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''Willwerathia'' is a
As a synziphosurine, ''Willwerathia'' is unusually large and so far the largest known synziphosurine, with largest
genus
Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of synziphosurine, a paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
group of horseshoe crab
Horseshoe crabs are marine and brackish water arthropods of the family Limulidae and the only living members of the order Xiphosura. Despite their name, they are not true crabs or crustaceans: they are chelicerates, most closely related to arachn ...
-like fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
chelicerate
The subphylum Chelicerata (from New Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. It contains the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, ticks, and mite ...
arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s. ''Willwerathia'' known only by one species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''Willwerathia laticeps'', discovered in deposits
A deposit account is a bank account maintained by a financial institution in which a customer can deposit and withdraw money. Deposit accounts can be savings accounts, Transaction account#Current accounts, current accounts or any of several othe ...
of the Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
period
Period may refer to:
Common uses
* Era, a length or span of time
* Full stop (or period), a punctuation mark
Arts, entertainment, and media
* Period (music), a concept in musical composition
* Periodic sentence (or rhetorical period), a concept ...
from the Klerf Formation
The Klerf Formation is an Early Devonian (Emsian) formation that includes a Lagerstätte in the Northern Eifel hills, at Willwerath near Prüm, Rhineland-Palatinate, Germany. In it ''Jaekelopterus rhenaniae'', a giant eurypterid was discovered. T ...
, in the Rhenish Slate Mountains
The Rhenish Massif, Rhine Massif or Rhenish Uplands (german: Rheinisches Schiefergebirge, : 'Rhenish Slate Uplands') is a geologic massif in western Germany, eastern Belgium, Luxembourg and northeastern France. It is drained centrally, south to n ...
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 ...
.
Morphology
As a synziphosurine, ''Willwerathia'' is unusually large and so far the largest known synziphosurine, with largest carapace
A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
measured about 90mm in width. Prosoma
The cephalothorax, also called prosoma in some groups, is a tagma of various arthropods, comprising the head and the thorax fused together, as distinct from the abdomen behind. (The terms ''prosoma'' and ''opisthosoma'' are equivalent to ''cepha ...
of ''Willwerathia'' covered by a vaulted carapace with pointed genal spines, recurved (M-shaped) ophthalmic ridges and pairs of dorsal nodes. Tergite
A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
s of the opisthosoma
The opisthosoma is the posterior part of the body in some arthropods, behind the prosoma (cephalothorax). It is a distinctive feature of the subphylum Chelicerata (arachnids, horseshoe crabs and others). Although it is similar in most respects to a ...
are either incomplete or disarticulated in available fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
materials, making it difficult to reveal the original number of opisthosomal segments. The opisthosoma of ''Willwerathia'' most likely compose of 10 segments, each expressed by a tergite that bore a median dorsal spine and a pair of tergopleurae (lateral extensions). The opisthosoma subdivided into a wider, most likely 7-segmented preabdomen and a narrower, 3-segmented postabdomen. tergite of the first opisthosomal segment is reduced in length while the remaining segments possess well-developed tergites with lateral nodes and posteriorly curved tergopleurae. The final segment terminated with a short, teardrop-shaped telson
The telson () is the posterior-most division of the body of an arthropod. Depending on the definition, the telson is either considered to be the final segment of the arthropod body, or an additional division that is not a true segment on accou ...
.
Paleoecology
''Willwerathia'' was most likely a bottom-dwellingpredator
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
. The marked articulation surfaces on each of the preabdominal segments suggest that ''Willwerathia'' capable to enroll itself in a way similar to '' Legrandella''. The environment in which ''Wilwerathia'' lived in was likely an estuarine
An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environment ...
to deltaic
A river delta is a landform shaped like a triangle, created by deposition of sediment that is carried by a river and enters slower-moving or stagnant water. This occurs where a river enters an ocean, sea, estuary, lake, reservoir, or (more rarel ...
one, and other animals like the largest eurypterid
Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 Myr, million yea ...
, ''Jaekelopterus
''Jaekelopterus'' is a genus of predatory eurypterid, a group of extinct aquatic arthropods. Fossils of ''Jaekelopterus'' have been discovered in deposits of Early Devonian age, from the Pragian and Emsian stages. There are two known species: th ...
'', are known from the same formation.
Classification
''Willwerathia'' was originally though to be aneurypterid
Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the Order (biology), order Eurypterida. The earliest known eurypterids date to the Darriwilian stage of the Ordovician period 467.3 Myr, million yea ...
(sea scorpion), with additional fossils described in 1998 reveal its synziphosurine affinities. In the redescription done by Anderson et al. 1998, ''Willwerathia'' had been grouped under the synziphosurine 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 ...
Weinberginidae alongside '' Weinbergina'' and '' Legrandella'', a classification which is not supported by phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis. ''Willwerathia'' was regarded as part of the monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
Xiphosura
Xiphosura () is an order of arthropods related to arachnids. They are more commonly known as horseshoe crabs (a name applied more specifically to the only extant family, Horseshoe crab, Limulidae). They first appeared in the Hirnantian (Late Ordo ...
''sensu stricto'' (true horseshoe crab) by Lamsdell 2013, but further phylogenetic analysis repeatedly resolving it within a clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
compose of Bunodids, Pseudoniscids and Dekatriata
Dekatriata is a clade of planatergan chelicerates including the groups Arachnida, Chasmataspidida, Eurypterid
Eurypterids, often informally called sea scorpions, are a group of extinct arthropods that form the order Eurypterida. The earliest ...
n (chasmataspidid
Chasmataspidids, sometime referred to as chasmataspids, are a group of extinct chelicerate arthropods that form the order Chasmataspidida. Chasmataspidids are probably related to horseshoe crabs ( Xiphosura) and/or sea scorpions ( Eurypterida), wi ...
s, eurypterids and arachnid
Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaroon ...
s).
References