''Pseudoangustidontus'' (meaning "false ''
Angustidontus''") is a genus of
hurdiid
Hurdiidae (synonymous with the previously named Peytoiidae) is an extinct cosmopolitan distribution, cosmopolitan Family (biology), family of radiodonts, a group of Crown group#Stem groups, stem-group marine arthropods, which lived during the Pal ...
(peytoiid)
radiodont
Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and were used for a variety ...
from the
Lower Ordovician
The Early Ordovician is the first epoch of the Ordovician period, corresponding to the Lower Ordovician series of the Ordovician system. It began after the Age 10 of the Furongian epoch of the Cambrian and lasted from to million years ago, unt ...
of
Morocco
Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It has coastlines on the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to Algeria–Morocc ...
.
This genus includes two described species, ''Pseudoangustidontus'' ''duplospineus'' and ''Pseudoangustidontus'' ''izidigua'', which both belong to the smaller
Aegirocassisinae
Aegirocassisinae is a subfamily of radiodonts (marine stem-arthropods) from the lower Paleozoic era. It belongs to the larger hurdiidae (peytoiid) family, which were the most diverse and long lasting radiodonts. The members of this subfamily are re ...
subfamily, however, a third unnamed species is also known. This animal is only known from the
Fezouata Formation
The Fezouata Formation or Fezouata Shale is a geological formation in Morocco which dates to the Ordovician, Early Ordovician. , a lower
Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
fossil site in Morocco that is of
''Konservat-Lagerstätten'' status, meaning that the fossils from this site are exceptionally well preserved.
This taxon was first described in 2006, but due to the fragmentary remains known at the time, its classification was debated, but with more complete fossils, this taxon was identified as a hurdiid radiodont in 2023.
Similarly to its larger relative,
''Aegirocassis benmoulae'', this radiodont is thought to have lived a primarily
filter feeding
Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters, food particles or smaller organisms (bacteria, microalgae and zooplanktons) suspended in water, typically by having the water pass over or through a spe ...
lifestyle, using its numerous
auxiliary spines to grab small organisms from the water.
Background
Located in the
Draa Valley of southeastern Morocco, the Fezouata Formation is a lower Ordovician (
Tremadocian
The Tremadocian is the lowest stage of Ordovician. Together with the later Floian Stage it forms the Lower Ordovician Epoch. The Tremadocian lasted from to million years ago. The base of the Tremadocian is defined as the First appearance datum ...
–
Floian
The Floian is the second stage of the Ordovician Period. It succeeds the Tremadocian with which it forms the Lower Ordovician series. It precedes the Dapingian Stage of the Middle Ordovician. The Floian extended from to million years ago. The ...
) aged
geologic formation
A geological formation, or simply formation, is a body of rock having a consistent set of physical characteristics ( lithology) that distinguishes it from adjacent bodies of rock, and which occupies a particular position in the layers of rock exp ...
well known for its exceptional preservation of
soft-bodied marine organisms.
The soft body producing layers of the formation, known as the Fezouata Lagerstätte, were first identified in the late 1990s, when Ben Moula,local fossil collector in the area, showed some uncovered specimens to a PhD student working in the area, and the soon to be discovered ecosystem would come to be known as the Fezouata Biota.
The formation is especially important, as it helped bridge the gap between
Cambrian
The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
-type and Ordovician-type
faunas, showing that many groups of organisms once thought to have gone extinct at the end of the Cambrian, survived the transition into the Ordovician.
The site is highly productive, with around 50 distinct taxa being represented from over 1,500 specimens, with the majority of said specimens coming from two small intervals of the formation around and thick, compared to the nearly thickness of the whole formation. During the Ordovician, the area that would become Morocco was situated much closer to the
South Pole
The South Pole, also known as the Geographic South Pole or Terrestrial South Pole, is the point in the Southern Hemisphere where the Earth's rotation, Earth's axis of rotation meets its surface. It is called the True South Pole to distinguish ...
then it is today, and the ecosystem of Fezouata lived in a
storm wave base roughly 50 and 150 metres (160 and 490 ft) underwater.
The organisms in the environment were preserved
in situ
is a Latin phrase meaning 'in place' or 'on site', derived from ' ('in') and ' ( ablative of ''situs'', ). The term typically refers to the examination or occurrence of a process within its original context, without relocation. The term is use ...
, and because of storms in the area helping to facilitate rapid sediment deposition, the majority of known organisms were
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
in nature.
Discovery and etymology
The fossil specimens belonging to ''Pseudoangustidontus,'' more specifically the species ''P. duplospineus'', were discovered in the upper strata of the Fezouata Formation, and were subsequently described by Van Roy & Tetlie, 2006.
However, because the fossils were found by local collectors, and were brought to the authors' attention via circulation, the exact locality of origin is uncertain, with several sites located near the town of
Zagora being possible candidates.
Because of its unique appearance and fragmentary state, the taxon would be considered problematic (in terms of its taxonomy), but would later be identified as a radiodont later on in 2023.
In that same year, a second species, ''P. izdigua'', was described based on differences in spine length compared to the
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 ...
, and is known from more complete remains (
endites,
frontal appendages, and a partial
h-element).There is also a currently unnamed species, but due to its fragmentary state, is unassignable aside from genus level classification.
The genus name, ''Pseudoangustidontus'', is derived from 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 ''pseudes'', meaning "false", and the genus name ''
Angustidontus'', an unrelated
pelagic
The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
crustacean
Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
from the
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 ...
-
Carboniferous
The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
periods, due to similarities between the
raptorial
In biology (specifically the anatomy of arthropods), the term ''raptorial'' implies much the same as ''predatory'' but most often refers to modifications of an arthropod leg, arthropod's foreleg that make it function for the grasping of prey whi ...
limbs of ''Angustidontus'', and the endites of ''P. duplospineus'', while still acknowledging the differences between the two taxa.
The type species name, ''duplospineus'', is derived from the
Latin
Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
words ''duplus'', meaning "double", and ''spineus'', meaning "spiny", or "thorny", highlighting the double pair of auxiliary spines seen in the species.
The other species name, ''izdigua'', is derived from the
Tamazight
The Berber languages, also known as the Amazigh languages or Tamazight, are a branch of the Afroasiatic language family. They comprise a group of closely related but mostly mutually unintelligible languages spoken by Berber communities, who ar ...
word ''izdigue'', which means "filter", highlighting the filter feeding nature of the taxon.
Description
File:20230828 Radiodonta frontal appendage Pseudoangustidontus duplospineus.png, Reconstruction of the frontal appendage of ''P. duplospineus''
File:20230828 Radiodonta frontal appendage Pseudoangustidontus izdigua.png, Reconstruction of the frontal appendage of ''P. izdigua''
File:20191229 Radiodonta frontal appendage Aegirocassis benmoulai Aegirocassis benmoulae.png, Reconstruction of the frontal appendage of ''Aegirocassis'' for comparison
''Pseudoangustidontus'' is only known from frontal appendage remains, with an exception of MGL 108047_1, 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 ...
of ''P. izidigua'', which preserved part of a frontal sclerite (H-element). ''P. duplospineus'' had paired auxiliary spines alternating in length short/long while the frontal appendages of ''P. izdigua'' bore endites with spines that are equal in length.
''P. duplospineus'' is differentiated from its close relative ''Aegirocassis'' due to the paired spines on its frontal appendages, with the shorter spines being equal in length compared to the width of the endites, and the longer spines being nearly twice as long compared to the endite width.
Classification
When described in 2006, the only known fossils of this genus were isolated endites that bore copious amounts of auxiliary spines. Because of this, the
taxonomy
image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy
Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...
of this stem-arthropod was debated, with possible suggestions of its affinity being with the
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 ...
s of the
megalograptidae family, radiodonts, or crustaceans such as ''Angustidontus''.
In a review article published in 2023, it was questionably placed in the Hurdiidae.
And in another paper that was released in the same year, and published by Potin, Gueriau & Daley, 2023, described newly found fossils, and found it to be closely related to ''Aegirocassis'', and both into the subfamily Aegirocassisinae.
The family that the Aegirocassisinae subfamily belongs to, the Hurdiidae, were the most diverse and long lasting lineage of radiodonts, usually characterized by subequal endites, a tetraradial
oral cone, and large frontal sclerites (which in some species reach, or exceed the total body length).
Paleoecology

''Pseudoangustidontus'' is speculated to be a filter feeder similar to ''Aegirocassis'', that used numerous auxiliary spines on its frontal appendages as mesh to catch microorganisms, similarly to the
baleen
Baleen is a filter feeder, filter-feeding system inside the mouths of baleen whales. To use baleen, the whale first opens its mouth underwater to take in water. The whale then pushes the water out, and animals such as krill are filtered by th ...
seen in various
cetacean
Cetacea (; , ) is an infraorder of aquatic mammals belonging to the order Artiodactyla that includes whales, dolphins and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively c ...
species. The spine and mesh size of the frontal appendages of ''Pseudoangustidontus'' is around 0.5 mm, slightly smaller than that of ''Aegirocassis'' (0.56 mm), both genera probably fed on
mesoplankton. However, a specimen of ''P. izdigua'' had an unidentified shell or carapace with a diameter about 3 mm trapped in its setae, suggesting it could have fed on even larger prey.
References
{{taxonbar, from=Q21368832, from2=Q3924863
Fossils of Morocco
Radiodonta
Fossil taxa described in 2006
Fezouata Formation fossils
Ordovician arthropods