Perimecturus

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Perimecturus
Temporal range: Early Carboniferous (Viséan to Serpukhovian), 345–324  Ma
Ar1050 Perimecturus rapax Unt Karbon Bear Gulch Montana USA.jpg
Fossil of P. rapax from the Bear Gulch Limestone
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Malacostraca
Order: Stomatopoda
Family:† Perimecturidae
Genus:† Perimecturus
Peach, 1908
Type species
† Anthrapalaemon parki
(= †Perimecturus parki)
Peach, 1882
Other species
  • †P. rapax
    Schram & Horner, 1978
Synonyms
Synonyms of Perimecturus
Synonyms of P. parki
  • Anthrapalaemon parkiPeach, 1882
  • Palaesquilla parkiPeach & Horne, 1903 (nomen nudum)
  • Perimecturus communisPeach, 1908
  • Perimecturus parki duplicicarinatusPeach, 1908
  • Perimecturus stockiPeach, 1908

Perimecturus is an extinct genus of mantis shrimp that lived during the Early Carboniferous period in what is now Scotland and the United States. The first known specimens were collected near the River Esk in Glencartholm, Scotland, and the genus was named in 1908 by Ben Peach, making it the second genus of Paleozoic mantis shrimp to be described (only after Archaeocaris ). While many species have been classified in the genus since then, taxonomic revisions in the late 20th and 21st centuries have reassigned most of these to different genera, leaving two named species currently assigned to this genus. The type species, P. parki, was first named in 1882 as a species of Anthrapalaemon and is known from the Viséan-aged Glencartholm Volcanic Beds of Scotland. Fossils of a later species, P. rapax, have been found in the Bear Gulch Limestone of Montana and were first described by Frederick Schram.

Unlike other Paleozoic mantis shrimps which had narrow, shrimp-like bodies, Perimecturus had a wide and flattened body more closely resembling that of a lobster. This condition is also seen in modern mantis shrimps, but Perimecturus would have developed this shape separately from them in an example of parallel evolution. Members of this genus had a wide carapace with five ridges, in addition to six other ridges running down the abdomen. The two species differ primarily in the tail fan, with P. rapax having a longer telson spike and lacking the bristles seen in P. parki. In addition, P. parki lacks the serrations on the telson and uropods that P. rapax has. Perimecturus is one of the largest Paleozoic mantis shrimps, with the largest known specimen of P. parki reaching a total length of 15 cm (5.9 in). However, most specimens are notably smaller than this.

Both Perimecturus species lived in fully marine habitats, and are believed to be intolerant of lower salinities. Like all other mantis shrimps, they would have been carnivores that handled prey with their raptorial thoracic appendages. Because the walking appendages are poorly preserved, it remains unclear whether Perimecturus would have been benthopelagic and swam just over the seabed like other early mantis shrimps.

Discovery and naming

Illustrations of the original Perimecturus parki fossils from Peach's 1882 paper, originally labeled as Anthrapalaemon parki Anthrapalaemon parki.jpg
Illustrations of the original Perimecturus parki fossils from Peach's 1882 paper, originally labeled as Anthrapalaemon parki

The first fossil of Perimecturus known to science was discovered by A. Macconochie, with additional specimens found by Walter Park. These specimens were collected near the River Esk in Glencartholm, Scotland, and turned in to the Geological Survey of Scotland. In 1882, British paleontologist Ben Peach became the first person to study these fossils, which he believed represented a new species of Anthrapalaemon , naming it Anthrapalaemon parki after Park. [1] Later, Peach would work alongside J. Horne in 1903 to move this species to a separate genus which they called Palaesquilla, however they did not formally rename it and thus Palaesquilla is deemed a nomen nudum. [2] A formal reassignment would be published by Peach in 1908, in which he erected the new genus Perimecturus with this species (now renamed as Perimecturus parki) as its type species. Furthermore, he also studied several other specimens which he named as a subspecies, P. parki duplicicarinatus, and additional species which he assigned to the genus, including P. stocki and P. communis. [3]

Perimecturus parki.jpg
Perimecturus parki duplicicarinatus.jpg
Perimecturus stocki.jpg
Perimecturus communis.jpg
Illustrations of P. parki fossils from Peach's 1908 paper, originally named as (in clockwise order from the top left) P. parki, P. parki duplicicarinatus, P. communis, and P. stocki respectively

In 1979, a review of the Carboniferous-aged malacostracans of Britain written by American paleontologist Frederick Schram was published. The review found that P. parki was the only valid species of Perimecturus named by Peach, and that P. p. duplicicarinatus (which Schram misspelled as duplicarinatus), P. stocki and P. communis were all junior synonyms of it. The specimen GSE 5896 was designated as the lectotype of this species. Schram stated that Peach's idea of the fossils as separate species was due to incorrect interpretation of the fossils preserved in different modes: the original P. parki holotype showed the animal in a top-down view, the P. p. duplicicarinatus specimen clearly preserved the longitudinal ridges, the P. stocki fossil is an individual in a slanted angle and the P. communis remains are displayed from the side. [4]

A second species of Perimecturus was erected by Schram in 1978 and named P. rapax, the specific name being a Latin word meaning "to grasp" (in reference to the animal's predatory lifestyle and raptorial appendages). The holotype for this species, UM 6141, is held in the collection of the University of Montana and was discovered in the Bear Gulch Limestone of Fergus County, Montana. [5] In 1985, David Factor and Rodney Feldmann published a redescription of P. rapax, after a detailed reanalysis of the fossils which found that Schram had misinterpreted some of their features. [6]

Reassigned species

Because it was the second Paleozoic mantis shrimp genus to be named (after Archaeocaris , which was named in 1872 but only recognized as one in 1962), [7] numerous species have been assigned to Perimecturus before better knowledge of early mantis shrimps was developed. The following species were formerly placed in Perimecturus but have since been moved to other genera:

Description

The body of Perimecturus is distinctively flattened, appearing wide and almost rectangular when viewed from above, and its abdomen could be held straight. [6] This makes its body partly lobster-like, differing from other Paleozoic mantis shrimps which have narrower, shrimp-like bodies with curved abdomens. While modern mantis shrimps also have a body shape more comparable to lobsters than to shrimp, Perimecturus would have evolved such an appearance separately from them, representing an example of parallel evolution. [11] With the largest specimen (GSE 5897) measuring 15 cm (5.9 in) in total length, P. parki is among the biggest Paleozoic mantis shrimps, though most specimens measure just 1.28–3.37 cm (0.50–1.33 in) in carapace length and 1.50–4.65 cm (0.59–1.83 in) in abdomen length. [4] P. rapax is a smaller species, reaching only 1.22–2.59 cm (0.48–1.02 in) in carapace length and 1.45–2.83 cm (0.57–1.11 in) in abdomen length. [5]

The rostrum is very broad and blunt, with a rounded tip in P. rapax, though it is slightly more pointed in P. parki. The antennae are short and flagellated, each with an elongated and pointed scaphocerite (exopod of the antenna). The compound eyes are stalked and oval in shape. The large, subrectangular carapace covers the entire thorax and has a marked furrow all around its margins, as well as short keels extending from the base of the antennae. The carapace has five prominent ridges (one down the middle of the carapace and two on either side) stretching across its latter half. [6] Though Schram (1978) claims P. parki has only four such ridges and P. rapax has only three, detailed reanalysis by Factor and Feldmann (1985) and Jenner et al. (1998) confirms both species actually have five. [5] [6] [12]

Like all mantis shrimps, Perimecturus had raptorial appendages attached to the thorax. Because the raptorial appendages of this genus are rather short and often retracted to overlap with each other, study of them has been difficult and their features often cannot be discerned. However, some P. parki specimens have shown that most segments of these appendages are approximately equal in length, except the fourth segment from the distal end which is notably shorter. [4]

Little is known about the thorax as it is completely covered by the carapace, though its hind segments at least are not fused with it. The abdomen is made up of six segments, each approximately the same length. A small backward-pointing spike is present at the edges of every abdominal segment. Schram (1978) wrongly stated that P. parki and P. rapax had four and three abdominal ridges respectively, believing they lined up with the carapace ridges. [5] Reanalysis has found that both species have six keels on each abdominal segment, forming six continuous ridges down the abdomen (two median ridges and four lateral ridges). The ridges run parallel to each other in the first four segments and begin to converge with each other at the fifth segment. Two small wart-like bumps are present on each abdominal segment, one located in each space between the median ridges and innermost lateral ridges. [6] [12]

While both species have biramous (two-branched) uropods, spines near the end of the outer margin of the uropodal exopod (outer branch), a subtriangular telson that tapers into a pointed spike at the end and small furcae (spikes forming the tail fork) flanking the base of this spike, other aspects of the tail fan differs greatly between the two Perimecturus species. In P. parki, this spike is less than half the length of the telson base, with delicate bristles around the edges of the telson and uropods. The exopod of the uropod and the hind part of the telson base are smooth. [4] Meanwhile, the telson spike of P. rapax is over half the length of the telson base, no bristles are present on the tail fan, the outer margin of the uropodal exopod is serrated, and the telson is serrated near the furcae. [6]

Classification

Perimecturus gives its name to the family Perimecturidae, of which it is the type genus. This family was established by Ben Peach in 1908, who placed it within the now defunct order Schizopoda, believing that perimecturids were intermediate forms between Lophogastridae and Anaspididae. [3] In 1962, the Perimecturidae family was first recognized as a group of early mantis shrimps by H. K. Brooks, reassigning it to the order Palaeostomatopoda (now delisted as a suborder and named Palaeostomatopodea). [7] With the advent of cladistic analyses, the palaeostomatopods as traditionally construed were first recovered as a paraphyletic grouping by Jenner et al. (1998), a finding later confirmed by Schram (2007). [8] [12] Currently, Palaeostomatopodea is still used in a paraphyletic sense to refer to the evolutionary grade. [13]

Several studies of conducted phylogenetic analyses on fossil mantis shrimps. While Jenner et al. (1998) found the genus to be monophyletic, some other authors have questioned its monophyly. [12] Schram (2007) and Haug et al. (2010) both recovered Perimecturus to be paraphyletic, the former finding P. rapax to be the sister taxon to Bairdops elegans within a monophyletic family Perimecturidae, while the latter suggests the family is a paraphyletic grade in which P. parki is the sister taxon to all other mantis shrimps except Archaeocaris and Bairdops. [8] [14] On the other hand, Smith et al. (2023) supports the idea that both Perimecturus and Perimecturidae are monophyletic. Their results are displayed in the cladogram below: [15]

Stomatopoda
Archaeostomatopods
Palaeostomatopods
Pseudosculdids

Paleobiology

Possessing raptorial appendages on its thorax for grabbing prey like all other mantis shrimps, Perimecturus is believed to have occupied a low trophic level as an active carnivore in its habitat. [6] It was proposed by Jenner et al. (1998) that palaeostomatopods like Perimecturus were scavengers that handled mostly dead prey since their raptorial appendages are small compared to those of extant mantis shrimps in the order Unipeltata, instead being closer in shape to the third to fifth pairs of thoracic appendages used by unipeltatans to manipulate food after it is caught. [12] In addition, the appendages of palaeostomatopods lack the click-joint mechanism seen in unipeltatans, which is formed by a specialized joint and muscles and allows them to extend quickly and catch prey. [16] Contrary to this, Haug and Haug (2021) suggest that at least some early mantis shrimps (namely Tyrannosculda , Tyrannophontes and Gorgonophontes ) had posterior thoracic appendages (i.e. the walking appendages) which were incapable of the wide stance needed to attack prey from the seabed. Therefore, they proposed that these early forms were benthopelagic predators that grabbed prey from above while swimming just over the seabed, unlike modern mantis shrimps which are bottom-dwelling. However, because the walking appendages of Perimecturus are not well-preserved, it remains unclear whether this genus would have also had such a lifestyle. [11]

Paleoenvironment

Based on the fact that both species are known exclusively from marine deposits, Perimecturus is believed to have been a fully marine animal with no adaptation towards lower salinity (unlike the related Bairdops which could tolerate a wider range of salinities). Remains of P. parki have only been collected from the Glencartholm Volcanic Beds, a site in Scotland that dates back to the Viséan stage of the Early Carboniferous period (around 345 million years ago), making it the older of the two known Perimecturus species. The Beds were deposited in a nearshore to fully marine environment, and P. parki is a commonly found crustacean in this locality, as are Bairdops, Anthracocaris , Belotelson and Anthracophausia . The most abundant crustaceans found at this site are Crangopsis , Pseudotealliocaris and Sairocaris , being known from even more specimens than the aforementioned genera. [17] Aside from crustaceans, fossils of other organisms are also known from the Beds. Fish are some of the most numerous, with over 200 specimens representing more than 30 species found in the area. Remains of plants, bivalves, xiphosurans and scorpions are also known from the deposits. [18]

All known specimens of P. rapax originate from the Bear Gulch Limestone in Montana, which was deposited around 324 million years ago during the Serpukhovian stage of the Carboniferous period. This site is believed to have been a marine bay oriented from northwest to southeast, and would have been located 10 to 12 degrees north of the equator. The bay was positioned at the boundary between a tropical belt in the south and an arid belt in the north, and had a monsoonal climate like that of the modern African Sahel with distinct wet summer and dry winter seasons. A very high fossil diversity is recorded from the Bear Gulch Limestone, with some of the most famous fossils being the many types of chondrichthyan fish (including petalodonts, holocephalans, symmoriiforms, Squatinactis and Thrinacodus ) that make up close to 60% of the known fish species diversity. [19] Other types of fish like actinopterygians, coelacanths and acanthodians are also known from this site, with the coelacanths being the most abundant. [20] Invertebrates represented in the area include various crustaceans, polychaete worms, cephalopods and bivalves. [5] [21] [22] [23]

Related Research Articles

<span class="mw-page-title-main">Mantis shrimp</span> Order of crustaceans

Mantis shrimp are carnivorous marine crustaceans of the order Stomatopoda. Stomatopods branched off from other members of the class Malacostraca around 340 million years ago. Mantis shrimp typically grow to around 10 cm (3.9 in) in length, while a few can reach up to 38 cm (15 in). A mantis shrimp's carapace covers only the rear part of the head and the first four segments of the thorax. Varieties range in colour from shades of brown to vivid colours, with more than 520 species of mantis shrimp known. They are among the most important predators in many shallow, tropical and subtropical marine habitats. However, despite being common, they are poorly understood, as many species spend most of their lives sheltering in burrows and holes.

<span class="mw-page-title-main">Malacostraca</span> Largest class of crustaceans

Malacostraca is the second largest of the six classes of pancrustaceans just behind hexapods, containing about 40,000 living species, divided among 16 orders. Its members, the malacostracans, display a great diversity of body forms and include crabs, lobsters, crayfish, shrimp, krill, prawns, woodlice, amphipods, mantis shrimp, tongue-eating lice and many other less familiar animals. They are abundant in all marine environments and have colonised freshwater and terrestrial habitats. They are segmented animals, united by a common body plan comprising 20 body segments, and divided into a head, thorax, and abdomen.

<span class="mw-page-title-main">Bear Gulch Limestone</span>

The Bear Gulch Limestone is a limestone-rich geological lens in central Montana, renowned for the quality of its late Mississippian-aged fossils. It is exposed over a number of outcrops northeast of the Big Snowy Mountains, and is often considered a component of the more widespread Heath Formation. The Bear Gulch Limestone reconstructs a diverse, though isolated, marine ecosystem which developed near the end of the Serpukhovian age. It is a lagerstätte, a particular type of rock unit with exceptional fossil preservation of both articulated skeletons and soft tissues. Bear Gulch fossils include a variety of fish, invertebrates, and algae occupying a number of different habitats within a preserved shallow bay.

<span class="mw-page-title-main">Thylacocephala</span> Extinct group of arthropods

The Thylacocephala are group of extinct probable mandibulate arthropods, that have been considered by some researchers as having possible crustacean affinities. As a class they have a short research history, having been erected in the early 1980s.

<i>Falcatus</i> Extinct genus of cartilaginous fishes

Falcatus is an extinct genus of falcatid chondrichthyan which lived during the early Carboniferous Period in Bear Gulch bay in what is now Montana.

<i>Tealliocaris</i> Extinct genus of crustaceans

Tealliocaris is an extinct genus of pygocephalomorphans that lived from the Late Devonian to Early Carboniferous periods in Europe and North America. The genus was established in 1908 by Ben Peach, and many species have been assigned to it since, though some are now invalid or have been reassigned to other genera. Although it can be inferred that T. loudonensis was originally intended as the type species of Tealliocaris, this species is now deemed a junior synonym of T. woodwardi, first described as a species of Anthrapalaemon in 1877, and thus T. woodwardi is currently deemed the type species of the genus.

Aeschronectida is an extinct order of mantis shrimp-like crustaceans which lived in the Mississippian subperiod in what is now Montana. They exclusively lived in the Carboniferous, or the age of amphibians. They have been found mostly in the U.S. and in the British Isles, in 1979 species were found in the Madera Formation in New Mexico. Aeschronectida was first identified appearing in Continental Europe in around 2014. While sharing similar characteristics to Stomatopoda, they lack certain physical characteristics of that taxon. The first species of Aeschronectida is accredited to Frederick R. Schram. They diverge substantially from typical hoplocaridan morphology by having more unmodified thoracopods. It's theorized that these thoracopods evolved to become more specialized, making them potential ancestors to Stomatopoda.

<span class="mw-page-title-main">Cyclida</span> Extinct order of crustaceans

Cyclida is an extinct order of crab-like fossil arthropods that lived from the Carboniferous to the Jurassic and possibly Cretaceous. Their classification is uncertain, but they are generally interpreted as crustaceans, likely belonging to the superclass Multicrustacea.

<span class="mw-page-title-main">Crustacean</span> Subphylum of arthropods

Crustaceans are a group of arthropods that are a part of the subphylum Crustacea, a large, diverse group of mainly aquatic arthropods including decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, opossum shrimps, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. The three classes Cephalocarida, Branchiopoda and Remipedia are more closely related to the hexapods than they are to any of the other crustaceans.

<i>Anderella</i> Extinct genus of chelicerate

Anderella is a genus of synziphosurine, a paraphyletic group of fossil chelicerate arthropods. Anderella was regarded as part of the clade Prosomapoda. Fossils of the single and type species, A. parva, have been discovered in deposits of the Carboniferous period in Montana, in the United States. Anderella is the first and so far the only Carboniferous synziphosurine being described, making it the youngest member of synziphosurines. Anderella is also one of the few synziphosurine genera with fossil showing evidence of appendages, but the details are obscure due to their poor preservation.

<i>Tyrannosculda</i> Extinct genus of mantis shrimp

Tyrannosculda is an extinct genus of mantis shrimp which lived during the Late Jurassic in southern Germany. It was named in 2021, with T. laurae as the type and only species. Several fossil specimens are known, representing various growth stages.

<i>Tyrannophontes</i> Extinct genus of mantis shrimp

Tyrannophontes is an extinct genus of mantis shrimp that lived during the late Carboniferous period in what is now the Mazon Creek fossil beds of Illinois. It is the only genus in the family Tyrannophontidae. The type species, T. theridion, was described in 1969 by Frederick Schram. A second, much larger species, T. gigantion, was also named by Schram in 2007. Two other species were formerly assigned to the genus, but have since been reclassified.

<i>Daidal</i> Extinct genus of mantis shrimp

Daidal is an extinct genus of mantis shrimp that lived during the Carboniferous period. It is the only genus in the family Daidalidae. Three species are currently placed within the genus. Fossils of the type species, D. acanthocercus, have been found in the Bear Gulch Limestone of Montana. A second species, D. pattoni, is known from the Lower Limestone Formation of Scotland, and the third species, D. schoellmanni, was discovered in Westphalia, Germany. The genus has been proposed to be polyphyletic, with D. pattoni possibly being an earlier diverging lineage, though more specimens and research are needed to confirm this.

<i>Sculda</i> Extinct genus of mantis shrimp

Sculda is an extinct genus of mantis shrimp known from the late Jurassic to late Cretaceous of Germany and Lebanon. Although several species have been assigned to it, some are now deemed dubious or moved to different genera. It was a moderate-sized crustacean, measuring no more than 50 mm (2.0 in) long. Sculda would have lived in a marine environment and been a predatory animal, likely smashing its prey with the widened segment of its raptorial appendages before cutting it with the sharp appendage tips.

<i>Gorgonophontes</i> Fossil genus of mantis shrimp

Gorgonophontes is an extinct genus of mantis shrimp that lived during the late Carboniferous period in what is now the United States and Belgium. It contains two named species. The type species, G. peleron, was described in 1984 by Frederick Schram based on 100 specimens found in Nebraska and Iowa. A second species, G. fraiponti, was first named from multiple specimens found near Liège in 1922 and later reassigned to the genus.

Chabardella is an extinct genus of mantis shrimp which lived during the Late Carboniferous in France. It was named in 2009, with C. spinosa as the type and only species.

<i>Bairdops</i> Fossil genus of mantis shrimp

Bairdops is an extinct genus of mantis shrimp that lived during the Early Carboniferous period in what is now Scotland and the United States. Two named species are currently assigned to it. The type species, B. elegans, has been collected from several Dinantian-aged localities in Scotland, and was first described in 1908 by British geologist Ben Peach as a species of Perimecturus. The generic name was coined decades later in 1979 by American paleontologist Frederick Schram, and honors William Baird. A later species, B. beargulchensis, was named in 1978 after the Serpukhovian-aged Bear Gulch Limestone of Montana where it was discovered. The two species were originally deemed close relatives based on their physical similarities, but several cladistic analyses published since 1998 have suggested the genus may be polyphyletic.

<i>Archaeocaris</i> Extinct genus of mantis shrimp

Archaeocaris is an extinct genus of mantis shrimp that lived in North America during the Early Carboniferous period. Though it was placed as a member of the family Perimecturidae until 2008, it is currently deemed the only genus in the family Archaeocarididae, and contains two species. The type species, A. vermiformis, was described by Fielding Bradford Meek in 1872 from specimens collected at the base of the Waverly Group in Kentucky. A second species, A. graffhami, was named by Harold Kelly Brooks in 1962 based on a fossil found in the Caney Shale of Oklahoma, with additional remains later found in the Pilot Shale of Nevada.

Nodosculda is an extinct genus of mantis shrimp that lived in North America during the late Albian stage of the Early Cretaceous period, between 105 and 100 million years ago. The only species is Nodosculda fisherorum, known from several specimens uncovered in the Paw Paw Formation of Texas.

Ursquilla is an extinct genus of mantis shrimp that lived in Israel and Jordan during the Campanian stage of the Late Cretaceous period. It contains a single species, Ursquilla yehoachi.

References

  1. ↑ Peach, Ben N. (1882). "IV.—On some new Crustaceans from the Lower Carboniferous Rocks of Eskdale and Liddesdale". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 30 (1): 73–91. doi:10.1017/S0080456800028982. ISSN   2053-5945. S2CID   129353541.
  2. ↑ Peach, B. N.; Horne, J. (1903). "The Canonbie Coalfield: its Geological Structure and Relations to the Carboniferous Rocks of the North of England and Central Scotland". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 40 (4): 835–878. doi:10.1017/S0080456800034815. ISSN   2053-5945. S2CID   128763435.
  3. 1 2 3 4 5 Peach, B. N. (1908). Monograph on the higher Crustacea of the Carboniferous rocks of Scotland. Glasgow: Printed for H.M. Stationery off., by J. Hedderwick & sons, ltd.
  4. 1 2 3 4 5 6 7 Schram, Frederick R. (1979). British Carboniferous Malacostraca. Chicago: Field Museum of Natural History.
  5. 1 2 3 4 5 Schram, Frederick R.; Horner, John (1978). "Crustacea of the Mississippian Bear Gulch Limestone of Central Montana". Journal of Paleontology. 52 (2): 394–406. ISSN   0022-3360. JSTOR   1303712.
  6. 1 2 3 4 5 6 7 Factor, David F.; Feldmann, Rodney M. (1985). "Systematics and paleoecology of malacostracan arthropods in the Bear Gulch Lilmestone (Namurian) of central Montana". Annals of the Carnegie Museum. 54: 319–356. doi: 10.5962/p.330776 . ISSN   0097-4463.
  7. 1 2 Brooks, Harold Kelly (1962). "The Paleozoic Eumalacostraca of North America". Bulletins of American Paleontology. 44 (202): 163–338.
  8. 1 2 3 Schram, Frederick R. (2007). "Paleozoic Proto-Mantis Shrimp Revisited". Journal of Paleontology. 81 (5): 895–916. Bibcode:2007JPal...81..895S. doi:10.1666/pleo05-075.1. ISSN   0022-3360. JSTOR   4498847. S2CID   85606671.
  9. ↑ van Straelen, Victor (1922). "Quelques eumalacostracés nouveaux du Westphalien inférieur d'Argenteau près Liége". Annales de la Société Géologique de Belgique (in French). 45: 35–40.
  10. ↑ Schöllmann, Lothar (2004). "Archaeostomatopodea (Malacostraca, Hoplocarida) aus dem Namur B (höheres Marsdenium, Karbon) von Hagen-Vorhalle (NRW, Deutschland) und eine Neudefinition einiger Arten und Famile Tyrannophontidaae". Geologie und Paläontologie in Westfalen. 62: 111–141.
  11. 1 2 Haug, Carolin; Haug, Joachim T. (2021-04-16). "A new fossil mantis shrimp and the convergent evolution of a lobster-like morphotype". PeerJ. 9: e11124. doi: 10.7717/peerj.11124 . ISSN   2167-8359. PMC   8054755 . PMID   33959413.
  12. 1 2 3 4 5 Jenner, Ronald A.; Hof, Cees H.J.; Schram, Frederick R. (1998). "Palaeo- and archaeostomatopods (Hoplocarida, Crustacea) from the Bear Gulch Limestone, Mississippian (Namurian), of central Montana". Contributions to Zoology. 67 (3): 155–185. doi:10.1163/18759866-06703001. ISSN   1383-4517.
  13. ↑ Schram, Frederick (2008). "An adjustment to the higher taxonomy of the fossil Stomatopoda". Crustaceana. 81 (6): 751–754. doi:10.1163/156854008784513429. ISSN   0011-216X. JSTOR   20111439.
  14. ↑ Haug, Joachim T; Haug, Carolin; Maas, Andreas; Kutschera, Verena; Waloszek, Dieter (2010). "Evolution of mantis shrimps (Stomatopoda, Malacostraca) in the light of new Mesozoic fossils". BMC Evolutionary Biology. 10 (1): 290. Bibcode:2010BMCEE..10..290H. doi: 10.1186/1471-2148-10-290 . ISSN   1471-2148. PMC   2955030 . PMID   20858249.
  15. ↑ Smith, C.P.A.; Aubier, P.; Charbonnier, S.; Laville, T.; Olivier, N.; Escarguel, G.; Jenks, J.F.; Bylund, K.G.; Fara, E.; Brayard, A. (2023-03-31). "Closing a major gap in mantis shrimp evolution – first fossils of Stomatopoda from the Triassic". Bulletin of Geosciences: 95–110. doi: 10.3140/bull.geosci.1864 . ISSN   1802-8225.
  16. ↑ Kunze, Janet C. (1981-05-12). "The functional morphology of stomatopod Crustacea". Philosophical Transactions of the Royal Society of London. B, Biological Sciences. 292 (1059): 255–328. Bibcode:1981RSPTB.292..255K. doi:10.1098/rstb.1981.0032. ISSN   0080-4622.
  17. ↑ Briggs, D. E. G.; Clarkson, E. N. K. (1989). "Environmental controls on the taphonomy and distribution of Carboniferous malacostracan crustaceans". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 80 (3–4): 293–301. Bibcode:1989EESTR..80..293B. doi:10.1017/S026359330002873X. ISSN   1755-6910. S2CID   129228575.
  18. ↑ Wood, Margaret (2017). "Glencartholm revisited: describing for the first time Stan Wood's discovery and excavation of Mumbie Quarry, adjacent to the important Palaeozoic fossil site of Glencartholm". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 108 (1): 47–54. Bibcode:2017EESTR.108...47W. doi:10.1017/S175569101800018X. ISSN   1755-6910. S2CID   134352553.
  19. ↑ Lund, Richard; Poplin, Cécile (1999). "Fish diversity of the Bear Gulch Limestone, Namurian, Lower Carboniferous of Montana, USA". Geobios. 32 (2): 285–295. Bibcode:1999Geobi..32..285L. doi:10.1016/S0016-6995(99)80042-4.
  20. ↑ Lund, Richard; Greenfest-Allen, Emily; Grogan, Eileen D. (2012-07-15). "Habitat and diversity of the Bear Gulch fish: Life in a 318 million year old marine Mississippian bay". Palaeogeography, Palaeoclimatology, Palaeoecology. 342–343: 1–16. Bibcode:2012PPP...342....1L. doi:10.1016/j.palaeo.2012.04.016.
  21. ↑ Thomas, Natalie (2004-01-01). The taphonomy of a carboniferous lagerstätte : the invertebrates of the Bear Gulch Limestone Member (thesis thesis). University of Leicester.
  22. ↑ Klug, Christian; Landman, Neil H.; Fuchs, Dirk; Mapes, Royal H.; Pohle, Alexander; Guériau, Pierre; Reguer, Solenn; Hoffmann, René (2019-07-31). "Anatomy and evolution of the first Coleoidea in the Carboniferous". Communications Biology. 2 (1): 280. doi:10.1038/s42003-019-0523-2. ISSN   2399-3642. PMC   6668408 . PMID   31372519.
  23. ↑ McRoberts, Christopher A.; Stanley, George D. (1989). "A unique bivalve–algae life assemblage from the Bear Gulch Limestone (Upper Mississippian) of central Montana". Journal of Paleontology. 63 (5): 578–581. Bibcode:1989JPal...63..578M. doi:10.1017/S0022336000041214. ISSN   0022-3360. S2CID   131030155.
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