Taxonomic revision of Coralliidae with descriptions of new species from New Caledonia and the Hawaiian Archipelago

Figures & data

Table I. Morphological characters and states used for identification in Coralliidae.

Characters	States
Colonial level
Branching pattern	Branches given off on all sides of stem, or ramification in one plane
Axis colour	Deep red, orange, dark pink, light pink, milk white, yellowish white, or pure white
Axis pits	Absent, slightly sunken, raindrop-shaped, or round with prominent beaded margin
Axis grooves	Absent or present
Thickness of cortex	Thin < 0.1 mm, or thick > 0.15 mm
Cortex colour	Deep red, orange, dark pink, light pink, milk white, yellowish white, or pure white
Polyp level
Type of autozooid	Contractile, or retractile with cortical mounds
Autozooid distribution	Distributed on all sides of branches, or along one surface of branches
Cortical mounds	Absent, conspicuous, inconspicuous
Colour of cortical mounds or autozooids	Deep red, orange, dark pink, light pink, milk white, yellowish white, or pure white
Siphonozooid	Non-visible, or in form of small verrucae
Sclerites
Long spindles	Absent or present
Short rods near oral space	Absent or present
Multiradiates
Crosses	Absent or present
6-radiates	Absent, asymmetrical, or symmetrical
7-radiates	Absent, asymmetrical, or symmetrical
8-radiates	Absent, asymmetrical, or symmetrical
Double clubs	Absent, rough or smooth surface
Irregular forms	Absent or present

Table II. List of coralliid species as a result of the taxonomic revision.

Corallium japonicum Kishinouye, 1903	H. halmaheirense (Hickson, 1907)	P. clavatum sp. nov.
C. medea Bayer, 1964	H. imperiale (Bayer, 1955)	P. elatius (Ridley, 1882)
C. nix Bayer, 1996	H. laauense (Bayer, 1956)	P. gotoense (Nonaka, Muzik & Iwasaki, 2012)
C. rubrum (Linnaeus, 1758)	H. maderense (Johnson, 1898)	P. inutile (Kishinouye, 1902)
C. salomonense (Thomson and Mackinnon, 1910)	H. niobe (Bayer, 1964)	P. johnsoni (Gray, 1860)
C. stylasteroides Ridley, 1882	H. regale (Bayer, 1956)	P. konojoi (Kishinouye, 1903)
C. tortuosum Bayer, 1956	H. reginae (Hickson, 1907)	P. niveum (Bayer, 1956)
Hemicorallium abyssale (Bayer, 1956)	H. sulcatum (Kishinouye, 1903)	P. norfolkicum sp. nov.
H. aurantiacum sp. nov.	H. tricolor (Johnson, 1899)	P. occultum (Tu, Altuna & Jeng, 2015a)*
H. bathyrubrum (Simpson & Watling, 2011)	H. taiwanicum (Tu, Dai & Jeng, 2012)	P. porcellanum (Pasternak, 1981)
H. bayeri (Simpson & Watling, 2011)	H. variabile (Thomson & Henderson, 1906)	P. pusillum (Kishinouye, 1903)
H. boshuense (Kishinouye, 1903)	Pleurocorallium bonsaiarborum sp. nov.	P. secundum (Dana, 1846)
H. ducale (Bayer, 1955)	P. borneense (Bayer, 1950)	P. thrinax (Bayer & Stefani in Bayer, 1996)
H. guttatum sp. nov.	P. carusrubrum (Tu, Dai & Jeng, 2012)	P. uchidai (Nonaka, Muzik & Iwasaki, 2012)

*Indicates generic reassignment in the present study.

Figure 1. Corallium stylasteroides. USNM 77450: (A) Different sides of colony with enlargements of (a) cortex, (b) autozooids and (c) main stem; (B) SEM of cortex; (C) SEM of axis; (D) SEM of cortical mound and retracted autozooid. Black arrows in B indicate siphonozooids.

Figure 2. Corallium stylasteroides.USNM 77450: (A) Cortical sclerites from the tip of the colony, (a) 6-radiates, (b) 8-radiates, (c) double clubs; (B) cortical sclerites from the base of the colony, (a) 7-radiates; (C) sclerites from cortical mound and autozooid.

Figure 3. (A) Hemicorallium reginae. Syntype, NBC COEL_02395; (B) Hemicorallium halmaheirense. Syntype, NBC COEL_02393, large fragment, white arrow indicates polychaetes in the middle of the branch; (C) Hemicorallium halmaheirense. Syntype, NBC COEL_02393, small fragment and enlargement of autozooids.

Figure 4. Hemicorallium reginae. Syntype, NBC COEL_02395: (A) Cortical sclerites, (a) crosses, (b) asymmetrical 6-radiates, (c) 7-radiates, (d) symmetrical 8-radiates, (e) double club, (f) irregular forms; (B) sclerites from autozooid, (a) long spindles.

Figure 5. Hemicorallium halmaheirense. Syntype, NBC COEL_02393: (A) Cortical sclerites, (a) spherical 8-radiate; (B) sclerites from autozooid.

Figure 6. Hemicorallium imperiale. Holotype, USNM 50110: (A) Front, back and enlargement of colony; (B) cortical sclerites, (a) elongated 8-radiates reaching 0.1 mm in length.

Figure 7. Hemicorallium aurantiacum sp. nov. (A)–(D) Holotype, MNHN-IK-2011-1580: (A) Front of colony; (B) enlargement of frontal colony, white arrows indicate the thickened cortex induced by polychaetes; (C) and (D) SEM of autozooids, white arrow indicates the apical pore of a siphonozooid; (E) paratype, USNM 1196454, front of colony.

Figure 8. Hemicorallium aurantiacum sp. nov. Holotype, MNHN-IK-2011-1580: (A) Cortical sclerites, (a) 6-radiates, (b) criss-cross, (c) 7-radiates, (d) spherical 8-radiates; (B) sclerites from autozooid, (a) spherical 8-radiates, (b) 7-radiates, (c) spiny spindles, (d) long spindles.

Figure 9. Hemicorallium ducale. Holotype, USNM 50111: (A) Front of colony; (B) SEM image of branch; (C) sclerites from autozooids. Branch image by F.M. Bayer courtesy of Stephen Cairns.

Figure 10. Hemicorallium guttatum sp. nov. Holotype, USNM 1072452 (A)–(C): (A) Front and back of colony; (B) SEM of branch and enlargement of autozooid; (C) SEM of the inner structure of autozooid and enlargement of partial cross-section of tentacle; (D) paratype, USNM 1116818; white arrows indicate the thickened cortex induced by polychaetes.

Figure 11. Hemicorallium guttatum sp. nov. Holotype, USNM 1072452: (A) Sclerites from cortex, (a) spherical 8-radiates; (B) sclerites from autozooid, (a) spherical 8-radiates.

Figure 12. (A) Pleurocorallium porcellanum. Holotype, IORAS IV-9-Alc-11-009; (B) Pleurocorallium kishinouyei. Holotype, USNM 94462; (C) Pleurocorallium kishinouyei. Paratype, USNM 94463.

Figure 13. Pleurocorallium porcellanum. Holotype, IORAS IV-9-Alc-11-009: (A) Cortical sclerites; (B) sclerites from autozooid.

Table III. Main characters of Pleurocorallium porcellanum (Pasternak, 1981) and Pleurocorallium kishinouyei (Bayer, 1996).

Characters	P. porcellanum		P. kishinouyei
Colonial form	Planar		Planar
Polyp arrangement	Biserial		Biserial
Diameter of cortical mound	ca. 2 mm		ca. 1.96–2 mm
Sclerites	Size range*	Ratio	Size range*	Ratio
6-radiate	0.024–0.080	0.14	0.042–0.102	0.15
7-radiate	0.052–0.111	0.06	0.064–0.129	0.05
8-radiate	0.044–0.128	0.70	0.059–0.128	0.69
Double clubs	–	0	–	0
Cross	0.052–0.114	0.10	0.052–0.088	0.12
Colour
Cortex	Yellowish white		Yellowish white
Axis	White		White
Distribution	Mid-Pacific Seamounts southwest off the Hawaiian Islands		Cross Seamount, west southwest off the Hawaiian Islands
Depth	1213–1292 m		1145 m

*Units in mm. Ratio refers to the portion in the cortex. The ratio of irregular-type sclerites is not listed in the table.

Figure 14. Pleurocorallium bonsaiarborum sp. nov. Holotype, MNHN-IK-2009-942: (A) Different sides of colony; (B) cortical sclerites; (C) sclerites from autozooid.

Figure 15. Pleurocorallium clavatum sp. nov. Holotype, USNM 1195223: (A) Front and back sides of colony, and enlargement of branch; (B) SEM image of branch and cross-section.

Figure 16. Pleurocorallium clavatum sp. nov. Holotype, USNM 1195223: (A) Cortical sclerites, (a) 6-radiates similar to the radiation warning symbol, (b) double clubs with smooth surface; (B) sclerites from autozooid, (a) rods in pharyngeal region.

Figure 17. Pleurocorallium borneense. Holotype, USNM 49325: (A) Front and back views of colony, and enlargement of branch; (B) sclerites from cortex, (a) double clubs with smooth surface.

Figure 18. Pleurocorallium norfolkicum sp. nov. Holotype, MNHN-IK-2011-1467 (A)–(C): (A) Front and back sides of colony; (B) close-up of cortical mounds and autozooids, white and black arrows indicate cortical mounds and openings of siphonozooids, respectively; (C) deep autozooid pits with beaded margins; (D) paratype, MNHN-IK-2011-1706.

Figure 19. Pleurocorallium norfolkicum sp. nov. Holotype, MNHN-IK-2011-1467: (A) Cortical sclerites from the tip of the colony, (a) symmetrical 6-radiates, (b) asymmetrical 6-radiates, (c) 7-radiate, (d) mushroom-like double clubs, (e) regular double clubs; (B) cortical sclerites from the base of the colony, (a) 7-radiates; (C) sclerites from cortical mounds and autozooid, (a) spherical 6-radiates, (b) small rods in pharyngeal region.

Figure 20. Multiple factor analysis (MFA) scatterplot of 26 Hemicorallium specimens based on both qualitative and quantitative morphological features.

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