''Bassipterus'' ("wing from

Bass Bass or Basses may refer to: Fish * Bass (fish), various saltwater and freshwater species Wood * Bass or basswood, the wood of the tilia americana tree Music * Bass (sound), describing low-frequency sound or one of several instruments in th ...

") is a

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 ...

eurypterid Eurypterids, often informally called sea scorpions, are a group of extinct marine arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period, 467.3 Myr, mil ...

, an extinct group of aquatic

arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...

s. ''Bassipterus'' is classified as part of 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 ...

Adelophthalmidae, the only

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

within the derived ("advanced")

Adelophthalmoidea Adelophthalmidae (the name deriving from the type genus ''Adelophthalmus'', meaning "no obvious Compound eyes, eyes") is a Family (biology), family of eurypterids, an extinct group of aquatic arthropods. Adelophthalmidae is the only family class ...

superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...

of eurypterids. Fossils of the single and

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

, ''B. virgnicus'', have been discovered in deposits of the Late

Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...

age in

West Virginia West Virginia is a mountainous U.S. state, state in the Southern United States, Southern and Mid-Atlantic (United States), Mid-Atlantic regions of the United States.The United States Census Bureau, Census Bureau and the Association of American ...

and

Maryland Maryland ( ) is a U.S. state, state in the Mid-Atlantic (United States), Mid-Atlantic region of the United States. It borders the states of Virginia to its south, West Virginia to its west, Pennsylvania to its north, and Delaware to its east ...

United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...

. The genus is named after Bass, where most of the fossils have been recovered. ''Bassipterus'' was a well-known basal ("primitive") genus that was distinguished from the more derived adelophthalmids by the specialization of its genital operculum (a plate-like segment which contains the genital aperture) and its long and narrow

eyes An eye is a sensory organ that allows an organism to perceive visual information. It detects light and converts it into electro-chemical impulses in neurons (neurones). It is part of an organism's visual system. In higher organisms, the ey ...

, being '' Pittsfordipterus''s closest relative.

Description

''Bassipterus'' is almost completely known, with the first to fifth pair of limbs and type B genital appendage being the only unknown body parts. The body had a lanceolate (lance-shaped) and streamlined shape. The

prosoma The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...

("head") was long, rounded and tray-shaped. The

carapace A carapace is a dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tortoises, the unde ...

(

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...

plate of the head) was surrounded by a narrow marginal rim, with

placed frontally, placed intramarginally (within the margin) and unusually long. PE 6139, a 15.2 millimeter (0.6 inch) long carapace, had eyes 4.5 mm (2 in) long and 2.2 mm (0.1 in) wide. The

ocelli A simple eye or ocellus (sometimes called a pigment pit) is a form of eye or an optical arrangement which has a single lens without the sort of elaborate retina that occurs in most vertebrates. These eyes are called "simple" to distinguish the ...

are preserved, and were placed in the center of the carapace, immediately behind the eyes. The

metastoma The metastoma is a ventral single plate located in the opisthosoma of non-arachnid dekatriatan chelicerates such as eurypterids, chasmataspidids and the genus '' Houia''. The metastoma located between the base of 6th prosomal appendage pair and ...

(a large plate part of the

abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...

) was long, truncated (shortened as by cutting it) at the base and cordate (heart-shaped) in the

anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

(frontal) part. One specimen preserves a metastoma of 8.3 mm (0.3 in) in length and 5 mm (0.2 in) in width. Of the

appendages An appendage (or outgrowth) is an external body part or natural prolongation that protrudes from an organism's body such as an arm or a leg. Protrusions from single-celled bacteria and archaea are known as cell-surface appendages or surface app ...

(limbs), the swimming legs (sixth pair of appendages) are the only known ones. These were of ''

Hughmilleria ''Hughmilleria'' is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of ''Hughmilleria'' have been discovered in deposits of the Silurian age in China and the United States. Classified as part of the basal family Hughmille ...

''-type (with enlarged seventh and eighth leg segments and the ninth very small), but in this

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), ...

, it was serrated along the anterior edge of the sixth to seventh joints. The terminal joint was a sharped spur. The

mesosoma The mesosoma is the middle part of the body, or tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the metasoma. It bears the legs, and, in the case of winged insects, the wings. Wasps, bees and a ...

(comprising

body segment Segmentation in biology is the division of some animal and plant body plans into a linear series of repetitive segments that may or may not be interconnected to each other. This article focuses on the segmentation of animal body plans, specifica ...

s 1 to 6) was tapered, with the first

tergite A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the Anatomical terms of location#Dorsal and ventral, dorsal ('upper') portion of an arthropod segment other than the head. The Anatomical terms of location#Anterior ...

(dorsal part of the segment) considerably narrower than the rest. The

metasoma The metasoma is the posterior part of the body, or tagma (biology), tagma, of arthropods whose body is composed of three parts, the other two being the prosoma and the mesosoma. In insects, it contains most of the digestive tract, respiratory sy ...

(comprising segments 7 to 12) followed the tapering of the mesosoma and ended in a long dirk-like

telson The telson () is the hindmost division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment (biology), segment of the arthropod body, or an additional division that is not a true segm ...

(the posteriormost division of the body). The telson was smooth in the ventral part, but carinated (with a keel) dorsally towards the end, which was a very sharped point. The specimen PE 6208 had a 32.5 mm (1.3 in) long telson. The

ornamentation An ornament is something used for decoration. Ornament may also refer to: Decoration * Ornament (art), any purely decorative element in architecture and the decorative arts * Ornamental turning * Biological ornament, a characteristic of animals ...

in ''Bassipterus'' is well known and developed. The carapace was covered by triangular scales. The tergites and abdominal plates had mucrones (sharped points) typical of Adelophthalmidae, but these were considerably larger and more prominent than in related genera. In some places, the mucrones touched each other and formed a rhombic ornamentation, while in others, they were so elongated that they resembled the linear striations present in the more derived ("advanced")

Adelophthalmus ''Adelophthalmus'' is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of ''Adelophthalmus'' have been discovered in deposits ranging in age from the Early Devonian to the Early Permian, which makes it the longest lived of ...

''. The

morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...

of the genital operculum (a plate-like segment which contains the genital aperture) allows an easy differentiation between ''Bassipterus'' and the other adelophthalmids. The type A genital appendage ( assumed to represent females) was long, reaching the second pair of abdominal plates, and was divided into three joints. The first had a short, triangular and hastate (with protruding lobes) portion followed by a tubular shaft that ended in two lateral triangular projections at the point of union with the following joint. The second joint was shorter and composed of three different areas. On both sides of the central shaft, two long narrow

sclerite A sclerite (Greek language, Greek , ', meaning "hardness, hard") is a hardened body part. In various branches of biology the term is applied to various structures, but not as a rule to vertebrate anatomical features such as bones and teeth. Instea ...

s (hardened parts), possibly plates, lengthened themselves at their ends. The third joint was very short and finished in two long points reminiscent of ''

Eurypterus ''Eurypterus'' ( ) is an extinct genus of eurypterid, a group of organisms commonly called "sea scorpions". The genus lived during the Silurian period, from around 432 to 418 million years ago. ''Eurypterus'' is by far the most well-studied and ...

'', a more basal genus. The appendage in its entirety was covered by triangular mucrones which pointed outwards. The genital operculum differentiates ''Bassipterus'' from other eurypterid genera and suggests a close relationship with '' Pittsfordipterus''.

History of research

''Bassipterus virginicus'' was described by the paleontologists Erik Norman Kjellesvig-Waering and Kenneth Edward Caster based on the

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

(PE 6201, an almost complete specimen), seven

paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...

s and dozens of fragments. Most of the fossils were found in the

Wills Creek Formation Wills Creek Formation is a mapped Silurian bedrock unit in Pennsylvania, Maryland, Virginia, and West Virginia. Description The Wills Creek is defined as a moderately well bedded greenish-gray shale containing local limestone and sandstone zon ...

, as well as an isolated carapace of the same

formation Formation may refer to: Linguistics * Back-formation, the process of creating a new lexeme by removing or affixes * Word formation, the creation of a new word by adding affixes Mathematics and science * Cave formation or speleothem, a secondary ...

Cumberland Cumberland ( ) is an area of North West England which was historically a county. The county was bordered by Northumberland to the north-east, County Durham to the east, Westmorland to the south-east, Lancashire to the south, and the Scottish ...

(previously referred to ''Hughmilleria''). The name ''Bassipterus'' is translated as "wing from Bass", with the first word of the name referring to the West Virginian

unincorporated community An unincorporated area is a parcel of land that is not governed by a local general-purpose municipal corporation. (At p. 178.) They may be governed or serviced by an encompassing unit (such as a county) or another branch of the state (such as th ...

of Bass and the last word composed of the

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 ...

word πτερόν (''

pteron In Classical architecture, a ''pteron'' (, 'wing') is an external colonnade around a building, especially an Ancient Greek temple. The '' pteroma'' or '' peristasis'' is the passage between the columns and the wall in a temple, the peristyle that ...

'', wing). Although Kjellesvig-Waering and Caster initially classified ''Bassipterus'' in the

Hughmilleriidae Hughmilleriidae (the name deriving from the type genus ''Hughmilleria'', which is named in honor of Scottish geologist Hugh Miller) is a family of eurypterids, an extinct group of aquatic arthropods. The hughmilleriids were the most basal member ...

, Victor P. Tollerton Jr. moved the genus along with others to the new family Adelophthalmidae. In 2004, the paleontologist Odd Erik Tetlie concluded that ''Bassipterus'' was a

synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...

of ''Parahughmilleria bellistriata'' and the more basal '' Stoermeropterus nodosus''. Although subsequent authors have followed this study, the most accepted

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

classification places ''Bassipterus'' together with ''Pittsfordipterus'', forming a basal clade.

Classification

''Bassipterus'' is classified as part of the family Adelophthalmidae, the only

("group") within the

. It was originally described as a hughmilleriid, but since then it has been considered closer to ''Adelophthalmus'' than to ''Hughmilleria''. ''Bassipterus'' and ''Pittsfordipterus'' form a basal ("primitive") group that differentiate them from the other adelophthalmids. This clade is backed by a pair of

synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...

(shared characteristics different from that of their

latest common ancestor A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...

); relatively long and narrow eyes and a complex termination of the genital appendage. Therefore, ''Pittsfordipterus'' is the

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

(closest relative) of ''Bassipterus''. ''Bassipterus'' was also similar to ''Parahughmilleria'', a derived adelophthalmid whose metastoma, telson and body were slightly differentiated. This has led some authors to synonymize ''Bassipterus'' with ''Parahughmilleria'', though this is not currently supported. The

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

below presents the inferred phylogenetic positions of most of the genera included in the three most derived superfamilies of the

Diploperculata Diploperculata is an infraorder of eurypterids, an extinct group of aquatic arthropods commonly known as "sea scorpions". The name, derived from Greek διπλόω ("double") and ''operculum'', refers to the distinguishing feature that unites the ...

infraorder of eurypterids (Adelophthalmoidea,

Pterygotioidea Pterygotioidea (the name deriving from the type genus ''Pterygotus'', meaning "winged one") is a superfamily (taxonomy), superfamily of eurypterids, an extinct group of aquatic arthropods. Pterygotioids were the most Derived trait, derived membe ...

and the waeringopteroids), as inferred by Odd Erik Tetlie and Markus Poschmann in 2008, based on the results of a 2008 analysis specifically pertaining to the Adelophthalmoidea and a preceding 2004 analysis.

Paleoecology

''Bassipterus'' fossils have been recovered from

deposits of the Late

Ludlow Ludlow ( ) is a market town and civil parish in Shropshire (district), Shropshire, England. It is located south of Shrewsbury and north of Hereford, on the A49 road (Great Britain), A49 road which bypasses the town. The town is near the conf ...

(

Ludfordian In the geologic timescale, the Ludfordian is the upper of two chronostratigraphic stages within the Ludlow Series. Its age is the late Silurian Period, and within both the Palaeozoic Era and Phanerozoic Eon. The rocks assigned to the Ludford ...

) and Early Pridoli

epochs In chronology and periodization, an epoch or reference epoch is an instant in time chosen as the origin of a particular calendar era. The "epoch" serves as a reference point from which time is measured. The moment of epoch is usually decided by ...

of the Wills Creek Formation of West Virginia and Maryland,

. In the Marylander part of the formation, fossils of other eurypterids have been found, such as ''Eurypterus remipes'' or '' Waeringopterus cumberlandicus'' together with an indeterminate species of the

ostracod Ostracods, or ostracodes, are a Class (biology), class of the crustacean, Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 33,000 species (only 13,000 of which are extant taxon, extant) have been identified,Brandão, S.N.; Antoni ...

'' Leperditia''. On the other hand, in the West Virginian section, remains of ''W. cumberlandicus'', '' Erettopterus exophthalmus'', ''Parahughmilleria bellistriata'' and ''Stoermeropterus nodosus'' have been associated, as well as unclassified

brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of 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 e ...

s and

bivalve Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...

s. ''Bassipterus'' lived in a

lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') an ...

al or restricted shallow

subtidal The neritic zone (or sublittoral zone) is the relatively shallow part of the ocean above the drop-off of the continental shelf, approximately in depth. From the point of view of marine biology it forms a relatively stable and well-illuminate ...

(the area where

sunlight Sunlight is the portion of the electromagnetic radiation which is emitted by the Sun (i.e. solar radiation) and received by the Earth, in particular the visible spectrum, visible light perceptible to the human eye as well as invisible infrare ...

reaches the bottom of the

ocean The ocean is the body of salt water that covers approximately 70.8% of Earth. The ocean is conventionally divided into large bodies of water, which are also referred to as ''oceans'' (the Pacific, Atlantic, Indian Ocean, Indian, Southern Ocean ...

) environment. The

lithology The lithology of a rock unit is a description of its physical characteristics visible at outcrop, in hand or core samples, or with low magnification microscopy. Physical characteristics include colour, texture, grain size, and composition. Lit ...

(description of the physical characteristics of the rocks) of the zone was composed of

argillaceous Clay minerals are hydrous aluminium phyllosilicates (e.g. kaolin, Al2 Si2 O5( OH)4), sometimes with variable amounts of iron, magnesium, alkali metals, alkaline earths, and other cations found on or near some planetary surfaces. Clay mineral ...

(composed of clay-like materials)

limestone Limestone is a type of carbonate rock, carbonate sedimentary rock which is the main source of the material Lime (material), lime. It is composed mostly of the minerals calcite and aragonite, which are different Polymorphism (materials science) ...

and

calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime (mineral), lime or being chalky. The term is used in a wide variety of Science, scientific disciplines. In zoology ''Calcare ...

(containing

calcium Calcium is a chemical element; it has symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar to it ...

)

shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...

References

{{Taxonbar, from=Q4868161 Adelophthalmoidea Silurian arthropods of North America Silurian eurypterids Eurypterids of North America Fossil taxa described in 1966

Description

History of research

Classification

Paleoecology

See also

References