Obaichthyidae
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Obaichthyidae (also known as "spiny gars" or "spike gars") is an extinct
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 ...
of ginglymodian
ray-finned fish Actinopterygii (; ), members of which are known as ray-finned fish or actinopterygians, is a class of bony fish that comprise over 50% of living vertebrate species. They are so called because of their lightly built fins made of webbings of sk ...
that lived in what is now
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
,
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
, and
southern Europe Southern Europe is also known as Mediterranean Europe, as its geography is marked by the Mediterranean Sea. Definitions of southern Europe include some or all of these countries and regions: Albania, Andorra, Bosnia and Herzegovina, Bulgaria, C ...
during the Upper
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period (
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
ages). They were close relatives of the modern gars of the family
Lepisosteidae Gars are an ancient group of ray-finned fish in the Family (biology), family Lepisosteidae. They comprise seven living species of fish in two genera that inhabit Fresh water, fresh, Brackish water, brackish, and occasionally marine waters of eas ...
, with the two groups making up the superfamily Lepisosteoidea. Though obaichthyids are generally similar to their extant relatives, they possessed a mobile
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 ...
along with a large posterior spine on their scales. The specializations of the skull seen in members of the family suggest that the fish were most likely feeding in a similar style to that seen in syngnathiforms, especially those specialized for pivot feeding. Though they are mostly found in fresh and
brackish water 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 estuary ...
deposits, obaichthyids also had a tolerance to saltwater similar to modern gars.


History

Even before these genera were placed into a single family, Obaichthys had been considered to be a sister to the rest of gars. The family was erected in 2010 by Lance Grande to include the genera '' Dentilepisosteus'' and '' Obaichthys''. ''
Afrocascudo ''Afrocascudo'' is a controversial genus of extinct Neopterygii, neopterygian fish, either an ancient loricariid catfish or a juvenile Obaichthyidae, obaichthyid Lepisosteiformes, lepisosteiform of the genus ''Obaichthys''. It is known from the L ...
'', initially described as the earliest known armored catfish, might represent a juvenile obaichthyid, possibly a junior synonym of ''Obaichthys'', though this has been disputed based on the complete ossification of the bones indicating full maturity and the absence of important holostean characters.


Anatomy


Skull

Just like gars today, the skull of obaichthyids were extremely long and are largely made up of the
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 ...
, with the length to width ratio of the
snout A snout is the protruding portion of an animal's face, consisting of its nose, mouth, and jaw. In many animals, the structure is called a muzzle, Rostrum (anatomy), rostrum, beak or proboscis. The wet furless surface around the nostrils of the n ...
being similar to ''Lepisosteus''. Unlike the skulls of modern relatives, members of the family have an inter- operculum; this bone was ancestrally present in gars as seen in genera like '' Araripelepidotes''. Some members, like ''Obaichthys'', have a large number of
ganoid A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as w ...
spines ornamenting the skull in contrast to other gars. One of the most notable features of the skull of obaichthyids in comparison to modern gars is the presence of a mobile
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 ...
. This bone is the largest element of the upper jaw and has a single row of short teeth. Along with this, members of this family lack the lacrimo-maxillary series that is so prevalent in the modern gars. Their lower jaw is made up of a reduced lower jaw which produces a prominent "
overbite Overbite is the extent of vertical ( superior-inferior) overlap of the maxillary central incisors over the mandibular central incisors, measured relative to the incisal ridges. The term overbite does not refer to a specific condition, nor is ...
" with teeth concentrated at the tip and a very small gape. Unlike those seen in the extant gars, the teeth of obaichthyids lack plicidentine, a structure suggested to be connected to the feeding style of the modern members. These differences in the feeding apparatus of obaichthyids suggests a different style of feeding than their modern relatives.


Postcrania

The overall body shape of obaichthyids is extremely similar that of modern gars, having a long body with the
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 ...
and anal fins being very far back on the body. Both families, along with other lepisosteoids, have opisthocoelous vertebrae. Unlike lepisosteids, at least some obaichthyids may have increased the number of vertebrae in the caudal region (tail), with ''Obaichthys decoratus'' having a total of 30; though this apparent elongation may be due to under- sampling, especially when compared to modern species. The main difference in the postcrania between the two groups is the shape and composition of their scales; unlike lepisosteids, obaichthyid scales have a spine projecting from their posterior end (rearwards) along with multiple spines along the margins of each scale. There is a layer of
dentin Dentin ( ) (American English) or dentine ( or ) (British English) () is a calcified tissue (biology), tissue of the body and, along with tooth enamel, enamel, cementum, and pulp (tooth), pulp, is one of the four major components of teeth. It i ...
e between the body of the scale and the ganoine layer; the placement of the dentine directly on top of bone is similar to what is seen in "
palaeoniscoid The Palaeonisciformes, commonly known as "palaeoniscoids" (also spelled "paleoniscoid", or alternatively "paleoniscids") are an extinct grouping of primitive ray-finned fish (Actinopterygii), spanning from the Silurian/Devonian to the Cretaceous. ...
" fish. This difference in scale morphology has caused the family to be referred to as either "spiny-" or "spike gars".


Classification

Obaichthyidae is thought to be the sister group of Lepisosteidae with the two groups making up Lepisosteoidea. Even so, it is not confirmed whether this family is
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
, as Arbarello (2012) found it to not be the case, though even within that study, it is agreed that members of the family are sister to one another. In this paper, it was defined as a
stem-based taxon Phylogenetic nomenclature is a method of nomenclature for taxon, taxa in biology that uses phylogenetics, phylogenetic definitions for taxon names as explained below. This contrasts with Biological classification, the traditional method, by which ...
containing all taxa more closely related to ''Obaichthys'' than to the genera ''
Lepisosteus ''Lepisosteus'' (from Greek ''lepis'' (), 'scale' and ''osteon'' (), 'bone') is a genus of gars in the family Gar, Lepisosteidae. It contains four extant species, found throughout eastern and central North America. It is one of two extant gar gen ...
'', '' Pliodetes'' or ''
Lepidotes ''Lepidotes'' (from , 'covered with scales') (previously known as ''Lepidotus'') is an extinct genus of Mesozoic ray-finned fish. It has long been considered a wastebasket taxon, characterised by "general features, such as thick rhomboid scales ...
''.


Evolutionary history

It is suggested that the family diverged from the modern gar lineage around 140 mya, which would have followed the separation of
North North is one of the four compass points or cardinal directions. It is the opposite of south and is perpendicular to east and west. ''North'' is a noun, adjective, or adverb indicating Direction (geometry), direction or geography. Etymology T ...
and
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
with the family likely originating in South America. A
radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or a material medium. This includes: * ''electromagnetic radiation'' consisting of photons, such as radio waves, microwaves, infr ...
between groups living on South America and
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
is seen due to the later opening of the
South Atlantic The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for ...
. This trend is also seen in a variety of other fish groups including mawsoniid coelacanths, vidalamiins, and cladocyclids. Though the family has mostly been found in
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
, a species of ''Obaichthys'' has been found in multiple sites in Europe.


Paleoenvironment

Like modern gars, they appear to have preferred
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. The term excludes seawater and brackish water, but it does include non-salty mi ...
&
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 ...
environments but were tolerant of marine conditions, allowing them to disperse across oceanic habitats. Also similar to their extant relatives, obaichthyids are mostly found in warmer environments, with the
range Range may refer to: Geography * Range (geographic), a chain of hills or mountains; a somewhat linear, complex mountainous or hilly area (cordillera, sierra) ** Mountain range, a group of mountains bordered by lowlands * Range, a term used to i ...
of the two groups overlapping at formations like the
Kem Kem The Kem Kem Group (commonly known as the Kem Kem beds) is a geological group (stratigraphy), group in the Kem Kem region of eastern Morocco, whose strata date back to the Cenomanian stage of the Late Cretaceous. Its strata are subdivided into two ...
.


Paleoecology

Due to various features of the mouth, it is unlikely that obaichthyids had a similar diet to modern gars with the family most likely not feeding on large prey. It is more likely that these fish fed mostly on
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s with a feeding style similar to what is seen in syngnathiforms, a method akin to pivot feeding.


References

Lepisosteiformes Prehistoric holostei Prehistoric ray-finned fish families Aptian first appearances Cenomanian extinctions {{Taxonbar, from=Q2008666