Independence of Evoked Vocal Responses from Stimulus Direction in Burrowing Frogs Eupsophus (Leptodactylidae)

Localization of a sound source is important for animals in mating contexts: females generally orient towards signalling males, and males can estimate the position and quality of potential rivals. In anurans, the effect of sound direction on evoked vocal responses has been studied in males of Rana catesbeiana, which alter their vocal responses depending on the location of the stimulus. The current study explored the effects of sound direction in Eupsophus calcaratus, a frog that calls from inside burrows having resonance that would hinder the localization of incoming sounds. The vocal responses of 11 males to synthetic imitations of the conspecific advertisement call broadcast from loudspeakers positioned in front, to the right and left from the burrow openings were similar in terms of call rate, duration and latency. The invariance of the vocal responses indicates that for burrowing male frogs engaged in chorusing behaviour, the specific location of an opponent does not alter the persistence of vocal activity.     

Western Burrowing Owls (Athene cunicularia hypugaea) Eavesdrop on Alarm Calls of Black‐Tailed Prairie Dogs (Cynomys ludovicianus)

Animals sharing a common habitat can indirectly receive information about their environment by observing information exchanges between other animals, a process known as eavesdropping. Animals that use an auditory alarm calling system are an important indirect information source for eavesdropping individuals in their environments. We investigated whether Western burrowing owls (Athene cunicularia hypugaea) nesting on black‐tailed prairie dog (Cynomys ludovicianus) colonies responded to broadcasts of prairie dog alarm calls. Western burrowing owls are closely associated with black‐tailed prairie dogs in Colorado and neighboring states on the Great Plains of the United States. Prairie dog burrows in active colonies can serve as nesting sites for Western burrowing owls, and prairie dogs may act as an alternative prey source for predators, potentially decreasing the burrowing owls' risk of predation through the dilution effect. Burrowing owls nesting on prairie dog colonies may also eavesdrop on prairie dog alarm calls, enhancing their survival and nesting success on prairie dog colonies. We performed broadcast experiments with three different sounds: a prairie dog alarm call, a biological control (cattle mooing), and a non‐biological control (an airplane engine), and characterized burrowing owl responses as either alert or relaxed. For each sound stimulus, we recorded the time to first alert response to broadcast sounds (latency) and also how frequently the target burrowing owl exhibited an alert response within the first ten seconds of the broadcast (intensity). Burrowing owls reacted more quickly to the prairie dog alarm than to the biological control. They significantly increased the intensity of alert behaviors in response to broadcasts of the alarm, but did not show an increased reaction to either the biological or the non‐biological control. Our results suggest that burrowing owls nesting on prairie dog colonies eavesdrop on, and increase their alert behaviors in response to, prairie dog alarm calls.     

Strategies of burrowing in soft muddy sediments by diverse polychaetes

Muddy sediments are elastic solids through which morphologically diverse animals extend burrows by fracture. Muddy sediments inhabited by burrowing infauna vary considerably in mechanical properties, however, and at high enough porosities, muds can be fluidized. In this study, we examined burrowing behaviors and mechanisms of burrow extension for three morphologically diverse polychaete species inhabiting soft muddy sediments. Worms burrowed in gelatin, a transparent analog for muddy sediments, and in natural sediments in a novel viewing box enabling visualization of behaviors and sediment responses. Individuals of Scalibregma inflatum and Sternaspis scutata can extend burrows by fracture, but both also extended burrows by plastic deformation and by combinations of fracture and plastic deformation. Mechanical responses of sediments corresponded to different burrowing behaviors in Scalibregma; direct peristalsis was used to extend burrows by fracture or a combination of plastic deformation and fracture, whereas a retrograde expansive peristaltic wave extended burrows by plastic deformation. Burrowing speeds differed between behaviors and sediment mechanical responses, with slower burrowing associated with plastic deformation. Sternaspis exhibited less variability in behavior and burrowing speed but did extend burrows by different mechanisms consistent with observations of Scalibregma. Individuals of Ophelina acuminata did not extend burrows by fracture; rather individuals plastically deformed sediments similarly to individuals of the related Armandia brevis. Our results extend the range of natural sediments in which burrowing by fracture has been observed, but the dependence of burrow extension mechanism on species, burrowing behavior, and burrowing speed highlights the need for better understanding of mechanical responses of sediments to burrowers.     

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

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

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

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

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

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

In situ measurement of calling metabolic rate in an Australian mole cricket, Gryllotalpa monanka

Examination of the energetics of sound production usually requires measurement of species that will produce normal calls under unnatural circumstances. Such measurements are potentially compromised by stress-related changes in calling input (through a reduction in calling effort) or output (through forced use of sub-optimal singing burrows). To determine if such measurements are indeed affected by abstraction from a natural setting, we measured the energetics of song production in undisturbed mole crickets Gryllotalpa monanka and employed a new approach where the animal's singing chamber replaces the respirometry chamber normally used in studies of this type. It was therefore possible to measure metabolic rate (MR) of calling crickets in situ for animals within self-constructed burrows under natural conditions. Calling MR measured under these conditions averaged 13.5-fold higher than standard MR and 2.2-fold higher than MR measured during burrowing in the lab. The calling MR of G. monanka was similar to that measured for other calling insects, and to endothermic insects, but was only 10% of that allometrically predicted for a similarly sized insect (0.89 g) during flight. A male mole cricket is estimated to consume 5.9 ml of oxygen during construction of a calling burrow and a 1-h calling bout; by comparison, a flying female would consume a similar volume in less than 6 min.     

In situ measurement of calling metabolic rate in an Australian mole cricket, Gryllotalpa monanka.

Examination of the energetics of sound production usually requires measurement of species that will produce normal calls under unnatural circumstances. Such measurements are potentially compromised by stress-related changes in calling input (through a reduction in calling effort) or output (through forced use of sub-optimal singing burrows). To determine if such measurements are indeed affected by abstraction from a natural setting, we measured the energetics of song production in undisturbed mole crickets Gryllotalpa monanka and employed a new approach where the animal's singing chamber replaces the respirometry chamber normally used in studies of this type. It was therefore possible to measure metabolic rate (MR) of calling crickets in situ for animals within self-constructed burrows under natural conditions. Calling MR measured under these conditions averaged 13.5-fold higher than standard MR and 2.2-fold higher than MR measured during burrowing in the lab. The calling MR of G. monanka was similar to that measured for other calling insects, and to endothermic insects, but was only 10% of that allometrically predicted for a similarly sized insect (0.89 g) during flight. A male mole cricket is estimated to consume 5.9 ml of oxygen during construction of a calling burrow and a 1-h calling bout; by comparison, a flying female would consume a similar volume in less than 6 min.     

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.     

Regulation of vocal amplitude by the blue-throated hummingbird, Lampornis clemenciae

Animals that rely on vocal communication must broadcast sound so that a perceptible signal is transmitted over an appropriate distance. We found that male blue-throated hummingbirds modified the amplitude of their vocalizations in response to both naturally occurring and experimenter-controlled changes in ambient noise levels. This phenomenon is known as the Lombard effect and may increase the efficiency of acoustic signalling. This study demonstrates the effect under natural field conditions and documents the first hummingbird species (Apodiformes: Trochilidae) to show this behaviour. We measured sound pressure levels (SPLs) of Serial Chip territorial advertisement calls across a natural range of ambient noise, primarily due to creeks within male territories. We found a significant correlation between the amplitude of Serial Chips and the amplitude of background noise. To test this relationship, we broadcast recordings of creek noise at high and low amplitudes while target individuals were producing Serial Chip vocalizations. We measured vocal SPLs before and during the playback. Individuals responded to changes in playback creek noise by changing the amplitude of Serial Chip production. We also measured transmission properties of Serial Chip calls through natural habitat to calculate the approximate amplitude of vocalizations at the position of the calling bird. We suggest that amplitude regulation of vocalizations contibutes to signal transmission distance along with the established relationships between singing behaviour, acoustic structure and habitat. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.     

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

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

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

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

Influence of age and environmental factors on burrow-making behavior of the short-tailed cricket,Anurogryllus muticus (De Geer) (Orthoptera: Gryllidae)

For the short-tailed cricket, Anurogryllus muticus, burrow-making behavior is essential. All nymphal instars construct burrows, but in the adult stage the rate of burrowing behavior is age dependent. Increases in photophase and light intensity stimulate burrowing, and the explicit negative phototaxis is correlated with the cricket's inability to exist under dry conditions. Ingestion of substrate during burrow construction may serve to acquire additional moisture. There is no evidence of burrow recognition, and crickets can construct a burrow when needed. The natural distribution of burrows at the plot investigated on Moorea supports the notion thatA. muticus builds burrows where the preferred food plantAlysicarpus vaginalis is most abundant. By minimizing the traveling distance to food sources when foraging they can retreat to their burrow again.     

Acoustic signals of a solitary subterranean rodent Ctenomys talarum (Rodentia: Ctenomyidae): physical characteristics and behavioural correlates

The purpose of this study was to analyse and describe vocalizations of a solitary subterranean rodent, Ctenomys talarum. In adult C. talarum five different sounds (four true vocalizations and one mechanical sound) were recorded during different behavioural contexts. Using data from the laboratory and literature, we classified these vocalizations as territorial, distress, and mating calls. We found that the vocalization range in C. talarum is shifted towards low frequencies, which transmit better in underground burrows and match well the hearing range described for other species of subterranean rodents. These low-frequency vocalizations, also found in other non-related subterranean rodents, may reflect an adaptation to the acoustic conditions of the habitat. Electronic Publication     

Female frog auditory sensitivity,male calls,and background noise: potential influences on the evolution of a peculiar matched filter

The matched filter hypothesis proposes that the tuning of females' auditory sensitivity matches the spectral energy distribution of males' signals. Such correspondence is expected to arise over evolutionary time, as it promotes conspecific information transfer and reduces interference from other sound sources. Our main objective was to determine the correspondence between the acoustic sensitivity of female frogs of Eupsophus roseus and the spectral characteristics of advertisement vocalizations produced by conspecific males. We also aimed to determine how auditory sensitivity is related to the characteristics of background noise. We analysed data on the auditory sensitivity of E. roseus females, and recordings of conspecific male vocalizations and of the acoustic environment during the breeding period of this species. Our results indicate a concordance between the auditory sensitivity of females and call spectra that would provide an appropriate detection of these signals. In addition, this matching is large relative to the correspondence between auditory sensitivity with the spectra of the abiotic and biotic background noise, with the last component being associated with calls of the related species Eupsophus vertebralis. This may be an adaptation of receivers confronting sound interference, which improves the capability of E. roseus to communicate sexually by means of acoustic signals. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 814–827.     

Hidden burrow plugs and their function in the tiger beetle, Cosmodela batesi (Coleoptera, Cicindelidae)

Tiger beetle larvae excavate and live in underground burrows, whose openings they sometimes plug with soil. This study documents the burrow plugging behavior of the tiger beetle, Cosmodela batesi (Fleutiaux), in the field. We also tested the function of burrow plugs in the laboratory. In the field, C. batesi more frequently made a plug when it rained. Most larvae made plugs inside their burrows (rather than at the soil surface), and the use of an endoscope was necessary to detect these sub-surface plugs. In the laboratory, flooding was simulated by artificially introducing water into specially-made arenas. Water filled the entire burrow when there was no plug, whereas plugged burrows maintained air chambers inside. When a plug was broken with a wire, burrows filled up with water. The burrowing and plugging behavior described in this study is likely an important adaptation of C. batesi to its habitat.     

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.     

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

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

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.     

When seasnake meets seabird: Ecosystem engineering,facilitation and competition

Ecosystem engineers such as burrowing seabirds can increase habitat availability for sympatric taxa – but only if the burrow's owner allows other species to use the newly created shelter site. Our studies on a small Pacific island suggest that an avian burrower (the wedge‐tailed shearwater Puffinus pacificus) is both a facilitator and a competitor for amphibious seasnakes. Video camera inspection of 102 burrows revealed frequent usage of these burrows as retreat sites by the snakes, with Laticauda laticaudata restricted to burrows <4 m from the water's edge, whereas Laticauda saintgironsi often used burrows further inland. Snakes never occupied burrows that contained adult shearwaters, suggesting active burrow defence by the birds. Model snakes that we inserted into burrows were attacked, especially on the head and upper body, and we found one snake pecked to death outside a burrow. Wedge‐tailed shearwaters act as facilitators, creating a thermally favourable microhabitat and substantially enhancing habitat suitability for snakes; but they are also competitors, aggressively competing with snakes for occupancy of the resource that has been created.     

Gecko phonotaxis to cricket calling song: A case of satellite predation

Insectivorous Mediterranean house geckos, Hemidactylus tursicus, are found in the field close to the burrows of calling male decorated crickets, Gryllodes supplicans. In playback experiments where adult geckos were presented with calls of these crickets or calls of frogs as controls, geckos exhibited positive phonotaxis to broadcast cricket calling song. This indicates that geckos orient to and approach male cricket calls, even though the crickets call from burrows where they are protected from the geckos. However, this behaviour enables the geckos to intercept and consume female crickets that also respond phonotactically to the cricket calls. Thus geckos act as ‘satellite predators’, a situation which may impose sex-biased mortality on female crickets.     

Behavior of the burrowing shrimp Alpheus macellarius in varying gravel substrate conditions

The behavior of the male snapping shrimp, Alpheus macellarius (Decapoda: Alpheidae), was studied in tank experiments with four levels of gravel content for five consecutive days. Markov-chain type analyses identified significant behavioral states that were grouped into six behavioral classes (Wandering, Surveying, Grooming and Foraging, Resting, Hidden). Significant trends in the durations and frequencies of the classes were found across days and periods, with shrimps mainly wandering and burrowing during the first day. Grooming, foraging and resting became more frequent during succeeding days. Shrimps built burrows in the morning and then groomed and foraged mostly in the afternoon. This periodicity implies a greater priority for the shrimp to construct and maintain a burrow rather than forage. Gravel content did not significantly influence burrowing behavior, but marked variations were noted in burrowing success, burrow structure and stability. Animals in 15% and 25% gravel substrates produced more burrows of greater complexity, which lasted longer than those of the shrimps in 0% and 5% gravel setups. Feeding was mainly sediment scavenging suggesting the predominance of a deposit-feeding trophic mode. Modifications in burrow construction and adaptations in burrowing and feeding indicate the ability of A. macellarius to respond to different environmental conditions.     

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

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