Rhenanida ("
Rhine
The Rhine ( ) is one of the List of rivers of Europe, major rivers in Europe. The river begins in the Swiss canton of Graubünden in the southeastern Swiss Alps. It forms part of the Swiss-Liechtenstein border, then part of the Austria–Swit ...
(fish)") is an
order
Order, ORDER or Orders may refer to:
* A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
...
of scaly
placoderm
Placoderms (from Ancient Greek πλάξ 'plax'', ''plakos'''Plate (animal anatomy), plate' and δέρμα 'derma'''skin') are vertebrate animals of the class (biology), class Placodermi, an extinct group of prehistoric fish known from Pal ...
s. Unlike most other placoderms, the rhenanids' armor was made up of a mosaic of unfused scales and tubercles. The patterns and components of this "mosaic" correspond to the plates of armor in other, more advanced placoderms, suggesting that the ancestral placoderm had armor made of unfused components, as well.
All rhenanids were flattened,
ray-like, bottom-dwelling
predator
Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
s that lived in marine environments.
Evolution
The rhenanids were once presumed to be the most primitive, or at least the closest to the ancestral placoderm, as their armor was made up of a mosaic of tubercles, as opposed to the solidified plates of "advanced" placoderms, such as
antiarchs and
arthrodire
Arthrodira (Greek for "jointed neck") is an Order (biology), order of extinct armored, jawed fishes of the class Placodermi that flourished in the Devonian period before their sudden extinction, surviving for about 50 million years and penetratin ...
s. However, comparing the skull anatomies of ''Jagorina pandora'' with those of antiarchs, the rhenanids are considered to be the sister group of the antiarchs (together with their respective
Acanthothoracid relatives).
Presence in the fossil record
The fossil record of Rhenanida is very sparse, with most fossils being isolated tubercles and skull fragments that are identified as being similar to ''
Gemuendina stuertzi'', the most well-known rhenanid, known from several specimens from the
Hunsruck slates. Given the rhenanids' worldwide distribution, this paupacity probably did not reflect a scarcity of living individuals (when the order was alive), but reflects the fact that rhenanid armor disintegrated into isolated fragments, and scattered soon after the owner's demise. Most fossils of rhenanids are from the Early
Devonian
The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
, primarily in the United States and Germany. The recently discovered ''
Nefudina qalibahensis'' is known from Northeastern
Saudi Arabia
Saudi Arabia, officially the Kingdom of Saudi Arabia (KSA), is a country in West Asia. Located in the centre of the Middle East, it covers the bulk of the Arabian Peninsula and has a land area of about , making it the List of Asian countries ...
. ''
Asterosteus stenocephalus'' is known from Mid Devonian
Ohio
Ohio ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Erie to the north, Pennsylvania to the east, West Virginia to the southeast, Kentucky to the southwest, Indiana to the ...
. Another species of rhenanid was ''
Bolivosteus chacomensis'', of the Lower to Middle Devonian
Malvinokaffric Fauna of Western
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 ...
, in what is now Bolivia, South America. The youngest rhenanid, ''
Jagorina pandora'' is known from Upper Devonian Germany.
Taxonomy
There are five recognized
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), ...
of rhenanids, in five
genera
Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
, ''Asterosteus stenocephalus'', ''Nefudina qalibahensis'', ''Gemuendina stuertzi'', ''Jagorina pandora'', and ''Bolivosteus chacomensis''. They are all placed within the family Asterosteidae, erected by Woodward in 1891: other families attributed to Rhenanida, i.e., Gemuendinidae and Jagorinidae, are considered synonyms.
A sixth genus, ''
Ohioaspis'', is of questionable status, as the first specimens were
ichthyoliths that were originally described as being tubercles from a new species of ''Asterosteus''. Later examinations of these tubercles have led to the formation of two camps of experts, one of which that believe the three recognized species of ''Ohioaspis'' were rhenanids, while the other suggests that they were actually some sort of
ostracoderm
Ostracodermi () or ostracoderms is an informal group of vertebrate animals that include all armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes and ...
agnatha
Agnatha (; ) or jawless fish is a paraphyletic infraphylum of animals in the subphylum Vertebrata of the phylum Chordata, characterized by the lack of jaws. The group consists of both extant taxon, living (Cyclostomi, cyclostomes such as hagfish ...
ns.
References
Paleos Rhenanida* Janvier, Philippe. ''Early Vertebrates'' Oxford, New York: Oxford University Press, 1998.
* Long, John A. ''The Rise of Fishes: 500 Million Years of Evolution'' Baltimore: The Johns Hopkins University Press, 1996.
*
{{Taxonbar, from=Q144336
Placoderm orders
Early Devonian first appearances