The palaeoecology of the Vombatidae: did giant wombats burrow?

Debate over the origins of burrowing in the Vombatidae has continued since the discovery of the remains of the largest of all wombats, Phascolonus gigas, in the nineteenth century. In this paper, we argue that the largest of the ancestors of extant wombats did not burrow due to physical and physiological limitations of burrows. Further, we suggest that the burrowing characteristics of the extant wombats were derived from an ancestor of similar body size (20–40 kg) that is presently not represented in the fossil record.     

Spatial ecology of the mulgara in arid Australia: impact of fire history on home range size and burrow use

Knowledge about the spatial ecology of small mammals in relation to fire history in arid zones in general and Australia in particular is limited. Here, we report data on the spatial ecology of the brush-tailed mulgara Dasycercus blythi in the hummock grasslands of Ulu r u – Kata Tju t a National Park during winter 2006, the beginning of the breeding season for this species. About 73% of the study area had been burnt in 2002 and spinifex cover was sparse. Mulgaras Marsupialia: Dasyuridae (six males and three females) were implanted with radio-transmitters and monitored daily for between 6 and 55 days. All mulgaras appeared to use defined home ranges, which overlapped extensively with those of several neighbours. Spatial overlap occurred between as well as within sexes. On average, males (25.5 ha) occupied significantly larger home ranges than females (10.8 ha). Mulgaras used a number of burrows within home ranges and several were used by more than one individual. Moreover, occasionally, two individuals used the same burrow simultaneously. Home ranges and burrows encompassed both mature spinifex Triodia basedowii and open regrowth areas and mulgaras did not exhibit a significant preference for either habitat type. However, three males were killed by introduced-predators and they all lived predominantly in the open regrowth area. We conclude that mulgaras do not select the dense cover of mature spinifex habitat, and might be subjected to increased risk from introduced predators, especially following fire.     

Hematologic and serum biochemical reference values for free-ranging northern hairy-nosed wombats

Hematologic and serum biochemistry values were determined for 31 adult (21 male and 10 female) and four subadult male northern hairy-nosed wombats (Lasiorhinus krefftii) from the only existing population in Epping Forest National Park, Australia. Blood samples were obtained from free-ranging northern hairy-nosed wombats during trapping for population census and health and reproductive assessment in 1999. Hematologic and biochemical values were compared between adult males and adult females, and between adult and subadult wombats. Values were also compared with those previously published for southern hairy-nosed (Lasiorhinus latifrons) and common (Vombatus ursinus) wombats. The values from this study were used to create reference intervals, and they make up the first comprehensive hematologic and biochemical study for this highly endangered species.     

Size as a Factor Influencing the Ownership of Copulation Burrows by the Ghost Crab (Ocypode ceratophthalmus)

On Cousin Island, Seychelles, male ghost crabs mostly constructed their copulation burrows around new moon. The burrows were sited below the high tide mark and therefore only lasted until washed away by the incoming tide. Large males were able to defeat small males in contests for burrows and so, to reduce the risk of eviction, small males built their burrows later on the ebbing tide than large males, after the period when fights for burrows were most frequent. There was also evidence that small males built their burrows on stretches of beach presumed to be inferior, because the burrow density was lower.     

Resources driving landscape‐scale distribution patterns of grazers in an African savanna

In order to effectively manage and conserve indigenous herbivores, a good understanding is needed of how resources drive their distribution patterns. This study employed a unique dataset to test a range of ecological theories and hypotheses on free‐ranging grazers. Using aerial census data collected over 14 yr across the 2 million ha Kruger National Park (South Africa), this study employs spatial autologistic regression models to explore the spatial relationships that exist between the distribution of eight indigenous grazer species and a set of resource variables. It was found that ecological theories relating to feeding guild, water‐dependence, allometric scaling, gut‐morphology and vulnerability to predation could explain most of the grazer distribution patterns observed in relation to surface‐water, forage quality, forage quantity and habitat openness. All the grazers studied were water‐dependent and occurred close to a permanent source of water in the dry season. This was ascribed to the lack of moisture in the diet of grazers during the dry season. Most ruminants’ distribution patterns were significantly associated with areas of high forage quality whereas hind‐gut fermentors were neutral towards forage quality. Average forage quantity was not a significant predictor of long‐term, landscape‐scale distribution patterns for any of the grazer species studied. Most small‐ and medium‐bodied grazers preferred open habitats above closed habitats, probably due to higher visibility and a lower predation risk. Large‐bodied grazers did not bias their distribution patterns towards open habitats. The way in which grazers distribute themselves with respect to different resources can potentially inform management actions on appropriate scales.     

Dispersal and social organization of the northern hairy-nosed wombat Lasiorhinus krefftii

Burrows of the northern hairy-nosed wombat were arranged in loose clusters, each of which was used in common by a group of up to 10 wombats. However, individual wombats rarely used the same burrows on the same days and feeding ranges tended to be separate within each sex. The incidence of breeding dispersal among females was quite high (at least 50%); dispersal distances ranged up to almost the full extent of the population's range. Dispersal by adult males was rare, and no juvenile males were observed to disperse.     

Distribution of life cycle stages of Sarcoptes scabiei var wombati and effects of severe mange on common wombats in Victoria

Seven female and three male common wombats (Vombatus ursinus) collected from forested areas of Victoria (Australia) over a 10 mo period, 10 April 1997 to 22 February 1998 had at least 30% of their skin affected by severe hyperkeratotic sarcoptic mange. Mangy wombats were grazing during the day, could be readily approached, were in poor body condition, and lacked subcutaneous fat. The anterolateral surface of the body was most heavily parasitised with Sarcoptes scabiei var wombati followed by the posterolateral surface, the dorsal region between the ears, the ears, ventral abdomen, medial aspect of the legs, axillary and inguinal areas, and the dorsal midline. Larvae were the most prevalent life-cycle stage followed by eggs, nymphs, females, and males. Mite numbers and the severity of clinical signs, namely thickness of scale crust and the degree of alopecia, were correlated and were symmetrical on each side of the body. Fissuring of crust and skin only occurred when scale crust was present. Bacterial infections occurred in three of 10 wombats within lymph nodes or the pleural cavity. Lymphoid depletion did not occur in lymph nodes or spleens and prescapular lymph nodes contained a greater amount of nuclear debris in germinal centres than non-mangy wombats. Seven wombats had fatty change in their livers. Gonads of mature wombats were not active or had minimal activity. Significant histopathological changes were not seen in the gastrointestinal tract, kidney, brain, myocardium, spleen, thyroid, reproductive tract, and gonads. Hematocrit, mean corpuscular volume, mean corpuscular hemoglobin, and concentrations of hemoglobin, lymphocytes, calcium, glucose, creatinine, total solids, total protein, albumin determined both colormetrically and electrophoretically, and globulins were significantly lower and concentrations of neutrophils, monocytes, phosphorus, urea, glutamate dehydrogenase, aspartate aminotransferase and creatine kinase were significantly higher in mangy versus captive wombats. Concentrations of erythrocytes, mean corpuscular hemoglobin, leucocytes, band neutrophils, eosinophils, nucleated erythrocytes, sodium, potassium, chloride, total bilirubin, alkaline phosphatase, and gamma glutamyltransferase for mangy wombats were not significantly different from that reported for captive wombats. Hematological and pathological changes in mangy wombats were consistent with anemia, inflammation, and changes seen with starvation.     

Nesting activity,breeding success and colony size for the Wedge-tailed Shearwater Puffinus pacificus on Heron Island

Abstract The Capricorn Group of islands in Australia's Great Barrier Reef Marine Park sustains one of the world's largest breeding populations of the Wedge-tailed Shearwater Puffinus pacificus. Heron Island, a 13.5 ha coral cay which supports tourist and research station leases as well as a national park, is the third largest nesting island in the group. Sample censuses of breeding burrows were conducted each year between 1985 and 1990 and a further survey was completed in 1993. These returned estimates of between 13 264±1387 and 16 337±1545 active burrows (Y±SE). Burrow densities within each of the habitats monitored showed no significant trends between years, although there were large differences in burrow density between habitats. There were roughly the same number of burrows in the developed (west) and national park (east) halves of the cay. A miniature video camera system (burrowscope), which allowed nesting chambers at the ends of burrows to be inspected, was used in 1989, 1990 and 1993. This demonstrated that around half the burrows were occupied by incubating birds. Variations were found in the distribution of incubating birds between habitats, although this did not remain constant between the years. In the 1993 season, breeding activity was traced from the burrow establishment to fledging stage. Fifty-one per cent of burrows were used for breeding (eggs laid), 77% of eggs hatched and 80% of chicks produced a fledgling. Overall breeding success for the island was estimated at 61%. In 1993 the area designated as Buildings was found to have significantly lower hatching success compared with natural habitats. Most mortality occurred at the egg stage; however, in the Fringe habitat, mortality was highest at the chick stage. Previous surveys have estimated the breeding population from burrow counts. It now appears that only about 30% of such burrows produce fledglings.     

Genetic marker investigation of the source and impact of predation on a highly endangered species

In September and October 2000, the remains of a number of apparently predated northern hairy-nosed wombats (Lasiorhinus krefftii) were discovered in Epping Forest National Park, the site of the only known population of this highly endangered species. Analysis of DNA recovered from six carcasses and a section of intestine found nearby was carried out using microsatellite and Y-specific primers. This identified seven individual wombats, the identity of three of which was inferred from a genotype database prepared from animals sampled during trapping programmes. Six victims were male and one female, suggesting that female-biased predation rates are unlikely to be the cause of the current male-biased population sex ratio. DNA isolated from four canid faeces found in the vicinity revealed three distinct canid microsatellite genotypes with very high probabilities of belonging to dingoes (Canis familiaris dingo). A wombat genotype matching that of one of the dead individuals was identified from scats of two of the dingoes. In addition, two macropod microsatellites were amplified from two dingo scats. These observations provided vital information regarding predation on northern hairy-nosed wombats, and prompted the permanent exclusion of dingoes from the park by the erection of a dingo-proof fence.     

Trace fossils and rhizoliths as a tool for sedimentological and palaeoenvironmental analysis of ancient continental evaporite successions

Recognition of evaporite formations from continental Tertiary basins of Spain provides evidence that trace fossils (including rhizoliths) can be abundant in some saline lake systems and their study helps in palaeoenvironmental interpretation of ancient continental evaporite sequences. Six main types of trace fossils have been distinguished and include: (1) networks of small rhizoliths; (2) large rhizoliths; (3) tangle-patterned small burrows; (4) isolated large burrows; (5) L-shaped traces; and (6) vertebrate tracks. Rhizoliths were related to both marginal areas of hypersaline lakes and lakes of moderately high saline waters. In these settings, pedoturbation resulted from colonization by grasses and bushes of distinct lake subenvironments. The activity of burrowing invertebrate faunas was especially intense in lakes of moderately concentrated brines from which gypsum was the main evaporite mineral deposited. A specific gypsum lithofacies (‘bioturbated gypsum deposits') forming thick, massive beds has a widespread occurrence in many of the basins. Tangle-patterned small burrows and minor isolated large burrows constitute the typical trace fossil types within the gypsum. The traces are interpreted as having been caused by burrowing insect larvae, probably chironomids, coleopterans and annelids. The behaviour of these organisms in recent lake environments yields information about the salinity range of lake waters from which gypsum precipitated. Concentration values averaging 100–150 g/l may be thus deduced though some organisms involved in the formation of the traces can tolerate higher salinities. The combined analysis of lithofacies and trace fossils from the lacustrine evaporite sequences contributes to the study of distinct saline lake subenvironments as well as changes in the sedimentary evolution of the lake systems. Consequently, trace fossils can provide valuable insight for palaeoenvironmental analysis of at least some evaporite formations that accumulated in continental settings.     

Socio-spatial Organization and Activity Distribution of the Alpine Marmot Marmota marmota: Preliminary Results

The behavioural ecology of the Alpine marmot Marmota marmota (Linné, 1758) was studied Jun.—Sep. 1990 in the Vanoise National Park (French Alps). We describe the socio-spatial structure of a high-altitude population, to give additional information on the formerly unclear marmot social organization. The social unit was the family group, with a common home range between 0.9 and 2.8 ha; a slight overlapping occurred (9 to 12.5%). Each home range had a central area of main burrows, where the hibernaculum was located, and peripheral areas. Space utilization and distribution of activities during the season were analysed. Some activities took place exclusively in the centre area while peripheral parts were used for foraging. This latter increased in Aug. and decreased in Sep., marmots tending to remain more and more at the centre area as the hibernation period approached.     

The Distribution of Grazing Pressure in Relation to Vegetation Resources in Semi-arid West Africa: The Role of Herding

In semi-arid West Africa, livestock are increasingly managed by sedentary producers in close proximity to expanding cropped lands. To evaluate the agricultural and environmental implications of this trend, a study was conducted to investigate the effect of grazing management on the spatial distribution of grazing pressure, the forage provided animals during the grazing period, and local herd-forage ratios across three agropastoral landscapes characterized by varying cultivation pressure. During the 19-month study period, data on herbaceous vegetation, livestock populations, and grazing itineraries were collected. These data were referenced to land units averaging 70 ha in area. Using this approach, each of 3,819 grazing itineraries was characterized as to: 1. the sum of the products of the palatable forage mass of a particular land unit and the time spent grazing by the herd within that unit (FAT, expressed in kg-hours ha⁻¹); and 2. the average palatable herbaceous forage mass encountered by livestock across the itinerary weighted by the time spent in the land units crossed (FA, expressed in kg ha⁻¹). The spatial dispersion of livestock grazing around human settlements was found to decline with a reduction in herding labor investment (herded>herd-release>free pasture). Multiple regression analyses of itinerary data demonstrate that both FAT and FA also decline with a reduction in herding labor investment. Herded and herd-release managed livestock were offered more palatable forage and grazed areas of higher forage availability than free-pastured animals. This supports arguments that as the investment of time and effort into herding declines, feed supply to livestock will decline and the potential for grazing-induced environmental change will increase.     

Phytomass utilization and deposition of feces by ungulates on steppe pastures of eastern Mongolia

The amount of phytomass removed by a complex of livestock (horses, cattle, sheep, and goats) and wild ungulates (Mongolian gazelle Procapra gutturosa Pall.) grazing in plain and mountain pastures of eastern Mongolia has been estimated by taking account of feces deposited by these animals. The results show that at an animal density of up to 30 head/km², the total annual amount of feces reaches 140 kg/ha (dry weight), with the greater part (up to 90 kg/ha) being deposited by horses. The contribution of Mongolian gazelles in some pastures reaches 20–40 kg/ha per year. Decomposition of feces proceeds very slowly, with the annual loss of their weight averaging only 9–12%. This is evidence for gradual accumulation of nondecomposed matter in the soil. The removal of phytomass by the complex of ungulates, calculated from the amount of feces with regard to their annual loss and forage digestibility, varies in different grazing areas from 240 to 400 kg/ha (25–60% of the total aboveground phytomass). The greatest amount of phytomass is utilized by horses, reaching 200 kg/ha (13%), and Mongolian gazelles utilize up to 86 kg/ha per year. In the growing season, ungulates remove no more than 11–16% of the total aboveground phytomass. It is concluded that the impact of total ungulate stock does not impair the productivity of vegetation in the pastures studied.     

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.     

Use of space by the African Striped Ground Squirrel Xerus erythropus

Five Striped Ground Squirrels Xerus erythropus were radio-tracked at Katumani, Kenya, for periods of up to 10 days. The results were used to explore their short-term use of space. Definitions of the terms 'home range' and 'core', and some aspects of the methodology of home-range analysis are discussed. In an area of low-productivity bushland, subject to intermittent drought, where their food supply was patchy and unpredictable, the squirrels travelled about 1.5 km per day, and occupied rather large home ranges averaging 12.4 ha. Levels of aggression between conspecifics was low, home ranges were not defended, and there was considerable range overlap. Nights were spent in burrows which were commonly shared with conspecifics. On most days a single relatively long exploratory excursion was made outside the home range, extending the average area of the total range to some 40 ha. Despite the presence of major environmental features which might have been expected to affect the squirrels' foraging behaviour, such as cultivated fields, home range use was very symmetrical, suggesting that essential resources (food, shade, burrows) were widely dispersed throughout the range. Squirrels foraged alone, harvesting small, scattered, high-quality food items, mainly vegetable. Activity was interrupted from time to time to retire to the shade to lose heat, and a midday rest was taken. Only part of the home range was used each day. A variable degree of central concentration of activity could be observed. The squirrels' use of space enabled them to exploit available food resources opportunistically and efficiently, and made them difficult to control effectively.     

ECOLOGY AND EVOLUTION OF MATING SYSTEMS OF FIDDLER CRABS (GENUS UCA)

1. General accounts of the natural history and behaviour of fiddler crabs suggest there exist two broad mating patterns in the genus. Most western and Indo-Pacific species mate on the surface of intertidal substrates near burrows females defend. The sexes associate only briefly during courtship and mating. In contrast, males of many American species court from and defend burrows to which females come for mating. Copulation occurs underground in burrows plugged at the surface; the sexes usually remain together for at least several hours. Here we summarize and contrast recent detailed field studies of the mating systems of U. pugilator, an American species, and U. vocans, a species widely distributed in the western and Indo-Pacific. We indicate how differences in the breeding ecology of these two species may account for basic differences in modes of sexual selection leading to the two broad mating patterns in the genus. 2. U. pugilator burrows in protected sandy substrates in the upper intertidal and supratidal zone. During ebb tide, nonbreeding crabs leave burrows they occupy during high tide to forage on food-rich substrates in the lower intertidal zone. Reproductively active males remain in the burrow zone where they fight for and defend burrows from which they court. Large males win most fights for burrows and tend to defend burrows high on the elevation gradient, especially during periods with relatively high tides. Females usually approach and descend the burrows of several males before choosing their mates by remaining in males' burrows. Males remain underground with their mates for 1–3 days until after they oviposit their eggs. Some males then emerge and leave their burrows while others sequester their mates in the chambers where mating and oviposition has occurred, dig new chambers and resume courtship, perhaps attracting additional females. In either case, females remain underground for approximately 2 weeks, finally emerging to release their planktonic larvae. Burrows that do not collapse due to tidal inundation or flooding by groundwater are best for breeding and usually are located relatively high on the elevation gradient. Females choose mates indirectly by preferring to breed in burrows that will remain intact while they oviposit and incubate their eggs. Large males mate more often than small males because they are better able to defend burrows at locations females prefer to breed. The mating system of U. pugilator may be classified as resource-defence polygyny. 3. U. vocans burrows in open muddy substrates in the mid- to lower intertidal zone. At a site near Chunda Bay, Australia, where the reproductive behaviour of this species has been studied in depth, both sexes feed near burrows they defend. Females tend to occupy their burrows for longer periods and move shorter distances than do males. Mating occurs on the surface near the burrows that females defend. Females accept both resident and wandering males as mates. They show no preference for mating with larger males. Female choice may be based on other male morphological or behavioural characteristics. Females oviposit their eggs either while on the surface or in their burrows. They produce relatively small clutches and are active on the surface throughout their breeding periods. Males fight both their neighbours and wandering males. Large males tend to win fights and defend burrows in areas where large females, which produce relatively many eggs, are most dense. Such areas may offer greater protection from predators than areas occupied by smaller females. Small males mate about as often as large males but may father fewer larvae. The mating system of U. vocans is resource-free and promiscuous. 4. The mating systems of U. pugilator and U. vocans differ fundamentally in that female U. pugilator require access to a specific microenvironment to breed successfully, while female U. vocans do not. We suggest this difference occurs because of contrasts in clutch sizes and the mobility and movement patterns of feeding females. Female U. pugilator produce relatively large clutches and probably experience more intense selection from factors that can cause egg loss and mortality than do U. oocans, which produce clutches of sufficiently small volume to be protected by their abdominal flaps. Hence, the range of suitable breeding environments for U. pugilator is small compared to that for U. vocans. In addition, U. pugilator burrows in areas that are relatively food-poor, leading to daily migrations to and from food-rich substrates in the lower intertidal zone, preventing female defence of an area suitable for both breeding and feeding. U. vocans, however, burrows in areas sufficiently rich to support feeding, leading to relatively low female mobility and defence of burrows that are also suitable breeding sites. 5. Adaptive radiation of the genus Uca in the Americas is manifest by trends toward smaller adult size, higher population densities, more frequent microgeographic sympatry and increased terrestriality, compared to species in the western and Indo-Pacific regions. We outline the general features of the selection mechanisms tying each of these trends to the evolution of resource—defence mating systems. Intraspecific variation in the courtship behaviour and site of mating in U. lactea and U. vocans supports our contention that resourse—defence behaviour tends to occur at high population densities. Additional data are needed to evaluate the other hypotheses critically.     

Spiders provide important resources for an endangered lizard

Conservation management requires knowledge of how a target species interacts with other species. Some relatively common species can modify the environment to the advantage of rarer, endangered species. Thus, local enhancement of those common species can positively influence remaining populations of the rarer species. The endangered pygmy bluetongue lizard Tiliqua adelaidensis inhabits burrows that are constructed by lycosid and mygalomorph spiders. We recorded 490 burrows in a 1 ha plot at the end of one season, and then observed at regular intervals the formation and loss of burrows, and the changes in occupancy status of each burrow over the next season. We found spiders in 94% of all newly constructed burrows and deduced that they had built the burrows. We found no evidence that lizards dug new burrows or deepened existing burrows. The numbers of both lizards and spiders in the burrows declined over the spring and summer, with lizards moving from their burrows more often early in the season than later. However, there was no strong trend for lizards to replace spiders in burrows. In fact, lizards tended to occupy deeper burrows than spiders, suggesting little negative impact of lizards on spiders. However, spiders had a positive impact on lizards by providing the refuge burrows central for lizard survival. Although lizards readily accept artificial burrows, long-term conservation for the lizards must include viable spider populations to maintain a supply of suitable burrow refuges.     

Small- and large-scale effect of the SW Atlantic burrowing crab Chasmagnathus granulatus on habitat use by migratory shorebirds

Here we address the question of whether the presence of the burrowing crabs Chasmagnathus granulatus affects small- and large-scale habitat use by migrant shorebirds. This crab is the dominant species in soft bare sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 42°S). They generate very extensive burrow beds in soft bottom intertidal areas. Our information shows that this burrowing crab affects the small-scale habitat use by shorebirds, given that shorebirds never walk through the funnel-shaped entrances of burrows. Given that crab burrow entrances occupy up to 40% of the intertidal area, there is a large decrease of available shorebird habitat in crab beds, restricting their activity to the spaces between the burrows. The southern migratory shorebird Charadrius falklandicus maximize the use of these areas by foraging closer to the burrows than the other bird species. Neotropical migrants, such as Calidris fuscicollis, Pluvialis squatarola and Tringa melanoleuca, used foraging paths that tended to maximize the distance from burrows, especially the distance to larger burrows. A field experiment showed that this was not necessarily due to a decrease in the availability of polychaetes near the crab burrows. A combination of landscape measurements and satellite images showed that crab beds covered up to 40% of the intertidal area of the Mar Chiquita coastal lagoon (37°40′S, Argentina), and nearly 100% of the intertidal area of the Bahia Blanca estuary (38°48′-39°25′S, Argentina). These two estuaries are located along the migratory flyway of Neotropical migratory shorebirds, but the Bahia Blanca estuary (area∼110,000 ha) shows a much lower shorebird diversity than Mar Chiquita (area∼4500 ha). The most common species in Bahia Blanca is the two-banded plover C. falklandicus, the species least affected by crabs at Mar Chiquita and which prefers to use high-density crab areas as foraging sites. The oystercatcher Haematopus palliatus was also most abundant in high-density crab areas, but they used these areas for resting. The abundances of preys varied during the study period and between the crab density areas, indicating that the use of these areas by birds is independent of crab density. However, burrowing crabs affect the depth distribution of polychaete and thus their availability to shorebirds. We suggest that this shorebirds-burrowing organism interaction could be generalized for other intertidal estuarine habitats.     

Underground mating in the fiddler crab Uca tetragonon: the association between female life history traits and male mating tactics

Brood size and other life-history traits of females affect male investment in mating. Female Uca tetragonon, producing relatively small broods, were attracted to the burrows of males for underground mating (UM) while carrying eggs. Most UM females released larvae and ovulated new broods during the pairing, averaging 3.9 days. While a female was incubating one brood, another brood was developing within the ovaries because the females were feeding adequately during incubation. These findings suggest that in U. tetragonon, a small-brood species, females increase the total number of broods produced by breeding continually. In contrast, in large-brood species, feeding by ovigerous females is relatively brief and not enough to prepare the next brood during incubation, inducing temporal separation between incubation and brood production. Unlike females in other ocypodids where females with large broods remain in the breeding burrows of males, most female U. tetragonon left the male after UM. Wandering in female U. tetragonon after the pairs separate may occur because their small broods are adequately protected by an abdominal flap. Relative brood size probably determines the vulnerability of the incubated broods to the females' surface behavior. Hence, male reproductive success in large-brood species may decrease greatly if males expel their mates after ovulation, although this is not necessarily so in small-brood species. Whether the male drives away the female or not may depend on which behavior within either small- or large-brood species yields the greater male reproductive success. In U. tetragonon some females extruded eggs in their own burrows after surface mating as well as in males' burrows after UM. It was unclear whether females chose a male with a larger burrow as an UM mate unlike several large-brood species. Burrows of both UM males and ovigerous females in U. tetragonon were relatively smaller than those in some large-brood species, indicating that incubation of small broods does not require large burrows. Rather than benefits of UM by female choice, wandering resulting from intersexual conflict, and sperm competition may explain why some females mate in males' burrows in this small-brood species.