The Bear Gulch Limestone is a
limestone
Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
-rich
geological lens in central
Montana
Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...
, renowned for the quality of its late
Mississippian-aged fossils. It is exposed over a number of outcrops northeast of the
Big Snowy Mountains
The Big Snowy Mountains ( ats, níichʔibííkʔa, lit=it is never summer) are a small mountain range south of Lewistown in Fergus County, Montana. Considerably east of and isolated from the main crest of the Northern Rockies, they are one of the ...
, and is often considered a component of the more widespread
Heath Formation. The Bear Gulch Limestone reconstructs a diverse, though isolated, marine ecosystem which developed near the end of the
Serpukhovian
The Serpukhovian is in the ICS geologic timescale the uppermost stage or youngest age of the Mississippian, the lower subsystem of the Carboniferous. The Serpukhovian age lasted from Ma to Ma. It is preceded by the Visean and is followed b ...
age. It is a
lagerstätte
A Lagerstätte (, from ''Lager'' 'storage, lair' '' Stätte'' 'place'; plural ''Lagerstätten'') is a sedimentary deposit that exhibits extraordinary fossils with exceptional preservation—sometimes including preserved soft tissues. These for ...
, a particular type of rock unit with exceptional fossil preservation of both articulated skeletons and
soft tissue
Soft tissue is all the tissue in the body that is not hardened by the processes of ossification or calcification such as bones and teeth. Soft tissue connects, surrounds or supports internal organs and bones, and includes muscle, tendons, ligam ...
s. Bear Gulch fossils include a variety of fish, invertebrates, and
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
occupying a number of different habitats within a preserved shallow bay.
Fish
Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
include a high diversity of unusual
chondrichthyans
Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. C ...
(cartilaginous fish)
and one of the oldest known
lamprey
Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s,
along with other
vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s.
Invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s include numerous fossils of
crustacean
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
worm
Worms are many different distantly related bilateral animals that typically have a long cylindrical tube-like body, no limbs, and no eyes (though not always).
Worms vary in size from microscopic to over in length for marine polychaete wor ...
s,
cephalopod
A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s, and
sea sponges
Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through ...
, which are concentrated in different parts of the bay basin. Shelled cephalopods are abundant, and the Bear Gulch Limestone also preserves some of the best early fossils of
coleoid
Subclass Coleoidea,
or Dibranchiata, is the grouping of cephalopods containing all the various taxa popularly thought of as "soft-bodied" or "shell-less" (i.e., octopuses, squid and cuttlefish). Unlike its extant sister group, Nautiloidea, whose ...
cephalopods.
Fossils of typical
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
(seabed-attached) organisms are rare, and those that do occur are restricted to reef-like sponge patches in eastern exposures. This indicates that the conditions of the bay responsible for the Bear Gulch Limestone were at odds with other marine areas nearby.
Some Bear Gulch fossils were preserved so rapidly and efficiently that mating behaviors,
internal organs,
coloration patterns,
gut content,
and even the pattern of blood vessels
could be observed in fossils. This exceptional preservation may be a consequence of the prevailing warm
monsoon
A monsoon () is traditionally a seasonal reversing wind accompanied by corresponding changes in precipitation but is now used to describe seasonal changes in atmospheric circulation and precipitation associated with annual latitudinal oscil ...
al climate, as storms could frequently and rapidly cover the seabed with oxygen-poor organic runoff from shallower areas.
Although up to 40 meters of sediment are found in the Bear Gulch Limestone,
biostratigraphic
Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of Bio ...
data suggests that the lens was emplaced in only 1000 years, a geological instant.
Geology
Geological setting
The Bear Gulch Limestone is commonly considered to be part of the
Heath Formation, the youngest formation in the
Big Snowy Group
The Big Snowy Group is a stratigraphical unit of Chesterian age in the Williston Basin.
It takes the name from Big Snowy Mountains in Montana, and was first described on the north slopes of the mountain by H.W. Smith in 1935.
Lithology
Subdiv ...
of central
Montana
Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...
.
Some authors instead consider the Bear Gulch Limestone to be an early member of the
Tyler Formation, a patchy but widespread unit of Carboniferous
limestone
Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
and terrestrial sediments.
Most of the Heath Formation is represented by
black shale
Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especiall ...
s and
marl
Marl is an earthy material rich in carbonate minerals, clays, and silt. When hardened into rock, this becomes marlstone. It is formed in marine or freshwater environments, often through the activities of algae.
Marl makes up the lower part o ...
s, indicative of
brackish
Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estuari ...
and salty
littoral
The littoral zone or nearshore is the part of a sea, lake, or river that is close to the shore. In coastal ecology, the littoral zone includes the intertidal zone extending from the high water mark (which is rarely inundated), to coastal areas ...
environments. It developed along a
transgressive sequence in a narrow saltwater seaway, known as the
Central Montana Trough or Big Snowy Trough.
This seaway flowed into the
Williston Basin
The Williston Basin is a large intracratonic sedimentary basin in eastern Montana, western North Dakota, South Dakota, southern Saskatchewan, and south-western Manitoba that is known for its rich deposits of petroleum and potash. The basin is a ...
, a shallow
inland sea
An inland sea (also known as an epeiric sea or an epicontinental sea) is a continental body of water which is very large and is either completely surrounded by dry land or connected to an ocean by a river, strait, or "arm of the sea". An inland se ...
further east. The Central Montana Trough would have also been linked to fully marine basins on the western coastline of
Laurussia
Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
,
but this connection may have been broken by the time of the Bear Gulch Limestone's deposition.
Many distinct limestone
lenses
A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements''), ...
(localized sediment packages) are developed in the Heath Formation. They overlap each other in an east-to-west sequence which extends over a distance of 160 km in the Central Montana Trough. The only exposed portions of the sequence are found at Potter Creek
Dome
A dome () is an architectural element similar to the hollow upper half of a sphere. There is significant overlap with the term cupola, which may also refer to a dome or a structure on top of a dome. The precise definition of a dome has been a m ...
, a small uplifted area northeast of the
Big Snowy Mountains
The Big Snowy Mountains ( ats, níichʔibííkʔa, lit=it is never summer) are a small mountain range south of Lewistown in Fergus County, Montana. Considerably east of and isolated from the main crest of the Northern Rockies, they are one of the ...
.
The last few limestone lenses form a large portion of the Upper Heath Formation,
which is sometimes termed the Bear Gulch Member in recognition of the most well-exposed and fossiliferous lens in the sequence.
This lens, the Bear Gulch Limestone, was also one of the last in the sequence, only succeeded by the Surenough Beds immediately west of it.
The Bear Gulch Limestone can be observed in numerous outcrops, spread out over an area of more than 50 km2 in
Fergus County, Montana
Fergus County is a county in the U.S. state of Montana. As of the 2020 census, the population was 11,446. Its county seat is Lewistown. The county was founded in 1885 and named for James Fergus, a Montana politician who was instrumental in cr ...
.
The creation of limestone lenses in the Heath Formation has been linked to tectonic activity extending the seaway by excavating
bays out of the surrounding land. As old bays are filled in and buried by sediment, faulting and seismic events form new bays in a long eastward to westward succession.
It may have taken a mere 1000 years for the bay responsible for the Bear Gulch Limestone lens to fill in completely, after only 25,000 years for the entire bay formation sequence to run its course across Montana.
The final limestone deposits in the area were succeeded by freshwater lake sediments of the
Cameron Creek Formation, the oldest unit of the early Pennsylvanian-age
Amsden Group.
Age
A wide variety of
biostratigraphic
Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of Bio ...
evidence places the Heath Formation in the upper part of the
Chesterian
The Mississippian ( , also known as Lower Carboniferous or Early Carboniferous) is a subperiod in the geologic timescale or a subsystem of the geologic record. It is the earlier of two subperiods of the Carboniferous period lasting from roughly ...
stage, near the end of the Mississippian subperiod (early
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
period). The upper Chesterian is the North American regional equivalent to the global Serpukhovian stage, as well as the lower part of the European
Namurian
The Namurian is a stage in the regional stratigraphy of northwest Europe with an age between roughly 326 and 313 Ma (million years ago). It is a subdivision of the Carboniferous system or period and the regional Silesian series. The Namurian is ...
stage. The Bear Gulch Limestone in particular is correlated with
ammonoid
Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish) ...
zone E2b (middle Arnsbergian) of the Namurian,
not far from the top of the Serpukhovian. Strata from zone E2b in
England
England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
and the
Czech Republic
The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
have been dated to around 324 million years ago.
Climate
During the time of deposition, the area was about 10-12 degrees north of the
equator
The equator is a circle of latitude, about in circumference, that divides Earth into the Northern and Southern hemispheres. It is an imaginary line located at 0 degrees latitude, halfway between the North and South poles. The term can als ...
, on the boundary between the arid
subtropics
The subtropical zones or subtropics are geographical and climate zones to the north and south of the tropics. Geographically part of the temperate zones of both hemispheres, they cover the middle latitudes from to approximately 35° north and ...
and tropical equatorial region. Cyclical deposition supports a climate model arguing that the overall climate was warm and
monsoon
A monsoon () is traditionally a seasonal reversing wind accompanied by corresponding changes in precipitation but is now used to describe seasonal changes in atmospheric circulation and precipitation associated with annual latitudinal oscil ...
al, with pronounced rainy and dry seasons. During the quiet
dry season
The dry season is a yearly period of low rainfall, especially in the tropics. The weather in the tropics is dominated by the tropical rain belt, which moves from the northern to the southern tropics and back over the course of the year. The te ...
, sedimentation was low and the basin would have been influenced by northeasterly
trade winds
The trade winds or easterlies are the permanent east-to-west prevailing winds that flow in the Earth's equatorial region. The trade winds blow mainly from the northeast in the Northern Hemisphere and from the southeast in the Southern Hemisph ...
,
tidal
Tidal is the adjectival form of tide.
Tidal may also refer to:
* ''Tidal'' (album), a 1996 album by Fiona Apple
* Tidal (king), a king involved in the Battle of the Vale of Siddim
* TidalCycles, a live coding environment for music
* Tidal (servic ...
currents, and
evaporation
Evaporation is a type of vaporization that occurs on the surface of a liquid as it changes into the gas phase. High concentration of the evaporating substance in the surrounding gas significantly slows down evaporation, such as when humidi ...
. Early in the
rainy season
The rainy season is the time of year when most of a region's average annual rainfall occurs.
Rainy Season may also refer to:
* ''Rainy Season'' (short story), a 1989 short horror story by Stephen King
* "Rainy Season", a 2018 song by Monni
* ''T ...
, warmer temperatures and reduced trade winds would have isolated the basin and increased its overall
salinity
Salinity () is the saltiness or amount of salt dissolved in a body of water, called saline water (see also soil salinity). It is usually measured in g/L or g/kg (grams of salt per liter/kilogram of water; the latter is dimensionless and equal ...
. As the rainy season progressed, heavy rainfall enhances the bay's horizontal salinity gradient, from the freshwater-influenced upper bay to the marine-influenced lower bay. Storms would also produce a shallow layer of freshwater, washing sediment and organic material from the basin margins down into deeper areas.
Sedimentology and ecosystems
Fine-grained
lithographic limestone
Lithographic limestone is hard limestone that is sufficiently fine-grained, homogeneous and defect free to be used for lithography.
Geologists use the term "lithographic texture" to refer to a grain size under 1/250 mm.
The term "sublithog ...
(
plattenkalk
Plattenkalk is a very finely grained limestone chemically precipitated in a stratified water column under conditions where bioturbation does not occur. The reasons for the quiet depositional environment and the processes of sediment accumulation v ...
) is predominant in the Bear Gulch lens, though
clay
Clay is a type of fine-grained natural soil material containing clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4).
Clays develop plasticity when wet, due to a molecular film of water surrounding the clay par ...
to
silt
Silt is granular material of a size between sand and clay and composed mostly of broken grains of quartz. Silt may occur as a soil (often mixed with sand or clay) or as sediment mixed in suspension with water. Silt usually has a floury feel when ...
-sized
siliclastic material also forms a significant portion of rock in some areas. Several
facies
In geology, a facies ( , ; same pronunciation and spelling in the plural) is a body of rock with specified characteristics, which can be any observable attribute of rocks (such as their overall appearance, composition, or condition of formatio ...
(sediment associations) are developed in different parts of the bay.
The thick and fossiliferous central basin facies develop along the main northwest-to-southeast axis of the bay, which is delimited by small
dewatering
Dewatering is the removal of water from a location. This may be done by wet classification, centrifugation, filtration, or similar solid-liquid separation processes, such as removal of residual liquid from a filter cake by a filter press as part ...
microfaults. Well-preserved fish and other
nekton
Nekton or necton (from the ) refers to the actively swimming aquatic organisms in a body of water. The term was proposed by German biologist Ernst Haeckel to differentiate between the active swimmers in a body of water, and the passive organisms ...
ic (free-swimming) animals are common, but
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
and typical
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
(seabed-living) animals are very rare. Central basin sedimentation was cyclical, with thin beds of pale
shale
Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especial ...
interbedded between thicker layers of dark, organic-rich
limestone
Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
with a distinct oily smell.
Each dark layer contains a sequence of sediments: dense, massive (
homogenous
Homogeneity and heterogeneity are concepts often used in the sciences and statistics relating to the uniformity of a substance or organism. A material or image that is homogeneous is uniform in composition or character (i.e. color, shape, siz ...
) beds followed by
laminae with
graded bedding
In geology, a graded bed is one characterized by a systematic change in grain or clast size from one side of the bed to the other. Most commonly this takes the form of normal grading, with coarser sediments at the base, which grade upward into pro ...
, which may be
bioturbated in the upper part of the layer. These small sequences, sometimes termed microturbidites, draw comparisons to the
Bouma sequence
300px, thumbnail, Turbidite from the Devonian-age Becke-Oese Sandstone">Devonian.html" ;"title="Turbidite from the Devonian">Turbidite from the Devonian-age Becke-Oese Sandstone, Germany showing a complete Bouma sequence.
The Bouma Sequence (afte ...
s of deep marine
turbidite
A turbidite is the geologic deposit of a turbidity current, which is a type of amalgamation of fluidal and sediment gravity flow responsible for distributing vast amounts of clastic sediment into the deep ocean.
Sequencing
Turbidites were ...
deposits. The pale shale layers of the central basin were likely slowly accumulated during the dry season, while the darker layers correspond to settling bay margin sediments washed into the basin during the rainy season.
Sedimentary
slumps along the central basin axis have been used to reconstruct flow within the basin, indicating that it deepened and drained to the northeast.
The Bear Gulch Limestone reached a thickness of 30 meters at its eastern outlet,
and total basin depth was approximately 40 meters.
At its eastern outlet and nearby sheltered alcoves, the central basin transitions into the lightly colored laminated sediments of the ''
Arborispongia''-
productid
Productida is an extinct order of brachiopods in the extinct class Strophomenata. Members of Productida first appeared during the Silurian. They represented the most abundant group of brachiopods during the Permian period, accounting for 45-70% ...
facies. They are characterized by reefs and patches of the arborescent (branching)
sponge
Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
''Arborispongia'', which is commonly covered with spiny productid
brachiopod
Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, w ...
s. Marine benthic organisms, such as algae,
bivalves
Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bival ...
,
bryozoa
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
ns, and
crinoid
Crinoids are marine animals that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their adult form are commonly called sea lilies, while the unstalked forms are called feather stars or comatulids, which are ...
s, make up a significant portion of the ecosystem in these few areas.
At the calm and shallow northwest rim of the basin, ''Arborispongia'' reefs, large
stromatolite
Stromatolites () or stromatoliths () are layered sedimentary formations (microbialite) that are created mainly by photosynthetic microorganisms such as cyanobacteria, sulfate-reducing bacteria, and Pseudomonadota (formerly proteobacteria). The ...
s, and plant debris can be observed. Salt crystal casts are abundant, indicative of
hypersalinity and high rates of evaporation in this area.
As the bay was filled in by sediments, similar conditions extended into the center of the basin. This results in deposition of the “shallow” or “marly” facies, a collection of very pale
micrite
Micrite is a limestone constituent formed of calcareous particles ranging in diameter up to four μm formed by the recrystallization of lime mud. Flügel, Erik, ''Microfacies of Carbonate Rocks: Analysis, Interpretation and Application,'' Springe ...
s with very little organic or siliciclastic material. They preserve various
algal laminations, diverse ''Arborispongia'' reefs, and local concentrations of
chert
Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a prec ...
and
gypsum
Gypsum is a soft sulfate mineral composed of calcium sulfate dihydrate, with the chemical formula . It is widely mined and is used as a fertilizer and as the main constituent in many forms of plaster, blackboard or sidewalk chalk, and drywall. ...
nodules.
The end of the bay's lifespan was indicated by a sequence of layers with leaf fragments, limestone conglomerates,
marl
Marl is an earthy material rich in carbonate minerals, clays, and silt. When hardened into rock, this becomes marlstone. It is formed in marine or freshwater environments, often through the activities of algae.
Marl makes up the lower part o ...
s, and finally fully freshwater sediments.
When the bay was in its heyday, the south edge of the central basin axis saw the development of the filamentous algal facies. These are mostly dark, organic-rich limestones and shales, similar to the central basin facies though with more siliciclastic silt and fewer microturbidites. As the name indicates, strands of filamentous algae are abundant.
Further beyond the main basin axis, the rocks are even more silty, not clearly bedded, and have a very dark coloration. These sediments, the marginal facies, have a high content of
peloids
Peloids are allochems that are composed of micrite, irrespective of size, shape, or origin. The two primary types of peloids are pellets and intraclasts
Intraclasts are irregularly shaped grains that form by syndepositional erosion of partially ...
, plant debris, and other organic material. They likely correspond to shallow, brackish areas with an influx of freshwater. Fossils of fully saltwater taxa are rare and poorly preserved relative to other environments, though organisms with a wider range of salinity tolerance (''
Acanthodes
''Acanthodes'' (from el, ἄκανθώδης , 'provided with spines') is an extinct genus of spiny shark. Fossils have been found in Europe, North America, Asia, and Australia. ''Acanthodes'' was most common in the Carboniferous and Early Perm ...
'',
gastropods
The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda ().
This class comprises snails and slugs from saltwater, from freshwater, and from land. Ther ...
, filamentous algae) are fairly common.
Preservation of fossils
The Bear Gulch Limestone is a
konservat lagerstätte, meaning that its fossils are uniquely well-preserved, including soft tissue details which offer rare insights into the biology of Carboniferous organisms.
The fine-grained sediments common in the formation allow for fossilized structures to retain fine resolution, as seen in equivalent plattenkalk-based lagerstätten throughout geological history. Most fossils are isolated and flattened into very thin films between sheet-like layers. ''Arborispongia'' assemblages,
cephalopod
A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
shells, and large vertebrate bones occasionally project through several thin layers, approaching three-dimensional preservation.
Vertebrate
Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
s typically come in the form of complete skeletons, and their fossils often preserve internal and external organic
pigments
A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compou ...
. Skin pigments can indicate color patterns and the extent of fins and other external structures, while internal pigments are used to outline the
liver
The liver is a major Organ (anatomy), organ only found in vertebrates which performs many essential biological functions such as detoxification of the organism, and the Protein biosynthesis, synthesis of proteins and biochemicals necessary for ...
,
spleen
The spleen is an organ found in almost all vertebrates. Similar in structure to a large lymph node, it acts primarily as a blood filter. The word spleen comes . , eyes, and other organs. In rare cases, even the structure of fragile
blood vessel
The blood vessels are the components of the circulatory system that transport blood throughout the human body. These vessels transport blood cells, nutrients, and oxygen to the tissues of the body. They also take waste and carbon dioxide away ...
s can be recognized from preserved blood pigments.
Uncommon gut contents indicate the diet of some Bear Gulch animals,
and
phosphatized Phosphatic fossilization has occurred in unusual circumstances to preserve some extremely high-resolution microfossils in which careful preparation can even reveal preserved cellular structures. Such microscopic fossils are only visible under the sc ...
muscles have been found in very rare situations.
Soft
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s are indicated by a variety of
molds, casts, and organic discolorations on rocks.
In the central basin, fossils of invertebrates with
calcareous
Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines.
In zoology
''Calcareous'' is used as an adje ...
shells are mostly dissolved, leaving only molds in the surrounding rock. On the other hand, invertebrates with
chitinous
Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
or
phosphate
In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid .
The phosphate or orthophosphate ion is derived from phospho ...
-rich shells become more common in the central basin. This may have been a consequence of acidity in the pore fluids of sediments, with less acidic fluids favoring the preservation of calcareous fossils and more acidic fluids favoring phosphatized fossils.
There is some uncertainty over how this exceptional preservation was achieved. Most fossils are complete and undecomposed animals, with no signs of disturbance from
scavenger
Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding b ...
s or strong currents. To prevent
decomposition
Decomposition or rot is the process by which dead organic substances are broken down into simpler organic or inorganic matter such as carbon dioxide, water, simple sugars and mineral salts. The process is a part of the nutrient cycle and is e ...
of fragile soft tissue in a warm environment, death and burial had to have been very rapid for the vast majority of articulated skeletons. Rare disarticulated fragments may correspond to large or buoyant carcasses which rise to the water surface to gradually decay and fall apart in a “bloat and float”
taphonomic
Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov t ...
process.
Fossils are dispersed throughout the Bear Gulch lens, rather than concentrated in specific fossil-rich beds (which would be expected if organisms were killed by
algal bloom
An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term ''algae'' encompas ...
s).
The cyclically deposited seabed of the central basin, though fossiliferous, is deprived of benthic invertebrates. Some authors have suggested that the deepest waters in the basin were too salty or oxygen-poor for most life.
However, bottom-dwelling fish are common in the central basin, suggesting that
anoxia (low oxygen) was not a persistent quality of bottom waters.
Nevertheless, many fish fossils are found with distended gills, favoring
asphyxiation
Asphyxia or asphyxiation is a condition of deficient supply of oxygen to the body which arises from abnormal breathing. Asphyxia causes generalized hypoxia, which affects primarily the tissues and organs. There are many circumstances that can i ...
as a cause of death.
One possible explanation for rapid asphyxiation and burial places blame on freshwater runoff during storms in the wet season. Heavy rainfall would bring a cascade of organic-rich marginal sediments into the central basin. As the sediments sink, they quickly absorb oxygen from the seawater, killing and burying basin organisms in only a matter of hours.
Paleobiota
The
ecosystem
An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
represented by the Bear Gulch Limestone was vibrant and well-represented by fossils, with many described and undescribed species of vertebrates and invertebrates among their ranks. Animals show a wide range of body types and show preferences to certain areas of the bay, indicating that the bay offered a wide range of habitats and
niches to be filled.
Many indicators of
paleoecology
Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...
and
paleobiology
Paleobiology (or palaeobiology) is an interdisciplinary field that combines the methods and findings found in both the earth sciences and the life sciences. Paleobiology is not to be confused with geobiology, which focuses more on the interactio ...
have been preserved, from associations between encrusting shelled organisms and algae,
to gut contents and other feeding traces,
and even some animals fossilized while mating.
Fish are diverse and abundant, with thousands of specimens representing approximately 150 species (as of 2015).
Many of these species are undescribed and unnamed, only mentioned through "code names"; unnamed species are not included in paleobiota lists here.
Chondrichthyans
The Bear Gulch Limestone is renowned for its unusual and ecologically diverse
chondrichthyans
Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. C ...
, the group of cartilaginous fish containing modern
shark
Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s,
rays
Ray may refer to:
Fish
* Ray (fish), any cartilaginous fish of the superorder Batoidea
* Ray (fish fin anatomy), a bony or horny spine on a fin
Science and mathematics
* Ray (geometry), half of a line proceeding from an initial point
* Ray (gra ...
, and
chimaera
Chimaeras are cartilaginous fish in the order Chimaeriformes , known informally as ghost sharks, rat fish, spookfish, or rabbit fish; the last three names are not to be confused with rattails, Opisthoproctidae, or Siganidae, respectively.
At ...
s. Over 80 species of chondrichthyans are known (as of 2012), representing close to 60% of the fish diversity (by species count) in the bay. This is a far cry from modern ecosystems, where chondrichthyans comprise only around 3% of fish species.
In the Bear Gulch Limestone, chondrichthyans possess a wide range of ecologies and body types, including
eel
Eels are ray-finned fish belonging to the order Anguilliformes (), which consists of eight suborders, 19 families, 111 genera, and about 800 species. Eels undergo considerable development from the early larval stage to the eventual adult stage ...
-like forms ("''
Thrinacoselache
''Thrinacodus'' is an extinct genus of basal elasmobranch, found worldwide from the Late Devonian-Lower Carboniferous. Most species are only known from their tricuspid teeth. ''T. gracia'', originally placed in the separate genus ''Thrinacoselach ...
''"), deep-bodied
durophages (
petalodontiforms), small active swimmers (most
holocephalans
Holocephali ("complete heads"), sometimes given the term Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. The earliest fossils are of teeth and come from the Devonian period. Little is known about these primit ...
), ray-like
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
forms (''
Squatinactis
''Squatinactis'' is a genus of extinct Elasmobranchii, elasmobranch Chondrichthyes known from the Carboniferous aged Bear Gulch Limestone, Bear gulch limestone in Montana. This fish was discovered in 1974 by Richard Lund. The Holotype, type speci ...
'') and more typical shark-like predators (
symoriiforms and
elasmobranchs
Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gil ...
).
Chondrichthyan diversity and abundance patterns are correlated, with the highest number of species and fossils found in the central basin facies and the lowest in the marginal facies. Large predatory chondrichthyans such as ''
Listracanthus
''Listracanthus'' is a genus of extinct chondrichthyan with uncertain affinities. Species of ''Listracanthus'' are known primarily from their tremendous, feather-like denticles, which range up to four inches in length. The denticles had a large ...
'' and
symoriiforms (except ''Falcatus'') make up a higher portion of fossils in the hypersaline upper bay, while ''
Falcatus
''Falcatus'' is an extinct genus of falcatid chondrichthyan which lived during the early Carboniferous Period in Bear Gulch bay in what is now Montana.
Description
This fish was quite small, only getting to around 25–30 cm or 10-12 i ...
'' and
holocephalans
Holocephali ("complete heads"), sometimes given the term Euchondrocephali, is a subclass of cartilaginous fish in the class Chondrichthyes. The earliest fossils are of teeth and come from the Devonian period. Little is known about these primit ...
are the most common fossils in the marginal and central basin facies. A sheltered reef habitat at the eastern edge of exposures has produced a high number of ''
Heteropetalus'' fossils. Regardless, each habitat observes significant species overlap with other habitats, and "common" chondrichthyans (>3 specimens) are not abundant to the point that they outnumber more rare species (<3 specimens), in terms of total specimen count.
Actinopterygians
Actinopterygians
Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species.
The ray-finned fishes are so called because their fins are webs of skin supported by bony or hor ...
(ray-finned bony fish) are also quite diverse, with about 50 described and undescribed species (as of 2012) making up more than a third of the Bear Gulch fish diversity. In terms of specimen count, they are the most abundant fish fossils in the Bear Gulch Limestone, representing approximately 80% of all fossil vertebrates.
However, they are more restricted than chondrichthyans in terms of
ecomorphology Ecomorphology or ecological morphology is the study of the relationship between the ecological role of an individual and its morphological adaptations. The term "morphological" here is in the anatomical context. Both the morphology and ecology exh ...
. Most have a small and generalized
fusiform
Fusiform means having a spindle-like shape that is wide in the middle and tapers at both ends. It is similar to the lemon-shape, but often implies a focal broadening of a structure that continues from one or both ends, such as an aneurysm on a b ...
(smoothly tapering) body shape, with only a few deviations towards more elongated or deep-bodied forms.
The most abundant actinopterygians include ''
Wendyichthys'' and three closely-related unnamed species (code names "Yogo", "Fub", and "Cop"). The relative abundance of actinopterygians drops off in the upper bay, though they are still fairly common. Schools of freshly hatched
larvae
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
The ...
have been found in isolated reef habitats near the central basin.
Other fish
Non-actinopterygian and non-chondrichthyan fish make up a fairly small component of the overall diversity, though their fossils are still common in some habitats. Preserved
sarcopterygians
Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includin ...
(lobe-finned fish) include
actinistia
The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
ns (
coelacanth
The coelacanths ( ) are fish belonging to the order Actinistia that includes two extant species in the genus ''Latimeria'': the West Indian Ocean coelacanth (''Latimeria chalumnae''), primarily found near the Comoro Islands off the east coast ...
s) and the
rhizodont
Rhizodontida is an extinct group of predatory tetrapodomorphs known from many areas of the world from the Givetian through to the Pennsylvanian - the earliest known species is about 377 million years ago (Mya), the latest around 310 Mya. Rhizodo ...
''
Strepsodus
''Strepsodus'' is a genus of rhizodont lobe-finned fish
Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. T ...
'', which have very different habitat preferences. Coelacanths, particularly fossils of the genus ''
Caridosuctor'', are abundant in the central basin and its margins, but are practically absent in the upper bay. ''Strepsodus'', on the other hand, is moderately common in the upper bay and very rare elsewhere. Neither are common in the sheltered eastern reef habitat. ''
Acanthodes
''Acanthodes'' (from el, ἄκανθώδης , 'provided with spines') is an extinct genus of spiny shark. Fossils have been found in Europe, North America, Asia, and Australia. ''Acanthodes'' was most common in the Carboniferous and Early Perm ...
'', the only
acanthodian
Acanthodii or acanthodians is an extinct class of gnathostomes (jawed fishes), typically considered a paraphyletic group. They are currently considered to represent a grade of various fish lineages leading up to the extant Chondrichthyes, which ...
known from the Bear Gulch Limestone, has a similar distribution pattern to ''Strepsodus'': common in the upper bay and nowhere else.
Lamprey
Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s, represented by ''
Hardistiella'', are too rare to estimate their habitat preference.
Arthropods
Arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s are diverse, the most abundant and speciose being various early
hoplocarids, relatives of modern stomatopods (
mantis shrimp
Mantis shrimp, or stomatopods, are carnivorous marine crustaceans of the order Stomatopoda (). Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around in lengt ...
s).
Other arthropods include an assortment of typical Carboniferous
crustacean
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s,
rare
xiphosura
Xiphosura () is an order of arthropods related to arachnids. They are more commonly known as horseshoe crabs (a name applied more specifically to the only extant family, Horseshoe crab, Limulidae). They first appeared in the Hirnantian (Late Ordo ...
ns (horseshoe crabs),
and a single "
synziphosurine" (''
Anderella'').
Trilobite
Trilobites (; meaning "three lobes") are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest-known groups of arthropods. The first appearance of trilobites in the fossil record defines the base of the At ...
s, though common in adjacent formations, are practically absent in the Bear Gulch Limestone.
Molluscs
Worms
Other invertebrates
See also
*
List of types of limestone
This is a list of types of limestone arranged according to location. It includes both formal stratigraphic unit names and less formal designations.
Africa Egypt
* Tura limestone, used for the Great Pyramid casing stones
* Mokattam limestone; ...
External links
Personal website of Richard Lund and Eileen Grogan, exhibiting the Bear Gulch Limestone and its fossils1998 Discovery Channel documentary "Prehistoric Sharks", featuring a segment on Bear Gulch (4:36-16:13)Segment of the 2020 PBS documentary "Prehistoric Road Trip", focusing on the Bear Gulch fossils (12:16-22:53)
References
{{reflist
Carboniferous Montana
Carboniferous paleontological sites
Lagerstätten
Limestone formations of the United States
Mississippian United States
Serpukhovian
Paleontology in Montana
Carboniferous System of North America