Supplementary Material Environmental influences on the Indo-Pacific octocoral Isis hippuris Linnaeus 1758: genetic fixation or capacity for plasticity?




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Supplementary Material
Environmental influences on the Indo-Pacific octocoral Isis hippuris Linnaeus 1758: genetic fixation or capacity for plasticity?

Sonia J. Rowley • Xavier Pochon • Les Watling


Supplementary material contents:

Systematic Summary: Isis Linnaeus 1758 (p. 2-7)

Supplementary Figure S1 (p. 3)

Supplementary Table S2 (p. 8)

Supplementary Figure S3 (p. 9)

Supplementary Table S4 (p. 10)

Supplementary Table S5 (p. 11)

Supplementary References (p. 12-13)


Systematic Summary: Isis Linnaeus 1758

Sub-Class OCTOCORALLIA

Order ALCYONACEA Lamouroux 1812

Sub-Order CALCAXONIA Grasshoff 1999

Family ISIDIDAE Lamouroux 1812

Sub-Family ISIDINAE Lamouroux 1812



Isis hippuris Linnaeus 1758

(Figure S1)



See Bayer & Stefani, 1987 for list of references [p. 55]

Type Material – Unfound, however ‘authentic’ specimens were collected and fully defined from Amboina, Indonesia (Milne-Edwards & Haime, 1857).

Diagnosis These arborescent colonies can be up to 1 m tall, planar or bushy with lateral or partially dichotomous branching but rarely anastomosing (net-like). Branch formations may also give a candelabrum appearance. The axis consists of alternating calcareous internodes that reduces to a fine rod through the non-scleritic and convex, dark proteinaceous (gorgonin) nodes, the former typically longer than the latter. Branching is internodal in single or multiple planes, the latter giving rise to the bushy appearance. Branch lengths and diameters are variable, and up to three short branches can arise per internode with some so close they appear nodal in highly branched colonies. The expanded calcareous cup-shaped base obliterates any trace of nodal composition particularly in older colonies. Calcareous internodes are sclerobastic (consisting of fused sclerites; but also see Nutting, 1910) with fibres arranged radially from a central core, akin to that of the sclerites bearing close resemblance to the Scleraxonia (Bayer, 1955). Internodes possess longitudinal grooves corresponding with 8 – 12 sinuous water vascular canals of ~1 mm diameter. Polyps distributed all around the branches are 0.5 – 1.25 mm apart and fully retractile to ~1.25 mm deep (and wide) into the thick coenenchyme. Such polyps bear eight lanceolate pinnate tentacles, and can possess up to three large (max. 1 mm diameter) round eggs, likely explaining the often swollen appearance of the branch tips as opposed to being a diagnostic feature (but see Nutting, 1910; Mai-Bao-Thu & Domante, 1971).



Figure S1. Isis Linnaeus, 1758 comparisons of (A) Isis hippuris Linnaeus, 1758 colony in Ellis & Solander, 1786; (B) sclerites of: i. I. hippuris and ii. Isis reticulata in Nutting 1910; (C) I. reticulata in Nutting, 1910; (D) Isis minorbrachyblasta Zou et al., 1991 colony and (E) sclerites. Note, images sourced from each citation respectively.

A diverse range of sclerites exists within the thick coenenchyme (Figure S3B, E). The surface layer consists of small warty clubs 0.08 x 0.001 mm [note: typographical error pg. 55, 2nd paragraph, Bayer & Stefani, 1987] typically bearing three large warts below the head wart (Figure S3B.i). Throughout the sub-surface layer some or all of the following are present in varying dimensions, asymmetry and commonly girdled: 6-, 7-, or 8-radiate capstans up to 0.19 mm in length, dumbbells/double heads (considered derivatives of 6-radiates; Bayer & Stefani, 1987) up to 0.32 mm long, warty or tuberculate spindles up to 0.25 mm long, and crosses. Very small e.g. rods of 0.07 x 0.01 mm (Bayer & Stefani, 1987), or no sclerites may be present within the polyp structures (Kölliker 1865). However smaller forms from those found within the coenenchyme (Simpson, 1906; Thomson & Simpson, 1909; Kükenthal, 1919; 1924) as well as small warty clubs with short handles ~0.055 x 0.045 mm located within the tentacles have been reported (Simpson, 1906; Thomson & Simpson, 1909).

Sclerites colourless and colonies typically light brown to mustard yellow, with slightly darker polyps.



Distribution – Central Indo-Pacific including Great Barrier Reef, Vanuatu, Papua New Guinea, Indonesia, Malaysia, Andaman, Philippines, Taiwan, Palau, South China Sea, Japan including Okinawa and the Ryukyu Islands.

Remarks – As evident from the description above, substantial phenotypic plasticity, from colony and branching structure to sclerite composition, exists in described specimens of I. hippuris (Wright & Studer, 1889; Simpson, 1906; Thomson & Simpson, 1909; Bayer & Stefani, 1987; Fabricius & Alderslade, 2001). Whether such plasticity is a consequence of environmental influence within and between its distributions, or significantly structured to be more than one species is unclear. Thus, for such a ubiquitous and well-known species it “has been very imperfectly described” (Thomson & Simpson, 1909) leading to “a slender basis on which to raise a superstructure of classification” (Wright & Studer, 1889). Clearly a revision of the Isis genus is required including thorough analyses of specimens throughout its geographic range, as such character trait variability may be ecologically dependent (Bayer & Stefani, 1987). Attempts, however, have been made to differentiate phenotypic patterns within the Isis genus that, even though somewhat tenuous, may equate to the morphotypes found within the WMNP (Rowley, 2014). Isis reticulata Nutting, 1910 and Isis minorbrachyblasta Zou, Huang & Wang, 1991 are therefore summarized below highlighting differences between the selected taxa.

Isis reticulata Nutting 1910

(Figure S1B)

See Mai-Bao-Thu & Domantay (1971) for list of references [p. 28]

Type Material – Syntypes: several fragments of varying sizes, ZMA COEL. no. 2721, Siboga Expedition, station 149 or 273 at Pulu Jedan, Aru Islands, Maluku, Indonesia, 13 meters on sand and shells. Fragment donated to State University of Iowa (van Soest, 1979). Specimens not located on request.

Diagnosis Slender colonies, typically arborescent with long slim terminal branches that are not swollen at the ends. Few very small polyps irregularly distributed around the branches, the latter occasionally anastomosing. Sclerites of the coenenchyme bear sharp rough warts symmetrically distributed around delicate spindles and clubs the latter 0.04 – 0.06 mm in length. Some spindles curved possessing large tubercules. Irregular radiates 0.06 x 0.03 mm to 0.2 x 0.1 mm in length and width respectively, smooth warty rods 0.1 - 0.15 mm long and occasional crosses present. No polyp sclerites reported.

Sclerites colourless, colony reddish brown with slightly darker polyps in alcohol. Also noted as “brownish white” by Mai-Bao-Thu & Domantay (1971).



Distribution I. reticulata has been documented in Indonesia (fragments from a single location 13 m depth), the Philippines (2 specimens and some fragments from a single location 12 - 15 m depth; Mai-Bao-Thu & Domantay, 1971), and Xisha Islands of China (single specimen and location; Zou, et al., 1991).

Remarks I. reticulata is thus differentiated from I. hippuris on the basis of planar versus bushy colonies, long thin sinuous branches without swollen ends versus short thick antler-like branches with swollen ends, and all sclerites of a smaller size with sharp rough warts in I. reticulata. Sclerite differences between I. hippuris and I. reticulata have been considered questionable owing to the huge diversity in form (Stiasny, 1940; Bayer & Stefani, 1987). However, Nutting (1910) observed smaller and more sharply warted sclerites further corroborated by Kükenthal (1924) and Mai-Bao-Thu & Domantay (1971), with illustrations showing marked asymmetry (Figure S1B.ii) contrary to that described. Curiously, Nutting (1910) noted I. reticulata having flaccid polyps if preserved when extended due to their lack of sclerites. However, in I. hippuris, polyp sclerites were “not being evident on account of their small size” (Nutting, 1910) with no further discussion, lending question to their presence at all (Simpson, 1906; Simpson & Thomson, 1909; Fabricius & Alderslade, 2001; but see Bayer & Stefani, 1987; Kölliker, 1865). Conflicting sclerite images between Nutting (1910) and Mai-Bao-Thu & Domantay (1971), in addition to regional differences between specimens of I. hippuris (see Bayer & Stefani, 1987) having some adherence to I. reticulata, lends further question to its validity as a taxon. Finally, colony and polyp colouration may be an artifact of preservation; Nutting’s ‘pink’ likely from buffered formalin used at that time (note: SJ Rowley, examined a ‘pink’ specimen from the Siboga expedition at the British Natural History Museum [BNHM. 1889.6.28.18], which adhered closely to the I. hippuris description above and not the proposed I. reticulata), and Mai-Bao-Thu and Domantay’s ‘white’ from endosymbiotic bleaching not uncommon with, in particular, damaged Isis specimens (e.g., Thomson & Simpson, 1909). In summary, the distinction between I. hippuris and I. reticulata is conflicting and unclear, requiring further investigation.

Isis minorbrachyblasta Zou, Huang & Wang 1991

(Figure S1C)



Type Material – Holotype (G85-001) and paratype (G87-031) from two locations of the Nansha Islands, China.

Diagnosis Colonies bushy with distal branches densely aggregated, themselves bearing tufts of branchlets no longer than 5 cm (ave. 3.5 cm). The short, fine branches arise from the scleritic internodes. Tiny polyps are equally distributed around the branches. Coenenchyme sclerites up to 0.140 x 0.091 mm being predominantly dumbbells and double heads with tubercules generally symmetrically arranged. Assortment of small clubs also present 0.06 x 0.025 mm with occasional crosses.

Sclerites colourless, colonies light brown in alcohol.



Distribution – Nansha Islands, China.

Remarks – Zou et al., (1991) state that the bushy non-planar colonies of I. minorbrachyblasta differ from the planar ones of I. hippuris and I. reticulata, in direct contrast to previous reports (e.g., Nutting, 1910; Mai-Bao-Thu & Domantay, 1971). Furthermore, the branches of I. minorbrachyblasta are fine, short and densely packed, whereby I. hippuris and I. reticulata are thick, short, dense, and fine, long, anastomosing and loosely packed respectively; thus I. minorbrachyblasta an intermediate between the two. Statistical significance between select morphological traits (branchlet and sclerite length and width) revealed differences among taxa were between I. minorbrachyblasta and I. reticulata. However, it is unclear what sclerites were used for comparative analyses, and n = 1 in all cases. Based on the information presented here, any appreciable difference in colony, branch and sclerite composition especially between I. minorbrachyblasta and I. hippuris (e.g. Nutting, 1910; Bayer & Stefani, 1987) is nebulous. Finally, Zou et al., (1991) propose I. minorbrachyblasta based on one or two specimens per taxon, somewhat unsatisfactory given both the nature of Isis phenotypic variability and analyses taken from a single region.

Table S2. Isis hippuris morphological traits summary table. All values expressed as metric or counts (± SE). Asterisk (*) indicate significantly (< P 0.05) informative traits selected for multivariate analyses. † Depicts low sample size.

Morphological Trait

Measures/Counts

Dimensions (± SE)

Morphological Trait

Measures/Counts

Dimensions (± SE)

Ridge 1

Sampela

Ridge 1

Sampela

Macromorphology (cm)










Micromorphology (mm)










Colony










Polyp










*H

Colony Height

48

49.74 ± 3.99

58.78 ± 3.10

*PD

Polyp Density

31,761

88.23 ± 2.6

100.21 ± 5.32

*W

Colony Mean Width

48

39.65 ± 2.24

58.03 ± 2.94

Pd

Polyp Depth

398

0.08 ± 0.001

0.08 ± 0.002

*w1

width 1

144

40.53 ± 3.24

65.31 ± 5.35

pD

Polyp Diameter

1920

0.04 ± 0.002

0.03 ± 0.001

*w2

width 2

144

57.02 ± 3.06

76.74 ± 3.6

ID

Inter-polyp Distance

1920

0.05 ± 0.002

0.05 ± 0.001

*w3

width 3

144

22.0 ± 1.35

31.86 ± 1.93

Cd

Canal diameter

1,920

0.02 ± 0.001

0.02 ± 0.002

*CS

Colony overhead Spread

48

45.5 ± 5.10

78.86 ± 3.10

C#

Canal#

1,677

8.17 ± 0.14

8.33 ± 0.21

*cs1

colony spread 1

48

52.06 ± 5.34

78.86 ± 3.92













*cs2

colony spread 2

48

38.94 ± 4.85

62.12 ± 3.99

Sclerites










B

Colony Base Width

18

3.47 ± 0.40

5.43 ± 0.64

*CL1

Club Length 1

960

0.073 ± 0.001

0.068 ± 0.001

*M

Colony Mid Branch Width

48

1.10 ± 0.08

1.53 ± 0.48

*CW1

Club Mean Width 1

48

0.021 ± 0.000

0.020 ± 0.000

*TW

Colony Tip Branch Width

960

0.63 ± 0.02

0.95 ± 0.05

c1w1

c1width 1

960

0.022 ± 0.001

0.021 ± 0.001
















c1w2

c1width 2

960

0.012 ± 0.000

0.011 ± 0.001

sub Colony










c1w3

c1width 3

960

0.030 ± 0.002

0.028 ± 0.001

*sH

sHeight

48

10.8 ± 0.33

11.55 ± 0.24

*CL2

Club Length 2

960

0.072 ± 0.000

0.068 ± 0.002

*sW

sMean Width

48

3.15 ± 0.21

2.66 ± 0.17

*CW2

Club Mean Width 2

48

0.033 ± 0.000

0.031 ± 0.001

sw1

swidth 1

144

2.50 ± 0.14

2.46 ± 0.15

c2w1

c2width 1

960

0.032 ± 0.001

0.030 ± 0.001

sw2

swidth 2

144

4.18 ± 0.28

3.37 ± 0.22

c2w2

c2width 2

960

0.016 ± 0.000

0.016 ± 0.001

sw3

swidth 3

144

2.76 ± 0.32

1.94 ± 0.23

c2w3

c2width 3

960

0.051 ± 0.001

0.047 ± 0.002

sML

sMean Mother Length

48

8.41 ± 0.33

9.77 ± 0.35

*CaL

Capstan Length

960

0.114 ± 0.004

0.101 ± 0.002

*sMW

sMean Mother Width

48

0.44 ± 0.02

0.55 ± 0.02

*CaW

Capstan Mean Width

48

0.075 ± 0.001

0.066 ± 0.001

mw1

mwidth 1

144

0.51 ± 0.01

0.62 ± 0.03

*caw1

cawidth 1

960

0.086 ± 0.004

0.076 ± 0.003

mw2

mwidth 2

144

0.48 ± 0.01

0.57 ± 0.02

*caw2

cawidth 2

960

0.041 ± 0.003

0.036 ± 0.020

mw3

mwidth 3

144

0.42 ± 0.01

0.54 ± 0.02

*caw3

cawidth 3

960

0.090 ± 0.003

0.080 ± 0.002

*sDL

sMean Daughter Branch Length

48

3.08 ± 0.21

4.44 ± 0.18

SL

Spindle Length

960

0.164 ± 0.003

0.162 ± 0.003

*sDW

sMean Daughter Branch Width

48

0.42 ± 0.01

0.48 ± 0.01

*SW

Spindle Mean Width

48

0.069 ± 0.001

0.065 ± 0.001

dw1

dwidth 1

480

0.44 ± 0.01

0.51 ± 0.01

sw1

swidth 1

960

0.038 ± 0.002

0.037 ± 0.002

dw2

dwidth 2

480

0.41 ± 0.01

0.49 ± 0.01

sw2

swidth 2

960

0.080 ± 0.003

0.076 ± 0.002

dw3

dwidth 3

480

0.38 ± 0.01

0.45 ± 0.01

*sw3

swidth 3

960

0.081 ± 0.003

0.077 ± 0.002

*MBW

Mean Branch Width

48

0.43 ± 0.01

0.52 ± 0.01
















*sTB#

sTotal Branch#

48

13.5 ± 1.10

10.83 ± 1.42

Total

59,328







sTBL

sTotal Branch Length

48

51.86 ± 3.49

52.57 ± 3.76
















*PA

Projected sub-colony Area

48

34.28 ± 2.65

30.79 ± 2.03

 




 

 

 

*PBA

Projected Branch Area

48

22.41 ± 1.69

26.96 ± 2.06
















*Po

Porosity

48

1.61 ± 0.10

1.19 ± 0.06














































Figure S3. Scanning electron micrographs showing sclerite diversity of Isis hippuris from (A - B) Ridge 1 and (C - F) Sampela within the WMNP. Inner coenenchyme spindles (A, C, D), surface capstans and clubs (B, E, F). Small rods at the end of both (A & F).
Table S4. ITS2 Accessions of octocoral outgroups used in the analyses.


Taxon

GenBank

Reference

[Group: Alcyoniinans]







Family: Alcyoniidae Lamouroux, 1812







Alcyonium digitatum Linnaeus, 1758

AF262347

McFadden et al., 2001

[Group: Scleraxonians]







Family: Coralliidae Lamouroux, 1812







Corallium rubrum Linnaeus, 1758

AF413059

Constantini et al., 2003

Corallium sp. 1

GQ358526

Herrera et al., 2010

Family: Paragorgiidae Kükenthal, 1916







Paragorgia kaupeka Sánchez, 2005

GQ293292

Herrera et al., 2010

Sibogagorgia cauliflora Herrera, Baco & Sánchez, 2010

GQ293288

Herrera et al., 2010

[Suborder: Holaxonians]







Family: Gorgoniidae Lamouroux, 1812







Africagorgia schoutedeni Stiasny, 1939

AY587533

Aguilar & Sánchez, 2007a

Gorgonia flabellum Linnaeus, 1758

AY587521

Aguilar & Sánchez, 2007a

Leptogorgia violacea Pallas, 1766

AY587527

Aguilar & Sánchez, 2007a

Lophogorgia [Synonym of Leptogorgia] euryale Bayer, 1952

AY587530

Aguilar & Sánchez, 2007a

Pacifigorgia stenobrochis Valenciennes, 1846

AY587531

Aguilar & Sánchez, 2007a

Pinnigorgia platysoma Nutting, 1910

AY587536

Aguilar & Sánchez, 2007a

Pseudopterogorgia [Synonym of Antillogorgia] bipinnata Verrill, 1864

AY587524

Aguilar & Sánchez, 2007a

Family: Plexauridae Gray, 1859







Eunicea tourneforti Milne Edwards & Haime, 1857

EF490982

Grajales et al., 2007

Muriceopsis bayeri Sánchez, 2001

AY587538

Aguilar & Sánchez, 2007a

[Suborder: Calcaxonians]







Family: Isididae Lamouroux, 1812







Acanella weberi Nutting, 1910

FJ790943

Dueñas & Sánchez, 2009

Acanella sp.

FJ790921

Dueñas & Sánchez, 2009

Isidella tentaculum Etnoyer, 2008

FJ790944

Dueñas & Sánchez, 2009

Keratoisis zelandica Grant, 1976

FJ790939

Dueñas & Sánchez, 2009

Lepidisis olapa Muzik, 1978

FJ790908

Dueñas & Sánchez, 2009

Family: Primnoidae Milne Edwards, 1857







Calyptrophora japonica Gray, 1866

EF090735

Aguilar & Sánchez, 2007b




 

 


T
Position/Site

Position/Site
able S5.
Isis hippuris ITS2 haplotype sequence view from the seven test sites within the WMNP. Each sequence represents haplotypes (in parentheses) present in each sample per site and GenBank accessions. Colour codes depict gaps (lilac), transitions (red), and transversions (yellow).


S1 (A) KP265677

S2 (A) KP265684

S3 (A) KP265683

S4 (A) KP265682

S5 (A) KP265681

S6 (A) KP265680

S7 (A) KP265679

S8 (A) KP265678

SG1 (D) KP265676

SG2 (D) KP265675

R1 (E) KP265690

R2 (E) KP265685

R3 (B) KP265694

R4 (B) KP265695

R5 (B) KP265696

R6 (B) KP265697

R7 (B) KP265698

R8 (E) KP265689

PK1 (C) KP265692

PK2 (C) KP265691

K1 (E) KP265688

K2 (E) KP265687

B3 (B) KP265699

B4 (B) KP265700

BB1 (B) KP265701

BB2 (B) KP265702

B1 (C) KP265693

B2 (E) KP265686




Supplementary References

Aguilar C, Sánchez JA. 2007a. Phylogenetic hypotheses of gorgoniid octocorals according to ITS2 and their predicted RNA secondary structures. Molecular Phylogenetics and Evolution 43(3): 774-786.

Aguilar C, Sánchez JA. 2007b. Molecular morphometrics: contribution of ITS2 sequences and predicted RNA secondary structures to octocoral systematics. Bulletin Marine Sciences 81(3): 335-349.

Bayer FM. 1955. Contributions to the nomenclature, systematics, and morphology of the Octocorallia. Proceedings of the United States National Museum 105(3357): 207-220, pl.8.

Bayer FM, Stefani J. 1987. Isididae (Gorgonacea) de Nouvelle-Calédonie--Nouvelle clé des genres de la famille. Bulletin of the Museum of Natural History Nature Paris, (4 sér.) 9 (section A) No. 1:47-106, pls. 1-30.

Constantini F, Tinti F, Abbiati M. 2003. Sistematica molecolare e filogenesi di Corallium rubrum. Biologia Marina Mediterranea 10: 73–75.

Dueñas LF, Sánchez JA. 2009. Character lability in deep-sea bamboo corals (Octocorallia, Isididae, Keratoisidinae). Marine Ecology Progress Series 397: 11-23.

Fabricius KE, Alderslade P. 2001. Soft corals and sea fans: a comprehensive guide to the tropical shallow-water general of the Central-West Pacific, the Indian Ocean and the Red Sea. AIMS (AIDAB), Townsville. pp. 264.

Grajales A, Aguilar C, Sánchez JA. 2007. Phylogenetic reconstruction using secondary structures of Internal Transcribed Spacer 2 (ITS2, rDNA): finding the molecular and morphological gap in Caribbean gorgonian corals. BMC Evolutionary Biology 7: 90.

Herrera S, Baco A, Sánchez JA. 2010. Molecular systematics of the bubblegum coral genera (Paragorgiidae, Octocorallia) and description of a new deep-sea species. Molecular Phylogenetics and Evolution 55(1): 123-135.

Kölliker RA. 1865. Die Bindesubstanz der Coelenteraten. Icones histologicae oder Atlas der vergleichenden Gewebelehre. Leipzig, Germany, pp. 87-181.

Kükenthal W. 1915. Das System der Seefedern. Zoologischer Anzeiger 45(6): 284-287.

Kükenthal W. 1919. Gorgonaria. Wissenschaft. Ergebn. Deutsch. Tiefsee-Exspedition auf dem Dampfer Valdivia 1898–1899, Band 13, 946 pp.

Kükenthal W. 1924. Coelenterata: Gorgonaria. Das Tierreich 47. Berlin: Walter de Gruyter and Co. pp.478.

Linnaeus C. 1758. Systema naturae per regna tria naturae :secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (in Latin) (10th ed.). Stockholm: Laurentius Salvius.

Mai-Bao-Thu F, Domantay JS. 1971. Taxonomic studies of the Philippine gorgonaceans in the collections of the University of Santo Tomas, Manila (cont’d). Acta Manilana 7: 3-77.

McFadden CS, Donahue R, Hadland BK, Weston R. 2001. A molecular phylogenetic analysis of reproductive trait evolution in the soft coral genus Alcyonium. Evolution 55: 54-67.

Milne-Edwards H, Haime J. 1857. Histoire naturelle des coralliaires, ou polypes proprement dits. Paris, Roret. pp. 326.

Nutting CC. 1910. The Gorgonacea of the Siboga Expedition V. The Isidae. Siboga- Expeditie Monograph, 13b2, 1–24.

Rowley SJ. 2014. Gorgonian responses to environmental change on coral reefs in SE Sulawesi, Indonesia. Doctoral thesis, Victoria University Wellington, New Zealand, pp. 213.

Simpson JJ. 1906. The structure of Isis hippuris, Linnaeus. Journal of the Linnean Society of London, Zoology 29(194): 421-434.

Stiasny G. 1940. Biological results of the Snellius Expedition. VII. Die Gorgonarien Sammlung.

Thomson JA, Simpson JJ. 1909. An account of the alcyonarians collected by the Royal Indian Marine Survey Ship Investigator in the Indian Ocean; with a report on the species of Dendronephthya by Henderson WD II. The alcyonarians of the littoral area. The Indian Museum, Calcutta.

Wright EP, Studer TH. 1889. Report on the Alcyonaria. Rep. Scient. Results Explor. Voyage Challenger. 31(1): 1-314.

Zou R, Huang B, Wang X. 1991. Studies on the gorgonians of China – I. Isis with one new species. Acta Oceanologica Sinica 10(4): 593-602.



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