Flagellopantopus
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''Flagellopantopus'' is an extinct genus of
pycnogonid Sea spiders are marine arthropods of the class Pycnogonida, hence they are also called pycnogonids (; named after ''Pycnogonum'', the type genus; with the suffix '). The class includes the only now-living order Pantopoda ( ‘all feet’), alongs ...
(sea spider)
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
known from the lower
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 ...
aged
Hunsrück Slate The Hunsrück () is a long, triangular, pronounced upland in Rhineland-Palatinate, Germany. It is bounded by the valleys of the Moselle-Saar (north-to-west), the Nahe (south), and the Rhine (east). It is continued by the Taunus mountains, past ...
. A single species is currently known, ''Flagellopantopus blocki'', which was described from the
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 Kaub Formation 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 ...
. ''F. blocki'' differs from other members of the Pycnogonida by possessing an extremely elongated
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 ...
, which was flagelliform in appearance. This animal was fairly large for a pycnogonid, with the whole body of the arthropod measuring around long, while the telson itself was around long. In its 2006 description, Poschmann and Dunlop placed ''Flagellopantopus'' into the order
Pantopoda Sea spiders are marine arthropods of the class Pycnogonida, hence they are also called pycnogonids (; named after ''Pycnogonum'', the type genus; with the suffix '). The class includes the only now-living order Pantopoda ( ‘all feet’), alongs ...
(which includes modern sea spiders). However, later publications have put doubt on this interpretation, and have suggested a new placement as a member of Pycnogonida ''
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''. It is currently assumed to have lived a
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". ...
lifestyle, as it lacks the flattened appendages and
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, ...
e that
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 ...
species of pycnogonids typically have.


Discovery and naming

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 ...
(and only known) specimen of this pycnogonid, NHMMZ PWL 2004/5024-LS, was first described by Poschmann in 2004 for Project Nahecaris, but was considered a ''
nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
'' due to the lack of a diagnosis. The specimen would later be given a proper diagnosis and description in 2006. The holotype specimen originates from the Kaub Formation, which is dated to the Emsian stage of the lower Devonian, but the exact locality of origin is uncertain. Poschmann and Dunlop (2006) state that the fossil was recovered from the Wingertshell Member, within the larger Obereschenbach quarry, near the
municipality A municipality is usually a single administrative division having municipal corporation, corporate status and powers of self-government or jurisdiction as granted by national and regional laws to which it is subordinate. The term ''municipality' ...
of Bundenbach. However, the administration housing the fossil, the Naturhistorisches Museum Mainz, states that the specimen originates from the Eschenbach-Bocksberg quarry. The arthropod's genus name, ''Flagellopantopus'', is derived from the Latin word ''flagella'', meaning "whip", which was named after the elongated telson possessed by the genus, and the pycnogonid order Pantopoda, which is derived from the Latin words ''pant'' and ''poda'', meaning "all feet". The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, ''blocki'', is in honour of Helmut Block, who donated the holotype for public viewing and research.


Description

''Flagellopantopus'' was relatively large compared to most other pycnogonids, with the main body (excluding telson) of the arthropod measuring around long. There are 5 dorsal tubercles located across the cephalon and trunk segments, with the first one represent ocular tubercle. By far the most distinguishing trait of this taxon was its extremely elongated
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 ...
(flagella), which measured around long, nearly three times longer than the arthropods main body. The telson was preceded by a 4-segmented abdomen, which possess a pair of long projections at the second abdominal segment. Like most other pycnogonids, this genus possessed four pairs of walking legs that had at least a arm-span. Each leg possess an annulated coxa 1 and immobile junction between segment 4-5 similar to ''
Palaeopantopus ''Palaeopantopus'' is a monotypic genus of fossil pycnogonid (sea spider), known only by one species, ''Palaeopantopus maucheri'' from the Devonian Hunsrück Slate of Germany. Description ''Palaeopantopus'' is known from 3 specimens, one o ...
'', while the distal regions (tibia and tarsal region) are not preserved. Coxa 1 of the first leg pair possess a long projection. All 3 pairs of cephalic appendages ( chelifores,
palps Pedipalps (commonly shortened to palps or palpi) are the secondary pair of forward appendages among chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicera ...
, ovigers) are evident, but most of the details are obscured by their poor preservation. Distal region of the oviger showing 6 segments (possibly patella, tibia, and 4 tarsal segments) and a terminal claw. The
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
of this taxon is partially obscured, and how it connected to the rest of the cephalic region is uncertain. The proboscis itself had a pyriform appearance and was roughly long, and had a width of around wide.


Classification

Although they are often colloquially known as "sea spiders", pycnogonids are not closely related to them and other
arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...
s. However, their placement as basal members of the
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 ...
subphylum means that they are more closely related to them then to other groups. ''Flagellopantopus'' is confidently classified within the Pycnogonida, however its further placement is uncertain. Both Poschmann (2004) and Poschmann and Dunlop (2006) placed this taxon as a relatively basal member of the Pantopoda order (which includes all
crown-group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
sea spiders) on the basis of its long telson, which is lost in more derived taxa. However, they classified this taxon as more derived than another contemporary pycnogonid, ''
Palaeoisopus problematicus ''Palaeoisopus'' is a genus of fossil pycnogonid (sea spider). The only known species is ''Palaeoisopus problematicus'' from the Lower Devonian Hunsrück Slate of Germany. It is characterized by several features unusual for a pycnogonid, such as ...
'', due to ''Flagellopantopus'' possessing a less well developed abdomen, which is a derived trait in crown-group pycnogonids. However, later studies, including Sabroux ''et al''. (2024) have suggested that not only was ''Flagellopantopus'' not a member of Pantopoda, but the entire order did not exist during the Devonian. They instead classified this taxon, and the other known Hunsrück pycnogonids into two different morphological groups separate from the Pantopoda.


Paleoecology

The Hunsrück Slate is deposited within the regions of
Hunsrück The Hunsrück () is a long, triangular, pronounced mountain range, upland in Rhineland-Palatinate, Germany. It is bounded by the valleys of the Moselle (river), Moselle-Saar (north-to-west), the Nahe (south), and the Rhine (east). It is continued ...
and
Taunus The Taunus () is a mountain range in Hesse and Rhineland-Palatinate, Germany, located north west of Frankfurt and north of Wiesbaden. The tallest peak in the range is '' Großer Feldberg'' at 878 m; other notable peaks are '' Kleiner Feldberg' ...
within the country of Germany. Although debates still stand about the exact depth of the ecosystem during the Devonian, most recent studies have suggested a maximum depth of around deep, within the
Epipelagic zone The photic zone (or euphotic zone, epipelagic zone, or sunlight zone) is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological ...
. Due to the fact that ''Flagellopantopus'' possessed chelifores, it is likely that it was not restricted to feeding on
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
organisms, which is common in modern representatives. It is also likely that it lived a fully benthic lifestyle, as it lacks the broad legs and long setae usually associated with swimming pycnogonids. The telson of this taxon most likely would've acted as a sensory structure, as it was probably too stiff to help with locomotion in the water column. A plethora of other organisms are known to lived alongside ''Flagellopantopus'', including other arthropods,
echinoderm An echinoderm () is any animal of the phylum Echinodermata (), which includes starfish, brittle stars, sea urchins, sand dollars and sea cucumbers, as well as the sessile sea lilies or "stone lilies". While bilaterally symmetrical as ...
s,
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 ...
s,
agnatha Agnatha (; ) or jawless fish is a paraphyletic infraphylum of animals in the subphylum Vertebrata of the phylum Chordata, characterized by the lack of jaws. The group consists of both extant taxon, living (Cyclostomi, cyclostomes such as hagfish ...
ns, and various other groups. Some notable examples of contemporary genera include the 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 ...
''
Schinderhannes bartelsi ''Schinderhannes bartelsi'' is a fossil species of hurdiid radiodont (anomalocaridid), known from a single specimen from the Lower Devonian Hunsrück Slate in Germany. Its discovery expanded the known range of radiodonts, the latest members of ...
'', the marrellomorphid arthropods '' Mimetaster hexagonalis'' and '' Vachonisia rogeri'', as well as the various placoderms of the bizarre
Pseudopetalichthyida Pseudopetalichthyida is an order of lightly armored placoderms known only from rare fossils in Lower Devonian strata in Hunsrück, Germany. Like '' Stensioella heintzi'', and the Rhenanida, the Pseudopetalichthids had armor made up of a mosaic o ...
order (which are only found at Hunsrück). Also found in the area are various other pycnogonids, including ''Palaeopantopus maucheri'', ''Pentapantopus vogteli'', ''Palaeothea devonica'', and the aforementioned ''Palaeoisopus problematicus''. The level of pycnogonid genera found at the Hunsrück Slate makes it the most diverse assemblage of fossil pycnogonids yet discovered.


Notes


References

{{Taxonbar, from=Q107252800 Pycnogonid genera Devonian arthropods of Europe Fossils of Germany Hunsrück Slate fossils Fossil taxa described in 2006 Monotypic prehistoric arthropod genera Monotypic chelicerate genera