Adelophthalmus

''Adelophthalmus'' is a genus of eurypterid, an extinct group of aquatic arthropods. Fossils of ''Adelophthalmus'' have been discovered in deposits ranging in age from the Early Devonian to the Early Permian, which makes it the longest lived of all known eurypterid genera, with a total temporal range of over 120 million years. ''Adelopththalmus'' was the final genus of the Eurypterina suborder of eurypterids and consisted the only known genus of swimming eurypterids from the Middle Devonian until its extinction during the Permian, after which the few surviving eurypterids were all walking forms of the suborder Stylonurina.

Fossils of ''Adelophthalmus'' have been described from four continents; North America, Europe, Asia and Australia, which indicates that ''Adelophthalmus'' might have had a nearly cosmopolitan (worldwide) distribution, one of few eurypterid genera to achieve one besides potentially ''Pterygotus''. The territorial expansion of ''Adelophthalmus'' had begun early, with representatives found in both Siberia and Australia during the Devonian, but it first gained its almost cosmopolitan distribution following the amalgamation of the supercontinent Pangaea during the Carboniferous and Permian.

The generic name ''Adelophthalmus'' means "no obvious eye", referencing that the holotype fossil seemingly represented an eyeless eurypterid, with a carapace (head plate) completely lacking any indication of eyes. Though this has caused much subsequent confusion, including the naming of several junior synonyms, the apparent eyelessness of the type specimen is treated by modern researchers as a preservational artifact, and not a feature that any species of ''Adelophthalmus'' would have possessed in life.

''Adelophthalmus'' was a genus of comparatively small eurypterids, with species ranging in size from 4 cm (1.6 in, ''A. douvillei'') to 32 cm (12.6 in, ''A. khakassicus''). As of 2020, ''Adelophthalmus'' is the most taxonomically diverse of all eurypterid genera, containing 33 species considered valid. This large amount of species, many named long ago, have prompted some researchers to designate ''Adelophthalmus'' as a "wastebasket taxon" with poorly known internal relationships and phylogeny. The genus as it is currently seen may form a monophyletic group (a group sharing a common ancestor) but might most appropriately be split into different genera along distinct clades formed within the current confines of the genus.

Description

Adelophthalmid eurypterids such as ''Adelophthalmus'' were small and streamlined nektonic (actively swimming) eurypterids with prominent cuticle sculptures (ornamentation consisting of small, minute, scales across their backs). These scales are perhaps the most distinguishing feature of the group, though similar scales have been reported in other eurypterid groups, most notably the pterygotids, as well.

Though the largest adelophthalmid, ''Adelophthalmus'' was, in comparison to larger apex predatory members of the group (such as ''Jaekelopterus''), a genus of relatively small eurypterids. The largest species of ''Adelophthalmus'' known, ''A. khakassicus'', reached a maximum length of approximately 32 cm (12.6 in). Many species were smaller, the smallest being the Permian ''A. douvillei'' at just 4 cm (1.6 in) in length. The genus as a whole does not appear to have fluctuated much in size over the course of its long evolutionary history, with "large" species occurring in the Devonian (''A. sievertsi'' at 18 cm, 7 in, and ''A. waterstoni'' at 15 cm, 6 in), the Carboniferous (the aforementioned ''A. mazonensis'', ''A. wilsoni'' at 20 cm, 7.9 in, and both ''A. granosus'' and ''A. zadrai'' at 15 cm, 6 in) and during the Permian (''A. luceroensis'' at 18 cm, 7 in). Most of the smaller species are known from the Carboniferous, when ''Adelophthalmus'' was the most abundant, including the "medium-sized" ''A. irinae'' (13 cm, 5.1 in) and ''A. moyseyi'' (12 cm, 4.7 in) and the smaller ''A. mansfieldi'', ''A. pennsylvanicus'' (both at 8 cm, 3.1 in), ''A. approximatus'' (7 cm, 2.8 in) and ''A. dumonti'' (6 cm, 2.4 in).

Like most eurypterids (with some exceptions, such as ''Slimonia'' and ''Rhinocarcinosoma''), the carapace (the segment covering the prosoma, the "head") of ''Adelophthalmus'' was parabolic in shape, with a narrow marginal rim (edge). The carapace was held in place with the aid of a small and hinged triangular "locking" mechanism placed anteriorly. The eyes were reniform (bean-shaped) and the small ocelli were located between, or slightly behind (depending on the species), the larger eyes. The metastoma (a large plate part of the abdomen) of ''Adelophthalmus'' was oval in shape, with the first opisthosomal (the opisthosoma refers to all segments after the carapace, essentially the abdomen) having a reduced length and being tapered laterally. The body of ''Adelophthalmus'' ended with a long and sharp styliform telson (the posteriormost segment, here in the shape of a spike). The feature that primarily distinguishes ''Adelophthalmus'' from other adelophthalmid eurypterids is its elongated body and the spurs present on its abdominal segments.

Table of species

The status of the 35 names (out of which two are synonyms) listed below follow a 2018 survey by German paleontologists Jason A. Dunlop and Denise Jekel and British paleontologist David Penney and size- and temporal ranges follow a 2009 study by American paleontologists James Lamsdell and Simon J. Braddy unless otherwise noted.'

History of research

Earliest discoveries

The first specimen of ''Adelophthalmus'' to be discovered was excavated in 1851 by German paleontologist Hermann Jordan in a railway shaft at Jägersfreude, near Saarbrücken in Germany. This specimen was described three years later in 1854 in the work ''Ueber die Crustaceen der Steinkohlenformation von Saarbrücken'' ("Of the crustaceans of the coal formation of Saarbrücken"), written by Jordan and Hermann von Meyer and featuring descriptions of several other arthropod taxa. The fossil was immediately recognized by Jordan as that of a eurypterid, with both the overall shape and form and the individual parts (particularly the head and the appendages) being very similar to those of '' Eurypterus'' which had been described in the United States in 1825, 29 years earlier. Among the differences noted between the specimens were the smaller size and later age of the Saarbrücken fossil and what Jordan and von Meyer perceived to be a complete lack of eyes.

Since the preserved carapace had no indication of there ever having been any eyes present, Jordan and von Meyer assumed that the animal would have been completely eyeless in life, with the original description of the fossil citing several cases in which eyeless forms occur in arthropod groups otherwise possessing eyes (such as in crustaceans and trilobites). This apparent eyelessness prompted the choice of name'', Adelophthalmus'', meaning "no obvious eye". The species name, ''granosus'', is derived from the latin ''grānōsus'' ("grainy" or "full of grains"), referring to the state of the fossil preservation having given some of the fossils a grainy texture. The type specimen, to this day the only specimen referred to ''A. granosus'', is today held in the arthropod paleontology collections of the Natural History Museum of Berlin under the specimen number MB.A. 890.

Though modern researchers tend to treat the assumed eyelessness as a preservational artifact and not a feature that ''A. granosus'' would have had in life, this issue was not resolved immediately which made the naming of subsequently discovered species confusing and problematic. ''Lepidoderma imhofi'', named in 1855 from Carboniferous-age deposits in Germany, shows definite eyes. The descriptor, Austrian paleontologist August Emanuel von Reuss, noted that ''Lepidoderma'' likely was synonymous with ''Adelophthalmus'', but ignored the rules of taxonomical priority and used his younger name due to it being based on material that he considered to be better preserved.' The name ''Lepidoderma'' derives from the Latin ''lepidus'' ("elegant" or "fine") and Ancient Greek δέρμα (ðerma, "skin").

In 1868, American paleontologists Fielding Bradford Meek and Amos Henry Worthen described ''Anthraconectes mazonensis'', ''Anthraconectes'' being designated a subgenus of ''Eurypterus'', based on fossils recovered in Carboniferous-age deposits at Mazon Creek in Illinois (the first species to be described from North America).' After examining the ''Adelophthalmus'' type specimen in 1934, German paleontologist Paul Guthörl remarked that ''Anthraconectes'' and ''Adelophthalmus'' were so similar that they would have been synonyms had ''Adelophthalmus'' possessed eyes. The name ''Polyzosternites'' was coined by German paleontologist Friedrich Goldenberg (who also named the species ''Polyzosternites raniceps'', today recognized as ''A. raniceps'') in 1873 to replace the name ''Adelophthalmus'' in regards to specimens described after the type specimen in the belief that the type of ''Adelophthalmus'' represented the fossil remains of a cockroach. ''Glyptoscorpius'' was erected to include some fossils from the Carboniferous of Scotland, including the species ''G. perornatus'' (designated as type, the type specimen consisting of only five tergites'), ''G. caledonicus'' and ''G. kidstoni'', by British geologist Ben Peach in 1882.' ''Glyptoscorpius'' would for a long time erroneously be considered to represent the fossil remains of a scorpion and not an eurypterid.'

The second species to be described from North America was ''A. pennsylvanicus'' (as ''Eurypterus pennsylvanicus''), by Meek and Worthen from the coal-measures of Venango County, Pennsylvania in 1877. That same year, American paleontologist James Hall described the species ''A. mansfieldi'' (under the name ''Eurypterus'' (''Dolichopterus'') ''mansfieldi'') based on fossils recovered in Cannelton, Pennsylvania.' In 1888, Hall described the species ''A. approximatus'' (as ''Eurypterus approximatus'') together with American paleontologist John Mason Clarke based on fossils also recovered from Pennsylvania.'

The English geologist Henry Woodward described the species ''Eurypterus wilsoni'' (=''Adelophthalmus wilsoni'') in 1888 based on a fossil recovered by an Edward Wilson of the Bristol Museum, naming the species in his honor. The only known specimen is composed of six body segments and Woodward noted that naming the species may have been slightly premature. He noted that the specimen possessed markings and spikes running alongside the abdomen in a similar way to ''A. mansfieldi'' (then classified as ''Eurypterus mansfieldi'').'

Portuguese paleontologist Pereira de Lima described the species ''Eurypterus douvillei'' (today seen as ''Adelophthalmus douvillei'') in 1890 based on fossils from Bussaco in Portugal.'

Twentieth century

In 1907, Henry Woodward described ''Eurypterus moyseyi'' (today recognized as ''Adelophthalmus moyseyi'') based on fossils recovered from Radstock, Somerset in England. Woodward compared the singular specimen of ''E. moyseyi'' to fossil specimens of ''A. mansfieldi'' from America, finding the spikes along the abdomen very similar, though noted that they were less prominent in ''E. moyseyi''. Woodward described very large fossil specimens, the carapace alone measuring 21 cm (8.3 in) and the seven associated body segments measuring an additional 25 cm (9.8) together.' Despite this, the latest available size estimates for ''A. moyseyi'' put the species at 12 cm (4.7 in) in length.

''A. nebraskensis'' was described as ''Eurypterus'' (''Anthraconectes'') ''nebraskensis'' in 1914 by American geologist Erwin H. Barbour based on fossils recovered in Nebraska, USA. The species was described alongside other fossils from the associated sediments, which helped reinforce the idea as ''Adelophthalmus'' (or ''Anthraconectes'') as a freshwater animal.

The species ''A. dumonti'', Carboniferous in age, was described by Belgian paleontologist Xavier Stainier in 1915 as ''Eurypterus dumonti''. The type specimen, a relatively complete fossil measuring just 3.3 centimetres (1.3 in) in length, was discovered through boring at a new coalfield in Campine, northern Belgium. Though the fossil had been slightly damaged, including the entire counterpart being fragmented, owing to careless usage of hammers and diamond bores during excavation, the fossil could nevertheless be studied in detail and compared to known eurypterid species. As Stainier considered every known Carboniferous eurypterid to be part of the genus ''Eurypterus'' (among them several species today recognized as ''Adelophthalmus'', such as the type species ''A. granosus'', ''A. imhofi'' and ''A. pennsylvanicus''), he classified the new Belgian eurypterid in that genus as well. He did note that the new species was very similar to species such as ''E. pennsylvanicus'' and especially ''E. mansfieldi'' (both seen as species of ''Adelophthalmus'' today). The species name ''dumonti'' honors the prominent Belgian geologist André Dumont.

The American geologist Amadeus William Grabau described the species ''Anthraconectes chinensis'' in 1920, based on fossils discovered in Zhaozezhuang, China.'

Canadian geologist Walter A. Bell described the species ''A. bradorensis'' in 1922 (as a species of ''Anthraconectes'') based on a single fossil recovered in New Campbelton in the Municipality of the County of Victoria, Canada, referring it to the genus due to similarities with the Scottish ''A. kidstoni'' and the American ''A. mansfieldi''.

1924 saw the description of the species ''Anthraconectes sellardsi'' by American paleontologist Carl Owen Dunbar based on two incomplete fossils and few other small fragments from Elmo in Kansas. The first specimen preserves the carapace and the first four tergites of the preabdomen, while the second preserves five preabdominal and three postabdominal tergites; this specimen represents twice the size of the first one.

The species ''A. oklahomensis'' was described by American paleontologist Carl E. Decker in 1938 based on Permian-age fossils in Oklahoma. Since the ''A. oklahomensis'' specimen was virtually identical to specimens of ''A. sellardsi'' of similar age and a similar stratigraphical horizon in Kansas, ''A. oklahomensis'' was designated a junior synonym of ''A. sellardsi'' by American geologist Carl Colton Branson, with the support of Decker, in 1959.

The type specimen of ''A. zadrai'', MB.A. 889, was collected in the Czech Republic in 1930 or 1931 and first mentioned in a manuscript by French Carboniferous worker Pierre Pruvost, who dubbed it "''Eurypterus (Anthraconectes) Zadrai''", but he did not formally describe the specimen or taxon. Pruvost had previous experience with the genus, having described the species ''Anthraconectes cambieri'' in 1930 based on fossils from Charleroi, Belgium. ''A. zadrai'' was first described formally in 1952 as ''Adelophthalmus zadrai'', at a point in time when the type specimen was seemingly lost. The specimen was rediscovered in Berlin under a different species name based on the original collector of the fossil (Dr. Palisa) and without any designation of it representing a type specimen. Pruvost was also honored through the naming of ''A. pruvosti'' (described as ''Lepidoderma pruvosti'' by Norwegian paleontologist Erik N. Kjellesvig-Waering in 1948 based on fossils discovered in Lens, France).'

1933 saw Ukrainian paleontologist Boris Isidorovich Chernyshev describe the species ''A. carbonarius'' based in one single specimen from the Donets in Ukraine. A new expedition in 2012 carried out by Russian paleontologist Evgeniy S. Shpinev and others in the respectively Russian and Ukrainian localities of Kakichev and Lomuvatka brought a number of well-preserved, presumably juvenile, fossils of ''A. carbonarius''. The exact identification of these fossils is not possible, but they are identified as ''A. carbonarius'' since there are no features showing the opposite. Another Belgian species, ''A. corneti'', was described by Pruvost in 1939 based on fossils from Quaregnon.'

All synonymous genera; ''Anthraconectes'', ''Glyptoscorpius'', ''Lepidoderma'' and ''Polyzosternites'', were subsumed into ''Adelophthalmus'' in studies during the middle twentieth century, notably that of Belgian paleontologist Fredrik Herman van Oyen (1956). Though most authors assign all described species to ''Adelophthalmus'', some, such as van Oyen in 1956, have considered ''Anthraconectes'' to potentially represent a distinct genus, citing that scorpions with similar dorsal anatomies can be quite different ventrally and that the same could be true for the Carboniferous ''Adelophthalmus'' where the ventral morphology is not yet known. A genus ''Anthraconectes'' of this nature would be problematic due to its classification depending on the preservational state of any given specimen.

''A. asturica'' was described as ''Lepidoderma asturica'' by Spanish paleontologist Bermudo Meléndez in 1971 based on fossils from d'Ablana in Spain.'

The species ''A. luceroensis'' was described by American paleontologists Barry S. Kues and Kenneth K. Kietzke in 1981 based on 150 fossil specimens recovered from the Madera Formation of New Mexico. The large amount of specimens recovered, representing individuals at various stages of development and ontogeny, allowed detailed studies to be performed on the ontogeny and intraspecific variation within ''Adelophthalmus''.

American paleontologist Roy E. Plotnick referred a species of ''Eurypterus'', ''E. lohesti'' (first described in 1889) to ''Adelophthalmus'' in 1983 (as ''A. lohesti''), but this classification is questionable as the morphology of the ''A. lohesti'' specimen is not consistent with that otherwise known of ''Adelophthalmus.'' The differences include ''A. lohesti'' having larger eyes, a wider carapace and what could possible by a median ridge on the carapace.

Twenty-first century

In 2004, the German paleontologist Markus Poschmann referred the species ''A. sievertsi,'' first described as part of the genus ''Rhenopterus'' by Norwegian paleontologist Leif Størmer in 1969 based on fossil remains from the Devonian Klerf Formation in Germany, to the genus. Poschmann also referred the species ''Rhenopterus waterstoni'' (described earlier in 2004 based on the singular specimen BMNH In 60174 from the Late Devonian of Australia) to ''Adelophthalmus.'' This species had previously not been assigned to the genus despite clear similarities to other species of ''Adelophthalmus'' partly due to there previously not being any solid evidence for the presence of ''Adelophthalmus'' as early as the Devonian.

''A. irinae'' was described in 2006 based on a fossil specimens (including the holotype, a prosoma, "head", with the specimen number PIN no. 5109/4) collected by the Krasnoyarsk Geological Expedition near Sakhapta, a village in the Nazarovsky District of the Krasnoyarsk Region of Russia. The fossils, from the Tournaisian Solomennyi Stan Formation, could confidently be assigned to ''Adelophthalmus'' based on their scalelike ornamentation, the position of their eyes and the shape of the carapace shortly after their excavation. The species is the first species of ''Adelophthalmus'' to be described from Russia and the first ever Carboniferous eurypterid known from the country. It is also one of few Carboniferous eurypterids found within the territory of the former Soviet Union, the only others being ''A. carbonarius'' from Ukraine and '' Unionopterus'' from Kazakhstan.

Shpinev described two new species of ''Adelophthalmus'' in 2012; ''A. kamyshtensis'' and ''A. dubius'' (the name deriving from the Latin ''dubius ='' "doubtful"), both based on fossils originally collected by Russian geologist Yuriy Fedorovich Pogonya-Stefanovich in 1960 in deposits 3 km southeast of the village of Kamyshta (which lent its name to ''A. kamyshtensis'') of the Republic of Khakassia, Russia and now housed at the Borissiak Paleontological Institute. Despite how poorly preserved these fossils are, several features (notably the parabolic carapace and the presence of spikes along the abdomen) place both species within ''Adelophthalmus''.

In 2013, ''A. piussii'' became the first eurypterid to be described from Italy. The specimen (specimen number MFSNgp 31681, housed at the Museo Friulano di Storia Naturale in Udine) was collected in the gravel bank of a small creek near the greater Bombaso creek, north of the village of Pontebba and consists of a carapace and seven opisthosomal segments on a large block of sandstone. The name of the species, ''piussii'', honors the collector of the type specimen, Stefano Piussi.

In 2018, Shpinev and Russian researcher A. N. Filimonov described a new species named ''A. khakassicus'' based in many well-preserved specimens. Found in the Ilemorovskaya Formation of Khakassia (hence the name) in 2014 by Filimonov, it represents the biggest species of the genus. The holotype, PM TGU 168/108, is composed of parts of the metasoma and a complete telson, with several other known paratypes. As ''A. khakassicus'' is known from similar stratigraphic levels to those of ''A. kamyshtensis'' and ''A. dubius'', it has been suggested that these three species could represent synonyms.

In 2020, Lamsdell, Victoria E. McCoy, Opal A. Perron-Felle and Melanie J. Hopkins described a new species of ''Adelophthalmus'' from the Tournaisian stage of (most likely) the Lydiennes Formation, in France. Its only known specimen, GLAHM A23113, is a nearly complete body only lacking the telson and preserved in phosphatic nodules. For this reason, it was called ''Adelophthalmus pyrrhae'', named after Pyrrha of Thessaly, a figure from Greek mythology who, together with her husband Deucalion, threw stones that transformed into babies to repopulate the world. The good preservation of ''A. pyrrhae'' allowed researchers to examine parts of its respiratory system, and after their study it was confirmed that even if they had a mostly aquatic lifestyle, the eurypterids could venture on to land for long periods.

Evolutionary history

Devonian

The adelophthalmids likely first appeared in the waters of the continent Baltica in the Late Silurian, being a part of a rapid diversification of swimming eurypterids (suborder Eurypterina) throughout the Silurian. Baltica would later collide with the continents Avalonia and Laurentia and form the landmass Euramerica, where most of basal adelophthalmid evolution took place in the Early Devonian. The earliest known species of ''Adelophthalmus'' is ''A. sievertsi'' from Early Devonian (Emsian) deposits of the Klerf Formation in Wilwerath (in the Rhineland-Palatinate), Germany, then part of Avalonia within Euramerica. ''A. sievertsi'' lived in near-shore environments, typically a varied and unstable habitat, which indicates that ''Adelophthalmus'' was eurytopic (capable of surviving in a wide range of environments).

Three other species from the Middle Devonian, ''A. khakassicus'', ''A. kamyshtensis'' and ''A. dubius'', are the earliest known species of ''Adelophthalmus'' from outside Europe, fossils of the three having been recovered from Khakassia in Russia. By the Late Devonian, ''Adelophthalmus'' had already become widespread, with the species ''A. waterstoni'' having been recovered from deposits of Frasnian (~382.2 to 372.2 million years old) age in the Gogo Formation of Western Australia, the only eurypterid with the exception of '' Acutiramus'' and ''Pterygotus'' known from the continent.

The only other species of ''Adelophthalmus'' known from the Devonian is the Famennian (latest Devonian) ''A. lohesti'', known from fossil deposits at Pont de Bonne in Liège, Belgium. Alongside a Famennian species of ''Hibbertopterus'', ''H. dewalquei'', ''A. lohesti'' represents the oldest known eurypterid hitherto discovered in Belgium. ''A. lohesti'' is however represented by a single fragmentary specimen whose identification as ''Adelophthalmus'' or even eurypterine at large is questionable, with it possibly representing a stylonurid eurypterid instead. Devonian specimens of ''Adelophthalmus'' have allegedly also been recovered from Siberia, which would mean that the range of the genus included water around all then existing continents.

The eurypterids were one of the groups most heavily affected by the Late Devonian extinction event, following a major decline in diversity during the Early Devonian, eurypterids were rare in marine environments by the Late Devonian. Of the 16 eurypterid families that had been alive at the beginning of the Devonian, only three persisted into the Carboniferous. All of these were non-marine groups. Whilst the suborder Stylonurina was relatively unscathed, adapting new strategies (such as sweep-feeding) to avoid competition, and came to diversify once more in the Carboniferous, the Eurypterina was rendered almost completely extinct, ''Adelophthalmus'' becoming the sole survivor of the entire suborder.

Carboniferous

Following the extinction of all other swimming eurypterids in the Devonian, ''Adelophthalmus'' became the most common of all eurypterids of the late Paleozoic, existing in far greater number than the surviving members of the Stylonurina, both in terms of the number of individuals and the number of species. ''Adelophthalmus'' experienced a rapid diversification through the Carboniferous, with 23 of its 33 species having been described from the Carboniferous alone, and reached its peak diversity in the Late Carboniferous.' This diversification did not lead to the evolution of any new genera–''Adelophthalmus'' remained the only genus of eurypterine eurypterids until the extinction of the group.

Already widespread and represented around all major landmasses in the Late Devonian, the amalgamation of Pangaea into a global supercontinent during the Carboniferous and Permian would allow the able swimmer ''Adelophthalmus'' to gain an almost worldwide distribution, with Carboniferous-age fossils of ''Adelophthalmus'' having been recovered from the United States, Spain, Belgium, Ukraine, China, Germany, the Czech Republic, Russia, England, Wales, Scotland, France and Italy.'

The Early Carboniferous saw the appearance of a few new species, notably ''A. approximatus'', the earliest record of ''Adelophthalmus'' in North America (although this species may have occurred as early as the Famennian stage, the last stage of the Devonian). The genus also spread to modern day Scotland (''A. perornatus'' recovered from fossil deposits of Early Carboniferous age in Glencartholm) and Asia (the Tournaisian-age ''A. irinae'' known from fossil deposits near Krasnoyarsk, Russia). The appearance of ''A. irinae'' is particularly notable as it represents the hitherto only known Carboniferous eurypterid in Russia. ''A. pyrrhae'' discovered in France is also known from this time. The Late Carboniferous would see the appearance of several more species in various places around the world. During the Bashkirian stage (from 323.2 to 315.2 million years ago), two species appeared in Belgium, ''A. cambieri'' from Charleroi and ''A. corneti'' from Quaregnon, and a third species, ''A. zadrai'' has been reported from deposits of Bashkirian age in Moravo-Silesia of the Czech Republic.

The abundance of the bivalve ''Anthracomya'' suggests strong evidence of freshwater deposition in the habitat of ''A. bradorensis'', a Radstockian (Upper Westphalian) species from Canada.

The Moscovian