Understanding the color variability and resolving taxonomic confusion in the sea cucumber Isostichopus badionotus (Echinodermata, Holothuroidea): a revision of the genus Isostichopus Author Borrero-Pérez, Giomar H. 987B3945-3A71-4DF0-B860-F23CC231F34C Santa Marta, Magdalena, Colombia. & Instituto de Investigaciones Marinas y Costeras - Invemar, Calle 25 No. 2 - 55, Playa Salguero, Rodadero, Santa Marta, Colombia. & Smithsonian Tropical Research Institute - STRI, BOX 0843 - 03092, Balboa, Panama. giomarborrero@gmail.com Author Solís-Marín, Francisco A. A2417F0D-CA2A-4BE2-A6F0-C8991F4B90EA Instituto de Ciencias del Mar y Limnología, Universidad Nacional Autónoma de México, 04510, Ciudad de Mexico, México. fasolis@cmarl.unam.mx Author Lessios, Harilaos B59566D2-3326-4026-B9B6-9AB12A3EB140 Smithsonian Tropical Research Institute - STRI, BOX 0843 - 03092, Balboa, Panama. lessiosh@si.edu text European Journal of Taxonomy 2024 2024-08-30 949 1 1 96 https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/2641/12173 journal article 302913 10.5852/ejt.2024.949.2641 34ae386a-671e-4dcb-8e86-8ecf0cfc346c 2118-9773 13710645 EA45BD5E-98F7-4229-A4FD-E377D6BC8591

Table 1 (continued on next six pages). Specimens of Isostichopus Deichmann, 1958 analyzed for the partial regions of cytochrome oxidase subunit 1 (COI-Fr1-Barcoding region, COI-Fr2) and 16S genes. Catalog number column includes voucher specimens or tissues from where the sequences were obtained. Field numbers used in phylogenetic trees correspond to the initial letter of the locality, the code and the color patterns in Fig. 1. Contractions in locality column: Mexico: Yuc = Yucatán; Belize: TC = Tobacco Cay; CBC = Carrie Bow Cay; Panama: BT = Bocas del Toro; Ga = Galeta; Ta = Taboguilla Island (Pacific); IP = Perlas Islands (Pacific); Colombia: SM = Santa Marta; MSM = Morro de Santa Marta; MPA = Morritos de Punta Aguja; Ne = Bahía Neguanje; LG = La Guajira; BA = Banco de las Ánimas; Prov = Providencia Island; Ch = Choco (Pacific); Ecuador: Gal = Galapagos; U.S. Virgin Islands: SCI = St. Croix; British Virgin Islands: GI = Guana Island; Curaçao: Cu = Curaçao; Cu-CMRS = Curaçao-Carmabi Marine Research Station; Brazil: SC = Santa Catalina; Cape Verde: SV = Sao Vicente Island; SI = Santiago Island; Senegal: DÎM = Dakar-Îles Madeleines; Liberia: K = Kligba, Rivercess County; AB = Artah Beach, Sasstow Grand Kru County; Ascension Island: EB = English Bay; G = Georgetown. Photo ID: figure number in the present paper or the process ID in the BOLD database. GenBank accession numbers in parentheses denote sequences by other authors. Astichopus multifidus (Sluiter, 1910), Stichopus horrens Selenka, 1867, and Apostichopus japonicus (Selenka, 1867) were used as outgroups in the phylogenetic analyses. An asterisk (*) in the species column indicates that the species ID in GenBank was changed from a previous name (shown in parentheses).

Species	Museum Catalog number	Field Number	Locality	Photo	GeneBank Acc. No. COI-Fr1	GeneBank Acc. No. COI-Fr2	GeneBank Acc. No. 16S	Reference
I. badionotus	TbBT0002	BT2-R	Panama-BT	8C’	–	–	MK477775	This study
I. badionotus	USNM 1659459	BT14-CH	Panama-BT	8A	–	MK477909	MK477776	This study
I. badionotus	TbBT0016	BT16-CH	Panama-BT	8B	–	–	MK477777	This study
I. badionotus	USNM 1659460	BT20-R	Panama-BT	1F–G	MK477881	–	MK477778	This study
I. badionotus	USNM 1659461	BT45-U	Panama-BT	8V	MK477882	MK477910	MK477779	This study
I. badionotus	USNM 1659462	BT48-U	Panama-BT	8W	MK477883	MK477911	MK477780	This study
I. badionotus	USNM 1659463	BT55-CH	Panama-BT	8C	MK477884	–	MK477781	This study
I. badionotus	USNM 1659464	BT62-CH	Panama-BT	8D	MK477885	–	MK477782	This study
I. badionotus	TbBT0067	BT67-BY	Panama-BT	–	–	MK477912	MK477783	This study
I. badionotus	USNM 1659465	BT69-BY	Panama-BT	–	MK477886	–	MK477784	This study
I. badionotus	TbBT0071	BT71-BY	Panama-BT	–	–	MK477913	MK477785	This study
I. badionotus	TbBT0073	BT73-BY	Panama-BT	–	–	MK477914	MK477786	This study
I. badionotus	MBMLP BT75	BT75-R	Panama-BT	–	MK477887	MK477915	MK477787	This study
I. badionotus	TbBT0083	BT83-R	Panama-BT	–	–	MK477916	MK477788	This study
I. badionotus	USNM 1659466	BT98-R	Panama-BT	8E’	MK477888	MK477917	MK477789	This study
I. badionotus	MBMLP BT106	BT106-CH	Panama-BT	8E	MK477889	–	MK477790	This study
I. badionotus	USNM 1659467	BT107-CH	Panama-BT	8F	MK477890	–	MK477791	This study
I. badionotus	USNM 1659454	BT111-CH	Panama-BT	8G	MK477891	MK477918	MK477792	This study
I. badionotus	USNM 1659455	BT112-U	Panama-BT	8X	MK477892	MK477919	MK477793	This study
I. badionotus	USNM 1659456	BT115-CH	Panama-BT	6H	MK477893	MK477920	MK477794	This study
I. badionotus	USNM 1659457	BT116-BY	Panama-BT	8L’	MK477894	MK477921	MK477795	This study
I. badionotus	USNM 1659458	BT118-CH	Panama-BT	8I	MK477895	MK477922	MK477796	This study
I. badionotus	INV TEJ1167	SM4-R	Colombia-SM	1H–I	–	–	MK477797	This study
I. badionotus	INV TEJ1168	SM5-R	Colombia-SM	8G’	–	–	MK477798	This study
I. badionotus	INV TEJ1170	SM7-R	Colombia-SM	8H’	–	MK477923	MK477799	This study
I. badionotus	INV TEJ1172	SM9-R	Colombia-SM	8H’ right	–	MK477924	MK477800	This study
I. badionotus	INV TEJ1173	SM10-R	Colombia-SM	8H’	–	MK477925	MK477801	This study
I. badionotus I. badionotus	INV TEJ1176 INV TEJ1178	SM13-BY SM15-R	Colombia-SM Colombia-SM	1J	– –	MK477926 –	MK477802 MK477803	This study This study
I. badionotus	INV EQU4572 INV TEJ1987	MSM16-U	Colombia-MSM	BECSC73-20	OL989848	–	–	This study
I. badionotus	INV EQU4573 INV TEJ1988	MPA17-R	Colombia-MPA	BECSC74-20	OL989849	–	–	This study
I. badionotus	INV TEJ1179	Ne1-CH	Colombia-Ne	–	–	MK477927	MK477804	This study
I. badionotus	INV TEJ1184	Ne6-CH	Colombia-Ne	8J	–	MK477928	MK477805	This study
I. badionotus	INV TEJ1193	Ne15-CH	Colombia-Ne	8K	–	–	MK477806	This study
I. badionotus	INV TEJ1194	Ne16-CH	Colombia-Ne	–	–	–	MK477807	This study
I. badionotus	INV TEJ1196	Ne18-CH	Colombia-Ne	8L	–	MK477929	MK477808	This study
I. badionotus	INV TEJ1267	GV93-CH	Colombia-LG	8M	–	MK477930	MK477809	This study
I. badionotus	INV TEJ1268	GV100-CH	Colombia-LG	8N	–	MK477931	MK477810	This study
I. badionotus	INV TEJ1292	GV155-U	Colombia-LG	8Y	–	–	MK477811	This study
I. badionotus	INV TEJ1294	GV157-R	Colombia-LG	8J’	–	MK477932	MK477812	This study
I. badionotus	INV TEJ1301	GV236-R	Colombia-LG	8K’	–	MK477933	MK477813	This study
I. badionotus	INV TEJ1306	GV250-CH	Colombia-LG	8O	–	MK477934	MK477814	This study
I. badionotus	INV TEJ1310	GV300-CH	Colombia-LG	8P	–	MK477935	MK477815	This study
I. badionotus	INV TEJ1125	Ma10-CH	Colombia-LG	1C–D	–	–	MK477817	This study
I. badionotus	INV EQU3046	EQU3046-CH	Colombia-LG	8Q	–	–	MK477816	This study
I. badionotus	INV EQU3089	EQU3089-R	Colombia-SM	BECSC49-20	–	–	OL989879	This study
I. badionotus	INV TEJ2052	Pr29-R	Colombia-Prov	BECSC71-20	OL989773	–	–	This study
I. badionotus	INV TEJ2056	Pr38-R	Colombia-Prov	BECSC72-20	OL989774	–	–	This study
I. badionotus		Cu1-BY	Curaçao-Cu	8M’	–	MK477940	MK477821	This study
I. badionotus		Cu2-U	Curaçao-Cu	8Z	–	MK477941	MK477822	This study
I. badionotus		Br1	Brazil-SC	–	–	MK477936	MK477818	This study
I. badionotus		Br6	Brazil-SC	–	–	MK477937	MK477819	This study
I. badionotus		Br9	Brazil-SC	–	–	MK477938	–	This study
I. badionotus		Br15	Brazil-SC	–	–	MK477939	MK477820	This study
I. maculatus phoenius	USNM 1659468	BT5-LSD	Panama-BT	17T	–	–	MK477823	This study
I. maculatus phoenius	MBMLP IpBT7	BT7-DW	Panama-BT	–	MK477896	MK477942	MK477824	This study
I. maculatus phoenius	USNM 1659473	BT21-DW	Panama-BT	–	MK477897	–	MK477825	This study
I. maculatus phoenius	USNM 1659474	BT22-DW	Panama-BT	17O	MK477898	MK477943	MK477826	This study
I. maculatus phoenius	USNM 1659475	BT23-DW	Panama-BT	17G	MK477899	–	MK477827	This study
I. maculatus phoenius	USNM 1659476	BT25-DW	Panama-BT	17F	MK477900	MK477944	MK477828	This study
I. maculatus phoenius	USNM 1659477	BT43-DW	Panama-BT	1ST/18A	MK477901	MK477945	MK477829	This study
I. maculatus phoenius	USNM 1659478	BT44-DW	Panama-BT	17E/18E	MK477902	MK477946	MK477830	This study
I. maculatus phoenius	USNM 1659479	BT53-LSD	Panama-BT	17A’	MK477903	MK477947	MK477831	This study
I. maculatus phoenius	USNM 1659469	BT131-DW	Panama-BT	17N/18H	MK477904	MK477948	MK477832	This study
I. maculatus phoenius	USNM 1659470	BT190-DW	Panama-BT	18L	MK477905	MK477949	MK477833	This study
I. maculatus phoenius	USNM 1659471	BT191-DW	Panama-BT	17M	MK477906	–	MK477834	This study
I. maculatus phoenius	USNM 1659472	BT192-DW	Panama-BT	17R/18I	MK477907	MK477950	MK477835	This study
I. maculatus phoenius	TpGa0008	Ga8-U	Panama-Ga	1K	–	MK477951	MK477836	This study
I. maculatus phoenius	TpGa0011	Ga11-LBD	Panama-Ga	17U	–	MK477952	MK477837	This study
I. maculatus phoenius	TpGa0209	Ga209-LSD	Panama-Ga	–	–	MK477953	MK477838	This study
I. maculatus phoenius	USNM 1659480	Ga223-LSD	Panama-Ga	17S	–	MK477954	MK477839	This study
I. maculatus phoenius	USNM 1659481	Ga224-DW	Panama-Ga	17I	–	MK477955	MK477840	This study
I. maculatus phoenius	INV EQU4144 INV TEJ1333	BA313-LBD	Colombia-BA	17V	–	MK477956	MK477841	This study
I. maculatus phoenius	INV EQU4146 INV TEJ1334	BA314-LBD	Colombia-BA	17W	–	MK477957	MK477842	This study
I. maculatus phoenius	INV TEJ1336	BA319-U	Colombia-BA	1L	–	MK477958	MK477843	This study
I. maculatus phoenius	INV TEJ1257	GV82-DW	Colombia-LG	17J	–	–	MK477844	This study
I. maculatus phoenius	INV TEJ1258	GV83-DW	Colombia-LG	17K	–	MK477959	MK477845	This study
I. maculatus phoenius	INV TEJ1259	GV84-DW	Colombia-LG	17P	–	MK477960	MK477846	This study
I. maculatus phoenius	INV TEJ1261	GV86-DW	Colombia-LG	17L	–	MK477961	MK477847	This study
I. maculatus phoenius	INV TEJ1265	GV90-LSD	Colombia-LG	17X	–	MK477962	MK477848	This study
I. maculatus phoenius	INV TEJ1269	GV108-DW	Colombia-LG	17Q	–	MK477963	MK477849	This study
I. maculatus phoenius	INV TEJ1272	GV117-LSD	Colombia-LG	17Y	–	MK477964	MK477850	This study
I. maculatus phoenius	INV TEJ1113	GCV1-LBD	Colombia-LG	–	–	–	MK477851	This study
I. maculatus phoenius	INV TEJ1115	GCV3-U	Colombia-LG	17C	–	–	MK477852	This study
I. maculatus maculatus		CV1-DW	Cape Verde-SVI	14A	–	MK477965	MK477853	This study
I. maculatus maculatus		CV7-LSD	Cape Verde-SI	14G	–	MK477971	MK477859	This study
I. maculatus maculatus		CV10-LSD	Cape Verde-SI	14E	–	MK477974	MK477862	This study
I. maculatus maculatus		ASI1-LSD	Ascension Is.-EB	–	–	MK477975	MK477863	This study
I. maculatus maculatus		ASI2-LSD	Ascension Is.-EB	–	–	MK477976	MK477864	This study
I. maculatus maculatus		ASI4-LSD	Ascension Is.-EB	–	–	MK477978	MK477866	This study
I. maculatus maculatus	USNM E16150	E16150-LSD	Ascension Is.-EB	12A	–	MK477980	MK477867	This study
I. maculatus maculatus	UF 16295	UF16295-LSD	Senegal-DÎM	14L	OM056886	–	–	Florida Museum, unpubl.
I. maculatus maculatus		FAOLIB08-LSD	Liberia-K	14N–O	MZ742160	–	MZ743890	González-Wangüemert et al. in prep.
I. maculatus maculatus		FAOLIB09-LSD	Liberia-K	14N–O	MZ742161	–	MZ743891	González-Wangüemert et al. in prep.
I. maculatus maculatus		FAOLIB10-LSD	Liberia-AB	14N–O	MZ742162	–	MZ743892	González-Wangüemert et al. in prep.
I. fuscus	TfTa0210	Ta210-R	Panama-Ta	21A	MK477908	MK477981	MK477869	This study
I. fuscus	USNM 1659483	Ta211-R	Panama-Ta	1L’–M’	–	MK477982	MK477870	This study
I. fuscus	USNM 1659482	IP322-R	Panamá-IP	21B	–	MK477983	MK477871	This study
I. fuscus	INV EQU4323	EQU4323-R	Colombia-Ch	21D–E	–	MK477984	MK477872	This study
I. fuscus	INV EQU4324	EQU4324-BR	Colombia-Ch	21D–E	–	MK477985	MK477873	This study
I. fuscus	INV EQU2746	EQU2746-R	Colombia-Ch	21C	–	–	MK477875	This study
I. fuscus	INV TEJ1960	Ri54C1-R	Colombia-Ch	–	–	MK477986	MK477874	This study
I. fuscus	INV TEJ1923	TEJ1923-R	Colombia- Ch	BECSC75-20	OL989775	–	–	This study
I. fuscus	INV EQU4229	EQU4229	Colombia- Ch	21M, BECSC52-20	–	–	OL989880	This study
I. fuscus	INV EQU4230	EQU4230-BU	Colombia- Ch	BECSC53-20	–	–	OL989881	This study
I. fuscus	UF 9112	Ga UF 9112	Ecuador-Ga		PP093756			Florida Museum, unpubl.
I. fuscus	UF 9651	Ga UF 9651	Ecuador-Ga		PP093755			Florida Museum, unpubl.
I. fuscus	UF 9672	Ga UF 9672	Ecuador-Ga		PP093757			Florida Museum, unpubl.
I. macroparentheses	USNM E40833	E40833	U.S. Virgin Islands-SCI	24G	–	MK477987	MK477876	This study
I. macroparentheses	USNM E47624	E47524	British Virgin Islands-GI	1Q’/24F	–	MK477988	MK477877	This study
I. macroparentheses	USNM 1659484	Be99	Belize-TC	1P’/24A–B	–	MK477989	MK477878	This study
I. macroparentheses	USNM 1659485	Be107	Belize-CBC	24C–D	–	MK477990	MK477879	This study
I. macroparentheses	ICMIL-UNAM 12000	MeUNAMYUC76	Mexico-Yuc		PP093693	–	–	Florida Museum-ICMIL-UNAM, unpubl
I. macroparentheses	UF20352	BCUR-0532	Curaçao-CMRS	24E	PP093694			Florida Museum, unpubl.
I. badionotus		UF4703	USA-Florida-GM, Off St. Petersburg		(KX874354)			Miller et al. (2017)
I. maculatus phoenius * (I. badionotus)		MHE133			(JN207564)		(JN207495)	Honey-Escandon et al. (2012)
I. maculatus phoenius * (Isostichopus sp.)		Isolate IF-2009			(FJ971400)		(FJ794474)	Wen et al. (2011)
I. maculatus phoenius * (Isostichopus sp.)		Isolate 1			(KX384008)		(KX383967)	Vergara et al. (2018)
I. maculatus phoenius * (Isostichopus sp.)		Isolate A5			(KX384001)		(KX383978)	Vergara et al. (2018)
I. maculatus phoenius * (Isostichopus sp.)		Isolate R3			(KX383999)		(KX383985)	Vergara et al. (2018)
I. maculatus phoenius * (Isostichopus sp.)		Isolate SPA2			(KX383995)		(KX383988)	Vergara et al. (2018)
I. badionotus		FAO 044			(EU848276)		(EU822435)	Uticke et al. (2010)
I. badionotus		Isolate 1H			(KX384010)		(KX383968)	Vergara et al. (2018)
I. badionotus		Isolate 3H			(KX384016)		(KX383972)	Vergara et al. (2018)
I. badionotus		Isolate Br2			(KX384011)		(KX393982)	Vergara et al. (2018)
I. fuscus		Isolate If1			(AF486424)			Uticke et al. (2010)
I. fuscus		Isolate If15			(AF486425)			Uticke et al. (2010)
Astichopus multifidus		AmuBT18	Panama-BT				MK477880	This study
A. multifidus		FAO 042			(EU848293)			Uticke et al. (2010)
A. multifidus * (I. macroparentheses)		AMNH A113281					(AY338415)	Kerr et al. (2005)
Stichopus horrens		y11			(NC014454)	(NC014454)	(NC014454)	Fan et al. (2011)
Apostichopus japonicus		clone 1			(NC012616)	(NC012616)		Sun et al. (2010)
A. japonicus		clone 1					(AY852278)	Sun et al. (2010)