Online Data Appendix 2 for “Woodland hypothesis for Devonian tetrapod
evolution” by G. J. Retallack
These reviews of selected Devonian and Carboniferous fossil vertebrate localities
outline the basis for paleoenvironmental interpretations.
Bundenbach, Germany. Slate quarries into the Hunsrückschiefer near
Bundenbach and Gemünden in the Eifel region of Germany are famous as the source of a
fish fauna of agnathans, placoderms and dipnoans of late Pragian to early Emsian age.
The slates are gray shales deposited in an open marine setting, as indicated by their
pyritized fauna of sponges, hydrozoans, corals, jellyfish, comb-jellies, snails, bivalves,
cephalopods, brachiopods, bryozoans, annelids, crustaceans, trilobites, horseshoe crabs
and echinoderms (Bartels et al. 1998).
Beartooth Butte, Wyoming. On the high eastern crest of Beartooth Butte a
paleochannel filled with red siltstone, sandstone and claystone breccia yields a diverse
fossil fish fauna of cephalaspids, ostracoderms, thelodonts, placoderms, acanthodians and
dipnoans, as well as rare eurypterids and spores of mid-late Emsian age (Elliott and
Johnson 1997; Tetlie 2007). Some fish and eurypterids are articulated, but most are
disarticulated in red paleosols with drab-haloed root traces (Fluvents or Alluvial Soils:
Retallack personal observations), of a stream set within a paleocanyon of Ordovician
dolomite.
Zachełmie, Poland. The upper Wojciechowie Formation in the disused
Zachełmie quarry in the Holy Cross Mountains of Poland has numerous yielded
footprints similar to the foot anatomy of aquatic tetrapods such as Ichthyostega.
Conodonts of the costatus zone (mid-Eifelian) were found 20 m above the trackways
which are in mud-cracked, red, stromatolitic dolomicrites with burrows of limited
diversity (ichnogenera Thalassinoides, Skolithos). These deposits lack definitive marine
fossils, and represent supratidal flats of a coastal estuary or lagoon (Niedźwiedzki et al.
2010), and are thus weakly developed paleosols (Fluvents or Alluvial Soils: Retallack et
al. 2009). The lower Wojciechowie Formation is similar to Lower-Middle Devonian nonmarine sediments of Germany (Schweitzer and Giesen 2002; Stets and Schäfer 2009).
The overlying Kowala Formation is a marine coral limestone (Narkiewicz and
Narkiewicz 2010).
Achanarras, Scotland. The Achanarras Limestone of the lower Caithness
Flagstones has yielded numerous articulated placoderms, dipnoans, acanthodians,
sarcopterygians and actinopterygians in a finely laminated lake beds (Rayner 1962). This
and other Scottish fossil fish localities appear to have been part of a large Orcadian lake,
dated by spores as equivalent in age to the Eifelian kockelianus conodont zone and the
Kačák event (Marshall et al. 2007).
Mt Crean, Antarctica. The Aztec Siltstone on the eastern and southern spurs of
Mt Crean exposes a sequence of paleosols between cross-bedded fluvial paleochannel
sandstones (McPherson 1980; Retallack 1997a) with fossil fish of the Givetian stage
(Long et al. 2008). Fragmentary jaws and scales of sarcopterygians are found along with
disarticulated remains of placoderms, sharks and acanthodians in three distinct
sedimentary settings: (1) atop green-red root-mottled paleosols with shallow calcareous
nodules (well drained Aridisols); (2) within light green-gray massive siltstones with root
Retallack: Pennsylvania trees and tetrapods
2
traces (gleyed Inceptisols); and (3) within dark gray finely laminated shales (lacustrine:
Retallack, 1997a).
Thurso, Scotland. Along Pennyland Shore west of Thurso in Caithness, northern
Scotland, laminated gray shales of the Upper Caithness Flags yield articulated
sarcopterygians and placoderms of the upper Givetian stage (Friend and Williams 1978).
These large Orcadian lake deposits have been quarried for many years without finding a
trace of marine influence (Miller 1867; Marshall et al. 2007).
Red Hill, Nevada. Quarries into Denay Limestone on the western slope of Red
Hill, 67 km north-northwest of Eureka, have yielded a variety of sarcopterygian, dipnoan,
acanthodian, placoderm and actinopterygian fish, some articulated (Murphy et al. 1976;
Reed 1985, 1986). The Denay Limestone contains brachiopods, tabulate corals and
conodonts of the upper Givetian disparils zone (Schultze 2005) below the fish bed. Along
with the fish, finely laminated, dark gray shales contain a restricted marine fauna,
including conulariids, hexactinellid sponges, algae, and land plant fragments. These
marine shales pass up-section into red paleosols of well drained coastal plains.
Howitt Spur, Victoria. A diverse fauna of sarcopterygian, actinopterygian,
acanthodian, placoderm fish and sharks of the lower Frasnian stage has been found from
the Snowy Plains Formation at the foot of the long eastern spur of Mt Howitt in the
Victorian Alps of southeastern Australia (Long 1991). Many of the fish are articulated in
laminated black shale, without marine fossils, and presumably lacustrine. The 52-cmthick, fish-bearing shales underlie trough cross-bedded quartz sandstone of a fluvial
paleochannel, and overlie 72 cm of ripple-marked fine sandstone above 66 cm of planarbedded fine sandstone, probably a lake margin beach deposit. These sandstones overly a
thin gray rooted claystone (Inceptisol) with root traces and wood fragments up to 6 cm
wide, which in turn overlies a coarse grained to conglomeratic sandstone, with trough
cross-bedding.
Miguasha, Quebec. Partial articulated skeletons of Elpisostege watsoni
(prototetrapod) are found with a fish fauna close to the Givetian-Frasnian boundary (ca.
385 Ma) in siderite nodules within laminated gray shales of the Escuminac Formation
near Miguasha, Quebec, Canada (Schultze and Cloutier 1996). Eurypterids could be
taken as evidence of limited marine influence as in a coastal lagoon, but conchostracans
are usually freshwater. Also found are sarcopterygian, heterostracan, anaspid,
acanthodian, placoderm and actinopterygian fish, as well as aquatic scorpions, tree trunks
up to 13 cm diameter (Callixylon) and leafy shoots (Archaeopteris halliana: Carroll et al.
1972). Without any definitive marine fossils, this deposit is best interpreted as a coastal
lake or lagoon.
Lode, Latvia. Panderichthys rhombolepis and Livoniana multidentata are upper
Givetian to lower Frasnian sarcopterygians (prototetrapods of Long and Gordon 2004)
from a greenish-gray smectite clay lens within clayey red beds (paleosols) and sandstones
of the Gauja Formation in Lode quarry, Latvia (Ahlberg et al. 2000; Boisvert 2005).
Panderichthys is represented by complete fish, but Livoniania only by fragments. Other
fish from the quarry include placoderms and other sarcopterygians. Nearby sandstones
include rare articulated brachiopods, but enclosure of the fish-bearing shale by massive
red siltstones (well drained paleosols?) and associated conchostracans are evidence of
largely freshwater parts of estuaries and lagoons (Kursš 1992; Upeniece and Upenieks
1992).
Retallack: Pennsylvania trees and tetrapods
3
Gogo, Western Australia. Articulated skeletons of sarcopterygians (including the
Gogonasus andrewsae which shares more limb and nasal features with prototetrapods
than other lobefin fish), placoderms, actinopterygians and lungfish from northwestern
Australia come from large calcareous nodules within lower Frasnian (Mesotaxis
falsiovalis conodont zone), black shales of the Gogo Formation (Young et al. 2010). This
open marine deposit includes marine ammonoids, nautiloids and conodonts. The Gogo
Formation accumulated offshore and laterally equivalent to coral-algal Sadler Limestone,
part of the fossil barrier reefs of the Canning Basin (Long et al. 2006).
Blind Fiord, Nunavut. The Fram Formation near Blind Fiord, southern Ellesmere
Island, has yielded a variety of lower Frasnian sarcopterygians, placoderms, and lungfish,
as well as articulated skeletons of Tiktaalik roseae, a sarcopterygian with simplified limb
structure (prototetrapod of Long and Gordon 2004; Shubin et al. 2006) and the first clear
indications of a neck (Daeschler et al. 2006). Partially articulated Tiktaalik skeletons were
clustered in a small area at the top of a clayey, deep-calcic, red paleosol, with drab-haloed
root traces (Miller et al. 2007; Shubin 2008), which discolor the skulls and skeletons.
Valentia Island, Ireland. The Valentia Slate Formation is a sequence of purple
sandstone and slates below Famennian fossil fish localities, and perhaps also
stratigraphically below lower Frasnian miospores. The coastal rock platform exposes
reveal several trackways, which include body drag marks and poorly differentiated digits
of presumed aquatic tetrapods. Nevertheless, the tracks are deeply impressed as if made
on land (Stossel 1995).
Genoa River, Victoria. The Combyingbar Formation exposed in the Genoa River,
Victoria, Australia, has yielded two distinct kinds of tetrapod trackways, a larger one with
outward-directed prints as in Acanthostega, and a smaller one with differently sized
manus and pes as in Tulerpeton. The flaggy sandstones with trackways include fossil
remains of trees (Archeopteris howitti, “Cordaites” australis) and other plants
(Barinophyton citrulliforme, “Sphenopteris” carnei), as well as desiccation cracks, as in
alluvial soils (Entisols). Associated placoderm and sarcopterygian fish are Frasnian
(Young 2007; Young et al. 2010).
Scat Craig, Scotland. This locality in the Edenkillie Beds near Elgin, Scotland is
well known for fossil fish (acanthodian, heterostracan, placoderm, sarcopterygian)
correlated with Baltic mid-upper Frasnian fish localities (Newman 2005; Blieck et al.
2007). Hugh Miller (1867, p.227) compared the Scat Craig locality with“ the loose
gravelly soil of an ancient graveyard”, and several observations support interpretation of
a woodland paleosol. The bone-bearing layer is mottled red and gray like drab-haloed
root traces, and has well mixed clay, sand and gravel, which would have been separated if
transported by water. Bones from Scat Craig are disarticulated and uncemented like bone
from paleosols at Hyner (Retallack et al. 2009). Disarticulated jaw elements of
Elginerpeton pancheni from Scat Craig are comparable with other Frasnian aquatic
tetrapods (Ahlberg 1995).
Velna-Ala, Latvia. Jaw fragments of Obruchevichthys gracilis are similar to those
of aquatic tetrapods such as Elginerpeton (Ahlberg 1995). These specimens from the
Lielvarde Member of the Ogre Formation along the Alava River near Velna-Ala, were
associated with disarticulated acanthodians, heterostracans, placoderms, and
sarcopterygians, of the upper Frasnian stage (correlated using fish with rhenana conodont
zone: Blieck et al. 2007). The fine-grained sandstones have gypsum cement in the lower
Retallack: Pennsylvania trees and tetrapods
4
part and calcareous cement in the upper part, and may have been deposited in a lagoon or
restricted marine bay (Blieck et al. 2007).
Novgorod, Russia. Jaw fragments of Obruchevichthys gracilis (Ahlberg 1995)
also may have come from the Hadsnezha beds along the Lovat River near Novgorod,
known for disarticulated fish considered upper Frasnian (equivalent to rhenana conodont
zone: Blieck et al. 2007). These fossils were collected in the 1860’s without detailed
locality or sedimentary records (Vorobyeva 1977; Kuršs 1992).
Plucki, Poland. Limestones at the Famennian-Frasnian boundary near the village
of Plucki, in the Holy Cross Mountains of Poland, include brachiopods and cephalopods
of a shallow marine environment. Black bituminous content of the limestones is evidence
of anoxia. Tetrapods are represented by rare shoulder girdle elements, but more details
are needed, because unillustrated and reported only in abstract (Szrek 2008).
Shixiagou, China. The Zhongning Formation in Shixiagou Gorge, near
Zhongning in Ningxia Hui, northwest China is the site of a tetrapod jaw fragment,
Sinostega pani, similar to Acanthostega (Zhu et al. 2002). The jaw was found along with
fossil trees (Leptophloem rhombicum, Sublepidodendron mirabile), and FamennianFrasnian placoderms and sarcopterygians, as well as a probable Frasnian galeaspid and
spores (Blieck et al. 2007). Observations pertinent to paleoenvironments are unavailable.
Canowindra, New South Wales. This mass kill of hundreds of articulated fish
(sarcopterygians, dipnoans, and placoderms) is preserved as sandstone casts within a
trough-cross bedded quartz sandstone with red claystone-breccia, in the Mandagery
Formation 7 km west of Canowindra, New South Wales, Australia (Johanson and
Ahlberg 2001; Johanson et al. 2003). The carcasses are preserved crammed together as if
crowded by a drying billabong (oxbow lake) of a freshwater stream (Johanson 1998). No
marine fossils are found at Canowindra, but horseshoe crabs (Kasibelinurus amicorum)
from the formation near Parkes (Pickett 1993) and conodonts and brachiopods from
correlative formations near Bathurst (Jones and Turner 2000), indicate a seashore to the
north and east, and age near the Frasnian-Famennian boundary.
Bunduburrah, New South Wales. A single complete jaw of Metaxygnathus
denticulus (Campbell and Bell 1977) is similar to those of Acanthostega and Ichthyostega
(aquatic tetrapods of Long and Gordon 2004). The locality in the Cloghnan Shale near
Bunduburrah homestead, south of Jemalong Gap and west of Forbes, New South Wales,
Australia, is stratigraphically above the Canowindra locality, and has a variety of
placoderms, sarcoptyerygians and the Famennian lungfish Soederberghia (Young 2007).
The disarticulated tetrapod jaw is from a bone breccia directly overlying a deep-calcic,
red paleosol with woody root traces and caliche nodules. Trough cross-bedded sandstone
paleochannels indicate a fluvial floodplain environment, and root traces and a paleosol
comparable with the Hyner pedotype of Pennsylvania are evidence of dry woodland.
Gornostayevka, Russia. The Gornostayevka industrial quarry, southwest of
Livny, in the Oryol region includes a lag of fragmentary fish bone with Jacubsonia
livnensis, a tetrapod similar to Acanthostega, from the Zadonskian Horizont (Lebedev
2004). Also found are lungfish, sarcopterygians, acanthodians, placoderms of the lower
Famennian stage (probably equivalent to crepida conodont zone). The coarse-grained
sandstone and limestones above and below the fish-bearing layer include rhynchonellid
brachiopods of a marine deltaic environment (Blieck et al. 2007).
Retallack: Pennsylvania trees and tetrapods
5
Ketleri and Pavāri, Latvia. Friable sand of the Pavāri Member of the Ketleri
Formation on the Ciecere River near the village of Pavāri has yielded the type specimen
of Ventastega curonica, a creature generally similar to Acanthostega (Ahlberg et al.
1994). Other remains of Ventastega come from the overlying Varkali Member of the
Ketleri Formation along the Venta River, near Ketleri village, 12 km southwest (Blieck et
al. 2007; Ahlberg et al. 2008). Both assemblages have similar sarcopterygians,
acanthodians, placoderms, and rare actinopterygians, but lack sharks and ptyctodonts.
Conodonts of the mid-Famennian marginifera to postera zones are found in the
underlying Zagare Formation, but these are separated by disconformity from the fish
beds, which contain Famennian fish and spores but no definitive marine fossils. The
Pavāri fish are fragments in sandstones with large scale cross bedding dipping at up to
37o, mud chip breccias, ripple marks, and soft-sediment slump structures, underlying
thick red-green claystones (paleosols?). The Ketleri fish are a bone lag in parallel crossbedded greenish-gray conglomeratic sandstone above the red-green claystones
(Lukševičs and Zupiņš 2004). Both sites appear to be deposits of coastal streams,
probably estuarine (Ahlberg et al. 2008).
Strud, Belgium. A jaw previously identified as a fish from a part of the Evieux
Formation with mid-upper Famennian spores (GF zone), now appears to have been a
tetrapod similar to Ichthyostega (Clément et al. 2004). It is from granule sandstone with
clasts of redeposited shale and paleosol, interpreted as a fluvial paleochannel by Blieck et
al. (2007).
Hyner, PA. A roadcut of Duncannon Member of the Catskill Formation north of
state route 120, 1 mile west of Hyner (N41.346183o W77.6800o) in Clinton County, has
yielded a jaw (Densignathus rowei), another jaw, cleithrum and scapulocoracoid
(Hynerpeton bassetti), and humerus (Tetrapoda sp. indet.) generally comparable with
those of aquatic tetrapods such as Acanthostega and Ichthyostega (Daeschler et al. 1994;
Daeschler 2000; Shubin et al. 2004). This prominent red outcrop has been called “Red
Hill” (by Woodrow et al. 1995), and should not be confused with another Devonian fish
locality with that name in Nevada (Murphy et al. 1976; Reed 1985, 1986). Associated
with the tetrapods near Hyner are sarcopterygians, lungfish, sharks, placoderms,
acanthodians, and actinopterygians (Davis et al. 2001; Daeschler et al. 2003; Friedman
and Daeschler 2006), as well as arachnids (Shear 2000), millipedes (Wilson et al. 2005),
and a variety of fossil plants (Cressler 2006), including spores which date the deposit as
upper Famennian (VH zone: Traverse 2003). Red paleosols are interbedded with thick
sandstones of meandering streams (Woodrow et al. 1995). There are six distinct
pedotypes among the 42 successive paleosols exposed at Hyner (Retallack et al. 2009):
red deep-calcic (Hyner), red shallow-calcic (Bucktail), red-green mottled and thin with
slickensides and no nodules (Farwell), red-green mottled and thin with relict bedding
(Gleasonton), red and thin with relict bedding (Renovo), and dark grey carbonaceous root
traces (Sproul).
Tetrapod bones (Densignathus rowei and Hynerpeton basseti) were disarticulated
and weathered on seasonally-dry alluvial soils (Farwell pedotype) in association with
disarticulated sarcopterygians (Hyneria lindae, Sauripterus sp. indet.), sharks (Ageleodus
pectinatus), placoderms (Groenlandaspis pennsylvanica, Turrisaspis elektior, Phyllolepis
rossimontana), actinopterygians (Limnomis delaneyi), and remains of trees
(Archaeopteris halliana) and aquatic plants (Otzinachsonia beerboweri). Farwell
Retallack: Pennsylvania trees and tetrapods
6
paleosols offer limited evidence of paleoclimate, but include tree fossils and are in a part
of the sequence deposited during the subhumid woodland part of the paleoclimatic cycle
(with Hyner pedotypes). Other paleosols of semi-arid shrublands (Bucktail), permanently
waterlogged swamps (Sproul), and early successional habitats (Renovo, Gleasonton)
have not yet yielded tetrapods (Retallack et al. 2009).
Gauss Halvø and Ymer Ø, Greenland. Fluvial deposits with upper Famennian
spores (between GF and LL zones) and fish fossils in East Greenland have yielded
tetrapods at two stratigraphic levels: Ichthyostega stensioei (perhaps also including I.
eigili and I. watsoni) in the Aina Dal Formation, and Acanthostega gunnari and
Ichthyostega sp. indet. (broad-headed), in the stratigraphically higher Britta Dal
Formation (Blieck et al. 2007). These two formations formed under a wetter climate than
the intervening Wimans Bjerg Formation, with playa lake deposits. Acanthostega
skeletons were found in a lens of fine silty sand, interpreted as a swale of a river point
bar, whereas Ichthyostega was preserved as partly mummified carcasses in coarse red to
fine-grained blackish-red sandstone floodplain deposits (Clack 2002), perhaps alluvial
soils (Entisols). The short limbs, polydactyly and flattened tails of both taxa suggest
aquatic adaptations, more marked in the smaller Acanthostega than in the larger
Ichthyostega (Clack 1988; 2006).
Andreyevka, Russia. An articulated skeleton of Tulerpeton curtum found in the
Khovanshchina Beds of the Zavolzkh Formation on the right bank of the Tresna River,
south of Andreyevka, near Tula, Russia, has the geologically oldest known hinged wrists
and ankles (Lebedev and Coates 1995). The skeleton was in a thin stromatolitic limestone
with brackish fossils (ostracods, bivalves, serpulid worms) as well as freshwater remains
(charophyte gyrogonites and axes). The ostracods indicate upper Famennian stage
(equivalent to praesulcata conodont zone), and associated fossil fish (sarcopterygians,
lungfish, placoderms) are also Famennian (Blieck et al. 2007). Limestones with fish
fragments underlie the stromatolitic layer, and it is overlain by 10-cm-thick fish bone
bed, interpreted as a lagoonal deposit in a tropical limestone coast (Lebedev 1990, 1992).
Nescopeck, PA. Tetrapod footprints (Hylopus hardingi) from the basal Mauch
Chunk Formation, of Ivorian age (ca. 347 Ma), were found on a loose slab of reddish
brown sandstone with large root traces in the banks of Nescopeck Creek under an
overpass of interstate highway I80 (N41.007600o W76.166433o), near Hazleton, Luzerne
County (Vrazo et al. 2007). The slab was presumably eroded from the creek bed because
there is not exposure there now. The slab is comparable to weakely developed paleosols
(Tigue pedotype) found in the uppermost Pocono Formation at Scranton (Fig. 5D), along
with deep-calcic gray paleosols (Dunmore pedotype) and remains of both lycopsid
(Lepidodendropsis scrobiniformis) and pteridosperm (Triphyllopteris lescurianam, T.
biloba) trees (Edmunds 1997). The footprint tracks are very different lithologically from
red siltstones of the lower Mauch Chunk Formation with shallow-calcic (Nescopeck
pedotype) paleosols in the nearby roadcut at least 20 m stratigraphically higher on the
westbound freeway 200 m to the south (Driese et al. 1993). Subhumid wooded
environments are most likely for the Nescopeck footprints.
Horton Bluff, Nova Scotia. Trackways and disarticulated limb bones of several
kinds of tetrapods from the upper Tournaisian, Horton Bluff Formation of Nova Scotia,
Canada, represent largely aquatic tetrapods generally similar to Ichthyostega, Tulerpeton
and Pederpes (Carroll 2009). The associated fish fauna includes placoderms(?), sharks,
Retallack: Pennsylvania trees and tetrapods
7
lungfishes, sarcopterygians, acanthodians, and actinopterygians. These remains were
recovered from sandstone paleochannels and weakly developed silty paleosols (Entisols),
but other stratigraphic levels have stump casts, fossil plants and non-calcareous grey
paleosols of lycopsid forests (Gleyed Inceptisols: Rygel et al. 2006).
Auchenreoch Glen, Scotland. Pederpes finneyae is an articulated skeleton from a
carbonate nodule near Dumbarton in the Ballagan Formation, dated as Ivorian (Tn3c)
from palynology. This skeleton has the short body and digits of a terrestrial tetrapod.
Associated with the specimen were worm tubes (Spirorbis), ostracods and fish
(sarcopterygian and actinopterygian), suggestive of a coastal lagoon paleoenvironment
(Clack and Finney 2005).
Middle Paddock, Queensland. Ossinodus pueri is known from terrestrial tetrapod
limb bones and skull fragments in a green conglomeratic sandstone paleochannel of the
Ducabrook Formation, dated by palynology as mid-Visean (3a), in Middle Paddock, 80
km southwest of Emerald, Queensland, Australia. The bone bed has been interpreted as
an estuarine channel lag overlying oolitic limestones of a saline lagoon (Warren and
Turner 2004).
Wardie, Scotland. Lethiscus stocki is represented by a single articulated skeleton
with the elongate limbless body of an eel or snake (Wellstead 1982), in a mid-Holkierian
locality known for articulated sharks, sarcopterygians, acanthodians, and actinopterygians
(Dineley and Metcalfe 1999). The articulated skeletons are in siderite nodules within
black shale of the Wardie Shales, near Granton Harbour, Scotland. There are no marine
invertebrates, but hints of marine influence in the diversity of sharks, and high organic
content may have come from coastal lake anoxia (Parnell 1998).
Delta, Iowa.A partly articulated skeleton of the terrestrial tetrapod Whatcheria
deltae is named for the nearby towns of What Cheer and Delta, in Iowa, USA (Lombard
and Bolt 1995). Along with another terrestrial amphibian (embolomore) and an aquatic
amphibian (colosteid), sharks, lungfish, acanthodians, sarcopterygians, and
actinopterygians, these remains are in a 50 cm thick bed of brownish green shale with
clasts of limestone like those of the underlying talus infill of a paleokarst depression in
the St Louis Limestone. This bed includes large root molds, ostracods and spirorbid
snails suggestive of a lagoon-margin paleosol, here interpreted as an Aquept. The age of
the paleosol is bracketed by enclosing marine rocks as upper Meramecian (Bolt et al.
1988), roughly upper Holkerian (Gradstein et al. 1984).
Rock Canyon, Nevada. A partial skeleton of a tetrapod (Antlerpeton clarkii) from
the upper Meramecian-lower Chesterian (Holkieran-Asbian), Diamond Peak Formation
in Rock Canyon, Eureka County, has robust pelvic girdle and sacral vertebrae of a
terrestrial animal (Thomson et al. 1998). It was found in non-marine red sandstone, but
observations pertinent to paleoenvironment are not available.
Broxburn, Scotland. The lower Asbian (NM spore zone) “Curly Shale”
(torbanite) of the Pumpherston Shale has yielded an articulated skeleton of
Palaeomolgophis scoticus (Andrews and Carroll 1991). These torbanites were produced
by freshwater algal blooms, and contain no marine fossils, other than ostracods. The sea
was not far away, judging from marine fossils in the Pumpherston Shell Bed lower in the
sequence (Parnell 1988).
Pentland, Scotland. The lower Asbian (NM spore zone) Dunnet Shale in the
Pentland Oil Works, near Edinburgh, has yielded a partial articulated skeleton of
Retallack: Pennsylvania trees and tetrapods
8
Aldelogyrinus simorhynchus, with a long back, short tail and limbs (Andrews and Carroll
1991), and a fish fauna including fossil sharks (Dick 1978). These oil shales were
deposited in freshwater lakes (Parnell 1988).
Straiton, Scotland. The lower Asbian (NM spore zone) Dunnet Shale at Straiton,
near Edinburgh, has yielded a headless articulated skeleton of an adelogyrinid, generally
comparable with the skeleton from Pentland (Andrews and Carroll 1991). These oil
shales were deposited in freshwater lakes (Parnell 1988).
Pitcorthie, Scotland. The upper Asbian, Anstruther Beds at Pitcorthie near
Abstruther in Fifeshire has yielded a variety of fossil fish (acanthodians, sarcopterygians,
actinopterygians, and sharks) and a skull table comparable with Dolichopareias disjectus
(Andrews and Carroll 1991). Also found were ostracods, and spores and cones of forest
plants (Lepidostrobus), but no definitive marine fossils. The Abstruther Beds probably
formed in a coastal lake (Love 1959; Wattison 1962).
Burdiehouse, Scotland. The upper Asbian Burdiehouse Limestone at
Burdiehouse has yielded an incomplete skull of Dolichopareias disjectus (Andrews and
Carroll 1991), as well as fossil plants (Sphenopteris, Lepidodendron), ostracods (Cypris
scotoburdigalensis), bivalves (“Unio” nuciformis), worm tubes (Spirorbis) and fish
(acanthodians, sarcopterygians, actinopterygians and sharks: Wattison 1962). Their
paleoenvironment has been interpreted as freshwater coastal lake (Love 1959).
Pottsville, PA. A variety of fossil footprints (Palaeosauropus primaevus,
Batrachichnus sp indet.) and body impressions confirming lack of thick scales have been
found in the middle Mauch Chunk Formation, 60 feet south from the southwest corner of
Center and Main Streets in Mount Carbon (N40.67444o W76.19111o), south of Pottsville,
Schuylkill County, and equivalent horizons on the other side of the valley (Lea 1849,
1855; Simpson et al. 2006; Lucas et al. 2007; Vrazo et al. 2007). These remains come
from green-gray, fine sandstones and siltstones, with mudcracks, raindrop impressions
(Fillmore et al. 2006), woody root traces (Carbon pedotype) and leaves of trees
(Adiantites antiquus: Jennings 1985), flanked by red paleosols with moderately deepcalcic horizons at the eastern end of Tumbling Run, Pottsville (at 840 m). The following
age model for this section (318.3 Ma top chron MMCn at 1094.8 m, 319.4 Ma base
MMCn at 1067.8 m, 320.3 Ma top chron MMBn at 1023.8 m, 323.8 Ma base chron
MMBn at 912.8 m, 328.8 Ma top chron MMA at 815.8 m (Opdyke and DiVenere 2004),
gives an age of these footprints as 328 Ma (Brigantian).
Cheese Bay, Scotland. Casineria kidderi from the upper Asbian of Cheese Bay,
Scotland, is known from an articulated small skeleton, found along with articulated
shrimp, hydroids and scorpions in laminated black shales of a freshwater to brackish
coastal lagoon. Long limbs and short back are indications that Casineria was a terrestrial
animal (Paton et al. 1999).
Gilmerton, Scotland. From a deep coal-mine at Gilmerton, near Edinburgh,
Scotland, the lower Brigantian, Gilmerton Ironstone has yielded two articulated
tetrapods. Pholidogaster pisciformis (Romer 1964) had limbs functional for terrestrial
walking, but Crassigyrinus scoticus had reduced limbs and a deep tail like an eel
(Panchen and Smithson 1990). These specimens were collected without observations
pertinent to reconstructing paleoenvironment.
East Kirkton, Scotland. A variety of articulated tetrapods have been found at East
Kirkton, near Bathgate, Scotland, in the lower Brigantian East Kirkton Limestone. The
Retallack: Pennsylvania trees and tetrapods
9
dark laminated limestones of this freshwater lake deposit have yielded fossil leaves and
wood, scorpions, myriapods, harvestmen, eurypterids and ostracods. Some of the
tetrapods such as Balanerpeton woodi, Silvanerpeton miripedes and Westlothiana lizziae
have the elongate bodies of aquatic creatures, but others such as Eldeeceon rolfei and
Eucritta melanolimnetes have the short back and strong limbs of terrestrial animals.
There are also limbless aistopods such as Ophiderpeton (Rolfe et al. 1994; Clack 2002).
Greer, West Virginia. A limestone quarry near Greer, 11 km southeast of
Morgantown, West Virginia, exposes a bone bed in the middle Chesterian (Pendleian:
Kammer and Lake 2001), Bickett Shale of the Bluefield Formation, Mauch Chunk Group
(Hotton 1970; Godfrey 1988). The Bickett Shale here is red and green mottled siltstones
and shales, with relict bedding, comparable with alluvial soils (Entisols). The shale lacks
marine fossils, but a coastal location is indicated by the overlying Reynolds Limestone
with brachiopods. The vertebrate fauna includes fragmentary fish (shark, acanthodian,
lungfish, sarcopterygian, actinopterygian) and both short-bodied (Proterogyrinus
scheelei, cf. Eoherpeton, cf. Eryops) and long bodied tetrapods (Greererpeton
burkemorani, cf. Crassigyrinus: Romer 1969; Godfrey 1989; Schultze and Bolt 1996).
Cowdenbeath, Scotland. Articulated specimens of Crassigyrinus scoticus have
the reduced limbs and elongate form of fully aquatic creatures, and were found in black
unbedded siltstone beneath the upper Pendleian, Lochgelly Blackband Ironstone in the
Dora opencast mine near Cowdenbeath. Other remains of acanthodian fish and terrestrial
tetrapods (Doragnathus woodi, Eoherpeton watsoni, Proterogyrinus pancheni) were
found in “a characteristic seat rock, a soft dark gray unbedded siltstone with rootlets”
(Panchen and Smithson 1990): here interpreted as a swamp soil (Histosol).
Loanhead, Scotland. A robust femur of Papposaurus traquairi, has been found
along with aquatic tetrapod skeletons (Spathicephalus mirus, Adelospondylus watsoni) in
the Rumbles Ironstone (“Loanhead no. 2 ironstone”) of the Pendleian stage (Panchen
1970). The laminated dark ironstones and fine-grained sandstones of these specimens
also include fossil fish, and are considered lacustrine (Andrews and Carroll 1991).
Fayetteville, Arkansas. Jaw fragments of Baphetes kansensis represent a large
terrestrial animal from the Baldwin Coal of the Bloyd Formation near Fayetteville,
Washington County, Arkansas, USA (Romer 1963). The coal includes a variety of
swamp forest plants, and overlies a gleyed paleosol (“underclay”of Simonds 1891). This
coal is Morrowan and early Pennsylvanian in age, or lower Namurian (Panchen 1970).
Green Creek, New Brunswick. A lower Namurian trackway (Pseudobradypus)
from the Enragé Formation, near Green Creek in the Maringouin Peninsula, New
Brunswick, Canada, was created by a terrestrial tetrapod. The footprints are at the top of a
10-12 cm thick brick-red mudrock, which caps the last fining upward cycle of this fluvial
formation (Wood and Miller 2007). This red mudrock is here interpreted as a well
drained, weakly developed, alluvial paleosol (Fluvent).
Tamaqua, PA. Footprints (Palaeosauropus sp., Vrazo et al. 2007) also have been
found only 20 m below the top of the Mauch Chunk Formation, in a large overgrown
outcrop adjacent to old railway bed, now a bicycle trail along the east side of state route
209, 1 mile south of Tamaqua (N40.782222o W76.96444o), Schuylkill County. They are
from fine-grained gray sandstone with sparse root traces as at Pottsville (Carbon
pedotype). A comparable stratigraphic level (1100 m) in the section at Pottsville has
Retallack: Pennsylvania trees and tetrapods
10
deep-calcic (woodland) paleosols (Henry pedotype), and using the paleomagnetic agemodel used above for Pottsville, can be dated at 319 Ma (late Serpukhovian).
Hawesville, Kentucky. Late Chesterian (lower Namurian), Menard Limestone in a
roadcut near Hawesville, Hancock County, Kentucky, has four separate facies with
vertebrate fossils. The marine Menard Limestone with shark teeth, contrasts with nonmarine overlying Palestine sandstone with fluvial sandstone, a red-green mottled paleosol
(Vertisol), and green-gray estuarine shale. Fragmentary sarcopterygian fish and
embolomere tetrapod bones were found the paleochannel sandstone, a variety of
terrestrial tetrapods (embolomere, watcheerid and temnospondyl) in the Vertisol paleosol,
and fish (sharks, sarcopterygians, acanthodians) and a partly articulated long-bodied
tetrapod (colosteid) in estuarine shales (Garcia et al. 2006).
Point Edward, Nova Scotia. The upper Chesterian (lower Namurian), Point
Edward Formation, on the north-eastern point of Sydney Harbour has yielded a variety of
vertebrates from “freshwater limestone and an overlying bed of red shale” (Carroll et al.
1972). The productive bed is gray-red mottled, with root traces, and micritic carbonate
nodules (like an Aridosol paleosol), and other beds show desiccation cracks like alluvial
soils (Entisols). Other horizons include remains of trees (Lepidodendron, Stigmaria),
freshwater bivalves (Carbonicola) and conchostracans (Leaia), and brackish ostracods
(Carbonita) and worm tubes (Spirorbis), indicative of forests and coastal lagoons nearby.
The vertebrate fauna includes fish (acanthodians, lungfish, sarcopterygians,
actinopterygians), and both short-bodied (“Pholiderpeton” bretonense) and long-bodied,
dorsoventrally flattened tetrapods (Spathicephalus pereger: Carroll et al. 1972).
Goreville, Illinois. Red sandy mudstones within gray mudstones of the lower
Morrowan stage (mid-Namurian) have yielded lungfish, terrestrial amphibians
(microsaur, embolomere) and aquatic (Greererpeton) tetrapods. This locality is a
sequence of well drained, moderately developed paleosols (Inceptisols) with root traces
of woody vegetation (Shultze and Bolt 1996).
Lehi, Utah. Utaherpeton franklini is an articulated skeleton of an aquatic tetrapod
in the lower Morrowan, Manning Canyon Shale. These lacustrine shales have been
quarried extensively for fossil plants and a megasecopteran insect of Namurian B age
(Carroll et al. 1991). No marine fossils have been found in the shales, but marine
limestones are interbedded in this cyclothemic sequence, which also includes purplegreen Inceptisols with root traces of woody plants.
Wuppertal, Germany. Gray shales with Namurian ammonoids Reticuloceras
bilingue (Namurian zone R2e), also include root traces and leaves of calamites, cordaites
and tree lycopsids are exposed in Ziegelei quarry on Schmiederstrasse in Hasslinghausen,
near Wuppertal, Germany. These coastal lagoonal deposits fringed by mangal and marsh
also yielded an articulated skeleton of the terrestrial anthracosaur Bruktererpeton fiebigi
(Boy and Bandel 1973).
Cape Linzee, Nova Scotia. TheYeadonian-Langsettian Port Hood Formation at
Cape Linzee has yielded an articulated tetrapod (Romeriscus “periallus”; Baird and
Carroll 1967; Laurin and Reisz 1992) from a 1.2 m thick paleochannel sandstone between
coal seams (Histosol paleosols). Scales of sarcopterygian fish (Rhizodopsis) are found in
other sandstones at the same locality. Other sandstone beds 250 m above this sandstone
have yielded a variety of pentadactyl trackways, made by terrestrial tetrapods on moist
Retallack: Pennsylvania trees and tetrapods
11
floodplain clay (Entisol). Some of these creatures were large, with feet some 12 cm long
(Keighley and Pickerill 1998).
Grand Anse, New Brunswick. Fossil footprints (cf. Pseudobradypus
longidactylus) from the Grand Anse Formation at Shepody Bay near Grand Anse are of
the Langsettian (upper Namurian) stage (Keighley et al. 2008). They are found in finegrained brown sandstones of riparian alluvial soils (Entisols), interbedded with red,
clayey, paleosols, 1 m thick (Falcon-Lang et al. 2007). These thick red paleosols
(Alfisols) have large, near-vertical, drab-haloed, root traces, similar to well drained
woodland paleosols (Retallack 1985, 1997a).
Jarrow, Ireland. Coal from the Jarrow Colliery of the Leinster Coalfield is
Langsettian (communis zone), and associated shales have been interpreted as deposits of
an oxbow lake (Hook and Ferm 1985). These shales have yielded a disarticulated skull of
a terrestrial tetrapod (Procochleosaurus jarrowensis Sequeira 1996), as well as other
tetrapods of uncertain affinities (Panchen 1970).
Joggins, Nova Scotia. Articulated skeletons of the terrestrial eutetrapod
Dendrerpeton acadianum (Holmes et al.1998) and a vertebral column of the reptile-like
Callignethodon are known from siderite nodules in the roof shales of a fossil Histosol
paleosol in the early Pennsylvanian (Langsettian) Joggins Formation of Nova Scotia.
Also found in this sequence were millipedes, arachnids, insects, ostracods, pterioid
bivalves and actinopterygian fish scales (Falcon-Lang et al. 2006). Fragmentary remains
of a variety of eutetrapods (Dendrerpeton acadianum, D. confusum, Callignethelon
watsoni, Asaphestera intermedium, Hylerpeton dawsoni, Leiocephalikon problematicum,
Ricnodon sp., Hylonomus lyelli, Archerpeton anthracos, Protoclepsydrops haplous,
Novascoticus multidens) are found within sandstone casts of trees at this locality (Milner
1996), from creatures trapped within dens during a forest fire. Other surfaces with
tetrapod footprints also are rooted with stump casts and were probably alluvial soils
(Fluvents, Calder et al. 2006).
REFERENCES CITED
Ahlberg, P.E.1995. Elginerpeton pancheni and the earliest tetrapod clade. Nature
373:420-475.
Ahlberg, P.E.; Lukševičs, E.; and Lebedev, O.A. 1994. The first tetrapod finds from the
Devonian (upper Famennian) of Latvia. Roy. Soc.London Phil. Trans. B343:303328.
Ahlberg, P.E.; Lukševičs, E.; and Mark-Kurik, E. 2000. A near-tetrapod from the Baltic
Middle Devonian. Paleontology 43:533-548.
Ahlberg, P.E.; Clack, J.A.; Lukševičs, E.; Blom, H.; and Zupiņš, I. 2008. Ventastega
curonica and the origin of tetrapod morphology. Nature 453:1199-1204.
Andrews, S.M.; and Carroll, R.L. 1991. The Order Adelospondyli: Carboniferous
lepospondyl amphibians: Roy. Soc. Edinburgh Trans. 82:239-275.
Baird, D.; and Carroll, R.L. 1967. Romeriscus, the oldest known reptile. Science 157:5659.
Banks, H.P.; Grierson, J.D.; and Bonamo, P.M. 1985. The flora of the Catskill clastic
wedge. In Woodrow, D.L., and Sevon, W.D., eds., The Catskill Delta, Geol. Soc.
America Spec. Pap. 201:125-141.
Retallack: Pennsylvania trees and tetrapods
12
Bartels, C., Briggs, D.E.G., and Brassel, G. 1998. The fossils of the Hunsrück Slate:
marine life in the Devonian. Cambridge, Cambridge University Press, 309 p.
Blieck, A.; Clement, G.; Blom, H.; Lelievre, H.; Luksevics, E.; Streel, M.; Thorez, J.; and
Young, G.C. 2007. The biostratigraphical and palaeogeographical framework of
the earliest diversification of tetrapods (Late Devonian). In Becker, R.T., and
Kirchgasser, W.T., eds., Devonian events and correlation. Geol. Soc. London
Spec. Publ. 278:219-235.
Boisvert, C.A. 2005. The pelvic fin and girdle of Panderichthys and the origin of tetrapod
locomotion. Nature 438:1145-1147.
Bolt, J.R.; McKay, R.M.; Witzke, B.J.; and McAdams, M.P. 1988. A new Lower
Carboniferous tetrapod locality in Iowa. Nature 333:768-770.
Boy, J.A.; and Bandel, K. 1973. Bruktererpeton fiebigi n. gen. n. sp. (Amphibia,
Gephyrostegida) der erste Tetrapode aus dem Rheinisch-Westfälischen Karbon
(Namur B; W.-Deutschland). Palaeontographica A145:39-77.
Calder, J.H.; Gibling, M.R.; Scott, A.C.; Davies, S.J.; and Hebert, B.L. 2006. A fossil
lycopsid forest succession in the classical Joggins section of Nova Scotia.:
paleoecology of a disturbance-prone Pennsylvanian wetland In Greb, S.F.,
DiMichele, W.A., and Gastaldo, R.A., eds., Wetlands through time. Geol. Soc.
America Spec. Pap. 399:169-195.
Campbell, K.S.W.; and Bell, M.W. 1977. A primitive amphibian from the late Devonian
of New South Wales. Alcheringa 1:369-381.
Carroll, R.L. 2009. The rise of amphibians: 365 million years of evolution. Baltimore,
Johns Hopkins University Press, 360 p.
Carroll, R.L.; Belt, E.S.; Dineley, D.L.; Baird, D.; and McGregor, D.C. 1972. Vertebrate
paleontology of eastern Canada. 24th Int. Geol. Congr. Montreal, Excursion
A59:1-113.
Carroll, R.L.; Bybee, P.; and Tidwell, W.D. 1991. The oldest microsaur (Amphibia). J.
Paleont. 65:314-322.
Clack, J.A. 1988. New material of Acanthostega from the Devonian of East Greenland.
Palaeontology 31:699-724.
Clack, J.A. 2002. Gaining ground: the origin and evolution of tetrapods. Bloomington,
Indiana University Press, 369 p.
Clack, J.A. 2006. The emergence of early tetrapods. Palaeogeogr. Palaeoclim. Palaeoec.
232:167-189.
Clack, J.A.; and Finney, S.M. 2005. Pederpes finneyae, an articulated tetrapod from the
Tournaisian of western Scotland. J. Syst. Palaeont. 2:311-346.
Clément, G.; Ahlberg, P.E.; Blieck, A.; Blom, H.; Clack, J.A.; Poty, E.; Thorez, J.; and
Janvier, P. 2004. Devonian tetrapod from western Europe. Nature 427:412-413.
Cressler, W.L. 2006. Plant paleoecology of the Late Devonian Red Hill locality, northcentral Pennsylvania, an Archaeopteris-dominated community and early tetrapod
site. In Greb, S.F., DiMichele, W.A., and Gastaldo, R.A., eds., Wetlands through
time. Geol. Soc. America Spec. Pap. 399:79-102.
Daeschler, E.B. 2000. Early tetrapod jaws from the Late Devonian of Pennsylvania,
USA. J. Paleont. 74:301-308.
Daeschler, E.B.; Cressler, W.; Thomson, K.S.; and Amaral, W.W. 1994. A Devonian
tetrapod from North America. Science 265:639-642.
Retallack: Pennsylvania trees and tetrapods
13
Daeschler, E.B.; Frumes, A.C.; and Mullison, F. 2003. Groenlandaspid placoderm fish
from the Late Devonian of North America. Austral. Mus. Rec. 55:45-60.
Daeschler, E.B.; Shubin, N.H.; and Jenkins, F.A. 2006. A Devonian tetrapod-like fish and
the evolution of the tetrapod body plan. Nature 444:753-757.
Davis, M.C.; Shubin, N.H.; and Daeschler, E.B. 2001. Immature rhizodontids from the
Devonian of North America. Mus. Comp. Zool. Bull. 156:171-187.
Dick, J.R.F. 1978. On the Carboniferous shark Tristychius arcuatus Agassiz from
Scotland. Roy. Soc.Edinburgh Trans. 70:1-108.
Dineley, D.J.; and Metcalfe, S.J. 1999. Fossil fishes of Great Britain. Peterborough, Joint
Nature Conservation Committee, 675 p.
Elliott, D.K.; and Johnson, H.G. 1997. Use of vertebrates to solve biostratigraphic
problems; examples from the Lower and Middle Devonian of western North
America. In Klapper, G., Murphy, M.A., and Talent, J.A., eds., Paleozoic
sequence stratigraphy, biostratigraphy, and biogeography; studies in honor of J.
Granville ('Jess') Johnson. Geol. Soc. America Spec. Pap. 321:179-188
Falcon-Lang, H.J.; Benton, M.J.; Braddy, S.J.; and Davies, S.J. 2006. The Pennsylvanian
tropical biome reconstructed from the Joggins Formation of Nova Scotia, Canada.
Geol. Soc. London J. 163:561-576.
Falcon-Lang, H.J.; Benton, M.J.; and Stimson, M. 2007. Ecology of early reptiles
inferred from Lower Pennsylvanian trackways. Geol. Soc. London J.164, 11131118.
Fillmore, D.L.; Simpson, D.G.; and Lucas, S.G. 2006. An investigation of the
Palaeosauropus primaevus type locality: an ichnofossil assemblage from the
Mississippian Mauch Chunk Formation, Schuylkill County, Pennsylvania: Geol.
Soc. America Abstr. 38(7):436.
Friedman, M.; and Daeschler, E.B. 2006. Late Devonian (Famennian) lungfishes from
the Catskill Formation of Pennsylvania, USA. Palaeontology 49:1167-1183.
Friend, P.F.; and Williams, B.P.J. 1978. A field guide to outcrop areas of the Devonian of
Scotland, the Welsh Borderland and South Wales. London, Palaeontological
Association, 106 p.
Garcia, W.J.; Storrs, G.W.; and Greb, S.F. 2006. The Hancock County tretrapod locality:
a new Mississippian (Chesterian) wetlands fauna from western Kentucky (USA).
In Greb, S.F., DiMichele, W.A., and Gastaldo, R.A., eds., Wetlands through time.
Geol. Soc. America Spec. Pap. 399:155-167.
Godfrey, S.J. 1988. Isolated tetrapod remains from the Carboniferous of West Virginia.
Kirtlandia 43:27-36.
Godfrey, S.J. 1989. Postcranial skeletal anatomy of the Carboniferous tetrapod
Greererpeton burkemorani Romer 1969. Roy. Soc. London Phil. Trans. B323:75133.
Goin, C.J.; and Goin, O.B. 1956. Further comments on the origin of the tetrapods.
Evolution 10:440-441.
Gradstein, F.M.; Ogg., J.G.; and Smith, A.G. 2004. A geologic time scale 2004.
Cambridge, Cambridge University Press, 589 p.
Holmes, R.; Carroll, R.L.; and Reisz, R.R. 1998. The first articulated skeleton of
Dendrerpeton acadianum (Temnospondyli, Dendrepetonidae) from the lower
Retallack: Pennsylvania trees and tetrapods
14
Pennsylvanian locality of Joggins, Nova Scotia, and a review of its relationships.
J. Verteb. Paleont.18:64-79.
Hook, R.W.; and Ferm, J.C. 1985. A depositional model for the Linton tetrapod
assemblages (Westphalian D, Upper Carboniferous) and its palaeoenvironmental
significance. Roy. Soc. London Phil. Trans. B311:101-109.
Hotton, N. 1970. Mauchchunkia bassa, gen. et sp. nov. an anthracosaur (Amphibia,
Labyrinthodonta) from the Upper Mississippian. Kirtlandia 12:1-38.
Jennings, J.R. 1985. Fossil plants from the Mauch Chunk Formation of Pennsylvania:
morphology of Adiantites antiquus. J. Paleont. 59:1146-1157.
Johanson, Z. 1998. The Upper Devonian fish Bothriolepis (Placodermi, Antiarchi) from
near Canowindra, New South Wales, Australia. Austral. Mus. Rec. 50:315-348.
Johanson, Z. and Ahlberg, P.E. 2001. Devonian rhizodontids and tristichopterids
(Sarcopterygii, Tetrapodomorpha) from East Gondwana. Roy. Soc. Edinburgh
Trans. Earth Sci. 92:43-74.
Johanson, Z.; Ahlberg, P.E.; and Ritchie, A. 2003. The braincase and palate of the
tetrapodomorph sarcopterygian Mandageria fairfaxi; morphological variability
near the fish-tetrapod transition. Palaeontology 46:271-293.
Jones, R.K.; and Turner, S. 2000. Late Devonian fauna from the Columbine Sandstone
(Coffee Hill Member), Gap Creek, central new South Wales. Cour. Forsch.
Senck. 223:523-531.
Kammer, T.W.; and Lake, A.M. 2001. Salinity ranges of Late Mississippian invertebrates
of the central Appalachian Basin. Southeast. Geol. 40:99-116.
Keighley, D.G.; and Pickerill, R.K. 1998. Systematic ichnology of the Mabou and
Cumberland groups (Carboniferous) of western Cape Breton Island, Eastern
Canada; 2, Surface markings. Atlantic Geol. 34:83-112.
Keighley, D.; Calder, J.; Park, A.; Pickerill, R.; and Waldron, J. 2008. Discussion on
ecology of earliest reptiles inferred from basal Pennsylvanian trackways. Geol.
Soc. London J. 165:983-987.
Kursš, V. 1992. Depositional environment and burial conditions of fish remains in Baltic
Middle Devonian. In Mark-Kurik, E., ed., Fossil fishes as living systems. Talinn,
Estonian Academy of Science, p. 251-260.
Laurin, M.; and Reisz, R.R. 1992. A reassessment of the Pennsylvanian tetrapod
Romeriscus. J. Verteb. Paleont. 12:524-527.
Lea, I. 1849. On the reptilian footprints found in the gorge of Sharp Mountain near
Pottsville, Pa. Amer. Phil. Soc. Proc. 5:91-94.
Lea, I. 1855. Fossil footmarks in the red sandstone of Pottsville. Philadelphia, Collins, 16
p.
Lebedev, O. 1990. Tulerpeton, l’animal a six droits. La Recherche 21:1274-1275.
Lebedev. O. 1992. The latest Devonian, Khovanian vertebrate assemblage of
Andreyevka-2 locality, Tula region, Russia. In Mark-Kurik, E., ed., Fossil fishes
as living systems. Talinn, Estonian Academy of Science, p. 265-272.
Lebedev, O.L. 2004. A new tetrapod Jakubsonia livnensis from the Early Famennian
(Devonian) of Russia and palaeoecological remarks on the Late Devonian
tetrapod habitats. Acta Universitatis Latviensis 679:79-98.
Retallack: Pennsylvania trees and tetrapods
15
Lebedev, O.A.; and Coates, M.I. 1995. The postcranial skeleton of Devonian tetrapod
Tulerpeton curtum Lebedev. Zool. J. Linnaean Soc. 114:307-348.
Lombard, R.E.; and Bolt, J.R. 1995. A new primitive tetrapod, Whatcheeria deltae, from
the Lower Carboniferous of Iowa. Palaeontology 38:471-194.
Long, J.A. 1991. The long history of Australian fossil fishes. In Vickers-Rich,
Monaghan, J.M., Baird, R.F., and Rich, T.H., eds., Vertebrate palaeontology in
Australasia. Melbourne, Monash University Press, p.336-428.
Long, J.A.; and Gordon, M.S. 2004. The greatest step in vertebrate history, a
paleobiological review of the fish-tetrapod transition. Physiol. Biochem. Zool.
77:700-719.
Long, J.A.; Young, G.C.; Holland, T.; Senden, T.J.; and Fitzgerald, E.M.G. 2006. An
exceptional Devonian fish from Australia sheds light on tetrapod origins. Nature
444:199-202.
Long, J.A.; Choo, B.; and Young, G.C. 2008. A new basal actinopterygian fish from the
Middle Devonian Aztec Siltstone of Antarctica. Antarctic Science 20:393-412.
Love, L.G. 1959. Assemblages of small spores from the lower Oil Shale Group of
Scotland. Roy. Soc.Edinburgh Proc. 47:99-126.
Lucas, S.G.; Fillmore, D.L.; and Simpson, E.L. 2007. Amphibian body impressions from
the Mississippian Mauch Chunk Formation, eastern Pennsylvania. Geol. Soc.
America Abstr. 39(6):400.
Lukševičs, E.; and Zupiņš, I. 2004. Sedimentology, fauna, and taphonomy of the Pavāri
site, Late Devonian of Latvia. Acta Universitatis Latviensis 679:99-119.
Marshall, J.E.A.; Astin, T.R.; Brown, J.F.; Mark-Kurik, E.; and Lazauskiene, J. 2007.
Recognizing the Kačák Event in the Devonian terrestrial environment and its
implications for understanding land-sea interactions. In Becker, R.T., and
Kirchgasser, W.T., eds., Devonian events and correlations. Geol. Soc. London
Spec. Publ. 278:133-155.
McPherson, J.G. 1980. Genesis of variegated redbeds in the fluvial Aztec Siltstone (Late
Devonian), southern Victoria Land, Antarctica. Sediment. Geol. 27:119-142.
Miller, H. 1867. The cruise of the Betsey. Boston, Gould and Lincoln, 524 p.
Miller, J.H.; Shubin, N.; Daeschler, E.B.; and Downs, J.P. 2007. Stratigraphic context of
Tiktaalik rosae (Late Devonian): paleoenvironment of the fish tetrapod transition.
Geol. Soc. America Abstr. 39(6):417.
Milner, A.R. 1996. A revision of the temnospondyl amphibians from the Upper
Carboniferous of Joggins, Nova Scotia. In Milner, A.R., ed., Studies on
Carboniferous and Permian Vertebrates. Spec. Pap. Palaeontology 52:81-103.
Mintz, J.S.; Driese, S.G.; and White, J.D. 2010. Environmental and ecological variability
of Middle Devonian (Givetian) forests in Appalachian Basin paleosols, New
York, United States. Palaios 25:85-96.
Murphy, M.A.; Morgan, T.G.; and Dineley, D.L. 1976. Asterolepis sp. from the upper
Devonian of central Nevada. J. Paleont. 50:467-471.
Narkiewicz, K.; and Narkiewicz, M. 2010. Mid-Devonian carbonate platform
development in the Holy Cross Mts ara (central Poland): new constraints from the
conodont bipennatus fauna. N. Jb. Geol. Paläont. Abh, 255:287-300.
Retallack: Pennsylvania trees and tetrapods
16
Niedźwiedzki, G.; Szrek, P.; Narkiewicz, K.; Narkiewicz, M.; and Ahlberg, P.E. 2010.
Tetrapod trackways from the early Middle Devonian period of Poland. Nature
463:43-48.
Newman, M.J. 2005. A systematic review of the placoderm genus Cosmacanthus, and a
description of acanthodian remains from the Upper Devonian of Scotland.
Palaeontology 48:1111-1116.
Opdyke, N.D., and DiVenere, V.J. 2004. The magnetic polarity stratigraphy of the Mauch
Chunk Formation, Pennsylvania. U.S. Nat. Acad. Sci. Proc.101:13423-13427.
Panchen, A.L. 1970. Encyclopedia of Paleoherpetology. Part 5A. Anthracosauria.
Stuttgart, Gustav Fischer, 84 p.
Panchen, A.L.; and Smithson, T.R. 1990. The pelvic girdle and hind limb of
Crassigyrinus scoticus (Lydekker) from the Scottish Carboniferous and the origin
of the tetrapod pelvic skeleton. Roy. Soc. Edinburgh Earth Sci. Trans. 81:31-44.
Parnell, J. 1988. Lacustrine petroleum source rocks in the Dinantian Oil Shale group,
Scotland: a review. In Fleet, A.J., Kelts, K., and Talbot, M.R., eds., Lacustrine
petroleum source rocks. Geol. Soc. London Spec. Publ. 40:235-246.
Paton, R.L.; Smithson, T.R.; and Clack, J.A. 1999. An amniote-like skeleton from the
Early Carboniferous of Scotland. Nature 398:508-513.
Pickett, J.W. 1993. A Late Devonian xiphosuran from near Parks, New South Wales. In
Jell, P.A., ed., Palaeontological studies in honour of Ken Campbell. Assoc.
Australas. Palaeont. Mem. 15:279-287.
Rayner, D.H. 1962. The Achanarras Limestone of the middle Old Red Sandstone,
Caithness, Scotland. Yorkshire Geol. Soc. Proc. 34:117-138.
Reed, J.W. 1985. Devonian dipnoans from Red Hill, Nevada. J. Paleont. 59:1181-1193.
Reed, J.W. 1986. The acanthodian genera Machaeracanthus and Persacanthus from the
Devonian of Red Hill, Nevada. Geobios 19: 409-419.
Retallack, G.J.; Hunt, R.R.; and White, T.E. 2009. Devonian palaeosols and tetrapod
habitats in Pennsylvania. Geol. Soc. London J. 166:1143-1156.
Rolfe, W.D.I.; Clarkson, E.N.K.; and Panchen, A.L., eds. 1994. Volcanism and early
terrestrial biota. Roy. Soc. Edinburgh Earth Sci. Trans. 84:1-464.
Romer, A.S. 1963. The larger embolomerous amphibians of the American Carboniferous.
Mus. Comp. Zool. Bull. 128:415-454.
Romer, A.S. 1964. The skeleton of the lower Carboniferous labyrinthodont Pholidogaster
pisciformis. Mus. Comp. Zool. Bull. 131:129-159.
Romer, A.S. 1969. A temnospondylous labyrinthodont from the Lower Carboniferous.
Kirtlandia 6:1-20.
Rygel, M.C.; Calder, J.H.; Gibling, M.R.; Gongras, M.K.; and Melrose, C.S.A. 2006.
Tournaisian forested wetlands in the Horton Group of Atlantic Canada. In Greb,
S.F., DiMichele, W.A., and Gastaldo, R.A., eds., Wetlands through time. Geol.
Soc. America Spec. Pap. 399:103-126.
Sarwar, G., and Friedman, G.M. 1995. Post-Devonian sediment cover over New York
State; evidence from fluid-inclusion, organic maturation, clay diagenesis and stable
isotope studies. Berlin, Springer, 113 p.
Schultz, C.H. 1999. The geology of Pennsylvania. Harrisburg, Geological Survey of
Pennsylvania, 888 p.
Retallack: Pennsylvania trees and tetrapods
17
Schultze, H.-P. 2005. The late Middle Devonian fish fauna of Red Hill Nevada, and its
paleobiogeographic connections. J. Verteb. Paleont. 25(suppl. 3):A111.
Schultze, H.-P.; and Bolt, J.R. 1996. The lungfish Tranodis and the tetrapod fauna from
the Upper Mississippian of North America. In Milner, A.R., ed., Studies on
Carboniferous and Permian vertebrates. Spec. Pap. Palaeontology 52:31-54.
Schultze, H.-P.; and Cloutier, R., ed. 1996. Devonian fishes and plants of Miguasha,
Quebec, Canada. Munich, Pfeil, 374 p.
Sequeira, S.E.K. 1996. A cochleosaurid amphibian from the Upper Carboniferous of
Ireland. In Milner, A.R., ed., Studies on Carboniferous and Permian Vertebrates.
Spec. Pap. Palaeontology 52:65-80.
Shear, W.A. 2000. Gigantocharinus szatmaryi, a new trigonotarbid arachnid from the
Late Devonian of North America (Chelicerata, Arachnida, Trigonotarbida). J.
Paleont. 74:25-31.
Shubin, N.H.; Daeschler, E.B.; and Coates, M.I. 2004. The early evolution of the tetrapod
humerus. Science 304:90-93.
Shubin, N.H.; Daeschler, E.B.; and Jenkins, F.A. 2006. The pectoral fin of Tiktaalik
roseae and the origin of the tetrapod limb. Nature 440:764 – 771.
Simonds, F.W. 1891. The geology of Washington County. Geol. Surv. Arkansas Ann.
Rept for 1888:1-153.
Simpson, E.L.; Lucas, S.G.; and Fillmore, D.L. 2006. Depositional facies of the tetrapod
footprint assemblage from the Mississippian Mauch Chunk Formation, Schuylkill
County, Pennsylvania. Geol. Soc. America Abstr. 38(2):64-65.
Stets, J., and Schäfer, A. 2009. The Siegenian delta: land-sea transitions at the northern
margin of the Rhenohercynian Basin. In Königshof, P., ed., Devonian Change:
Case Studies in Palaeogeography and Palaeoecology. Geol. Soc. Lond. Spec.
Publ. 314:37-72.
Stössel, I. 1995. The discovery of a new Devonian tetrapod trackway in SW Ireland.
Geol. Soc.London J.152:407-413.
Szrek, P. 2008. Vertebrates from the Upper Kellwasser Limestone, Frasnian-Famennian
boundary beds (Upper Devonian of the Holy Cross Mountains (Poland). J. Verteb.
Paleont. 28(suppl. 2):150A.
Tetlie, O.E. 2007. Like father, like son? Not amongst the eurypterids (Chelicerata) from
Beartooth Butte, Wyoming. J. Paleont. 81:1423-1431.
Thomson, K.S.; Shubin, N.S.; and Poole, F.G. 1998. A problematic early tetrapod from
the Mississippian of Nevada. J. Verteb. Paleont. 18:315-320.
Traverse, A. 2003. Dating the earliest tetrapods: a Catskill palynological problem in
Pennsylvania. Cour. Forsch. Senck. 241:19-29.
Upeniece, I., and Upenieks, J. 1992. Young Upper Devonian antiarch (Asterolepis)
individuals from the Lode quarry, Latvia. In Mark-Kurik, E., ed., Fossil fishes as
living systems. Talinn, Estonian Academy of Science, p. 167-176.
Vorobeyeva, E.I. 1977. Morfologiya i osobennosti evolyushii kistepernykh ryab
(Morphology and nature of the evolution of crossopterygian fishes). Trudy
Paleont. Inst. Akad. Nauk 163:1-238.
Vrazo, M.B.; Benton, M.J.; and Daeschler, E.B. 2007. Tetrapod tracks from the Mauch
Chunk Formation (middle to upper Mississippian) of Pennsylvania, U.S.A. Acad.
Nat. Sci. Philadelphia Proc.156:199-209.
Retallack: Pennsylvania trees and tetrapods
18
Warren, A.; and Turner, S. 2004. The first stem tetrapod from the Lower Carboniferous
of Gondwana. Palaeontology 47:151-184.
Wattison, A. 1962. Temporary exposures in the Calciferous Sandstone Measures of East
Fife. Edinburgh Geol. Soc. Trans. 19:133-138.
Wellstead, C.F. 1982. A Lower Carboniferous aïstopod amphibian from Scotland.
Palaeontology 25:193-208.
Wilson, H.M.; Daeschler, E.B.; and Desbiens, S. 2005. New flat-backed Archipolypoda
millipedes from the Upper Devonian of North America. J. Paleont. 79:737-743.
Wood, N.S.; and Miller, R.F. 2007. A Mississippian trackway (Pseudobradypus ichnsp)
from the Enragé Formation, New Brunswick, Canada. Atlantic Geol. 43:80-86.
Woodrow, D.L.; Robinson, R.A.J.; Prave, A.R.; Traverse, A.; Daeschler, E.B.; Rowe,
N.D.; and Delaney, N.A. 1995. Stratigraphic, sedimentologic and temporal
framework of Red Hill (Upper Devonian Catskill Formation), near Hyner, Clinton
County, Pennsylvania, site of the oldest amphibian known from North America.
Ann. Field Conf. Pennsylvania Geologists Gdbk 60:1-8.
Young, G.C. 2007. Devonian formations, vertebrate faunas and age control on the far
south coast of New South Wales and adjacent Victoria. Austral. J. Earth
Sci.54:991-1008.
Young, G.C.; Burrow, C.J.; Long, J.A.; Turner, S.; and Choo, B. 2010. Devonian
macrovertebrate assemblages and biogeography of east Gondwana (Australasia,
Antarctica). palaeoworld 19:55-74.
Zhu, M.; Ahlberg, P.E.; Zhao, W.; and Jia, L. 2002. First Devonian tetrapod from Asia.
Nature 420:760.
Zimmer, C. 1998. At the water’s edge. New York, Free Press, 290 p.