''Tanazios'' is a genus of
Silurian
The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...
stem-
mandibulate
The clade Mandibulata constitutes one of the major subdivisions of the phylum Arthropoda, alongside Chelicerata. Mandibulates include the crustaceans, myriapods (centipedes and millipedes, among others), and all true insects. The name "Mandibulat ...
from the
Coalbrookdale Formation
Coalbrookdale Formation, earlier known as Wenlock Shale or Wenlock Shale Formation and also referred to as Herefordshire Lagerstätte in palaeontology, is a fossil-rich deposit ('' Konservat-Lagerstätte'') in Powys and Herefordshire at the Engl ...
of England.
Description
''Tanazios'' has a head shield that is semicircular in shape, divided into a fringe and an axial (central) region. The fringe margins meet somewhat abruptly at the front of the shield, with a long projection pointing from their back corners and six pairs of horn-like structures at the sides. The axial region is semi-ovoid in shape, around 3 times wider than long, and bears a wide furrow running down its length. At the sides there is a narrow ledge which pinches out towards the back and continues as the inner trunk pleurae. No eyes seem to be present. The
hypostome is subrectangular and extends from the antennulae to the mandibular gnathobases, with an anterior margin bearing four small spines and a convex posterior margin with a lip-like structure which may be the
labrum.
The antennulae are very short and
uniramous
The arthropod leg is a form of jointed appendage of arthropods, usually used for walking. Many of the terms used for arthropod leg segments (called podomeres) are of Latin origin, and may be confused with terms for bones: ''coxa'' (meaning hip, : ...
, with the antennae being around five times longer and more ovoid in cross-section. The antennae are divided into three sections based on angle changes in the podomeres (segments). While there seems to be a pair of setae-like structures near the tip of the antennae on 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 ...
specimen, this feature is absent in others and may be a preservational artefact. The
mandible
In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla).
The jawbone i ...
is also uniramous, with its large
coxa bearing a sharply raised
gnathobase with an incisor-like protrusion. The basipod is subrectangular (rectangular with rounded corners), and narrows distally (away from the body axis). The endopod is rather small and like the antennae is split into three sections. The first two are fairly short, with the third being as long as both combined and tapering into a finger-like extension.
[
The fourth and fifth appendages are somewhat similar to both each other and the trunk appendages, both being biramous. The limb base is subrectangular with large subtriangular areas on the dorsal side (likely muscle insertion sites) alongside a flange-like endite projecting backwards. The limb base also bears an epipodite on its dorsal surface, likely alongside one more ventrally. The endopod is long and ribbon-like at its base, but becomes more finger-like towards its tip. It too is composed of three sections, with the first two only comprising one segment and the other being much longer. The exopod is similar (though shorter) and more ribbon-like. The endopod also has an unusual membrane running from its middle down to the limb base.][
The trunk is composed of at least sixty-four segments alongside a small terminal piece, with a ]telson
The telson () is the hindmost division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment (biology), segment of the arthropod body, or an additional division that is not a true segm ...
and caudal furcae at the terminal end. The anterior ten become smaller anteriorly and the posterior twenty become smaller posteriorly, with the last few being tiny. Each segment bears a pair of biramous appendages except possibly the last four (although they are too poorly preserved to tell), with a narrow groove running down the trunk’s center. The axial region is around 80% the trunk’s width, with the head groove continuing onto the first three segments, with a median spine on segments 22 to 62 (the last two instead have small median nodes). The pleural region has a very small ledge, with descending facets on all except the first segment. The pleurae are somewhat rounded at their corners with slight overlap posteriorly. The last three or four trunk segments either lack pleurae or have them, but are too small to be preserved.[
Behind the terminal axial piece a small telson is preserved, extending into two triangular flaps (one dorsal, one ventral) with two caudal furcae, ovoid in cross-section, extending from the back parts of this telson. While these are incomplete, the preserved length is around 8% of the body length. Near the back of the telson a small opening flanked by two conical structures is preserved; this is likely the ]anus
In mammals, invertebrates and most fish, the anus (: anuses or ani; from Latin, 'ring' or 'circle') is the external body orifice at the ''exit'' end of the digestive tract (bowel), i.e. the opposite end from the mouth. Its function is to facil ...
.[
]
Affinity
''Tanazios'' somewhat resembles both the enigmatic arthropod ''Wingertshellicus
''Wingertshellicus'' is an extinct genus of arthropod that has been found in Hunsrück Slate, that is located in the Rhenish Massif in Germany, and lived about 405 million years ago, during the Lower Emsian (part of the Lower Devonian).
Etymolo ...
'' and modern remipede
Remipedia is a class of blind crustaceans, closely related to hexapods, found in coastal aquifers which contain saline groundwater, with populations identified in almost every ocean basin so far explored, including in Australia, the Caribbean Se ...
s in morphology, however the limb arrangement of the head places it as a basal pancrustacea
Pancrustacea is the clade that comprises all crustaceans and all hexapods (insects and relatives). This grouping is contrary to the Atelocerata hypothesis, in which Hexapoda and Myriapoda are sister taxa, and Crustacea are only more distantl ...
n. More specifically, it either falls as a basal member of the clade containing both Pancrustacea and Phosphatocopina
Phosphatocopina (alternatively Phosphatocopida) is an extinct group of bivalved arthropods known from the Cambrian period. They are generally sub-milimetric to a few millimetres in size. They are typically only known from isolated carapaces, but ...
(Labrophora) or as an even more basal member allied with various Orsten
The Orsten fauna are fossilized organisms preserved in the Orsten lagerstätte of Cambrian (Late Miaolingian to Furongian) rocks, notably at Kinnekulle and on the island of Öland, all in Sweden.
The initial site, discovered in 1975 by Klaus M� ...
arthropods, although likely the closest to the crown-group of these due to having a uniramous mandible.[
]
Palaeoecology
The Coalbrookdale Formation where ''Tanazios'' was found was deposited in waters over 200m deep, with the lack of light at these depths likely explaining the lack of eyes on the fossil as it being a benthic animal. Its many trunk appendages probably helped it swim, with the caudal furcae as stabilisers, with epipodites acting as respiratory organs and/or helping move water to other respiratory organs. Alongside this, the groove in the middle of the belly likely helped transfer food up to the mouth, with the enditic membranes possibly helping contain this food. The pleural facets and articulating half-rings suggest it was likely able to flex its trunk up and down, with the spines probably being defensive in nature. Despite having large gnathobases on its mandibles, its blindness and lack of elongated sensory organs suggests it was likely a scavenger
Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...
.[
]
Etymology
''Tanazios'' is a combination of the words "tanaos", meaning "long", "zoon", meaning "animal" and "pelagios", meaning "of the sea". The species name ''dokeron'' is a combination of "dodeka" ("twelve"), "kerouchos" ("horned") and "epikranon" ("helmet").
Classification
''Tanazios'' was originally considered as stem-group 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 ...
. According to phylogenic analysis in Pulsipher ''et al.'' (2022), it is considered as stem-mandibulate instead.
References
{{Taxonbar, from=Q123453090
Silurian arthropods of Europe
Fossils of England
Enigmatic arthropod taxa
Homerian