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Pseudastacus
Temporal range: Sinemurian–Cenomanian
Fossil of Pseudastacus pustulosus
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Malacostraca
Order: Decapoda
Suborder: Pleocyemata
Family: †Stenochiridae
Genus: †Pseudastacus
Oppel, 1861
Type species
†Bolina pustulosa
Münster, 1839
Species
  • †P. lemovices Charbonnier & Audo, 2020
  • †P. minor? Fraas, 1878
  • †P. mucronatus Phillips, 1835
  • †P. pusillus? Van Straelen, 1925
  • †P. pustulosus Münster, 1839
Synonyms
synonyms of Pseudastacus
  • Alvis Münster, 1840
  • Bolina Münster, 1839
synonyms of P. pustulosus
  • Alvis octopus Münster, 1840
  • Bolina pustulosa Münster, 1839
  • Pseudastacus muensteri Oppel, 1861
synonyms of P. mucronatus

Pseudastacus (meaning "false Astacus", in comparison to the extant crayfish genus) is an extinct genus of decapod crustaceans that lived during the Jurassic period in Europe, and possibly the Cretaceous period in Lebanon. Many species have been assigned to it, though the placement of some species remain uncertain and others have been reassigned to different genera. Fossils attributable to this genus were first described by Georg zu Münster in 1839 under the name Bolina pustulosa, but the generic name was changed in 1861 after Albert Oppel noted that it was preoccupied. The genus has been placed into different families by numerous authors, historically being assigned to Nephropidae or Protastacidae. Currently, it is believed to be a member of Stenochiridae.

Not exceeding 6 cm (2.4 in) in total length, Pseudastacus was a small animal. Members of this genus have a crayfish-like build, possessing long antennae, a triangular rostrum and a frontmost pair of appendages enlarged into long and narrow pincers. Deep grooves are present on the carapace, which is around the same length as the abdomen. The carapace is usually uneven, with either small tubercles or pits across the surface. Sexual dimorphism is known in Pseudastacus, with the pincers of the females being more elongated than those of the males. There is evidence of possible gregarious behavior in the form of multiple individuals preserved alongside each other, possibly killed in a mass mortality event.

With the oldest known record dating to the Sinemurian age of the early Jurassic, and possible species surviving into the Cenomanian stage of the late Cretaceous, Pseudastacus has a long temporal range and was a widespread taxon. Fossils of this animal were first found in the Solnhofen Limestone of Germany, but have also been recorded from France, England and Lebanon. All species in this genus lived in marine environments.

Discovery and naming

Figure from Münster (1839) labelled as Bolina pustulosa
Fossil of Pseudastacus pustulosus as illustrated in Münster's 1840 paper, originally labelled as Alvis octopus

Fossils of Pseudastacus had been described prior to the naming of this genus, under other names which are currently invalid. In 1839, Georg zu Münster established the genus Bolina to include two species, B. pustulosa (the type species) and B. angusta, both of which are based on specimens collected from the Solnhofen Limestone. The generic name references the nymph Bolina from Greek mythology.[1] A year later, Münster described several fossils from the Solnhofen Limestone he believed to represent isopods, and erected the genus Alvis to contain the single species A. octopus, naming it after the dwarf Alvíss from Norse mythology.[2]

In 1861, Albert Oppel noted that the name Bolina was preoccupied by a genus of cnidarian, and thus the crustacean named by Münster had to be renamed. Oppel placed B. pustulosa and B. angusta into two new genera, Pseudastacus and Stenochirus respectively. Now renamed as Pseudastacus pustulosus and Stenochirus angustus, the two species became the type species of their own respective genera. The name Pseudastacus means "false Astacus", referencing its resemblance to the modern crayfish genus.[3] Oppel declared that 10 specimens known at the time represented Pseudastacus pustulosus, of which one was from the Redenbacher collection and the remaining nine were from the collection of the Palaeontological Museum, Munich. In addition, he identified one specimen (BSPG AS I 672) housed in the Palaeontological Museum as a second species of the genus which he named Pseudastacus muensteri.[4] In 2006, Garassino and Schweigert reviewed the decapod fossils from Solnhofen and found that four of the P. pustulosus specimens from Oppel's collection were still present, and that P. muensteri is a junior synonym of P. pustulosus.[5]

Valid species

Several species have been assigned to the genus Pseudastacus, though the placement of some species remains uncertain or tentative. In addition, some have since been moved into different genera after they were discovered not to be closely related to the type species. In 2020, Charbonnier and Denis published a study including a summary of recognized stenochirid species, which covered the reclassification of former Pseudastacus species and left the following as members of the genus:[6]

Illustration of the P. mucronatus type specimen
  • P. mucronatus was originally named as Astacus mucronatus by John Phillips in 1835. The type specimen was extracted from the Speeton Clay Formation in Yorkshire, England, and is a fragment of the pincer. The chela is very large, with alternating large and small tubercles on the inner margins.[7] This is unlike the narrower and longer pincers of other Pseudastacus species, and the specimen may be referrable to Hoploparia dentata.[8]
  • P. minor was described by Oscar Fraas in 1878 from a specimen found in Cenomanian-aged deposits in Lebanon.[9] This specimen is now lost and only the original illustration remains, which shows features unlike any other Pseudastacus species: the rostrum is extremely long, there is an additional abdomen segment, the clawed limbs are placed further back and the general pincer shape is different. Its placement in this genus is thus uncertain.[6]
  • P. pusillus is based on a fossil from the Bajocian-aged deposits of May-sur-Orne, France described in 1925 by Victor van Straelen.[10] The fossil was destroyed in World War II and it is difficult to tell from the original line drawing of the specimen whether this species truly belongs to Pseudastacus.[6]
  • P. lemovices was named in 2020 based on five specimens preserved in a slab of Sinemurian-aged limestone, collected from Chauffour-sur-Vell, France. The specific name honors the Lemovices, a Gallic tribe that lived near this locality. It is the oldest known species of the family Stenochiridae.[6]

Reassigned species

The following species were formerly placed in Pseudastacus, but have now been moved to different genera.

Description

Reconstructions of P. pustulosus

Pseudastacus is a small invertebrate, with the carapace of P. lemovices reaching a length of 11 mm (0.43 in) (excluding the rostrum) and a height of 6.5 mm (0.26 in).[6] The known specimens of P. pustulosus range from 4–6 cm (1.6–2.4 in) in total length.[5]

Members of this genus often have an uneven carapace surface, with some species (such as P. pustulosus) having tubercles and others (such as P. lemovices) having pits distributed uniformly across the carapace surface. Individuals with smoother carapaces are also documented, though this may be due to abrasion. Grooves are present on the carapace, including a deep, arch-shaped cervical groove that stretches across the top and sides of the carapace, and an additional groove behind it on either side. The rostrum is triangular and elongated, with three lateral spines.[6][5] The carapace and head are separated by an arch-shaped incision. A pair of long antennae and two pairs of shorter antennules extend from the head, with the outer antennules being slightly narrower and more pointed than the outer pair.[2] A pair of compound eyes are attached to the head by short eye stalks.[5]

The first three pairs of appendages terminate with chelae (pincers), and the appendages furthest front are particularly long and enlarged. The abdomen is around the length of the carapace, with the frontmost segment being the smallest. The pereiopods (walking legs) on the thorax decrease in size the further back they are placed, the pair furthest front being largest and longest. The uropods are equal in length, with a ridge down the middle and long setae on the margins.[6]

Classification

In the centuries since it was first discovered, Pseudastacus has been placed in a wide variety of families by many different authors. For many decades, the genus was thought to be a member of Nephropidae (the lobster family), as first reported by Victor van Straelen in 1925.[10] This placement was followed by subsequent authors such as Beurlen (1928), Glaessner (1929), and Chong & Förster (1976).[15][16][17] In 1983, Henning Albrecht erected the family Protastacidae and moved Pseudastacus into it, whereas Tshudy & Babcock (1997) included the genus into their newly-established family Chilenophoberidae.[18][19] Although Garassino & Schweigert (2006) continued to place Pseudastacus in Proastacidae following Albrecht (1983), other authors in the 2000s would place it in Chilenophoberidae based on the more recent findings of Tshudy & Babcock (1997).[5][20][21]

In 2013, Karasawa et al. recovered Pseudastacus as the sister taxon to Stenochirus, making Chilenophoberidae a paraphyletic group. The family was therefore synonymized with Stenochiridae. The following cladogram shows the placement of Pseudastacus within Stenochiridae according to the study:[22]

Stenochiridae

Palaeobiology

Sexual dimorphism

Illustrations of male (left) and female (center and right) P. pustulosus

Albert Oppel noticed that Pseudastacus fossils from the Solnhofen Limestone could be distinguished into two morphs; aside from those most similar to the P. pustulosus type specimen, there were also some with smaller bodies and longer, more slender claws. Oppel believed the latter morph to be a separate species which in 1862 he named P. muensteri.[4] Over a century later, Garassino and Guenter (2006) found that specimens of P. muensteri were essentially identical to P. pustulosus aside from the claw form. In addition, they noted that in fossil glypheids and the extant Neoglyphea inopinata, the females possess longer clawed limbs than the males. Based on this, they declared P. muensteri as a junior synonym of P. pustulosus, actually representing female specimens of the species.[5]

Social behavior

The type series of P. lemovices is made up of five individuals preserved together in a single limestone slab, possibly indicating the species exhibited gregarious behaviour, with this group being killed in a mass mortality event (perhaps caused by temperature changes or lack of oxygen).[6] Evidence of gregarious behaviour is also known in other fossil lobsters, as well as in extant species.[23][24]

Palaeoenvironment

Early Jurassic

Pseudastacus is believed to have first evolved during the early Jurassic, with P. lemovices being the oldest member of the genus currently known. The five known specimens of this species were preserved in a single limestone slab collected from a garden in Chauffour-sur-Vell, France. This sediment in this locality represents a marine environment dating back to the Sinemurian age, and the general area has been specifically dated to the late Sinemurian based on the presence of the green alga Palaeodasycladus mediterraneus in a regional bed.[6]

Late Jurassic

Pseudastacus pustulosus, the type species of the genus, is also known from the most specimens. All known remains of this species were collected from the Solnhofen Limestone of Bavaria, Germany, which dates to the Tithonian age of the late Jurassic period. During the time of deposition, the European continent was partly inundated, forming a dry, tropical archipelago at the edge of the Tethys Ocean. The Solnhofen Limestone would have been laid down in a lagoonal environment cut off from the main ocean by reefs.[25] A coastal habitat is further confirmed by the fossil content of the area, which includes numerous marine species that P. pustulosus would have lived alongside. These include cephalopods (such as ammonoids and belemnites),[26] crinoids (such as Saccocoma),[27] other crustaceans (including eryonids, axiids, glypheids, mantis shrimps, and the closely related Stenochirus),[5] fish (such as pycnodonts, pachycormids, aspidorhynchids and caturids)[28] and marine reptiles (such as turtles, ichthyosaurs and metriorhynchids).[29][30][31] Remains of terrestrial animals, though rarer, are also present and represent species that would have lived on the islands surrounded by the lagoons, including dinosaurs (such as Archaeopteryx and Compsognathus),[32] lizards (such as Ardeosaurus, Bavarisaurus and Schoenesmahl),[33] and pterosaurs.[34]

Cretaceous

Two Pseudastacus species, P. mucronatus and P. minor, originate from deposits dating to the Cretaceous period, though their assignment to this genus remains uncertain. These two species did not coexist, being from different stages of the Cretaceous as well as different locations. Known remains of P. mucronatus have been collected from the Speeton Clay Formation in England, which extends from the Berriasian to Aptian ages of the early Cretaceous.[35] The formation was a marine environment that initially was deposited during a period of marine transgression, which later transitioned into an event of marine regression, correlating to sequences of warmer and cooler temperatures. This is reflected in the formation's interbedded layers of mudstones and clays.[36] The Speeton Clay Formation preserves fossilized remains of various marine animals, with those of belemnites being the most abundant.[37] Ammonites, crustaceans, and the teeth of sharks and rays (including Cretorectolobus, Spathobatis, Dasyatis and Synechodus) are also commonly recorded from these deposits.[7][38][39]

Known from a single (currently missing) specimen from the Cenomanian-aged marine deposits of Lebanon, P. minor would be the geologically youngest species of Pseudastacus, assuming it does belong to the genus. During this age, Lebanon was located on a large carbonate platform mostly submerged in the Neotethys Ocean, and located near the northeastern edge of the Afro-Arabian continent.[40] Plant fossils from Cenomanian Lebanese deposits (including gymnosperms and deciduous angiosperms) indicate a similar climate to the modern-day Mediterranean Basin, and are similar to floral assemblages from contemporary Crimea, North America and Central Europe.[41] The paleontological sites of Lebanon have yielded many well-preserved fossils, including a wide variety of fish, crustaceans and even octopuses.[42][43][44] Terrestrial insects and reptiles (including pterosaurs and squamates) are also represented in the fossil finds from these deposits.[40][45][46]

References

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