Bringing light into deep-sea biodiversity: a systematic revision and molecular phylogeny of the genus Scaphander Montfort, 1810 (Gastropoda: Cephalaspidea), with a focus on the Indo-Pacific Author Siegwald, Justine Department of Natural History, University Museum of Bergen, University of Bergen, PB 7800 5020 Bergen, Norway justine.siegwald@gmail.com Author Malaquias, Manuel António E. Department of Natural History, University Museum of Bergen, University of Bergen, PB 7800 5020 Bergen, Norway text Zoological Journal of the Linnean Society 2025 2024-05-17 203 1 60 journal article 308540 10.1093/zoolinnean/zlad201 4cc80347-d129-4cf6-9b65-be01da03fbf8 0024-4082 15006324 A9AFDD7-B8BE-47EB-9676-77488EE78A24

Table 1. List of specimens for sequence analyses, with sampling localities, voucher numbers, and GenBank and BOLD accession numbers (numbers marked in bold are novel sequences generated for the present study).

Species	Voucher	Code	Locality	COI	12S	16S	18S	28S
Eoscaphander fragilis Habe, 1952	ZMBN 131874	SJ149	Off Hachinohe, Aomori, Honshu Island, Japan	MZ473267	MZ478726	MZ478672	MZ479109	MZ479064
Nipponoscaphander japonicus (Adams, 1862)	ZMBN 127895	YK4376	Off Misaki Marine Biological Station, Kanagawa, Japan	MZ473268	MZ478728	MZ478673	MZ479111	MZ479065
Sabatia bathymophila (Dall, 1881)	MNHN-IM-2013-67211	SJ184	South Plain, Walters Shoals, Indian Ocean	MZ473274	MZ478733	MZ478679	MZ479119	MZ479069
Scaphander amygdalus	NIWA 30258	st31	New Zealand	MZ473299	MZ478758	MZ478704	MZ479145
Scaphander amygdalus	NIWA 30374 A	SJ42	Off Western New Zealand	MZ473300	MZ478759	MZ478705	MZ479146
Scaphander amygdalus	AM C.563070	SJ77	Hunter Commonwealth Marine Reserve, New South Wales, Australia	OR552922		OR557483	OR559122	OR555862
Scaphander amygdalus	AM C.519351	SJ78	Central Eastern Commonwealth Marine Reserve, New South Wales, Australia	OR552923		OR557484	OR559123	OR555863
Scaphander amygdalus	NIWA 30512	SJ200	New Zealand	OR552924	OR557468
Scaphander amygdalus	NIWA 30469	SJ203	New Zealand	OR552925	OR557469
Scaphander amygdalus	NIWA 30235	SJ216	New Zealand	OR552926
Scaphander amygdalus	NIWA 30291	SJ217	New Zealand	OR552927
Scaphander amygdalus	NIWA 48567	SJ218	New Zealand	OR552928
Scaphander cancellatus Martens, 1902	MNHN-IM-2007-35413	NP7	Off Balicasag Island, Bohol Sea, Philippines	BOLD PHILI001-10
Scaphander cancellatus Martens, 1902	MNHN-IM-2009-4339	SC33	Bohol Sea, Philippines	KC351574	MZ478768	KC351539	MZ479155	KC351556
Scaphander cancellatus Martens, 1902	MNHN-IM-2013-52478	SJ18	New Georgia, Solomon Islands	OR552929
Scaphander cancellatus Martens, 1902	MNHN-IM-2009-6678	SJ20	East of San Cristobal, Solomon Islands	OR552930		OR557485
Scaphander cancellatus Martens, 1902	MNHN-IM-2013-52472	SJ29	Solomon Islands	OR552931		OR557486
Scaphander cancellatus Martens, 1902	MNHN-IM-2007-35412	SJ35	Bohol Sea, Philippines	MZ473309	MZ478769	MZ478714	MZ479156	MZ479103
Scaphander cancellatus Martens, 1902	MNHN-IM-2019-7925	SJ198	Tanimbar Islands, Indonesia	OR552932
Scaphander cornus	AM C.563069	SJ15	Tasmania/Victoria, Australia	MZ473302	MZ478762	MZ478707	MZ479148	MZ479097
Scaphander cornus	AM C.519368	SJ38	East of Cape St George, New South Wales, Australia	MZ473303	MZ478763	MZ478708	MZ479149	MZ479098
Scaphander cornus	AM C.590959	SJ205	Bass Strait, Tasmania/Victoria, Australia	OR552933	OR557470
Scaphander cornus	AM C.590968	SJ221	Off Cape Howe, Victoria, Australia	OR552934
Scaphander darius Marcus & Marcus, 1967	MZSP 29016	SC21	Brazil	KC351560		KC351521
Scaphander gracilis Watson, 1883	DBUA 1630	SJ104	Azores	MZ457932	MZ478742	MZ478691	MZ479130	MZ479081
Scaphander grandis (Minichev, 1967)	MIMB 36540	CH207	Kuril-Kamchatka Abyssal Plain	MK952767		MK952768		MK952769
Scaphander grandis (Minichev, 1967)	ZSM Mol 20150062	st2	Kuril-Kamchatka Abyssal Plain	MZ473286	MZ478743	MZ478692	MZ479131	MZ479082
Scaphander grandis (Minichev, 1967)	ZSM Mol 20210096	st3	Kuril-Kamchatka Abyssal Plain	MZ473287	MZ478744	MZ478693	MZ479132	MZ479083
Scaphander grandis (Minichev, 1967)	ZSM Mol 20210053	SJ68	Kuril-Kamchatka Abyssal Plain	OR552935	OR557471	OR557488		OR555864
Scaphander interruptus Dall, 1890b	NIWA 30427 A	SJ199	Off Western New Zealand	MZ473297	MZ478756	MZ478702	MZ479143	MZ479092
Scaphander interruptus Dall, 1890b	AM C.563068	SJ201	Off Cape Howe, Victoria, Australia	MZ473298	MZ478757	MZ478703	MZ479144	MZ479093
Scaphander interruptus Dall, 1890b	AM C.590961	SJ219	Bass Strait, Tasmania/Victoria, Australia	OR552936
Scaphander interruptus Dall, 1890b	SCBUCN-2837	SJ236	South of Concepción, Chile	OR552937	OR557472	OR557489	OR559124	OR555865
Scaphander cf. lignarius 1	ZMBN 88000	SC37	Bergen, Norway	KC351563	MZ478745	KC351526	MH933326	KC351545
Scaphander cf. lignarius 1	ZMBN 127893	SJ49	Cadiz, Spain	MZ473288	MZ478746	MZ478694	MZ479133	MZ479084
Scaphander cf. lignarius 2	ZMBN 127869	SJ5	Vigo, Spain	MZ473289	MZ478747	MZ478695	MZ479134	MZ479085
Scaphander cf. lignarius 2	ZMBN 127870	SJ50	Cadiz, Spain	MZ473290	MZ478748	MZ478696	MZ479135	MZ479086
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	ZMBN 127881	st11	Off Mar del Plata, Argentina	MN433681	MZ478749	MN450225	MZ479136	MN450265
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	ZMBN 127882	st17	Off Mar del Plata, Argentina	MN433676		MN450226	OR559125
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	AM C.590963	SJ14	Freycinet Commonwealth Marine Reserve, Tasmania, Australia	OR552938	OR557473	OR557490	OR559126	OR555866
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	ZMBN 127882	SJ36	Off Mar del Plata, Argentina	MN433682		MN450221		MN450263
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	AM C.482252	SJ39	Hunter Commonwealth Marine Reserve, New South Wales, Australia	OR552939	OR557474	OR557491	OR559127	OR555867
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	AM C.519366	SJ74	Freycinet Commonwealth Marine Reserve, Tasmania, Australia	MZ473291	MZ478750	MZ478697	MZ479137	MZ479087
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	AM C.590966	SJ75	Jervis Commonwealth Marine Reserve, New South Wales, Australia	OR552940		OR557492	OR559128	OR555868
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	MACN-In 42431	SJ101	Off Mar del Plata, Argentina	MN433680		MN450223
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	MACN-In 42432	SJ102	Off Mar del Plata, Argentina	OR552941
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	AM C.590967	SJ220	East Gippsland Commonwealth Marine Reserve, Victoria, Australia	OR552942
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	AM C.594398	SJ225	Bass Strait, Tasmania/Victoria, Australia	OR552943
Scaphander meridionalis Siegwald, Pastorino, Oskars & Malaquias, 2020	AM C.594399	SJ226	Bass Strait, Tasmania/Victoria, Australia	OR552944
Scaphander mundus Watson, 1883	MNHN-IM-2013-61614	NP1	Southwest of DongSha, South China Sea	OR552945
Scaphander mundus Watson, 1883	MNHN-IM-2013-61612	NP2	Southwest of DongSha, South China Sea	OR552946
Scaphander mundus Watson, 1883	MNHN-IM-2013-50091	NP3	Southwest of Taiwan, South China Sea	OR552947
Scaphander mundus Watson, 1883	MNHN-IM-2013-50090	NP4	Southwest of Taiwan, South China Sea	OR552948
Scaphander mundus Watson, 1883	MNHN-IM-2013-50089	NP5	Southwest of Taiwan, South China Sea	OR552949
Scaphander mundus Watson, 1883	MNHN-IM-2013-50039	NP6	Southwest of Taiwan, South China Sea	OR552950
Scaphander mundus Watson, 1883	MNHN-IM-2009-4319	SC29	Philippines	KC351565	OR557475	KC351529	OR559129	KC351547
Scaphander mundus Watson, 1883	MNHN-IM-2009-4318	SC31	Philippines	KC731429	MZ478751	KC351528	MZ479138	KC351546
Scaphander mundus Watson, 1883	MNHN-IM-2013-44305	st15	Off Taiping Island, South China Sea	OR552951	OR557476
Scaphander mundus Watson, 1883	MNHN-IM-2013-44310	st28	Off Taiping Island, South China Sea	OR552952	OR557477		OR559130	OR555869
Scaphander mundus Watson, 1883	AM C.519372	SJ12	East of Bundaberg, Queensland, Australia	MZ473292	MZ478752	MZ478698	MZ479139	MZ479088
Scaphander nobilis Verrill, 1884	ZMBN 127875	SJ67	Azores	MN433679		MN450222
Scaphander nobilis Verrill, 1884	MNHN-IM-2013-67215	SJ183	South Plain, Walters Shoals, Indian Ocean	MZ473294	MZ478753	MZ478699	MZ479140	MZ479089
Scaphander obnubilus	AM C.519273	SJ40	East of Cape St George, New South Wales, Australia	MZ473305	MZ478765	MZ478710	MZ479151	MZ479100
Scaphander obnubilus	AM C.482192	SJ76	Northwest of Flinders Island, Tasmania, Australia	MZ473306	MZ478766	MZ478711	MZ479152	MZ479101
Scaphander obnubilus	AM C.590969	SJ223	Bass Strait, Tasmania/Victoria, Australia	OR552953
Scaphander otagoensis Dell, 1956	MNZ 301800/1.1	st6	New Zealand	MZ473295	MZ478754	MZ478700	MZ479141	MZ479090
Scaphander otagoensis Dell, 1956	NIWA 63032 A	SJ79	South of the North Island, New Zealand	MZ473296	MZ478755	MZ478701	MZ479142	MZ479091
Scaphander otagoensis Dell, 1956	MNZ M.301800/1.2	SJ109	New Zealand	OR552954
Scaphander otagoensis Dell, 1956	NIWA 30182	SJ202	New Zealand	OR552955	OR557478
Scaphander otagoensis Dell, 1956	NIWA 63804 A	SJ204	New Zealand	OR552956	OR557479
Scaphander otagoensis Dell, 1956	MNZ M.301817/1.1	SJ227	New Zealand	OR552957
Scaphander otagoensis Dell, 1956	MNZ M.301817/1.2	SJ228	New Zealand	OR552958
Scaphander punctostriatus (Mighels & Adams, 1842)	ZMBN 88006	SC34	Norway	KC351571	MZ478760	KC351536	MH933325	KC351553
Scaphander punctostriatus (Mighels & Adams, 1842)	DBUA 1629	SJ105	Azores	MZ473301	MZ478761	MZ478706	MZ479147	MZ479096
Scaphander punctostriatus (Mighels & Adams, 1842)	ZMBN 127899	SJ207	Gulf of St. Lawrence, Québec, Canada	OR552959	OR557480	OR557493	OR559131	OR555870
Scaphander semicallus	MNHN-IM-2013-52464	st16	Inhambane, Mozambique	MZ473304	MZ478764	MZ478709	MZ479150	MZ479099
Scaphander sibogae Schepman, 1913	MNHN-IM-2013-18549	st14	East of Kotakot, Papua New Guinea	MZ473284	MZ478740	MZ478689	MZ479128	MZ479079
Scaphander sibogae Schepman, 1913	MNHN-IM-2019-7922	SJ17	New Georgia, Solomon Islands	OR552960
Scaphander sibogae Schepman, 1913	MNHN-IM-2013-52471	SJ45	Northwest of Isabel, Solomon Islands	MZ473285	MZ478741	MZ478690	MZ479129	MZ479080
Scaphander sibogae Schepman, 1913	MNHN-IM-2019-7918	SJ168	Big Bay, Santa, Vanuatu	OR552961
Scaphander sibogae Schepman, 1913	MNHN-IM-2013-52477	SJ188	New Georgia, Solomon Islands	OR552962
Scaphander sibogae Schepman, 1913	MNHN-IM-2013-58331	SJ189	New Ireland, Papua New Guinea	OR552963
Scaphander solomonensis	MNHN-IM-2013-52480	st21	Northwest of Choiseul, Solomon Islands	MZ473307		MZ478712	MZ479153
Scaphander solomonensis	MNHN-IM-2019-7923	SJ163	Northwest of Isabel, Solomon Islands	OR552964
Scaphander solomonensis	MNHN-IM-2013-52483	SJ166	Northwest of Choiseul, Solomon Islands	MZ473308	MZ478767	MZ478713	MZ479154	MZ479102
Scaphander teramachii (Habe, 1954)	ZMBN 131888	SJ150	West of Takara Island, East China Sea, Japan	MZ473310	OR557481	MZ478715	OR559132	OR555871
Scaphander teramachii (Habe, 1954)	ZMBN 131891	SJ151	Tosa Bay, Kochi, Shikoku Island, Japan	MZ473311	MZ478770	MZ478716	MZ479157	MZ479104
Scaphander teramachii (Habe, 1954)	ZMBN 131888	SJ229	West of Takara Island, East China Sea, Japan	OR552966
Scaphander teramachii (Habe, 1954)	ZMBN 131888	SJ230	West of Takara Island, East China Sea, Japan	OR552967
Scaphander teramachii (Habe, 1954)	ZMBN 131891	SJ231	Tosa Bay, Kochi, Shikoku Island, Japan	OR552968
Scaphander teramachii (Habe, 1954)	ZMBN 131891	SJ232	Tosa Bay, Kochi, Shikoku Island, Japan	OR552969
Scaphander watsoni Dall, 1881	MNHN-IM-2013-56523	SJ19	French Guyana	MZ473312	MZ478771	MZ478717	MZ479158	MZ479105
Scaphander watsoni Dall, 1881	MNHN-IM-2013-60242	SJ58	North of Grande-Terre, Guadeloupe	MZ473313	MZ478772	MZ478718	MZ479159	MZ479106