Laboratory observations on the burrowing of the snake blenny, Lumpenus lampretaeformis (Walbaum), in soft sediment

The Lumpenus lampretaeformis used in this study were caught on soft sediments in Irvine Bay and Ardmucknish Bay (West Coast of Scotland). In aquaria the fish is capable of constructing and maintaining its own burrow in soft sediments. The burrow has a'Y'configuration which may be made complex by the construction of additional tunnels as the original system begins to decay. The cross sections of these burrows are distinctive in that they have an oval shape in the main tunnel and a rounded cross section in the subsidiary (side) tunnel. Burrowing can be relatively rapid, a burrow with three entrances being constructed within 12 h.     

Burrow Architecture of the Ghost Crab Ocypode ceratophthalma on a Sandy Shore in Hong Kong

The ghost crab Ocypode ceratophthalma (Pallas) creates burrows of variety shapes at different ages. Juveniles (mean carapace length 11 mm) produced shallow J-shaped burrows, which incline vertically into the substratum (mean depth 160 mm). Larger crabs (17–25 mm carapace length) have Y-shaped and spiral burrows (mean depth 361 mm). These Y-shaped burrows have a primary arm, which extends to the surface forming the opening, and a secondary arm which terminates in a blind spherical ending. The two arms join in a single shaft and end with a chamber at the base. The secondary arms and chambers are believed to be used for mating or as a refuge from predation. The spiral burrows have spiral single channel ending in a chamber. Older crabs (mean carapace length 32.6 mm) had simple, straight single tube burrows, which inclined into the substratum at mean of 73° and had a mean depth of 320 mm. During summer daytime periods, the burrows shelter the crabs from heat and desiccation stress. The sand surface temperature at the burrow opening was ~48 °C but temperatures inside the burrows can drop to 32 °C at a depth of 250 mm. Variation in the burrow architecture with crab age appears to be related to the crab’s behaviour. Juvenile crabs have smaller gill areas and move out of the burrows regularly to renew their respiratory water and, as a result, they do not need a deep burrow. Larger crabs, in contrast, can tolerate prolonged periods without renewing their respiratory water and therefore create deeper and more complex burrows for mating and refuges.     

Passive irrigation and functional morphology of crustacean burrows in a tropical mangrove swamp

Flushing measurements and a resin cast of a burrow inhabited by Sesarma messa and Alpheus cf macklay were taken from a Rhizophoraspp. forest. The burrow had 9 openings and occupied a swamp surface area of 0.64 m². Passive irrigation of the burrow was investigated by recording change in conductivity of burrow water in a chamber 45 cm below the swamp surface during tidal inundation of the swamp. The chamber was completely flushed within approximately one hour, i.e. by a single tidal event. Burrow morphology was determined by means of resin casting. The investigated burrow was of discrete structure, with an overall depth of 1.2 m and a total volume of 68 l, i.e. ca. 9% of the volume of swamp soil. The below ground surface area of chambers and tunnels was 3.8 m². The mean and maximum chamber/tunnel diameter was 7 cm and 11 cm respectively. The soil in the close vicinity of the burrow was extensively penetrated by roots, and any two parts of the burrow were located no further than 20 cm away from each other. By reducing diffusion distances within the soil and by being well flushed, the burrows provide an efficient mechanism for removal of excess salt accumulated in the soil around mangrove roots due to exclusion.     

The underground life of the oriental mole cricket: an analysis of burrow morphology

The underground life of the oriental mole cricket Gryllotalpa orientalis has been investigated by studying the structure of its burrows under different environmental situations and in different seasons. The different uses of different burrow types and their advantages and disadvantages have been examined. The total length, number of tunnels and combination of burrow types varied from a simple tunnel to a more complex one with branches at various angles to the surface, burrow types being divided roughly into shallow horizontal or deep vertical ones. In horizontal burrows, the branching structure was well developed in various directions. It is notable that the vertical burrows of G. orientalis were occupied by only one individual. Both vertical and horizontal burrows were used for foraging: vertical burrows for plants with subterranean stems and horizontal burrows for creeping plants. Vertical burrows were also used for hiding from predators, resting, moulting and overwintering, whereas horizontal burrows were used for escaping from predators and as mating routes. Egg chambers were constructed beside horizontal burrows, and calling burrows were constructed as part of horizontal burrows. Based on their current requirements, mole crickets continuously modify their burrow structures or change burrowing sites.     

Accurate quantification of the influence of benthic macro- and meio-fauna on the geometric properties of estuarine muds by micro computer tomography

This paper describes the novel application of high resolution micro computer tomography (microCT) to the quantification of the properties of marine biogenic structures. CT scanning has been used to examine sediments in the past but the resolution of most previous techniques has been dependent upon commercial medical CT scanners which only have a slice thickness of ≥ 0.625 mm. In addition adequate software has not previously been available to rapidly quantify all the properties of biogenic structures. The microCT technique developed here used a standard core sample of estuarine sediment and new software was developed to calculate the axial variation of the following burrow parameters: number, diameter, volume, surface area and density. The increased resolution has resulted in the first quantification of meiofaunal burrow structures.A test core has shown, as an example, that the total volume of burrows created by macrofaunal organisms decreased from 827 mm³, within the top 15 mm of the core, to 204.2 mm³ at a depth 60 -75 mm within the core. Total burrow surface area decreased from 1883 mm² to 512 mm², for these depth ranges, respectively and burrow diameter ranged from 2.37-2.58 mm, remaining fairly constant between depths. Meiofaunal burrow structures decreased from 1.3-0.1 mm³ within the top 6 mm of the core with burrow surface area decreasing from 33.52-3.4 mm². Again, burrow diameter remained relatively constant, ranging from 0.23-0.25 mm.Quantification to this resolution is required to identify the impact of infaunal organisms on factors such as oxygen penetration, vertical and horizontal (across burrow walls) gradients in redox conditions and chemical/nutrient speciation and flux. The quantification of these burrow properties will improve the ability to examine the interrelationships between chemical, physical and biological processes and their role in ecological functioning. The present study indicates that there is potential for further development of this software to allow more detailed analysis of burrow structures and surface features including parameters such as burrow length, shape and sediment surface roughness.     

Microscale oxygen distribution in various invertebrate burrow walls

Profiles of dissolved oxygen were measured in pore waters of unburrowed sediment and the burrow walls of seven invertebrate dwellings. Burrows studied include those of Corophium volutator, Heteromastus filiformis, Arenicola marina, Saccoglossus bromophenolosus, Clymenella sp., Hemigrapsus oregonensis and Cirriformia luxuriosa all from mudflats in Willapa Bay, Washington. These animals comprise a range of burrow architectures ranging from simple, unlined burrows to more complex, mucous lined burrows. Oxygen penetrated unburrowed sediment between depths of 0.4–2.6 mm, whereas oxygen penetrated the burrow walls from 0.3 mm to 2.3 mm. Three groups of burrows are recognized based on the oxygen diffusive properties relative to the unburrowed sediment including those that: (1) slightly impeded oxygen penetration, (2) clearly inhibited oxygen penetration, and (3) enhanced oxygen penetration. Differences in the diffusive properties of the burrow wall are related to the burrow microstructure and presumably the microbial communities living within the burrow microenvironment. The results of this study suggest that burrow shape and burrow‐wall architecture may play an important role in controlling the diffusion of oxygen, and possibly of other dissolved gases (i.e. CO₂, H₂S). The results further demonstrate that simplified assumptions (i.e. that bioturbation uniformly enhances oxygen diffusion into suboxic and anoxic sediments), while requisite for numerical modelling, are not necessarily representative of field data.     

Burrow architecture and digging activity in the Cape dune mole rat

While females are traditionally thought to invest more time and energy into parental care than males, males often invest more resources into searching and displaying for mates, obtaining mates and in male–male conflict. Solitary subterranean mammals perform these activities in a particularly challenging niche, necessitating energetically expensive burrowing to both search for mates and forage for food. This restriction presumably affects males more than females as the former are thought to dig longer tunnels that cover greater distances to search for females. We excavated burrow systems of male and female Cape dune mole rats Bathyergus suillus the, largest truly subterranean mammal, to investigate whether male burrows differ from those of females in ways that reflect mate searching by males. We consider burrow architecture (length, internal dimensions, fractal dimension of tunnel systems, number of nesting chambers and mole mounds on the surface) in relation to mating strategy. Males excavated significantly longer burrow systems with higher fractal dimensions and larger burrow areas than females. Male burrow systems were also significantly farther from one another than females were from other females' burrow systems. However, no sex differences were evident in tunnel cross-sectional area, mass of soil excavated per mound, number of mounds produced per unit burrow length or mass of soil excavated per burrow system. Hence, while males may use their habitat differently from females, they do not appear to differ in the dimensions of the tunnels they create. Thus, exploration and use of the habitat differs between the sexes, which may be a consequence of sex differences in mating behaviour and greater demands for food.     

The burrow system of the African ice rat Otomys sloggetti robertsi

We studied the architecture of the burrow system of the African ice rat Otomys sloggetti robertsi, a non–hibernating, diurnal murid rodent endemic to the sub–alpine and alpine regions of the southern African Drakensberg and Maluti mountains. In our study site we found ice rat burrows in two substrates (organic and mineral soils). The structure of the burrow system was similar in both soil types, comprising several interlinking tunnels, numerous burrow entrances and 1–2 nest chambers. However, the surface area of the burrow systems in organic soils was larger, the tunnels were deeper, and some of the systems contained two levels, all of which was contrary to our assumption that digging would be more difficult in the compact organic soils. Ice rats occur in colonies of up to 17 individuals, and the collected efforts of several individuals are required for constructing complex burrow systems. The burrow structure is similar to those of two arid–adapted relatives, Parotomys brantsii and Parotomys littledalei, suggesting that the burrow architecture among these three taxa may reflect the similar functions of burrows in extreme environments. For ice rats, burrows could provide a suitable microhabitat in which to escape adverse environmental conditions, particularly during winter. Moreover, ice rat burrows contained far fewer nest chambers than those of both Parotomys species, indicating that members in a colony share nest chambers, thereby facilitating huddling. Finally, the extensive interlinking tunnels may provide underground routes to aboveground feeding sites, thereby reducing exposure to adverse conditions.     

Intertidal burrows of the air-breathing eel goby, <Emphasis Type="Italic">Odontamblyopus lacepedii</Emphasis> (Gobiidae: Amblyopinae)

Odontamblyopus lacepedii inhabits burrows in mudflats and breathes air at the surface opening. Investigations of the intertidal burrows using resin casting demonstrated a highly branched burrow system. The burrows are composed primarily of branching patterns of interconnected tunnels and shafts that communicate into two to seven surface openings. Bulbous chambers (i.e., dilated portions of the burrow) at branching sections of the tunnels or shafts are common features of the burrow. The presence of these chambers accords the fish adequate space to maneuver inside the burrow, and thus constant access to the surface. The combination of all burrow characteristics and previously reported variability in air breathing patterns are ostensibly of selective value for aerial predator avoidance during air breathing in O. lacepedii.     

Identification versus counting protocols as sources of uncertainty in diatom-based ecological status assessments

The heterogeneity of oxygen distribution in a Hediste diversicolor burrow environment was investigated in a laboratory experiment using a 6-mm thick tank equipped with oxygen planar optodes. The two-dimensional oxygen distribution in a complete burrow was monitored every 2 min for 4 h. Oxygen concentrations fluctuated over a scale of minutes in the burrow lumen and wall (up to 2 mm) reflecting the balance between worm ventilation activity and oxygen consumption. The magnitude of the three surrounding micro-horizons (oxic, oscillating and anoxic) induced by the intermittent worm ventilation was spatially and temporally variable within the structure. Oxygen variations appeared to be controlled by distance from the sediment–water interface and the direction of water circulation. Moreover, there was an apparent ‘buffer effect’, induced by the proximity to the overlying water, which reduced the variations of lumen and wall oxygen in the upper part of the structure. These results highlight the heterogeneous distribution and dynamics of oxygen associated with H. diversicolor burrows and ventilation activity. They also highlight the necessity of integrating this complexity into the current burrow-base models in order to estimate the ecological importance of burrowing species in coastal ecosystems.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Burrow use by Tibetan Ground Tits Pseudopodoces humilis: coping with life at high altitudes

The Tibetan Ground Tit Pseudopodoces humilis is a high-altitude passerine endemic to the Tibetan Plateau. A 4-year study in alpine meadows in Northern Tibet at 4300 m asl demonstrated that rather than using Pika Ochotona spp. holes as previously reported, the birds excavated one nest burrow themselves in spring and another after breeding, which they used for roosting in winter. Both the nesting and winter-roosting burrows comprised a straight tunnel leading to an ellipsoid chamber. There were no significant differences in placement or structural characteristics between the two types of burrows, except that winter-roosting burrows had a significantly smaller entrance diameter. Most burrows were 100–160 cm long and their chambers 20–40 cm deep. Tibetan Ground Tits tended to maximize the thermal benefits of their burrows by adjusting their spatial and structural characteristics in response to local solar radiation and wind regimes. Burrows tended to be oriented towards the sun and away from prevailing winds, presumably to maintain burrow temperature. Longer tunnels could function to save heat from solar radiation or reduce wind-disturbance, while shorter tunnels allow chambers to be warmed sooner in situations where wind potentially reduced soil temperatures. The thermal benefits to the birds of burrow architecture are likely to play a crucial role throughout the year in these extreme alpine environments.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Burrow morphology and utilization of the goby (Parapocryptes serperaster) in the Mekong Delta,Vietnam

Some fish species living in mudflats construct burrows for dwelling and hiding. The goby Parapocryptes serperaster is a burrowing fish in mudflats of many estuaries in South East Asia. This study was carried out in the Mekong Delta, Vietnam, to examine burrow morphology and usage by this species. Morphology of the burrows constructed by P. serperaster was investigated by resin castings in situ to obtain the physical structure and configuration of each burrow. Fish from the burrows were caught and measured before burrow casts were made. Fish burrows comprised several openings, a few branching tunnels and multi-bulbous chambers. The surface openings were circular, and the shapes of branching tunnels were nearly round. The burrows had interconnected tunnels and various short cul-de-sac side branches. The burrow structure differed between fish sizes, but burrow dimensions were positively correlated with fish size, indicating that larger fish can make larger and more sophisticated burrow. The burrow structure and dimensions were not different between the dry and wet seasons. Laboratory observations showed that P. serperaster used body movements to dig burrows in the sediment. Burrows could provide a low-tide retreat and protection from predators, but were not used for spawning and feeding for this goby species. This study indicates that the burrowing activity of gobies is an important adaptation for living in shallow and muddy habitats.     

Burrow structure and foraging costs in the fossorial rodent,Thomomys bottae

Summary A model for calculating the energy cost of burrowing by fossorial rodents is presented and used to examine the energetics of foraging by burrowing. The pocket gopher Thomomys bottae (Rodentia: Geomyidae) digs burrows for access to food. Feeding tunnels of Thomomys are broken into segments by laterals to the surface that are used to dispose of excavated soil. Energy cost of burrowing depends on both soil type and on burrow structure, defined by the length of burrow segments, angle of ascent of laterals, depth of feeding tunnels, and burrow diameter. In a desert scrub habitat, Thomomys adjust burrow segment length to minimize cost of burrowing. Observed segment lengths (mean=1.33 m) closely approximate the minimum-cost segment length of 1.22 m. Minimizing energy expended per meter of tunnel constructed maximizes efficiency of foraging by burrowing in the desert scrub. Burrow diameter and cost of burrowing increase with body size, while benefits do not, so foraging by burrowing becomes less enconomical as body size increases. Maximum possible body size of fossorial mammals depends on habitat productivity and energy cost of burrowing in local soils.     

Influence of soil compaction on tunnel network construction by the eastern subterranean termite (Isoptera: Rhinotermitidae)

Eastern subterranean termites, Reticulitermes flavipes (Kollar), workers were introduced into arenas containing low, moderate, and high compaction builder's sand (1.05 g/cm3, 1.18 g/cm3 or 1.35 g/cm3 bulk densities, respectively), and they immediately began tunneling. Termites built the tunnel network significantly fastest in soil of low compaction compared with moderately or highly compacted soil. In soil of low compaction, 221.67 +/- 4.73 cm of total tunnel distance was constructed in 1 d compared with only 96 cm of tunneling in highly compacted soil. At 14 d, total tunnel distance averaged 216.83 +/- 4.56 cm in soil of low compaction compared with 169.70 +/- 4.10 and 181.18 +/- 6.13 cm in moderately and highly compacted soil, respectively. Decreases in total tunnel distance between 1 and 14 d were caused by backfilling of seldom-used tunnels. Termites did the majority of tunneling during the first day of introduction into arenas. In soil of low and moderate compaction, termites essentially constructed the entire tunnel network within the first day, only modifying it by backfilling or maintaining tunnels. In highly compacted soil, 53% of the final tunnel network was constructed during the first day, 87% was constructed by the third day, and 97% was constructed by the seventh day. Soil compaction did not affect the number of primary tunnels or the number and diameter of secondary tunnels. The angle between the secondary tunnel and primary tunnel also was not significantly affected by soil compaction. However, the number of secondary tunnels in soil of low compaction (5.89 +/- 0.51) was significantly greater than in moderately (2.74 +/- 0.36) and highly (3.58 +/- 0.59) compacted soils.     

The burrow ofDasypus hybridus (Cingulata: Dasypodidae)

Dasypus hybridus (Desmarest, 1804) inhabits open fields in southern South America. Burrows ofD. hybridus were cylindrical with a conical end. They had an entrance and a single tunnel without branches. Three locations of the burrows in the terrain were detected: in banks of dried waterways, near rocks, and in open field. The orientation of 29 burrow mouths was not random and, although they did not point to any cardinal point, in particular there was a strong tendency to avoid the south quadrant. In a sample area of 90 × 140 m, burrow mouths were arranged in a random spatial pattern with a density of 25.4 burrows per ha. Great variation in burrow length was found (118.8 ± 105.69 cm, CV = 89.0%). Tunnels may be used as refuges and/or for thermoregulation. The orientation of burrow mouths can also be related to thermoregulation as tunnels are covered from dominant winds, and, for many hours every day, the temperature at the mouths can be influenced by insolation. The possible existence of shelter-burrows and resting-burrows is discussed.     

Behavioural and physiological adaptations to a burrowing lifestyle in the snake blenny, Lumpenus lampretaeformis, and the red band-fish, Cepola rubescens

Observations have been made on the mode of burrow construction in the snake blenny, Lumpenus lampretaeformis , under laboratory conditions. It appears that head probing and lateral oscillations of the body are principally responsible for the excavation of the burrow which is completed within 24 h. The burrow structure has been analysed in detail, showing a mean depth of 7.2 cm with a maximum observed length of 73 cm, with most systems between 20 and 35 cm in length. Initially linear burrows with two openings are usually provided with a small side tunnel, giving the system a characteristic Y-shape.
Burrow irrigation was investigated for the first time in L. lampretaeformis. The mean duration of burrow irrigation, by flexions of the tail of the fish, was 21 s with over 13 min h⁻¹ spent in irrigating the burrow. The mean water displacement per irrigation period was 3.1 ml. The PO ₂ and PCO ₂ were measured in both surface water and within the burrow system of L. lampretaeformis. Surface water values for PO ₂ were high (> 150 Torr) and PCO ₂ low (<0.4 Torr). Hypoxic and hypercapnic conditions were measured in the burrow system itself, with PO ₂ values ranging between 57 and 129 Torr and PCO ₂ rising to > 1.3 Torr in some burrows.
A comparative study of Cepola rubescens burrows indicated similar surface water PO ₂ and PCO ₂ values as in L. lampretaeformis. Burrow water PO ₂ values ranged between 60 and 94 Torr, with PCO ₂ values as high as 1.5 Torr being recorded. These results are discussed in relation to the adaptation of both species to a burrowing lifestyle.     

Burrow morphology and dynamics of mudshrimp in Asian soft shores

Mudshrimps are important soft shore bioturbators but research on the ecology of tropical species has received less attention when compared with their temperate counterparts. The mudshrimp Austinogebia edulis is common on Asian soft shores and lives in burrows for its entire adult life. Epoxy resin casting of A. edulis burrows showed that they were approximately Y-shaped, with an upper U-part and the lower central shaft part. The burrows had two openings extending to the surface; the mean distance between the two openings was 11.0 cm in Hong Kong and 26.4 cm in Taiwan. Openings of the burrows had small chimneys. The tunnels of the burrows were circular, narrow and with a smooth surface (tunnel diameter corresponded to shrimp carapace width). Each burrow was inhabited by a single shrimp and burrows were inter-connected during the mating and reproductive season. Each burrow had four to 12 spherical chambers, which were free of detritus. The chambers were thought to be used for suspension feeding, current generation and as turning points. The depth of burrows was up to 1.1 m. Multivariate analysis on various burrow parameters showed that burrows collected on a mud flat in Taiwan were deeper, had a wider distance between the openings and a larger volume than burrows collected from a sandy shore in Hong Kong, suggesting that burrow architecture is variable between shore types. Burrow architecture, however, did not vary between tidal levels, seasons and shrimp density on the shores in Hong Kong, indicating that the burrows were quite stable within the substratum and were not affected by environmental and biological factors.     

Comparative studies of the branchial morphology, gill area and gill ultrastructure of some thalassinidean mud-shrimps (Crustacea: Decapoda: Thalassinidea)

The morphology, gill area and branchial formulae of six thalassinidean decapods ( Calocaris macandreae, Jaxea nocturna, Callianassa subterranea, Upogebia stellata, U. deltaura and U. pusilla ) are reported. Additionally, the rarely-encountered Axius stirhynchus receives brief attention. Gill formulae are presented; the simplest arrangements are found in the Callianassidae and Upogebiidae. The deeper-burrowing, deposit-feeding species that are regularly exposed to prolonged periods of hypoxia, i.e. Callianassa subterranea and Jaxea nocturna , had significantly larger gill areas than thalassinideans that occupied more oxygenated burrows ( Upogebia spp., Calocaris macandreae ). The increase in gill area was a result of flattening of the trichobranchiate gill filaments giving rise to a phylloid gill form. It is suggested that the efficiency of gas transfer, and hence diffusing capacity, was enhanced in the phylloid gill by the larger gill area and, because of the reduced cuticle thickness, by the shorter water-haemolymph diffusion distance. The increased diffusion capacity of the phylloid gill is interpreted as a functional adaptation to the more severe physicochemical burrow water conditions experienced by Callianassa subterranea and Jaxea nocturna .