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The nomenclature, identity and synonyms of Cancer vocans minor Herbst, 1782 and Gelasimus caerulens Adams, 1847 (Decapoda, Brachyura, Ocypodidae)

In: Crustaceana
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  • 1 Department of Life Science and Research Center for Global Change Biology, National Chung Hsing University, 250, Kuo Kuang Road, Taichung 402, Taiwan
  • | 2 Lee Kong Chian Natural History Museum, National University of Singapore, 2 Conservatory Drive, Singapore 117377, Republic of Singapore
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Abstract

The nomenclature of two ocypodid names, Cancer vocans minor Herbst, 1782 and Gelasimus caerulens Adams, 1847, is discussed, because their identities have remained unresolved. The former is identified with Gelasimus annulipes H. Milne Edwards, 1837, and the latter with Mesuca (Latuca) paradussumieri Bott, 1973. To conserve the current and widespread usage of both junior synonyms, Article 23.9 of the Zoological Code is applied to reverse precedence. Neotypes are also designated for both species.

Abstract

The nomenclature of two ocypodid names, Cancer vocans minor Herbst, 1782 and Gelasimus caerulens Adams, 1847, is discussed, because their identities have remained unresolved. The former is identified with Gelasimus annulipes H. Milne Edwards, 1837, and the latter with Mesuca (Latuca) paradussumieri Bott, 1973. To conserve the current and widespread usage of both junior synonyms, Article 23.9 of the Zoological Code is applied to reverse precedence. Neotypes are also designated for both species.

Introduction

Shih et al. (2016b) revised the classification of the fiddler crabs and showed that the genus Uca Leach, 1814 was paraphyletic, belonging to two subfamilies. All Indo-West Pacific and some Atlantic-East Pacific fiddler crabs were assigned to nine genera in the subfamily Gelasiminae Miers, 1886, with 97 species recognized (Shih et al., 2016, 2018, 2019; Rosenberg, 2019; Shih & Poupin, 2020).

In this note, we deal with the identities of two names that have been missed or remain unresolved in gelasimine taxonomy. The identity of Cancer vocans minor Herbst, 1782, has been uncertain and the name has been associated with different species; it is here shown to be a synonym of Gelasimus annulipes H. Milne Edwards, 1837. Another name, Gelasimus caerulens Adams, 1847, has not been used since the original description; it is almost certainly conspecific with Mesuca (Latuca) paradussumieri Bott, 1973. Cancer vocans minor Herbst, 1782, and Gelasimus caerulens Adams, 1847, however, are rarely used names, and for them to replace the widely used Gelasimus (currently Austruca) annulipes H. Milne Edwards, 1837, and Mesuca (Latuca) (currently Tubuca) paradussumieri Bott, 1973, respectively, does not serve the cause of taxonomic stability. As such, we apply Article 23.9 of the current Zoological Code (ICZN, 1999) to reverse precedence to conserve both junior names. To ensure long-term stability for the names, we also designate neotypes for Cancer vocans minor Herbst, 1782, and Gelasimus caerulens Adams, 1847.

Specimens examined were deposited in the Zoological Collections of the Department of Life Science, National Chung Hsing University, Taichung, Taiwan (NCHUZOOL); the Senckenberg Museum, Frankfurt am Main, Germany (SMF); and the Zoological Reference Collection of the Lee Kong Chian Natural History Museum, National University of Singapore (ZRC). Measurements provided are of carapace width (CW) and carapace length (CL) in millimetres, respectively. Rosenberg (2019) proposed new subgenera for Austruca Bott, 1973, and Tubuca Bott, 1973, but we are not convinced that a subgeneric system is necessary for these taxa at the moment, at least based on the available evidence (Shih & Poupin, 2020). Until stronger morphological and genetic data shows otherwise, we follow the classification for the Ocypodidae proposed by Shih et al. (2016b) for the time being.

Taxonomy

Superfamily Ocypodoidea Rafinesque, 1815 Family Ocypodidae Rafinesque, 1815 Cancer vocans minor Herbst, 1782

Herbst (1782: 81) stated that “Es giebt von dem C. vocans 2 Arten” [= “There are two species of C. vocans”] and he divided Cancer vocans Linnaeus, 1758, into two taxa of subspecific rank: Cancer vocans minor and Cancer vocans major (cf. Herbst, 1782: 81-83).

The identity of Cancer vocans major Herbst, 1782, was resolved following the designation of the type specimen of Gelasimus platydactylus H. Milne Edwards, 1837, as the neotype of Cancer vocans major Herbst, 1782, by Holthuis (1979a: 248-252). This neotype designation was confirmed by the International Commission on Zoological Nomenclature in Opinion 1262 (ICZN, 1983: 200). This fixed the identity of Cancer vocans major, and the name is in current use, in the combination Uca major (Herbst, 1782) (see Shih et al., 2016b: 149).

The identity of Cancer vocans minor Herbst, 1782, however, has not been clarified. Since 1899, only six authors have used the name Cancer vocans minor Herbst, 1782. Stebbing (1917: 16) discussed the tentative placement by H. Milne Edwards (1852: 145) of Cancer vocans minor in the synonymy of C. vocans Linnaeus, 1758 (currently Gelasimus vocans (Linnaeus, 1758); see Shih et al., 2016b: 151). Maccagno (1928: 44) considered C. vocans minor Herbst, 1782, to be a possible synonym of Ocypoda pugilator Bosc, 1801 (currently Leptuca pugilator (Bosc, 1801); see Shih et al., 2016b: 153). Alcock (1900: 353), Boone (1934: 197) and Estampador (1959: 100) listed C. vocans minor Herbst, 1782, as a synonym of Gelasimus annulipes H. Milne Edwards, 1837 (currently Austruca annulipes (H. Milne Edwards, 1837; see Shih et al., 2016b: 153). Most recently, however, Crane (1975: 326) mistakenly attributed authorship of C. vocans minor to Linnaeus (1758) and remarked that its identity was “[i]ndeterminate” and that “[s]pecimen apparently not extant”. None of these authors considered C. vocans minor Herbst, 1782, to be the valid name for any of their putative synonyms, even though this name has priority.

In describing C. vocans minor, Herbst (1782: 81, 82) wrote: “Es giebt von dem C. vocans 2 Arten; die gegenwärtige um die Hälfte kleinere Art ist die Linnéische. Die in der Müllerschen Uebersetzung des Linnéischen Natursystems Tab. 34. fig. 2. 3. gegebene Abbildung hat den Fehler, dass der Schild hinten zu spitz zulaüft [sic!], da er doch fast viereckig ist. Die Rumphische Abbildung ist besserDie Farbe is überall gelbbraun, mit einigen violetschwarzen Schattirungen. Die Abbildung ist nach der Natur.” [There are two species of C. vocans; the smaller species under discussion is the Linnaean species. The figure given in [the] Müller’s (1775) translation of the Linnaean Systema naturae [1775] plate 34, figures 2, 3 [reproduced as fig. 1H, I, herein] is wrong because the angle of the margins of the carapace towards the abdomen is too obtuse, since the carapace is almost square. Rumphius’s [1705, pl. 10, fig. E; reproduced as fig. 1F herein] figure is better … The colour is golden-brown overall, with shades of violet-black. The figure is drawn from nature].

Fig. 1.
Fig. 1.

Illustrations of the syntypes of Cancer vocans minor Herbst, 1782. A, Syntype from Herbst’s collection but now no longer extant, as figured in Herbst (1782, pl. 1, fig. 10); B, C, syntypes figured by Marcgravi (1648: 184, 185); D, E, syntypes figured by Jonstonus (1650, pl. 9, figs. 7, 12); F, syntype figured by Rumphius (1705, pl. 10, fig. E); G, syntype figured by Petiver (1767, pl. 78, fig. 5); H, I, two specimens illustrated by Müller (1775, pl. 34, figs. 2, 3) that Herbst (1782: 81) considered to be have been misidentified as Cancer vocans Linnaeus, 1758. [Images digitized from works no longer in copyright: A, from the Museum für Naturkunde Berlin, Germany; B-I, from the Biodiversity Heritage Library.]

Citation: Crustaceana 94, 2 (2021) ; 10.1163/15685403-bja10077

His last sentence indicates that Herbst had at least one specimen of C. vocans minor when he described it. In addition, Herbst (1782: 81) also referred to the material figured by: (a) Rumphius (1705, pl. 10, fig. E; fig. 1F herein); (b) Marcgravi (1648, fig. on p. 184, fig. on p. 185; figs. 1B, 1C herein; see also discussion below); (c) Jonstonus (1650, pl. 9, figs. 7, 12; fig. 1D, E herein); and (d) Petiver (1767, pl. 78, fig. 5; fig. 1G herein).

Herbst (1782: 81) also referred to “pl. 78” of “Piso brasil.” (i.e., Marcgravi, 1648; see Holthuis, 1991: 7, 8), but no such plate is found in that work. Holthuis (1991: 7-15) discussed that Marcgravi (1648) was actually based on a set of plates that was deposited in Berlin, sometime before World War II. Herbst would have had access to these plates during his time, before they were removed from Berlin during World War II (Holthuis, 1991: 10). These plates were not available for study, but Herbst (1782: 81) could have been referring to either or both of the two species of fiddler crabs figured by Marcgravi (1648, fig. on p. 184, fig. on p. 185; fig. 1B, C herein). These last two figures appear to have been copied by Jonstonus (1650, pl. 9, figs. 7, 12; fig. 1D, E herein; see Holthuis, 1991: 15).

Herbst (1782: 81) obviously included several taxa of fiddler crabs under the name C. vocans minor. The figures cited above have been identified with at least four currently recognized taxa of fiddler crabs: (a) the figures of Marcgravi (1648, fig. on p. 184; fig. 1B herein) and Jonstonus (1650, pl. 9, fig. 12; fig. 1E herein) are identifiable with Gelasimus maracoani Latreille, 1803 (see Holthuis, 1991: 69, 70) (currently Uca maracoani (Latreille, 1803); see Shih et al., 2016b: 149); (b) the figures of Marcgravi (1648, fig. on p. 185; fig. 1C herein) and Jonstonus (1650, pl. 9, fig. 7; fig. 1D herein) are identifiable with Uca thayeri Rathbun, 1900 (see Holthuis, 1991: 69, 70) (currently Leptuca thayeri (Rathbun, 1900); see Shih et al., 2016b: 153); (c) the figure of Rumphius (1705, pl. 10, fig. E; fig. 1F herein) has been identified with C. vocans Linnaeus, 1758 (e.g., Holthuis, 1959: 68, 115, 116) (currently Gelasimus vocans (Linnaeus, 1758); see Shih et al., 2016b: 151), and with Gelasimus platydactylus H. Milne Edwards, 1837 (e.g., Bott, 1973a: 314) (currently Uca major (Herbst, 1782); as discussed under Cancer vocans major above); and (d) the specimen figured by Herbst (1782, pl. 1, fig. 10; fig. 1A herein), which represents a taxon distinct from those listed above. As the specimen figured in Herbst (1782, pl. 1, fig. 10) was the only actual specimen available to Herbst, this would have been the most suitable specimen to select as the type for C. vocans minor. Unfortunately, the type specimen of C. vocans minor figured in Herbst (1782, pl. 1, fig. 10) is not in the Herbst Collection at the Museum für Naturkunde Berlin, Germany (ZMB) (see also Sakai, 1999) and is almost certainly lost. With the crustacean curator of ZMB, Oliver Coleman, the second and third authors have searched for this specimen and we are confident it is no longer extant.

Nonetheless, the specimen figured in Herbst (1782, pl. 1, fig. 10) shows a species that is currently known as Austruca annulipes (H. Milne Edwards, 1837) (see Shih et al., 2016b: 153). This species was first described as Gelasimus annulipes by H. Milne Edwards (1837: 55, pl. 18, figs. 10-13; fig. 2A herein) for material from “mer des Indes” (= Indian Ocean). Material from Java as well as specimens originally identified as “Gelasimus marionis Desmarest, 1823” by H. Milne Edwards (1837: 53, 54) were later referred to a new species, Gelasimus perplexus, by H. Milne Edwards (1852: 150, pl. 4, fig. 18; fig. 2D herein; see Shih & Poupin, 2020). Gelasimus marionis Desmarest, 1823 is currently a synonym of G. vocans (Shih et al., 2016b: 151). The lectotype of Gelasimus annulipes, collected from an unspecified location in “Indian seas” (“mer des Indes”), is extant and deposited in the collection of the Muséum national d’Histoire naturelle, Paris (MNHN-IU-2008-10637, ex B11854; Naderloo et al., 2016).

Fig. 2.
Fig. 2.

Austruca annulipes (H. Milne Edwards, 1837). A, Gelasimus annulipes H. Milne Edwards, 1837 (after H. Milne Edwards, 1837, pl. 18, fig. 10); B, male, CW 14.7 mm, NCHUZOOL 15072, Chalong Bay, Phuket, Thailand; C, B, Gelasimus annulipes H. Milne Edwards, 1837 (after H. Milne Edwards 1852, pl. 4, fig. 15b); D, Gelasimus perplexus H. Milne Edwards, 1852 (after H. Milne Edwards, 1852, pl. 4, fig. 18a); E, F, outer and inner surface of major cheliped (CW 19,2.0 mm, NCHUZOOL 15075, Jeram, Selangor, Malaysia); G, H, male (CW 14.9 mm, NCHUZOOL 15077, Mersing, Johor, Malaysia.

Citation: Crustaceana 94, 2 (2021) ; 10.1163/15685403-bja10077

The figure of C. vocans minor (pl. 1, fig. 10; fig. 1A herein) is inaccurate because the specimen was depicted as being right-handed, but the chela figured must be that of a left cheliped considering the way the dactylus was drawn. Either it is a composite figure based on two specimens, the artist took liberties with how the chela was oriented when drawing it, or it was due to a printer’s error. Disregarding this inaccuracy, the illustrated characters can be associated with species belonging to the Austruca lactea complex, notably in the broad front and subquadrate carapace (cf. fig. 2A, B, G); and the palm of the major chela with the outer surface smooth and the inner surface with a high and serrated oblique ridge (Crane, 1975; fig. 2F herein). In addition, the small predistal pollex tooth and the relatively narrow proximal half of the major dactylus also show that it is A. annulipes (fig. 2F, H; see Shih & Poupin, 2020). The closely related A. perplexa (H. Milne Edwards, 1852), which is a senior synonym of Uca annulipes var. orientalis Nobili, 1901, differs from A. annulipes in possessing a large and wide predistal pollex tooth and a proportionately higher proximal part of the major dactylus (Crane, 1975; Nobili, 1901; Shih & Poupin, 2020). Biogeographically, A. annulipes has a wider distribution in West Pacific and eastern Indian Ocean, than other members of the A. lactea complex (see Crane, 1975; Naderloo et al., 2010; Shih & Poupin, 2020).

Identifying G. annulipes with C. vocans minor would cause the latter to replace the former as the valid name according to the Principle of Priority (Article 23 of the International Code of Zoological Nomenclature, hereafter the Code, ICZN, 1999: 24). The replacement of the name Gelasimus annulipes with Cancer vocans minor, would, however, cause unnecessary nomenclatural destabilization as the former is in current and widespread use for a well-known and widely distributed species of fiddler crab (see references cited below).

Invoking Article 23.9 of the Code (ICZN, 1999: 27-29) allows a reversal of precedence of a junior synonym when the senior synonym has not been used as a valid name after 1899 (Article 23.9.1.1) and the junior synonym “has been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years” (Article 23.9.1.2).

The species-group name Cancer vocans minor has been used in only six post-1899 publications (Alcock, 1900: 353; Stebbing, 1917: 16; Maccagno, 1928: 44; Boone, 1934: 197; Estampador, 1959: 100; Crane, 1975: 326), and none of these publications considered the name to be valid. Since Cancer vocans minor has not been used as valid name since 1899, Article 23.9.1.1 of the Code is fulfilled.

Over the past 50 years, 31 publications by 51 authors who used the name Gelasimus annulipes H. Milne Edwards, 1837 as the valid name for the taxon it denotes, viz., Jones & Morton, 1994: 35, 36; Rosenberg, 2001: 843, 844, 860, 866; 2019: 731, 734; Jaroensutasinee & Jaroensutasinee, 2003: 183; 2004: 533, 534, 536, 539-548; Bunnuang et al., 2005: 35, 41; Beinlich & Von Hagen, 2006: 14, 18-20, 22, 24, 26; Koga, 2007: 5-8; Naiyanetr, 2007: 131; Ng et al., 2008: 241; Barnes, 2010: 252; Nabout et al., 2010: 104; Shih et al., 2010: 1, 3, 6, 13, 16, 17; 2012: 32, 33, 43; 2013a: 187; 2013b: 643-645; 2015a: 181; 2015b: 475, 486, 493; 2016a: 69, 77, 79; 2016b: 152, 153; 2018: 53; 2019: [2], [3], [7], [8], [13], [17], [19]; Masunari, 2012: 1161, 1163, 1164; Shih, 2012: 84; Mengersen & Schmid, 2013: 139; Christy & Wada, 2015: 440, 441; Mandal et al., 2015: 28, 29; Mokhtari et al., 2015: [1], [2], [5], [6], [9], [11], [13], [14], [15]; 2016: 2; Sasaki, 2019: 12424; Pardo et al., 2020: 139, 148, 150, 151; thereby fulfilling Article 23.9.1.2 of the Code.

As both requirements of Article 23.9.1 are met, and in accordance with Article 23.9.2, the name Gelasimus annulipes H. Milne Edwards, 1837 (herein declared a “nomen protectum”), is considered a valid name and takes precedence over the senior subjective synonym Cancer vocans minor Herbst, 1782 (herein declared a “nomen oblitum”). As such, Gelasimus annulipes H. Milne Edwards, 1837, remains the valid name for the taxon under consideration in the combination Austruca annulipes (H. Milne Edwards, 1837).

In addition, to ensure stability in gelasimine nomenclature, the lectotype of Gelasimus annulipes H. Milne Edwards, 1837 (MNHN-IU-2008-10637, ex B11854), is here designated as the neotype for Cancer vocans minor Herbst, 1782. This makes both names objective synonyms.

Gelasimus caerulens Adams, 1847

At the conclusion of the voyage of H. M. S. “Samarang”, a narrative of the voyage was published by the commanding officer (Belcher, 1847). Arthur Adams, who was surgeon-naturalist aboard the “Samarang”, wrote a section within this narrative entitled ‘Notes on the natural history of the islands’ which also appeared as a separate publication (see Low et al., 2020: 282, 283). Low et al. (2020: 286) discussed the authorship of the new names established in this publication and indicated that it should be cited as Adams (1847).

While the “Samarang” was in Borneo, Adams (1847: 437, 438) made the following observations regarding a fiddler crab:

“The muddy banks of the Batang-Lupar, Sarãwak, and many other rivers of Borneo, are covered at low water by numerous handsome species of Gelasimus, among the number of which is an undescribed species which I have named G. caerulens, from the beautiful blue colour of its carapace. I have seen the black mud in many parts assume a quite brilliant blue tinge during the heat of the day, at low water, when these crustaceans come forth to feed.”

The description of “the beautiful blue colour of its carapace” and the allusion to the species aggregating and causing “the black mud in many parts” to “assume a quite brilliant blue tinge during the heat of the day, at low water, when these crustaceans come forth to feed” fulfils the requirement that a new name “must be accompanied by a description of a definition of the taxon that it denotes” (Article 12.1 of the Code, ICZN, 1999: 16), making the name Gelasimus caerulens available. As discussed in Low et al. (2020: 286), the author of Gelasimus caerulens is Adams (1847).

From the description, Gelasimus caerulens Adams, 1847, is probably what is today known as Mesuca (Latuca) paradussumieri Bott, 1973, currently Tubuca paradussumieri (Bott, 1973) (see Shih et al., 2016b: 159). This species was first described by Bott (1973b: 317, 320-322, fig. 10) based on material from “Bengalen” (fig. 3A, B). Bott (1973b: 321) stated that its distribution encompasses the “Küsten Vorderindiens und der großen Sunda-Inseln” and he also examined additional material from “Java, Soetji”, “Penang”, “Sumatra, Deli” and “Sumatra, Balawan”.

Fig. 3.
Fig. 3.

Tubuca paradussumieri (Bott, 1973). A, B, Dorsal and frontal views of holotype male (35.6 × 21.5 mm, SMF 5650, Bengalen, Indien [India]); C, D, colour in life. C, Neotype male (31.8 × 19.8 mm, ZRC 2020.0067, Buntal, Kuching, Sarawak, Malaysia); D, male, Ranong, Thailand (specimen not collected).

Citation: Crustaceana 94, 2 (2021) ; 10.1163/15685403-bja10077

This identification was mentioned in Shih et al. (2016b: 159) and Low et al. (2020: 294, 296) but its taxonomy was not resolved. Shih et al. (2016b) had noted that Gelasimus caerulens Adams, 1847, should be regarded a “nomen oblitum” and Tubuca dussumieri (H. Milne Edwards, 1852), a “nomen protectum”. However, this identification is not correct and the taxon should be Mesuca (Latuca) paradussumieri Bott, 1973, instead (see below).

As far as known, there are no extant types of Gelasimus caerulens. The second and third authors, with Paul Clark, have searched the dry and rehydrated collections in the Natural History Museum, London and have not been able to find specimens (see also Low et al., 2020: 291). If Adams or the crew did collect material of the species, the specimen(s) are no longer extant. In addition, there were no records of G. caerulens from around Sarawak in the decades after 1847. For example, Adams & White (1848-1849) only reported G. porcellanus White, 1847 [now Austruca annulipes (H. Milne Edwards, 1837)] and G. forcipatus Adams & White, 1849 from Borneo; and G. arcuatus De Haan, 1835 [now Tubuca arcuata (De Haan, 1835), but its identification needs further confirmation] was the only fiddler crab species from Borneo reported by Miers (1880).

In Borneo, four species of fiddler crabs may display blue on the carapace in life: Tubuca bellator (White, 1847), T. dussumieri, T. paradussumieri and G. vocans (Crane, 1975; this study). The colouration of the carapace of T. bellator is brown with variable markings of white and/or blue, especially for displaying males (Crane, 1975: 66; Shih et al., 2016, fig. 10H), but it occurs in Borneo only on the island of Labuan, which is further north (Crane, 1975: 66, map 3; Tweedie, 1950: 357, as Uca angustifrons). Similarly, while T. dussumieri does possess smaller areas of blue patches on the carapace of females and juveniles (Crane, 1975: 33; Nagai & Nomura, 1988: 51; Shih, 1994, fig. 42; Ho, 1996, figs. 3-5; Ng et al., 2008, fig. 188), it only occurs around southern Borneo (Crane, 1975: map 18). There is no record of these latter two species from near or adjacent areas of Batang Lupar, Sarawak, the type locality of G. caerulens (see Crane, 1975; pers. obs.). As a result, T. bellator and T. dussumieri are unlikely to be G. caerulens.

In northern Borneo, including Sarawak, a blue colouration can be observed in T. paradussumieri and G. vocans, during their display phase in hot weather and especially in young crabs (this study, see below). Furthermore, the typical habitat of T. paradussumieri are mudflats, specifically those composed of finer sediments (figs. 3D, 4 herein; Jones & Morton, 1994: 18; Kwok & Tang, 2006: 4; Shih et al., 2015: 271), which agrees well with the “muddy banks” and “black mud” described in Adams (1847). In contrast, the typical habitat of G. vocans is always muddy sand, often on coarser sediments (fig. 5A, C, D herein; Crane, 1975: 88; Lim et al., 2005: 112-113; Shih et al., 2015: 222).

In addition, smaller specimens of T. paradussumieri are uniformly brilliant cobalt blue, without spots; and the blue of T. paradussumieri varies from royal blue to turquoise blue (Crane, 1975: 33; fig. 4 herein), which agrees with the specific name “caerulens”, although most large individuals only have brown or dark brown carapaces and pereiopods (fig. 3C, D). The blue of G. vocans (and other species of the G. vocans complex), however, varies from pale blue to bluish-white (Crane, 1975: 87; fig. 5 herein).

Fig. 4.
Fig. 4.

Blue colour in life of Tubuca paradussumieri (Bott, 1973). A-D, Male; E, probably juvenile male; F, female. A, B, F, Bako National Park, Kuching, Sarawak, Malaysia (courtesy of Bernard Dupont); C, Phuket, Thailand; D, Can Gio, Ho Chi Minh City, Vietnam; E, Sungai Rayu, Kuching, Sarawak, Malaysia. None of these specimens collected.

Citation: Crustaceana 94, 2 (2021) ; 10.1163/15685403-bja10077

Fig. 5.
Fig. 5.

Blue colour in life of the Gelasimus vocans complex. A, G. vocans (Linnaeus, 1758), female, Santubong, Kuching, Sarawak, Malaysia; B, G. vocans, male (CW 17.9 mm, NCHUZOOL 15078, Mersing, Johor, Malaysia); C, G. vomeris (McNeill, 1920), males, Australia (courtesy of Patricia Backwell); D, G. jocelynae (Shih, Naruse & Ng, 2010), male and female, Dongsha Island, Taiwan. A, C, D, specimens not collected.

Citation: Crustaceana 94, 2 (2021) ; 10.1163/15685403-bja10077

On the available evidence presented here, G. caerulens and T. paradussumieri are considered synonymous. The holotype of Mesuca (Latuca) paradussumieri, from Bengal, is extant and is deposited in the Senckenberg Museum, Frankfurt am Main collection (SMF 5650; fig. 3A, B; see Bott, 1973b). Identifying Mesuca (Latuca) paradussumieri with G. caerulens would cause the latter to replace the former as the valid name according to the Principle of Priority (Article 23 of the Code, ICZN, 1999: 24). The replacement of the name Mesuca (Latuca) paradussumieri with G. caerulens would, however cause nomenclatural destabilization as the former is in current usage, and is a well-known and widely distributed species (see references cited below).

Invoking Article 23.9 of the Code (ICZN, 1999: 27-29) allows a reversal of precedence of a junior synonym when the senior synonym has not been used as a valid name after 1899 (Article 23.9.1.1) and the junior synonym “has been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years” (Article 23.9.1.2).

The species-group name G. caerulens has only appeared in two post-1899 publications (viz., Shih et al., 2016b: 159; Low et al., 2020: 294, 296). Both these publications considered G. caerulens to be a synonym of T. dussumieri or T. paradussumieri, but did not use the species of Adams (1847) as the valid name. The name G. caerulens has thus not been used as valid name since 1899, thereby fulfilling Article 23.9.1.1 of the Code.

Over the past 50 years, 33 publications by 59 different authors have used Mesuca (Latuca) paradussumieri as a valid name, viz., Jones & Morton, 1994: 9, 11, 15, 18, 20, 22, 25, 30, 32-36; Rosenberg, 2001: 844, 845, 860, 866; 2002: 162; 2019: 732, 735; Jaroensutasinee & Jaroensutasinee, 2003: 177-184; 2004: 533, 534, 536, 540-549; Weis & Weis, 2004: 56; Bunnuang et al., 2005: 35, 38; Beinlich & Von Hagen, 2006: 25; Kwok & Tang, 2006: 1, 4, 5; Takeda, 2006: 526; Koga, 2007: 3-5, 7; Naiyanetr, 2007: 132; Ng et al., 2008: 241; 2017: 127; Barnes, 2010: 252; Nabout et al., 2010: 104; Shih et al., 2010: 1, 3, 5, 6, 10-12, 16, 17; 2012: 32, 33, 43; 2015a: 180, 266, 270-274; 2015b: 475, 493; 2016a: 57, 74-78; 2016b: 159, 274; 2018: 42, 49, 53, 56; 2019: [7]; Masunari, 2012: 1155; Shih, 2012: 84; Mengersen & Schmid, 2013: 140; Christy & Wada, 2015: 457; Mandal et al., 2015: 28, 29; Mokhtari et al., 2015: [1], [2], [4], [6], [7], [11]-[14]; 2016: 1-6, 8, 9; Sasaki, 2019: 12521; Pardo et al., 2020: 142; thereby fulfilling Article 23.9.1.2 of the Code.

As both requirements of Article 23.9.1 are met, and in accordance with Article 23.9.2, the name Mesuca (Latuca) paradussumieri Bott, 1973 (herein declared a “nomen protectum”), is here considered to have precedence over the senior subjective synonym Gelasimus caerulens Adams, 1847 (herein declared a “nomen oblitum”). As such, the valid name for the taxon under consideration remains Tubuca paradussumieri (Bott, 1973).

To ensure taxonomic stability in this species, which is common in the Indo-West Pacific, a male specimen (31.8 × 19.8 mm, ZRC 2020.0067) recently collected from Buntal, Kuching, Sarawak, Malaysia (01°41.511′N 110°22.379′E) is herein selected as the neotype of Gelasimus caerulens Adams, 1847. This location is only about 70-80 km northwest from the original type locality of Batang Lupar and the specimen is morphologically identical to the holotype of Mesuca (Latuca) paradussumieri Bott, 1973.

3

Corresponding author; e-mail: htshih@dragon.nchu.edu.tw

Acknowledgements

Oliver Coleman (Museum für Naturkunde Berlin, Germany) generously hosted two of the authors (MEYL and PKLN) during a research visit in December 2015 and arranged for the image of Cancer vocans minor from Herbst (1782, pl. 1, fig. 10) to be digitized and used herein. Thanks are also due to Paul Clark (Natural History Museum, London) for helping us search for Adams’ types. The other images used in this article were digitized by the Biodiversity Heritage Library (http://www.biodiversitylibrary.org/). We are also grateful to Bernard Dupont and Patricia Backwell for providing pictures used in this study. This study was partially supported by a grant from the Ministry of Science and Technology (MOST 108-2621-B-005-002-MY3), Executive Yuan, Taiwan, to HTS. We acknowledge the help of two anonymous reviewers and of the editor, J. C. von Vaupel Klein, with the manuscript.

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