Lunaspis Broili
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''Lunaspis'' is an
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus of armor-plated petalichthyid
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 ...
fish that lived in shallow marine environments of the
Early Devonian The Early Devonian is the first of three Epoch (geology), epochs comprising the Devonian period, corresponding to the Lower Devonian Series (stratigraphy), series. It lasted from and began with the Lochkovian Stage , which was followed by the Pr ...
period, from approximately 409.1 to 402.5 million year ago.Fossilworks: Gateway to the Paleobiology database. Retrieved March 6, 2017, from Fossilworks, http://www.fossilworks.org/cgi-bin/bridge.pl?page=paleodb Fossils have been found in
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
and
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
. There are three different identified species of within the genus ''Lunaspis'': ''L. broilii, L. heroldi, and L. prumiensis.'' Like many other petalichthyid fish, ''Lunaspis'' are flat and have elongated pectoral spines, shortened thoracic armor, and dorsally oriented eyes. ''Lunaspis'' has been studied mainly by German, Australian, and Chinese palaeontologists because of where it is most commonly encountered. The tails are long and thin and resemble a whip, somewhat like
extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...
skates and
rays Ray or RAY may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), the bony or horny spine on ray-finned fish Science and mathematics * Half-line (geometry) or ray, half of a line split at an ...
.


Description

A typical individual of ''Lunaspis'' was a flattened fish with a short trunk and long, curved cornual plates. These long, spine-like plates give the suggestion of a crescent moon, hence the generic name (moon-shield). The nostrils and the anterior part of the head shield around the orbits is covered by a number of tiny scales, as is the elongated trunk. ''Lunaspis'' were marine bottom dwelling creatures, like many other
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 ...
placoderm fish, and were
nekton Nekton or necton (from the ) is any aquatic organism that can actively and persistently propel itself through a water column (i.e. swimming) without touching the bottom. Nektons generally have powerful tails and appendages (e.g. fins, pleopods, ...
ic carnivores. The body shape of ''Lunaspis'' is also similar to other petalichthyid fish, compressed dorsoventrally.More about Placoderms. (2002). Retrieved March 6, 2017, from Devonian Times, http://www.devoniantimes.org/who/pages/placoderm.html Its body consists of a short trunk shield and elongated cronual plates (the wing-like structures that make up the crescent shape of the moon-like head). The rest of the body, which excludes those parts covered in the enlarged plates of armor, are covered in tiny scales, as is the anterior region of the face near the orbits and nostrils. ''Lunaspis'' also has two traits that are thought to be primitive, or ancestral: large submarginal plates and the occurrence of a ventral pit on the median dorsal plate.


Bony armor

According to specimens examined from the Taemas-Wee Jasper region of New South Wales in Australia, both ''L. broilii'' and ''L. herolfi'' have tubercles of the median dorsal plate coalesce into a distinct median dorsal ridge (MDR). The fusion of the tubercles is unique to ''Lunaspis'' in the order Petalichthyida, where other devonian fish in this order have ornament tubercles that are separately occurring. The maximum height of the ridge is reached before the posterior margin of the median dorsal plate. The spinal plates are curved and are the wings off of the center part of the bony armoring. They have many small spines along the anterior ridge of spinal plates. The ornament ridges on the bony plates make it easily distinguishable from other petalichtyid fish. These ornament ridges are widely spaced and continuous in the ''Lunaspis'', across the three species; in ''L. broilii'', the ornaments are more densely packed than ''L. heroldi'' and ''L. prumiensis''.Young, G. C. (1985). Further petalichthyid remains (placoderm fished, Early Devonian) from the Taemas-Wee Jasper region, new South Wales. BMR Journal of Australian Geology and Geophysics, 9, 121–131.


Discovery

The generic named is a compound word combining two different Greek words: “Luna” and “aspis”. In Greek, the word “luna” means The Moon, and the word “aspis” means Round Shield. Together, luna and asp signify “moon shield”; appropriately named as the long projections from the head shield make a crescent moon-like shape. The first specimens of ''L. broilii'' were discovered and described through the collaboration of two German palaeontologists, Walter R. Gross and Ferdinand Broili. Broili discovered ''L. heroldi'' in 1929 in Germany. In Bundenbach, Germany 1937, Broili then discovered another fossil specimen that resembled ''Lunaspis'' ''heroldi''. At first he put into the same species as ''L. heroldi'', but noticed a difference in morphology from the previously discovered species. Near where Broili had discovered his specimen, another palaeontologist by the name of Gross was working with another specimen of ''Lunaspis'' which he originally thought to be ''Lunaspis'' ''prumiensis''. Gross was working with fragments of the cranial exoskeletal bones of the unidentified ''Lunaspis'' species. The palaeontologists connected soon after and compared their specimens with one another and worked together to discover the differences that existed among the existing species within the genus and were able to come to a conclusion. According to the spinal and anterior ventro-lateral anatomy of the specimens they had collected, they determined that the specimen that both of them had found was a different one than had yet been identified. This new species within the same genus as ''L. heroldi'' and ''L. prumiensis'' was named ''Lunaspis broilii''. ''L. Broilii'' is very commonly larger than ''L. heroldi''. The specimens of ''L. broilii'' were originally only found in
Emsian The Emsian is one of three faunal stages in the Early Devonian Epoch. It lasted from 410.62 ±1.95 million years ago to 393.47 ±0.99 million years ago. It was preceded by the Pragian Stage and followed by the Eifelian Stage. It is named after ...
-aged Hunsruck Slate of
Bundenbach Bundenbach is an '' Ortsgemeinde'' – a municipality belonging to a ''Verbandsgemeinde'', a kind of collective municipality – in the Birkenfeld district of Rhineland-Palatinate, Germany. It belongs to the ''Verbandsgemeinde'' Herrstein-Rhaune ...
, Germany,Lehmann, W. M. (1951). Neue Beobachtungen an Lunaspis. Neues Jahrbuch Fuer Geologie Und Palaeontologie. Abhandlungen, 94(1), 93-100. but in 1980, Liu Shifan found specimens that are most likely identifiable as ''L. broilii''. Specimens of ''L. broilii'' have been more recently found in Reefton, New Zealand, and have been identified as closely related, if not the same as those found in New South Wales, Australia. They have been found to have the same concentrically tuberculated ridges which is typical of ''Lunaspis''.Macadie, I. (2007). A Placoderm Fish Plate from the Lower Devonian of Reefton, New Zealand. Records of Canterbury Museum, 21, 21–26. Specimens of ''L. heroldi'' are also found in similarly aged marine strata in China and Australia.


Classification

''Lunaspis'' was originally placed within the family Acanthaspidae, but according to Walter Gross's findings in 1937, the type genus of Acanthaspidae, ''Acanthaspis'', was synonymized with ''Lunaspis'', and the entire family merged into the family Macropetalichthyidae. ''Lunaspis'' is currently placed among seven other Macropetalichthyids, and sister to two other
families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...
within
Petalichthyida Petalichthyida is an Extinction (biology), extinct Order (biology), order of small, flattened placoderm fish. They are typified by their splayed pectoral fins, exaggerated lateral spines, flattened bodies, and numerous tubercles that decorated a ...
.


Paleoecology

Specimens from the Reefton, New Zealand site were found in the Waitah
Outlier
near th
Adam Mudstone
east of Reefton. The specimen from the Waitahu Outlier was found among a multitude of invertebrate fossils,
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,
bivalves Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed by a calcified exoskeleton consis ...
, orthocene
nautiloid Nautiloids are a group of cephalopods (Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and species rich, with over 2,500 recorded species. Th ...
s,
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
ns, and isolated
crinoid Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in their adult form are commonly called sea lilies, while the unstalked forms, called feather stars or comatulids, are ...
stem ossicles. The Adam Mudstone spans from 407.6 to 393.3 million years ago. The Adam Mudstone formation is marine and composed of silty mudstone and calcareous siltstone. "The stratigraphic sequence in the Inangahua and Waitahu Outliers is essentially the same, and includes thick sandstones at the bottom and top of the succession, with an alternation of nearshore limestone and offshore mudstone formations in between. The Waitahu sequence is more continuous and is not broken by tectonic slides".Bradshaw, M. (1995). Stratigraphy and structure of the Lower Devonian rocks of the Waitahu and Orlando Outliers, near Reefton, New Zealand, and their relationship to the Inangahua Outlier. New Zealand Journal of Geology and Geophysics, 38, 81–92.


References

{{Taxonbar, from=Q3267453 Petalichthyida Placoderms of Asia Placoderms of Australia Placoderms of Europe Placoderm genera Hunsrück Slate fossils