron experiment, roots occurred more frequently in burrows and soil cracks than in the soil matrix. The roots of Achillea millefolium L. preferentially occupied earthworm burrows, where nutrient availability was presumably higher than in cracks due to earthworm excreta. In contrast, the roots of Campanula rotundifolia L. were less likely to occur in burrows. This shift in root behaviour was associated with a 30% decline in the overall biomass of C. rotundifolia when earthworms were present. Our results indicate earthworm impacts on plant foraging can occur indirectly via physical and chemical changes to the soil and directly via root consumption or abrasion and thus may be one factor influencing plant growth and community change following earthworm invasion. More generally, this work demonstrates the potential for interactions to occur between the foraging behaviours of plants and soil animals and emphasizes the importance of integrating behavioural understanding in foraging studies involving plants.     

Application of morphologic burrow interpretations to discern continental burrow architects: Lungfish or crayfish?

A methodology for trace fossil identification using burrowing signatures is tested by evaluating ancient and modern lungfish and crayfish burrows and comparing them to previously undescribed burrows in a stratigraphic interval thought to contain both lungfish and crayfish burrows. Permian burrows that bear skeletal remains of the lungfish Gnathorhiza, from museum collections, were evaluated to identify unique burrow morphologies that could be used to distinguish lungfish from crayfish burrows when fossil remains are absent. The lungfish burrows were evaluated for details of the burrowing mechanism preserved in the burrow morphologies together forming burrowing signatures and were compared to new burrows in the Chinle Formation of western Colorado to test the methodology of using burrow signatures to identify unknown burrows.

Permian lungfish aestivation burrows show simple, nearly vertical, unbranched architectures and relatively smooth surficial morphologies with characteristic quasi‐horizontal striae on the burrow walls and vertical striae on the bulbous terminus. Burrow lengths do not exceed 0.5 m. In contrast, modern and ancient crayfish burrows exhibit simple to highly complex architectures with highly textured surficial morphologies. Burrow lengths may reach 4 to 5 m.

Burrow morphologies unlike those identified in Gnathorhiza aestivation burrows were found in four burrow groups from museum collections. Two of these groups exhibit simple architectures and horizontal striae that were greater in sinuosity and magnitude, respectively. One of these burrows contains the remains of Lysoro‐phus, but the burrow surface reveals no reliable surficial characteristics. It is not clear whether Lysorophus truly burrowed or merely occupied a pre‐existing structure. The other two groups exhibit surficial morphologies similar to those found on modern and ancient crayfish burrows and may provide evidence of freshwater crayfish in the Permian.

Burrows from the Upper Triassic Chinle Formation in western Colorado exhibit simple to moderately complex architectural morphologies, ranging from predominantly vertical, unbranched, with little or no chamber development to predominantly vertical, few branches, and with minor chamber development. Surficial burrow morphologies are moderate to highly textured. The burrows have scrape marks, scratch marks, mud and lag‐liners, knobby surfaces, pleopod striae, and body impressions.

Although no fossil remains of the burrowing organism were found within or associated with the Chinle burrows from western Colorado, the similarity of architectural and surficial burrow morphologies to those in the Chinle of Canyonlands, Utah and to modern crayfish burrows, clearly indicates that the Colorado burrows are the product of burrowing crayfish rather than lungfish. Evaluation of burrowing signatures preserved in the architectural and surficial burrow morphologies is a very useful tool to compare and contrast Chinle burrows from different regions on the Colorado Plateau. Documentation of crayfish burrows in the Chinle of Utah and Colorado strongly suggests that other large‐diameter Chinle burrows elsewhere on the Colorado Plateau and in stratigraphically equivalent units may also be the product of crayfish activity.     

Rhombomys opimus contribution to the “fertile island” effect of tamarisk mounds in Junggar Basin

Fertile islands are created and maintained by a combination of physical and biologically mediated processes. Plants have been shown to be very important in the formation of fertile islands, and recent research indicates that the activities of burrowing animals have a significant influence on the physiochemical properties of soil that can promote the development of fertile islands. In this paper, we chose tamarisk (Tamarix spp.)??a constructive species that grows in the oasis-desert ecotone, and the great gerbil (Rhombomys opimus)??a widely distributed rodent in Central Asia to study the influence of great gerbils on soil nutrient dynamics in tamarisk mounds. Results indicate that fertile islands exist in tamarisk mounds without burrows of the great gerbil. However, the great gerbil??s burrowing activities promote the fertile island effect in tamarisk mounds: soil nutrients under shrubs with great gerbil activity were significantly higher than inter-mound areas in both surface soil and deep soil from 15 to 50?cm. Available nitrogen in mounds with rodent burrows was over twice as high as in tamarisk mounds without burrows at the same depth. The great gerbil??s burrowing activities promote the concentration of soil nutrients in the tamarisk mounds.     

ECHINOID AND CRUSTACEAN BURROWS AND THEIR DIAGENETIC SIGNIFICANCE IN THE MAASTRICHTIAN-DANIAN OF STEVNS KLINT, DENMARK

The increase in species and specimens of fossils in the uppermost part of the Maastrichtian White Chalk is interpreted as a result of reduced depth. The absence of bryozoans, brachiopods, and regular echinoids in the Cerithium Limestone indicates sedimentation in tidal pools. After sedimentation of the Cerithium Limestone, burrowing activity followed. A burrow of Brissopneustes danicus similar to burrows of the recent Echinocardium cordatum is described. Callianassa and its burrows are found in the Upper Danian calcarenite but not in the Lower Danian or Maastrichtian of Denmark. The dominant type of burrows along the Maastrichtian-Danian boundary has presumably been formed by the crustacean Ctenocheles.
The early post-Maastrichtian burrowing activity was succeeded by (1) induration of the bottom sediment and a slight abrasion (2) dissolution of aragonite shells and siliceous sponges, (3) offshore sedimentation and filling of the burrows with Lower Danian chalk mud, bryozoan fragments and other fossil remains, and (4) settling in the deeper part of the soft chalk sediment and precipitation of flint in or around burrows near the surface of the sediment.     

Colonization and substratum preference of an introduced burrowing crustacean in a temperate estuary

Habitat use in marine invertebrates is often influenced by multiple abiotic and biotic factors. Substratum composition is one factor known to have a dramatic effect on habitat selection. The Australasian burrowing isopod (Sphaeroma quoianum, H. Milne Edwards 1840) is a common introduced species in many estuaries on the Pacific coast of North America. S. quoianum burrows into a variety of firm substrata including marsh banks (composed of peat, clay, and/or mud), wood, friable rock, and Styrofoam floats. In some areas, isopods achieve high densities and may accelerate the rate of shoreline erosion and damage marine structures; thus, understanding the substratum preference of this species may be important for conservation and management efforts. Field experiments were conducted in Coos Bay, Oregon to examine substratum preference, burrowing rates, and the life stage of colonizers. In three experimental trials (Fall 2005, Spring 2006, Fall 2006), replicates of four intertidal substrata (marsh banks, decayed wood, sandstone, Styrofoam) were deployed near intertidal populations of S. quoianum. The numbers of burrows created in each substratum were enumerated weekly or daily (depending on trial). After the trials were completed, the total numbers of isopods inhabiting each substratum were counted. In weeks, S. quoianum extensively burrowed the substrata but exhibited a distinct preference for decayed wood. Significantly more isopods were present in wood than the other substrata at the end of the experiments and rates of burrowing were greatest in wood, although significance varied across time in one trial. Nearly 90% of colonizing isopods were under 5 mm in length suggesting that juvenile isopods primarily colonize intertidal substrata. Differences between burrow densities measured in the field and the results from these preference trials may indicate other factors, such as relative availability of substrata, recruitment and dispersal limitations, and possible gregarious behavior also influence local isopod densities.     

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.     

Manganese as an ecological factor in salt marshes

The plant available manganese concentration (Mn²⁺) of salt-marsh sediments was compared to that of acidic and neutral soils. The mean soil-manganese concentration was higher in the top 1 cm of salt-marsh soil than in the neutral soil and comparable to that of the acidic soil (0–5 cm). A peak in the soil-manganese concentration in the upper marsh was observed one week after the spring tide but this effect was not evident in the lower marsh. Despite these differences, there was no correlation between mean manganese concentration and position on the marsh.The response to manganese of salt-marsh halophytes was studied by measuring growth and root elongation in a range of Mn²⁺ concentrations with and without sodium chloride. Although there was a differential response to manganese between salt-marsh species, manganese resistance was not related to position on the marsh. Most of the species investigated were tolerant of Mn²⁺ at concentrations higher than normally recommended for plant growth. Moreover a salt-marsh ecotype of Festuca rubra was found to have a higher manganese resistance than an inland ecotype of the same species.When sodium chloride was included in the growth medium, salt-marsh plants had a greatly increased resistance to manganese associated with a reduced uptake. This effect is reflected in the tissue-manganese concentration which was lower than in Deschampsia flexuosa although both groups of plants were exposed to a similar range of Mn²⁺ concentrations. It is concluded that sodium chloride markedly reduces the phytotoxicity of manganese in salt marshes.Nomenclature following Clapham, Tutin & Warburg (1968). Flora of the British Isles.The work was carried out while one of us (C. E. Singer) was in receipt of an SERC studentship, which is gratefully acknowledged.     

A comparison of crayfish burrow morphologies: Triassic and Holocene fossil,paleo‐ and neo‐ichnological evidence,and the identification of their burrowing signatures

The architectural and surficial morphologies of crayfish burrows from the Upper Triassic Chinle Formation and Holocene sediments were compared in order to determine: 1) if Triassic burrows could truly be attributed to crayfish activity; 2) how comparable the burrowing mechanisms are; and 3) whether or not a common set of burrowing signatures could be identified for both ancient and modern freshwater crayfish. Materials used in this study include burrows from the members of the Upper Triassic Chinle Formation, casts of modern burrows constructed by Procambarus clarkii Hobbs and Procambarus acutus acutus (Girard) in the laboratory, and casts of naturally constructed modern burrows of Cambarus diogenes di‐ogenes (Girard).

Triassic and Holocene crayfish burrow morphologies exhibit simple to complex architectures, varying degrees of branching, chamber, and chimney development. They also exhibit relatively textured surficial morphologies (bioglyphs) such as scrape and scratch marks, mud‐ and lag‐liners, knobby and hummocky surfaces, pleopod striae, and body impressions. Holocene crayfish construct distinctive burrows due to their conservative limb arrangement, functional morphology, and behavior with respect to environmental stimuli. Similarities between Holocene and Triassic crayfish burrows suggest that extant and Triassic crayfish employed identical burrowing mechanisms. Features of the surficial and architectural morphologies impart a distinctive signature to burrows of both ancient and modern freshwater burrowing crayfish.

Burrowing signatures of crayfish can be used to identify new and previously misinterpreted continental trace fossils. These are useful in studies of the paleohydrogeology, paleoclimatology and paleoecol‐ogy of burrow‐bearing deposits.     

The impact of plateau zokor<Emphasis Type="Italic">Myospalax fontanierii</Emphasis> burrows on alpine meadow vegetation on the Qinghai-Xizang (Tibetan) plateau

The impact of burrows constructed by plateau zokorsMyospalax fontanierii (Milne-Edwards, 1867) on alpine meadow vegetation on the Qinghai-Xizang (Tibetan) plateau was investigated. Plant samples taken from quadrats directly over active zokor burrows, back-filled burrows, adjacent burrow controls, and random sites from a field, in which no burrows or mounds occurred were compared. The biomass of plants (below- and above-ground) directly over shallow active burrows was significantly lower than on control plots. This reduction in biomass was not significantly different than that between deep active burrows and control plots. There were no significant differences between above- and below-ground plant biomass on areas perpendicular to active burrows when compared to random sites. Back-filling soil in burrows could promote the growth of above-ground monocotyledonous plants. However, the burrowing activities of zokors had a negative effect on biomass of dicotyledonous 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.     

Ethological and environmental significance of Bergaueria hemispherica from the Late Cretaceous of the Bagh Group,Western India

The probable trace maker of B. hemispherica is the fact that one encounters different age groups of sea anemone burrows extended at variable depths in soft, unconsolidated, non-fluidized mud to get stability. It occurs as densely packed, vertical to subvertical, usually unornamented, occasionally showing faint, thin, ring-like structures, allochemic sandstone-filled cylinders with hemispherical base extending to variable depths in the shale. The diameter-to-height ratio calculated for 44 burrow specimens shows that the diameters of most of the specimen are smaller than their height. The probable trace maker of B. hemispherica is the fact that one encounters different age groups of sea anemones, which extended their column at variable depth in soft, unconsolidated, non-fluidized mud to get stability. The inclined nature of paired burrows towards each other suggests social aggression while unpaired inclined burrows suggest swaying in search of food. The monodominant occurrence of B. hemispherica as pre-storm colonization of r-selected organisms (sea anemones) suggest sstressed environment and simultaneous vacation of the burrower reflects rapid sedimentation due to high-energy storm events. Further, storm and inter-storm events deposited sandy allochemic limestone and shale series, respectively, but did not form an identical condition for the colonization of the sea anemone. The species B. hemispherica of the Bagh Group revealed physicochemical parameters (energy conditions, turbidity, sedimentation rate, bathymetry, suspended organic matter, substrate consistency, oxygen, and salinity) of the Late Cretaceous transgressive sea.     

Location of long‐term communal burrows of a threatened arid‐zone lizard in relation to soil and vegetation

The great desert skink (Liopholis kintorei) of the Egerniinae subfamily (Reptilia: Scincidae) is a communal burrowing lizard that inhabits arid spinifex grasslands in central Australia. Great desert skink activity is centred in and around the burrows which are inhabited for many years. However, it is not known whether skinks select burrow sites with specific attributes or how continuing occupancy of burrows is influenced by the surrounding habitat; especially post‐fire, when plant cover is reduced. Here, we test whether great desert skink burrows in areas burnt 2 years previously and in longer unburnt areas are associated with particular habitat attributes, and whether there are differences between occupied and recently abandoned burrow sites. Vegetation composition, cover and soil surface characteristics at 56 established great desert skink burrows, including occupied and recently unoccupied burrows, were compared with 56 random nearby non‐burrow control sites. Burrow sites had higher plant cover compared with the surrounding landscape in both recently burnt and longer unburnt areas and were more likely to be associated with the presence of shrubs. Soil stability and infiltration were also higher at burrow sites. However, we found no evidence that burrows with lower cover were more likely to be abandoned. Our results suggest that great desert skinks may actively select high cover areas for burrow construction, although differences between burrow and control sites may also partly reflect local changes to plant cover and composition and soil properties resulting from burrow construction and long‐term habitation of a site. Further research should determine if burrows with shrubs or higher plant cover provide greater protection from predators, more structural stability for burrow construction, increased prey abundance or other benefits. We recommend that maintenance of areas with relatively higher plant cover be prioritized when managing great desert skink habitat.     

Population dynamics of annual Salicornia species in the tidal salt marshes of the Oosterschelde,The Netherlands

1. The population dynamics of two Salicornia species from the Bergen op Zoom salt marsh (south-west Netherlands) was examined. Based on the results of several field studies three preliminary life tables were constructed, two for S. procumbens agg. populations growing respectively on the mud flats and in the salt marsh, and one for S. europaea agg. living in the upper marsh.
2. The life cycles are described and quantified in terms of eight phases and the transition probabilities between them, starting from a notional individual representative of each population.
3. The models depicting the life cycle of S. procumbens show a mean offspring number of 4.26 individuals per parent for the mud-flat population and 0.18 for the salt-marsh population. The S. europaea model gives an output of 0.44 individuals per parent. These results reflect the fluctuations in population size observed in sample plots over the years 1976–78.
4. Comparison of the transition probabilities reveals that on the mud flats most S. procumbens individuals die during pollination and seed germination, while the population in the salt marsh proper is thinned especially during the seed phase in winter time and during the growth from established seedlings to maturation. S. europaea behaves in a similar but less pronounced way to S. procumbens in the salt marsh.
5. Probabilities for one flower or one seed to produce a mature flowering plant were calculated, and were compared with those found in the literature. They are roughly of the same order of magnitude as the probabilities for other annual species, but much higher than those reported for biennial species.

     

Disturbance,emigration, and refugia: How the mud snail,Ilyanassa obsoleta (Say), affects the habitat distribution of an epifaunal amphipod,Microdeutopus gryllotalpa (Costa)

In the presence of the mud snail Ilyanassa obsoleta (Say), the tubicolous amphipod Microdeutopus gryllotalpa (Costa) emigrates to snail-free sediments, as demonstrated in laboratory and field experiments. Emigration occurs predominantly in the dark when the amphipod is most active. Unlike crevices, the thickness of sediments in which the amphipod is established offers no protection from snail disturbance. Emigration is shown to be caused by the disturbance generated by the snail's plowing and burrowing across the sediment surface, and not a response to a reduction in the shared microfloral food supply. The crawling and burrowing of the smaller mud snail, Hydrobia totteni Morrison, does not disturb Microdeutopus, supporting the hypothesis that relative body sizes affects the ability of bioturbators/burrowers to disturb tube-dwellers. As the burrows of Microdeutopus extend only ≈2 cm below the sediment surface, thick mud layers do not offer any refuge from Ilyanassa. However, very small solid surfaces (≈1–2 mm in relief) to which the amphipods build tubes do provide some protection from Ilyanassa. In soft-sediment benthic communities, such small structures may provide significant refuge for small epifauna and shallow burrowing infauna escaping from small-scale, biogenic disturbance.     

Effects of plateau pika (Ochotona curzoniae) on net ecosystem carbon exchange of grassland in the Three Rivers Headwaters region, Qinghai-Tibet, China

Background and aim

Because the indigenous burrowing lagomorph plateau pika (Ochotona curzoniae) is considered to have negative ecological impacts on alpine meadow steppe grasslands of the Headwaters Region of the Yellow, Yangtze and Mekong Rivers we investigated its effects on ecosystem productivity and soil properties, and especially net ecosystem carbon flux.

Methods

We measured net ecosystem CO₂ exchange (NEE) and its components gross ecosystem productivity (GEP) and ecosystem respiration (ER) at peak aboveground biomass by the chamber method with reference to plant and soil characteristics of areas of alpine meadow steppe with different densities of pika burrows.

Results

Higher burrow density decreased NEE, GEP and ER. Above-ground biomass, species number, plant cover and leaf area index decreased with increasing pika density. Higher burrow density was associated with lower soil moisture and higher soil temperature. Responses of NEE were related to changes of abiotic and biotic factors affecting its two components. NEE was positively related to soil moisture, soil ammonium nitrogen, plant cover, leaf area index and above-ground biomass but was negatively correlated with higher soil nitrate nitrogen.

Conclusion

Decrease of NEE by plateau pika may reduce the carbon sink balance of Qinghai-Tibet plateau grassland. Such effects may be influenced by grazing pressure from domestic livestock, population levels of natural predators, and climate change.     

Meandering worms: mechanics of undulatory burrowing in muds

Recent work has shown that muddy sediments are elastic solids through which animals extend burrows by fracture, whereas non-cohesive granular sands fluidize around some burrowers. These different mechanical responses are reflected in the morphologies and behaviours of their respective inhabitants. However, Armandia brevis, a mud-burrowing opheliid polychaete, lacks an expansible anterior consistent with fracturing mud, and instead uses undulatory movements similar to those of sandfish lizards that fluidize desert sands. Here, we show that A. brevis neither fractures nor fluidizes sediments, but instead uses a third mechanism, plastically rearranging sediment grains to create a burrow. The curvature of the undulating body fits meander geometry used to describe rivers, and changes in curvature driven by muscle contraction are similar for swimming and burrowing worms, indicating that the same gait is used in both sediments and water. Large calculated friction forces for undulatory burrowers suggest that sediment mechanics affect undulatory and peristaltic burrowers differently; undulatory burrowing may be more effective for small worms that live in sediments not compacted or cohesive enough to extend burrows by fracture.