Sociality in New World hystricognath rodents is linked to predators and burrow digging

The importance of predation and burrow digging in explainingthe evolution of sociality is generally unclear. We focusedon New World hystricognath rodents to evaluate three key predictionsof the predation hypothesis. First, large-bodied surface-dwellingspecies will be more vulnerable because they are more detectable;thus sociality should be associated with body size. Second,surface-dwelling, diurnal species would be more vulnerable topredators than nocturnal species; thus sociality should be associatedwith the evolution of diurnality. Third, species living in openhabitats will be more vulnerable; thus sociality should evolvein species living in open habitats. Regarding the importanceof burrows, we tested if species that dig burrows can benefitfrom communal labor; thus, sociality should be associated withburrow digging. All traits had significant phylogenetic signal,thus comparative analyses should explicitly address this. Ina comparative analysis on independent contrasts we found thatsociality was correlated with body size (larger species weremore social), diurnality (diurnal species were more social),and burrowing (burrowing species were more social), but we foundno effect of overhead plant cover of habitat on sociality inhystricognath rodents. Somewhat different results were foundwhen we analyzed the raw data. Taken together, our results providesupport for a link between predation risk, burrow digging, andsociality in this group.     

Is communal burrowing or burrow sharing a benefit of group living in the lesser cavy <Emphasis Type="Italic">Microcavia australis</Emphasis>?

Burrow systems play an important role in the life of rodents in arid environments. The objectives of this study were to examine the hypothesis that group living is beneficial to the semifossorial rodent, and determine whether Microcavia australis (Geoffroy and d’Orbigny, 1833) burrows communally and/or shares burrow systems. I related the structure of burrow systems to the number of cavies inhabiting them, in two habitats with different soil hardness and different plant cover (El Leoncito and Ñacuñán). El Leoncito has a harsh climate, with lower plant density and softer soil than Ñacñuán. A total of 18 burrow systems were characterized at Ñacuñán, and 12 at El Leoncito. Social groups at El Leoncito have a higher number of individuals than at Ñacuñán, but the structure of burrow systems in softer soil is narrower (small area size), with fewer holes, less slope and depth of galleries, and with no relationship between the number of holes and burrow area. Therefore, considering the development of the burrow system as an indicator of the cost of burrowing, I conclude that communal burrowing to reduce the energetic cost of burrowing per capita is not the primary cause of cavy sociality. M. australis were not active diggers, because digging behaviour was rarely recorded at either site. Burrow systems of cavies persisted over the years of study, occupied by the same cavies and new offspring, and digging new burrow systems and tunnels was a relatively rare event at both sites. Under the burrow-sharing hypothesis, sociality could prevail in M. australis that regularly dig to build and maintain a burrow system which they use for a long time.     

The impact of digging on the evolution of the rodent mandible

There are two main (but not mutually exclusive) methods by which subterranean rodents construct burrows: chisel-tooth digging, where large incisors are used to dig through soil; and scratch digging, where forelimbs and claws are used to dig instead of incisors. A previous study by the authors showed that upper incisors of chisel-tooth diggers were better adapted to dig but the overall cranial morphology within the rodent sample was not significantly different. This study analyzed the lower incisors and mandibles of the specimens used in the previous study to show the impact of chisel-tooth digging on the rodent mandible. We compared lower incisors and mandibular shape of chisel-tooth digging rodents with nonchisel-tooth digging rodents to see if there were morphological differences between the two groups. The shape of incisors was quantified using incisor radius of curvature and second moment of area (SMA). Mandibular shape was quantified using landmark based geometric morphometrics. We found that lower incisor shape was strongly influenced by digging group using a Generalized Phylogenetic ancova (analysis of covariance). A phylogenetic Procrustes anova (analysis of variance) showed that mandibular shape of chisel-tooth digging rodents was also significantly different from nonchisel-tooth digging rodents. The phylogenetic signal of incisor radius of curvature was weak, whereas that of incisor SMA and mandibular shape was significant. This is despite the analyses revealing significant differences in the shape of both mandibles and incisors between digging groups. In conclusion, we showed that although the mandible and incisor of rodents are influenced by function, there is also a degree of phylogenetic affinity that shapes the rodent mandibular apparatus.     

Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.     

Cape ground squirrels as ecosystem engineers: modifying habitat for plants,small mammals and beetles in Namib Desert grasslands

Burrowing and foraging of semi‐fossorial rodents can affect species distribution and composition. Ground squirrels dig large burrow systems for refuge from predators and temperature extremes. Burrowing and foraging around burrows by squirrels may affect habitat and resource distributions for other organisms. We examined the impact of Cape ground squirrels (Xerus inauris) on vegetation, small mammals and beetles during winter and summer in grasslands on the edge of the Namib Desert. At each burrow system and paired control site without burrows, we estimated plant cover and height using quadrats (N = 8 paired sites), small mammal abundance and species richness using mark‐recapture techniques (N = 8 paired sites) and beetle abundance and species richness using pitfall traps (N = 6 paired sites, winter only). Squirrel burrowing and foraging activities resulted in lower plant cover and height, higher small mammal abundance and lower beetle abundance and species richness. Squirrels also reduced more plant cover in winter compared to summer, but had no effect on small mammal species richness. Furthermore, plant cover and height were higher in summer, whereas small mammal abundance and species richness were higher in winter. Our results suggest that Cape ground squirrels are important ecosystem engineers that influence plant and animal communities in the Namib Desert grasslands.     

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.     

Home-Range Dynamics in a Solitary Subterranean Rodent

Despite an important role of subterranean rodents as ecosystem engineers, their belowground mobility is poorly documented. It is supposed that their underground burrow systems, once established, are relatively stable because of high-energy costs of digging. We chose the silvery mole-rat, Heliophobius argenteocinereus (Bathyergidae, Rodentia) from mesic Afrotropics as a representative of solitary subterranean rodents to investigate how, and how fast these rodents process their established burrow systems. We combined radio-tracking of individual animals with subsequent mapping of their burrow systems, and we developed a new method for assessing the rate of burrowing. Mole-rats continuously rebuilt their burrow systems; they excavated approx. 0.7 m of new tunnels per day and backfilled on average 64% of all tunnels. On average, every 32 d they established a new nest. They often completely backfilled newly excavated peripheral burrows, while other parts of their burrow systems were more permanent. Their home-ranges were dynamic and continuously shifted in space. Burrow system processing continued even in the advanced dry season, when soil is difficult to work.     

Paternal effort and its relation to mating success in the European ground squirrel

The necessity for parental care is a driving force for determining mating systems and social organization. The European ground squirrel, Spermophilus citellus, is a polygynous, gregarious species in which male parental behaviour would not be expected. We had observed males digging in litter burrows that were later occupied by females and their offspring. Males never stayed overnight within these burrows. To determine whether this was some kind of paternal effort we tested the following hypotheses: (1) that male burrowing behaviour was directed towards the male's own offspring or towards the pregnant or lactating mother of the male's offspring; (2) that this behaviour had costs in terms of condition, decreased survival or fecundity; and (3) that it benefited offspring condition or survival. All three assumptions were met. Males worked on the litter burrow of their copulatory partners. Thus, this behaviour was directed towards the male's potential offspring. Male burrowing costs were seen in decreased foraging time and increased body mass loss. Offspring benefits were evident in increased mass at natal emergence. We conclude that male digging at litter burrows can be considered as paternal effort. Lastly, we considered the effects of polygyny on this male parental effort by comparing mating effort, mating success and paternal effort. High mating success was associated with high mating effort and low paternal effort. Moderate to low mating success was associated with lower mating effort and higher paternal effort, indicating a trade-off between the two.     

One House Two Families: Petrel Squatters Get a Sniff of Low‐Cost Breeding Opportunities

Burrowing is a widespread nesting behaviour, found in vertebrates and invertebrates. It is particularly common in small procellariiform seabirds such as blue petrels (Halobaena caerulea) and Antarctic prions (Pachyptila desolata), two closely related petrel species. However, digging a burrow is costly and alternative strategies may evolve. Accordingly, blue petrel males can adopt two alternative nesting strategies: digging a new burrow or squatting in an empty one. Importantly, a blue petrel squatter arriving at the colony to breed is more likely to find empty Antarctic prion burrows than empty blue petrel burrows, since the former species only start breeding a month later. However, squatting in a prion’s burrow is risky for blue petrels as the legitimate owner very often returns and claims the burrow back, thus ruining the squatter’s breeding attempt. We present here results of a survey of two sympatric colonies of blue petrels and Antarctic prions on Kerguelen Island. Our data show that blue petrel squatters preferentially occupy blue petrel empty burrows. To investigate potential underlying mechanisms behind this preference, we used a simple Y‐maze design to show that blue petrels can discriminate and prefer their specific odour over the prion odour. Our results confirm the existence of alternative burrowing strategies in blue petrels and suggest that squatters could use olfaction to avoid the less suitable Antarctic prion burrows.     

Habitat selection by the Spiny-tailed lizard (Uromastyx aegyptia): A view from spatial analysis

Many factors affect the habitat selection for animal species, which in turn may greatly affect their distribution in different ecosystems. Understanding the processes that affect habitat selection is also critical for guiding and managing conservation initiatives. Our study aimed to assess the habitat selection by free-ranging Spiny-tailed lizard (Uromastyx aegyptia) by analyzing a geospatial data connecting its burrow parameters to different habitat characteristics within selected sites in Hail region, Saudi Arabia. We examined evidence and patterns of significant spatial clustering for (366) active burrows by linking their parameters (burrow entrance size, burrow entrance width and burrow entrance height), their reference geographical locations and, two habitat characteristics defined by soil type and vegetation cover. The objective of the analysis was to increase the understanding on the burrows aggregation process in the space and, to describe its possible relation to other spatial habitat configurations. Analysis of distances based on the Nearest Neighbor Index (NNI) and hotspots detection in Nearest neighbor hierarchical clustering (Nnh) suggested twelve (12) spatial clusters located within the study area. In addition, a spatial ordinary least square (OLS) and Poisson regression models revealed significant effects of soil type and vegetation cover on burrow parameters (OLS, p < 0.05; Poisson, p < 0.001), which indicate a strong association between burrows parameters and habitats characteristics. Findings from the study also suggest that other factors such as elevations, highways, and human settlement concentration spots could possibly play a major role in defining burrow spatial aggregation and furthermore have a significant impact on habitat selection.     

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 control of nest depth in a digger wasp (Sphex ichneumoneus L.)

Golden digger wasps, Sphex ichneumoneus L. (Sphecidae), are a solitary, ground-nesting species that dig burrows to particular depths in the soil. I develop and evaluate alternative hypeses about the mechanisms controlling digging behaviour. By altering the wasps' burrows as thare digging, I show that they are not digging for some prescribed length of time nor are they diggingtil they reach some suitable environmental characteristic deep in the soil. Rather, they appear be digging until they reach a particular tunnel length, making corrections if the tunnel is too shallor too deep. This distance can be altered somewhat by surface environmental conditions.     

Evolution of African mole-rat sociality: burrow architecture, rainfall and foraging in colonies of the cooperatively breeding Fukomys mechowii

African mole rats (Bathyergidae) offer an excellent system with which to test theories relating to the evolution and maintenance of sociality in mammals. The aridity food distribution hypothesis (AFDH) suggests that, within the bathyergids, sociality has evolved in response to patterns of rainfall, its effects on food distribution, and the subsequent costs and risks of foraging and dispersal. Here, in the first detailed study of burrow architecture in a social mole-rat species, with data from 32 burrows, we show that in the giant mole-rat Fukomys mechowii burrow fractal dimension increases with colony size and is higher during the rainy season than during the dry season. The mass of food in the burrow increases with fractal dimension and is higher during the rainy season than during the dry season. These results link for the first time colony size, burrow architecture, rainfall and foraging success and provide support for two assumptions of the AFDH, namely that (1) in arid conditions burrowing may be severely constrained by the high costs of digging; and (2) the potential risks of failing to locate food may be mitigated by increases in colony size.     

Environmental factors affecting the spawning burrow selection by the gobiid Valenciennea longipinnis

Spawning burrow selection by the longfinned goby Valenciennea longipinnis was studied in the near-shore moat on coral reefs, Okinawa, Japan. The gobies make several burrows within their home range, and spawn in one of them. To examine the factors important for spawning burrow selection three characteristics were investigated: current strength, burrow length and effect of underground water on the burrow. Among the burrows, pairs tended to spawn in a larger burrow irrespective of their body sizes. Most of the other non-spawning burrows were too small for a pair to stay together, because hard substrata may prevent the fish from excavating and shaping the burrow as they like. Moreover, pairs preferred to spawn in burrows where the underground water was oozing out, probably because the male's parental burden will decrease due to the higher dissolved oxygen concentration in such burrows. Although current strength may affect a water-exchange in a V. longipinnis burrow in relation to water-exchange function of a mound, it did not affect the spawning burrow selection because of the smaller velocity difference among the burrows relative to the daily fluctuation of tidal current.     

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.     

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.     

An assessment of the breeding range,colony sizes and population of the Westland petrel (Procellaria westlandica)

Abstract

The Westland petrel (Procellaria westlandica) is an endemic New Zealand species and one of the very few burrowing seabird species still breeding on mainland New Zealand. It nests only on a series of coastal ridgelines near to Punakaiki on the West Coast of the South Island. Between 2002 and 2005, surveys were undertaken at 28 of the 29 known colonies. The area occupied by the colonies was 73 ha; most colonies had fewer than 50 burrows, but six colonies had 201–500 burrows and four colonies had more than 1000 burrows. We find that the current breeding range of Westland petrel and the location of individual colonies are similar to those reported in both the 1950s and 1970s. Based on total burrow counts at 28 colonies and burrow occupancy rates determined by annual monitoring, the annual breeding population is estimated to be between 2954 and 5137 breeding pairs.     

Grouping increases the ability of the social rodent,Octodon degus,to detect predators when using exposed microhabitats

We examined the hypothesis that a main benefit of group‐living in the hystricognath rodent, Octodon degus (common degu), is to decrease individual risk of predation. During a first series of field observations, we contrasted group size of degus when using covered microhabitats with that of degus using exposed patches. During a second set of field observations, we assessed how distance to detection and to escape by degus varied with group size upon the approach of a potential human predator. Degus in exposed patches formed larger groups than degus in covered microhabitats. After excluding the influence of nearest burrow to focal subjects, we found that degus of larger groups detected an approaching human predator at a greater distance than degus of smaller groups. Likewise, degus of larger groups escaped to nearby burrows at a greater distance from the approaching predator than degus of smaller groups. All these pieces of evidence support the predatory risk hypothesis according to which group‐living in degus functions to reduce the risk of predation.     

Burrow limitations and group living in the communally rearing rodent, Octodon degus

Group living is thought to evolve whenever individuals attain a net fitness advantage due to reduced predation risk or enhanced foraging efficiency, but also when individuals are forced to remain in groups, which often occurs during high-density conditions due to limitations of critical resources for independent breeding. The influence of ecological limitations on sociality has been studied little in species in which reproduction is more evenly shared among group members. Previous studies in the caviomorph rodent Octodon degus (a New World hystricognath) revealed no evidence that group living confers an advantage and suggest that burrow limitations influence formation of social groups. Our objective was to examine the relevance of ecological limitations on sociality in these rodents. Our 4-year study revealed no association between degu density and use of burrow systems. The frequency with which burrow systems were used by degus was not related to the quality of these structures; only in 1 of the 4 years did the frequency of burrow use decrease with decreasing abundance of food. Neither the number of females per group nor total group size (related measures of degu sociality) changed with yearly density of degus. Although the number of males within social groups was lower in 2008, this variation was not related clearly to varying density. The percentage of females in social groups that bred was close to 99% and did not change across years of varying density. Our results suggest that sociality in degus is not the consequence of burrow limitations during breeding. Whether habitat limitations contribute to variation in vertebrate social systems is discussed.     

Notes on the relationship between the burrowing capacity, size and shoulder anatomy of some eastern Asiatic moles

Talpa micrura leucura and T. m. malayana make superficial burrows through tropical forest floor litter. They do not dig deeply into the soil or produce mole heaps as does Talpa europaea. Although some comparatively minor points of difference between the shoulder girdles of the eastern and European species are of sufficient interest to be described, it is suggested that it is the greater size and strength of Talpa europaea that enables it to perform the heavy manual labour of deep digging which T. m. leucura and T. m. malayana are not powerful enough to undertake.