Belinurus Bronn, 1839 (Chelicerata, Xiphosura) has priority over Bellinurus Pictet, 1846

Abstract

In the first half of the nineteenth century, a marked shift occurred in our understanding and treatment of the chelicerate fossil record, with the differentiation and recognition of entirely extinct genera for the first time. At the heart of this taxonomic revolution were the Eurypterida (sea scorpions) and Xiphosura (horseshoe crabs), although both groups were in fact considered crustaceans until Lankester's (1881) seminal comparative anatomical study of the extant xiphosuran Limulus Müller, 1785 and modern scorpions. The oldest available eurypterid genus is Eurypterus deKay, 1825; the oldest available fossil arachnid genus name is that of the scorpion Cyclophthalmus Corda, 1835. However, there has been considerable historical confusion over the oldest available fossil xiphosuran genus name, which has been recognized alternately as Belinurus König (with a publication date of either 1820 or 1851) or the synonymous Bellinurus Pictet, 1846. Most recent treatments (e.g., Selden and Siveter, 1987; Anderson and Selden, 1997; Anderson et al., 1997; Lamsdell, 2016, 2021; Bicknell and Pates, 2020) have favored Bellinurus Pictet, 1846 as the available name; however, Haug and Haug (2020) recently argued that Belinurus König, 1820 is valid and has priority, a position then followed by Lamsdell (2020), prompting a reinvestigation of the taxonomic history of the genus. Upon review, it is clear that neither of the previously recognized authorities for Belinurus are accurate and that the two candidate type species for each genus are, in fact, synonyms. Given the convoluted and at times almost illogical history of the competing names, along with the most recent controversy as to which has priority, we present a complete history of the treatment of the genus to resolve the issue.

Full text

Belinurus Bronn, 1839 (Chelicerata, Xiphosura) has priority over
Bellinurus Pictet, 1846
James C. Lamsdell1and Matthew E. Clapham2
1
Department of Geology and Geography, West Virginia University, Morgantown, West Virginia 26501, USA <james.lamsdell@mail.wvu.edu>
2
Department of Earth and Planetary Sciences, UC Santa Cruz, Santa Cruz, California 95064, USA <mclapham@ucsc.edu>
In the first half of the nineteenth century, a marked shift occurred
in our understanding and treatment of the chelicerate fossil
record, with the differentiation and recognition of entirely
extinct genera for the first time. At the heart of this taxonomic
revolution were the Eurypterida (sea scorpions) and Xiphosura
(horseshoe crabs), although both groups were in fact considered
crustaceans until Lankester’s(1881) seminal comparative ana-
tomical study of the extant xiphosuran Limulus Müller, 1785
and modern scorpions. The oldest available eurypterid genus
is Eurypterus deKay, 1825; the oldest available fossil arachnid
genus name is that of the scorpion Cyclophthalmus Corda,
1835. However, there has been considerable historical confusion
over the oldest available fossil xiphosuran genus name, which
has been recognized alternately as Belinurus König (with a pub-
lication date of either 1820 or 1851) or the synonymous Belli-
nurus Pictet, 1846. Most recent treatments (e.g., Selden and
Siveter, 1987; Anderson and Selden, 1997; Anderson et al.,
1997; Lamsdell, 2016,2021; Bicknell and Pates, 2020)have
favored Bellinurus Pictet, 1846 as the available name; however,
Haug and Haug (2020) recently argued that Belinurus König,
1820 is valid and has priority, a position then followed by Lams-
dell (2020), prompting a reinvestigation of the taxonomic history
of the genus. Upon review, it is clear that neither of the previously
recognized authorities for Belinurus are accurate and that the two
candidate type species for each genus are, in fact, synonyms.
Given the convoluted and at times almost illogical history of
the competing names, along with the most recent controversy
as to which has priority, we present a complete history of the
treatment of the genus to resolve the issue.
The issue arises from the partial publication of König’s
(1825)Icones Fossilium Sectiles. This volume (Centuria
Prima) was bound and published, comprising one hundred
figures across eight plates with associated descriptions. Planned
further volumes of Icones Fossilium Sectiles beyond the first
were never published (Urban, 1851); however, it is apparent
that a limited number of lithographs for the plates for the second
and third volumes were produced and shared by König among
interested parties (Woodward, 1830). Among these plates (fig-
ure 230, on plate 18) was a single specimen of a small horseshoe
crab labeled simply Belinurus bellulus.
The informal proliferation of these plates created a scenario
in which the proposed name was widely known among research-
ers but explicitly recognized as unpublished (see Woodward,
1830) and presumably therefore unavailable. Buckland (1837),
in his contribution to the Bridgewater Treatise series, figured
and described a new horseshoe crab species as Limulus trilobi-
toides, explicitly stating that this new species was the same as
König’s unpublished Belinurus bellulus and Martin’s(1809)
Entomolithus Monoculites?Lunatus (a non-binomial name
that has been suppressed by the International Commission on
Zoological Nomenclature along with all other names within Pet-
rificata Derbiensia). Subsequently, Louis Agassiz, in his Ger-
man translation of Buckland’s(1839)Bridgewater Treatise,
followed Buckland’s description of Limulus trilobitoides with
a note that the genus Bellinurus König [sic] is deserving of rec-
ognition as a distinct genus from Limulus (pl. 46; it is worth not-
ing that this is the first recorded occurrence of the alternative
Bellinurus spelling and that it follows an accurate transcription
of the spelling of Belinurus from Buckland’s text on the same
page). In the same year, a summary list of fossil horseshoe
crabs appeared in the Neues Jahrbuch für Mineralogie, Geogno-
sie, Geologie und Petrefaktenkunde, which was edited by Karl
von Leonhard and Heinrich Bronn. The summary itself is not
attributed to either editor; however, the article stated that it
was done at the request of van der Hoeven. In turn, van der Hoe-
ven (1838) stated that he had been in contact with Bronn regard-
ing the status of the fossil xiphosuran species, and so Bronn
(1839) can be reliably ascribed as the author of the article. In
it, Bronn directly assigned the species Entomolithus Monocu-
lites? Lunatus Martin, 1809 (as “Entomolites monoculites”)
and Limulus trilobitoides Buckland to the genus Belinurus
König, with no reference to B. bellulus König. However, the
species reappeared in Morris’s(1843)A Catalogue of British
Fossils, where it was listed as a synonym of Limulus trilobi-
toides Buckland.
Later, in his Traité élémentaire de paléontologie, Pictet
(1846) included the genus Bellinurus König—once again mis-
spelled—to which he assigned the species Limulus trilobitoides,
with the erroneous taxonomic authority of König (notably, there
was again no reference to B. bellulus König), although the
second edition listed B. bellulus König as a senior synonym of
Limulus trilobitoides (Pictet, 1854). Matters were further com-
plicated by Baily (1859a–c), who in a series of papers utilized
the name Bellinurus König, recognizing B. bellulus König as
a synonym of B. trilobitoides (Buckland) and proposing the
genus Steropis to accommodate a variety of species, including
B. trilobitoides (Buckland). Baily (1863) later recognized Ster-
opis as a junior synonym of Belinurus König, 1820 (potentially
the year that the unpublished lithographs were first made avail-
able), stating that Pictet (1846) had made the genus name
Journal of Paleontology, page 1 of 4
Copyright © The Author(s), 2021. Published by Cambridge University Press on behalf of The
Paleontological Society. This is an Open Access article, distributed under the terms of the
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permits unrestricted re-use, distribution, and reproduction in any medium, provided the
original work is properly cited.
0022-3360/21/1937-2337
doi: 10.1017/jpa.2021.53
1
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available. Baily also here treated B. bellulus König, 1820 as a
valid and available species for the first time and considered it
to have priority over B. trilobitoides (Buckland). Henry Wood-
ward, in his extensive works on British fossil chelicerates, was
inconsistent in his treatment of the genus, at first attributing
Baily (1863) as the authority for Belinurus König with B. trilo-
bitoides (Buckland) as the type species (Woodward, 1867)
before later following Baily (1863) in recognizing Bellinurus
König, 1820 with B. bellulus König, 1820 as the valid type spe-
cies (Woodward, 1872,1907).
Moving into the twentieth century, there appeared to be a
consensus that Belinurus König, 1820 was the accurate name
and authority, with B. trilobitoides (Buckland) being a junior
synonym of the type species B. bellulus König (Dix and Pringle,
1929; Eller, 1938; Størmer, 1952). Uncertainties regarding the
taxonomic history of the species continued, however, with
Størmer (1952) erroneously listing Woodward (1866–1878)as
the taxonomic authority for B. bellulus. Størmer (1955) later
compounded this error in the Treatise on Invertebrate Paleon-
tology, which continued to recognize Belinurus König, 1820,
but incorrectly listing Meek and Worthen (1865) as the authority
of the junior synonym Bellinurus.
This situation changed dramatically with the publication of
Morris’s(1980)Catalogue of the type and figured specimens of
fossil Crustacea (excl. Ostracoda), Chelicerata, Myriapoda and
Pycnogonida in the British Museum (Natural History). In it,
Morris reported that the second part of König’s work had
become available only after his death and that the earliest the
genus and species could date to is 1851. This would render Bel-
linurus, made available by Pictet in 1846, as having priority over
Belinurus König, c. 1851. Similarly, B. trilobitoides (Buckland,
1837) would have priority over B. bellulus König, c. 1851. The
next treatment of the group, by Selden and Siveter (1987), fol-
lowed Morris in recognizing Bellinurus Pictet as having priority
but considered B. bellulus König, c. 1851 the valid type species.
Subsequent workers all recognized Bellinurus Pictet, 1846 as
the appropriate taxon name and authority (Schultka, 1994;
Anderson and Selden, 1997; Anderson et al., 1997; Lamsdell,
2016,2021; Bicknell and Pates, 2020). Very few of these treat-
ments considered the issue of the appropriate type species,
although Bicknell and Pates (2020) listed both B. bellulus
(ascribed to Pictet, 1846) and B. trilobitoides (Buckland) as dis-
tinct, valid species. Then Haug and Haug (2020) argued on the
basis of an available scan of Icones Fossilium Sectiles from the
Biodiversity Heritage Library that König’s plates were pub-
lished in 1820 and that Belinurus König had priority over Belli-
nurus Pictet, which Lamsdell (2020) then followed in a
comprehensive revision of xiphosurid taxonomy.
With such a turbulent taxonomic history, the question
remains: what is the correct formulation of the genus name,
and what is the correct taxonomic authority? To determine
this, it must first be ascertained whether the plates for König’s
second volume were published before 1851, or if not, which
publication first made Belinurus available and whether it pre-
or post-dated Pictet’s making Bellinurus available in 1846.
With regard to the publication of König’s plates, all contempor-
ary sources (Woodward, 1830; Urban, 1851) are clear in stating
that the later volumes of Icones Fossilium Sectiles were not pub-
lished and the plates were not publicly available. The fact that
the plates for the second and third volumes were distributed
and appended to bindings of the first volume after König’s
death, as stated by Sherborn (1902) and Lang et al. (1940),
explains the undated binding from the Natural History Museum,
London, that led Haug and Haug (2020) to believe that these
plates were published before 1851. The matter then becomes
when Belinurus and Bellinurus each first became available.
Although Buckland (1837) stated his Limulus trilobitoides is
the same as König’sBelinurus bellulus, he did so with reference
to the unpublished plates and so did not make the genus or spe-
cies available as per Article 12.3 of the International Code of
Zoological Nomenclature (International Commission on Zoo-
logical Nomenclature, 1999). Similarly, Agassiz’s note in his
translation of Buckland (1839) was the first reference to Belli-
nurus in the literature but did not include an assignation of an
available species name and so did not make that spelling of
the genus available. However, Bronn’s(1839) summary of fossil
horseshoe crabs explicitly listed Belinurus König as a valid
genus including the valid species Limulus trilobitoides Buck-
land by indication. This satisfies Article 12.2.5 of the Inter-
national Code of Zoological Nomenclature (International
Commission on Zoological Nomenclature, 1999) and is suffi-
cient for the genus name to become available, whereby its taxo-
nomic authority is Bronn (1839) following Article 50.1
(International Commission on Zoological Nomenclature,
1999). Bellinurus was first made available by Pictet, and so
Belinurus Bronn, 1839 is the valid senior synonym of Bellinurus
Pictet, 1846. Pictet, however, made no reference to B. bellulus,
which Morris (1980) stated was first made available by the dis-
tribution of König’s plates in or after 1851. If the plates were dis-
tributed after 1854, B. bellulus would be attributable to Pictet
(1854), who figured a specimen alongside a diagnosis of the
species. Irrespective as to whether the species was made avail-
able in 1851 or 1854, as B. trilobitoides (Buckland, 1839) and
B. bellulus König, c. 1851/Pictet, 1854 are subjective synonyms,
B. trilobitoides has seniority and is the valid name.
To clarify these issues for future paleontologists, we present
a revised systematic paleontology for Belinurus Bronn, 1839,
including all currently valid species assignable to the genus.
Systematic paleontology
Xiphosurida Latreille, 1802
Belinurina von Zittel in von Zittel and Eastman, 1913
Family Belinuridae von Zittel in von Zittel and Eastman, 1913
(= Euproopidae Eller, 1938; = Liomesaspidae Raymond, 1944)
Genus Belinurus Bronn, 1839
(= Bellinurus Pictet, 1846;=Steropis Baily, 1859a)
Type species.—Belinurus trilobitoides (Buckland, 1837)
(lectotype: BNMH 34889; paralectotype: BMNH 46393)
(= Belinurus bellulus König, c. 1851/Pictet, 1854) from the
clay-ironstone of the Coalbrookdale Coal Measures, Telford,
Shropshire, by subsequent designation.
Other species.—Belinurus carwayensis Dix and Pringle, 1929;
Belinurus concinnus Dix and Pringle, 1929;Belinurus
grandaevus Jones and Woodward, 1899;Belinurus
kiltorkensis Baily, 1869;Belinurus morgani Dix and Pringle,
Journal of Paleontology:1–42
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1930;Belinurus pustulosus Dix and Pringle, 1929;Belinurus
silesiacus (Roemer, 1883); Belinurus sustai (Prantl and Pr
̌ibyl,
1956); Belinurus trechmanni Woodward, 1918.
Diagnosis.—Belinurid with axis of first thoracetron tergite
medially inflated; thoracetron ovoid to semicircular in outline;
thoracetron fixed tergopleural spines elongate, needle-like
(after Lamsdell, 2020).
Occurrence.—Carboniferous: Canada, Czech Republic,
Germany, and United Kingdom.
Remarks.—Morris (1980) listed two syntypes for Belinurus
trilobitoides, one figured by Buckland (1837) and the other
not. No lectotype has been subsequently designated, and so
here we select Buckland’sfigured specimen, BMNH 34889,
as the lectotype for the species following Article 74.1 of the
International Code of Zoological Nomenclature (International
Commission on Zoological Nomenclature, 1999). According
to Morris, this is also the specimen figured by König (c.
1851). There is no indication as to the identity of the
specimen figured by Pictet (1854).
Acknowledgments
JCL thanks A. Baumgartner and D. Lamsdell for reading
through an early draft of the manuscript and helping ensure
the convoluted taxonomic history was presented in an under-
standable format. We thank R. Plotnick and C. Haug for their
reviews of the manuscript.
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