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''Seppo'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
spider Spiders (order (biology), order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude spider silk, silk. They are the largest order of arachnids and ran ...
s, possibly of the superfamily
Palpimanoidea The Palpimanoidea or palpimanoids, also known as assassin spiders, are a group of araneomorph spiders, originally treated as a superfamily. As with many such groups, its circumscription has varied. , the following five families were included: * ...
, that lived about 180 million years ago, in the Early Jurassic (Lower
Toarcian The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...
) of what is now
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
. The sole species ''Seppo koponeni'' is known from a single fossil from
Grimmen Grimmen (; ) is a town in Vorpommern-Rügen, a district in the States of Germany, Bundesland Mecklenburg-Vorpommern, Germany. Prior to 2011, when district reforms were made in Mecklenburg-Vorpommern, it was the capital of the now bygone Nordvorpom ...
, Germany. With the scorpion '' Liassoscorpionides'', it is one of the two only known arachnids from the Lower Jurassic of
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 ...
. ''Seppo'' is the first unequivocal Early Jurassic spider, and was recovered from the Grimmen Formation.


Description

The spider was described from a single female specimen. It is unknown if was an adult. The
carapace A carapace is a 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 tortoises, the unde ...
is unknown, and besides that has preserved bowed converging sides with a curved posterior margin with straight posterior border of the labium, with row of at least 12 peg teeth along the cheliceral furrow, no true teeth, scattered setae on anterior surface, and slender
pedipalps Pedipalps (commonly shortened to palps or palpi) are the secondary pair of forward appendages among Chelicerata, chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to ...
. Legs are preserved, the first and second being much longer than the third and fourth. All are well covered in setae and bristles, especially on the tibiae and metatarsi of leg I. It most likely belongs to the
Palpimanoidea The Palpimanoidea or palpimanoids, also known as assassin spiders, are a group of araneomorph spiders, originally treated as a superfamily. As with many such groups, its circumscription has varied. , the following five families were included: * ...
, based on the presence of cheliceral peg teeth.


Discovery

The single known specimen was found on locality known for its fossil insects in
Grimmen Grimmen (; ) is a town in Vorpommern-Rügen, a district in the States of Germany, Bundesland Mecklenburg-Vorpommern, Germany. Prior to 2011, when district reforms were made in Mecklenburg-Vorpommern, it was the capital of the now bygone Nordvorpom ...
, near
Greifswald Greifswald (), officially the University and Hanseatic City of Greifswald (, Low German: ''Griepswoold'') is the fourth-largest city in the German state of Mecklenburg-Western Pomerania after Rostock, Schwerin and Neubrandenburg. In 2021 it surpa ...
, at the north of
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 ...
. It was reported in 2003, when Ansorge did a recompilation of insect taxa on the
Toarcian The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...
strata of Germany and
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
. The specimen was recovered from the Falciferum zone (Exaratum subzone), c. 180 Ma, and presented as "
Araneae Spiders ( order Araneae) are air-breathing arthropods that have eight limbs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
gen. et sp. nov." (
Chelicerata The subphylum Chelicerata (from Neo-Latin, , ) constitutes one of the major subdivisions of the phylum Arthropoda. Chelicerates include the sea spiders, horseshoe crabs, and arachnids (including harvestmen, scorpions, spiders, solifuges, tic ...
). It was recovered from a fragment of calcareous nodule from the grey-green
claystone Mudrocks are a class of fine-grained siliciclastic sedimentary rocks. The varying types of mudrocks include siltstone, claystone, mudstone and shale. Most of the particles of which the stone is composed are less than and are too small to ...
, recovered from the closed since the 90's clay pit of Klein Lehmhagen, near
Grimmen Grimmen (; ) is a town in Vorpommern-Rügen, a district in the States of Germany, Bundesland Mecklenburg-Vorpommern, Germany. Prior to 2011, when district reforms were made in Mecklenburg-Vorpommern, it was the capital of the now bygone Nordvorpom ...
, Western
Pomerania Pomerania ( ; ; ; ) is a historical region on the southern shore of the Baltic Sea in Central Europe, split between Poland and Germany. The central and eastern part belongs to the West Pomeranian Voivodeship, West Pomeranian, Pomeranian Voivod ...
, Germany. It is curious for being one of the rare few examples of spiders found on calcium carbonate (along with others from the
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
limestone of the
Isle of Wight The Isle of Wight (Help:IPA/English, /waɪt/ Help:Pronunciation respelling key, ''WYTE'') is an island off the south coast of England which, together with its surrounding uninhabited islets and Skerry, skerries, is also a ceremonial county. T ...
,
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
). The specimen was found with some parts preserved as external moulds, and these show
seta In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...
tion and spination. This is an exception, since most of the specimen is an internal mould of calcium carbonate. The specimen was labeled with the number MB.A 2966, and deposited in the
Museum für Naturkunde The Natural History Museum () is a natural history museum located in Berlin, Germany. It exhibits a vast range of specimens from various segments of natural history and in such domain it is one of three major museums in Germany alongside Naturm ...
,
Berlin Berlin ( ; ) is the Capital of Germany, capital and largest city of Germany, by both area and List of cities in Germany by population, population. With 3.7 million inhabitants, it has the List of cities in the European Union by population withi ...
, being named in 2014 by Paul A. Selden & Jason A. Dunlop ''Seppo koponeni'', after the Finnish
arachnologist Arachnology is the scientific study of arachnids, which comprise spiders and related invertebrates such as scorpions, pseudoscorpions, harvestmen, ticks, and mites. Those who study spiders and other arachnids are arachnologists. More narrowly ...
, to celebrate his 70th birthday.


Phylogeny

The genus was described as the oldest
Palpimanoidea The Palpimanoidea or palpimanoids, also known as assassin spiders, are a group of araneomorph spiders, originally treated as a superfamily. As with many such groups, its circumscription has varied. , the following five families were included: * ...
or the nearest sister taxon to the group. The exact familiar placement of the genus is unclear, as it was assigned to the family based on the presence of peg teeth, a
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
thought to be unique to the superfamily. However, recent revisions of modern spiders have shown that this character is also present in the Mimetidae and other
Entelegynae The Entelegynae or entelegynes are a subgroup of araneomorph spiders, the largest of the two main groups into which the araneomorphs were traditionally divided. Females have a genital plate ( epigynum) and a "flow through" fertilization system; ...
families. Early araneomorphs such as those in the clade
Synspermiata Synspermiata is a clade of araneomorph spiders, comprising most of the former " haplogynes". They are united by having simpler genitalia than other araneomorph spiders, lacking a cribellum, and sharing an evolutionary history of synspermia – a ...
and the families
Filistatidae Crevice weaver spiders (Filistatidae) comprise cribellate spiders with features that have been regarded as " primitive" for araneomorph spiders. They are weavers of funnel or tube webs. The family contains 18 genera and more than 120 described s ...
,
Austrochilidae Austrochilidae is a small spider family with nine species in two genera. ''Austrochilus'' and ''Thaida'' are endemic (ecology), endemic to the Andean forest of central and southern Chile and adjacent Argentina. Taxonomy , two genus, genera are p ...
and
Leptonetidae Leptonetidae is a family of small spiders adapted to live in dark and moist places such as caves. The family is relatively primitive having diverged around the Middle Jurassic period. They were first described by Eugène Simon in 1890. Distingu ...
lack it, locating the genus as either a stem Palpimanoidea or a stem Entelegynae.


Ecology

''Seppo koponeni'' is one of the only two arachnids ever to have been found in the
Toarcian The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...
rocks of north Germany, outnumbered by several thousand insect specimens at several localities. The different between the number of insects as opposed to arachnids has not been studied in depth. A possibility is that because insects can fly over water, they fall into it far more easily than spiders. It is unclear how the spider ended on a marine clay deposit, far from the land, although several theories have been suggested:
ballooning Ballooning may refer to: * Hot air ballooning * Balloon (aeronautics) * Ballooning (spider) * Ballooning degeneration, a disease * Memory ballooning In computing, memory ballooning is a technique that is used to eliminate the need to overcommit ...
is a possibility as a method of transport, perhaps helped by severe storms, hurricanes, and tornadoes. Another possibility might be that it was carried out to sea on floating vegetation, as wood remains have been recovered in the deposit. On nearby land, ground dwellers were represented by arachnids, such as ''Seppo'' and scorpions, but also by
Gryllidae The family Gryllidae contains the subfamilies and genera which entomologists now term true crickets. Having long, whip-like antennae, they belong to the Orthopteran suborder Ensifera, which has been greatly reduced in the last 100 years (''e.g ...
, such as '' Protogryllus dobbertinensis'',
grylloblattodea Grylloblattidae, commonly known as the icebugs or ice crawlers, is a family of extremophile (psychrophile) and wingless insects that live in the cold on top of mountains and the edges of glaciers. It is the only member of Grylloblattodea, which ...
ns such as '' Nele jurassica'', and dermapterans. These arthropods never moved far from where they lived and are generally very rare in the rocks. ''Seppo'' shows a rather unusual morphology, with large and porrect
chelicerae The chelicerae () are the arthropod mouthparts, mouthparts of the subphylum Chelicerata, an arthropod group that includes arachnids, horseshoe crabs, and sea spiders. Commonly referred to as "jaws", chelicerae may be shaped as either articulated ...
and a robust leg I, contrasted with a short leg III. The robust and well-armed first legs, directed forwards, give the impression that they were prey capture appendages, a morphology typical of a sit-and-wait predator, while the short third legs are more typical of web spiders, especially orbweavers, but also palpimanoids. Short third legs are not usually found on spiders that are substrate dwellers, which have more equal legs. ''Seppo'' was probably not a habitual ground dweller. The armoured front legs related to capturing dangerous prey are typical of many extant palpimanoids that are araneophagous.


References

{{Taxonbar, from1=Q20722585, from2=Q24238763 Mesozoic arachnids
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...
Fossils of Germany Fossil taxa described in 2014 Jurassic arachnids
A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species or languages). It is one of the mo ...