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''Boii'' is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
microsaur Microsauria ("small lizards") is an extinct, possibly polyphyletic order of tetrapods from the late Carboniferous and early Permian periods. It is the most diverse and species-rich group of lepospondyls. Recently, Microsauria has been consider ...
within the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
Tuditanidae Tuditanidae is an extinct family of tuditanomorph microsaurs. Fossils have been found from Nova Scotia, Ohio, and the Czech Republic and are Late Carboniferous in age. Tuditanids were medium-sized terrestrial microsaurs that resembled lizards. ...
. It was found in
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 ...
coal from mines near the community of Kounov in 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 ...
. The only remains of the genus consist of a crushed skull, shoulder girdle bones, and
scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number w ...
, which were similar to microsaurian elements originally referred to ''
Asaphestera ''Asaphestera'' is an extinct genus of a tetrapod described on the basis of fossils from the Carboniferous of the Joggins locality in Nova Scotia, Canada. It was originally described as an undetermined lepospondyl and subsequently classified as a ...
''. ''Boii'' can be characterized by its heavily sculptured skull, thin ventral plate of the clavicles, and a larger number of fangs on the roof of the mouth. For many years the type and only known
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''Boii crassidens'', was considered to be a species of ''
Sparodus ''Sparodus'' is an extinct genus of microsaur within the family Gymnarthridae. See also * Prehistoric amphibian * List of prehistoric amphibians This list of prehistoric amphibians is an attempt to create a comprehensive listing of all genera ...
'', until 1966 when Robert Carroll assigned it to its own genus.


History

The specimen now designated as ''Boii crassidens'' was first described by Antonin Frič, one of the most notable paleontologists in the late 19th century region of
Bohemia Bohemia ( ; cs, Čechy ; ; hsb, Čěska; szl, Czechy) is the westernmost and largest historical region of the Czech Republic. Bohemia can also refer to a wider area consisting of the historical Lands of the Bohemian Crown ruled by the Bohem ...
(now part of the Czech Republic). Frič made many contributions to knowledge of Carboniferous tetrapods during his lifetime, including a particular article published in "''Sitzungsberichte der königlichen Böhmischen Gesellschaft der Wissenschaften in Prague''", the most notable bohemian scientific journal of its day. The article, which was published in 1876 within volume 1875 of the journal, was a list of Carboniferous animals he and his associates recently discovered at gaskohle (
bituminous coal Bituminous coal, or black coal, is a type of coal containing a tar-like substance called bitumen or asphalt. Its coloration can be black or sometimes dark brown; often there are well-defined bands of bright and dull material within the seams. It ...
) mines near the localities of
Nýřany Nýřany (; german: Nürschan) is a town in Plzeň-North District in the Plzeň Region of the Czech Republic. It has about 6,900 inhabitants. Administrative parts Villages of Doubrava and Kamenný Újezd are administrative parts of Nýřany. Geo ...
and Kounová. His list included short preliminary descriptions for many new genera and species of tetrapods, including ''
Microbrachis ''Microbrachis'' is an extinct genus of lepospondyl amphibian from the Carboniferous Kladno Formation of the Czech Republic. Description ''Microbrachis'' was an elongated, salamander-like creature, about long, with over 40 vertebrae ins ...
,
Branchiosaurus ''Branchiosaurus'' (from el, βράγχιον , 'gill' and el, σαῦρος , 'lizard') is a genus of small, lightly built early prehistoric amphibians. Fossils have been discovered in strata dating from the late Pennsylvanian Epoch to the ...
,
Hyloplesion ''Hyloplesion'' is an extinct genus of microbrachomorph microsaur. It is the type and only genus within the family Hyloplesiontidae. Fossils have been found from the Czech Republic near the towns of Plzeň, Nýřany, and Třemošná, and da ...
'' (at that time called ''Stelliosaurus''), and ''
Sparodus ''Sparodus'' is an extinct genus of microsaur within the family Gymnarthridae. See also * Prehistoric amphibian * List of prehistoric amphibians This list of prehistoric amphibians is an attempt to create a comprehensive listing of all genera ...
.'' Most of the "Saurier" (reptiles, amphibians, or other tetrapods) he discussed in the article hailed from the Nýřany mines, with only two diagnostic examples found within the Kounová mines. One of these was a supposed amphibian he designated as ''"
Labyrinthodon ''Mastodonsaurus'' (meaning "teat tooth lizard") is an extinct genus of temnospondyl amphibian from the Middle Triassic of Europe. It belongs to a Triassic group of temnospondyls called Capitosauria, characterized by their large body size and pre ...
" shwarzenbergii'', which is now known to not be an amphibian at all, but rather an early
synapsid Synapsids + (, 'arch') > () "having a fused arch"; synonymous with ''theropsids'' (Greek, "beast-face") are one of the two major groups of animals that evolved from basal amniotes, the other being the sauropsids, the group that includes reptil ...
named '' Macromerion''. The other specimen, which is now known as ''Boii crassidens'', he named ''Batrachocephalus crassidens''. This specimen consisted of a crushed skull, preserved on a slab and
counterslab A compression fossil is a fossil preserved in sedimentary rock that has undergone physical compression. While it is uncommon to find animals preserved as good compression fossils, it is very common to find plants preserved this way. The reason f ...
of coal. Jaw bones, scales, and shoulder girdle elements were preserved on the slabs as well. It is currently held at the
National Museum A national museum is a museum maintained and funded by a national government. In many countries it denotes a museum run by the central government, while other museums are run by regional or local governments. In other countries a much greater numb ...
in
Prague Prague ( ; cs, Praha ; german: Prag, ; la, Praga) is the capital and largest city in the Czech Republic, and the historical capital of Bohemia. On the Vltava river, Prague is home to about 1.3 million people. The city has a temperate ...
, with the designation ČGH 83. This article was only a preliminary review of the creatures Frič and his associates discovered. A more elaborate description was published in 1883 as part of a personal monograph focusing solely on the creatures discovered at these mines. This description renamed ''Batrachocephalus crassidens'' to ''Sparodus crassidens'', as the genus name Batrachocephalus''' was already taken by '' Batrachocephalus mino'', an
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
n species of
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...
. Frič also assigned an isolated
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
(upper jaw bone) with designation ČGH 124 to this species, although it was quite a bit larger than the maxilla of the crushed skull. In a 1966 review of microsaurs published by Robert Carroll, ''"Sparodus" crassidens'' was not found to be a member of the genus ''Sparodus''. Instead, it was re-evaluated as belonging to a family of early microsaurs known as the Tuditanidae. This prompted Carroll to create a new genus name for the specimens previously considered to belong to ''"Sparodus" crassidens''. The new name he found was ''Boii crassidens'', named after the
Boii The Boii (Latin plural, singular ''Boius''; grc, Βόιοι) were a Celtic tribe of the later Iron Age, attested at various times in Cisalpine Gaul (Northern Italy), Pannonia (Hungary), parts of Bavaria, in and around Bohemia (after whom the ...
tribe which inhabited the area of Bohemia during the time of the
Roman Republic The Roman Republic ( la, Res publica Romana ) was a form of government of Rome and the era of the classical Roman civilization when it was run through public representation of the Roman people. Beginning with the overthrow of the Roman Kin ...
.


Description


Skull

The crushed skull was pressed between two plates of coal which preserved the outer impressions of bones on both the underside and upper side of the specimen. Although not all of the bones were preserved, the outer impressions helped to reconstruct the structure of these missing bones. The impressions were used to reconstruct the skull and lower jaws, while the skull itself (which preserved the palate better than the impressions) was removed and encased in Canada balsam. The skull was initially believed to have been stout, approximately as long as it was wide. This 'frog-like' skull is responsible for the original genus name "''Batrachocephalus''", which is
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
for "frog head". However, later reconstructions from Robert Carroll have interpreted this wide shape as a result of the crushing the skull experienced, with the skull actually being narrow and somewhat triangular, more similar in shape to that of a lizard rather than a frog. The orbits (eye holes) are roughly midway down the length of the skull. Unlike the fairly smooth skulls of most microsaurs, the skull of ''Boii'' is covered with numerous grooves and ridges which radiate from the middle of their respective bones. The maxillary and
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
ry bones forming the edge of the snout contained many conical teeth, about 30 per each side of the upper jaw. This number is somewhat larger than that of the eponymous tuditanid '' Tuditanus''. These marginal (edge) teeth in general are slightly larger towards the front of the skull, although only to a small extent. The palate (roof of the mouth) also possesses teeth. The
palatine A palatine or palatinus (in Latin; plural ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman times.
bones, positioned right next to the maxillae, possess a fair number of teeth. The most notable assortment of palatal teeth are a bundle of large fangs preserved midway down the skull. Large palatal fangs are also shared by ''Sparodus'', explaining how ''Boii crassidens'' was once considered to be part of that genus. Frič (1883) considered these fangs to have grown out of the
vomer The vomer (; lat, vomer, lit=ploughshare) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxill ...
s (from the front of the skull) or possibly the
parasphenoid The parasphenoid is a bone which can be found in the cranium of many vertebrates. It is an unpaired dermal bone which lies at the midline of the roof of the mouth. In many reptiles (including birds), it fuses to the endochondral (cartilage-derived) ...
(from the back of the skull), but Carroll (1966) reconstructed the fangs as being part of the long
pterygoid bone The pterygoid is a paired bone forming part of the palate of many vertebrates, behind the palatine bone In anatomy, the palatine bones () are two irregular bones of the facial skeleton in many animal species, located above the uvula in the th ...
s, which were originally reported as being toothless. The dentaries (main bones of the lower jaws) are also preserved and covered with teeth similar to those of the upper jaws. A row of small pits run from the
symphysis A symphysis (, pl. symphyses) is a fibrocartilaginous fusion between two bones. It is a type of cartilaginous joint, specifically a secondary cartilaginous joint. # A symphysis is an amphiarthrosis, a slightly movable joint. # A growing together ...
(chin) along the upper portion of the outer face of the bones.


Other bones

Some bones of the shoulder girdle are known. A pair of V-shaped bones are preserved behind the skull, which Frič (1883) identified as coracoids. However, research by Carroll has indicated that microsaurs did not possess coracoid bones, and that the bones identified by Frič were actually
clavicle The clavicle, or collarbone, is a slender, S-shaped long bone approximately 6 inches (15 cm) long that serves as a strut between the shoulder blade and the sternum (breastbone). There are two clavicles, one on the left and one on the rig ...
s. Clavicles possess two blade-like regions pointing away from each other at right angles when seen from the front. The lower regions point inwards and laying along the chest and the upper regions point upwards along the sides of the body. The lower regions (a.k.a. ventral plates) of ''Boii'''s clavicles are very thin, akin to those of a reptile. As typical for many tetrapods, ''Boii'' also possessed an
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals ( marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In th ...
which was positioned behind the clavicles at the center of the chest. However, the only remnant of this bone in the specimen is a large yet indistinct impression. A single arm bone was also present on the specimen, although paleontologists disagree whether it was a
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
(according to Frič, 1883) or a
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
or
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
(according to Carroll, 1966). ''Boii'' possessed large scales, as preserved in the only known specimen. The scales of the back of the animal were tile-like with rounded corners. They tightly overlapped in alternating rows, and each possessed a bulging rear edge. The scales of the belly of the animal were also overlapping. However, these belly scales were wider than the back scales, and their rows were stacked so frequently that only a small portion of each individual scale is visible.


Classification

Frič (1883) considered ''Boii crassidens'' to be a member of the family
Branchiosauridae Branchiosauridae is an extinct family of small amphibamiform temnospondyls with external gills and an overall juvenile appearance. The family has been characterized by hundreds of well-preserved specimens from the Permo-Carboniferous of Middle Eu ...
, along with other species of ''Sparodus''. Branchiosaurids, a group of small gilled
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...
, are now believed to be only distant relatives of ''Sparodus'' and ''Boii''. In 1894,
John William Dawson Sir John William Dawson (1820–1899) was a Canadian geologist and university administrator. Life and work John William Dawson was born on 13 October 1820 in Pictou, Nova Scotia, where he attended and graduated from Pictou Academy. Of Scotti ...
listed the three ''Sparodus'' species as microsaurs rather than branchiosaurids. He did not explicitly note each individual species (instead clumping them as "''Sparodus'' sp."), because he was unsure whether they were all valid members of the same genus. Dawson's suspicions were rectified in 1966, when Carroll split ''Boii crassidens'' off of ''Sparodus''. The skull structure of ''Boii'' was considered to be very similar to that ''
Asaphestera ''Asaphestera'' is an extinct genus of a tetrapod described on the basis of fossils from the Carboniferous of the Joggins locality in Nova Scotia, Canada. It was originally described as an undetermined lepospondyl and subsequently classified as a ...
'' (which itself was similar to '' Tuditanus), s''o he placed ''Boii'' along with ''Asaphestera'' in Tuditanidae. Tuditanids were basal microsaurs, which did not evolve the unusual adaptations of more advanced families of microsaurs. They were terrestrial, lizard-like creatures with well-developed legs and jaw joints set about as far back as the neck joint. Although ''Boii crassidens'' was quite old by microsaur standards, it is one of the last known species of tuditanids. Carroll (1966) suggested that it may have been descended from ''Asaphestera'', supposedly one of the earliest known microsaurs to have evolved. However, as of 2020 ''Asaphestera'' has been recognized as a chimeric taxon, based on specimens of a potential eothyridid along with a newly-named microsaur, '' Steenerpeton''.


External links


A low-quality photograph of the type specimen, from "Carboniferous and Permian faunas and their occurrence in the limnic basins of the Czech Republic" by Stanislav Stamberg and Jarislav Zajic (2008)


References

{{Taxonbar, from=Q11415431 Microsauria Carboniferous amphibians of Europe Fossil taxa described in 1966 Taxa named by Robert L. Carroll