ArticlePDF Available



Abstract and Figures

The Kinney Brick Quarry, located in the Manzanita Mountains of central New Mexico, is a world-famous fossil locality in deposits of a marine embayment of Late Pennsylvanian age. The quantity and quality of fossil preservation identify Kinney as a Konservat Lagerstätte. This volume presents the results of recent research on the Kinney rocks and fossils, as well as new research based on older collections. Here, we provide a review of previous work and of the context within which to understand the Kinney Quarry Lagerstätte and the articles in this volume. A new look at Kinney, the environment, the animals, plants, and ichnofauna preserved there, was initiated by a controlled excavation carried out in 2014. This excavation revealed additional, more refined information about the sedimentology of the Kinney deposits, and additional information about the distribution of organisms during the period of accumulation.
Content may be subject to copyright.
Lucas, S. G., DiMichele, W. A. and Allen, B. D., eds., 2021, Kinney Brick Quarry Lagerstätte. New Mexico Museum of Natural History and Science Bulletin 84.
1New Mexico Museum of Natural History, 1801 Mountain Road N.W., Albuquerque, New Mexico 87104; email:;
2Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560;
3New Mexico Bureau of Geology and Mineral Resources, 801 Leroy Place, Socorro, New Mexico 87801
Abstract—The Kinney Brick Quarry, located in the Manzanita Mountains of central New Mexico, is a
world-famous fossil locality in deposits of a marine embayment of Late Pennsylvanian age. The quantity
and quality of fossil preservation identify Kinney as a Konservat Lagerstätte. This volume presents
the results of recent research on the Kinney rocks and fossils, as well as new research based on older
collections. Here, we provide a review of previous work and of the context within which to understand
the Kinney Quarry Lagerstätte and the articles in this volume. A new look at Kinney, the environment,
the animals, plants, and ichnofauna preserved there, was initiated by a controlled excavation carried
out in 2014. This excavation revealed additional, more rened information about the sedimentology of
the Kinney deposits, and additional information about the distribution of organisms during the period
of accumulation.
The Kinney Brick Quarry (Figs. 1-3), located in the
Manzanita Mountains of central New Mexico, is a clay pit
actively mined for the making of bricks at the Kinney Brick
Company plant in Albuquerque, New Mexico. The quarry is
also a world-famous fossil locality. These fossils come from
deposits of a marine embayment of Late Pennsylvanian age
and are remarkable for their diversity, abundance and quality of
preservation, which includes the preservation in some cases of
soft tissues that normally do not readily fossilize, and a variety
of large and exceptionally complete plant remains not well
known from correlative deposits. The quantity and quality of
preservation identify Kinney as a Lagerstätte (Lucas and Huber,
1991; Kues and Lucas, 1992; Zidek, 1992a).
Scientically signicant fossils were discovered at Kinney
by students studying at the University of New Mexico in the
early 1960s. In 1992, the rst 30 years of research at Kinney
were brought together in an edited volume that detailed the
stratigraphy, age, sedimentology and paleontology, among other
aspects of the Lagerstätte (Zidek, 1992a). The next two decades
saw only sporadic research on the Kinney Lagerstätte. In 2009,
renewed interest in the stratigraphic position and age of the
Kinney deposit ultimately led to the rst controlled excavation
at Kinney, in April 2014. This renewed research interest and the
controlled excavation have produced a wealth of new data on the
Kinney biota and its preservational environment. This volume
presents the results of this recent research, as well as new
research based on older collections. Here, we provide a review
of previous work and of the context within which to understand
the Kinney Quarry Lagerstätte and the articles in this volume.
The Kinney Brick Company, originally owned by the
Kinney family, began to quarry clay and manufacture bricks
in 1928. Though the exact date is not certain, quarrying at
the current clay pit in Pennsylvanian strata in the Manzanita
Mountains began sometime after World War II, in the early 1950s
according to Kelley and Northrop (1975), or in 1946 according
to Elston (1957). In the 1980s, the family sold the company to
Robert Jurgena and Gordon Skarsgard, and they sold it to Ralph
Homan in 1996. Currently (2018 gures), about 9,000 tons
of clay are extracted from the quarry each year to make about
8-9 million bricks at the company’s plant in Albuquerque (R.
Homan, pers. comm., 2020).
Fossils were discovered at the Kinney Brick Quarry in the
early 1960s by two University of New Mexico (UNM) students
who went on to careers in paleontology, Sidney R. Ash (1928-
2019) and John P. Bradbury (1936-2005). Ash discovered fossil
insects and plants at Kinney in 1961, and Bradbury found the
rst fossil shes there in 1963. These discoveries were reported
to Charles B. Read (1907-1979), a paleobotanist who was in
charge of the U.S. Geological Survey’s oce in Albuquerque.
Read visited the site and made some preliminary collections. He
then contacted Smithsonian Curator David H. Dunkle (1911-
1984), who collected at Kinney in 1964, and U. S. Geological
Survey paleobotanist Sergius H. Mamay (1921-2008), who
collected there in 1967 and 1969 (Fig. 3).
Other members of the paleontological community were
soon alerted to the Kinney Brick Quarry as a source of important
fossils. In 1971, two high school students living in Albuquerque,
Neil Lafon and Thomas Lehman (both went on to careers in
geology), found a fossil amphibian at Kinney that was sent to
David Berman, a curator at the Carnegie Museum of Natural
History in Pittsburgh. Berman (1973) named the amphibian
Lafonius lehmani after its discoverers and went on to make a
substantial collection at Kinney now housed at the Carnegie
The rst scientic publications about Kinney paleontology
thus appeared in the 1970s and 1980s (Berman, 1973; Zidek,
1975; Schram and Schram, 1979; Mamay, 1981; Ash and Tidwell,
1982; Kues, 1985). In the late 1980s, the University of Kansas
and the New Mexico Museum of Natural History (NMMNH)
FIGURE 1. Index map and generalized stratigraphy showing
location of the Kinney Brick Quarry in central New Mexico.
FIGURE 2. Photograph of part of the Kinney Brick Quarry in 2012. The oor of the quarry (lower right) exposes the basal limestone
overlain by the primary fossil-producing interval of mostly dark gray shale. The higher wall of the quarry exposes delta-front and
channel sandstones (compare to Figure 7).
both began collecting programs at Kinney. The Kansas collectors
focused on the fossil shes at Kinney and brought in geological
collaborators who studied the sedimentation of the deposits at
the quarry. The NMMNH collections were made primarily by
Phillip Huber, then a student at UNM working with one of us
(SGL) (Fig. 3).
However, by 1990, no comprehensive publication on
the Kinney Lagerstätte had appeared. In that year, Huber, and
Jiri Zidek of the New Mexico Bureau of Mines and Mineral
Resources suggested to one of us (SGL) that a special session
on Kinney be held at the 1991 meeting of the Rocky Mountain
Section/South-central Section of the Geological Society of
America (GSA) in Albuquerque. That session was held and
brought together much research on Kinney (Archer and Clark,
1991; Feldman et al., 1991; Gottfried, 1991; Huber and Lucas,
1991; Lehman, 1991; Lorenz and Lucas, 1991; Lucas, 1991;
Mamay and Mapes, 1991; Mapes, 1991; Shear et al., 1991;
Willard, 1991; Zidek, 1991). The GSA session became the
basis of the volume edited by Zidek (1992a); it published many
new data on Kinney and represented the rst synthesis of the
stratigraphy, sedimentology and paleontology of the Lagerstätte.
Subsequent collecting at Kinney was sporadic for nearly
two decades, mostly by NMMNH volunteers. A eldtrip of
the New Mexico Geological Society visited the quarry in 1999
(Lucas et al., 1999), and, some years later, a eldtrip took place
during the Carboniferous-Permian transition conference ran by
the NMMNH in 2013 (Lucas et al., 2013a). Research interest in
the quarry was renewed in 2009. The completion of a detailed
study of the Pennsylvanian stratigraphy and biostratigraphy in
the Cerros de Amado of Socorro County (Lucas et al., 2009)
raised questions about the stratigraphic position and age of
the Kinney deposit. Fieldwork commenced and expanded to
re-evaluate the entire Pennsylvanian section exposed in the
Manzano and Manzanita mountains (Lucas et al., 2011, 2013a,
b, 2014, 2016; Vachard et al., 2012, 2013; Allen and Lucas,
2018). This research placed Kinney in a dierent stratigraphic
position and assigned it an older age than that of most workers
who contributed to the 1992 volume (Lucas et al., 2011) (Fig. 4).
Additional research on the shes at the quarry was undertaken
by another UNM student, Sally Williams, who, in collaboration
with SGL studied the taphonomy of the Kinney shes (Williams
and Lucas, 2013).
The most recent phase of collecting at Kinney took place
in 2014, when a controlled excavation was completed over the
course of two weeks (Fig. 3). Previous collecting at Kinney
had been by splitting rock to collect the most complete or
interesting fossils. The controlled excavation delineated a 3 x 2
meter grid and collected it layer by layer through the lower 3 m,
recording the detailed stratigraphic and spatial positions of all
the fossils found. This has allowed a much better understanding
of the taphonomy of the fossils, especially of the plant fossils.
The results of that excavation and the analysis of the collected
material are a major component of this volume.
In a UNM masters thesis, Stukey (1967) made the rst
attempt to establish the stratigraphic position and age of the
Kinney Brick Quarry. He located Kinney stratigraphically low in
the “arkosic limestone member” of the Madera Formation (Fig.
4). However, Charles Read claimed to have found what was
then considered a Permian index fossil, the plant Callipteris, at
Kinney (this claim has never been substantiated or replicated).
Thus, Stukey (1967), and later Kelley and Northrop (1975, p.
49), considered that Kinney might be of Permian age and in
strata equivalent to the lower Permian (Wolfcampian) Bursum
In the 1960s and 1970s, Donald Myers of the U. S.
Geological Survey revised the Pennsylvanian lithostratigraphy
in the Manzano and Manzanita mountains and also developed a
fusulinid biostratigraphy of those strata (e.g., Myers, 1973, 1982,
1988a,b). Myers and McKay (1976) mapped the geology of the
Kinney Brick Quarry and the surrounding area. Their mapping
placed the Kinney Brick Quarry in their Pine Shadow Member
of the Wild Cow Formation, a unit of early-middle Virgilian
age based on fusulinid biostratigraphy (Myers, 1988a, b). Note,
however, that Myers knew of no fusulinids from Kinney and
the immediately surrounding area, and he never mentioned the
fossil deposit in any of his publications. Thus, most workers
assigned the Kinney Lagerstätte a Virgilian age (see articles in
Zidek, 1992a). An exception was Huber (1992), who regarded
Kinney as Missourian based on the presence of the chonetid
brachiopod Chonetinella emingi.
Lucas et al. (2011, 2014, 2016), Vachard et al.
(2012, 2013) and Allen and Lucas (2018) restudied the
Pennsylvanian stratigraphy and biostratigraphy in the
Manzano and Manzanita mountains. They rejected Myers’
Pennsylvanian lithostratigraphic nomenclature, and replaced his
lithostratigraphic names by the 1940s nomenclature proposed
by Thompson (1942) and Kelley and Wood (1946), as modied
by Rejas (1965) and Lucas et al. (2009) (Fig. 4). This revised
lithostratigraphy located Kinney in the lower part of the Tinajas
Member of the Atrasado Formation (Figs. 4-5), strata of
Missourian age to the south of the Manzano Mountains. Indeed,
fusulinids from a bed a few meters below the stratigraphic level
of the Kinney fossil deposit and conodonts from the sh bed at
Kinney indicate an early Missourian age (Lucas et al., 2011).
The Pennsylvanian strata at the Kinney Brick Quarry were
deposited in the northeastern portion of the Orogrande basin,
one of the depositional basins of the Ancestral Rocky Mountains
in New Mexico (Fig. 6). At Kinney, the quarrying operation has
exposed about 30 m of the Tinajas Member of the Atrasado
Formation (Fig. 7).
The depositional setting of Kinney has long been interpreted
to be that of a shallow marine embayment (often referred to as an
“estuary” or a “lagoon”) fed by a river delta (Archer and Clark,
1992; Feldman et al., 1992; Lorenz et al., 1992). Lorenz et al.
(1992) identied several distinct depositional environments in
the strata exposed at Kinney that make up a regressive sequence
in which limestone grades up through prodelta and deltaic
clastics to a capping delta-plain facies (Fig. 7).
In this volume, Schneider et al. re-evaluate sedimentation
at the Kinney Quarry. The depositional environment of the
FIGURE 3. Google Earth image of the Kinney Brick Quarry in 2020 showing locations of the three principal excavations, by
Mamay in the late 1960s, by Huber in the late 1980s and the controlled excavation of 2014.
brackish-marine laminated mudstones at Kinney was previously
interpreted as a tide-dominated estuary (Archer and Clark, 1992;
Feldman et al., 1992) or as a non-tidally inuenced prodelta
(Huber, 1992). However, the implied rapid deposition of tidal
deposits contradicts some paleobiological and taphonomic
observations. Schneider et al. interpret the depositional
environment of the laminated mudstone at Kinney as a tidally
modulated bayll sequence controlled by several factors:
(1) an embayed shoreline led to tidal amplication; (2) the
embayed coastline protected the environment from storm-wave
inuence; (3) the prograding bayhead delta supplied nutrients
to the embayment and resulted in increasingly brackish-water
conditions; (4) restricted circulation, poor mixing and elevated
bioproductivity resulted in dysoxic to anoxic bottom water
conditions; and (5) the main sediment input occurred during
seasonal river discharge into the embayment. Two superimposed
orders of lamination are observed in the mudstones. Thicker
packages of laminae representing seasonal river discharge
commonly exhibit internal laminae caused by waxing/waning
ow related to tidal acceleration and deceleration of the river
and the associated sediment plume entering the basin. Poorly
oxygenated bottom water and the resulting lack of infaunal
activity led to the unique preservation of both fossils and
lamination structure in the Kinney Brick Quarry mudstones.
The basal limestone at Kinney (Fig. 7, units 1-2) represents
deposition in a nearshore marine environment that received some
input of freshwater and clastic sediments. Note its lithology
(especially the high black-clay content of this micrite) and
unusual fauna (some stenohaline brachiopods and other groups,
but dominated by euryhaline taxa, such as the inarticulate
brachiopod Lingula and the bivalves Myalina and Solemya).
The black-clay content, terrestrial plant debris, and euryhaline
elements of the fauna (especially abundant Lingula) are
consistent with deposition near the shoreline with a signicant
freshwater input.
The overlying highly fossiliferous shales (Fig. 7, units 3-4)
were deposited in a calm marine embayment with signicant
freshwater input. Uniform and ne grain size, ne lamination
and lack of bioturbation, dark colors, and preservation of soft-
bodied forms suggest deposition in quiet, oxygen-poor waters
with restricted circulation. A mixed hygromorphic-xeromorphic
plant assemblage, dominated by pteridosperm remains, and
FIGURE 4. Evolution of Pennsylvanian stratigraphic nomenclature in the Manzanita Mountains showing changing ideas about the
stratigraphic position of the Kinney Brick Quarry (from Lucas et al., 2011).
freshwater faunal elements (especially conchostracans and
a few salamander-sized amphibians) suggest low salinity.
Overlying shales (Fig. 7, units 5-7) represent a similar facies,
but probably with a greater freshwater inuence. Dominant
fossils are Dunbarella, a euryhaline bivalve, and terrestrial
plants, particularly conifers.
Overlying silty shales (Fig. 7, units 8-9) represent increased
sedimentation rates in a prodelta environment, brought about
by the onset of signicant uvial discharge. Plant remains
are few, and the ora is conifer-dominated, with sparse, small
Dunbarella. Most of the relatively xeromorphic elements of the
oral assemblage may have been oated a short distance into
an environment characterized by frequent shallow ponding and
deposition on a surface better drained than the underlying shales.
Overlying laminated and ripple-laminated sandstone
ledges and intercalated shales and claystones (Fig. 7, units
10-14) are interpreted as delta front, distributary mouth bars,
and associated deposits. Unit 15 is a shale that shows marine
inuence indicated by the presence of Lingula and Myalina.
This unit, and the overlying uvial sandstone/conglomerate,
may be the base of another transgressive sequence. Thus, the
stratigraphic sequence at the Kinney Brick Quarry mostly reects
shoreline progradation, created by the progressive construction
(progradation) of a clastic delta (Fig. 7). Lateral shifts in the
accumulation of sediments from the delta probably fromed an
embayment, isolated from normal marine conditions as the
clastic wedge developed and extended seaward. Clastic input
in the embayment was initially restricted to clay-size particles.
Eventually, the embayment was lled by silty shales from an
advancing delta plain on which sand was later deposited. The
onset of a subsequent transgression is documented by the highest
strata in the quarry section.
Fossils documented from the Kinney Brick Quarry are
palynomorphs, a diverse, macroora consisting of plants typical
of a range of substrate moisture, a shelly marine invertebrate
assemblage that includes a few ammonoids but is dominated
by brachiopods and the pectinacean bivalve Dunbarella,
syncarid and hoplocarid crustaceans, conchostracans, ostracods,
eurypterids, trilobites, terrestrial arthropods (mostly diplopods
and insects), arachnids, conodonts, a diverse assemblage of
shes (mostly acanthodians and palaeoniscoids) and amphibians,
as well as microbially induced sedimentary structures (MISS),
insect and pathogen damage to vegetation and bromalites (mostly
regurgitalites and coprolites). Most of the documentation of
these fossils is published in Zidek (1992a) and this volume.
Willard (1991, 1992) presented the only published work
on palynonomrphs from Kinney. She recovered abundant
assemblages of cordaitealean conifer and pteridosperm pollen as
well as a diversity of spores and identied distinct palynomorph
assemblages from two dierent, spatially separate collections
made at the quarry. Thus, the collections made by Mamay at the
northern end of the clay pit (Fig. 3) indicate a local macroora
composed mostly of cordaitaleans and conifers. To the south, at
the site of the collections made by Huber (Fig. 3), the assemblage
is dominated by spores of pteridophytes, primarily fern spores,
with a background of conifers, cordaitaleans, and a spectrum
of typically wet-substrate taxa (pteridosperms, sphenopsids,
lycopsids, and marattialean tree ferns). Willard (1992)
interpreted the assemblage from the Huber site to be drawn
from a wetter habitat than that of the Mamay site, and suggested
that this may reect dierent distances from the uvial source.
However, the dierences also may reect sampling of dierent
temporal-stratigraphic horizons in the Kinney strata.
Kietzke and Kaesler (1992, g. 6A-C) illustrated two
charophyte gyrogonites from Kinney but did not identify them.
These gyrogonites display characteristic features of Palaeochara,
including their oblate spheroidal shape, the presence of six,
sinistrally spiralled cells, a round and not protruding base and a
small apical beak (cf. Lucas and Johnson, 2016). Palaeochara
is a well-known charophyte that has a stratigraphic range of
Mississippian-early Permian (Lucas and Johnson, 2016; Lucas,
2018). Generally freshwater denizens, these gyrogonites were
likely washed into the Kinney deposit.
Among the most abundant and striking fossils from Kinney
are those of the macroora, which has been the subject of several
FIGURE 5. Geologic map of the area around the Kinney Brick
Quarry (KBQ) (from Allen in this volume).
FIGURE 6. Paleotectonic map of late Paleozoic New Mexico showing basins and uplifts of the Ancestral Rocky Mountain orogeny
and the location of the Kinney Brick Quarry.
FIGURE 7. (facing page) Summary diagram of the paleontology, stratigraphy, depositional environments and sea-level changes at
the Kinney Brick Quarry (from Schneider et al. in this volume).
publications (Mamay, 1981, 1990, 1992; Mamay and Mapes,
1991, 1992; DiMichele et al., 2013). A list of Kinney plant taxa
(Table 1) is taken from DiMichele et al. (2013). In this volume,
DiMichele et al., present an analysis of the taphonomy of the
Kinney plants. The ora, sorted by individual fossiliferous
beds, corresponding to the controlled excavation, is illustrated
by Donovan et al. A particularly large walchian conifer branch
found in one of the beds of the exacavation is described and
illustrated by Looy and Duijnstee.
The Kinney Quarry ora is a characteristic “mixed”
assemblage, typical of a type that characterized the Late
Pennsylvanian and early Permian of western Pangea. By “mixed”
is meant that it includes elements typically characterized as
hygromorphic, mesomorphic, and xeromorphic, intimately
associated within the host strata and each other. Presumably, the
parent plants from which the fossil remains were drawn lived
in environmentally distinct microhabitats in a heterogeneous
landscape. On average, medullosan pteridosperms are
conspicuous, particularly Neuropteris ovata, a widespread and
well characterized species, and a prominent component through
the entire plant-bearing interval. Other important elements
include the calamitalean sphenopsid Phyllotheca, and a variety
of other calamitalean remains, the coniferophytes Walchia and
Dicranophyllum, the likely pteridosperm Sphenopteridium
manzanitanum (Sphenopteris germanica), and the pteridosperm
Mixoneura (Odontopteris) subcrenulata. Marattialean fern
foliage of various forms also is occasionally abundant at some
horizons and locations in the quarry.
Plant fossils were collected from six separate horizons. In
all instances, the remains are allochthonous, which may have
allowed for the mixing of plant remains, drawn from parent
plants growing in close proximity, but in dierent microhabitats
within the shoreline landscape. Some trends were noted in both
composition and preservation from the lowest to the highest
plant-bearing beds. First, plant-fragment size increases from
the bottom to the top of the deposit, within the nearshore
depositional environment (Beds 2 through 5). In fact, in Bed 5,
some very large, complete representatives of walchian conifer
branch systems were found (see Looy and Duijnstee), as well
as the Sphenopteris germanica plant. This nding is concordant
Schultze (2013) suggested that the Kinney amphibians
were salinity tolerant, perhaps marine tetrapods. However,
Werneburg et al. (2013) documented freshwater ostracods as
the gut contents (consumulite) of the holotype of the Kinney
amphibian Milnerpeton huberi. This does not support a marine
habitus for this amphibian.
Schram and Schram (1979) documented two new shrimp
species from Kinney, the syncarid Uronectes kinniensis and the
hoplocarid Aenigmacaris minima. They judged these taxa to be
indicative of a lagoonal environment.
In this volume, Lerner and Lucas re-evaluate the Kinney
syncarid fossils and a better preserved collection of syncarids
from the Tinajas Member in the Cerros de Amado of Socorro
County, New Mexico (cf. Lerner et al., 2009). They assign the
Socorro County syncarids to Palaeocaris secretanae, which is
the rst report of P. secretanae from North America. Uronectes
kinniensis is endemic to the Kinney Lagerstätte, and Lerner and
Lucas add 14 recently collected topotypes to the sparse record
of this species.
Diplopoda and Myriapoda
Shear et al. (1991, 1992) documented a few diplopod
(millipede) specimens from Kinney but judged them to be too
incomplete to be identied precisely. They also documented
a myriapod specimen from Kinney, and considered it to be a
possible centipede.
Most of the insect fossils from Kinney are of blattoids
(cockroaches). Shear et al. (1992) illustrated a single blattoid
wing. Other insects are much less common at Kinney, but some
have been described by Carpenter (1970) and Shear et al. (1992).
These include the lycocercid Madera mamayi, the caloneurod
Pseudobiella fasciata and an indeterminate monuran described
by Carpenter (1970), and a possible brodiid megasecopteran
illustrated by Lucas and Huber (1991, g. 5D) and Shear et
al. (1992, gs. 4A-B, 5A). Much more research remains to be
undertaken on the non-blattoid insects from Kinney.
In this volume, Schneider et al. present a study of about
41 blattoid fossils from Kinney. The family Phyloblattidae is
represented by Phyloblatta occidentalis and the Anthracoblattina
ensifera-gigantea group. Only one specimen belonging to the
Family Necymylacridae is present, and it is identied as the large-
winged Necymylacris scudderi. Members of the Mylacridae are
rare; Opsiomylacris thevenini is represented by one specimen,
and three specimens are identied as Neorthroblattina germari.
Representatives of the family Spiloblattinidae are more
common and pertain to the species Syscioblatta allegheniensis,
Sysciophlebia sp. form KBQ, and a new taxon that Schneider et
al. name Kinneyblatta huberi gen. et sp. nov.
The Kinney Brick Quarry insect fauna is of typical
Euramerican composition, but an unusually high number of
specimens, 23 out of 41, which is 56% of the blattoid insect
remains, are represented by articulated specimens. This contrasts
with the coal-seam roof-shale entomofaunas from Europe, and
may be explained by the taphonomic conditions at Kinney.
Dysoxic to anoxic conditions in the sediment and in the bottom
waters prevented the oxidation of organic remains, as well as
preventing bioturbation and the existence of benthic scavengers.
Nectic predators and necrophageous animals such as shes
and the nectobenthic eurypterids were extremely rare. A high
sedimentation rate caused by sediment plumes during seasonal
river oods resulted in fast burial. Because of the extreme rarity
of terrestrial arthropods, the lack of freshwater aquatic insects,
and the dominance of ying adult insects, river transport seems
with the sedimentological model, in which the sediments were
formed in progressively more proximal environments. Second,
there is no signicant size sorting by taxonomic group within
any one of the sampling levels, nor are there any obvious
dierences in the quality of preservation. Additionally, the fossil
plant components are thoroughly intermixed, regardless of
inferred microhabitat preferences of the parents. Third, the ora
of Beds 2 and 3 diers in its pattern of dominance from the ora
of Beds 4 and 5. The lower beds were dominated by or greatly
enriched in Neuropteris ovata, Phyllotheca sp., Walchia sp., and,
to a lesser extent, Dicranophyllum sp., and marattialean foliage.
This is a mixed assemblage with hygromorphic to mesomorphic
taxa as the most abundant elements. Beds 4 and 5, in contrast,
are characterized by abundant Sphenopteridium manzanitanum
(Sphenopteris germanica), Walchia sp., Dicranophyllum, and
a continuing signicant presence of Neuropteris ovata, with
a noticeable component of marattialean fern foliage. This
represents a subtle change to a more xeromorphic ora. Bed 6
continues this trend to a more xeromorphic fossil assemblage,
and is dominated by walchian conifers.
In this volume, Looy and Dujinstee describe a foliated
walchian conifer branch system from the Kinney deposits. The
specimen’s large size (101 cm) and three orders of branching
are unique among specimens of late Paleozoic, Euramerican
walchian conifers. Analysis of the morphological characteristics
of the specimen’s leaves indicates that it does not t well within
existing taxonomic categories for Pennsylvanian walchians.
There seems to be a strong relationship between leaf-
morphological characteristics and position within the branch
system, and leaf measurements produce a suite of allometric
relationships that govern the observed variation in leaf shape.
These allometric relationships are signicant as a new type of
gross-morphology-based taxonomic characteristic, potentially
of much greater diagnostic value than the highly variable
leaf-measurement ranges commonly employed in walchian
Kozur et al. (1992) presented a preliminary report on
the conchostracans from Kinney. They assigned them to
Pseudestheria sp. Given that conchostracans are primarily
freshwater organisms, Kozur et al. (1992) suggested that some
accumulations of conchostracans on selected bedding planes at
Kinney are mass death assemblages that reect a rise in salinity
that killed the crustaceans.
In this volume, Scholze et al. present a more detailed analysis
of the Kinney conchostracans. The conchostracans from Kinney
are assigned to Pseudestheria sp. a. Pseudestheria limbata.
At Kinney, the occurrence of conchostracans in the marine
to brackish-marine beds is regarded as allochthonous. Thus,
Scholze et al. conclude that the conchostracans were, together
with other freshwater organisms, such as smooth shelled
ostracods, syncarid shrimps, temnospondyl amphibians, and, of
course, the remains of terrestrial plants, most likely washed in
from nearshore freshwater and terrestrial habitats during ood
events caused by heavy rainfalls under a seasonal climate.
Kietzke and Kaesler (1992) documented a low diversity of
ostracods from Kinney that they grouped into three assemblages:
(1) a marine assemblage dominated by Paraparchites from basal
strata at the quarry; (2) an overlying assemblage characterized
by Geisina and judged to represent brackish water; and (3) a
stratigraphically higher freshwater/brackish water assemblage
dominated by Darwinula and Carbonita?. Freshwater forms
also were documented at Kinney by Werneburg et al. (2013) as
gut contents of the amphibian Milnerpeton huberi.
TABLE 1. List of Kinney plant taxa (from DiMichele et al.,
to have played a small role in the input of insects into the Kinney
embayment. Hence, Schneider et al. posit a mainly wind-driven
origin for the very high percentage of nearly complete insect
body fossils at Kinney.
Kues (1985) documented two eurypterids found at Kinney
and identied them as Adelopthalmus luceroensis, a species
originally described from late Virgilian strata of the Red Tanks
Member of the Bursum Formation at Carrizo Arroyo in Valencia
County, central New Mexico.
During the 2014 controlled excavation, a single trilobite
pygidium was found in the sh bed (bed 3) at Kinney.
Dunlop et al. (2014) described a new species of spider-
like trigonotarbid, Pleophrynus hawesi, from Kinney. In this
volume, Selden, in the broader context of a review of Paleozoic
spiders, describes a new specimen of fossil spider from the
Kinney Brick Quarry as Protolycosa suazoi n. sp., in the family
Vermiform Fossils
Vermiform fossils of uncertain anity, possibly annelid
worms or onycophorans, have been reported from Kinney by
Hannibal (1992) and Lerner et al. (2004). They merit further
Kelley and Northrop (1975) mentioned an unpublished
undergraduate study by Burton, who, in 1964, and again later,
extracted about 500 conodonts from the Kinney quarry and
considered them to be of Virgilian age. Krukowski (1992)
identied some conodonts from Kinney as Adetognathus lautus
and Idiognathodus delicatus.. He judged the sample inadequate
for an age assignment.
Barrick (in Lucas et al., 2011) documented conodonts
from the “sh bed” (bed 3) at Kinney. The Kinney conodont
fauna they reported is characterized by Idiognathodus
corrugatus and I. cherryvalensis, which suggest an assignment
to the Idiognathodus confragus Zone of the North America
Midcontinent region (Dennis cyclothem; middle Missourian).
In this volume, Rosscoe and Barrick re-evaluate the
Kinney conodont fauna based on a much larger sample
than was previously available. Two conodont faunas were
recovered; one from the sh bed in the Kinney Brick Quarry
and one from a stratigraphically lower fusulinid marker bed
from nearby outcrops. Both faunas are characteristic of the
lower part of the Missourian Stage (Kasimovian). The fusulinid
marker bed conodont fauna correlates with the diverse fauna
of the Hushpuckney Shale from the Swope cyclothem in the
Midcontinent Basin (Idiognathodus cancellosus Zone). Species
of the fusulinid genus Triticites occur with the Swope-equivalent
conodonts in the fusulinid marker bed, indicating that Triticites
appeared in New Mexico very early in Missourian time. The
Kinney Brick Quarry sh-bed conodont fauna correlates with
the low diversity fauna of the younger minor Mound Valley
cyclothem (base of I. confragus Zone).
Kues (1992a) documented the fossil assemblage from
the basal limestone at the Kinney Quarry that includes the
brachiopods Lingula (very abundant) and a few specimens
of Chonetinella, Linoproductus, Composita and Derbya. He
inferred that these were brachiopods that lived in an environment
of uctuating salinity and restricted circulation.
From the basal limestone bed at Kinney, Kues (1992a)
documented bivalves that are common fossils of Solemya,
Myalina and Dunbarella and rare fossils of Streblochondria?,
Clinopisthia, Leptodesma and Parallelodon? Some living
Solemya prefer low oxygen settings with large amounts of
dissolved organic matter (Pojeta, 1988), and Myalina and
Dunbarella are well known to have been euryhaline. This
ts the interpretation of the basal limestone at Kinney having
been deposited in an oxygen depleted environment with poor
circulation. Myalina is also present in the shale immediately
above the basal limestone at Kinney.
Dunbarella is a well known pectinacean bivalve found in
a range of fully marine, brackish and freshwater settings (e. g.,
Johnson, 1962; Murphy, 1967). It is the most obvious and most
abundant animal fossil at the Kinney Brick Quarry, particularly
in shale beds in the quarry section. Clark (1978) rst drew
attention to the close association of many Dunbarella shells with
plant matter at Kinney, and the fact that some Dunbarella were
evidently attached to plant stems (also see Mamay, 1981, 1990;
Lucas and Huber, 1991). Kues (1992b) presented a detailed
study of the Kinney Dunbarella and concluded that they were
r-strategists that proliferated rapidly to achieve large size and
extensive numbers, and experienced seasonal mortality due to
uctuations in salinity and sediment inux.
From the basal limestone, Kues (1992a) reported a few
poorly preserved gastropods that he assigned to Euphemites,
Glabrocingulum (most common) and an unidentied form. This
included an undetermined taxon of high-spired gastropod, also
found in the overlying sh bed.
Kietzke and Kaesler (1992, g. 6H-L) illustrated two
specimens they identied as Spirorbis sp., but such fossils are
now correctly identied as those of microconchiod gastropods.
Microconchids may indicate some degree of brackish water,
although that is subject to debate (Gierlowski-Kordesch and
Cassle, 2015; Gierlowski-Kordesch et al., 2016; Zatoń et al.,
2016). The mixed marine, brackish and freshwater nature of
the Kinney deposits means that the salinity preferences of the
Kinney microconchids cannot be readily resolved.
The basal limestone at the Kinney quarry contains ammonoids
and a few straight and coiled nautiloids (Mapes, 1991; Mapes
and Boardman, 1992; Kues, 1992a). All of the ammonoids were
assigned to Prothalassoceras kingorum Miller, and one of the
straight nautiloids was identied as Pseudorthoceras knoxense
(McChesney) by Mapes and Boardman (1992). One of the
ammonoid specimens shows exceptional preservation of a thick
carbon lm inferred to represent the mandibles (aptychi) and
stomach contents. Mapes and Boardman (1992) considered the
ammonoids to have lived in a “restricted environment” (not a
normal marine environment) in the Kinney embayment.
The Kinney deposit yields a diverse assemblage of shes,
many complete and superbly preserved. As Hodnett and Lucas
(2015) noted, this is one of the best preserved and most diverse
Pennsylvanian sh assemblages in the American Southwest.
The taxa recognized can be divided into acanthodians,
chondrichthyans, actinopterygians and sarcopterygians. In this
volume, Hodnett and Lucas review the Kinney sh fauna to
identify 31 distinct sh taxa, including the rst records of a new
ctenacanthiform shark, two hybodontiforms, two holocephalans,
three actinopterygians, and a megalichthyoform sarcopterygian
(Table 2). This is a mixed salinity sh assemblage found almost
exclusively in bed 3 at the Kinney Quarry.
Zidek (1975) rst described acanthodians from Kinney,
and later (Zidek, 1992b) named A. jurgenai, the single species
of acanthodian found at the Kinney Quarry. As Williams and
Lucas (2013) noted, A. kinneyi is the second most common sh
at Kinney, typically found as incomplete skeletons of young
individuals of various ontogenetic stages.
Zidek (1975, 1992b) rst published on the shark fossils from
Kinney, which were few in number and mostly isolated teeth
and a dermal spine. He assigned them to ve taxa: Peripristus
a. P. semicircularis, Symmorium reniforme, ?Listracanthus,
Orthacanthus huberi (named by Zidek, 1992b) and Cobelodus
aculeatus. Hodnett and Lucas (2015) questioned some of these
identications but only revised one, changing Symomorium to
Glikmanius. Of these sharks, the best known is the symmoriid
Cobelodus,” including a remarkably complete specimen that
contains a well-preserved cololite. Williams and Lucas (2013)
regarded the sharks as known from fossils allochthonous to the
Kinney embayment or representing occasional marine visitors
to the embayment.
Collecting at Kinney since 2013 has substantially
augmented the record of chondrichthyans from Kinney so that
Hodnett and Lucas (2015, and in this volume) listed 11 species
of chondrichthyans, several of which are new and yet unnamed
taxa (Table 2). This increased diversity and the presence of
complete specimens may necessitate a reassessment of the
conclusion that none of the shark fossils represent shes that
lived in the Kinney estuary.
Particularly signicant is a 2.5-meter long skeleton of a new
ctenacanth that Hodnett et al. in this volume name Dracopristis
homanorum gen. et sp. nov. It represents the most complete
ctenacanth found in North America. The morphology of
Dracopristis suggests it specialized in the benthic environment
as a slow moving ambush predator in the marine embayment at
Actinopterygian shes from the Kinney quarry are a diverse
group of about 16 taxa (Table 2). Most common is the deep-bodied
Platysomus” schultzei (Zidek, 1992b). Another well known,
deep-bodied sh from Kinney is “Amphicentrum” jurgenai.
Other well-described taxa are Schizolepis manzanitaensis,
Tanyrhinichthys mcallesteri, Pyritocephalus lowneyae and an
aduelliform, cf. Bourbonella (Gottfried, 1987a, b, 1992; Huber,
1992; Williams and Lucas, 2013; Hodnett and Lucas, 2015;
Stack et al., 2021). Less studied are various “paleoniscoids”
that Bardack (1992) reviewed and assigned to six morphotypes.
These shes are abundant at Kinney and in need of further study.
A single lungsh toothplate assigned to Sagenodus hlavini
(Zidek, 1975; Kemp, 1996), a rhizodont and an undescribed
coelacanth species are the sarcopterygian record from Kinney
(Schultze, 1992; Hodnett and Lucas, 2015).
Tetrapods from Kinney are a small assemblage of
amphibians: the lepospondyl Brachydectes?, the “amphibamid”
Milnerpeton, the trimerorhachid Lafonius and a new dvinosaurian
named by Werneburg et al. in this volume (Berman, 1973; Hunt
et al., 1992, 1996; Werneburg et al., 2013). This is an assemblage
of primarily aquatic tetrapods.
In this volume, Werneburg et al. make two separate
contributions to knowledge of the Kinney amphibians. In
one, a new early adult specimen of Milnererpeton huberi
brings new insights to the ontogenetic development of the
Werneburg et al. in their second contribution name the
new dvinosaurian Bermanerpeton kinneyi. Nine unique
characters diagnose Bermanerpeton kinneyi, many shared
with branchiosaurids and larval “amphibamids.” Otherwise,
Bermanerpeton is clearly a dvinosaurian. The recorded prey
in the intestines and stomach of Bermanerpeton consists of
dierent arthropods, shes and amphibians. In the consumulite,
ostracods with smooth shells belong to the freshwater/brackish
water ostracod ?Carbonita. Bermanerpeton was thus not marine
adapted, but rather a freshwater animal, either washed into the
TABLE 2. The sh assemblage of the Kinney Brick Quarry (from Hodnett and Lucas in this volume).
Acanthodes kinneyi
Cobelodus sp.”
Orthacanthus sp.
Family Indeterminate
Glikmanius occidentalis
Dracopristis homanorum
Family Indeterminate
Hybodontiform indeterminate 1.
Hybodontiform indeterminate 2.
Peripristis sp.
Family Chondrechelyidae
Chondrechelyid indeterminate
?Family Myriacanthidae
?Myriacanthid indeterminate
Subclass Indeterminate
Order Indeterminate
Family Listracanthidae
Ancanthorhachis sp.
Family indeterminate
Indeterminate Chondrichthyan
Rhadinichthyid indeterminate (Bardack’s type 3)
Elonichthyid indeterminate 1
(Bardack’s type 5)
Elonichthyid indeterminate 2
Pyritocephalus lowneyae
Family Indeterminate
Schizolepis manzanitaensis
Tanyrhinichthys mcallisteri
“Palaeoniscoid” indeterminate 1
“Palaeoniscoid” indeterminate 2
“Palaeoniscoid” indeterminate 3
“Palaeoniscoid” indeterminate 4
“Palaeoniscoid” indeterminate 5
“Palaeoniscoid” indeterminate 6
Platysomus” schultzei
Platysomid indet.
Amphicentrum” jurgenai
cf. Bourbonnella sp.
Gen. et. sp indet
Sagenodus hlavini
Family Indeterminate
Indeterminate Megalichthyid
Kinney embayment or living there when freshwater conditions
Trace Fossils
Microbially-induced sedimentary structures
Microbially-induced sedimentary structures (MISS) reect
the inuence of microbial biolms and mats on sedimentation
(e.g., Noke, 2010). Little studied before the 2000s, MISS
is now recognized as an important ichnological aspect of the
sedimentary record.
Recognition of MISS at Kinney only began in 2019 when
one of us (SGL) identied what are possible syneresis fractures
mediated by microbial activity on some bedding planes in
the quarry strata. In this volume, Schneider et al. discuss this
MISS, interpreting the polygonal networks as subaqueous MISS
generated by growth and expansion of microbial mats.
Arthropod herbivory
Like MISS, the study of arthropod damage on fossilized
vegetation is of recent vintage, having really begun during
the 1990s (Lucas, 2016). In this volume, Donovan and Lucas
document damage on Kinney plant fossils due to arthropod
herbivory (these are considered trace fossils) and by pathogens
(these are not generally considered trace fossils: Bertling et
al., 2006). The highest diversity of damage is on medullosan
pteridosperms. Donovan and Lucas record insect and pathogen
damage on 2254 fossil plant foliage specimens, describe all
damage by host plant, and analyzed damage diversity and
frequency. They nd low damage diversity, with nine damage
types in two functional feeding groups, including external foliage
feeding (hole feeding, margin feeding, surface feeding), piercing
and sucking, as well as oviposition and pathogen damage.
Insect damage was associated with both drought-tolerant and
wetland components of the ora, suggesting herbivorous insects
had colonized multiple microhabitats across the landscape.
Medullosan pteridosperms, including Neurodontopteris
auriculata, Neuropteris ovata, and Mixoneura subcrenulata, are
associated with the highest damage diversity at Kinney, which
provides further evidence for a general preference for seed
plants during the early proliferation of insect herbivory.
Eggs are not generally considered trace fossils (Bertling et
al., 2006), but we discuss them here for convenience. Mamay
(1994) identied small (up to 2 mm diameter) compressed,
spherical bodies attached to pteridosperm foliage at Kinney as
sh eggs. In this volume, Lucas et al. re-evaluate these fossils
as gastropod eggs. They are spherical rings of carbon around
host-sediment-lled cavities, or carbon-lm-coated spheres.
Attached to pteridosperm foliage, these eggs display denite
evidence of desiccation, indicating that they were almost
certainly laid subaerially and thus not by shes. The Kinney
eggs are remarkably similar to eggs of Devonian, Jurassic
and Cretaceous age attributed to gastropods, and also t well
within the range of modern gastropod egg morphology. Thus,
gastropods, not shes, likely produced the Kinney eggs.
The term bromalite refers to “anally or orally derived ejecta
and in situ intestinal matter” including “coprolites, cololites and
regurgitalites” (Hunt, 1992, p. 221; also see Hunt and Lucas,
2012). Hunt (1992) coined this widely used term in his rst
study of coprolites from Kinney. These were about 20 specimens
of coprolites and a cololite in the body of the shark Cobelodus
(also see Zidek, 1992b). These trace fossils were attributed to
sh producers.
Hunt et al. (2012) presented another study of Kinney
bromalites based on a larger sample. They assigned the bromalites
to seven morphotypes, including one that was the basis of a new
ichnotaxon, Conchobromus kinneyensis. This ichnotaxon refers
to coprolites with a groundmass of conchostracan shells, likely
made by acanthodian shes.
In this volume, Hunt and Lucas review the more than 100
bromalites from Kinney. They name two new ichnogenera
and three new ichnospecies of non-evisceralite consumulites:
Werneburgichnus kinneyensis and W. varius from branchiosaur-
like amphibians, and Chondripilula zideki from chondrichthyans.
New ichnogenera and ichnospecies of non-consumulite
bromalites named by Hunt and Lucas are Huberobromus ovatus,
Maculacoprus ateri, Virgacoprus brevis, and Kinneybromus
jurgenai. Conchobromus kinneyensis is also present, as are
various unnamed morphotypes of coprolites. The Kinney
bromalite ichnofauna is signicant because: (1) it contains
the most studied bromalites of any Paleozoic ichnofauna and
includes the highest number of named ichnotaxa; (2) its study
stimulated the development of a synthetic nomenclature, with
the introduction of the terms bromalite and regurgitalite; (3) it
includes the rst named non-evisceralite consumulite taxa; and
(4) the Kinney ichnofauna provides a reference for bromalites in
shallow marine embayment paleoenvironments.
The Kinney Brick Quarry presents classic aspects of
sedimentary geology and paleontology that makes it an ideal
teaching tool for science educators. To that purpose, in this
volume Burton presents a eld trip to enhance the learning
experience by placing the students in a real-world environment
at Kinney. This eldtrip demonstrates in the eld the results
of earth-change processes that can only be talked about in the
classroom environment. Burton aligns the content of his eld
guide to the applicable National Next Generation Standards for
Public Education.
The Kinney Lagerstätte is signicant in several ways.
Perhaps foremost are the many taxa rst discovered at Kinney
FIGURE 8. Pennsylvanian Lagerstätten (after Schultze and
Maples, 1992).
and the exceptional preservation of many of its fossils that
provide unique morphology not known otherwise. Recent work
indicates that such discoveries will continue at Kinney, and it
will long remain an important source of new morphology and
new taxa.
As noted above, and described in associated contributions
to this volume, the Kinney ora consists of an intimately
intermixed assemblage of plants typical of high soil moisture,
tolerant of only short periods of drought, and forms that are
considered drought-tolerant. Such a “mixed” assemblage is
most likely to be drawn from a landscape characterized by
habitat, even microhabitat, heterogeneity. The extremes of
heterogeneity indicated by the Kinney ora would be unlikely
to be found on a delta plain and associated oodplain under
a humid climate, with relatively high rainfall, nearly equably
distributed throughout the year. Rather, the regional climate
almost certainly was strongly seasonal. The rationale for this
interpretation is explained in detail in papers by DiMichele
et al. (2020) and Bashforth et al. (2021). Seasonal drought
magnies microhabitat dierences that would be masked under
a higher volume, more equably distributed rainfall regime. In
a nearshore to shoreline setting, like the Kinney Quarry, the
opportunity for the close proximity of standing water and better
drained microhabitats is great. We suggest, therefore, that the
parent plants of the fossil ora populated a complex, spatially
and environmentally variable terrestrial environment, and lived
within close proximity of one another
The animal fossils at Kinney are a mixture of taxa that
lived in the embayment (most of the invertebrates and shes),
those washed in from terrestrial/freshwater environments (the
insects and amphibians) and marine visitors to the estuary (the
sharks). If these fossils fully capture the diversity that lived in
the Late Pennsylvanian embayment, then that diversity was
low compared to modern analogues (e.g., Williams and Lucas,
2013), either a result of taphonomic bias and/or a Pennsylvanian
biota of lower diversity than the Modern world.
Schultze and Maples (1992) compared the Kinney
Lagerstätte to other Pennsylvanian Lagerstätten (Fig. 8) to
conclude that Kinney is most similar to the Lagerstätten at
the Garnett, Hamilton and Robinson localities in Kansas (also
see Maples and Schultze, 1988). These Lagerstätten were
characterized by Schultze and Maples (1992) as nearshore
marine fossil assemblages that accumulated along tidally
inuenced coastlines or in estuaries. Kinney is now known to be
older than Hamilton and Robinson, which are both of Virgilian
age. Indeed, Kinney lls a temporal gap in the Pennsylvanian
Lagerstätten between Desmoinesian localities such as Linton
and Mazon Creek and the late Missourian Garnett locality. In
contrast, from the perspective of the vegetation, Kinney falls
among those oras with an abundance of both drought-tolerant
and drought-intolerant taxa. Many so-called mixed oras have
been documented in both the Pennsylvanian and early Permian,
summarized in some detail in Bashforth et al. (2021). Many of
these, especially those with a major tree-fern component, are
found well into the early Permian, up into the Leonardian (e.g.,
Emily Irish – Koll and DiMichele, 2020; Montgomery Ranch
- Simon et al., 2018). In contrast, Garnett is heavily dominated
by conifers, with few pteridophyte, or even medullosan
pteridosperm elements (Winston, 1983), and Hamilton is
similarly highly conifer dominated, but with pteridosperms
and calamitaleans, but no signicant tree-ferns. The Hamilton
ora also is composed both of adpressions and anatomically
preserved remains in a carbonate matrix (Rothwell and Mapes,
1988). These oristic and preservational dierences suggest
dissimilarities in the various Lagerstätte, perhaps in prevailing
climatic conditions.
We are grateful to the owners of the Kinney Brick Quarry, in
particular the late Robert Jurgena, and the current owners, Ralph
and Jeanette Homan, for their willingness to allow collecting
and other research at the quarry over many years. Our thanks
also to all the contributors to this volume, which was completed
during the COVID pandemic of 2020-2021 under conditions
that slowed but did not prevent its completion. Adrian Hunt and
Joerg Schneider provided helpful reviews of the manuscript.
Allen, B.D. and Lucas, S.G., 2018, The Late Pennsylvanian (Missourian)
index fusulinid Eowaeringella in the Manzanita Mountains of
central New Mexico: New Mexico Geology, v. 40, p. 35-44.
Archer, A.W. and Clark, G.W., 1991, Tempo and mode of deposition
in the Dunbarella bed: reconciling its paleoecology and sediment
cyclicity: Geological Society of America, Abstracts with Programs,
v. 23, no. 4, p. 3.
Archer, A.W. and Clark, G.W., 1992, Depositional environment of
the Dunbarella beds: an exercise in paleoecology and sediment
cyclicity: New Mexico Bureau Mines and Mineral Resources,
Bulletin 138, p. 27-36.
Ash, S.A. and Tidwell, W.D., 1982, Notes on the upper Paleozoic
plants of central New Mexico: New Mexico Geological Society,
Guidebook 33, p. 245-248.
Bardack, D., 1992, Late Pennsylvanian paleonisciform sh from the
Kinney Quarry, New Mexico: New Mexico Bureau Mines and
Mineral Resources, Bulletin 138, p. 197-203.
Bashforth, A.R., DiMichele, W.A., Eble, C.F., Falcon-Lang, H.J., Looy,
C.V. and Lucas, S.G., 2021, The environmental implications of
upper Paleozoic plant-fossil assemblages with mixtures of wetland
and drought-tolerant taxa in tropical Pangea. Geobios, in press.
Berman, D.S., 1973, A trimerorhachid amphibian from the Upper
Pennsylvanian of New Mexico: Journal of Paleontology, v. 47, p.
Bertling, M., Braddy, S.J., Bromley, R.G., Demathieu, G.R., Genise,
J., Mikuláš, R., Nielsen, J.K., Nielsen, K.S. S., Rindsberg, A.K.,
Schlirf, M., Uchman, A., 2006, Names for trace fossils: a uniform
approach: Lethaia, v. 39, p. 265-286.
Carpenter, F.M., 1970, Fossil insects from New Mexico: Psyche, v. 77,
p. 400-412.
Clark, G.R., II, 1978, Byssate scallops in a Late Pennsylvanian lagoon:
Geological Society of America, Abstracts with Programs, v. 10,
p. 380.
DiMichele, W.A., Bashforth, A.R., Falcon-Lang, H.J. and Lucas, S.G.,
2020, Uplands, lowlands, and climate: taphonomic megabiases
and the apparent rise of a xeromorphic, drought-tolerant ora
during the Pennsylvanian-Permian transition: Palaeogeography,
Palaeoclimatology, Palaeoecology, v. 559, 109965.
DiMichele, W.A., Wagner, R.H., Bashforth, A.R. and Álvarez-
Vázquez, C., 2013, An update on the ora of the Kinney Quarry
of central New Mexico (Upper Pennsylvanian), its preservational
and environmental signicance: New Mexico Museum of Natural
History and Science, Bulletin 59, p. 289-325.
Dunlop, J. A., Wang, Y., Selden, P.A. and Krautz, P., 2014, A
trigonotarbid arachnid from the Pennsylvanian Atrasado
Formation of the Kinney Brick Quarry, New Mexico: University
of Kansas Paleontological Contributions, no. 9, 6 p.
Elston, W.E., 1957, Summary of the mineral resources of Bernalillo,
Santa Fe, and Sandoval Counties, New Mexico: New Mexico
Bureau of Mines and Mineral Resources, Bulletin 81, 81 p.
Feldman, H.R., Archer, A.W., Kvale, E.P. and Maples, C.G., 1991,
Origin and taphonomy of the Kinney Brick Company Quarry
sh bed: implications for a Carboniferous Lagerstätten model:
Geological Society of America, Abstracts with Programs, v. 23,
no. 4, p. 21.
Feldman, H.R., Archer, A.W., West, R.R. and Maples, C.G., 1992,
The Kinney Brick Company Quarry: preliminary analysis using
an estuarine depositional model: New Mexico Bureau Mines and
Mineral Resources, Bulletin 138, p. 21-26.
Gierlowski-Kordesch, E.H., and C.F. Cassle. 2015. The ‘Spirorbis’
problem revisited: Sedimentology and biology of microconchids
in marine-nonmarine transitions: Earth-Science Reviews, v. 148,
p. 209-227.
Gierlowski-Kordesch, E.H., H.J. Falcon-Lang, and C.F. Cassle, 2016,
Reply to comment on the paper of Gierlowski-Kordesch and Cassle
“The ‘Spirorbis’ problem revisited: sedimentology and biology of
microconchids in marine-nonmarine transitions” [Earth-Science
Reviews, 148 (2015):209-227]: Earth-Science Reviews, v. 152, p.
Gottfried, M.D., 1987a, A Pennsylvanian aeduelliform (Osteichthyes,
Actinopterygii) from North America with comments on
aeduelliform relationships: Paläontologische Zeitschrift, v. 61, p.
Gottfried, M.D., 1987b, A new long-snouted actinopterygian sh from
the Pennsylvanian of north-central New Mexico: New Mexico
Journal of Science, v. 27, p. 7-19.
Gottfried, M.D., 1991, Unusual lower actinopterygian shes from
the Kinney Quarry fauna, Upper Pennsylvanian, New Mexico:
Geological Society of America, Abstracts with Programs, v. 23,
no. 4, p. 26.
Gottfried, M.D., 1992, A new deep scaled “paleoniscoid” from the
Kinney Quarry, Late Pennsylvanian of New Mexico: New Mexico
Bureau Mines and Mineral Resources, Bulletin 138, p. 189-196.
Hannibal, J.T., 1992, Enigmatic vermiform fossils from Upper
Pennsylvanian rocks at the Kinney Brick Quarry, New Mexico:
New Mexico Bureau Mines and Mineral Resources, Bulletin 138,
p. 119-122.
Hodnett, J.P. and Lucas, S.G., 2015, Paleozoic shes of New Mexico:
A review: New Mexico Museum of Natural History and Science,
Bulletin 68, p. 51-63.
Huber, P., 1992, Pyritocepalus lowneyae n. sp., the youngest
Haplolepiformes (Pisces: Actinopterygii) from the Pennsylvanian
of central New Mexico: New Mexico Bureau Mines and Mineral
Resources, Bulletin 138, p. 183-187.
Huber, P. and Lucas, S.G., 1991, Youngest haplolepiform (Osteichthyes:
Actinopterygii), from the Pennsylvanian (early Virgilian) of New
Mexico: Geological Society of America, Abstracts with Programs,
v. 23, no. 4, p. 34.
Hunt, A.P., 1992, Late Pennsylvanian coprolites from the Kinney Brick
Quarry, central New Mexico, with notes on the classication and
the utility of coprolites: New Mexico Bureau Mines and Mineral
Resources, Bulletin 138, p. 221-229.
Hunt, A.P. and Lucas, S.G., 2012, Classication of vertebrate coprolites
and related trace fossils: New Mexico Museum of Natural History
and Science, Bulletin 57, p. 137-146.
Hunt, A.P., Lucas, S.G. and Berman, D.S., 1992, The Late Pennsylvanian
amphibian fauna of the Kinney Quarry, central New Mexico: New
Mexico Bureau Mines and Mineral Resources, Bulletin 138, p.
Hunt, A.P., Lucas, S.G. and Berman, D.S., 1996, A new amphibamid
(Amphibia: Temnospondyli) from the Late Pennsylvanian (Middle
Stephanian) of central New Mexico, USA: Paläontologische
Zeitschrift, v. 70, p. 555
Hunt, A.P., Lucas, S.G., Spielmann, J.A., Suazo, T.L. and Cantrell,
A.K., 2012, A re-evaluation of Late Pennsylvanian bromalites
from the Kinney Brick Quarry Lagerstätte, New Mexico, USA:
New Mexico Museum of Natural History and Science, Bulletin
57, p. 185-192.
Johnson, R.G., 1962, Interspecic association in Pennsylvanian fossil
assemblages: Journal of Geology, v. 70, p. 32-55.
Kelley, V.C. and Northrop, S.A., 1975, Geology of Sandia Mountains
and vicinity, New Mexico: New Mexico Bureau of Mines and
Mineral Resources, Memoir 29, 136 p.
Kelley, V.C. and Wood, G.H., Jr., 1946, Lucero uplift, Valencia, Socorro
and Bernalillo Counties, New Mexico: U. S. Geological Survey,
Oil and Gas Investigations Preliminary Map 47, scale 1:63,360.
Kemp, A., 1996, Sagenodus (Proceratodus) carlinvillensis (Romer and
Smith 1934), (Osteichthyes: Dipnoi), short ridge anomaly and
classication of dipnoans: Journal of Vertebrate Paleontology, v.
16, p. 16-19.
Kietzke, K.K. and Kaesler, R.L., 1992, Late Pennsylvanian Ostracoda
from the Kinney Brick Quarry, Bernalillo County, New Mexico,
with notes on other microfossils: New Mexico Bureau Mines and
Mineral Resources, Bulletin 138, p. 127-133.
Koll, R.A. and DiMichele, W.A., 2021, Dominance-diversity
architecture of a mixed hygromorphic-to-xeromorphic ora
from a botanically rich locality in western equatorial Pangea
(lower Permian Emily Irish site, Texas, USA: Palaeogeography,
Palaeoclimatology, Palaeoecology, v. 563, 110132.
Kozur, H., Lucas, S.G. and Hunt, A.P., 1992, Preliminary report on
Late Pennsylvanian Conchostraca from the Kinney Brick Quarry,
Manzanita Mountains, New Mexico: New Mexico Bureau Mines
and Mineral Resources, Bulletin 138, p. 123-126.
Krukowski, S.T., 1992, Conodont platform elements from the Madera
Formation (Pennsylvanian) at the Kinney Brick Company Quarry,
Manzanita Mountains, New Mexico: New Mexico Bureau Mines
and Mineral Resources, Bulletin 138, p. 143-144.
Kues, B.S., 1985, Eurypterids from the Wild Cow Formation (Upper
Pennsylvanian), Manzano Mountains, New Mexico: New Mexico
Journal of Science, v. 25, p. 23-31.
Kues, B.S., 1992a, A Late Pennsylvanian restricted-marine fauna from
the Kinney Quarry, Manzanita Mountains, New Mexico: New
Mexico Bureau Mines and Mineral Resources, Bulletin 138, p.
Kues, B.S., 1992b, The bivalve Dunbarella in marine and nonmarine
facies of the Upper Pennsylvanian sequence at the Kinney Quarry,
Manzanita Mountains, New Mexico: New Mexico Bureau Mines
and Mineral Resources, Bulletin 138, p. 99-111.
Kues, B.S. and Lucas, S.G., 1992, Overview of Upper Pennsylvanian
stratigraphy and paleontology of the Kinney Brick Quarry,
Manzanita Mountains, New Mexico: New Mexico Bureau Mines
and Mineral Resources, Bulletin 138, p. 1-11.
Lehman, T.M., 1991, Sedimentology of the Kinney Brick Company
Quarry section, New Mexico: Geological Society of America,
Abstracts with Programs, v. 23, no. 4, p. 41.
Lerner, A.J, Hannibal, J.T. and Lucas, S.G., 2004, A probable annelid
from the Upper Pennsylvanian (Virgilian) Atrasado Formation
(Madera Group) of central New Mexico: Geological Society of
America, Abstracts with Programs, v. 36, no. 4, p. 5.
Lerner, A.J., Lucas, S.G., Spielmann, J.A., Krainer, K., DiMichele, W.A.,
Chaney, D.S., Schneider, J.W., Nelson, W.J. and Ivanov, A.B.,
2009, The biota and paleoecology of the Upper Pennsylvanian
(Missourian) Tinajas locality, Socorro County, New Mexico: New
Mexico Geological Society, Guidebook 60, p. 267–280.
Lorenz, J.C. and Lucas, S.G., 1991, Sedimentation patterns in
Pennsylvanian strata at the Kinney Brick Quarry, Bernalillo
County, NM: Geological Society of America, Abstracts with
Programs, v. 23, no. 4, p. 44.
Lorenz, J.C., Smith, G.S. and Lucas, S.G., 1992, Sedimentation patterns
in Pennsylvanian strata at the Kinney Brick Quarry, Bernallilo
County, New Mexico: New Mexico Bureau Mines and Mineral
Resources, Bulletin 138, p. 13-19.
Lucas, S.G., 1991, Late Pennsylvanian stratigraphy, paleontology and
depositional environments, Kinney Brick Quarry, Manzanita
Mountains, New Mexico: Geological Society of America,
Abstracts with Programs, v. 23, no. 4, p. 44.
Lucas, S.G., 2016, Two new, substrate-controlled nonmarine
ichnofacies: Ichnos, v. 23, p. 248-261.
Lucas, S.G., 2018, Permian-Triassic charophytes: Distribution,
biostratigraphy and biotic events: Journal of Earth Science, v. 29,
p. 778-793.
Lucas, S.G. and Huber, P., 1991, Late Pennsylvanian stratigraphy and
paleontology of the Kinney Brick Quarry, Manzanita Mountains,
New Mexico: New Mexico Bureau of Mines and Mineral
Resources, Bulletin 137, p. 79-86.
Lucas, S.G. and Johnson, G.D., 2016, Palaeochara from the lower
Permian of Oklahoma: New Mexico Museum of Natural History
and Science, Bulletin 74, p. 141-143.
Lucas, S.G., Krainer, K. and Barrick, J.E., 2009, Pennsylvanian
stratigraphy and conodont biostratigraphy in the Cerros de Amado,
Socorro County, New Mexico: New Mexico Geological Society,
Guidebook 60, p. 183-212.
Lucas, S.G., Krainer, K. and Vachard, D., 2016, The Pennsylvanian
section at Priest Canyon, southern Manzano Mountains, New
Mexico: New Mexico Geological Society, Guidebook 67, p. 275-
Lucas, S.G., Krainer, K., Allen, B.D. and Vachard, D., 2014, The
Pennsylvanian section at Cedro Peak: A reference section in
the Manzanita Mountains, central New Mexico: New Mexico
Geology, v. 36, p. 3-24.
Lucas, S.G., Read, A., Karlstrom, K.E., Estep, J.W., Kues, B.S.,
Anderson, O.J., Smith, G.A. and Pazzaglia, F.J., 1999, Second-
day trip 1 road log, from Albuquerque to Tijeras, Cedar Crest and
Sandia Crest: New Mexico Geological Society, Guidebook 50, p.
Lucas, S.G., Allen, B.D., Krainer, K., Barrick, J., Vachard, D.,
Schneider, J.W., William, A., DiMichele, W.A. and Bashforth,
A.R., 2011, Precise age and biostratigraphic signicance of the
Kinney Brick Quarry Lagerstätte, Pennsylvanian of New Mexico,
USA: Stratigraphy, v. 8, p. 7-27.
Lucas, S.G., Allen, B.D., Krainer, K. and Spielmann, J.A., 2013a, Road
log from Albuquerque to Upper Pennsylvanian strata in the Kinney
Brick Quarry, Manzanita Mountains, New Mexico: New Mexico
Museum of Natural History and Science, Bulletin 59, p. 1-10.
Lucas, S.G., Krainer, K., Allen, B.D. and Vachard, D., 2013b,
Pennsylvanian stratigraphy and biostratigraphy at Cedro Peak,
Manzanita Mountains, New Mexico, USA: A brief summary: New
Mexico Museum of Natural History and Science, Bulletin 59, p.
Mamay, S.H., 1981, An unusual new species of Dicranophyllum
Grand’Eury from the Virgilian (Upper Pennsylvanian) of New
Mexico, U.S.A.: The Palaeobotanist, v. 28-29, p. 86-92.
Mamay, S.H., 1990, Charliea manzanitana, n. gen. n. sp., and
other enigmatic parallel-veined foliar forms from the Upper
Pennsylvanian of New Mexico and Texas: American Journal of
Botany, v. 77, p. 858-866.
Mamay, S.H., 1992, Sphenopteridium and Telangiopsis in a
Diplopteridium-like association from the Virgilian (Upper
Pennsylvanian) of New Mexico: American Journal of Botany, v.
79, p. 1092-1101.
Mamay, S.H., 1994, Fossil eggs of probable piscine origin preserved on
Pennsylvanian Sphenopteridium foliage from the Kinney Quarry,
central New Mexico: Journal of Vertebrate Paleontology, v. 14, p.
Mamay, S.H. and Mapes, G., 1991, Early Virgilian plant megafossils
from the Kinney Quarry, Manzanita Mountains, central New
Mexico: Geological Society of America, Abstracts with Programs,
v. 23, no. 4, p. 45.
Mamay, S.H. and Mapes, G., 1992, Early Virgilian plant megafossils
from the Kinney Brick Company Quarry, Manzanita Mountains,
New Mexico: New Mexico Bureau Mines and Mineral Resources,
Bulletin 138, p. 61-86.
Mapes, R.H., 1991, The late Pennsylvanian (Virgilian) cephalopods
from the Kinney Quarry, Manzanita Mountains, central New
Mexico: Geological Society of America, Abstracts with Programs,
v. 23, no. 4, p. 45.
Mapes, R.H. and Boardman, D.R., II, 1992, Late Pennsylvanian
cephalopods from the Kinney Quarry, Manzanita Mountains,
New Mexico: New Mexico Bureau Mines and Mineral Resources,
Bulletin 138, p. 113-118.
Maples, C.G. and Schultze, H.-P., 1988, Preliminary comparison of
the Pennsylvanian assemblage of Hamilton, Kansas, with marine
and nonmarine contemporaneous assemblages: Kansas Geological
Survey, Guidebook Series 6, p. 253-273.
Murphy, J.L., 1967, R.P. Stevens’ pelecypod species from the
Brush Creek Shale Member (Conemaugh) of Ohio: Journal of
Paleontology, v. 41, p.1498-1504.
Myers, D.A., 1973, The upper Paleozoic Madera Group in the Manzano
Mountains, New Mexico: U.S. Geological Survey, Bulletin 1372-
F, 13 p.
Myers, D. A., 1982, Stratigraphic summary of Pennsylvanian and lower
Permian rocks, Manzano Mountains, New Mexico: New Mexico
Geological Society, Guidebook 33, p. 233-237.
Myers, D.A., 1988a, Stratigraphic distribution of some Pennsylvanian
fusulinids from the Sandia Formation and the Los Moyos
Limestone, Manzano Mountains, New Mexico: U. S. Geological
Survey, Professional Paper 1446-A, 21 pp.
Myers, D.A., 1988b, Stratigraphic distribution of some Pennsylvanian
fusulinids from the Wild Cow and Bursum formations, Manzano
Mountains, New Mexico: U. S. Geological Survey, Professional
Paper 1446-B, 42 p.
Myers, D.A., and McKay, E.J., 1976, Geologic map of the north end
of the Manzano Mountains, Tijeras and Sedillo quadrangles,
Bernalillo County, New Mexico: U. S. Geological Survey,
Miscellaneous Investigation Series Map I-968, scale 1:24,000.
Noke, N., 2010, Geobiology of microbial mats in sandy deposits from
the Archean to Today: Berlin, Springer-Verlag, 194 p.
Pojeta, J., Jr., 1988, The origin and Paleozoic diversication of
solemydoid pelecypods: New Mexico Bureau of Mines and
Mineral Resources, Memoir 44, p. 201-271.
Rejas, A., 1965, Geology of the Cerros de Amado area, Socorro County,
New Mexico: [M.S. thesis]: Socorro, New Mexico Institute of
Mining and Technology, 128 p.
Rothwell, G.W. and Mapes, G., 1988, Vegetation of a Paleozoic conifer
community; in: Mapes, G. and Mapes R.H., eds., Regional Geology
and Paleontology of Upper Paleozoic Hamilton Quarry Area in
Southeastern Kansas. Kansas Geological Survey Guidebook 6, p.
Schram, F.R. and Schram, J.M., 1979, Some shrimp of the Madera
Formation (Pennsylvanian) Manzanita Mountains, New Mexico:
Journal of Paleontology, v. 53, p. 169-174.
Schultze, H-P., 1992, Coelacanth sh (Actinista, Sarcopterygii) From
the Late Pennsylvanian of the Kinney Brick Company Quarry,
New Mexico: New Mexico Bureau Mines and Mineral Resources,
Bulletin 138, p. 205-209.
Schultze, H-P., 2013, The paleoenvironment at the transition from
piscine to tetrapod sarcopterygians: New Mexico Museum of
Natural History and Science, Bulletin 60, p. 373-397.
Schultze, H-P. and Maples, C. G., 1992, Comparison of the Late
Pennsylvanian faunal assemblage of Kinney Brick Company
Quarry, New Mexico, with other Pennsylvanian Lagerstätten:
New Mexico Bureau Mines and Mineral Resources, Bulletin 138,
p. 231-242.
Shear, W.A., Hannibal, J.T. and Kukalová-Peck, J., 1991, Terrestrial
arthropods (insects and myriapods) from Upper Pennsylvanian
rocks at the Kinney Brick Quarry, central New Mexico: Geological
Society of America, Abstracts with Programs, v. 23, no. 4, p. 93.
Shear, W.A., Hannibal, J.T. and Kukalová-Peck, J., 1992, Terrestrial
arthropods from Upper Pennsylvanian rocks at the Kinney Brick
Quarry, New Mexico: New Mexico Bureau Mines and Mineral
Resources, Bulletin 138, p. 135-141.
Simon, S.S., Gibling, M.R., DiMichele, W.A., Chaney, D. and Koll, R.,
2018, An exhumed ne‐grained meandering channel in the lower
Permian Clear Fork Formation, north‐central Texas: Processes of
mud accumulation and the role of vegetation in channel dynamics:
International Association of Sedimentologists Special Publication
48, p. 149-172.
Stack, J., Hodnett, J-P., Lucas, S.G. and Sallan, L., 2021, Tanyrhinichthys
mcallisteri, a long-rostrumed Pennsylvanian ray-nned sh
(Actinopterygii) and the simultaneous appearance of novel
ecomorphologies in Late Palaeozoic shes: Zoological Journal of
the Linnean Society, v. 191, p. 347-374.
Stukey, A.H., 1967, Stratigraphic relations of Pennsylvanian-Permian
strata, Manzanita Mountains, New Mexico [M.S. thesis]:
Albuquerque, University of New Mexico, 64 p.
Thompson, M.L., 1942, Pennsylvanian System in New Mexico: New
Mexico Bureau of Mines and Mineral Resources, Bulletin 17, 92
Vachard, D., Krainer, K. and Lucas, S.G., 2012, Pennsylvanian
(Carboniferous) calcareous microfossils from Cedro Peak (New
Mexico, USA). Part 1: Algae and Microproblematica: Annales de
Paléontologie, v. 98, p. 225-252
Vachard, D., Krainer, K. and Lucas, S.G., 2013, Pennsylvanian
(Carboniferous) calcareous microfossils from Cedro Peak (New
Mexico, USA). Part 2: Smaller foraminifers and fusulinids:
Annales de Paléontologie, v. 99, p. 1-42.
Werneburg, R., Schneider, J.W. and Lucas, S.G., 2013, The
dissorophoid Milnererpeton huberi (Temnospondyli) from the Late
Pennsylvanian Kinney Brick Quarry in New Mexico restudied –
paleontology, paleoenvironment, and age: New Mexico Museum
of Natural History and Science, Bulletin 59, p. 349-370.
Willard, D.A., 1991, Early Virgilian palynooras from the Kinney
Quarry, Manzanita Mountains, central New Mexico: Geological
Society of America, Abstracts with Programs, v. 23, no. 4, p. 105.
Willard, D.A., 1992, Early Virgilian palynooras from the Kinney
Quarry, Manzanita Mountains, New Mexico: New Mexico Bureau
of Mines and Mineral Resources, Bulletin 138, p. 49–60.
Williams, S.C. and Lucas, S.G., 2013, Taphonomy and paleoecology of
Pennsylvanian shes from the Kinney Brick Quarry, New Mexico,
USA: New Mexico Museum of Natural History and Science,
Bulletin 39, p. 371-389.
Winston, R.B., 1983. A Late Pennsylvanian upland ora in Kansas:
systematics and environmental implications: Review of
Palaeobotany and Palynology, v. 40, p. 5-31.
Zatoń, M., Wilson, M.A. and Vinn, O., 2016, Comment on the paper
of Gierlowski-Kordesch and Cassle “The ‘Spirorbis’ problem
revisited: Sedimentology and biology of microconchids in marine–
nonmarine transitions” [Earth-Science Reviews, 148 (2015): 209–
227]: Earth-Science Reviews, v. 152, p. 198-200.
Zidek, J., 1975, Some shes of the Wild Cow Formation (Pennsylvanian)
Manzanita Mountains, New Mexico: New Mexico Bureau of
Mines and Mineral Resources, Circular 135, p. 1-22.
Zidek, J., 1991, Acanthodian, chondrichthyan, and platysomid shes
of the Kinney Quarry, Manzanita Mountains, New Mexico
(Pennsylvanian: Virgilian): Geological Society of America,
Abstracts with Programs, v. 23, no. 4, p. 109.
Zidek, J., ed., 1992a, Geology and Paleontology of the Kinney Brick
Quarry, Late Pennsylvanian, central New Mexico: New Mexico
Bureau Mines and Mineral Resources, Bulletin 138, 242 p.
Zidek, J., 1992b, Late Pennsylvanian Chondrichthyes, Acanthodii, and
deep bodied Actinopterygii from the Kinney Quarry, Manzanita
Mountains, New Mexico: New Mexico Bureau Mines and Mineral
Resources, Bulletin 138, p. 145-182.
... Shelly marine invertebrates, mainly brachiopods and bivalves (Dunbarella), crustaceans (syncarids, hoplocarids, conchostracans and ostracodes), conodonts, fishes (including coprolites and regurgitalites) lived in the estuary, while terrestrial myriapods, insects (flying adult Blattodea), arachnids (Dunlop et al. 2014;Selden 2021) and amphibians were washed in by seasonal river floods, and preserved in the anoxic muds. This Lagerstätte has recently received renewed interest (Lucas et al. 2021b). It has produced three previously documented eurypterids (Adelophthalmus) (Kues 1985;Braddy et al. 2021), but this is the first report of a Hibbertopterus. ...
Hibbertopterus lamsdelli sp. nov., from the Late Carboniferous Kinney Quarry Lagerstätte of New Mexico (USA), is a large (ca. 1.1 m long) stylonurid eurypterid (sea-scorpion; Chelicerata), similar to H. scouleri from Scotland but with less serrate segment margins, a wider pretelson, shorter telson (tail-spine), and more parallel ventral keels. It is only the fourth, yet most reliable record of an American hibbertopterid. A taxonomic reassessment of Hibbertopterus regards Dunsopterus and Vernonopterus (but not Cyrtoctenus) as synonyms. Hibbertopterids were aquatic (benthic) scavengers and microphagous sweep-feeders, but their trackways indicate that they were capable of brief terrestrial, seasonal nuptial walks, despite their large size; Hibbertopterus had walking legs with spinose extensions at the base (Laden) to spread their load, and the ventral keels on their telson functioned like sled rails to reduce body drag. Hibbertopterids were interpreted as moving into freshwater during the Late Palaeozoic, but a trackway from the middle Permian Collingham Formation (Ecca Group) of South Africa may be from a marine setting, though further analysis is needed to fully evaluate this possibility.
... Среди всех пермских местонахождений рыб Европейской России только Кичкасс [15] и Шихово-Чирки [16] могут сравниться по количеству и качеству материала. По сохранности комплекса ископаемых остатков Куеда-Ключики приближается к такому известному пресноводному лагерштетту, как Кинней Брик Кверри (Kinney Brick Quarry) [17] чем 10 скелетами каждый. Характерной чертой ориктокомплекса является наличие ювенильных особей лучепёрых рыб, редко встречающихся в других пермских местонахождениях. ...
Full-text available
The Kueda-Klyuchiki locality of Permian vertebrates located in the south of the Perm Region in the Kuedinsky District was discovered in 2005 and quickly became known among paleontologists. The locality contains a rich complex of exceptionally preserved plants and vertebrates (fish, amphibians, and reptiles). Their remains are confined to limestones of various textures, which form two bone-bearing beds in the section of the quarry. However, the age of rocks remains a controversial issue. This research aims to analyze and compare various stratigraphic markers and discuss a number of existing stratigraphic inconsistencies in an attempt to arrive at a general conclusion about the age of the locality. According to geological survey data on a scale of 1:200 000, the Buraevo Sequence (Sheshmian For-mation, Ufimian Stage) was identified in the territory of the quarry. At the boundary of the Sheshmian and Belebey Formations, a mineralogical “boundary” is observed, which is manifested in a decrease in the proportion of minerals of the epidote group and an increase in the content of black ore minerals. Terrigenous rocks of the Kueda-Klyuchiki section are close to the values of the Sheshmian Formation, in terms of the amount of epidote and black ore minerals in the heavy fraction. The results of the geo-chemical study indicate that the bitumoids of the Kueda-Klyuchiki and Posad outcrops (Kazanian Stage) developed in different paleogeographic and geomorphological conditions. The bitumoids of the Kazanian outcrop Posad formed through the sapropelic organic matter accumulated in the coastal-marine reducing conditions, while the organic matter of the Kueda-Klyuchiki outcrop formed in the relatively deep-water conditions. The composition of the ichthyological assemblage indicates the Upper Kazanian age of the section. The fish orictocenosis includes Kazanichthys viatkensis, Acropholis kamensis, Acropholis stensioei, Palaeoniscum kasanense, Platysomus biarmicus, Kargalichthys efremovi, Kargalichthys pritokensis, Platysomidae gen. indet., Palaeonisci ordo indet. sp. 1, Palaeonisci ordo indet. sp. 2. Its composition indicates that Kueda-Klyuchiki has higher stratigraphic position than most Upper Kazanian localities, which is confirmed by the absence of terminal Kazanian species. However, the composition of the ichthyofauna in Kueda-Klyuchiki differs significantly from that of the early Kazanian localities, typical of the Kazanichthys golyushermensis ichthycomplex. The orictocomplex of fish from the Kueda-Klyuchiki locality is most similar to the two Upper Kazanian localities – Aksakovo (Pechishchi beds) and Shikhovo-Chirki (Verkhnyi Uslon beds). The complex of amphibians includes temnospondyls (presumably, Melosauridae), discosauriscid (Discosauriscidae gen. ind.) and kotlassiid (Leptoropha sp.) seymouriamorphs, diapsids, and therapsid reptiles. To clarify the age of the Kueda-Klyuchiki outcrop, the results of the mineralogical analysis of the heavy fraction of terrigenous rocks and the geochemical characteristics of the dispersed organic matter from the carbonate-clayey rocks of the section are presented. The results obtained can serve as a starting point for further multidisciplinary research.
Full-text available
Actinopterygii makes up half of living vertebrate diversity, and study of fossil members during their Palaeozoic rise to dominance has a long history of descriptive work. Although research interest into Palaeozoic actinopterygians has increased in recent years, broader patterns of diversity and diversity dynamics remain critically understudied. Past studies have investigated macroevolutionary trends in Palaeozoic actinopterygians in a piecemeal fashion, variably using existing compendia of vertebrates or literature‐based searches. Here, we present a comprehensive occurrence‐based dataset of actinopterygians spanning the whole of the Palaeozoic. We use this to produce the first through‐Palaeozoic trends in genus and species counts for Actinopterygii. Diversity through time generally tracks metrics for sampling, while major taxonomic problems pervading the Palaeozoic actinopterygian record obscure diversity trends. Many described species are concentrated in several particularly problematic ‘waste‐basket’ genera, hiding considerable morphological and taxonomic diversity. This taxonomic confusion also feeds into a limited understanding of phylogenetic relationships. A heavy sampling bias towards Europe and North America exists in both occurrence databases and available phylogenetic matrices, with other regions underrepresented despite yielding important data. Scrutiny of the extent to which spatial biases influence the actinopterygian record is lacking, as is research on other forms of bias. Low richness in some time periods may be linked to geological biases, while the effects of taphonomic biases on Palaeozoic actinopterygians have not yet been investigated. Efforts are already underway both to redescribe poorly defined taxa and to describe taxa from underrepresented regions, helping to address taxonomic issues and accuracy of occurrence data. New methods of sampling standardisation utilising up‐to‐date occurrence databases will be critical in teasing apart biological changes in diversity and those resulting from bias. Lastly, continued phylogenetic work will enable the use of phylogenetic comparative methods to elucidate the origins of actinopterygian biogeography and subsequent patterns of radiation throughout their rise to dominate aquatic faunas.
Full-text available
Ancient fine‐grained meandering channels are under‐represented in the literature and their formative processes are rarely explored. The Montgomery Ranch 3 site of the Clear Fork Formation of Texas contains an exhumed fine‐grained point bar that migrated for at least 50 m within a channel 2 m deep and 36 m wide. The point bar comprises thick inclined layers of unstratified mudstone intercalated with thin layers of fine‐grained, ripple cross‐laminated sandstone, with dips averaging nearly 16o. Rill casts and swept ripples on the sandstone surfaces indicate declining water levels. Petrographic analysis of the mudstone shows silt and clay laid down from suspension, but sand‐sized mud aggregates transported as bedload (present at other sites in the formation) were not observed. The sandstone beds are attributed to lateral accretion on the point bar during periods of sustained flow, whereas the mudstone beds are attributed to oblique accretion as fine sediment draped the bar during waning and low‐flow periods. Sandstone and mudstone units are composite units from numerous flow events and their alternation may reflect secular variation in flood frequency and intensity. In an associated abandoned‐channel fill, weakly laminated mudstone with desiccation cracks contains leaves and seeds of Evolsonia texana, marattialean foliage and Taeniopteris sp., with root traces penetrating the leaves. Some taxa preferred high water tables and humid conditions, whereas others were dryland colonisers. This apparent discrepancy may reflect the persistence of wetter channel reaches within an otherwise dry setting. Despite the scarcity of preserved plant fossils, vegetation was probably sufficiently widespread to promote bank strength and local sediment accumulation.
Full-text available
A new eophrynid trigonotarbid (Arachnida: Trigonotarbida: Eophrynidae) from the Pennsylvanian (Kasimovian) Astrasado Formation of the Kinney Brick Quarry, New Mexico is described. This fossil – the first arachnid to be recorded from the Astrasado Formation – is preserved primarily as a dorsal opisthosoma. Pleophrynus hawesi sp. nov. diagnostically preserves evidence for three pairs of posterior opisthosomal spines rather than the two usually seen in other eophrynids. A comparison of opisthosomal tuberculation patterns among the best known eophrynid species is included. At ca. 304.0–306.5 Ma, our new taxon represents one of the youngest records of Eophrynidae, while the Kinney Brick Quarry is only the twelfth site in North America to yield trigonotarbid arachnids; compared to more than 60 such localities in Europe.
Upper Pennsylvanian (Virgilian) rocks of a lagoonal deposit in the Manzanita Mountains, north-central New Mexico, contain a rich biota of both plants and animals. The plants are mostly typical Pennsylvanian genera, but the assemblage is dominated by a new species of Dicranophyllum (D. readii), characterized by its unusually long leaves. The leaves are slender, consistently twice-bifurcate, and judging from the largest fragments, reached lengths of 75 cm or slightly more. Inasmuch as Dicranophyllum is very rare in North America, this large-leaved new species lends considerable interest to the New Mexico flora and indicates that palaeobotanical exploration in the south-western United States should prove continuingly productive.
We evaluate the influences of elevation and climate on the spatio-temporal distribution of wetland and dryland biomes during the Pennsylvanian and early Permian in tropical Pangea. The longstanding ‘‘upland model” places drought-tolerant vegetation in elevated habitats, where slope and drainage created moisture-limited substrates under a humid climate that simultaneously promoted peat accumulation in contemporaneous lowlands. Upland plants were periodically transported to, and buried in, lowlands. Rare preservation of dryland vegetation thus reflects its general absence in basins, and taphonomic vagaries of long-distance transport. The alternative ‘‘climate model” proposes that drought-tolerant plants dominated tropical habitats when climate was seasonally dry, with wetland vegetation reduced to scattered refugia. Environmental changes attending glacial-interglacial cycles caused alternating wetter-drier conditions, and the relative abundance of wetland versus dryland biomes in basinal lowlands thus varied with climatic oscillations. The paucity of drought-tolerant plants reflects a preservational megabias against habitats with seasonal moisture deficits. The environmental signal of ‘‘mixed” plant-fossil assemblages, comprising taxa characteristic of both wetland and dryland biomes, may help resolve these debates. We review key Pennsylvanian and lower Permian mixed assemblages from tropical Euramerican Pangea, and interpret their original habitats and climatic contexts based on multidisciplinary lines of evidence, including sedimentology, taphonomy, physiology, and paleoecology. Evaluations also consider patterns of vegetational distribution and taphonomy in modern tropical environments. We suggest that even a cursory view of current tropical plant distribution exposes flaws in the upland model. Where tropical climate is sufficiently humid to support peat swamps, slopes and elevated habitats do not host drought-tolerant vegetation, but are occupied by plants similar to those in lowland settings. This occurs because equable, high precipitation strongly dampens water-table variation across entire landscapes. Furthermore, taphonomic studies indicate that most plant-fossil assemblages record vegetation living near the burial site. Fossil floras thus reflect environmental conditions near their growth site, excluding an upland origin for most occurrences of drought-tolerant taxa. Conversely, the climate model is consistent with modern tropical vegetational distribution and soundly explains late Paleozoic floristic patterns. When Pangean tropical lowlands experienced seasonally dry conditions, plants tolerant of moisture deficits dominated most habitats, whereas wetland vegetation was restricted to wetter sites with greater preservation potential. This occurred because topographic variations are magnified under seasonal precipitation regimes, creating a complex habitat mosaic with wetland patches in a landscape subject to seasonal drought. Accordingly, we propose that a macrofloral assemblage with even rare drought-tolerant taxa indicates seasonality in the broader landscape. At larger spatio-temporal scales, disagreement also persists about whether tectonic uplift or long-term climatic drying was the primary driver of changes in late Paleozoic floristic patterns and areal extent of tropical peat swamps. We argue that tectonic activity alone cannot explain the drastic reduction in peat swamps or coincident changes in dominance-diversity of wetland vegetation. Rates of plant dispersal and evolution far outpace that of mountain building, and peat-forming wetlands persisted in elevated habitats well into the Late Pennsylvanian. Therefore, progressive late Paleozoic aridification was the most probable driver of changing floral patterns and the distribution of wetland and dryland biomes in tropical Pangea.
Here we describe the Leonardian-age flora from “Emily Irish” (Russell Ranch), a small oxbow lake deposit, which is believed to be one of the most extensively collected, single-excavation floras of this age from any place in the world, including ca. 5200 hand specimens housed at the US National Museum of Natural History. The flora is dominated by a mixture of xeromorphic and meso-hygromorphic elements. The xeromorphs include conifers, taeniopterids, cordaitaleans, and noeggerathialeans. The mesomorphic-to-hygromorphic elements include marattialean tree ferns, sphenopsids, lycopsids, callipterids, and medullosan pteridosperms. This co-dominance of hygromorphic and xeromorphic elements is made especially conspicuous by the intimate co-mingling of plant remains from each group on the same hand specimens, and by the relative preservation-state of the macrofossils, which does not differ between the two groups. These observations suggest that they inhabited the same landscape immediately surrounding the depositional environment, and were likely only differentiated by microhabitat. We suggest that the Emily Irish flora grew under seasonally dry climate, but that areas adjacent to the lake and feeder streams retained sufficient moisture to support patches of taxa requiring consistently high soil moisture. These patches, thus, were embedded within a landscape of periodic moisture deficit that supported vegetation composed mostly of xeromorphic elements. This is more likely than requiring the xeromorphic taxa to be transported to the site from extrabasinal habitats, the nearest of which were hundreds of kilometers away at the time the deposit formed.
The Late Mississippian and Pennsylvanian have been referred to as the Coal Age due to enormous paleotropical peat accumulations (coal beds). Numerous fossil floras have been collected from these coals, and their associated seat-earth paleosols and roof-shales, over more than two centuries, leading to the inference of vast swampy wetlands covering the Pangean tropics during the Pennsylvanian. In contrast, the Permian tropics are characterized as more arid, with sparser and more heterogeneous vegetation than inferred for the Pennsylvanian. In the tropics, the Pennsylvanian to Permian transition has been described as a changeover from a pteridophytedominated “Paleophytic flora”, to a seed-plant dominated “Mesophytic flora. This view notwithstanding, floras dominated by xeromorphic seed plants also are well known from the Pennsylvanian tropics. Some authors have characterized these plants as being occupants of uplands, subsequently transported into basinal-lowland, preservational environments. In this model, uplands are well drained, causing areas of drought under otherwise everwet climates. In this paper, we present an alternative interpretation: that the apparent transition in Pennsylvanian-Permian tropical vegetation reflects two types of taphonomic megabias. First is a preservational megabias, strongly favoring the vegetation of humid climates over that of seasonally dry climates. Accordingly, tropical-plant preservational potential fluctuated in concert with Late Paleozoic Ice Age glacial-interglacial oscillations, and contemporaneous sea-level and climatic changes. Second is an analytical megabias, strongly favoring the discovery and collection of the wetland biome from Pennsylvanian strata, overlooking the less frequently and more poorly preserved drought-tolerant biome. By Permian times, vast wetlands, and their fossil record, had largely disappeared from central Pangea (although continuing in Cathaysia), making drought-tolerant vegetation more “visible” to searchers, without changing its preservational circumstances. We demonstrate that the upland model is untenable, being inconsistent with the principles of plant biogeography and with geological aspects of the fossil record.
The Carboniferous radiation of fishes was marked by the convergent appearance of then-novel but now common ecomorphologies resulting from changes in the relative proportions of traits, including elongation of the front of the skull (rostrum). The earliest ray-finned fishes (Actinopterygii) with elongate rostra are poorly known, obscuring the earliest appearances of a now widespread feature in actinopterygians. We redescribe Tanyrhinichthys mcallisteri, a long-rostrumed actinopterygian from the Upper Pennsylvanian (Missourian) of the Kinney Brick Quarry, New Mexico. Tanyrhinichthys has a lengthened rostrum bearing a sensory canal, ventrally inserted paired fins, posteriorly placed median fins unequal in size and shape, and a heterocercal caudal fin. Tanyrhinichthys shares these features with sturgeons, but lacks chondrostean synapomorphies, indicating convergence on a bottom-feeding lifestyle. Elongate rostra evolved independently in two lineages of bottom-dwelling, freshwater actinopterygians in the Late Pennsylvanian of Euramerica, as well as in at least one North American chondrichthyan (Bandringa rayi). The near-simultaneous appearance of novel ecomorphologies among multiple, distantly related lineages of actinopterygians and chondrichthyans was common during the Carboniferous radiation of fishes. This may reflect global shifts in marine and freshwater ecosystems and environments during the Carboniferous favouring such ecomorphologies, or it may have been contingent on the plasticity of early actinopterygians and chondrichthyans.
Permian charophytes are known from the Ukraine, Russia, Kazakhstan, Germany, Saudi Arabia, China, the USA, Brazil, Paraguay and India. Most of these records are of Middle- Late Permian Age and are the basis of local biostratigraphic zonation in southern Russia and China. Development of a robust Permian charophyte biostratigraphy will require a more extensive record. Triassic charophytes are known from Germany, Sweden, Poland, Slovenia, Bulgaria, the Ukraine, Russia, Morocco, Congo, the USA, Argentina, Kazakhstan and China. This encompasses records from all Triassic stages and has been the basis of detailed biostratigraphic zonation in southern Russia-Kazakhstan-eastern Europe. Permian and Triassic charophyte biostratigraphy at the level of genus does not provide detailed correlations beyond local or regional schemes. Nevertheless, it does identify some important evolutionary datums that constrain the timing of important biotic events in the Permian-Triassic evolutionary history of the Charophyta, including: (1) Early Permian extinction of the Palaeocharaceae; (2) Late Permian extinction of the “Trochiliscales” (Moellerinales); (3) Carboniferous origin of the paraphyletic Porocharaceae, soon followed during the Permian by the origin of the multicellular basal plate; and (4) an important generic turnover of charophytes across the Triassic-Jurassic boundary, though there are insufficient data to identify this as a mass extinction.
Charliea is a new genus (type-species: C. manzanitana), based on pinnately compound leaf material from the richly fossiliferous Virgilian (Upper Pennsylvanian) shales of the Kinney Brick Company quarry near Albuquerque, New Mexico. In several features Charliea resembles Russellites or a zamioid cycad. It has linear-oblong pinnae with broad, oblique attachment and a truncate tip, which is deeply incised to form two to four nearly equal lobes. The venation is simple, parallel, and sparingly dichotomous, each vein ending at the distal margin. The Kinney beds also contain Plagiozamites planchardi, another zamioid form with parallel-veined pinnae, differing from Charliea chiefly in having rounded tips and veins ending in the denticulate margins. An unnamed third form (genus B) in the Kinney beds has long, narrow pinnae with parallel veins and blunt tips; this strongly resembles the Mesozoic conifer Podozamites, but may just as well represent a cycadophyte. Another unnamed taxon (genus A), from an Upper Pennsylvanian deposit in Jack County, Texas, resembles genus B or Russellites in general shape and venation, but the critical distal margins are unknown. In their single-ordered parallel venation, these four foliar types contrast sharply with the two-ordered pinnate venation of most Pennsylvanian fern-like leaves, and seem to foreshadow Mesozoic morphologies. This tendency toward precocious evolution of parallel-veined foliar form in North America is also expressed by a single occurrence of the Asiatic, Permian genus Tingia in the Lower Pennsylvanian of Utah, and by the presence of the predominantly Triassic cycadeoid genus Pterophyllum in the Lower Permian of Texas.