Biodiversity Data Journal :
Taxonomy & Inventories
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Corresponding author: Katarzyna Vončina (katarzyna.voncina@senckenberg.de), Julia D. Sigwart (julia.sigwart@senckenberg.de)
Academic editor: Andrew Davinack
Received: 14 Jul 2023 | Accepted: 28 Sep 2023 | Published: 28 Nov 2023
© 2023 Katarzyna Vončina, Nina Mikkelsen, Christine Morrow, Rory Ang, Julia Sigwart
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:
Vončina K, Mikkelsen NT, Morrow C, Ang R, Sigwart JD (2023) Clarification of the taxonomic status of Acanthochitona discrepans (Brown, 1827) with new data for the North-East Atlantic Acanthochitona (Polyplacophora, Acanthochitonidae). Biodiversity Data Journal 11: e109554. https://doi.org/10.3897/BDJ.11.e109554
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The genus Acanthochitona can be easily distinguished from other chitons by having eighteen tufts of bristles on the dorsal side of the densely spiculose girdle. In the North-East Atlantic, five species of this genus have been recognised so far: A. crinita (Pennant, 1777), A. discrepans (Brown, 1827), A. fascicularis (Linnaeus, 1767), A. oblonga Leloup, 1968 and A. pilosa Schmidt-Petersen, Schwabe et Haszprunar, 2015. The nomenclature of A. crinita, A. discrepans and A. fascicularis was confused for a very long time until Kaas (1985) designated type specimens for them and provided a brief key. However, his work lacked detailed descriptions of the three species and some authors doubted that A. discrepans constitutes a separate species. Subsequently, the taxonomic status of A. discrepans has remained unclear.
Here, we implemented an integrative approach which combined morphology and molecular evidence to show that Acanthochitona discrepans is, indeed, a valid species and we present re-descriptions for A. crinita, A. discrepans and A. fascicularis.
Acanthochitona, geographic ranges, integrative taxonomy, North-East Atlantic chitons
The class Polyplacophora, also known as chitons, comprise one of the eight extant classes of molluscs. These exclusively marine animals can be found worldwide, from the intertidal zone to the deep sea (
In the North-East Atlantic, five species of Acanthochitona have been recognised so far: A. crinita (Pennant, 1777), A. discrepans (Brown, 1827), A. fascicularis (Linnaeus, 1767), A. oblonga Leloup, 1981 and A. pilosa Schmidt-Petersen, Schwabe et Haszprunar, 2015 (
In the current study, we used morphological and molecular data of Acanthochitona from the North-East Atlantic using freshly-collected specimens and museum material. An integrative approach showed that A. discrepans is a valid species; additionally, we included re-descriptions for A. crinita, A. discrepans and A. fascicularis.
Specimens for SEM analysis were dissected and then the valves, girdle and radula were transferred on to a glass cube with bleach and left until the rest of the tissue had dissolved. Prepared parts were gold-sputtered and examined in CamScan "CS 24" from Cambridge Instruments available at the Senckenberg Research Institute in Frankfurt. All figures were assembled in Adobe Photoshop CS6.
For DNA barcoding, a small piece of tissue from the chiton foot was sampled. DNA from the specimens ZSM20040263 and ZSM20150336 was extracted using QIAamp DNA Micro Kit (QIAGEN). Specimens SMF 373026–36, ZMBN140293–6, ZMBN146755 were sent to BOLD for the extraction and sequencing (The Barcode of Life Data System, https://www.boldsystems.org/). Specimens ZMBN140331–33, ZMBN138238, SMF 363941–3, SMF 360506 and SMF 373024–5 were extracted using the Qiagen DNeasy kit and amplified using TaKaRa Taq HS. For extraction procedures, the manufacturers’ protocols were followed. The cytochrome oxidase subunit I (COI primers LCO1490 and HCO2198;
The new sequences are publicly available in The Barcode of Life Data System (BOLD, https://www.boldsystems.org) and GenBank (https://www.ncbi.nlm.nih.gov/genbank/); see "Materials" sections of each species for their accession numbers (as "associatedSequences"). The list of DNA sequences downloaded from GenBank, the alignment and genetic distances are available as Suppl. materials
Chiton crinitus: Pennant, 1777 -
non Chiton crinitus: Sowerby II, 1840a -
Chiton onyx: Spengler, 1797 -
non Chiton onyx: Morch, 1870 -
Chiton fascicularis: Brown, 1827 -
non Chiton fascicularis: Linnaeus, 1767 -
Chiton fascicularis var. minor: Philippi, 1836 -
Acanthochaetes vulgaris: Leach, 1852 -
Chiton fascicularis var. attenuata: Jeffreys, 1865 -
Acanthochites aeneus: di Monterosato, 1878a -
non Acanthochites aeneus: Risso, 1826 -
Acanthochites (ton) adansoni: de Rochebrune, 1881a -
Acanthochites (tori) bouvieri: de Rochebrune, 1881a -
Anisochiton (Acanthochites) fascicularis var. violacea: Pallary, 1902 -
Acanthochites fascicularis vars var. lutescens, cinnabrina et fusca: Dautzenberg & Durouchoux, 1906 -
Acanthochitona crinitus: Winckworth, 1926 -
Acanthochiton fascicularis: Leloup, 1936 -
non Chiton fascicularis: Linnaeus, 1767 -
Acanthochiton gracilis: Leloup, 1968 -
non Chiton gracilis Jeffreys, 1859 -
Acanthochitona crinita: Kaas, 1985 -
Type material. Neotype: RSMNH 1978.052.02601, Royal Scottish Museum of Natural History, Edinburgh, Scotland. Designated by
Type locality. Scotland, sea near Aberdeen (
(See
Head valve slightly wider than long, almost semicircular, five rays barely raised, apex clearly visible (Fig.
Tegmentum covered with densely packed to widely separated (density from 23 to 38 granules per 1 mm2, mean = 31, n = 8 specimens) granules of variable shape, from oval to more or less elongate drop-shaped; granules raised, flat-topped, with one macroaesthete subcentral surrounded by 8–16 posteromedially located microaesthetes (Fig.
Articulamentum thick, apophyses large, broadly rectangular, rounded at anterolateral margins, separated by a wide sinus; insertion plates wide, continuous with apophyses, slits moderately deep (slit formula: 5/1/2) (Fig.
Girdle wide, fleshy, usually brighter than tegmentum, deeply encroaching sutural areas. Highly variable colour of the tegmentum and perinotum; usually from creamy-white to olive-green and brown, with different degrees of blotches in which white, yellow and brown variously combined; in some specimens, colours can be very bright, for example, bright yellow and orange. (Fig.
Radula central tooth elongated, almost straight at the top and slightly keeled near base; first lateral tooth wing-shaped and smaller than the central; second (major) lateral elongated, with accessory plate tricuspidate, outer denticle slightly shorter than the others; cusps pointed (Fig.
Gills merobranchial, 13–15 ctenidia on each side.
Acanthochitona crinita is well known for its high variability of morphological features. However, it can be distinguished from other NE Atlantic relatives on the basis of the below-mentioned morphological characters. A. crinita can be distinguished from A. discrepans (Brown, 1827), based on the number, size and arrangement of aesthetes on tegmental granules (one macroaesthete subcentral surrounded by larger, posteromedially-located 8–16 microaesthetes in A. crinita vs. 1–2 macroaesthetes: a single macroaesthete located in posterior third, second, if present, located centrally, surrounded by very small, posteriomedially located 26–40 microaesthetes in A. discrepans), dorsal spicules (longer, thicker spicules up to 100 x 15 µm in A. crinita vs. smaller, thinner spicules up to 68 x 7.5 µm in A. discrepans), tuft needles (longer spicules up to 340 x 20 µm, surrounded by a smaller number of very short and thin needles in A. crinita vs. longer spicules up to 1000 µm x 60 µm, surrounded by a large number of somewhat shorter and much thinner needles in A. discrepans). A. crinita differs from A. fascicularis (Linnaeus, 1767) by the shape of the intermediate valves (ellipsoidal in A. crinita vs. triangular in A. fascicularis), jugal area (hardly raised, not sharply separated from the latero-pleural areas in A. crinita vs. raised, sharply separated from the latero-pleural areas in A. fascicularis), shape of dorsal granules (moderately to widely apart, oval to elongated oval in A. crinita vs. small, round, densely packed granules with the incision in the middle in A. fascicularis), density of tegmental granules (23–38 granules per 1 mm2, mean = 31 in A. crinita vs. 42–70 granules per 1 mm2, mean = 52 in A. fascicularis), number of microaesthetes (8–16 microaesthetes in A. crinita vs. 0–6 microaesthetes in A. fascicularis), number of bristles in the sutural tufts (20–46 in A. crinita vs. 55-120 in A. fascicularis). It differs from A. oblonga Leloup, 1981 by the shape of dorsal granules (moderately apart, oval to elongated oval in A. crinita vs. widely apart and very much elongated in A. oblonga), number and arrangement of microaesthetes (8–16 microaesthetes located posteromedially in A. crinita vs. 6–9 microaesthetes, located mainly in the central area in A. oblonga). It can be easily distinguished from A. pilosa Schmidt-Petersen, Schwabe et Haszprunar, 2015 by the shape of the IV valve (valve with the hind margin concave at both sides of the pronounced apex in A. crinita vs. valve with triangular posterior margin with no apex in A. pilosa), density of tegmental granules (moderately to widely apart in A. crinita vs. densely packed in A. pilosa), size of microaesthetes on the tegmental granules (bigger microaesthetes in A. crinita vs. smaller, ca. ½ of the diameter, in A. pilosa), large dorsal spicules (not always present, sparse, curved spicules of size up to 260 x 40 µm in A. crinita vs. always present, dense and straight spicules of size up to 320 × 60-62 µm in A. pilosa).
Acanthochitona crinita is a widely distributed species in North-East Atlantic. The species inhabits the seas from of north of Scotland (locus typicus), south of England (molecular data and SEM photos from this study), north of France and Spain, west coast of Portugal (molecular data from GenBank), the Azores (molecular data and SEM photos from this study) and, most likely, the Mediterranean Sea (literature data).
Chiton discrepans: Brown, 1827 -
non Chiton discrepans: Sowerby II, 1840a: 2 [in synonymy of C. crinitus (non Pennant)] et mult. auct.
Chiton fascicularis: Brown, 1827 -
non Chiton fascicularis: Linnaeus, 1767 -
Chiton gracilis: Jeffreys, 1859 -
Chiton fascicularis var. gracilis: Jeffreys, 1865 -
Acanthochitona discrepans: Winckworth, 1926 -
Chiton gracile: Warén, 1980 -
Acanthochtona crinita: Kaas, 1985 -
Acanthochitona discrepans: Kaas, 1985 -
Type material. Lectotype: TENBM 1983.4588/1, Tenby Museum, Tenby, Pembroke, Wales. Designated by
Type locality. Wales, Pembroke, Tenby (
Animal small to medium size (in examined material, BL: 11–22 mm, BW: 5–11 mm, BL/BW ratio: 2.1), outline oval, moderately elevated (elevation ratio 0.41), semi-carinated, side slopes flat to slightly raised, apices not prominent; girdle wide, spiculose (Fig.
Head valve slightly wider than long, almost semicircular, anterior slope slightly convex, posteriormost margin straight without a notch (Fig.
Tegmentum uniformly covered with rather densely distributed (density from 24 to 50 granules per 1 mm2, mean = 37, n = 8 specimens), oval to elongated granules, arranged in quincunx order, except for the jugal area of intermediate valves; the granules raised, flat topped, to slightly concave (Fig.
Articulamentum well developed, solid, slightly pinkish ventrally beneath tegmentum. Apophyses large, separated at jugum, broadly rectangular to wing-shaped, trapezoidal in tail valve (direct dorsal view can create appearance of triangular outline); insertion plates wide, continuous with apophyses, slits deep, extending proximally into shallow dorsal channels (slit formula: 5/1/2) (Fig.
Girdle wide, fleshy, leathery looking, brighter than tegmentum, deeply encroaching sutural areas. Colour from beige or light yellow to pale brown with brown and olive-green streaks and blotches, in some specimens prominent dark green bands with fuzzy boundaries located near valves between the sutural tufts; sutural tufts with translucent or brownish spicules; base of sutural tufts creamy-yellow or brown (Fig.
Radula central tooth elongated, without cusp, almost straight at the top, keeled near base; first lateral tooth wing-shaped and wrapping around central; second (major) lateral massive with accessory plate tricuspidate with outer denticle shorter than the others; cusps pointed and triangular in outline (Fig.
Gills merobranchial, composed of 9–13 ctenidia on each side reaching around half body length.
This species can be distinguished from other Acanthochitona species from NE Atlantic on the basis of a set of morphological characters. It differs from A. crinita (Pennant, 1777) by the number, size and arrangement of aesthetes on tegmental granules (1–2 macroaesthetes - a single macroaesthete located in posterior third, second, if present, located centrally, surrounded by very small, posteriomedially located 26–40 microaesthetes in A. discrepans vs. one macroaesthete subcentral surrounded by larger, posteromedially located 8–16 microaesthetes in A. crinita), dorsal spicules (smaller, thinner spicules up to 68 x 7.5 µm in A. discrepans vs. longer, thicker spicules up to 100 x 15 µm in A. crinita), tuft needles (longer spicules up to 1000 µm x 60 µm, surrounded by a large number of somewhat shorter and much thinner needles in A. discrepans vs. longer spicules up to 340 x 20 µm, surrounded by a smaller number of very short and thin needles in A. crinita). Acanthochitona discrepans can be distinguished from A. fascicularis (Linnaeus, 1767) by the shape of the intermediate valves (ellipsoidal in A. discrepans vs. triangular in A. fascicularis), jugal area (hardly raised, not sharply separated from the latero-pleural areas in A. discrepans vs. raised, sharply separated from the latero-pleural areas in A. fascicularis), shape of dorsal granules (decidedly apart, oval to elongated oval in A. discrepans vs small, round, densely packed granules with the incision in the middle in A. fascicularis), density of tegmental granules (24–50 granules per 1 mm2, mean = 37 in A. discrepans vs. 42–70 granules per 1 mm2, mean = 52 in A. fascicularis), number of microaesthetes (26–40 microaesthetes in A. discrepans vs. 0–6 microaesthetes in A. fascicularis), number of bristles in the sutural tufts (15–40 in A. discrepans vs. 55-120 in A. fascicularis). Acanthochitona discrepans differs from A. oblonga Leloup, 1981 by the shape of dorsal granules (rather densely distributed, oval to elongated granules, arranged in quincunx order in A. discrepans vs. widely apart and very much elongated in A. oblonga), number and arrangement of microaesthetes (26–40 microaesthetes located posteromedially in A. discrepans vs. 6–9 microaesthetes located mainly in the central area in A. oblonga). It can be distinguished from A. pilosa Schmidt-Petersen, Schwabe et Haszprunar, 2015 on the basis of the shape of the IV valve (very slightly-beaked valve with hind margin slightly concave at both sides of the apex in A. discrepans vs. valve with triangular posterior margin with no apex in A. pilosa), number of microaesthetes on the tegmental granules (26–40 microaesthetes in A. discrepans vs. around 14 microaesthetes in A. pilosa), large dorsal spicules (only sometimes present and very sparse large dorsal spicules in A. discrepans vs. always present, dense and straight spicules in A. pilosa).
This species seems to have a more northern distribution than Acanthochitona crinita. The range of species is from Norway (molecular data and SEM photos from this study), through the north coast of Ireland (molecular data and SEM photos from this study) to the south of Wales (locus typicus). The literature records from more southern regions must be treated with caution as they probably represent other misidentified Acanthochitona species.
The original locus typicus of Acanthochitona discrepans (Brown, 1827) is in Tenby, Pembroke in Wales.
Chiton fascicularis: Linnaeus, 1767 -
non C. fascicularis: Sowerby, 1840a -
Acanthochites communis: Risso, 1826 -
Chiton fascicularis var. major: Philippi, 1836 -
Chiton crinitus: Sowerby, 1840: figs. 88-93; 1840a: 2.
non C. crinitus: Pennant, 1777 -
Chiton discrepans: Sowerby, 1840a: 2 (in synonymy of C. crinitus; Sowerby, non Pennant, 1777); Forbes & Hanley, 1849 -
non C. discrepans: Brown, 1827 -
Acanthochites carinatus: H. Adams & Angas, 1864
non Acanthochites carinatus: Risso, 1826 -
Chiton fascicularis var. rubra: Issel, 1870 -
Acanthochites discrepans var. minorflava: di Monterosato, 1878a -
Acanthochites hamatus: de Rochebrune, 1882 -
Anisochiton discrepans vars elongata, marmorata, nigrolineata: Dautzenberg, 1893 -
Acanthochites discrepans var. albina: Dautzenberg & Durouchoux, 1900 -
Anisochiton (Acanthochites) discrepans var. viridis: Pallary, 1902 -
Acanthochites discrepans var. violaceolimbata: Dautzenberg & Durouchoux, 1906 -
Acanthochiton(a) communis: Winckworth, 1926 -
Acanthochiton heterochaetus: Bergenhayn, 1931 -
Acanthochiton discrepans var. angustivalvus: Bergenhayn, 1931 -
Acanthochiton communis f. barashi: Leloup, 1969 -
Acanthochitona bonairensis: Kaas, 1972 -
Acanthochitona fascicularis: Kaas, 1985 -
Type material. Neotype: MNHN-IM-2000-5923, Muséum National d’Histoire Naturelle, Paris, France. Designated by Kaas (1985) (material not seen).
Type locality. Algeria, coast of Barbary (
(See
Head valve much wider than long, semicircular, with five barely-raised rays, posterior margin almost straight, no notch and apex (Fig.
Tegmentum uniformly and thickly covered (density from 42 to 70 granules per 1 mm2, mean = 52, n = 7 specimens) by small, round granules arranged in arched lines on the valves; each granule with 0–6 aesthetes randomly distributed, without a clear separation between macro- and microaesthetes; some granules with an incision in the middle of the granule posterior margin (Fig.
Articulamentum well-developed; apophyses wide, rectangular, separated at jugum, rounded at anterolateral margins; insertion plates wide, continuous with apophyses, slits deep, extending dorsally into shallow channels (slit formula: 5/1/2) (Fig.
Girdle very wide and fleshy, deeply encroaching sutural areas. Colours ranging from creamy-yellow to olive-green and brown, with different degrees of blotches in which white, green and brown variously combined; sometimes yellow or orange (Fig.
Central radula teeth elongated, almost straight at the top and keeled near base; first lateral tooth wing-shaped and smaller than the central; second (major) lateral elongated with accessory plate tricuspidate with central denticle longer than the others; cusps pointed and triangular in outline (Fig.
Gills merobranchial, 15-17 ctenidia on each side.
Acanthochtiona fascicularis, can be easily distinguished from other NE Atlantic relatives by the triangular shape of valves, raised and clearly separated jugal area, small and round, densely-packed tegmental granules and small number of aesthetes (0–6) without clear separation between micro- and macroaesthetes. For comparison, see the Remarks section for A. crinita and A. discrepans.
This species is widely distributed in the North-East Atlantic, with a range from the west coast of Ireland (molecular data and SEM photos from this study), south to the Azores (molecular data and SEM photos from this study) and the Mediterranean Sea (literature records and GenBank sequences).
Phylogenetics results
The consensus tree obtained through Bayesian Inference is shown in Fig.
A total of 32 new COI sequences and 17 sequences downloaded from GenBank of a length of 618 bp were used for the phylogenetic analysis. No stop codons or gaps in the alignment were present. Uncorrected pairwise distances in COI sequences between the three analysed species were as follows: 14.7–15.7% between Acanthochitona fascicularis and A. crinita; 13.6–14.7% between A. fascicularis and A. discrepans; 10.8–11.3% between A. crinita and A. discrepans (uncorrected pairwise distances in COI between all sequences included in the phylogenetic analysis can be found in Suppl. material
Acanthochitona crinita, A. discrepans and A. fascicularis were confused for a very long time, but do constitute three distinct species. They can be separated, based on the morphology and differ also in their geographic distribution. Acanthochitona fascicularis has a very wide distribution, it inhabits seas from the west coast of Ireland to the Mediterranean Sea, A. crinita occurs from Scotland to the west coast of Portugal and, most likely, in the Mediterranean Sea. A. discrepans represents a more northern species than A. crinita, which has a range from northern Norway, along the north Irish coasts to the south of Wales; the ranges of A. crinita and A. discrepans probably overlap in Scotland and Ireland.
Taxonomy of Acanthochitona has been mostly based on the shape and distribution of tegmental granules combined with the girdle ornamentation; however, these characters in isolation tend to be variable and are not diagnostic (e.g.
The three NE Atlantic Acanthochitona require careful observation in order to be separated with confidence. Acanthochitona fascicularis is the most distinct species, with its large size, very broad girdle and small, densely-arranged granules. Acanthochitona crinita and A. discrepans are superficially similar and confident separation should be made, based on SEM photos or DNA barcodes. Assessment of detailed granule structure is only practicable with SEM. Girdle characters, although less reliably diagnostic, can be useful in field identification. The girdle of A. fascicularis is much broader than in the two other species and densely covered with very short thin spicules, but with spines of different lengths interspersed; A. crinita usually has the girdle covered with two kind of spines: short, thick spines and much longer, thicker and curved spines between them; A. discrepans has the cover of very short, densely-arranged spicules which give the girdle a more leathery look.
Acanthochitona crinita, A. discrepans and A. fascicularis are difficult to tell apart and past records must be treated with caution; however, with our new data, there is an emergent pattern in their distributions that helps to clarify how these species are separated. Acanthochitona fascicularis can be found from the west coasts of Ireland to the Azores and the Mediterranean Sea (
Acanthochitona crinita has always been challenging for taxonomists. It remains difficult to assess its geographic ranges because it is highly variable and apparently genuinely occupies a very broad range, overlapping with other Acanthochitona species. The confirmed range extends from the north of Scotland, along the coast of England, France, Spain and Portugal to the Azores. There are dozens of records of this species from North-East Atlantic: Iceland, Norway, Scotland, Ireland, Mediterranean Sea, Black Sea, Cape Verde Archipelago and Sao Tome and Principe Islands, but most of them are questionable (e.g.
The very wide apparent range of Acanthochitona crinita (from Scotland to the west coast of Africa as far south as São Tomé and Príncipe Islands), its well-known high polymorphism, different descriptions by different authors and confused nomenclature may be a sign of the existence of a species complex in North-East Atlantic. In the past, a priori assumption of its high variability has led to assigning all morphotypes that did not fit the description of A. fascicularis to A. crinita and, hence, the longstanding confusion about the validity of A. discrepans. Some authors provided records without further description (e.g.
The three Acanthochitona species considered herein are very variably coloured, but we noticed that specimens of multiple Acanthochitona species from Ireland, N. Ireland and England are usually more brownish or greenish, whereas those from Norway and the Azores are more diversely and vividly coloured. This could reflect the local environment in context of the substratum. Richness and variability of the background colour has been associated with higher colour polymorphism in other chiton populations; random colours of chitons on a complex and mixed colour background is thought to help avoid predation by visual predators (
Northern Ireland is located at a junction between southern warm waters and northern cold waters, which results in very high marine biodiversity in this region. Unfortunately, ongoing climate changes have led to shifting ranges in many marine animals (e.g.
We are especially grateful to our colleagues: Bernard Picton (National Museum of Northern Ireland) for help with the collection of specimens and providing us with photographs (uploaded to BOLD database), Enrico Schwabe (Bavarian State Collection of Zoology) who provided us with Acanthochitona samples and Jessie Dermody (University of Exeter) who accompanied RA in the search for Acanthochitona crinita in England. An anonymous reviewer is kindly thanked for the comments and corrections on a draft version of this paper. Funding to NTM was provided by grant #26-19 from the Norwegian Taxonomy Initiative. This is contribution #4 from the Senckenberg Ocean Species Alliance (SOSA).