Boii (genus)
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''Boii'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
microsaur Microsauria 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 considered paraphyletic, as s ...
within the
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Tuditanidae Tuditanidae is an extinct family (biology), family of microsaurian tetrapods. 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 rese ...
. It was found in
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
coal from mines near the community of Kounov in the
Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Central Europe. The country is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the south ...
. 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 ...
, which were similar to microsaurian elements originally referred to ''
Asaphestera ''Asaphestera'' is an extinct genus of a synapsid 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 ...
''. ''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 A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
, ''Boii crassidens'', was considered to be a species of ''
Sparodus ''Sparodus'' is an extinct genus of microsaur within the family Gymnarthridae Gymnarthridae is an extinct family of the group Recumbirostra. Gymnarthrids are known from Europe and North America and existed from the Late Carboniferous through t ...
'', 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 ( ; ; ) is the westernmost and largest historical region of the Czech Republic. In a narrow, geographic sense, it roughly encompasses the territories of present-day Czechia that fall within the Elbe River's drainage basin, but historic ...
(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 coal seam, ...
) mines near the localities of
Nýřany Nýřany (; ) is a town in Plzeň-North District in the Plzeň Region of the Czech Republic. It has about 7,000 inhabitants. Administrative division Nýřany consists of three municipal parts (in brackets population according to the 2021 census): ...
and Kounová. His list included short preliminary descriptions for many new genera and species of tetrapods, including ''
Microbrachis ''Microbrachis'' is an extinct genus of microsaurian tetrapod from the Carboniferous Kladno Formation of the Czech Republic. Description ''Microbrachis'' was an elongated, salamander-like creature, about long, with over 40 vertebrae instea ...
,
Branchiosaurus ''Branchiosaurus'' (from , 'gill' and , 'lizard') is a genus of small, prehistoric amphibians. Fossils have been discovered in strata dating from the late Pennsylvanian Epoch to the Permian Period. The taxa may be invalid; the material refe ...
,
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 Gymnarthridae is an extinct family of the group Recumbirostra. Gymnarthrids are known from Europe and North America and existed from the Late Carboniferous through t ...
.'' 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, large ...
" shwarzenbergii'', which is now known to not be an amphibian at all, but rather an early
synapsid Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida (which includes all extant rept ...
named ''
Macromerion ''Macromerion'' (Macro- is Greek/Latin for “large”) is an extinct genus of non-mammalian synapsids, specifically Pelycosaurs, in the family Sphenacodontidae from Late Carboniferous deposits in the Czech Republic.Paton, R. L. 1974. ''Lower Per ...
''. 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 fo ...
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 can be 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 the United States, most nati ...
in
Prague Prague ( ; ) is the capital and List of cities and towns in the Czech Republic, largest city of the Czech Republic and the historical capital of Bohemia. Prague, located on the Vltava River, has a population of about 1.4 million, while its P ...
, 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 Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...
n species of
catfish Catfish (or catfishes; order (biology), order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Catfish are common name, named for their prominent barbel (anatomy), barbels, which resemble a cat's whiskers, though not ...
. Frič also assigned an isolated
maxilla In vertebrates, the maxilla (: maxillae ) 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 two maxil ...
(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 language, Latin plural, singular ''Boius''; ) were a Celts, Celtic tribe of the later Iron Age, attested at various times in Cisalpine Gaul (present-day Northern Italy), Pannonia (present-day Austria and Hungary), present-day Ba ...
tribe which inhabited the area of Bohemia during the time of the
Roman Republic The Roman Republic ( ) was the era of Ancient Rome, classical Roman civilisation beginning with Overthrow of the Roman monarchy, the overthrow of the Roman Kingdom (traditionally dated to 509 BC) and ending in 27 BC with the establis ...
.


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 Canada balsam, also called Canada turpentine or balsam of fir, is the oleoresin of the balsam fir tree (''Abies balsamea'') of boreal North America. The resin, dissolved in essential oils, is a viscous, sticky, colourless or yellowish liquid t ...
. 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: 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 of all kno ...
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 mammals h ...
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 ''Tuditanus'' is an extinct genus of tuditanid microsaur from the Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years ...
''. 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 (Latin; : ''palatini''; cf. derivative spellings below) is a high-level official attached to imperial or royal courts in Europe since Roman Empire, 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 (; ) 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 maxillary bones. The vomer forms ...
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 (; derived from the Latin ''palatum'') are two irregular bones of the facial skeleton in many animal specie ...
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 (, : 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 o ...
(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 A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is not ...
. 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, collarbone, or keybone is a slender, S-shaped long bone approximately long that serves as a strut between the scapula, shoulder blade and the sternum (breastbone). There are two clavicles, one on each side of the body. The clavic ...
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 ...
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 (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
(according to Frič, 1883) or a
radius In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
or
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
(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 ...
, along with other species of ''Sparodus''. Branchiosaurids, a group of small gilled
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
, 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 Sco ...
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 synapsid 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 ...
'' (which itself was similar to ''
Tuditanus ''Tuditanus'' is an extinct genus of tuditanid microsaur from the Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years ...
), 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