Bored and Burrowed: An Unusual Echinoid Steinkern from the Type Maastrichtian (Upper Cretaceous,Belgium)

A test of the irregular echinoid Hemipneustes striatoradiatus (Leske) was dissolved in dilute hydrochloric acid to expose the flint steinkern. This revealed slender, string-like borings that were mainly limited to ambulacra (Talpina isp. cf. T. canna (Price)); less common pouch-like borings limited to part of one ambulacrum only (Rogerella isp. cf. R. pattei (Saint Seine)); and slender, sinuous to contorted burrows, Planolites montanus Richter. Ingress to the test (borers) and sediment infill (burrowers) was presumably through the ambulacral pore pairs of the dead test. Borings and burrows were probably not coeval. Such infestations of fossil echinoids may be locally common, but their identification is controlled by the difficulty of seeing features of both the test and the infill.     

Cutting-edge technology: burrows lined with sponge bioclasts from the Upper Cretaceous of Denmark

Abstract

Many tracemakers use different materials to line their burrows. Koptichnus rasmussenae n. igen. n. isp. is lined with cuboid fragments of siliceous sponges, interpreted as evidence of harvesting and trimming material to reinforce the burrow wall. The act of trimming, as evidenced in the polyhedral faces, is considered to be behaviourally significant. The tracemaker was evidently a lobster-like crustacean.     

In deep water: a crinoid–brachiopod association in the Upper Oligocene of Antigua,West Indies

Extant brachiopods and stalked crinoids are found together in the deeper waters of the Caribbean Sea. Analogous brachiopod/crinoid associations have been reported from diverse palaeoenvironments in the Neogene of the region. Studied examples include the Pleistocene of Jamaica (deeper water fore reef), and the Miocene of Jamaica (island slope chalks), Barbados (accretionary prism) and Carriacou (turbiditic siliciclastic shelf). Comparison with analogous modern environments indicates deposition in 150+m water depth. This association has now been extended back into the Late Oligocene. Crinoids and brachiopods both occur in the Antigua Formation of Antigua; both occur high in the formation, implying deeper water in this retrograde succession. They have received little attention from systematists, although the brachiopods Cistellarcula dubia Cooper and Tichosina foresti Cooper have previously been described from the Antigua Formation; to these, we add Cistellarcula sp., Argyrotheca sp. and Tichosina sp. At Half Moon Bay in southeast Antigua, high in the Antigua Formation, we have found columnals of isocrinid crinoids (cf. Isocrinus sp.) associated with rare brachiopods (Terebratulina sp.) in island slope deposits. These taxa provide independent evidence for the deeper water aspect of this part of the Antigua Formation, in beds that also yield large, thin‐walled fossil sponges.     

Scolicia shirahamensis isp. nov.: a triple-corded scolicia and its ichnological implications

Abstract

Peculiar meniscate burrows with three sediment cords occur in early to middle Miocene tidal-flat deposits of southwestern Japan. Two of the cords are situated at the bottom and the other is at its center. Detailed observations of the burrow structures and comparative neoichnological studies of modern spatangoid burrows in a tidal flat revealed that the former two were true drainage tubes and the latter was fecal in origin. The trace fossil was thus assigned to the ichnogenus Scolicia. Based on these findings, a new ichnospecies Scolicia shirahamensis isp. nov. has been described here. The central sediment cord is seemingly identical to the drainage tube of the ichnogenus Bichordites, another ichnogenus that has been commonly ascribed to a fossil spatangoid burrow, similar to Scolicia. Careless ichnogeneric identification of a spatangoid burrow, based only on the central sediment cord, therefore, may produce an incorrect identification.     

The feeding activity of Echinostrephus molaris (de Blainville) in the central red sea

Echinostrephus molaris (de Blainville) is a small Indo‐pacific echinoid which burrows in coral reef limestone. Normally individuals do not leave the burrows so they cannot graze on algae growing around the burrow mouth. E. molaris catches floating algal particles with its long aboral spines. When a particle touches one of these organs or a tube‐foot in the area, the surrounding spines converge and grip it. Captured fragments are lowered to the test by further tube‐foot and spine action and are then passed across the ambitus towards the mouth. They are held by the oral tube‐feet and the shorter curved oral spines which aid ingestion. The behavioural and structural modifications shown for this habit are discussed. Burrowing and particle collecting have allowed E. molaris to occupy a particular niche on the reef. A similar method of food gathering is reported for Echinometra mathaei (de Blainville).     

Burrow construction and behavior of tilefish,Lopholatilus chamaeleonticeps,in Hudson Submarine Canyon

Synopsis During 22 daylight submersible dives in August 1979 numerous juvenile and adult tilefish, Lopholatilus chamaeleonticeps, were observed in and around vertical burrows in the clay substrate of portions of Hudson submarine canyon in depths from 110–230 m. The size and shape of the burrows varied considerably with the smallest juveniles occupying simple vertical shafts in the substrate. Larger fish were found in much larger burrows (up to 4–5 m in diameter and at least 2–3 m deep) that were funnel shaped in cross-section with the upper conical portions containing numerous smaller burrows of associated crabs. The range of burrow sizes observed suggests a regular sequence of burrow construction by tilefish and the associated crabs. Both juvenile and adult tilefish swam into the burrows head first and exited tail first. This behavior, which would preclude the possibility of ambushing prey, and evidence of predation by sharks and other tilefish, suggests that the burrow is a refuge from predators.Tilefish burrows appear to serve as a focus for biological activity. Species associated with the burrows included galatheid crabs, Cancer sp., Acanthocarpus alexandri, Homarus americanus, Heliocolenus dactylopterus and Conger oceanicus. Tilefish may play an important role in structuring outer continental shelf communities. They physically shape their environment and probably have significant biological interactions with the species that associate with their burrows.     

Microvertebrates preserved in mammal burrows from the Holocene of the Argentine Pampas: a taphonomic and paleoecological approach

Microvertebrates are a major component of many assemblages recovered from the Quaternary of the Argentine Pampas. The main goal of this paper is to analyse the taphonomic history of a Holocene microfossil bonebed, recovered from the infilling of a burrow. Evidences suggest the plains vizcacha Lagostomus maximus as the putative producer of the burrow. The assemblage includes individuals belonging to different taxa of mammals (marsupials and rodents) and reptiles (snakes). Taphonomic features suggest that the accumulation inside the burrow was related to flooding processes in the plain. The burrow was a natural trap that favoured the accumulation and preservation of remains corresponding to individuals from different sources. According to the taphonomic evidence, some individuals (Lagostomus maximus, Lestodelphys halli and Serpentes indet.) died inside the burrow, whereas others (Microcavia australis, Reithrodon auritus and Ctenomys sp.) died outside the burrow, and after a time of being exposed on the surface their remains were transported by surface run-offs into the burrow. The record of Lestodelphys halli and Serpentes indet. in the burrow produced by Lagostomus maximus could be related to a circumstantial use. Mammal burrows are a significant taphonomic mode for the late Cenozoic of the Argentine Pampas.     

Review of “Marine Hard Substrates: Colonization and Evolution,” a Thematic Session at the Geological Society of America Annual Meeting,Denver, USA,November 7–10, 2004

Planolites, although strictly a junior synonym of Palaeophycus, can be retained as a valid ichnotaxon on the basis of stability of nomenclature. As emended by Pemberton and Frey (1982) and consistently adopted by later authors, Planolites is diagnosed primarily on the basis that it is an unlined simple burrow, whereas Palaeophycus is a lined burrow. Other ichnotaxobases and ethological considerations are, without exception, subordinate to this directive.     

Ichnologic Note A New Ichnospecies of Cardioichnus from the Cretaceous (Albian) of New Mexico

A new ichnospecies of Cardioichnus, Cardioichnus biloba isp. nov., is described and documented from shallow marine strata of Cretaceous (Late Albian) age in southern New Mexico, USA. It is a heart-shaped resting trace which, unlike all other ichnospecies of Cardioichnus, is epichnial and increases in width to a bilobate trace, often with a raised anterior terminus. Only the posterior end of this trace is associated with Bichordites burrows, and the anterior end is bilobate and undisturbed. This represents a resting position which differs from other known species of this cubichnium, which are hypichnial and subquadrate in symmetry. The tracemaker is a spatangoid echinoid, most likely Heteraster d'Orbigny, 1853.     

Internal Disorder: Post-Mortem Burrows within the Tests of the Holasteroid Echinoid Echinocorys Leske (Upper Cretaceous to Paleocene)

A test of the Lower Paleocene holasteroid echinoid Echinocorys ex. gr. scutata Leske from Göynük, Turkey, preserves a Planolites beverleyensis (Billings) burrow close to the internal surface. Identification of such a specimen is fortuitous; only where the test has broken cleanly away from the internal mould can such burrows be exposed, if present at all. A second specimen, from the Upper Cretaceous of Norfolk, England, bears a sinuous boring identified as Planolites montanus Richter.     

Rogerella isp. Infesting the Pore Pairs of Hemipneustes striatoradiatus (Leske) (Echinoidea: Upper Cretaceous,Belgium)

An otherwise well-preserved test of the holasteroid echinoid Hemipneustes striatoradiatus (Leske) from the Emael Member, Maastricht Formation (Maastrichtian, Upper Cretaceous) of Belgium, is infested by encrusting bivalves and foraminiferans and the boring Rogerella isp. In this specimen, Rogerella preferentially infested and modified the ambulacral pore pairs of the echinoid close to the apex. This was not a commensal association. The echinoid test shows no growth deformations in response to this invasion; pore pairs are locally strongly infested; and encrusting invertebrates testify to the long post-mortem residence time of the test on the sea floor. Rather, the pore pairs of the dead echinoid were crannies attractive to settling larvae of acrothoracian barnacles, the producers of Rogerella.     

Stable deep-sea macrobenthic trace maker associations in disturbed environments from the Eocene Lefkara Formation,Cyprus

Ichnological analysis of Eocene deep-pelagic whitish chalky calcilutites interstratified with high-energy calcarenite beds in the lower part of the Petra Tou Romiou section (southern Cyprus) was conducted to interpret the development and evolution of the trace maker associations during calcilutite deposition after high-energy episodes. The trace fossil assemblage from the chalky calcilutites consists of Chondrites isp. (Chondrites intricatus and Chondrites targionii), Planolites isp., Taenidium isp., Thalassinoides isp., and Zoophycos isp., typical of the Zoophycos ichnofacies. A composite chalky ichnofabric reveals a multi-tiered association of burrowing animals: the uppermost tier determines a mottled background, the upper tier shows the highest trace fossil abundance and diversity (Planolites, Taenidium and Thalassinoides), the middle tier features Zoophycos and large Chondrites, and the deepest tier consists mainly/exclusively of small Chondrites. This ichnofabric has an autocomposite character, associated with bioturbation by a single ichnocoenosis and gradual upward migration of the tiered macrobenthic community as the pelagic calcilutite sedimentation slowly progresses. There are no changes in the trace fossil assemblage between or within calcilutite intervals, regardless of the associated calcarenite beds. This supports a stable, mature, background calcilutite trace maker association, which recovers shortly after the deposition of high-energy calcarenites. In turn, there would have been a rapid re-establishment of paleoenvironmental conditions during pelagic calcilutite accumulation after episodic deposition of any calcarenitic material.     

Microscale oxygen distribution in various invertebrate burrow walls

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

Burrow depth,carbon dioxide and reproductive success in Sand Martins Riparia riparia

Capsule: Carbon dioxide (CO₂) concentrations in the burrows of Sand Martins Riparia riparia increase with depth but have no detectable impact on fledging success.

Aims: To investigate whether burrow depth and CO₂ concentrations influence reproductive success in Sand Martins.

Methods: We monitored two Sand Martin colonies along the River Lune, Lancashire, UK, to investigate the effect of burrow depth on reproductive success. We also measured CO₂ levels in a sample of burrows to test whether burrow depth predicts CO₂ concentration, and to test for a relationship between CO₂ concentration and breeding success.

Results: Burrow depth was significantly correlated with fledging success, but the correlation was positive in first broods and negative in second broods. The highest CO₂ concentration recorded was 73?650?ppm and the mean concentration across burrows was 31?757?ppm. However, while CO₂ concentrations were positively correlated with burrow depth after controlling for the number and age of nestlings, they were not correlated with reproductive success.

Conclusion: There are reproductive costs associated with deeper burrows in second broods, but these could not be attributed to CO₂ concentrations despite the exceptionally high levels recorded. This study highlights the need for further investigation into gas exchange and the potential impacts of, or adaptations to, CO₂ accumulation in avian burrows.     

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

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

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.     

Three-dimensional morphology and palaeobiology of the trace fossil Dactyloidites jordii nov. isp. from the Carboniferous of England

The diverse ichnological assemblage from the outcrops near Howick (Northumberland, United Kingdom) is exceptionally well-preserved. Among these Carboniferous ichnotaxa is a new ichnospecies of Dactyloidites. Specimens were collected and processed using high-resolution serial grinding and photography to produce an accurate and precise three-dimensional model of these new burrows in full colour. The model produced in association with petrographic thin sections and field observations is used as the basis for comparison between Dactyloidites jordii nov. isp. and other ichnospecies of Dactyloidites. The current taxonomic status of the ichnogenus is examined and reviewed. D. jordii nov. isp. is a broadly bisymmetrical, stellate to palmate burrow composed of numerous long, narrow rays that exhibit three orders of branching arranged into tiered galleries radiating from a central shaft. The trace maker is suggested to be a vermiform organism with an adaptive burrowing strategy that facilitates alteration of its burrow construction to accommodate suboptimal sediment conditions. The adaptive nature of trace-making organisms and the inherent anisotropy of many burrowed media highlight the need for ichnologists to provide a type series rather than a single holotype, in order to capture the inherent range of common burrow morphologies.     

Burrow defense behaviors in a sand-bubbler crab, Scopimera globosa, in relation to body size and prior residence

The burrow defense behaviors in a sand-bubbler crab, Scopimera globosa, living on a tidal flat, were experimentally examined. Body size and prior residence influenced the results of struggles for the burrows, and large individuals or the burrow owners won in most cases when the intruders were not significantly larger than the owners. Most large owners defended their burrows by directly fighting their opponents. On the other hand, small owners defended their burrows in three different ways. (1) Owners fought directly against same-sized or smaller intruders. For larger intruders, (2) most owners returned to their burrows when the owner was nearer to the burrow than the intruder (returning behavior), and (3) owners sat motionless when the intruder was nearer to the burrow than the owner (sitting behavior). Success ratios of the three types of burrow defense were 38.2%, 88.5%, and 100%, respectively. It was considered that sitting behavior of the cryptically colored S. globosa has evolved because intruders cannot see motionless owners and consequently cannot detect the owner's burrow. Received: October 6, 2000 / Accepted: January 22, 2001     

New Ichnospecies of Cardioichnus from the Miocene of the Guadalquivir Basin,Huelva, Spain

A new ichnospecies of Cardioichnus, Cardioichnus reniformis isp. nov., is documented and described from marine strata of the Late Miocene of southwestern Spain. It is a bilobate resting trace with a heart‐shaped outline and is related to the work of a wedge‐shaped irregular echinoid of the spatangoid group.     

Diet of the killifish Rivulus marmoratus collected from land crab burrows,with further ecological notes

Synopsis One-hundred eleven specimens of the cyprinodont fish Rivulus marmoratus were trapped from burrows of the great land crab Cardisoma guanhumi in east central Florida and the entire alimentary tract removed for analysis of food items. Three categories of burrows (inactive, active and sealed) were sampled to examine possible relationships between burrow type and presence of R. marmoratus. Forty percent of the guts examined contained no food items, while the remaining 60% contained items of both terrestrial and aquatic origin. Capture rates among burrow categories did not vary significantly, and most collections consisted of 1 or 2 individuals per burrow. Mean standard length of fish within a burrow declined when more than 3 specimens were captured from a burrow. Supplemental observations on reproductive status and burrow water temperature are noted.