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.     

Factors Affecting Mating Tactics in the Fiddler Crab, Uca vocans hesperiae

The mating strategies of male fiddler crabs are variable and highly flexible within species. In this study I examine three types of mating strategy used by individual male Uca vocans hesperaie. The most common strategy, termed a ‘standard gambit’, where males approached females at their burrow entrance and initiated courtship, accounted for 63% of mating attempts and 75% of successful matings. The rarest strategy (4% of mating attempts) was the ‘dig out’, where males attempted to mate with females whose burrows they had excavated. This strategy accounted for 19% of successful matings. ‘Herding’ behaviour which involved a male attempting to herd a female into a burrow and mate, contributed 33% of mating attempts but were generally unsuccessful, accounting for only 2.6% of successful matings. Males used more than one strategy during the study period. Smaller males used the standard gambit strategy more often than herding or dig outs while larger males used the herding strategy more often. There was no relationship between male size and mating success and males did not preferentially mate with females of a certain size. The predominant strategy adopted by males over the lunar cycle depended on female behaviour. Herding behaviour was induced by female wandering which escalated at full moon. Standard gambits were the commonest strategy adopted at and around new moon. The low success rate of male mating attempts (16%) indicates a reluctance by females to mate multiply. This may lead to conflict between the sexes because in fiddler crabs there is last male sperm precedence.     

Sound transmission and burrow characteristics of the subterranean rodentCtenomys talarum (Rodentia: Ctenomyidae)

Males of tuco-tucoCtenomys talarum Thomas, 1898 use particular burrow’s entrances to emit their territorial vocalization. Therefore we studied the internal structure of these entrances and the possible effect on the emission and propagation of airborne sounds. Externally, the burrow entrances used by tuco-tucos males to vocalize were characterized by the absence of sand mounds around their openings. Internally, most of the burrow’s entrances consisted of a main, relatively straight, tunnel of 30–40 cm length, with a diameter of 5.7–6.4 cm. After passing through the burrow’s entrance, the low-frequency components of an artificial signal played back inside the tunnel were not only less attenuated but also amplified (measured at 10–30 cm from the burrow opening). Therefore, the emission of territorial vocalizations inside the particular burrow’s entrances may be considered as a complex adaptative behavior, in which burrow structure improves the signal emission and propagation. Moreover, this work also showed thatC. talarum’s territorial vocalization seems to be adequate for long, inter-burrow communication, since its physical characteristics (high amplitude and low main frequency) are concomitant with the frequencies that are better transmitted in the natural habitat of this species of subterranean rodent.     

Hunting Behavior in the Ctenizidae

The most primitive way of hunting in Mygalomorph spiders seemsto be the free roaming and catching of encountered prey Theraphosidae.The trap-door spiders Ctenizidae, Actinopodidae and Barychelidawhich are entirely sedentary, lie in wait behind the trap-doorand leap at prey that happens to pass close to the door. Somespecies spin radial silk threads outside the door which functionas stumbleor signal-lines, and some Australian species use grassblades and other litter in the same way. A further evolutionarystep leads to species which do not build a trap-door but crowntheir burrow by a funnel-shaped web. The Dipluridae finallyare real web-builders, which depend on a sheet-web to catchtheir prey. Generally they do not dig a burrow but hide in asmall retreat from which a funnel-web leads to the net. With a few exceptions Ctenizidae are entirely nocturnal. Theirrhythm of activity has been analyzed. The "Zeitgeber" is thedaylight during the last half hour before sunset. Most trapdoorspiders never leave their burrow during their whole life. Theyneed three to four years from hatching to become adults. Adultmales die during or at the end of the restricted mating-season;they take no food when adult. Females, which undergo post-adultmolts, can probably live for 15–20 years. Nemesia caementaria, like most other species, hunts during thewhole night. The mean time of activity is about Si/o hours,consisting of periods of lying in wait and periods of intermediaterests. The spiders make an average of three leaps per nightto catch a prey, but only about 10% of all bounds are successful.A hungry animal, lurking in vain, shows unmotivated leaps. The effects of light, moisture, and temperature on hunting activityare analyzed. Ctenizidae hunt during autumn, winter, and springbut interrupt their activity in summer for an estivation whichlasts usually two months. The females of some species capture the male after mating andeat him; others never attack him. This difference in behaviorlias repercussions on reproduction. Among certain species the young nymphs of the third instar refuseall food until they have made their own burrow and can huntby themselves. The young of other species stay with the motherfor one year and leave her at the fifth or sixth instar to maketheir own burrow. The young of Nemesia caementaria can remainin the burrow of the mother until they are almost adult, i.e.,for two or three years, and feed on the prey the mother hascaught. The behavior of Ctenizidae can be grouped. Some species lurkbehind the closed or almost closed trap-door. Another groupopens the trap-door and puts the pedipalpi and the two anteriorpairs of legs radially out onto the rim of the burrow, whilethe cephalothorax is hidden behind the trap-door. A third groupspins threads of silk radially about the entrance and uses themas signal-lines. A fourth group can come out and pursue theprey, then drags it to the trap-door and into the burrow. The nature of the prey depends on the bio tope. The Ctenizidaefeed almost exclusively on insects, mainly ants and beetles;in the laboratory crickets are accepted as prey. Ctenizidae have no tarsal organ. They have different types oftrichobothria, transversal and longitudinal slit-organs, andlyriform organs. With these three kinds of sense-organs, thefunctions of which are not clearly understood, the trap-doorspiders are able to perceive the approach of prey, to judgeits distance from the trap-door, and to locate it in direction.They seem to have an organ of smell, since certain groups ofinsects are repulsive to them. Sight is not used for hunting.     

Snake-directed Behavior by Snake Naive and Experienced California Ground Squirrels in a Simulated Burrow

Snake naive and experienced California ground squirrels (Spermophilus beecheyi) were video taped while interacting with either a gopher snake or rattlesnake in a simulated burrow dimly illuminated with red light. Using nonvisually guided behavior, naive and experienced squirrels reacted to snakes in qualitatively similar ways, and behaved more defensively toward snakes than toward a control stimulus (white rat). The squirrels alternately interacted with the snake and attempted to escape from the burrow, which had a sealed entrance. Interaction with the snake included cautious approach in elongate postures, prolonged investigation of adjacent alleys before entering them, kicking sand at the snake, frequent tooth chattering, occasional calling, and building burrow plugs out of sand. These they packed by butting with their heads. When permitted to escape from the burrow, they turned just outside the entrance to tail flag, kick sand, scent mark, and finally plug the burrow. Since visual cues were not available, olfactory and auditory stimuli from the snake appeared to mediate snake-directed behavior in the burrow.     

Courtship tactics by male Ilyoplax pusilla (Brachyura, Dotillidae)

Mating in the dotillid crab Ilyoplax pusilla occurs after the female enters the male’s burrow in the tidal flat. Males use two tactics to cause females to enter their burrows for mating: the male either directs claw waving to the female (courting-wave display), to which the females responds by following the male to his burrow, or the male runs rapidly away from, then back toward, his burrow (dash-out-back display), which startles the female into his burrow. Males more often used the courting-wave than the dash-out-back display, but mating success did not differ between the two tactics. Male use of either tactic was influenced by date, female density and male size; the courting-wave display was used by larger males, later in the breeding period, and under higher female density.     

Extrinsic and Intrinsic Factors Influencing the Emerging and Burrowing Behaviors of Reproductively Active Adults of the White Grub Beetle, Dasylepida ishigakiensis

The subtropical scarab beetle, Dasylepida ishigakiensis, has a two-year life cycle. This study showed the time of adult emergence from the soil relative to the time of dusk and the presence of the female sex pheromone. Beetles collected on Miyako Island were transported to Tsukuba where they were immediately placed under natural day lengths in February. They exhibited two emergence peaks that corresponded to the times of dusk in Tsukuba and on the island, respectively. Males emerged precociously if a lure containing synthetic female sex pheromone was placed in their container, whereas the females’ behavior was unaffected. Previous observations that mated females dig deeper in the soil than virgin females, males or mated males were confirmed. To explore the underlying mechanism controlling the behavioral change associated with mating, liquid material derived from the male accessory glands, seminal vesicles and female bursa copulatrix was injected into beetles, but without any significant influence on burrowing behavior. No significant influence was also observed in beetles injected with anisomycin, an inhibitor of protein synthesis related to memory in other animals.     

Burrow fractal dimension and foraging success in subterranean rodents: a simulation

For animals that forage underground, the success with whichfood items are located may be closely related to burrow architecture.Fractal dimension, which describes how a burrow explores thesurrounding area in a way that is independent of burrow length,is an obvious choice for a single metric describing burrow shape.Although it is often assumed that burrows of high fractal dimensionwill be associated with greater foraging success, this has notpreviously been demonstrated. In this study, we use computersimulations to study the success with which burrows of differentfractal dimensions locate randomly distributed food items. Inaddition, we examine the effect of different patterns of fooddistribution (in particular the patchiness with which food itemsare distributed) and consider how using different criteria forlocating food items affects the relationship between fractaldimension and foraging success. We conclude that, under a widerange of plausible assumptions about the ways in which subterraneanrodents forage, burrows of high fractal dimension are more successfulat locating food items than burrows of lower fractal dimension.     

The dispersal behaviour of the phoretic mite Poecilochirus carabi (Mesostigmata,Parasitidae): adaptation to the breeding biology of its carrier Necrophorus vespilloides (Coleoptera,Silphidae)

Summary When the phoretic mite Poecilochirus carabi reproduces in the brood chamber of its carrier Necrophorus vespilloides, a beetle with biparental brood care, the first deuteronymphs of the new mite generation aggregate on the male beetle. They do not use sex-specific traits to discriminate between male and female beetles in the brood chamber, but traits that are related to the beetles' behaviour and may be displayed by both parent beetles. When the male beetle departs, it carries virtually all deuteronymphs then present in the brood chamber. Deuteronymphs that develop later congregate on the female, which leaves the crypt some days after the male. Only those deuteronymphs that miss the female's departure disperse on the beetle larvae, meaning they have to wait in their pupal chambers until the beetles have completed their development. On average, 86% of the deuteronymphs leave the brood chamber on the parent beetles, thereby gaining the advantage of an early departure. As soon as their carrier arrives at one of the beetles' meeting places, the deuteronymphs can transfer between the beetles present. Choice experiments revealed that the deuteronymphs tend to even out density differences between congregating carriers, and prefer sexually mature to immature beetles. Therefore, transferring between beetles results in a dispersion of deuteronymphs on the sexually mature beetles of the population.     

Choosing a mate in a high predation environment: Female preference in the fiddler crab Uca terpsichores

The interplay between a receiver's sensory system and a sender's courtship signals is fundamental to the operation of sexual selection. Male courtship signals that match a female receiver's preexisting perceptual biases can be favored yet the message they communicate is not always clear. Do they simply beacon the male's location or also indicate his quality? We explored this question in a species of fiddler crab Uca terpsichores that courts under elevated predation risk and that mates and breeds underground in the safety of males' burrows. Sexually receptive females leave their own burrows and are thereby exposed to avian predators as they sequentially approach several courting males before they choose one. Males court by waving their single greatly enlarge claw and sometimes by building a sand hood next to their burrow entrance. Hoods are attractive because they elicit a risk‐reducing orientation behavior in females, and it has been suggested that claw waving may also serve primarily to orient the female to the male. If the wave communicates male quality, then females should discriminate mates on the basis of variation in elements of the wave, as has been shown for other fiddler crabs. Alternatively, variation in elements of the claw waving display may have little effect on the display's utility as a beacon of the location of the male and his burrow. We filmed courting males and females under natural conditions as females responded to claw waving and chose mates. Analysis of the fine‐scale courtship elements between the males that females rejected and those they chose revealed no differences. When predation risk during courtship is high, males' courtship displays may serve primarily to guide females to safe mating and breeding sites and not as indicators of male quality apart from their roles as beacons.     

Feral Brown rats, Rattus norvegicus, in South Georgia (South Atlantic Ocean)

Brown rats were introduced to the sub-antarctic island of South Georgia probably around 1800. They are now widespread and abundant, particularly on the north coast. The population is divided into discrete units by the rugged topography of the island, particularly the many glaciers. Rats are found mostly in dense stands of coastal tussock grass which provides both shelter and food. They dig burrows in the tussock stools and make nest chambers in the leaf canopy. Tussock, which is rich in carbohydrate, forms the main part of their diet throughout the year. Perimylopid beetles are regularly eaten, and the rats forage on the sea-shore. Carrion is eaten where available and the rats prey on ground-nesting birds. Breeding is probably seasonal, as litters were found only from December to February. The rats have adapted successfully to the rigorous South Georgia climate, but are dependent on tussock grass for their survival. They have made rather little impact on the vegetation. Dove prions, diving petrels and some large petrels are preyed on but breeding colonies of these birds can coexist with rats. The Antarctic pipit rarely if ever nests in rat-infested areas. No management procedures would be possible.     

Location before emergence of the female bee, Centris pallida, by its male (Hymenoptera: Anthophoridae)

Large numbers of males of the bee Centris pallida Fox have been observed patrolling areas in which females are emerging. Males locate specific sites at which a buried bee is about to emerge and dig down to meet the other individual, male or female. If it is a female, mating is initiated when she scrambles into the excavation pit created by the male. Males fight intensely with one another for possession of digging sites and for unburied virgin females. Experiments indicate that males locate conspecifics beneath the surface on the basis of extremely non-specific olfactory cues; they are capable of locating buried honey bees and other insects. The evolution of digging behaviour is traced to selection favouring males that are first to reach a virgin female which will mate just once in her lifetime. A number of examples are given of other insects that have evolved similar abilities, apparently in response to similar selection pressures.     

Aggregation in a Desert Tenebrionid Beetle: A Cost/Benefit Analysis

Parastizopus armaticeps (Coleoptera: Tenebrionidae), a nocturnal fossorial detritivore inhabiting southern Kalahari dunes, aggregates in burrows during the day. Group size increases during drought but 25% of beetles are still found alone or in pairs. During drought, beetles from large groups leave burrows after sunset synchronously and carlier than pairs and single animals and earlier than beetles of any group size after rain. Detritus from the beetles' major foodplant is scarce and food competition high. Beetles emerging early preferentially select and carry high-quality transportable items into burrows to eat (forage); late-emerging ones feed on the low-quality large twigs on the surface. Foraging is shown to be a strategy to secure food items against surface competitors, not one to reduce body water loss during surface exposure. The costs and benefits of group vs. solitary lifestyles and alternate hypotheses for early and synchronous emergence were tested experimentally. Grouped beetles had lower body water loss rates but, due to competition with burrow mates, higher feeding costs than single ones. It is hunger that advances and thus synchronizes emergence time, not social facilitation. Field data support a model predicting that, for maximal benefits, beetles should alternate between solitary and group life at optimal time intervals.     

Structure and function of the large pronotal horn of the sand‐living anthicid beetle Mecynotarsus tenuipes

Large insect horns function as antipredator armaments, digging implements and intraspecific combat weapons. The sand‐living anthicid beetle Mecynotarsus tenuipes possesses a large horn on the pronotum. Allometric relationships between body size and horn size did not show either a slope of more than 1 or sexual dimorphism, suggesting another function of the horn other than sexual selection via combat. Behavioral observation of individuals using a microvideo camera indicated that the horn is used to dig and move forward in loose sand. Only the horned M. tenuipes could dig into sand, in contrast to the hornless anthicid beetles Stricticollis valgipes and Clavicollis fugiens, which could not dig. When moving in sand, M. tenuipes joins its pronotal horn and head to form a conical shape, with which it pierces into the sand. Then, it opens its horn and head outward to create a space in the sand for forward motion. Although it can dig deeply into sand by repeating these behaviors sequentially, digging speed tends to slow with depth, probably because the weight of the substrate increases.     

Pre-ovipositional maternal care alleviates food stress of offspring in the flower beetle <Emphasis Type="Italic">Dicronocephalus wallichii</Emphasis>

Unlike most other flower beetles, females of Dicronocephalus wallichii exhibit nesting behaviour. The female constructs a burrow in the soil, cuts dead plant leaves into small pieces to provision the nest, and then lays one egg inside the nest. Hatched larvae have been thought to feed on the nest materials prepared by their mothers, but the effects of pre-ovipositional care on larval performance have not been tested. The hatched larvae were found to stay in the nest for 15–30 days until they consumed the nest materials. We examined whether the presence of provisioned nests enhanced larval performance under both benign and food-stress conditions. With high-nutrient soil, larval survival rate and growth speed were not affected by the presence of provisioned nests. By contrast, with low-nutrient soil, mortality of the larvae was much higher in the absence than in the presence of provisioned nests. The growth speed of larvae with nests located in low-nutrient soil was as high as those reared in high-nutrient soil. These results indicate that females alleviate the food stress of larvae during their initial developmental stage by constructing provisioned nests.     

Resource distribution and its effect on the mating system of a longhorned beetle,Perarthrus linsleyi (Coleoptera: Cerambycidae)

Summary Adults of Perarthrus linsleyi feed on flowers of creosote bush, Larrea tridentata. Mating also occurs on the flowers and foliage of this plant. This food resource is widely and evenly distributed in space, and is usually abundant. The spatial distribution of the beetles bore no relationship to the spatial distribution of flowers among creosote bushes, nor were female beetles distributed predictably with respect to flower distribution. Males moved widely over the area under study, and moved much more frequently than females. Males actively searched for females, and mounted and began copulation attempts without preliminary courtship. Males did not engage in aggressive defense of females, creosote flowers, or creosote bushes. This species exhibits a scramble competition mating system. The spatial distribution of the food resource is a primary factor in the evolution of the mating system of Perarthrus linsleyi.     

BODY WEIGHTS IN LAYSAN ALBATROSSES DIOMEDEA IMMUTABILIS

Male Laysan Albatrosses are heavier (3300 gm.) than females (3000 gm.) upon arrival on Midway. This difference between the sexes persists except at egg-laying and during certain periods of incubation. Weights of males and females are essentially the same at egg-laying, perhaps because of the presence of the 300 gm. egg in the female, but mostly because the males have lost weight in the pre-egg phase. Losses in weight during the long, continuous incubation spans are about 18–25% of the body weight. Males show a net loss of about 10% during the entire incubation period; there is no statistically significant change in the weights of the females. Weights of both sexes decline in parallel from the end of incubation to mid-May. Thereafter, they increase as attentiveness to the young is less and the parents have more time to forage at sea. Nevertheless, the body weights of both sexes at the end of the breeding season (July) are down 10–45%. Decreased body weights of breeding birds cannot be attributed solely to the rigours of parental care; former breeders and juveniles not yet of breeding age show a similar decline in weight. It is suggested that the normal pattern of yearly breeding by adult Laysan Albatrosses may be broken by failure to add weight (fat) before the breeding season. Former breeders consistently weigh 15–20% less than current breeders. Nestlings normally show a gradual increase in body weight from hatching in late January until mid-May, at which time they may be significantly heavier than either parent. As a result of the increased energy requirements of locomotion and wing exercise and decreased food supplied by parents, nestlings lose about 30% of the May weight before they finally go to sea in Jufy A possible “mutation” for small body weight is discussed.     

Sexual Dimorphism,Fighting Success and Mating Tactics of Male Onymacris plana Péringuey (Coleoptera: Tenebrionidae) in the Namib Desert

Sexually dimorphic characters of Onymacris plana, a dune-living, solitary tenebrionid beetle of the Namib Desert, were tested for their roles in male-male fighting over females. Males were smaller than females but had extraordinarily wide elytra, with great variance in this characteristic. In males, but not in females, elytra width increased with body length at an allometric scale. Male beetles were often aggressive towards each other, especially when mating or guarding females after mating or waiting for females at shady spots. Interactions were less intense when contesting over females on the open surface, where these fast-running beetles often overran each other in their attempts to retain their positions behind females until the females retreated into the sand, where mating took place. Winners of intrasexual fights and the successful mates of females tended to have longer bodies and wider elytra than the losers. Sexual selection appears to be the best explanation of the allometric scaling of the lateral extensions of male elytra. Sexual selection may furthermore contribute to other characteristics, such as large body length and long legs, that have ecological and ecophysiological significance.     

Sex-role reversal in the tidewater goby, Eucyclogobius newberryi

Species in which females compete more intensely than males for access to mates are uncommon. Sex-role reversal in fishes has been documented only in species in which males bear eggs, such as pipefish and a mouth brooding cardinalfish. I investigated the reproductive behavior of the tidewater goby, Eucyclogobius newberryi (Gobiidae), to determine whether and to what degree this species is sex-role reversed. Males constructed and defended burrows for spawning in sand. Both sexes initiated courtship, but the female's breeding coloration was more striking. The intensity of sexual aggression was greater among females than among males. The female laid her entire clutch with a single male, and the male accepted only one clutch per brooding cycle. Both sexes spawned repeatedly (up to 12 times in aquaria), but fish did not form permanent pairs. Males cared for eggs in the burrow 9–11 days until hatching, and rarely if ever emerged to feed. Many aspects of male behavior (nest construction and defense, courtship, and parental care) were typical of most gobiids. On the other hand, female behavior (black nuptial coloration and intense female-female competition) was unusual, not only for gobiids but for animals in general. I therefore concluded that the tidewater goby is moderately sex-role reversed. Its sexual behavior is apparently unique among fishes because it is the only reported case of sex-role reversal in teleost males that do not bear eggs or developing young. This revised version was published online in July 2006 with corrections to the Cover Date.     

The role of hood-building in defining territories and limiting combat in fiddler crabs

In Panama, courting males of three species of fiddler crabs sometimes construct semi-domes of mud (=hoods) over their burrow entrances. Males that do not construct hoods court in a circular area surrounding their burrow entrance. Males with hoods restrict their general activity and courtship to the semi-circular area in front of the hood opening. For Uca latimanus hoods appear to be an integral part of the courtship ritual and are constructed by all courting males. In the other two species, U. musica terpsichores and U. beebei, hood-building is associated with high population density and the presence of hoods reduces the frequency of combat among neighbouring males.