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One name to rule them all: Belinurus trilobitoides (Buckland, 1837) is senior synonym to fourteen named species

Published online by Cambridge University Press:  25 August 2021

James C. Lamsdell Open the ORCID record for James C. Lamsdell [Opens in a new window]
James C. Lamsdell*
Affiliation:
Department of Geology and Geography, West Virginia University, Morgantown, West Virginia 26501, USA
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Extract

One of the oldest fossil horseshoe crabs figured in the literature is Entomolithus lunatus Martin, 1809, a Carboniferous species included in his Petrificata Derbiensia. While the species has generally been included within the genus Belinurus Bronn, 1839, it was recently used as the type species of the new genus Parabelinurus Lamsdell, 2020. However, recent investigation as to the appropriate authority for Belinurus (see Lamsdell and Clapham, 2021) revealed that all the names in Petrificata Derbiensia were suppressed in Opinion 231 of the International Commission on Zoological Nomenclature (1954) for being consistently nonbinomial under Article 11.4 of the International Code of Zoological Nomenclature (ICZN) (International Commission on Zoological Nomenclature, 1999). Despite the validation of several species names for anthozoans, brachiopods, and cephalopods described in Petrificata Derbiensia in subsequent rulings (International Commission on Zoological Nomenclature, 1956a, b), Belinurus lunatus has not been the subject of any subsequent Commission ruling or opinion, and so its use in Petrificata Derbiensia remains suppressed. The Belinurus lunatus species name was used in several subsequent publications during the 1800s, none of which made the name available under ICZN article 11.5; Parkinson (1811) is also suppressed for being nonbinomial, while Woodward (1830), Buckland (1837), Bronn (1839), and Baily (1859) refer to the species only as a synonym of Belinurus trilobitoides (Buckland, 1837) through citation to the suppressed Pretificata Derbiensia. The first author to make Belinurus lunatus an available name was Baldwin (1905), who used the name in reference to a new figured specimen from Sparth Bottoms, Rochdale, UK, but again as an explicit junior synonym of Belinurus trilobitoides (Buckland, 1837). Therefore, it was not until Eller (1938) treated B. lunatus as a distinct species from B. trilobitoides that B. lunatus became an available name as per ICZN Article 11.6.1 under the authorship of Baldwin (1905) following ICZN Article 50.7.

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Journal of Paleontology , Volume 96 , Issue 1 , January 2022 , pp. 237 - 241
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One of the oldest fossil horseshoe crabs figured in the literature is Entomolithus lunatus Martin, Reference Martin1809, a Carboniferous species included in his Petrificata Derbiensia. While the species has generally been included within the genus Belinurus Bronn, Reference Bronn1839, it was recently used as the type species of the new genus Parabelinurus Lamsdell, Reference Lamsdell2020. However, recent investigation as to the appropriate authority for Belinurus (see Lamsdell and Clapham, Reference Lamsdell and Clapham2021) revealed that all the names in Petrificata Derbiensia were suppressed in Opinion 231 of the International Commission on Zoological Nomenclature (1954) for being consistently nonbinomial under Article 11.4 of the International Code of Zoological Nomenclature (ICZN) (International Commission on Zoological Nomenclature, 1999). Despite the validation of several species names for anthozoans, brachiopods, and cephalopods described in Petrificata Derbiensia in subsequent rulings (International Commission on Zoological Nomenclature, 1956a, b), Belinurus lunatus has not been the subject of any subsequent Commission ruling or opinion, and so its use in Petrificata Derbiensia remains suppressed. The Belinurus lunatus species name was used in several subsequent publications during the 1800s, none of which made the name available under ICZN article 11.5; Parkinson (Reference Parkinson1811) is also suppressed for being nonbinomial, while Woodward (Reference Woodward1830), Buckland (Reference Buckland1837), Bronn (Reference Bronn1839), and Baily (Reference Baily1859) refer to the species only as a synonym of Belinurus trilobitoides (Buckland, Reference Buckland1837) through citation to the suppressed Pretificata Derbiensia. The first author to make Belinurus lunatus an available name was Baldwin (Reference Baldwin1905), who used the name in reference to a new figured specimen from Sparth Bottoms, Rochdale, UK, but again as an explicit junior synonym of Belinurus trilobitoides (Buckland, Reference Buckland1837). Therefore, it was not until Eller (Reference Eller1938) treated B. lunatus as a distinct species from B. trilobitoides that B. lunatus became an available name as per ICZN Article 11.6.1 under the authorship of Baldwin (Reference Baldwin1905) following ICZN Article 50.7.

When Eller (Reference Eller1938) made B. lunatus available, he did so seemingly unwittingly, with no reference to distinguishing characteristics. Belinurus lunatus is furthermore noticeably absent from otherwise comprehensive species lists (e.g., Raymond, Reference Raymond1944; Morris, Reference Morris1980). Prantl and Přibyl (Reference Prantl and Přibyl1955) correctly listed B. lunatus as a synonym of B. bellulus (itself a junior synonym of B. trilobitoides); Filipiak and Krawczyński (Reference Filipiak and Krawczyński1996) deferred to (but did not reiterate) Prantl and Přibyl's (Reference Prantl and Přibyl1955) synonymy, but incorrectly retained B. lunatus under Martin (Reference Martin1809) as the senior species. Belinurus lunatus, therefore, is and always has been a junior synonym of B. trilobitoides.

Lamsdell (Reference Lamsdell2020) and Bicknell and Pates (Reference Bicknell and Pates2020) both erroneously included B. lunatus as a valid species, referring particularly to Filipiak and Krawczyński's (Reference Filipiak and Krawczyński1996) treatment of the species. In coding the species for phylogenetic analysis, Lamsdell (Reference Lamsdell2016, Reference Lamsdell2020) referred extensively to Filipiak and Krawczyński's (Reference Filipiak and Krawczyński1996) material, which incidentally comprises predominantly large individuals. On the basis of the resulting tree topology, Lamsdell proposed the new genus Parabelinurus, with P. lunatus (Martin) as the type species. As P. lunatus and B. trilobitoides are synonyms, Parabelinurus Lamsdell, Reference Lamsdell2020 and Belinurus Bronn, Reference Bronn1839 are objective synonyms. Thirteen other Belinurus species have also been proposed as synonyms with B. trilobitoides, with their constituent specimens representing taphonomic or ontogenetic variants (Fisher, Reference Fisher1975; Anderson, Reference Anderson1996; Haug and Haug, Reference Haug and Haug2020), including the type species of further genera considered valid by Lamsdell (Reference Lamsdell2020), Koenigiella Raymond, Reference Raymond1944 and Macrobelinurus Lamsdell, Reference Lamsdell2020. To clarify this issue for future researchers, a full taxonomic history of Belinurus trilobitoides (Buckland, Reference Buckland1837), including all synonymies, is presented here.

Systematic paleontology

Belinurus Bronn, Reference Bronn1839
(= Bellinurus Pictet, Reference Pictet1846; = Steropis Baily, Reference Baily1859; = Koenigiella Raymond, Reference Raymond1944; = Macrobelinurus Lamsdell, Reference Lamsdell2020; = Parabelinurus Lamsdell, Reference Lamsdell2020)

Type species

Limulus trilobitoides (Buckland, Reference Buckland1837), by subsequent designation (= Belinurus bellulus König, c.Reference König1851/Pictet, Reference Pictet1854; = Steropis arcuatus Baily, Reference Baily1859; = Belinurus reginae Baily, Reference Baily1863; = Belinurus koenigianus Woodward, Reference Woodward1872; = Bellinurus grandaevus Jones and Woodward, Reference Jones and Woodward1899; = Parabelinurus lunatus (Baldwin, Reference Baldwin1905); = Bellinurus baldwini Woodward, Reference Woodward1907; = Bellinurus longicaudatus Woodward, Reference Woodward1907; = Bellinurus trechmanni Woodward, Reference Woodward1918; = Belinurus concinnus Dix and Pringle, Reference Dix and Pringle1929; = Belinurus carwayensis Dix and Pringle, Reference Dix and Pringle1929; = Belinurus truemani Dix and Pringle, Reference Dix and Pringle1929; = Belinurus pustulosus Dix and Pringle, Reference Dix and Pringle1929; = Belinurus morgani Dix and Pringle, Reference Dix and Pringle1930).

Other species

?Belinurus iswariensis (Chernyshev, Reference Chernyshev1928); ?Belinurus kiltorcanensis Baily, Reference Baily1869; Belinurus lacoei (Packard, Reference Packard1885); ?Belinurus metschensis (Chernyshev, Reference Chernyshev1928); Belinurus silesiacus (Roemer, Reference Roemer1883); ?Belinurus stepanovi (Chernyshev, Reference Chernyshev1928); Belinurus sustai (Prantl and Přibyl, Reference Prantl and Přibyl1955).

Diagnosis

Belinurid with ophthalmic spines positioned at posterior of ophthalmic ridges; axis of first thoracetron tergite medially inflated; thoracetron ovoid to semicircular in outline; thoracetron fixed tergopleural spines elongate, needle-like; conical opisthosomal boss present (after Lamsdell, Reference Lamsdell2020).

Remarks

The correct authorship and spelling of Belinurus was recently clarified by Lamsdell and Clapham (Reference Lamsdell and Clapham2021), who demonstrated that Belinurus Bronn, Reference Bronn1839 was the correct spelling and attribution as opposed to Bellinurus Pictet, Reference Pictet1846.

?Belinurus kiltorcanensis is known only from an isolated carapace that exhibits ophthalmic ridges similar to those in Bellinuroopsis and may not be a Belinurus. The Belinurus species described by Chernyshev (Reference Chernyshev1928) need restudy to ascertain their placement within the genus.

Belinurus trilobitoides (Buckland, Reference Buckland1837)
 Figure 1
Reference Martin1809

‘Entomolithus Monoculites? (lunatus)’; Martin, p. 191, pl. 45, fig. 4.

Reference Parkinson1811

‘Monoculites lunatus’; Parkinson, p. 275, pl. 18, fig. 18.

Reference Woodward1830

‘Entomolithus Derbyensis lunatus’; Woodward, p. 8.

Reference Buckland1837

Limulus trilobitoides Buckland, p. 77, pl. 46″, fig. 3.

Reference Bronn1839

‘Belinurus monoculites’; Bronn, p. 489.

Reference Bronn1839

Belinurus trilobitoides; Bronn, p. 489.

Reference Prestwich1840

Limulus trilobitoides; Prestwich, p. 491, pl. 41, fig. 8.

Reference Morris1843

Limulus trilobitoides; Morris, p. 75.

Reference Mantell1850

Limulus trilobitoides; Mantell, p. 156, pl. 68, fig. 15.

c.Reference König1851

Belinurus bellulus König, pl. 28, fig. 230.

Reference Pictet1854

Bellinurus bellulus; Pictet, p. 538, pl. 46, fig. 23.

Reference Baily1859

Steropis arcuatus Baily, p. 90.

Reference Baily1859

Steropis trilobitoides; Baily, p. 91.

Reference Baily1859

‘Steropis monoculus’; Baily, p. 91.

Reference Baily1859

Steropis bellulus; Baily, p. 91.

Reference Baily1863

Belinurus reginae Baily, p. 110, pl. 5, fig. 1A–D.

Reference Baily1863

Belinurus arcuatus; Baily, p. 111, pl. 5, fig. 2A–C.

Reference Woodward1867

Belinurus trilobitoides; Woodward, p. 32.

Reference Woodward1867

Belinurus reginae; Woodward, p. 32, pl. 1, fig. 1.

Reference Woodward1867

Belinurus arcuatus; Woodward, p. 32.

Reference Woodward1872

Limulus trilobitoides; Woodward, p. 439.

Reference Woodward1872

Bellinurus bellulus; Woodward, p. 439.

Reference Woodward1872

Bellinurus reginae; Woodward, p. 439.

Reference Woodward1872

Bellinurus arcuatus; Woodward, p. 439.

Reference Woodward1872

Bellinurus koenigianus Woodward, p. 439, pl. 10, fig. 8.

Reference Woodward1878

Bellinurus bellulus; Woodward, p. 239, pl. 31, fig. 3a–c.

Reference Woodward1878

Bellinurus reginae; Woodward, p. 240, pl. 31, fig. 1a–d.

Reference Woodward1878

Bellinurus arcuatus; Woodward, p. 241, pl. 31, fig. 2a, b.

Reference Woodward1878

Bellinurus koenigianus; Woodward, p. 243, pl. 31, figs. 3c, 4.

Reference Jones and Woodward1899

Bellinurus grandaevus Jones and Woodward, p. 388, pl. 15, figs. 2, 3.

Reference Baldwin1903

Belinurus bellulus; Baldwin, p. 198.

Reference Baldwin1905

Belinurus lunatus Baldwin, p. 136, fig. 2.

Reference Parker1907

Belinurus lunatus; Parker, p. 44.

Reference Woodward1907

Bellinurus baldwini Woodward, p. 540, fig. 1.

Reference Woodward1907

Bellinurus longicaudatus Woodward, p. 451, fig. 2.

Reference Parker1908

Belinurus baldwini; Parker, p. 71.

Reference Parker1908

Belinurus longicaudatus; Parker, p. 72.

Reference Parker1909

Belinurus baldwini; Parker, p. 6.

Reference Parker1909

Belinurus longicaudatus; Parker, p. 6.

Reference Pruvost1911

Belinurus reginae; Pruvost, p. 299, pl. 7, fig. 4, 4a.

Reference Woodward1918

Bellinurus trechmanni Woodward, p. 462, fig. 5.

Reference Dix and Pringle1929

Belinurus concinnus Dix and Pringle, p. 92, fig. 1.

Reference Dix and Pringle1929

Belinurus carwayensis Dix and Pringle, p. 93, fig. 2.

Reference Dix and Pringle1929

Belinurus truemani Dix and Pringle, p. 94, fig. 3.

Reference Dix and Pringle1929

Belinurus pustulosus Dix and Pringle, p. 95, fig. 4.

Reference Dix and Pringle1929

Belinurus bellulus; Dix and Pringle, p. 97, fig. 5.

Reference Dix and Pringle1929

Belinurus arcuatus; Dix and Pringle, p. 98, fig. 6.

Reference Dix and Pringle1929

Belinurus cf. B. arcuatus; Dix and Pringle, p. 99, fig. 7.

Reference Dix and Pringle1929

Belinurus cf. B. koenigianus; Dix and Pringle, p. 100, fig. 8.

Reference Pruvost1930

Belinurus reginae; Pruvost, p. 197, pl. 12, fig. 5.

Reference Pruvost1930

Belinurus lunatus; Pruvost, p. 198, pl. 12, figs. 7, 8.

Reference Pruvost1930

Belinurus koenigi; Pruvost, p. 199, pl. 12, fig. 6.

Reference Dix and Pringle1930

Belinurus morgani Dix and Pringle, p. 137, fig. 1.

Reference Dix and Pringle1930

Belinurus cf. B. truemani; Dix and Pringle, p. 138, fig. 2.

Reference Dix and Pringle1930

Belinurus bellulus; Dix and Pringle, p. 139, fig. 3.

Reference Dix and Pringle1930

Belinurus konigianus; Dix and Pringle, p. 141.

Reference Dix and Pringle1930

Belinurus reginae; Dix and Pringle, p. 141.

Reference Eller1938

Belinurus bellulus; Eller, p. 132, p. 10, figs. 3–10.

Reference Eller1938

Belinurus koenigianus; Eller, p. 132, pl. 9, fig. 3, pl 11., figs. 1, 2, 8.

Reference Eller1938

Belinurus grandaevus; Eller, p. 132, pl.12, figs. 7, 8.

Reference Eller1938

Belinurus pustulosus; Eller, p. 133, pl. 12, fig. 9.

Reference Eller1938

Belinurus reginae; Eller, p. 133, pl. 10, figs. 1, 2.

Reference Eller1938

Belinurus arcuatus; Eller, p. 133, pl. 11, figs. 3–6.

Reference Eller1938

Belinurus lunatus; Eller, p. 133, pl. 14, fig. 3.

Reference Eller1938

Belinurus baldwini; Eller, p. 133, pl. 14, fig. 1.

Reference Eller1938

Belinurus longicaudatus; Eller, p. 133, pl. 14, fig. 2.

Reference Eller1938

Belinurus truemanni; Eller, p. 134, pl. 11, fig. 7.

Reference Eller1938

Belinurus concinnus; Eller, p. 134, pl. 12, fig. 5.

Reference Eller1938

Belinurus carwayensis; Eller, p. 134, pl. 12, fig. 6.

Reference Eller1938

Belinurus morgani; Eller, p. 134, pl. 12, fig. 2.

Reference Eller1938

Belinurus trechmanni; Eller, pl. 12, fig. 1.

Reference Raymond1944

Belinurus concinnus; Raymond, p. 480.

Reference Raymond1944

Belinurus grandaevus; Raymond, p. 480.

Reference Raymond1944

Belinurus bellulus; Raymond, p. 480.

Reference Raymond1944

Belinurus truemani; Raymond, p. 480.

Reference Raymond1944

Belinurus morgani; Raymond, p. 480.

Reference Raymond1944

Belinurus pustulosus; Raymond, p. 480.

Reference Raymond1944

Koenigiella reginae; Raymond, p. 480.

Reference Raymond1944

Koenigiella arcuata; Raymond, p. 480.

Reference Raymond1944

Koenigiella koenigiana; Raymond, p. 480.

Reference van der Heide1951

Belinurus reginae; van der Heide, pl. 7, figs. 2, 5.

Reference Størmer1952

Belinurus reginae; Størmer, p. 635.

Reference Størmer1952

Belinurus bellulus; Størmer, p. 635.

Reference Størmer and Moore1955

Belinurus regina; Størmer, p. 20, fig. 13.1a.

Reference Størmer and Moore1955

Belinurus arcuatus; Størmer, p. 20, fig. 13.1b.

Reference Størmer and Moore1955

Belinurus baldwini; Størmer, p. 20, fig. 13.1c.

Reference Prantl and Přibyl1955

Belinurus bellulus; Prantl and Přibyl, p. 385, pl. 1, figs. 1, 2.

Reference Copeland1957

Belinurus reginae; Copeland, p. 48, pl. 16, figs. 2, 9, 10.

Reference Copeland1957

Belinurus grandaevus; Copeland, p. 48, pl. 16, figs. 1, 3–8.

Reference Bergström1975

Belinurus koenigianus; Bergström, p. 294, pl. 1, fig. 5.

Reference Morris1980

Belinurus bellulus; Morris, p. 31.

Reference Morris1980

Bellinurus baldwini; Morris, p. 31.

Reference Morris1980

Bellinurus koenigianus; Morris, p. 31.

Reference Morris1980

Bellinurus longicaudatus; Morris, p. 31.

Reference Morris1980

Bellinurus trilobitoides; Morris, p. 31.

Reference Fisher1981

Belinurus koenigianus; Fisher, p. 51, fig. 3A.

Reference Fisher, Eldredge and Stanley1984

Belinurus reginae; Fisher, p. 199.

Reference Fisher, Eldredge and Stanley1984

Belinurus bellulus; Fisher, p. 199.

Reference Fisher, Eldredge and Stanley1984

Belinurus koenigianus; Fisher, p. 199.

Reference Selden and Siveter1987

Bellinurus koenigianus; Selden and Siveter, p. 384.

Reference Schultka1994

Bellinurus grandaevus; Schultka, p. 347.

Reference Schultka1994

Bellinurus arcutus; Schultka, p. 347.

Reference Schultka1994

Bellinurus reginae; Schultka, p. 347.

Reference Schultka1994

Bellinurus lunatus; Schultka, p. 347.

Reference Schultka1994

Bellinurus concinnus; Schultka, p. 347.

Reference Schultka1994

Bellinurus longicaudatus; Schultka, p. 347.

Reference Schultka1994

Bellinurus koenigianus; Schultka, p. 347.

Reference Schultka1994

Bellinurus baldwinii; Schultka, p. 347.

Reference Schultka1994

Bellinurus bellulus; Schultka, p. 347.

Reference Schultka1994

Bellinurus carwayensis; Schultka, p. 347.

Reference Schultka1994

Bellinurus morganii; Schultka, p. 347.

Reference Schultka1994

Bellinurus truemanii; Schultka, p. 347.

Reference Schultka1994

Bellinurus trechmannii; Schultka, p. 347.

Reference Filipiak and Krawczyński1996

Bellinurus lunatus; Filipiak and Krawczyński, p. 420, fig. 4C–H.

Reference Anderson and Selden1997

Bellinurus truemani; Anderson and Selden, p. 20.

Reference Anderson and Selden1997

Bellinurus morgani; Anderson and Selden, p. 20.

Reference Anderson and Selden1997

Bellinurus koenigianus; Anderson and Selden, p. 20.

Reference Anderson and Selden1997

Bellinurus trilobitoides; Anderson and Selden, p. 20, fig. 1.

Reference Anderson and Selden1997

Bellinurus arcuatus; Anderson and Selden, p. 20.

Reference Anderson, Dunlop, Horrocks, Winkelmann and Eager1997

Bellinurus trilobitoides; Anderson et al., p. 203, fig. 4a, b.

Reference Lamsdell2016

Bellinurus arcuatus; Lamsdell, p. 182.

Reference Lamsdell2016

Bellinurus bellulus; Lamsdell, p. 182.

Reference Lamsdell2016

Bellinurus lunatus; Lamsdell, p. 182.

Reference Lamsdell2016

Bellinurus reginae; Lamsdell, p. 182.

Reference Lamsdell2016

Bellinurus trilobitoides; Lamsdell, p. 182.

Reference Lamsdell2016

Bellinurus truemanii; Lamsdell, p. 182.

Reference Bicknell and Pates2020

Bellinurus arcuatus; Bicknell and Pates, p. 18, fig. 13C.

Reference Bicknell and Pates2020

Bellinurus baldwini; Bicknell and Pates, p. 18, fig. 13E.

Reference Bicknell and Pates2020

Bellinurus bellulus; Bicknell and Pates, p. 18, fig. 13D.

Reference Bicknell and Pates2020

Bellinurus carwayensis; Bicknell and Pates, p. 18, fig. 13F.

Reference Bicknell and Pates2020

Bellinurus concinnus; Bicknell and Pates, p. 18, fig. 14B.

Reference Bicknell and Pates2020

Bellinurus koenigianus; Bicknell and Pates, p. 18, fig. 14E.

Reference Bicknell and Pates2020

Bellinurus longicaudatus; Bicknell and Pates, p. 18, fig. 15C.

Reference Bicknell and Pates2020

Bellinurus lunatus; Bicknell and Pates, p. 18, fig. 15A, B.

Reference Bicknell and Pates2020

Bellinurus pustulosus; Bicknell and Pates, p. 18, fig. 16D.

Reference Bicknell and Pates2020

Bellinurus reginae; Bicknell and Pates, p. 19, fig. 16C, E.

Reference Bicknell and Pates2020

Bellinurus trechmanni; Bicknell and Pates, p. 19, fig. 17B.

Reference Bicknell and Pates2020

Bellinurus trilobitoides; Bicknell and Pates, p. 19, fig. 17D.

Reference Bicknell and Pates2020

Bellinurus truemani; Bicknell and Pates, p. 19, fig. 17C.

Reference Haug and Haug2020

‘Belinurus sp.’; Haug and Haug, figs. 1I–VI, 3, 4, 5, 8a–f, 10a–c.

Reference Lamsdell2020

Belinurus bellulus; Lamsdell, p. 13.

Reference Lamsdell2020

Belinurus carwayensis; Lamsdell, p. 13, fig. 1G.

Reference Lamsdell2020

Belinurus concinnus; Lamsdell, p. 13.

Reference Lamsdell2020

Belinurus grandaevus; Lamsdell, p. 13.

Reference Lamsdell2020

Belinurus morgani; Lamsdell, p. 13, fig. 1J.

Reference Lamsdell2020

Belinurus pustulosus; Lamsdell, p. 13.

Reference Lamsdell2020

Belinurus trechmanni; Lamsdell, p. 14.

Reference Lamsdell2020

Belinurus trilobitoides; Lamsdell, p. 14.

Reference Lamsdell2020

Koenigiella reginae; Lamsdell, p. 14.

Reference Lamsdell2020

Koenigiella baldwini; Lamsdell, p. 15.

Reference Lamsdell2020

Koenigiella koenigianus; Lamsdell, p. 15.

Reference Lamsdell2020

Koenigiella longicaudatus; Lamsdell, p. 15.

Reference Lamsdell2020

Koenigiella truemani; Lamsdell, p. 15.

Reference Lamsdell2020

Macrobelinurus arcuatus; Lamsdell, p. 15.

Reference Lamsdell2020

Parabelinurus lunatus; Lamsdell, p. 15.

Figure 1. Specimens of Belinurus trilobitoides previously considered to represent distinct species and genera. (1) MM LL.111267a, assigned to Belinurus trilobitoides. (2) NHMUK PI. I. 2754, previously assigned to Belinurus lunatus. (3) GSC 12803, previously assigned to Belinurus reginae. (4) NHMUK PI. In. 18572, holotype of Belinurus baldwini. (5) AM F29886, previously assigned to Belinurus arcuatus. Scale bars = 5 mm. Institutional abbreviations: AM = Australian Museum, Sydney, Australia; GSC = Geological Survey of Canada, Ottawa, Canada; MMUP = Manchester Museum, Manchester, UK; NHMUK = Natural History Museum, London, UK. (1, 2, 4, 5) Images reproduced from Bicknell and Pates (Reference Bicknell and Pates2020) under a CC BY 4.0 license.

Lectotype

Complete individual preserving dorsal exoskeleton, comprising prosomal carapace, thoracetron, and telson (BNMH 34889) from the Carboniferous clay ironstone of the Coalbrookdale Coal Measures, Telford, Shropshire (Buckland, Reference Buckland1837, pl. 46″, fig. 3).

Remarks

Fifteen of the 22 species historically included within Belinurus have been considered synonyms after critical evaluation by a number of researchers (Fisher, Reference Fisher1975; Anderson, Reference Anderson1996), with many regularly co-occurring (e.g., B. reginae and B. grandaevus in Nova Scotia, Canada (Copeland, Reference Copeland1957); B. arcuatus and B. reginae in Leinster, Ireland (Baily, Reference Baily1863); B. trilobitoides, B. lunatus, B. longicaudatus, and B. baldwini in Rochdale, UK (Eller, Reference Eller1938); B. morgani, B. trilobitoides, and B. reginae in Neath, Wales (Dix and Pringle, Reference Dix and Pringle1930)). Haug and Haug (Reference Haug and Haug2020), in reconstructing an ontogenetic series of Belinurus specimens based on material held in the Natural History Museum, London (BMNH), did not refer to the previous species assignments of individual specimens but included numerous specimens referred to B. trilobitoides (BMNH 18357, 18565, 18571, 36188, 13897, 13898, 13958, In41494, 46421) along with the holotypes of B. baldwini (BMNH In18572), B. koenigianus (BMNH In59227), and B. trechmanni (BMNH In18487) as different ontogenetic stages of the same species.

Most of the 14 junior synonyms of B. trilobitoides were diagnosed on the basis of dubious morphological criteria that are now known to be the result of taphonomic or ontogenetic processes, a conclusion supported by recent increases in our understanding of horseshoe crab decay (Babcock and Chang, Reference Babcock and Chang1997) and development (Haug et al., Reference Haug, Van Roy, Leipner, Funch, Rudkin, Schöllman and Haug2012; Haug and Rötzer, Reference Haug and Rötzer2018; Tashman et al., Reference Tashman, Feldmann and Schweitzer2019; Haug and Haug, Reference Haug and Haug2020; Lamsdell, Reference Lamsdell2021). Belinurus arcuatus (the type species of Macrobelinurus Lamsdell, Reference Lamsdell2020), known from an isolated carapace, was diagnosed on the basis of the presence of a “facial suture” that is actually a crease in a taphonomically deformed specimen that otherwise exhibits no clear differences from specimens of B. trilobitoides. Belinurus reginae (type species of Koenigiella Raymond, Reference Raymond1944) was diagnosed solely on the occurrence of long thoracetron epimera, a trait known to vary throughout ontogeny. Belinurus koenigianus was diagnosed on the basis of the lack of carapace spines (which are actually broken away from the specimen rather than absent) and the proportions of the thoracetron, which is laterally compressed. No diagnosis was presented for B. grandaevus, which appears to have been named solely due to its geographic occurrence, and the available material shows no obvious differences from B. trilobitoides. Baldwin (Reference Baldwin1905) explicitly named B. lunatus as a junior synonym of B. trilobitoides and as such provided no diagnostic differences. Belinurus baldwini was defined as a species on the basis of carapace dimensions; however, the type specimen is taphonomically distorted and no other characters separate it from B. trilobitoides. Belinurus longicaudatus was diagnosed solely on its possession of a long telson although telson length is known to vary between individuals and through ontogeny. Belinurus trechmanni is known from a small specimen and diagnosed on the basis of genal spine length and thoracetron shape, both of which change through ontogeny. Finally, Dix and Pringle (Reference Dix and Pringle1929, Reference Dix and Pringle1930) named B. concinnus, B. carwayensis, B. truemanni, and B. morgani on the basis of their possession of different numbers of free segments in the thoracetron, an interpretation of segment articulation now recognized as erroneous (Anderson and Selden, Reference Anderson and Selden1997). Belinurus truemanni was further diagnosed by the possession of curved genal spines (the form of which is due to compression), B. concinnus by a long headshield (when the specimen is tectonically distorted), and B. morgani by granulation on the prosoma (which is also known from B. trilobitoides). The final synonymous species, B. pustulosus, was diagnosed on the basis of a pustulose cuticular ornament, which again does not serve to distinguish the specimens from B. trilobitoides. With none of these species having valid diagnostic traits to separate them from B. trilobitoides, the weight of evidence suggests that all are considered synonyms, in keeping with the conclusions of Fisher (Reference Fisher1975), Anderson (Reference Anderson1996), and Haug and Haug (Reference Haug and Haug2020).

Acknowledgments

I am grateful to L. Babcock and C. Schweitzer for their reviews of the manuscript. I thank R. Bicknell for providing a copy of Prantl and Přibyl (Reference Prantl and Přibyl1955).

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Figure 0

Figure 1. Specimens of Belinurus trilobitoides previously considered to represent distinct species and genera. (1) MM LL.111267a, assigned to Belinurus trilobitoides. (2) NHMUK PI. I. 2754, previously assigned to Belinurus lunatus. (3) GSC 12803, previously assigned to Belinurus reginae. (4) NHMUK PI. In. 18572, holotype of Belinurus baldwini. (5) AM F29886, previously assigned to Belinurus arcuatus. Scale bars = 5 mm. Institutional abbreviations: AM = Australian Museum, Sydney, Australia; GSC = Geological Survey of Canada, Ottawa, Canada; MMUP = Manchester Museum, Manchester, UK; NHMUK = Natural History Museum, London, UK. (1, 2, 4, 5) Images reproduced from Bicknell and Pates (2020) under a CC BY 4.0 license.