The Mississippian Tetrapod Footprint Ichnogenus Palaeosauropus: Extramorphological Variation and Ichnotaxonomy

Palaeosauropus primaevus is a tetrapod footprint ichnotaxon first described from the Upper Mississippian (Visean) Mauch Chunk Formation near Pottsville, Pennsylvania, United States. Our relocation of the type locality and stratigraphic horizon of P. primaevus, a long-available but unstudied collection of tetrapod footprints from these strata, and our new collections allow a much fuller characterization of this ichnotaxon and the range of extramorphological variation encompassed by it. P. primaevus is characterized as the footprints of a quadruped with a pentadactyl pes and a tetradactyl manus, in which the pes frequently oversteps the manus and with which tail drags are common. In the manus, all digits are relatively broad and have rounded tips, digit III is longest, and digit IV is more widely separated from digit III than the other digits are from each other. The pes has five digits that are also wide and blunt-tipped, digit IV is longest, and digit V projects nearly laterally. P. primaevus is the track of a relatively large temnospondyl (～400 mm gleno-acetabular length) and documents the Mississippian presence of such large amphibians long before their body fossil record. Palaeosauropus also occurs in Mississippian strata in Indiana and is distinguished from the geologically younger but similar temnospondyl footprint ichnogenus Limnopus by its relatively narrower manus and pes that lack broad and rounded sole impressions.     

The Fish Swimming Trace Undichna from the Mississippian Mauch Chunk Formation,Eastern Pennsylvania

Undichna, the swimming trace of fish, was recovered from the middle member of the Mauch Chunk Formation in eastern Pennsylvania in strata of Late Mississippian (Visean) age. These traces represent the only evidence of fish known from the Mauch Chunk Formation and the Carboniferous of Eastern Pennsylvania. The Mauch Chunk Formation in the study area is characterized by an ephemeral fluvial depositional environment that yields a diverse invertebrate ichnoassemblage of the Scoyenia ichnofacies and a tetrapod footprint assemblage dominated by the tracks of temnospondyl amphibians. The Undichna specimens are preserved on fine-grained, mudstone-draped, rippled sandstone. Two specimens, assigned to U. Britannica, consist of pairs of well-defined, narrowly incised, sinusoidal wave traces that are out-of-phase. Two specimens, assigned to U. quina, consist of two pairs of in-phase sinusoidal waves intersected by an additional single wave with a greater amplitude. Two specimens, each composed of a single wave, are assigned to U. unisulca.

The slabs of rock with the Undichna traces also preserve tetrapod undertracks assigned to Batrachichnus, Matthewichnus, and Hylopus. This trace fossil association indicates the presence of small fish, some with caudal and/or anal fins and some with pectoral, pelvic, and anal and/or caudal fins, that touched the sediment during periods of intermittent flooding of subaerial surfaces walked upon by early tetrapods. Examples of Undichna from Mississippian or older deposits, especially from a fluvial paleoenvironment, are rare; few have been reported from the Devonian and the Lower Carboniferous.     

Isaac Lea's Palaeosauropus (= “Sauropus”) primaevus: A Review of His Discovery

In 1849, Isaac Lea named Sauropus primaevus for footprints from Mount Carbon, Pennsylvania, USA, then the oldest fossil vertebrate footprints reported. In 1902, O. P. Hay constructed a new ichnogenus Palaeosauropus for this ichnospecies. Palaeosauropus has been one of the most frequently reported Mississippian footprint ichnogenera in North America and remains a valid ichnotaxon. The holotype of Palaeosauropus (= “Sauropus”) primaevus (referred to as P. primaevus), consisting of a single manus/pes pair, is described and illustrated in Lea (1853) Lea, I. 1853. On the fossil foot-marks in the Red Sandstones of Pottsville, Schuylkill County, Penna. Transactions of the American Philosophical Society, 10(new series): 307–315.  [Google Scholar] and is housed at the Academy of Natural Sciences in Philadelphia, Pennsylvania (ANS9752). Lea's large specimen of P. primaevus (approximately 86 cm by 53 cm), that included a trackway of six manus/pes pairs, described and illustrated in 1853 and 1855, was a combination of ANS9752 and a second specimen represented by a plaster cast housed at the National Museum of Natural History (USNM487148). Historical documents and examination of the Mauch Chunk Formation at Mount Carbon, Pennsylvania, enabled the identification of Lea's tracksite, originally reported to be a few hundred feet (about 75 m) from the former Mount Carbon Hotel. Our forensic evidence indicates the type locality for P. primaevus is approximately 90 m south from the southwest corner of Centre and Main Streets in Mount Carbon, Pennsylvania, with geographical coordinates of N 40° 40' 25.7”, W 76° 11' 14.9”. The type locality is within the middle member of the Mauch Chunk Formation, a fluvial sequence of late Mississippian (Visean) Age.     

The stapes of Edops craigi and ear evolution in the lissamphibian stem group

The Lower Permian temnospondyl Edops craigi exemplifies an early and plesiomorphic condition of the single ear ossicle or stapes among the temnospondyls, the probable stem group of lissamphibians. In Edops, the 11-cm-long bone is more massive than in other temnospondyls, has a distinct neck, a dorsal crest and incompletely subdivided footplate and ventral process. Despite a range of invariances, temnospondyl stapes were much more diverse than previously conceived. A survey of described stapes gives insight into character evolution of the ear ossicle in the lissamphibian stem group. These include alternative patterns of paedomorphosis, proportional size change, morphology of tympanic region and reorientation of the auditory apparatus.     

A Temnospondyl Trackway from the Early Mesozoic of Western Gondwana and Its Implications for Basal Tetrapod Locomotion

Background

Temnospondyls are one of the earliest radiations of limbed vertebrates. Skeletal remains of more than 190 genera have been identified from late Paleozoic and early Mesozoic rocks. Paleozoic temnospondyls comprise mainly small to medium sized forms of diverse habits ranging from fully aquatic to fully terrestrial. Accordingly, their ichnological record includes tracks described from many Laurasian localities. Mesozoic temnospondyls, in contrast, include mostly medium to large aquatic or semi-aquatic forms. Exceedingly few fossil tracks or trackways have been attributed to Mesozoic temnospondyls, and as a consequence very little is known of their locomotor capabilities on land.

Methodology/Principal Findings

We report a ca. 200 Ma trackway, Episcopopus ventrosus, from Lesotho, southern Africa that was made by a 3.5 m-long animal. This relatively long trackway records the trackmaker dragging its body along a wet substrate using only the tips of its digits, which in the manus left characteristic drag marks. Based on detailed mapping, casting, and laser scanning of the best-preserved part of the trackway, we identified synapomorphies (e.g., tetradactyl manus, pentadactyl pes) and symplesiomorphies (e.g., absence of claws) in the Episcopopus trackway that indicate a temnospondyl trackmaker.

Conclusions/Significance

Our analysis shows that the Episcopopus trackmaker progressed with a sprawling posture, using a lateral-sequence walk. Its forelimbs were the major propulsive elements and there was little lateral bending of the trunk. We suggest this locomotor style, which differs dramatically from the hindlimb-driven locomotion of salamanders and other extant terrestrial tetrapods can be explained by the forwardly shifted center of mass resulting from the relatively large heads and heavily pectoral girdles of temnospondyls.     

Scorpionid Resting Trace from The Lower Permian of Southern New Mexico,USA

We describe an arthropod body impression associated with arthropod trackways of the ichnogenus Stiaria from the Lower Permian (upper Wolfcampian) Robledo Mountains Formation (Hueco Group) in the Prehistoric Trackways National Monument of southern New Mexico. The probable producer of these traces was a scorpion, and we name the likely scorpionid resting trace Alacranichnus braddyi, new ichnogenus and ichnospecies. There are no prior reports of scorpionid body impressions from terrestrial settings in the fossil record.     

The otic region of Doleserpeton (Temnospondyli) and its implications for the evolutionary origin of frogs

There is increasing evidence that the Palaeozoic temnospondyl amphibians had a frog‐like tympanic hearing system. For this reason, the otic region of Doleserpeton is described and compared with modern anurans. The otic capsules are expanded laterally and ventrally relative to other temnospondyls. The opisthotic has a bulbous ventral region resembling the ventrolateral ledge in modern frogs. Two lateral processes are located on the paroccipital process. Comparison with the condition in modern anurans with a tympanic hearing system shows that this may have been the attachment site for the tympanic annulus. Parts of the osseous labyrinth are also described. The inner ear shows numerous features resembling the condition found in frogs. These include strong evidence for the presence of a lissamphibian‐type perilymphatic duct most closely resembling that of anurans. This is the first time such a perilymphatic system has been described in any Palaezoic form. The posterior part of the braincase shows a jugular foramen closely associated with the perilymphatic foramen, as in anurans. Although the distribution of these traits among other temnospondyl groups remains little known, the sum of the evidence points to affinities between anurans and temnospondyls, and adds to the evidence for a close relationship between anurans and the Permian amphibamid Doleserpeton. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 738–751.     

PETRIFIED PLANTS FROM THE UPPER MISSISSIPPIAN OF NORTH AMERICA. I: THE SEED RHYNCHOSPERMA GEN. N.

Petrified seeds of Upper Mississippian age (Chester Series) are described from sediments from northwestern Arkansas. Specimens of Rhynchosperma quinnii gen. et sp. n. are radially symmetrical and range in length from 1.2 to 2.2 cm and 1.0 to 1.3 cm at the greatest diameter. The seeds are ovate with a tapered apex having 8–10 integumentary ribs and a broadly rounded chalaza. The integument is two-parted, consisting of an outer presumably soft tissue with secretory canals and an inner layer constructed of longitudinally oriented thick-walled cells. Nucellus and integument are confluent, but they diverge near the seed midlevel with the free portion of the nucellus forming a dome-shaped pollen-receiving structure. Additional features such as vasculature and the structure of well-preserved megagametophytes are described. The petrified specimens are compared with taxa instituted for casts and impressions which include Rhynchogonium and Boroviczia. The ovulate versus cupulate nature of these genera which has remained uncertain until now appears to be settled in favor of an ovular identity.     

The braincase and middle ear region of Dendrerpeton acadianum (Tetrapoda: Temnospondyli)

Dendrerpeton acadianum from the Westphalian A (Upper Carboniferous) of Joggins, Nova Scotia, is a phylogenetically and chronologically early temnospondyl. Its external cranial anatomy has been used previously to suggest the presence of a tympanic membrane, and thus of an ear adapted to the perception of airborne sound. However, supporting evidence provided by stapedial and braincase morphology has so far been lacking. The braincase and middle ear region have remained almost wholly unknown. CT scanning and 3-D computer reconstruction of BMNH R.436 have been used to shed light on these important areas. Both stapes prove to be present in the specimen; the right stapes is distorted, but the left stapes lies inside the cranial cavity and is perfectly preserved. The latter resembles the stapes of the relatively few other temnospondyls in which the bone has been described and is most similar to that of Doleserpeton . The morphology and orientation of the stapes provide strong evidence for the presence of an ear adapted to the perception of airborne sound, with similarities to the extant anuran condition. The reconstructed braincase shows a high degree of similarity to that of other adequately known temnospondyls. This gives supporting evidence that D. acadianum is correctly placed in the temnospondyl phylogeny and thus demonstrates one of the earliest hearing systems adapted to the perception of airborne sound that can be homologized with the extant anuran condition.  © 2005 The Trustees of the Natural History Museum, Zoological Journal of the Linnean Society , 2005, 143 , 577−597.     

Multiple-Rusophycus (Arthropod Ichnofossil) Assemblages and Their Significance

Rusophycus is an ichnogenus comprising shallow burrows generally attributed to trilobites and other arthropods. The paleoethological interpretation of these structures is not conclusively known; workers variously have attributed the ichnogenus to feeding, resting, hiding/escape, hunting or nesting behaviors. Rusophycus morphology varies from unornamented, bilobed forms to forms that preserve details of ventral anatomy of the trace maker. Rusophycus occurs as single or multiple impressions. Some examples of multiple-Rusophycus assemblages are clearly the result of the activity of a single animal (e.g., serially arranged impressions of the same size), but others represent the activity of several individuals. Associated traces representing multiple individuals are especially interesting, for they may give evidence of complex behavior, for example, interactions between the trace makers. Some of the multiple-trace assemblages show alignment (congruent anterior-posterior orientation of the individuals) suggesting rheotactic behavior; other assemblages comprise randomly oriented traces. The difference between the aligned and randomly oriented assemblages may reflect differences in the current energy and amount of available food, and may also suggest the following feeding modes for these benthic-feeding arthropods: (1) alignment with head into the current in resource-poor environments, (2) orientation with head at an oblique angle to the current in high-energy, resource-poor environments, and (3) random orientation in low-energy, resource-rich environments.     

Oldest fossil basidiomycete clamp connections

A rachis of the fossil filicalean fern Botryopteris antiqua containing abundant septate hyphae with clamp connections is preserved in a late Visean (Mississippian; ~330 Ma) chert from Esnost (Autun Basin) in central France. Largely unbranched tubular hyphae pass from cell to cell, but may sometimes produce a branch from a clamp connection. Other clamp-bearing hyphae occur clustered in individual cells or small groups of adjacent host cells. These hyphae may be tubular, catenulate with numerous hyphal swellings, or they may display a combination of both. The Visean hyphae with clamp connections predate Palaeancistrus martinii, the heretofore oldest direct fossil evidence of Basidiomycota, by some 25 Ma.     

THE POSTCRANIUM OF ARCHEGOSAURUS DECHENI, AND A PHYLOGENETIC ANALYSIS OF TEMNOSPONDYL POSTCRANIA

Abstract:  The gharial-like Archegosaurus decheni from the Permian/Carboniferous boundary of south-west Germany is one of the best known temnospondyls. Based largely on new material, we restudied the postcranial anatomy of this species, including ontogenetic aspects. A. decheni has 24 presacral vertebrae, and the length of the deep tail exceeds the length of the rest of the body. Neural spines are moderately high and slowly become differentiated during ontogeny. The intercentra start to ossify very late. Distal uncinate processes are developed on the anterior ribs in adult specimens. Only the ventral portion of the scapula is ossified. The slender ilium and the ischium are not co-ossified, and the pubis remained cartilaginous. Among stereospondylomorph temnospondyls, the very short and rudimentarily developed humerus exhibits a unique morphology. Carpals and tarsals start to ossify only in the largest specimens. The poorly ossified postcranium indicates that A. decheni was primarily an aquatic temnospondyl. We undertook a phylogenetic study of A. decheni and 16 other temnospondyls, based exclusively on postcranial characters. We analysed 52 characters, obtaining a single most parsimonious tree that agrees in many aspects with cranium-based analyses: Trimerorhachis and Dvinosaurus form a basal clade (Dvinosauria), followed by monophyletic euskelians (dissorophoids plus eryopids) and finally the stereospondylomorphs, within which A. decheni is nested above Sclerocephalus . Among stereospondyls, trematosaurids and metoposaurids form a clade, whereas the chigutisaurid Siderops is nested with capitosauroids. The primitive condition of Temnospondyli is still not adequately understood, especially the degree of terrestriality of the earliest temnospondyls.     

The evolution of metamorphosis in temnospondyls

Temnospondyls, possible relatives of extant amphibians and crudely similar to recent salamanders, are known from larval, neotenic and metamorphosed stages. Here, ontogenetic data of various temnospondyl taxa are analysed in order to recognize metamorphosis. Here, metamorphosis is strictly defined as a shift from an aquatic to a terrestrial existence. Following a check-list of criteria, the most likely metamorphosis-induced changes are proved in three temnospondyl lineages: eryopids, zatrachydids and dissorophoids. In a few other, unrelated taxa, terrestrial adults are known but no larval or metamorphosing forms. The distribution of metamorphosis among the Temnospondyli does not strictly correlate with phylogeny, which highlights the widespread occurrence of neoteny. In each group, characteristic patterns of metamorphosis are described and compared. Among temnospondyls, dissorophoids had the most intensive type of metamorphosis, characterized by a condensed ontogeny and a relatively small body size. The result was a distinct transformed morphotype with far-reaching terrestrial adaptations.     

The origin of modern amphibians: a re‐evaluation

There are currently three competing hypotheses seeking to explain the evolutionary origins of modern amphibians. The lepospondyl hypothesis holds that the lysorophian lepospondyls constitute the sister taxon to all lissamphibians. The temnospondyl hypothesis suggests that modern amphibians are most closely related to the dissorophoid temnospondyls. Finally, the polyphyletic hypothesis posits that the modern amphibian orders have separate evolutionary origins from among different groups of Palaeozoic tetrapods. Here, we review the character matrices used in previous studies. These data sets differ significantly in choice of characters. Therefore, we built a matrix based on data from all three hypotheses and analysed key taxa phylogenetically using both Bayesian inference and parsimony. Uncorrected, the supermatrix yielded inconclusive results, demonstrating the presence of at least two phylogenetic optima. When the data were corrected according to new observations on Doleserpeton, Eocaecilia, and other fossil forms, the phylogeny supported the temnospondyl hypothesis of lissamphibian origins. This conclusion is also supported by a careful study of character changes in the individual lineages. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 457–469.     

THE MORPHOLOGY OF STIGMARIA STELLATA

Stigmaria stellata Goeppert is a lycopod underground system occurring in Upper Mississippian and equivalent age rocks of Europe and North America. This taxon has previously been based on impressions exhibiting radiating ridges and furrows around each lateral appendage scar and numerous polyhedral projections on the remainder of the axis. Anatomically preserved specimens from the Chester Series (Upper Mississippian) of Illinois reveal that the distinctive surface pattern of this species results from polyhedral wedges of thick-walled cells in the outer cortex. Decortication produces a smooth outer surface that is indistinguishable from that of the much more abundant Stigmaria ficoides. The structure of S. stellata is quite similar to the structure of other petrified stigmarians, but the following are some of the anatomical characters that distinguish it: (1) presence of abundant polyhedral wedges of thick-walled cells in the outer cortex; (2) absence of secondary cortex; (3) very tall rays associated with appendage traces that remain confluent with the secondary xylem to its outer margin; (4) the absence of a connective in the lateral appendages. The anatomical characters of Stigmaria stellata confirm it as a taxon of at least specific rank.     

Reinterpretation of crinoid-platyceratid interaction

The well-known association of platyceratid gastropods with crinoids has traditionally been considered an example of coprophagous commensalism. The Occurrence of several crinoid 'stands' ( Platycrinites sp.) from closely spaced bedding surfaces in the upper Mississippian Wymps Gap Limestone member of the Mauch Chunk Formation of southwestern Pennsylvania encourages reinterpretation of this relationship. Crinoid calyces were collected from five separate clusters, two of which contained associated platyceratids. 'Infested' crinoids either died prematurely or were stunted, compared to crinoids in the 'uninfested' clusters. Platyceratid attachment apparently had an adverse effect upon crinoid growth. Serial acetate peels show positioning of the gastropod on the crinoid tegmen over a highly developed anal tube, or chimney. However, the terminus of the anal tube abuts the gastropod's shell and is poorly situated for fecal ingestion by the snail. We suggest that the snail probably pursued another trophic strategy, perhaps taking advantage of aerosol filtration by the crinoid and elevation above the substrate.     

The cranial morphology of Greererpeton burkemorani Romer (Amphibia: Temnospondyli)

The skull of Greererpeton burkemorani Romer, a temnospondyl amphibian from the Upper Mississippian at Greer, West Virginia is described. A detailed account of the stapes of a Mississippian amphibian is given for the first time and its function is discussed. It is suggested that the stapes formed the principal element of support for the back of the braincase and resisted potential dislocation of the otico-occipital region from the skull roof during contraction of the hypaxial musculature.
Greererpeton is included in the Colosteidae and an amended diagnosis of the family is given. Erpetosaurus differs from Colosteus, Greererpeton and Pholidogaster in the pattern of bones in the skull roof and palate, the dentition and the otic region and, consequently, it is removed from the Colosteidae. The Temnospondyli are considered to be a monophyletic group characterized by the development of a connection between the dorsal portion of the occipital arch, the exoccipital bones, and the skull roof. The loxommatids are removed from the Temnospondyli as they retain the plesiomorphic condition of braincase attachment which relies exclusively on derivatives of the auditory capsules.
On the basis of similarities in the structure of the braincase, palate and manus it is suggested that microsaurs are the collateral descendants (sister group) of temnospondyls. This relationship may account for the large number of similarities in the three living groups of Amphibia: Anura are generally believed to have descended from temnospondyls, while the Urodela and Apoda are often considered to have descended from microsaurs. These systematic conclusions endorse the recent suggestions that neither the Lepospondyli nor the Labyrinthodontia are natural groups, and both terms should be abandoned.     

Redescription and Phylogenetic Analysis of the Mandible of an Enigmatic Pennsylvanian (Late Carboniferous) Tetrapod from Nova Scotia,and the Lability of Meckelian Jaw Ossification

The lower jaw of an unidentified Pennsylvanian (Late Carboniferous) tetrapod from Nova Scotia – the “Parrsboro jaw”- is redescribed in the light of recent tetrapod discoveries and work on evolution of tetrapod mandibular morphology and placed for the first time in a numerical cladistics analysis. All phylogenetic analyses place the jaw in a crownward polytomy of baphetids, temnospondyls, and embolomeres. Several features resemble baphetids and temnospondyls including dermal ornamentation, absence of coronoid teeth, and presence of coronoid shagreen. Dentary dentition is most similar to Baphetes. An adsymphysial toothplate may not preclude temnospondyl affinity. An apparent large exomeckelian fenestra, with the dorsal foraminal margins formed by an unossified element, echoes the morphology of the stem tetrapod Sigournea and is unusually primitive given the other features of the jaw. The jaw may thus provide an example of an intermediate stage in Meckelian element evolution.     

The evolution of the scalation pattern in temnospondyl amphibians

In most Palaeozoic temnospondyls, thin round-oval scales covering the flanks and the back of the trunk can be distinguished from ventral, elongate gastral scales arranged in a chevron pattern. The extensive growth series of the temnospondyl Sclerocephalus reveals that the morphology of the gastral scales in small larvae corresponds to the round-oval scales of the rest of the body. During subsequent ontogeny, the gastral scales differentiate and attain a spindle-shaped morphology. The tapering end of each gastral scale fits into a dorsal groove on the medial adjacent scale. This arrangement allowed telescoping of the scales and thus provided a high degree of flexibility. In the ontogenetically most advanced specimens of Sclerocephalus the gastral scales attain a rhomboid outline, and the articulation by well-defined facets has reduced the flexibility between them. In most temnospondyls, the gastral scales retain the 'juvenile' spindle-shape or the 'larval' round-oval shape, which can be interpreted as a paedomorphic trait. This suggests that the different types of gastral scales in temnospondyls, as well as the scales of the back and the flanks, can be traced back to the same Anlage of round-oval scales that differentiated early in ontogeny. In the Mesozoic, a complete reduction of dermal scalation occurred independently in distinct dissorophoid, capitosauroid, and trematosauroid temnospondyls. This reduction was probably the result of several factors unique to each group, such as cutaneous respiration, the demand for greater mobility, and the decreased importance of belly protection in fully aquatic temnospondyls.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 815–834.