Holonematidae is an
extinct
Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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 ...
of relatively large
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 ...
placoderms
Placodermi (from Greek πλάξ 'plate' and δέρμα 'skin', literally 'plate-skinned') is a class of armoured prehistoric fish, known from fossils, which lived from the Silurian to the end of the Devonian period. Their head and thorax were ...
from the Early to Late Devonian. Almost all fossil specimens are of armor fragments, though, all have distinctive ornamentation, often of unique arrangements and patterns of tubercles, that are diagnostic of the family. The trunkshield is very elongated, giving the armor an overall "barrel" like appearance.
Due to pebbles found inside articulated specimens of the Frasnian species, ''Holonema westolli'', holonematids are thought to have been specialized
herbivore
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s that grazed on a form of horn-shaped, stony algae called "onycholite," snipping off the tips with their odd snouts.
Genera
''Artesonema''
A poorly known genus from the Givetian-aged Blacourt Formation of
Boulonnais, France.
[LELIEVRE, n.; GOUJET, D., BLIECK, A. & JANVIER, P. 1986. ''Poissons du DEvonien du Boutonnais (France)''. - n:BRICE, D. d.Le DEvonien de Ferques, Bas -Boulonnais (N. France) - Biostratigraphie du PalEozoique 7: 503-522, Brest.] The holotype of ''
Artesonema'' is an anterodorsolateral plate from an adult that has an ornamentation of tubercles normally characteristic of juvenile holonematids.
[Otto, Markus. "New finds of vertebrates in the Middle Devonian Brandenberg Group (Sauerland, Northwest Germany)." Paläontologische Zeitschrift 72, no. 1-2 (1998): 117-134.]
''Belemnacanthus''
''
Belemnacanthus
''Belemnacanthus giganteus'' ("Gigantic arrow-spine") is a large, extinct, barrel-shaped holonematid arthrodire placoderm from Givetian-aged strata of Middle Devonian Eifel, Germany. ''B. giganteus'' is known only from the holotype, a portion ...
'' is from the Givetian of
Eifel
The Eifel (; lb, Äifel, ) is a low mountain range in western Germany and eastern Belgium. It occupies parts of southwestern North Rhine-Westphalia, northwestern Rhineland-Palatinate and the southern area of the German-speaking Community of ...
,
Germany
Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. It has been previously described as being an
elasmobranch
Elasmobranchii () is a subclass of Chondrichthyes or cartilaginous fish, including sharks (superorder Selachii), rays, skates, and sawfish (superorder Batoidea). Members of this subclass are characterised by having five to seven pairs of gil ...
, an
agnatha
Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct ( conodonts and ostracoderms) species. Among recent animals, cyclosto ...
n, and an
antiarch
Antiarchi ("opposite anus") is an order of heavily armored placoderms. The antiarchs form the second-most successful group of placoderms after the arthrodires in terms of numbers of species and range of environments. The order's name was coi ...
. The genus is known only from the holotype: a 37 cm long portion of a median dorsal plate, suggesting a very large animal. The median dorsal plate has a long crest running down the median of the dorsal surface. The outer surface has an ornamentation of long ridges radiating from a point beyond the preserved portions.
''Bimbianga''
If ''
Bimbianga'' is a holonematid, then it is the earliest genus, dating from the
Emsian
The Emsian is one of three faunal stages in the Early Devonian Epoch. It lasted from 407.6 ± 2.6 million years ago to 393.3 ± 1.2 million years ago. It was preceded by the Pragian Stage and followed by the Eifelian Stage. It is named after the ...
-aged Taemas Limestone of
Wee Jasper, New South Wales
Wee Jasper is a hamlet in the Goodradigbee valley at the western foot of the Brindabella Ranges, near Burrinjuck Dam in New South Wales, Australia in Yass Valley Shire. It is located about 90 km north-west of Canberra and 60 km south ...
. ''Bimiangas placement within Holonematidae is questionable, as the morphology of its plates and ornamentation are very similar to those of the arthrodire ''
incertae sedis
' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' ''
Aspidichthys''.
[YOUNG, Gavin C. "Early Devonian arthrodire remains (Placodermi,? Holonematidae) from the Burrinjuck area, New South Wales, Australia." Geodiversitas 27, no. 2 (2005).]
''Deirosteus''
''
Deirosteus'' is another poorly known genus. The ornamentation consists of acute ridges between broad interspaces often decorated with large tubercles. The genus is very similar to ''Holonema'', and it is hard to distinguish between the two. Fossils are known from the Frasnian of
New York
New York most commonly refers to:
* New York City, the most populous city in the United States, located in the state of New York
* New York (state), a state in the northeastern United States
New York may also refer to:
Film and television
* '' ...
, and possibly
Iran
Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
, and from the
Givetian
The Givetian is one of two faunal stages in the Middle Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Eifelian Stage and followed by the Frasnian Stage. It is named after the town of Givet in Fra ...
of
Estonia
Estonia, formally the Republic of Estonia, is a country by the Baltic Sea in Northern Europe. It is bordered to the north by the Gulf of Finland across from Finland, to the west by the sea across from Sweden, to the south by Latvia, a ...
,
Russia
Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
and
Belgium
Belgium, ; french: Belgique ; german: Belgien officially the Kingdom of Belgium, is a country in Northwestern Europe. The country is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeast, France to th ...
.
''Deveonema''
''
Deveonema'' is known from a fragmentary median dorsal plate from
Frasnian
The Frasnian is one of two faunal stages in the Late Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Givetian Stage and followed by the Famennian Stage.
Major reef-building was under way during th ...
-aged strata within the
Holy Cross Mountains
Sacred describes something that is dedicated or set apart for the service or worship of a deity; is considered worthy of spiritual respect or devotion; or inspires awe or reverence among believers. The property is often ascribed to objects (a ...
of
Poland
Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...
. The median dorsal suggests a very small holonematid. The dermal surface is ornamented with a pattern of small tubercles, usually separate but sometimes in clumps of two or three. Along the center, the tubercles are arranged in longitudinal rows.
''Gyroplacosteus''
''
Gyroplacosteus'' is a genus known from various Frasnian deposits throughout Eastern Europe. The ornamentation is formed from coarse tubercles that are often fused together as sinuous ridges in various, usually complicated patterns.
''Holonema''
''
Holonema'' is the best known genus, as at least one species, ''H. westolli'', is known from complete, articulated specimens. ''Holonema'' is also the most widespread genus, as the various species are found in Eifelian to Frasnian-aged deposits in the United States, Scotland, Europe, Western Asia, and Australia. The typical ornamentation consists of patterns of alternating rows of tubercles and ridges, with narrow interspaces.
''Megaloplax''
''
Megaloplax'' is a genus based on an incomplete median dorsal plate from Frasnian-aged deposits in the
Ural Mountains
The Ural Mountains ( ; rus, Ура́льские го́ры, r=Uralskiye gory, p=ʊˈralʲskʲɪjə ˈɡorɨ; ba, Урал тауҙары) or simply the Urals, are a mountain range that runs approximately from north to south through western ...
.
''Rhenonema''
''
Rhenonema'' has a high crest along the median ridge of its median dorsal plate. The few fragments of armor are from Givetian-aged strata in
Gerolstein
Gerolstein () is a town in the Vulkaneifel district of Rhineland-Palatinate, Germany. Gerolstein is a local municipality of the ''Verbandsgemeinde Gerolstein''. It has been approved as a '' Luftkurort'' (spa town).
History
As early as the Ston ...
, Germany.
''Tropidosteus''
''
Tropidosteus'' is a genus known from the Givetian-aged Crinoidenmergel of the
Rheinland
The Rhineland (german: Rheinland; french: Rhénanie; nl, Rijnland; ksh, Rhingland; Latinised name: ''Rhenania'') is a loosely defined area of Western Germany along the Rhine, chiefly its middle section.
Term
Historically, the Rhinelands ...
. The median dorsal plate has a large, hump-like arch. Because it lacks ornamentation along the median edge of the median dorsal, Denison 1978 questions its placement within Holonematidae (as all other genera with known median dorsals have ornamentation along the median edge). Otherwise, the dermal surface is covered in very small tubercles.
References
Bibliography
*
* Case, Gerald Ramon.
A Pictorial Guide to Fossils
* Long, John.
Swimming In Stone
{{Taxonbar, from=Q16880282
Placoderm families
Early Devonian first appearances
Late Devonian animals
Late Devonian extinctions