Research Article |
Corresponding author: Chirasak Sutcharit ( jirasak4@yahoo.com ) Academic editor: Thierry Backeljau
© 2021 Parin Jirapatrasilp, Jonathan D. Ablett, Somsak Panha, Chirasak Sutcharit.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Jirapatrasilp P, Ablett JD, Panha S, Sutcharit C (2021) Clarification on the name-bearing type designation of several cyclophorid species (Mollusca, Gastropoda) by H. H. Godwin-Austen (1915). ZooKeys 1049: 43-66. https://doi.org/10.3897/zookeys.1049.66842
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The type series boundary and the name-bearing type designation of each cyclophorid taxon originally described by Godwin-Austen are clarified based on an interpretation that complies with the ICZN. Previous statuses of type specimens designated by previous authors are reconsidered. Lectotypes of Spiraculum oakesi Godwin-Austen, 1915, Spiraculum kempi Godwin-Austen, 1915, Pterocyclos aborensis Godwin-Austen, 1915, Pterocyclos miriensis Godwin-Austen, 1915, Pterocyclos brahmakundensis Godwin-Austen, 1915, Spiraculum luyorensis Godwin-Austen, 1915, Spiraculum putaoensis Godwin-Austen, 1915, and Theobaldius oakesi Godwin-Austen, 1915 are here designated to stabilize the existing nomenclature. In addition, the type specimens of Pterocyclos miriensis and Theobaldius oakesi are photographed and figured for the first time.
Cyclophoridae, hypodigm, ICZN, NHM, nomenclature, NZSI, onomatophore
The phylogenetic analyses of the operculated land snail genus Cyclophorus (Caenogastropoda: Cyclophoridae) from Thailand uncovered a high degree of intra- and interspecific morphological variation and a wide distribution of the genus (
The Natural History Museum in London (hereafter the NHM) holds the type specimens of 42 nominal Cyclophorus species (
Of the type specimens housed in the NHM, the cyclophorid taxa originally described in the “Zoological Results of the Abor Expedition” by
In this article, we review the type status of all cyclophorid taxa originally described by
When
Article 72.4.1. The type series of a nominal species-group taxon consists of all the specimens included by the author in the new nominal taxon (whether directly or by bibliographic reference), except any that the author expressly excludes from the type series [Art. 72.4.6], or refers to as distinct variants (e.g. by name, letter or number), or doubtfully attributes to the taxon.
Article 72.4.1.1. For a nominal species or subspecies established before 2000, any evidence, published or unpublished, may be taken into account to determine what specimens constitute the type series.
Article 72.4.6. If an author when establishing a nominal species-group taxon nominates either “syntypes” (by that term, or by use of one of the terms “cotypes” or “types” alone), or “holotype and paratypes” used together (or by use of the term “type” together with “allotype” or “cotypes”), and also lists other specimens, the separate mention of the latter expressly excludes them from the type series.
Article 72.4.7. The mere citation of “Type” or equivalent expression, in a published work other than that in which the nominal species-group taxon is established, or in an unpublished catalogue of a museum, or on a label, is not necessarily evidence that a specimen is or is fixed as any of the kinds of types referred to in this Chapter.
Article 73.2. Syntypes are specimens of a type series that collectively constitute the name-bearing type. They may have been expressly designated as syntypes (see Article 73.2.1 for acceptable terms); for a nominal species-group taxon established before 2000 [Art. 72.3] all the specimens of the type series are automatically syntypes if neither a holotype [Art. 72.1] nor a lectotype [Art. 74] has been fixed. When a nominal species-group taxon has syntypes, all have equal status in nomenclature as components of the name-bearing type.
According to Art. 72.4.1.1, additional evidence found within or outside the original descriptions, either published or unpublished, may be considered when determining which specimens constitute a type series. Yet, if we do so for the type material of
These two possible interpretations of the type series are rooted in the ambiguous usage of the “type” terminology in the 19th century. The term “type” had been given three operational functions by
The ICZN regulates the nomenclatural rules but is not involved in “restricting the freedom of taxonomic thought or actions” (
The problem of defining
Original labels bearing Godwin-Austen’s handwriting of A Cyclophorus aborensis NHMUK 1903.7.1.3051 with the word “Co-Type” B Cyclophorus (Glossostylus) koboensis NHMUK 1903.7.1.3579 with the word “Co-Type”, and C Spiraculum minimum NHMUK 1903.7.1.3145 with the word “Typical” in different handwriting.
Consequently, following Art. 72.4.1. the type series cannot be restricted to only the first lot in the type paragraph, and this action corresponds well to the hypodigm as recognized by
Article 73.1.1. If an author when establishing a new nominal species-group taxon states in the original publication that one specimen, and only one, is the holotype, or “the type”, or uses some equivalent expression, that specimen is the holotype fixed by original designation.
Article 73.1.2. If the nominal species-group taxon is based on a single specimen, either so stated or implied in the original publication, that specimen is the holotype fixed by monotypy (see Recommendation 73F). If the taxon was established before 2000 evidence derived from outside the work itself may be taken into account [Art. 72.4.1.1] to help identify the specimen.
Recommendation 73F. Avoidance of assumption of holotype. Where no holotype or syntype was fixed for a nominal species-group taxon established before 2000, and when it is possible that the nominal species-group taxon was based on more than one specimen, an author should proceed as though syntypes may exist and, where appropriate, should designate a lectotype rather than assume a holotype (see also Article 74.6).
The indication whether the holotype designation of each taxon comply with an “equivalent expression” is a crucial point because if the original author explicitly designated only one onomatophore (= holotype), that holotype designation would comply with the Code and there would be no contradiction in accepting a type (series) as the original hypodigm. Unfortunately,
We identified four ways of type annotation by
The second way of type annotation by
The third way of
Finally, the fourth way of type annotation occurs in the original description of Cyclophorus aborensis. Two specimens were figured in the plate but only one specimen was marked as “Type” in the plate caption (Fig.
Although
The order of taxa below follows that of
Current status of type series, other non-type materials, type locality, and original onomatophores of cyclophorid taxa in
Taxon | Type series | Type locality | Original onomatophores recognized by |
Remarks |
---|---|---|---|---|
1. Cyclophorus aborensis | Holotype NZSI M.6010/1. Paratypes NZSI M.6009/1, NHMUK 1903.7.1.3048, NHMUK 1903.7.1.3051 | Rotung, 2000 ft., near Egar stream | NZSI “No. 6010/1” (1 shell) | Invalid lectotype and paralectotype designation by |
2. Cyclophorus (Glossostylus) bapuensis | Lectotype NHMUK 1903.7.1.3108/1. Paralectotypes NHMUK 1903.7.1.3108/2–3 | Abor Hills, vicinity of Bapu | NHMUK 1903.7.1.3108 (4 shells) | Valid lectotype and paralectotype designation by |
3. Cyclophorus (Glossostylus) sidiensis | Syntypes NZSI M.6002, NZSI M.6001, NHMUK 1903.7.1.3095 | On Sidi River, Abor Hills; Rotung; Tsanpu Valley | NZSI “No. 6002” | – |
4. Cyclophorus (Glossostylus) koboensis | Holotype NZSI M.6015/1. Paratypes NZSI M.6019–20, NHMUK 1903.7.1.3045, NHMUK 1903.7.1.3117, NHMUK 1903.7.1.3579 | Abor Hills, Kobo, on right bank of Tsanspu or Brahmaputra River | NZSI “No. 6015” (1 shell) | Invalid lectotype and paralectotype designation by |
5. Spiraculum oakesi | Lectotype (design. nov.) NHMUK 1903.7.1.3081/1. Paralectotypes NHMUK 1903.7.1.3081/2–5, NZSI | Abor Hills | NHMUK 1903.7.1.3081 (5 shells) | – |
6. Spiraculum kempi | Lectotype (design. nov.) NHMUK 1903.7.1.3105/1. Paralectotypes NHMUK 1903.7.1.3105/2, NHMUK 1903.7.1.3047, NZSI | Abor Hills | NHMUK 1903.7.1.3105 (2 shells) | – |
7. Spiraculum planum | Syntypes NZSI M.5992, NZSI M.5992a | Upper Rotung, Abor Hills; Yembung | NZSI “No. 5992” | – |
8. Pterocyclos aborensis | Lectotype (design. nov.) NHMUK 1903.7.1.3104/1. Paralectotypes NHMUK 1903.7.1.3104/2–3, NHMUK 1903.7.1.3046, NHMUK 1903.7.1.3050 | Abor Hills | NHMUK 1903.7.1.3104 (3 shells) | – |
9. Pterocyclos miriensis | Lectotype (design. nov.) NHMUK 1903.7.1.3580/1 (Fig. |
Miri Hills | NHMUK 1903.7.1.3580 (4 shells) | – |
Paralectotypes NHMUK 1903.7.1.3580/2–4 | ||||
10. Pterocyclos spiramentum | Holotype NHMUK 1903.7.1.3082 | Abor Hills | NHMUK 1903.7.1.3082 (1 shell) | – |
11. Pterocyclos brahmakundensis | Lectotype (design. nov.) NHMUK 1903.7.1.713/1. Paralectotypes NHMUK 1903.7.1.713/2–3 | Brahmakund, Eastern Assam | NHMUK 1903.7.1.713 (3 shells) | – |
12. Spiraculum luyorensis | Lectotype (design. nov.) NHMUK 1903.7.1.3530/1 | Luyor, Abor Hills | NHMUK 1903.7.1.3530 (1 shell) | – |
13. Spiraculum putaoensis | Lectotype (design. nov.) NHMUK 1903.7.1.3598/1. Paralectotypes NHMUK 1903.7.1.3598/2–3 | Putao, Upper Burma | NHMUK 1903.7.1.3598 (3 shells) | – |
14. Spiraculum minimum | Lectotype NHMUK 1903.7.1.3145/1. Paralectotypes NHMUK 1903.7.1.3145/2–3, NHMUK 1903.7.1.3147, NZSI M.6142, NZSI M.6143 | Sibbum, Abor Hills | NZSI “No. 6142–43” (2 shells) | Valid lectotype and paralectotype designation by |
15. Theobaldius oakesi | Lectotype (design. nov.) NHMUK 1903.7.1.3083/1 (Fig. |
Tsanspu Valley, Abor Hills | NHMUK 1903.7.1.3083 (2 shells) | – |
Paralectotype NHMUK 1903.7.1.3083/2 (Fig. |
Cyclophorus aborensis
Godwin-Austen, 1915: 494, pl. 38, fig. 1, 1a–c.
Holotype
NZSI M.6010/1. Paratypes NZSI M.6009/1 (1 shell) from Rotung, 2000 ft., near Egar stream; NHMUK 1903.7.1.3048 (2 shells;
NHMUK 1903.7.1.3049 (2 shells) from Rami Dambang, Abor.
Specimen “No. 6010/1 in Ind. Mus.” (NZSI M.6010/1) is deemed the holotype fixed by original designation as explained above. All specimens in the remaining lots are paratypes, except for lot “No. 3049 B.M.”, which
Record in the NHM Register of Godwin-Austen Collection in Godwin-Austen’s handwriting of A No. 3051 Cyclophorus aborensis B No. 3108 Cyclophorus (Glossostylus) bapuensis C No. 3095 Cyclophorus (Glossostylus) sidiensis D No. 3579 Cyclophorus (Glossostylus) koboensis E No. 3105 Spiraculum kempi F No. 3047 Spiraculum kempi, and G No. 3530 Spiraculum luyorensis.
Cyclophorus (Glossostylus) bapuensis
Godwin-Austen, 1915: 494, 495, pl. 38, fig. 2, 2a–c.
Lectotype
NHMUK 1903.7.1.3108/1 (
Only one specimen lot, “No. 3108 Brit. Mus.”, was mentioned in the “Type” paragraph of the original description and was labelled as “Type” in Godwin-Austen’s handwriting (
Cyclophorus (Glossostylus) sidiensis Godwin-Austen, 1915: 495, pl. 38, fig. 3, 3a–c.
Syntypes NZSI M.6002 from Sidi River, Abor Hills; NZSI M.6001 from Rotung; NHMUK 1903.7.1.3095 (3 shells) from Tsanpu Valley.
The type series of this species comprises three specimen lots. Without any explicit holotype designation or equivalent expression, all specimens in these three lots are syntypes. However, only specimen lot “No. 6002 Ind. Mus.” was mentioned in the “Type” paragraph, and one specimen from this lot was figured with the annotation “Type” in the plate caption. This specimen lot is, therefore, the onomatophore as originally intended by
Cyclophorus (Glossostylus) koboensis
Godwin-Austen, 1915: 495, 496, pl. 38, fig. 4, 4a–d.
Holotype
NZSI M.6015/1 (
NHMUK 1903.7.1.3581 (1 shell) from between N. 28°15'–29°15'L. 94°50'–95°10'.
Specimen “No. 6015 Ind. Mus.” (NZSI M.6015/1) is deemed the holotype fixed by original designation as explained above. All specimens in the remaining lots are paratypes, except for lot “No. 3581 B.M.” of which
Spiraculum oakesi Godwin-Austen, 1915: 496, pl. 39, fig. 3, 3a.
Pearsonia oakesi
–
Lectotype
(design. nov.) NHMUK 1903.7.1.3081/1 (
The type series of this species comprises lot “No. 3081 Brit. Mus.” and two uncatalogued specimens in the Indian Museum. Without any explicit holotype designation or equivalent expression, all specimens are syntypes. However, only specimen lot “No. 3081 Brit. Mus.” was mentioned in the “Type” paragraph, and one specimen from this lot was figured with the annotation “Type” in the plate caption. The author explicitly indicated that five specimens were examined, and type lot NHMUK 1903.7.1.3081 accordingly contains five specimens with a label in Godwin-Austen’s handwriting stating “Type”. We hereby designate the specimen from lot NHMUK 1903.7.1.3081 which is figured in the original description and in
Spiraculum kempi Godwin-Austen, 1915: 496, 497, pl. 39, figs 4, 4a, 5, 5a.
Pearsonia kempi
–
Lectotype
(design. nov.) NHMUK 1903.7.1.3105/1 (
The type series of this species comprises two specimen lots, “No. 3105 Brit. Mus.” and “No. 3047 B.M.”, and two uncatalogued specimens in the Indian Museum. Without any explicit holotype designation or equivalent expression, and given that the Register of Godwin-Austen Collection explicitly states that each lot contains two specimens (Fig.
Spiraculum planum Godwin-Austen, 1915: 497, pl. 39, fig. 6, 6a, b.
Syntypes NZSI M.5992 from Upper Rotung, Abor Hills; NZSI M.5992a from Yembung.
NHMUK 1903.7.1.3596 (1 shell) from the Miri Hills.
The type series of this species comprises two specimen lots, except for lot “No. 3596 B.M.” that
Pterocyclos aborensis
Godwin-Austen, 1915: 498, pl. 39, fig. 1, 1a.
Lectotype
(design. nov.) NHMUK 1903.7.1.3104/1 (
The type series of this species comprises three specimen lots. Without any explicit holotype designation or equivalent expression, all specimens in these lots are syntypes. However, only specimen lot “No. 3104 Brit. Mus.” from Abor Hills was mentioned in the “Type” paragraph, one figured specimen from this lot was marked as “Type” in the plate caption, and this lot was labelled as “Type” in Godwin-Austen’s handwriting (Fig.
Pterocyclos miriensis Godwin-Austen, 1915: 498, pl. 39, fig. 2, 2a–c.
Lectotype
(design. nov.) NHMUK 1903.7.1.3580/1 (Fig.
Pterocyclos spiramentum
Godwin-Austen, 1915: 498, 499, pl. 40, fig. 4, 4a, b.
Holotype
NHMUK 1903.7.1.3082 (
Pterocyclos brahmakundensis
Godwin-Austen, 1915: 499, 500, with text figure 1.
Lectotype
(design. nov.) NHMUK 1903.7.1.713/1 (
? Spiraculum luyorensis Godwin-Austen, 1915: 500, pl. 40, fig. 5, 5a, b.
Pearsonia luyorensis
–
Lectotype
(design. nov.) NHMUK 1903.7.1.3530/1 (
Only one specimen lot, “No. 3530 Brit. Mus.”, was mentioned in the original description and associated with the “Type” paragraph. Although this type lot contains only one specimen and the Register of Godwin-Austen Collection reveals that there is only one specimen in this lot (Fig.
Spiraculum putaoensis Godwin-Austen, 1915: 500, 501, pl. 40, fig. 3, 3a, b.
Pearsonia putaoensis
–
Lectotype
(design. nov.) NHMUK 1903.7.1.3598/1 (
Spiraculum minimum Godwin-Austen, 1915: 501, 502, pl. 40, fig. 2, 2a–c.
Pearsonia minima
–
Lectotype
NHMUK 1903.7.1.3145/1 (
The type series of this species comprises four specimen lots. Only two specimen lots, “No. 6142–43 Ind. Mus.”, which were explicitly stated to contain two specimens from “Jeku, Abor Hills”, were mentioned in the “Type” paragraph, and one of these two specimens was figured and marked as “Type” in the plate caption. In contrast, the remaining lots were mentioned in the body of the text of subsequent paragraphs. This could mean that
Cyclophorus oakesi Godwin-Austen, 1915: 502, pl. 40, fig. 1, 1a.
Theobaldius oakesi
–
Lectotype
(design. nov.) NHMUK 1903.7.1.3083/1 (Fig.
We thank F. Naggs and H. Taylor (NHM, London) for photographing the type material and allowing the authors to examine the material housed in the type collections. We also thank F. Welter-Schultes, G. Rosenberg, A. Breure, T. Backeljau and an anonymous reviewer for the most insightful and valuable comments that greatly improved the manuscript. This project was funded through grants received from Ratchadapisek Somphot Fund for Postdoctoral Fellowship, Chulalongkorn University to PJ; the Darwin Initiative Project no. 14–653, the TRF Strategic Basic Research DBG 6080011 (2017–2020) and The Thailand Research Fund (TRF-DPG628001).