European Journal of Taxonomy 218: 1-52

http://dx.doi.org/10.5852/ejt.2016.218

www. europeanj ournaloftaxonomy. eu

2016 • Galea H R.

ISSN 2118-9773

This work is licensed under a Creative Commons Attribution 3.0 License.

Monograph

urn:lsid:zoobank.org:pub:A4D7AA38-D18F-4604-A5E0-D965637BD9F8

Notes on some sertulariid hydroids (Cnidaria: Hydrozoa) from the

tropical western Pacific, with descriptions of nine new species

Horia R. GALEA

Hydrozoan Research Laboratory, 405 Chemin des Gatiers, 83170 Tourves, France.

E-mail: horia.galea.@gmall .com

urn:lsid:zoobank.org:author:DE5AC672-0243-46F2-A910-AFF4E91A4C5D

Abstract. Forty-three species of sertulariid hydroids (Cnidaria: Hydrozoa: Sertulariidae), collected

from the tropical western Pacific (Taiwan, Philippines, New Caledonia, French Polynesia, Vanuatu,

Fiji, Tonga, Solomon Islands) during various expeditions of the French Tropical Deep-Sea Benthos

program, are discussed. Of these, nine are new to science: Gonaxia nova sp. nov., G. plumularioides

sp. nov., Sertularella folliformis sp. nov., Se. plicata sp. nov., Se. pseudocatena sp. nov., Se. splendida

sp. nov., Se. tronconica sp. nov., Se. tubulosa sp. nov., and Symplectoscyphus paucicatillus sp. nov.

The subspecies Symplectoscyphus johnstoni (Gray, 1843) tropicus Vervoort, 1993 is raised to species

but, in order to avoid the secondary homonymy with Sy. tropicus (Hartlaub, 1901), the replacement

name, Sy fasciculatus nom. nov., is introduced. The male and female gonothecae of Diphasia cristata

Billard, 1920, the male gonothecae of Gonaxia elegans Vervoort, 1993, as well as the female gonothecae

of Salacia macer Vervoort & Watson, 2003, are described for the first time. Additional notes on the

morphology of several other species are provided. All taxa are illustrated, in most cases using figures

drawn at the same scale, so as to highlight the differences between related species.

Key words. Deep water, new species, thecate hydroids, Sertulariidae.

Galea H.R. 2016. Notes on some sertulariid hydroids (Cnidaria: Hydrozoa) from the tropical western Pacific, with

descriptions of nine new species. European Journal of Taxonomy 218: 1-52. http://dx.doi.org/10.5852/eit.2016.218

Introduction

The present report is a continuation of an earlier study dealing with some new sertulariid and syntheciid

hydroids (Galea 2015 a), and completes a series of previous reports dealing with members of the families

Sertulariidae Lamouroux, 1812 (Vervoort 1993), Halopterididae Millard, 1962 (AnsinAgis et al. 2009),

Lafoeidae Hincks, 1868 (specifically, the genus Acryptolaria Norman, 1875) (Pena Cantero & Vervoort

2010), and Kirchenpaueriidae Stechow, 1921 (AnsinAgis et al. 2014) from the tropical western Pacific,

obtained during numerous expeditions undertaken conjointly by the Institut de Recherche pour le

Developpement (IRD, formerly ORSTOM) and the Museum national d'Histoire naturelle (MNHN) of

Paris (Bouchet et al. 2008).

The sertulariid hydroids dealt with herein belong to the following ten genera: Caminothujaria von

Campenhausen, 1896, Diphasia L. Agassiz, 1862, Dynamena Lamouroux, 1812, Geminella Billard,

1925b, Gonaxia Vervoort, 1993, Idiellana Cotton & Godfrey, 1942, Salacia Lamouroux, 1816,

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Sertularella Gray, 1848, Sertularia Linnaeus, 1758 and Symplectoscyphus Marktanner-Turneretscher,

1890. Although most species have already been dealt with by Vervoort (1993), additional notes on them

are provided, along with new data on the morphology of other taxa, including descriptions of nine new

hy droids.

Material and methods

The methods of study were described in detail by Galea (2007, 2008). The list of stations is given in

Table 1. Station numbers, as indicated in the text, are preceded by a two-letter prefix referring to the

sampling gear used to secure the material, either a Charcot dredge (CC), a beam trawl (CP), a rocky-

bottom dredge (DR) or a Warren dredge (DW). The material is deposited in the collections of MNHN.

The species dealt with herein are listed according to their morphological similarities, notably the shape

of their hydrothecae. Most figures, representing portions of colonies, hydro- and gonothecae, are drawn

to the same scale and grouped onto the same plates, so as to facilitate comparisons between related

species.

Results

Phylum Cnidaria Verrill, 1865

Class Hydrozoa Owen, 1843

Subclass Hydroidolina Collins & Marques, 2004

Order Leptothecata Cornelius, 1992

Family Sertulariidae Lamouroux, 1812

Genus Caminothujaria von Campenhausen, 1896

Caminothujaria molukkana von Campenhausen, 1896

Fig. 1A-B

Caminothujaria molukkana von Campenhausen, 1896: 103.

Caminothujaria molukkana - Schuchert 2015: 338, fig. 12.

Material examined

MUSORSTOM 3: Stn. DR117, numerous sterile fragments from an originally large colony (size

undeterminable) (MNHN-IK-2012-16500); Stn. CP121, a 2.3 cm high, sterile colony (MNHN-

IK-2012-16501); Stn. DR126, an originally 11.5 cm high colony, now broken into two parts, one

gonotheca (MNHN-IK-2012-16502).

MUSORSTOM 8: Stn. CP 1001, several stems and fragments up to 5.3 cm high, all sterile (MNHN-

IK-2012-16503); Stn. DW1021, numerous sterile stems and fragments, 3-10.5 cm high, all sterile

(MNHN-IK-2012-16504); Stn. CPI 104, an originally large but sterile colony, now fragmented, largest

fragment ca 8 cm high (MNHN-IK-2012-16505).

BORDAU2: Stn. CP 1582, ca 4 cm high, fertile colony composed of three stems (MNHN-IK-2012-16506).

SALOMON 1: Stn. DW1756, three sterile stems 2.5-5 cm high (MNHN-IK-2012-16507); Stn. DW1767,

a ca 12 cm high, sterile colony (MNHN-IK-2012-16508).

Remarks

A list of synonyms is provided by Schuchert (2015) and, for a re description, see Hirohito (1995).

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Table 1 . List of stations.

Campaign

DR117

CP121

DR126

CP134

CP34

DW37

DW38

DW66

CP69

DW46

CC174

DW205

DW221

DW423

DW40

DW48

DW55

DW57

DW39

DW127

DW716

CP737

CP804

CP806

DW809

DW810

CP813

CP814

CP821

CP823

DW829

DW830

DW902

CP906

DW914

DW923

CP1001

DW1019

DW1021

CP1083

CP1084

CP1104

CP 1131

DW1204

CP1265

CP 1447

CP1448

DW13

DW1513 1 Jun. 2000

CP1582 13 Jun. 2000

DW1595 16 Jun. 2000

DW1 27 Jul. 2000

DW1659 20Jun. 2001

DW1666 20Jun. 2001

DW1667 21 Jun. 2001

DW1699 24Jun. 2001

DW1704 25Jun. 2001

DW1712 26Jun. 2001

CP1719 26Jun. 2001

DW1722 26Jun. 2011

DW118 31 Jul. 2001

DW1741 23 Sep. 2001

DW1756 26 Sep. 2001

DW1767 28 Sep. 2001

DW1854 7 Oct. 2001

92-97

12°31' S

120°39' E

73-84

12°08' S

121°17'E

266

11°49' S

121°22'E

92-95

i2°or s

121 °57' E

710

23°11.88' S

167° 11.30' E

350

22°59.99' S

167°15.65 , E

360

22°59.74' S

167°15.31' E

505-515

24°55.43' S

168°21.67' E

1220-1225

23°51.38' S

167°58.68’E

570-610

22°53.05' S

167°17.08'E

385

19°00.30' S

163°18.50' E

160

22°38.50' S

167°06.80' E

535-560

22°58.60' S

167°36.80' E

280

20°25.85 f S

166°40.50'E

260

24°46.20' S

168°08.70'E

245

24°46.20 f S

168°08.70'E

260

23 o 21.40 , S

168°04.50' E

210

23°21.00' S

168°21.00' S

305

22°20.50' S

168°43.50'E

205-215

19°06' S

163°22'E

290-308

22°40' S

167°12'E

357-400

23°03.42' S

166°59.97' E

244-278

23°41.40' S

168°00.42'E

308-312

23°42.31' S

168°00.52'E

650-730

23°39.39 f S

167°58.94'E

850-900

23°40.45' S

167°58.83'E

410-415

23°45.23' S

168°16.57' E

444-530

23°47.60' S

168°17.10 , E

864-880

23°19.16' S

167°58.63'E

980-1000

23°22.76' S

167°51.60' E

386-390

23°21.37 f S

168°01.84' E

361-365

23°19.75' S

168°01.45'E

341-351

19°00.84 f S

163°14.83'E

339-350

19°01.07' S

163°14.5EE

600-616

18°48.79' S

163°15.23'E

502-470

18°51.51' S

163°24.17’ E

150-250

18°48.97' S

168°59.83' E

397-430

17°38.25' S

168°33.9E E

124-130

17°42.75' S

168°37.00' E

397-439

15°51.91' S

167°19.82'E

207-280

15°50.29' S

167°17.88'E

125-129

15°03.83' S

167 o 06.96'E

140-175

15°38.41' S

167°03.52'E

60-62

9°52.6' S

139°03.2' W

90-92

9°20.4' S

140°07.3' W

420-513

16°45.23' S

179°59.13'E

410-500

16°45.04' S

179°58.97'E

400

23°45’ S

168° 17' E

190-221

21°19' S

175°0E w

79-82

18°41' S

174°03' W

523-806

19°03'S

174°19 f W

23°38. F N

119°50.9'E

467-449

23°38 f S

167°42'E

469-860

23°42'S

167°43'E

237-250

23°41' S

168°0E E

581-600

24°40' S

168°39'E

420-400

23°47' S

168°17'E

180-250

23°22' S

168°03'E

391-407

23°21' S

168°02'E

540

23°16' S

168°01’E

128-136

24°56.3'N

122°01.5' E

557-655

11 °29 r S

159°57' E

497-511

08°52'S

159°50'E

98-200

08°19' S

160°40 f E

229-260

09°46' S

160°53 f E

Station

Date

Depth (m)

Latitude

Longitude

3 Jun. 1985

4 Jun. 1985

5 Jun. 1985

29 Aug. 1985

30 Aug. 1985

3 Sep. 1985

30 Aug. 1985

17 Sep. 1985

27 Sep. 1985

29 Sep. 1985

16 Feb. 1989

7 Mar. 1989

8 Mar. 1989

9 Mar. 1989

8 Jun. 1989

4 Mar. 1990

10 May 1993

13 May 1993

27 Nov. 1993

28 Nov. 1993

29 Nov. 1993

4 Aug. 1994

5 Aug. 1994

6 Aug. 1994

25 Sep. 1994

28 Sep. 1994

5 Oct. 1994

7 Oct. 1994

11 Oct. 1994

28 Aug. 1997

3 Sep. 1997

4 Mar. 1999

12 Aug. 1999

MUSORSTOM 3

BIOCAL

MUSORSTOM 4

MUSORSTOM 6

SMIB4

VOLSMAR

SMIB6

BATHUS 2

BATHUS 3

BATHUS 4

MUSORSTOM 8

MUSORSTOM 9

BORDAU1

LITHIST

BORDAU 2

TAIWAN 1

NORFOLK 1

TAIWAN 2

SALOMON 1

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Geographical distribution

The known area of distribution ranges from Indonesia and Tuvalu in the south, through Japan in the

north, and encompasses the Philippines and the South China Sea (Schuchert 2015).

Genus Diphasia L. Agassiz, 1862

Diphasia cristata Billard, 1920

Fig. 1C-J

Diphasia cristata Billard, 1920: 147, fig. If.

Diphasia cristata - Billard 1925b: 218, fig. 57, pi. 3 fig. 32. — Hirohito 1983: 38, fig. 15; 1995: 161,

fig. 51a-c.

Material examined

SMIB 4: Stn. DW57, several sterile stems and fragments, up to 2.5 cm high (MNHN-IK-2012-16509).

SMIB 6: Stn. DW127, a female colony composed of several stems up to 4 cm high, epizoic on Gonaxia

sp. (MNHN-IK-2012-16510).

MUSORSTOM 8: Stn. DW1021, small female colony composed of a few stems, up to 2 cm high,

epizoic on Caminothujana molukkana (MNHN-IK-2012-16511).

BORDAU 1: Stn. CP 1447, small female colony composed of several stems up to 2.4 cm high (MNHN-

IK-2012-16512); Stn. CP 1448, dense, fertile colony (or colonies?), up to 4 cm high, on dead antipatharian

(MNHN-IK-2012-16513).

BORDAU 2: Stn. DW1513, male and female colonies with stems up to 1.5 cm high, on antipatharian

and aglaopheniid hydroid (MNHN-IK-2012-16514).

Remarks

This species is immediately recognizable through the presence of five prominent, longitudinal, perisarcal

crests running over the whole body of hydrothecae, as well as of an additional one found on the dorsal

side of the intemodes. For the most recent description, see Hirohito (1995). It should be noted that

the oblique node demarcating the junction between the basal, ahydrothecate part and the main, distal,

hydrothecate part of the stems and side branches is not always present in the material examined here.

The gonothecae of this species, hitherto unknown, are described from the present material. Male and

female gonothecae present, each sex borne on different stems, but, due to their high density and the

numerous branches and anastomoses of the stolon, it is impossible to determine if it is a single colony

or whether several colonies are represented. The male gonothecae (Fig. II—J) are 1505-1580 pm long,

ovoid (480-530 pm wide in middle), and provided with numerous upturned spines (width including the

spines 1000-1170 pm). The aperture occurs on the apex of a short neck region and is surrounded by a

few, short spines. The female gonothecae (Fig. 1F-H) are comparatively larger (2985-3160 pm long),

have a rhomboidal cross-section (940-1010 pm wide), forming four crests at each angle; each one

is provided with a succession of moderately long, upturned spines (width of gonotheca including the

spines ca 1300 pm). A central, piriform chamber is attached to the walls of the gonotheca through four

perisarcal connections, whose position alternates with that of the crests. The aperture is situated apically,

and is small and quadrangular.

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Geographical distribution

Previously known from the Philippines (Billard 1920, 1925b) and Japan (Hirohito 1983, 1995). The

present material originates from New Caledonia, Vanuatu, Fiji, and Tonga.

Dipltasia orientalis Billard, 1920

Fig. IK

Diphasia orientalis Billard, 1920: 146, fig. Id.

Diphasia orientalis -Billard 1925b: 212, figs 52-53. — Gibbons & Ryland 1989: 407, fig. 26.

Fig. 1. — A-B. Caminothujaria molukkana von Campenhausen, 1896 (BORDAU 2, Stn. CP 1582).

A. Fragment of internode with verticil of hydrothecae. B. Gonotheca. — C-J. Diphasia cristata Billard,

1920 (BORDAU 1, Stn. CP1448). C. Pair of hydrothecae, frontal view. D. Same, dorsal view. E. Cross

section through basal part of hydrothecal pair, showing perisarcal crests. F. Female gonotheca. G. Same,

apical view. H. Same, cross section. I. Male gonotheca. J. Same, apical view. — K. Diphasia orientalis

Billard, 1920 (MUSORSTOM 3, Stn. DR117), internode with pair of hydrothecae. — L-O. Dynamena

heterodonta (Jarvis, 1922) (MUSORSTOM 9). L. Pair of hydrothecae in material from Stn. DW1204.

M. Same sample, hydrothecal aperture, showing internal, submarginal cusps. N-O. Isolated (N) or

stacked (O) pairs of hydrothecae in material from Stn. CP 1265. — P. Geminella ceramensis Billard,

1925a (MUSORSTOM 8, Stn. CP 1084), internode with pair of hydrothecae. Scale bars: A-D, K-L,

N-P = 500 pm; E = 400 pm; F-J = 1 mm ; M = 100 pm.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Material examined

MUSORSTOM 3: Stn. DR117, two infertile, unbranched stems, 9 and 12 mm high, detached from

substrate (MNHN-IK-2012-16515); Stn. CP121, two infertile, unbranched stems, 13 and 22 mm high,

detached from substrate (MNHN-IK-2012-16516).

Remarks

For a thorough description of this species, see Billard (1925b).

Geographical distribution

Indonesia (Billard 1925b), Fiji (Gibbons & Ryland 1989), Philippines (present report).

Genus Dynamena Lamouroux, 1812

Dynamena heterodonta (Jarvis, 1922)

Fig. 1L-0

Pasythea heterodonta Jarvis, 1922: 344, pi. 24, figs 11-12.

Dynamena heterodonta -Billard 1925b: 198, fig. 44. — Redier 1966: 86. — Vervoort & Vasseur 1977:

36, figs 16-17.

Sertularia gracilis - Billard 1905: 334 (not Sertularia gracilis FI assail, 1848: 2223).

non Dynamena quadridentata f. heterodonta - Vannucci 1951a: 83; 1951b: 108, 112, 114 [= D. quadri-

dentata (Ellis & Solander, 1786)].

Material examined

MUSORSTOM 9: Stn. DW1204, small colony composed of four infertile stems, 6-12 cm high, detached

from substrate (MNHN-IK-2012-16517); Stn. DW1265, sterile colony with stems up to 2.6 cm high,

epizoic on Synthecium sp. (MNHN-IK-2012-16518).

Remarks

The sterile, unbranched stems, divided here and there by oblique nodes, each of which comprises a varied

number of isolated hydrothecal pairs, occasionally forming apically stacked groups of two, the lack of

evidence of an abcauline caecum in retracted hydranths, the two-cusped hydrothecae with a pleated

adcauline flap and a semicircular abcauline flap, and the inconstant presence of internal, submarginal

cusps in the hydrothecae suggest that they belong, with little doubt, to Jarvis’ (1922) species.

Vervoort & Vasseur (1977) provided a reexamination of the type material, as well as a discussion

regarding its differences with/), quadridentata (Ellis & Solander, 1786).

Geographical distribution

St. Brandon (Cargados Carajos) (Jarvis 1922), Indonesia (Billard 1925b), New Caledonia (Redier 1966),

French Polynesia [Billard (1905), as Sertularia gracilis Hassall, 1848; Vervoort & Vasseur (1977);

present study].

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Genus Geminella Billard, 1925b

Geminella ceramensis (Billard, 1925a)

Fig. IP

Sertularella ceramensis Billard, 1925a: 649.

Geminella ceramensis - Schuchert 2003: 174, fig. 31.

Material examined

MUSORSTOM 8: Stn. CP 1084, small, sterile colony, less than 1 cm high, with tangled stems fixed on

calcareous substrate (MNHN-IK-2012-16519); Stn. CPI 131, two sterile stems 0.5 cm high, detached

from substrate (MNHN-IK-2012-16520).

Remarks

For a redescription of this distinctive species, see Schuchert (2003), who also provided a synonymy.

Geographical distribution

Indonesia, Philippines, New Caledonia (Schuchert 2003), and Vanuatu (present study).

Genus Gonaxia Vervoort, 1993

Gonaxia amphorifera Vervoort, 1993

Figs 2A, 3A-B

Gonaxia amphorifera Vervoort, 1993: 117, figs 3f, 4, 5, 6a-b, 8a.

Material examined

SMIB 6: Stn. DW127, two colonies, one unbranched and 4.5 cm high, the other forked twice and 9 cm

high, both bear male gonothecae (MNHN-IK-2012-16521).

Remarks

The difficulty of distinguishing G. amphorifera from G. ampullacea Vervoort, 1993 without their female

gonothecae is notorious. Since the present material bears only male gonothecae, attention was focused

on the hydrothecal features, in an attempt to separate them specifically. Although Vervoort (1993)

emphasized several distinguishing characters, the distinction was still laborious and, eventually, two

features proved useful. It was found that in the present species the free, distal part of the hydrocladial

hydrothecae was constantly set at about 90° with the axis of the cladia, regardless the position of the

thecae (Fig. 2A) while, in the following species, that angle varied considerably (Fig. 2B) between the

proximal hydrothecae (where it was of 80-90°) and the distalmost ones (usually reaching 45°). The size

of the hydrothecae was comparatively smaller in the present material (Fig. 3 A) than in that assigned to

G. ampullacea Vervoort, 1993 (Fig. 3C). I believe that the lack of a distinct inflation of the hydrothecae

invoked by Vervoort is misleading.

Some hydrothecae exhibit incipient lateral diverticuli (Fig. 3A). Their structure is discussed under

G. aff. similis Vervoort, 1993.

Geographical distribution

New Caledonia (Vervoort 1993; present study).

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Gonaxia ampullacea Vervoort, 1993

Figs 2B, 3C-D

Gonaxia ampullacea Vervoort, 1993: 121, figs 6c, 7a-c, 9.

Material examined

BATHUS 4: DW914, a 3.5 cm high colony with unbranched stem, bearing male gonothecae (MNHN-

IK-2012-16522).

Fig. 2. — A. Gonaxia amphorifera Vervoort, 1993 (SMIB 6, Stn. DW127), portion of stem with

cladium. — B. Gonaxia ampullacea Vervoort, 1993 (BATHUS 4, Stn. DW914), portion of stem with

cladia. — C. Gonaxia compacta Vervoort, 1993 (BATHUS 4, Stn. DW923), portion of stem with

cladium. — D. Gonaxia intermedia Vervoort, 1993 (BATHUS 4, Stn. CP906), portion of stem with

cladium. — E. Gonaxia sinuosa Vervoort, 1993 (SMIB 4, Stn. DW40), portion of stem with cladia.

Scale bar: A-E = 1 mm.

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Remarks

As stated above, the larger size and the more acute angle formed by the free part of the hydrocladial

hydrothecae with the axis of cladia were used to assign this material to G. ampullacea. As in

G. amphorifera , some hydrothecae exhibit incipient lateral diverticuli (Fig. 3C).

Geographical distribution

New Caledonia (Vervoort 1993; present study).

Gonaxia comp acta Vervoort, 1993

Figs 2C, 3E-F

Gonaxia compacta Vervoort, 1993: 135, figs 8c, 14a-b, 15a.

Material examined

BATHUS 4: Stn. DW923, a single, 6 cm high colony bearing an abortive gonotheca, as well as a row

of incipient gonothecae, highly obscured by the accessory tubes of the stem (MNHN-IK-2012-16523).

Remarks

As already stated by Vervoort (1993), the hydrothecae of this species bear an obvious resemblance to

those of G. amphorifera , G. ampullacea , and G. sinuosa , but it could be readily distinguished from them

on the account of the absence of a pit formed by the free adaxial wall with the internode (Fig. 3E). Only

an abortive gonotheca, arising from within the first hydrotheca of a cladium (Fig. 3F), was found. Several

successive, elongate masses among the accessory tubes of the stem suggest the imminent fonnation of

normal gonothecae.

Geographical distribution

New Caledonia (Vervoort 1993; present study).

Gonaxia intermedia Vervoort, 1993

Figs 2D, 3G-K

Gonaxia intermedia Vervoort, 1993: 157, figs 23e, 24d, 25a-c, 26a.

Material examined

BATHUS 4: Stn. DW902, several colonies and fragments, up to 9.5 cm high, mostly fertile, bearing

male gonothecae (MNHN-IK-2012-16524); Stn. CP906, a total of twelve fertile colonies (10 male and

2 female), 4-6 cm high (MNHN-IK-2012-16525).

Remarks

In some colonies, the length of cladia reaches as much as 2.8 cm, and each may bear up to 116 hydrothecae.

Aberrant secondary cladia, as those occurring in specimens from Stn. DW902, are given off from either

within some cladial hydrothecae (Fig. 3J), or from short, auxiliary cladial tubes running parallel to the

proximal parts of the axes of cladia.

Both male and female colonies are present in the material in hand. As stated by Vervoort (1993), the

gonothecae of this species resemble those of G. amphorifera and G. ampullacea through their general

shape and nearly fully adnate condition. They seem to originate from below the axillar hydrothecae. The

distal part of a gonotheca overlaps the proximal part of the following one, though their bodies never

fuse with each other. Their apertures (50-65 pm wide in males, and 250-290 pm wide in females)

European Journal of Taxonomy 218 : 1-52 ( 2016 )

are mounted on raised cones situated near the distal ends of their bodies. In the male colonies, the

gonothecae are so obscured by the numerous auxiliary tubes running parallel to the stem, that only the

tips of their neck regions protrude from the strongly fascicled mass which composes the stems. In one

male specimen (from Stn. CP906) with otherwise normal gonothecae, an elongate, abortive gonotheca,

with two distal apertures, is given off from the lumen of a cladial hydrotheca The occurrence of such

Fig. 3. — A-B Gonaxia amphorifera Vervoort, 1993 (SMIB6, Stn. DW127). A. Two successive

cladial internodes with their hydrothecae. B. Portion of stem with male gonothecae. — C-D. Gonaxia

ampullacea Vervoort, 1993 (BATHUS 4, Stn. DW914). C. Two successive cladial internodes with their

hydrothecae. D. Portion of stem with male gonothecae. — E-F. Gonaxia compacta Vervoort, 1993

(BATHUS 4, Stn. DW923). E. Two successive cladial internodes with their hydrothecae. F. Abortive

gonotheca (of undetermined sex) originating from within a hydrotheca. — G-K. Gonaxia intermedia

Vervoort, 1993 (BATHUS 4). G. Two successive cladial internodes with their hydrothecae (Stn. CP906).

H. Portion of stem with male gonothecae (Stn. CP906). I. Portion of stem with female gonothecae (Stn.

CP906). J-K. Abortive gonothecae arising from hydrothecal lumina (Stn. DW902). — L-N. Gonaxia

sinuosa Vervoort, 1993 (SMIB 4, DW40). L. Two successive cladial intemodes with their hydrothecae.

M-N. Two female gonothecae. Scale bars: A, C, E, G, T = 300 pm; B, D, F, H-K, M-N = 1 mm.

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

gonothecae is considerable in the material from Stn. DW902, and specimens with one (Fig. 3K), two

(Fig. 3J), or exceptionally three apertures could be seen. These so called abortive gonothecae may prove

fully functional, as they are provided with apertures. However, their lumina are empty in the specimens

in hand, and this assumption could not be checked at this stage.

Geographical distribution

New Caledonia (Vervoort 1993; present study).

Gonaxia sinuosa Vervoort, 1993

Figs 2E, 3F-N

Gonaxia sinuosa Vervoort, 1993: 179, figs 31c-e, 35b, 37a-b, 38a.

Material examined

SMIB 4: Stn. DW40, an originally 8 cm high colony with simple stem, now broken into two parts,

bearing male gonothecae (MNHN-IK-2012-16526); Stn. DW48, an originally 5.5 cm high, sterile

colony with simple stem, now broken into two unequal parts (MNHN-IK-2012-16527).

BATHUS 2: Stn. DW716, six colonies and fragments, 2.3-9.1 cm high, with simple or branched

stems, two of which are sterile, two others bear male gonothecae, and the remaining two carry female

gonothecae (MNHN-IK-2012-16528).

BATHUS 3: Stn. CP804, eleven colonies and fragments, 1.5—4.9 cm high, all with simple stems, of

which four are sterile, three bear male gonothecae, and the remaining four bear female gonothecae

(MNHN-IK-2012-16529); Stn. DW829, two colonies with simple stems, 3.8 and 2.4 cm high, the latter

devoid of its basal part, and bearing three incipient gonothecae (MNHN-IK-2012-16530).

Remarks

The shape of the hydrothecae (Fig. 3F), as well as the condition and morphology of the gonothecae

(Fig. 3M-N) are characteristic. One stem from BATHUS 2, Stn. DW716 is fully fertile (male) and bears

its gonothecae in two parallel rows, each one originating from below the base of an axillar hydrotheca.

The female stems examined here bear a lesser number of gonothecae.

Geographical distribution

New Caledonia (Vervoort 1993; present study), Norfolk Ridge (Vervoort 1993).

Gonaxia crassicaulis Vervoort, 1993

Fig. 4A

Gonaxia crassicaulis Vervoort, 1993: 145, figs 18c-d, 19a-b.

Material examined

MUSORSTOM 4: Stn. DW205, a cladium, 2.5 cm long (MNHN-IK-2012-16531).

NORFOFK 1: Stn. DW1712, an infertile colony, 3.8 cm high (MNHN-IK-2012-16532).

Remarks

According to Vervoort (1993), G. crassicaulis is easily recognizable through the characteristic

development of the perisarc, forming thick, internal, longitudinal ridges, connecting the adnate adaxial

walls of the hydrothecae in each row (Fig. 4A).

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Fig. 4. - A Gonaxia crassicaulis Vervoort, 1993 (MUSORSTOM 4, Stn. DW205), portion of

cladium. — B-D. Gonaxia elegans Vervoort, 1993 (BATHUS 3, Stn. DW809). B. Portion of stem and

basal part of a cladium. C. Two consecutive internodes with their hydrothecae. D. Portion of stem with

male gonothecae. — E-F. Gonaxia cf. errans Vervoort, 1993 (BATHUS 2, Stn. CP737). E. Distalmost

portion of stem with basal parts of four hydrocladia. F. Two consecutive cladial intemodes with their

hydrothecae. — G-J. Gonaxia plumularioides sp. nov. (BIOCAL, Stn. CP34). G. Distalmost part of a

stem, showing the structure of internodes. H. Two consecutive cladial internodes with their hydrothecae.

I. Stem hydrotheca. J. Axillar hydrotheca. Scale bars: A-B, D-E, G = 1 mm; C, F, H-J = 300 pm.

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Also characteristic is the presence of large, internal, semi-circular “thickenings” of the perisarc (Fig. 4A),

arching over the basal foramina, when the hydrothecae are seen laterally. These structures, one situated

in the “frontal” aspect of the hydrothecae, the other on their “dorsal” side, correspond in fact to the

attachment points of the hydrothecal bases to the lateral walls of the thecae which, in this species,

are raised above the level of the foramina. When the hydrothecae are seen facing their apertures, the

hydrothecal bases appear U-shaped, the foramina being situated at the bottom of this concavity.

Geographical distribution

New Caledonia (present study), Norfolk Ridge (Vervoort 1993).

Gonaxia elegans Vervoort, 1993

Fig. 4B-D

Gonaxia elegans Vervoort, 1993: 153, figs 19c, 21b-c, 22.

Material examined

BATHUS 3: Stn. DW809, a 4.4 cm high colony devoid of hydrorhiza and bearing female gonothecae,

as well as an originally 5.4 cm high colony (now broken into two parts) bearing male gonothecae and

several broken cladia originating from both colonies (MNHN-IK-2012-16533).

Remarks

This species is immediately recognizable through the characteristic position and shape of its hydrothecae.

The 5.4 cm high colony is evidently male when compared to the smaller one, which bears gonothecae

possessing much wider apertures. Only the female gonothecae were described by Vervoort (1993) but,

upon a careful inspection of the present material, it should be stated that they do not arise from the

accessory tubes on the hydrocaulus, but from the main tube itself, as this is indisputably seen in the

monosiphonic part of the stem. In addition, in this colony at least, nearly all accessory tubes are found

on the “dorsal” aspect of the stem, while the row of gonothecae is situated on its “front”.

The male gonothecae, observed here for the first time, are similar in shape to the female ones, and also

arise from the main tube of the stem, but in both the frontal and dorsal aspects of the colony. They are

given off closely and in a single row (Fig. 4D), are about half adnate, and their free parts are shifted

alternately left and right; their bases are obscured by the numerous accessory tubes running up the stem.

They are long (1.5-2 mm), tubular (480-520 pm wide), with small, circular apertures (100-120 pm

wide) mounted on the summit of a conical, distal neck region.

Geographical distribution

New Caledonia (Vervoort 1993; present study), Norfolk Ridge (Vervoort 1993).

Gonaxia cf. errans Vervoort, 1993

Fig. 4E-F

Gonaxia errans Vervoort, 1993: 154, figs 23a-d, 24a-c.

Material examined

BATHUS 2: Stn. CP737, two stem fragments, 13 and 9 mm high, the former corresponding to the middle

part of a colony, while the latter is likely the distalmost end of (another?) colony; both are sterile; several

broken cladia are also present (MNHN-IK-2012-16534).

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Remarks

The pattern of cladial arrangement on the stem in this scarce material seems to follow a rather peculiar

sequence: each equivalent of internode (the nodes are inconspicuous) has a proximal apophysis with its

associated axillar hydrotheca, two alternate hydrothecae above, as well as a second, distal apophysis (on

the side opposite to the former) with its associated axillar hydrotheca; the same sequence is repeated on

the following “internode” but, there, it is the opposite image of the preceding one. In the distalmost part

of one of the available colony fragments, two successive cladia occur on the same side, and are separated

on the opposite side by a single stem hydrotheca. According to Vervoort (1993), the cladial arrangement

is varied, with “zero to three hydrothecae between two successive apophyses”, which is apparently the

case in the present material.

The shape of the hydrothecae also fits the original description of G. errans , though they are comparatively

shorter in the material in hand (free adaxial side 335-365 pm, adnate adaxial side 280-300 pm, abaxial

side 490-510 pm), but seem to have almost the same width (maximum width 205-220 pm, diameter

at aperture 165-175 pm) as in Vervoort’s samples. Consequently, the present material is provisionally

assigned to G. errans.

Geographical distribution

New Caledonia (present study), Norfolk Ridge (Vervoort 1993).

Gonaxia plumularioides sp. nov.

um:lsid:zoobank.org:act:6A07FF84-EAA6-4AE6-8755-B450B17A28EB

Fig. 4G-J; Table 2

Diagnosis

Gonaxia with slender, fascicled, branched or unbranched stems; cladia monosiphonic. Equivalents of

internodes composed of a proximal hydrocladial apophysis and its associated axillary hydrotheca, two

alternate hydrothecae above, and a second, distal hydrocladial apophysis and its associated hydrotheca.

Hydrothecae tubular, slightly curved up- and outwards, comparatively slender and more immersed on

stem than on cladia. Gonothecae unknown.

Etymology

From the Latin pluma , meaning feather, and the Greek eiSog, meaning likeness, with reference to their

plumose appearance, recalling the branching pattern displayed by some hydroids belonging to the

superfamily Plumularioidea.

Material examined

Holotype

BIOCAL: Stn. CP34, a 2.8 cm high, distal fragment of a branch or a stem, itself branched twice, sterile

(MNHN-IK-2012-16535).

Paratype

BIOCAL: Stn. CP34, a 7.7 cm high, unbranched colony devoid of its hydrorhiza, as well as a 1 cm high

fragment corresponding to the distalmost part of a colony; both are sterile (MNHN-IK-2012-16536).

Description

Colonies attaining at least 8 cm high, composed of either simple or ramified stems of slender

appearance, despite being fascicled; accessory tubes on both “frontal” and “dorsal” sides of stems;

the latter uniformly grading to monosiphonic distally. Side branches, when present, arising only in

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Table 2. Measurements (in pm) of hydrothecae of Gonaxia plumularioides sp. nov. and G. nova sp. nov.

G. plumularioides sp. nov.

G. nova sp. nov.

Caulinar

hydrotheca

Axillar

hydrotheca

Cladial

hydrotheca

Caulinar

hydrotheca

Axillar

hydrotheca

Cladial

hydro theca

Free adaxial side

85-125

75-140

180-210

555-575

565-630

580-615

Adnate adaxial side

355-380

360-440

315-350

220-245

175-220

175-205

Abaxial side

315-345

195-245

395M40

600-630

505-575

620-665

Maximum width

110-125

125-145

195-205

260-270

270-290

250-260

Diameter at rim

80-90

80-95

160-170

160-165

150-185

165-175

“frontal” aspect of the stem or higher order branches; up to 2 nd order branching observed. Division into

internodes indistinct, but equivalents of internodes composed of the regular sequence: a first, basal

apophysis (60-90 pm long) and its associated axillar hydrotheca, two additional, alternate hydrothecae

above, and a second axillar apophysis with its associated hydrotheca. Following internode with same

sequence, but its proximal apophysis on side opposite to the distal apophysis of preceding internode.

Cladia leaving axis at 75-80° from upper part of stem or branches; two successive cladia on same side

of stem are distant of 2200-2310 pm; strictly monosiphonic, alternate, up to 1 cm long, comprising

a succession of up to 23 alternate hydrothecae. Division into internodes indistinct; equivalent of first

cladial internode comparatively longer (580-705 pm) than subsequent ones (480-515 pm); intemodes

short, accommodating a single hydrotheca distally. Hydrothecae widely spaced on stem (570-740 pm),

less so on cladia (470-530 pm); tubular, slightly curving up- and outwards, free part angled at 50° to

the axis of internode; stem hydrothecae slightly shifted on frontally; the two rows of cladial hydrothecae

coplanar; less than half adnate on cladia, but considerably immersed into the stem internodes; free

adaxial side nearly straight, adnate counterpart curved, ending basally in perisarc peg; abaxial side

convex, with inflexion point in middle; aperture with three low cusps separated by shallow embayments;

opercular apparatus lost. Gonothecae unknown.

Remarks

The branching pattern of the stems of G. plumularioides sp. nov. resembles that seen in some plumularioid

hy droids that build three-dimensional colonies (e.g., Macrorhynchia philippina Kirchenpauer, 1872,

M. allmani (Nutting, 1900), Plumularia spiralis Billard, 1911, etc.), and is consequently different from

that observed in its congeners (e.g., G. amphorifera, G. sinuosa , G. nova sp. nov., see description below),

which generally build planar colonies.

The regular structure of the stem internodes is also peculiar, and is only occasionally observed in the

material assigned herein to G. errans, a species known for its inconstant structure of the stem (Vervoort

1993).

The shape of the hydrocladial hydrothecae resembles that of other congeners: 1) G. complexa Vervoort,

1993, but in this species the hydrothecae have an exceedingly long part free from internode; 2) G. elegans ,

but its hydrothecae have a conspicuously straight axis instead of being convex (compare Fig. 4C and

4H); 3) G. errans , whose hydrothecae have, however, a much longer free part (compare Fig. 4F with

Fig. 4H); 4) G. intercalata Vervoort & Watson, 2003, but its hydrothecae are huge compared to those

of the new species; 5) G. stricta Vervoort, 1993, but here the hydrothecae have a much longer free

part, and the cladial internodes are marked by internal, oblique perisarcal thickenings (Vervoort 1993);

6) G. tasmanica Watson & Vervoort, 2001, but its hydrothecae are comparatively larger.

Geographical distribution

New Caledonia.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Gonaxici nova sp. nov.

um:lsid:zoobank.org:act:6C6AA611-2E5C-40B9-9502-31888388A7E9

Figs 5A-B, 6A-E; Table 2

Diagnosis

Gonaxia with either branched or unbranched, rigid, fascicled stems arising from disc-shaped hydrorhiza.

Intemodes slightly geniculate, comprising a latero-distal cladial apophysis with its associated hydrotheca,

and two alternate hydrothecae below. Cladia alternate, composed of moderately long, geniculate

internodes. Hydrothecae arranged at a wide angle to the stem; long, tubular, slightly arcuate, tumid

proximally, distally tubular, adnate for % their length. Gonothecae originating on both sides of the stem

from below axillar hydrothecae; adnate for about half their length; erect portion tubular, ending in rather

long neck region bearing the aperture distally.

Etymology

From the Latin novus , -a, -um (and the Grek vsof), meaning new, indicating that we are dealing with a

new, undescribed species.

Material examined

Holotype

MUSORSTOM 8: Stn. CP 1083, a 4.2 cm high, unbranched colony bearing male gonothecae (MNHN-

IK-2012-16537).

Paratypes

MUSORSTOM 8: Stn. CP1083, eight colonies or fragments with branched or unbranched stems, of

which four bear female gonothecae and range between 2.6 and 5.0 cm in height, while the remaining

four bear male gonothecae and range between 2.8 and 3.9 cm in height (MNHN-IK-2012-16538).

Description

Colonies composed ofbranched or unbranched, thick, rigid, fascicled stems arising from rhizoid hydrorhiza

firmly attached to substrate. Nodes inconspicuous, even in the monosiphonic parts, but equivalents of

internodes 1580-2000 pm long, slightly geniculate, composed of a latero-distal hydrocladial apophysis

with its associated hydrotheca, and two additional, alternate hydrothecae below. Hydrocladia strictly

monosiphonic, arranged at approximately right angles to the stem; up to 1.2 cm long, nodes indistinct

(60-70 pm wide); equivalents of internodes moderately long, geniculate, each carrying a hydrotheca

distally; first internode comparatively longer (1020-1240 pm) than subsequent ones (590-760 pm);

up to 20 hydrothecae per cladium. Hydrothecae long, tubular, facing outwards at nearly right angles to

their corresponding internodes; slightly tumid basally, walls parallel distally; adnate for about % th their

length; free adaxial side slightly convex proximally, then straight; abaxial wall with noticeable convexity

in middle; aperture triangular in frontal view; margin with three conspicuous cusps (two latero-adaxial

and one abaxial) separated by rather deep embayments; operculum composed of three triangular flaps

forming a low roof. Gonothecae on both sides of the stem, originating from below axillar hydrothecae;

about half adnate, strongly appressed proximally to the stem; distal half erect, tubular, ending in

moderately long, tronconical collar, bearing the aperture distally (small in male, Fig. 6E; comparatively

larger in female, Fig. 6D); perisarc of abaxial wall of fused part finely and densely striated (Fig. 6E-F);

individual gonothecae either solitary (towards apex of stem) or fused together through auxiliary tubes

given off from one or both ends of their adnate parts; in highly polysiphonic parts of stem, adnate parts

of gonothecae obscured by apposition of auxiliary tubes; length of adnate part 1250-1340 pm (S) and

1720-1970 pm (§), and of free part 1540-1910 pm (c?) and 1640-1720 pm (§); width of free part

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Fig. 5. — A-B Gonaxia nova sp. nov. (MUSORSTOM 8, Stn. CP 1083). A. Distalmost, monosiphonic

part of a colony. B. Portion of stem with basal part of a cladium. — C-G. Gonaxia scalariformis

Vervoort, 1993, portions of stems and basal parts of cladia from BIOCAL, Stn. DW46 (C = colony

#1; D = colony #2; E = colony #3), Stn. DW66 (F), and NORFOLK 1, Stn. DW1699 (G), showing

discrete differences in shape and size. — H. Gonaxia aff. similis Vervoort, 1993 (MUSORSTOM 6, Stn.

DW423), portion of stem with cladium. Scale bars: A = 1 mm; B-H = 500 pm.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

565-590 pm (c?) and 620-690 pm (§); length of neck region 260-365 pm (c?) and 165-245 p ($);

diameter at aperture 90-105 pm (c?) and 245-260 pm (5).

Remarks

The hydrothecae of this species resemble those of several congeners: 1) G. bulbifera Vervoort, 1993, but

this species has proportionally smaller hydro- and gonothecae (about two times smaller), its hydrothecal

axes are straight instead of being curved, its caulinar apophyses are conspicuously inflated, and its

cladial internodes are collinear instead of being slightly geniculate; 2) G. elegans , but in this species

the hydrothecae are much more closely-set, and the cladial internodes are strictly collinear (compare

Fig. 5B with Fig. 4B); 3) G. perplexa Vervoort, 1993, but in this species the hydrothecae are more

approximated and have straight axes, and the gonothecae are nearly fully adnate, their bodies being

likely fused to each other, and their apertures are characteristically shifted outward from the stem; 4-7)

G. complexa Vervoort, 1993, G. persimilis Vervoort, 1993, G. similis, and G. scalariformis, but these

species are immediately distinguished through their free gonothecae, originating perpendicularly to the

stems; 8) G. stricta Vervoort, 1993, but in this species the hydrothecae have straight axes and they are

more approximated on cladia.

Geographical distribution

Vanuatu.

Gonaxia scalariformis Vervoort, 1993

Figs 5C-G, 6F-R

Gonaxia scalariformis Vervoort, 1993: 173, figs 31b, 32, 33c-d, 35a.

Material examined

BIOCAF: Stn. DW46, three colonies (#1-3) slightly differing in the size of their hydrothecae: colony

#1 is 3.5 cm high, devoid of its hydrorhiza, and bears two female gonothecae (MNHN-IK-2012-16539),

colony #2 is 2.8 cm high, devoid of its hydrorhiza, and bears three female gonothecae (MNHN-

IK-2012-16540), and colony #3 is 2.2 cm high and broken in two parts, the basal one e xhi biting re mn ants

of hydrorhiza; both parts bear five male gonothecae each (MNHN-IK-2012-16541); Stn. DW66, a 1.6

cm high colony devoid of both its hydrorhiza and apical part, with two abortive gonothecae (MNHN-

IK-2012-16542); Stn. CP69, a 1.4 cm high colony devoid of its distalmost part, with a single female

gonotheca (MNHN-IK-2012-16543).

NORFOFK 1: Stn. DW1699, a 1.7 cm high colony devoid of both its hydrorhiza and upper part, bearing

two female gonothecae (MNHN-IK-2012-16544).

Fig. 6. (previous page) — A-E. Gonaxia nova sp. nov. (MUSORSTOM 8, Stn. CP 1083). A. Axillar

hydrotheca. B. Caulinar hydrotheca. C. Cladial hydrotheca. D. Female gonothecae, lateral view. E. Male

gonotheca, slightly oblique view. — F-R. Gonaxia scalariformis Vervoort, 1993. F. Axillar hydrotheca

(BIOCAF, Stn. DW46, colony # 1). G. Cladial hydrotheca (BIOCAF, Stn. DW46, colony # 1). H. Caulinar

and axillar hydrothecae (BIOCAF, Stn. DW66). I-M. Normal gonothecae from BIOCAF, Stn. DW46

(I = colony #1; J-K = colony #2; F-M = colony #3). N-O. Abortive gonotheca from BIOCAF, Stn.

DW66. P-R. Normal gonothecae from BIOCAF, Stn. CP69 (P) and NORFOFK 1, Stn. DW1699 (Q-

R). — S-X. Gonaxia aft similis Vervoort, 1993 (MUSORSTOM 6, Stn. DW423). S. Cauline hydrotheca.

T. Axillar hydrotheca. U. Cladial hydrotheca. V. Fateral view of a hydrotheca, showing diverticulum.

W. Same hydrotheca viewed from above, showing minute foramen for the passage of coenosarc, and

lateral diverticuli. X. Same hydrotheca in apical view, showing lateral diverticuli and basal foramen.

Scale bars: A-C, F-H, S-U = 300 pm; D-E, I-R = 1 mm; V-X = 200 pm.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Remarks

All available specimens have lightly fascicled stems basally, and none is ramified. Although the

arrangement and general shape of the hydrothecae perfectly match the description of the holotype,

noticeable variations in size could be observed. Thus, the colony #3 from sample BIOCAL, Stn. DW46

has the slenderest hydrothecae, while those from NORFOLK 1, Stn. DW1699 are the widest ones. In

the sample from BIOCAL, Stn. DW66, incipient antero-dorsal diverticuli are seen in most hydrothecae

(Fig. 5F). Gonothecae of both sexes are present in the available material. Although they conform to

the original description, the male gonothecae in the colony #3 from BIOCAL, Stn. DW46 are arranged

on both anterior and posterior sides of the stem, and some exhibit bifid (Fig. 6M) or trifid apices. The

female gonothecae from BIOCAL, Stn. DW66 are likely abortive, as they originate from within the

hydrothecae (Fig. 6N-0); in addition, they are not fully formed, as their distal ends are missing; their

sex is indeterminate. The female gonothecae from NORFOLK 1, Stn. DW1699 (Fig. 6Q-R) are the

longest with respect to those present in the other samples.

Geographical distribution

New Caledonia and Norfolk Ridge (Vervoort 1993; present study).

Gonaxia aff. similis Vervoort, 1993

Figs 5H, 6S-X

Gonaxia similis Vervoort, 1993: 174, figs 33e-f, 34, 36a, 39a.

Material examined

MUSORSTOM 6: Stn. DW423, a 2.7 cm high, sterile colony (MNHN-IK-2012-16545).

Remarks

The present specimen is likely part of the material in RMNH-Coel. 25839 (also from MUSORSTOM 6,

Stn. DW423) studied by Vervoort (1993).

According to Vervoort, his material had gonothecae adnate to the stem (p. 174, fig. 34c), not arranged

perpendicularly to it, as in the majority of samples assigned by him to G. similis. Consequently, the

material in hand may belong to a separate species.

One hydrocladium in the present specimen is branched, and the secondary cladium is borne on a short,

quadrangular apophysis originating from below the hydrothecal base of the first internode of a cladium.

Not mentioned by Vervoort in his sample RMNH-Coel. 25839 is the constant presence of well-developed

antero-posterior diverticuli within the hydrothecae (Figs 5H, 6S-V). Such variably developed diverticuli

also occur in other congeners discussed herein, viz. G. amphorifera (Fig. 3 A), G. ampullacea (Fig. 3C),

and G. scalariformis (Fig. 5F). In the present material, their development is considerable, and extends

well on each lateral side of the hydrotheca. A frontal view of the free adaxial wall of the hydrotheca

(Fig. 6W), and a frontal view of the aperture (Fig. 6X) show two empty lateral “pockets” which narrow

considerably the hydrothecal lumen; the foramen for the passage of the hydranth within the hydrotheca

is minute and horseshoe-shaped.

Geographical distribution

New Caledonia (Vervoort 1993; present study).

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Genus Idiellcma Cotton & Godfrey, 1942

Idiellana pristis (Lamouroux, 1816)

Fig. 7A

Idiapristis Lamouroux, 1816: 199, pi. 5, fig. 5.

Idiellana pristis - Schuchert 2003: 175, fig. 32.

Material examined

MUSORSTOM 3: Stn. CP 121, sterile colony composed of two stems, 1.7 and 3.7 cm high (MNHN-

IK-2012-16546); Stn. CP134, sterile colony composed of two stems, 1.5 and 2.7 cm high (MNHN-

IK-2012-16547).

MUSORSTOM 8: Stn. DW1021, two sterile stems, 3.5 and 5 cm high (MNHN-IK-2012-16548); Stn.

CP1131, several sterile stems, 1.5-8.5 cm high (MNHN-IK-2012-16549).

TAIWAN 1: Stn. DW1, four sterile stems, 0.7-1.4 cm high, on gravel (MNHN-IK-2012-16550).

Remarks

For the latest redescription and a list of synonyms, refer to Schuchert (2003).

Geographical distribution

Circumglobal in tropical and subtropical seas (Precker & Lawn 2010).

Genus Salacia Lamouroux, 1816

Salacia macer Vervoort & Watson, 2003

Fig. 7B-G

Salacia macer Vervoort & Watson, 2003: 154, figs 34g, 35a.

Material examined

BATHUS 3: Stn. DW809, single sterile stem, 2.4 cm high, partly damaged (several cladia missing)

(MNHN-IK-2012-16551).

NORFOLK 1: Stn. DW1659, four stems 3—4.5 cm high, two of which bear female gonothecae (MNHN-

IK-2012-16552).

Remarks

The slender habit of these colonies, their regularly-pinnate branching pattern and their division into

intemodes, each of which bears a single pair of strictly opposite hydrothecae, are diagnostic (Vervoort

& Watson 2003). The gonothecae, not previously described, are present in this material, and are female.

They are borne on either the stem or cladia, and are arranged perpendicularly from below one hydrotheca

of a pair. Urn-shaped, smooth-walled, 3280-3565 pm long and 1110-1170 pm wide, tapering basally

into a short pedicel; distally truncate, aperture circular, 590-615 pm wide, borne on a short, 565-575 pm

wide neck region. Possibly up to 100 oocytes per gonotheca, though their number is difficult to ascertain.

Geographical distribution

Norfolk Ridge (Vervoort & Watson 2003; present study).

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Fig. 7. (continued on next page) —A. Idiellanapristis (Lamouroux, 1816) (TAIWAN 1, Stn. DW1), stem

internode and basal part of cladium. — B-G. Salacia macer Vervoort & Watson, 2003 (NORFOLK 1,

Stn. DW1659). B. Stem internode and basal part of cladium. C. Hydrothecae, frontal view. D. Same,

dorsal view. E. Hydrothecal aperture, frontal view. F. Hydrothecal aperture, oblique view, showing

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Salacia sibogae Billard, 1924

Fig. 7H

Salacia sibogae Billard, 1924: 64, fig. lb-c.

Salacia sibogae - Schuchert 2003: 177, fig. 34.

Material examined

MUSORSTOM 3: Stn. DR126, four small (0.4-1.1 cm high), sterile stems on axis of dead gorgonian

(MNHN-IK-2012-16553).

Remarks

A recent redescription of this species and a list of synonyms are available in Schuchert (2003).

Geographical distribution

Previosuly known from Indonesia and Japan (Schuchert 2003). The present material originates from the

Philippines.

Genus Sertularella Gray, 1848

Sertularella acutidentata Billard, 1919

Fig. 71

Sertularella acutidentata Billard, 1919: 20, figs le, 2.

Sertularella acutidentata acutidentata - Vervoort & Watson 2003: 154, fig. 35b-e.

Material examined

TAIWAN 2: Stn. DW118, two infertile colonies, 7 and 8 cm high (MNHN-IK-2012-16554).

Remarks

For a redescription of this species, refer to Hirohito (1995). Its synonymy is given by Vervoort & Watson

(2003).

Geographical distribution

Indonesia, Japan, Philippines, New Caledonia, and north of New Zealand (Vervoort & Watson 2003).

operculum. G. Female gonotheca. — H. Salacia sibogae Billard, 1924 (MUSORSTOM 3, Stn. DR126),

portion of stem and basal part of cladium. — I. Sertularella acutidentata Billard, 1919 (TAIWAN 2,

Stn. DW118), portion of stem with basal part of side branch. — J. Sertularella areyi Nutting, 1904

(BATHUS 3, Stn. CP813), portion of fertile stem. — K-L. Sertularella diaphana (Allman, 1885)

(MUSORSTOM 8, Stn. CPI 104). K. Two hydrothecae. L. Gonotheca. — M. Sertularella mirabilis

Jaderholm, 1896 (TAIWAN 2, Stn. DW118), portion of colony. — N-Q. Sertularella novaecaledoniae

Vervoort, 1993 (MUSORSTOM 4, Stn. DW221). N. Portion of stem and proximal part of a branch.

O-P. Hydrothecae. Q. Portion of stem with gonothecae. — R. Sertularella quadridens (Bale, 1884)

(MUSORSTOM 3, Stn. CP121), portion of stem and proximal part of a side branch. Scale bars: A-B =

500 pm; C-F = 400 pm; G-N, Q-R = 1 mm; O-P = 300 pm.

European Journal of Taxonomy 218: 1-52 (2016)

Sertularella areyi Nutting, 1904

Fig. 7J

Sertularella areyi Nutting, 1904: 83, pi. 17, fig. 6.

Sertularella areyi - Vervoort & Watson 2003: 156, fig. 35f-l.

Material examined

MUSORSTOM 4: Stn. CC174, several stems, up to 1.4 cm high, all but one sterile (MNHN-

IK-2012-16555).

BATHUS 3: Stn. CP813, fertile colonies with stems up to 1.4 cm high on two fragments of gorgonian

(MNF1N-IK-2012-16556); Stn. CP814, two infertile colonies with stems up to 1.5 cm high, one on dead

gorgonian, the other on sponge (MNHN-IK-2012-16557).

LITHIST: Stn. DW13, small colony composed of a few sterile stems, up to 1.2 cm high, growing on

Acryptolaria sp. (MNF1N-IK-2012-16558).

BORDAU 2: Stn. DW1513, small colony composed of several sterile, up to 5 mm high, stems growing

on G. ceramensis (MNHN-IK-2012-16559).

NORFOLK 1: Stn. DW1704, small colony composed of a restricted number of stems, some fertile and

up to 1 cm high, epizoic on Zygophylax sp., Sertularella helenae Vervoort, 1993, and tube of coronate

medusa (MNHN-IK-2012-16560).

Remarks

This very distinctive species needs no additional comments on morphology. Its synonymy, as well as a

redescription, are provided by Vervoort & Watson (2003).

Geographical distribution

Tropical and subtropical parts of the Atlantic and Indo-Pacific (Vervoort & Watson 2003).

Sertularella diaphana (Allman, 1885)

Fig. 7K-L

Thuiaria diaphana Allman, 1885: 145, pi. 18, figs 1-3.

Sertularella diaphana - Vervoort & Watson 2003: 159, fig. 36g-h.

Material examined

MUSORSTOM 8: Stn. CPI 104, colony composed of several fertile (male gonothecae) stems, up to 6 cm

high, epizoic on stem of aglaopheniid hydroid (MNHN-IK-2012-16561).

Remarks

For lists of synonyms and descriptions, see Calder (1991), Vervoort (1933) and Galea (2010).

Geographical distribution

Circumglobal in tropical and subtropical seas (Vervoort & Watson 2003).

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Sertulcirella mirabilis Jaderholm, 1896

Fig. 7M

Sertularella mirabilis Jaderholm, 1896: 9, pi. 2, fig. 1.

Sertularella mirabilis - Schuchert 2015: 339, fig. 13.

Material examined

TAIWAN 2: Stn. DW118, a 3.2 cm high, sterile colony (MNHN-IK-2012-16562).

Remarks

Hirohito (1995) provided a thorough description of this peculiar species. Its synonymy is given by

Schuchert (2015).

Geographical distribution

From the South China Sea to Korea and Japan (Schuchert 2015).

Sertularella novaecaledoniae Vervoort, 1993

Fig. 7N-Q

Sertularella novaecaledoniae Vervoort, 1993: 225, figs 50, 51aA, 52a.

Material examined

MUSORSTOM 4: Stn. DW221, a 4 cm high, fertile colony (MNHN-IK-2012-16563).

Remarks

The present material, represented by a colony with flabellate appearance, with fascicled stem and side

branches divided into very short intemodes, each carrying a tubular hydrotheca adnate for about half

its length, conforms to the description of the type given by Vervoort (1993). The gonothecae have

undulated walls, and their apical part is flattened (Fig. 7Q).

Geographical distribution

New Caledonia (present study), Norfolk Ridge (Vervoort 1993).

Sertularella quadridens (Bale, 1884)

Fig. 7R

Thuiaria quadridens Bale, 1884: 119, pi. 7, figs 5-6.

Sertularella quadridens - Schuchert 2003: 185, fig. 41.

Material examined

MUSORSTOM 3: Stn. DR117, a 18 mm high, sterile colony (MNHN-IK-2012-16564); Stn. CP121,

a complete colony, 3 cm high, as well as four fragments, 1.4-3.2 cm high, all sterile (MNHN-

IK-2012-16565).

Remarks

For a list of synonyms and a recent redescription, see Schuchert (2003).

European Journal of Taxonomy 218: 1-52 (2016)

Geographical distribution

Widely distributed throughout the tropical Indo-Pacific (Precker & Lawn 2010).

Sertularella folliformis sp. nov.

um:lsid:zoobank.org:act:7A654AE7-6B4E-465C-9F5A-lD48528466E5

Fig. 8A-G; Table 3

Diagnosis

Sertularella species with flabellate colonies; branching irregular; stem and branches strongly polysiphonic

basally, grading to monosiphonic distally; internodes short, slightly geniculate to collinear; hydrothecae

exceedingly long, tubular, curved downwards, with 8-10 broad, complete annuli and four internal,

submarginal cusps. Gonothecae large, elongated-ovoid, distally truncate.

Etymology

From the Latin words follis, meaning bellows, and forma , meaning shape, with reference to the plicate

condition of the hydrothecal wall, recalling the bellows of an accordion.

Material examined

Holotype

BATHUS 3: Stn. DW809, a 4.5 x 4.5 cm sterile colony and four smaller fragments, three of which bear

one gonotheca, and the fourth two gonothecae (MNHN-IK-2012-16566).

Paratype

BATHUS 3: Stn. DW809, a few colonies and fragments, largest one 7.5 x 3.5 cm, all sterile (MNHN-

IK-2012-16567).

Description

Colonies flabellate, attaining 7.5 cm high, arising from rhizoid stolons firmly attached to substrate.

Branching irregular, with up to 3 rd order branches arranged roughly in one plane. Stems and side branches

polysiphonic, main stem reaching as much as 2 mm wide basally. Both stem and branches divided into

internodes by slightly-marked nodes slanting in alternate directions. Internodes short, slightly geniculate

to collinear, each carrying a hydrotheca in its distal half. Side branches originate laterally below the

bases of stem hydrothecae; first intemode comparatively longer than subsequent ones. Hydrothecae

exceedingly long, tubular, slightly curved downward, adnate for % th or less their length, tapering

imperceptibly towards aperture, walls provided with 8-10 broad, complete annuli; margin occasionally

renovated; rim provided with four short cusps separated by shallow embayments; four submarginal,

intrathecal projections of perisarc, two latero-adaxial and two latero-abaxial. Gonotheca originating

from axil of hydrotheca, in front and/or backside of colony; elongated-ovoid, tapering abruptly below,

distally truncate, wall smooth to slightly undulate.

Remarks

The hydrothecal shape is diagnostic in this species. There are few congeners which approach

this condition, viz. S. catena (Allman, 1888) and S. pseudocatena sp. nov. (Fig. 9J), but both have

much shorter hydrothecae, and their transverse ribs do not extend over the abaxial wall. Three other

species, namely S. helenae Vervoort, 1993 (Fig. 8H), S. paucicostata Vervoort, 1993 (Fig. 81), and

S. pseudocostata Vervoort, 1993, exhibit comparatively more prominent hydrothecal ridges. The eastern

Pacific S. exilis Fraser, 1938 has isodiametric hydrothecae “regularly curved upward and then outward”

(Fraser 1938), but their surface is reportedly smooth. In addition, it is “an especially diminutive species”

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Table 3. Measurements (in pm) of Sertularella folliformis sp.

nov. and S. plicata sp. nov.

S. folliformis sp. nov.

S. plicata sp. nov.

Internodes

- length (in general)

680-865

660-890

- length (1 st cladial internode)

1360-2405

1365-1450

- diameter at node

325M00

125-145

Hydrotheca

- free adaxial side

1025-1235

365-440

- adnate adaxial side

375—415

190-225

- abcauline side

1170-1295

450M95

- maximum width

320-345

255-275

- diameter at rim

290-305

170-180

Gonotheca

- length

2245-2345

—

- maximum width

780-850

—

- apical width

445M55

—

(Calder et al. 2009). Sertularella whitei Rees & Vervoort, 1987 also has exceedingly long, outwardly-

curved hydrothecae, but their surface is entirely smooth (Rees & Vervoort 1987).

Geographical distribution

New Caledonia.

Sertularella helenae Vervoort, 1993

Fig. 8H

Sertularella helenae Vervoort, 1993: 218, fig. 47b-e.

Material examined

BIOCAL: Stn. DW37, several sterile stems, up to 2.6 cm high, and smaller fragments (MNHN-

IK-2012-16568); Stn. DW38, single, sterile colony 3 cm high and 4.7 cm wide (MNHN-IK-2012-16569).

BATHUS 3: Stn. CP804, a 2.5 cm high and 3.8 cm wide, sterile colony (MNHN-IK-2012-16570);

Stn. CP821, a 3.4 cm high, sterile colony (MNHN-IK-2012-16571); Stn. DW830, four sterile colony

fragments, 0.7-1.5 cm high (MNHN-IK-2012-16572).

NORFOLK 1: Stn. CP1719, a 4.2 cm high, sterile colony (MNHN-IK-2012-16573).

SALOMON 1: Stn. DW1741, two sterile colonies, 2.2 and 2.6 cm high (MNHN-IK-2012-16574).

Remarks

This very distinctive species needs no additional comments on morphology. For a thorough description,

see Vervoort (1993). The gonothecae still remain to be discovered.

Geographical distribution

New Caledonia (Vervoort 1993; present study), Norfolk Ridge and Solomon Islands (present study).

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Sertularella paucicostata Vervoort, 1993

Fig. 81

Sertularella paucicostata Vervoort, 1993: 227, figs 51b-f, j.

Material examined

MUSORSTOM 3: Stn. DW809, several infertile stems, some branched, up to 1.3 cm high, epizoic on

Sertularella folliformis sp. nov. (MNHN-IK-2012-16575).

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Remarks

Surprisingly, Vervoort (1993) compared his new species with Sertularella costata Leloup, 1940, which

is a much smaller and delicate hy droid (compare Fig. 81 and 8J), as illustrated by specimens of this

species from the Andaman Sea, Thailand belonging to my private collection.

Sertularella costata is a rather poorly known species, with a few records around the world. Hirohito

(1995) provided succinct measurements of the hydrothecae, while Calder et al. (2003) gave the first

description and measurements of the gonotheca. The dimensions of the hydrothecae in the Thailand

material are as follows: abcauline side 315-360 pm, free adcauline side 265-280 pm, adnate adcauline

side 100-110 pm, maximum width 155-170 pm, diameter at aperture 130-140 pm. The external wall

is provided with 10-13 transverse ridges, and three internal, submarginal cusps are present within the

hydrothecae (Fig. 8K). Through the size of its hydrothecae, the material from Thailand comes close to

the Japanese specimens examined by Hirohito and, through the number of external ridges, it approaches

that of the Galapagos specimens, which is apparently 10-11 (Calder et al. 2003: fig. 14). In contrast,

about 20 ridges ornament the hydrothecal walls in material from Japan (Leloup 1940; Hirohito 1995).

In my opinion, S. paucicostata most closely resembles both S. helenae and S. pseudocostata Vervoort,

1993. However, unlike S. paucicostata , S. helenae forms fascicled colonies with shorter internodes, and

its hydrothecae are smaller (compare Fig. 81 and 8H) and provided with numerous perpendicular creases

between the transverse ridges. According to Vervoort (1993), the hydrothecae of S. pseudocostata are

comparatively larger and their outer wall has frilled ribs.

Geographical distribution

New Caledonia, Norfolk Ridge (Vervoort 1993), and the Philippines (present study).

Sertularellaplicata sp. nov.

urn:lsid:zoobank.org:act:5CBEB7F6-12F7-403C-AC8D-58AE01BllAC7

Fig. 8F-0; Table 3

Diagnosis

Small Sertularella with monosiphonic stems; side branches given off irregularly in one plane. Internodes

moderately long and geniculate. Hydrothecae piriform, with 8-10 conspicuous, transverse ribs; aperture

on short, narrow neck facing outwards; 3 or 5 internal, submarginal, lamellar cusps. Gonothecae

unknown.

Fig. 8. (previous page) —A-G. Sertularellafolliformis sp. nov. (BATHUS 3, Stn. DW809). A. Holotype.

B. Paratype. C. Portion of stem with proximal part of a side branch. D. Successive internodes with

their hydrothecae. E. Hydrothecal aperture, frontal view, showing internal cusps. F. Portion of stem

with gonotheca, showing insertion site. G. Gonotheca. — H. Sertularella helenae Vervoort, 1993

(SAFOMON A, Stn. DW1714), two successive internodes with their hydrothecae. — I. Sertularella

paucicostata Vervoort, 1993 (BATHUS 3, Stn. DW809), two consecutive intemodes with their

hydrothecae, and comparison with Sertularella costata Leloup, 1940 (J-K). — L-O. Sertularella plicata

sp. nov. (MUSORSTOM 3, Stn. CP 121). L. Portion of stem and proximal part of a side branch. M. Two

consecutive internodes with their hydrothecae. N. More detailed view of the hydrotheca, to show its

wrinkled surface. O. Hydrothecal aperture in frontal view, showing internal cusps. — P-S. Sertularella

aff. sinensis Jaderholm, 1896. P. Portion of stem (BATHUS 3, Stn. DW829). Q. Hydrotheca (BORDAU

2, Stn. DW1595). R. Frontal view of three hydrothecal apertures, to show variation in the number of

internal cusps. S. Gonotheca (BORDAU 2, Stn. DW1595). Scale bars: A-B = 1 cm; C, F-G, F, P =

1 mm; D, H-J, M, Q, S = 500 pm; E, K, O, R = 100 pm; N = 200 pm.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Etymology

From the Latin plico , - atum , -are (and the Greek izlt/yo), meaning “pleat, pleated, to pleat”, to characterize

the appearance of the external surface of the hydrothecal wall.

Material examined

Holotype

MUSORSTOM 3: Stn. CP121, colony composed of several sterile stems up to 1 cm high, epizoic on

dead gorgonian (MNHN-IK-2012-16576).

Description

Colony composed of up to 1 cm high, monosiphonic stems arising from creeping, branching hydrorhiza.

Basal parts of stems of varied length and ahydrothecate, perisarc smooth; above, stems divided into

regular, moderately long, geniculate intemodes, by means of slightly oblique nodes slanting in alternate

directions; each intemode with one, distal hydrotheca. Side branches arising irregularly from below the

bases of stem hydrothecae and in the same plane as the stem; from one to several hydrothecae between

two successive side branches; up to second order branching observed; structure identical to that of

stem, except for the first internode, which is comparatively longer than following ones; distal-most,

tendril-like internodes occasionally anastomose with each other or with neighboring stems. Hydrothecae

swollen basally, then narrowing gradually distally to a short neck region projecting the aperture slightly

outwards; adaxial side less than l A adnate; walls with 8-10 conspicuous transverse ridges encircling

the theca; numerous creases, perpendicular to the ridges, give the hydrotheca a wrinkled appearance;

aperture rhomboidal, rim with four blunt cusps separated by shallow embayments; opercular apparatus

composed of four triangular valves forming a low roof; 3 or 5 internal, submarginal, lamellar cusps

(two latero-adaxial and one adaxial and, in some hydrothecae, two additional, latero-adaxial ones).

Gonothecae not observed.

Remarks

Hydrothecae with such deep, transverse ridges are present in a number of congeners, but several distinctive

features allow their separation from the new species: 1) S. areyi Nutting, 1904, but its hydrothecae

are provided with only 2-3 transverse ridges (Fig. 7J); 2) S. costata Leloup, 1940 is a much smaller

hydroid (compare Fig. 8J and 8M); 3) S. crenulata Nutting, 1905 possesses a considerable number

of closely-set, transverse ridges, and its hydrothecae are tronconical (Vervoort 1993); 4-6) S. helenae

Vervoort, 1993 (Fig. 8H), S. paucicostata Vervoort, 1993 (Fig. 81) and S. pseudocostata Vervoort, 1993,

but their hydrothecae are larger and more tubular; 7) S. fraseri Galea, 2010 is a more delicate hydroid

and its hydrothecae are provided with fewer ridges; 8) S. quadrata Nutting, 1895 has densely-striated,

exceedingly long and tubular hydrothecae, with characteristically squared cross-section (Nutting 1904);

9) the hydrothecae of S. sinensis Jaderholm, 1896 are only half adnate, and the number of ridges they

carry reaches as many as 15 in the northern hemisphere (Naumov 1969); 10) S. striata Stechow, 1923,

but in this species the internodes are comparatively shorter, almost collinear, and the hydrothecae are

less fusiform and only about half adnate (Millard 1964).

Geographical distribution

Philippines.

Sertularella aff. sinensis Jaderholm, 1896

Fig. 8P-S; Table 4

Sertularella sinensis Jaderholm, 1896: 11, pi. 2, figs 2-3.

Sertularella sinensis - Vervoort & Watson 2003: 111 , fig. 41d-h.

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Table 4. Measurements (in pm) of Sertularella sinensis

Jaderholm, 1896 from various

sources.

Present study

Naumov (1969)

Vervoort(1993)

Vervoort & Watson (2003)

(CHALCAL 2,

Stn. DW80)

(NZOl Stn.

K825, slide 2270)

(NMNZ BS437,

slide 2977)

Internode

- length

655-865

—

- diameter at node

180-215

—

Hydro theca

- free adaxial side

480-540

230-270

250-310

345-390

395M25

- adnate adaxial side

315-335

250-270

265-310

345-375

280-360

- abcauline side

555-590

350-370

295-355

425-460

420M30

- maximum width

335-360

—

250-275

310-330

280-295

- diameter at aperture

205-220

150-170

170-185

145-165

195-215

- number of ridges

19-21

11-15

10-12

16-20

ca 18

Material examined

BATHUS 3: Stn. DW829, a sterile, polysiphonic, flabellate colony 3 cm high and 3.8 cm wide, broken

off from its 2.3 cm long basal part (MNHN-IK-2012-16577).

BORDAU 2: Stn. DW1595, a single fertile stem, 6.7 cm high, polysiphonic basally, with only two side

branches left, one of which is incomplete (MNHN-IK-2012-16578).

Remarks

The specimen from BORDAU 2 comprises a tall, erect, straight stem with only two side branches left,

while numerous lateral apophyses indicate the presence of pre-existent branches. The upper part of the

specimen from BATHUS 3 is flabellate, and this structure most probably arose upon the breakage of the

apical part of the initial stem and the subsequent profuse development of several distal side branches,

which divided up to four times. Though of different appearance, both specimens bear hydrothecae of the

same shape and size, and are believed to be conspecific.

The identification of the present material is provisional, owing to multiple discrepancies exhibited by

the specimens from a wide geographical area assigned to S. sinensis by various authors. Among the

points of difference, the following ones are noteworthy: 1) the habit of the colonies is apparently strictly

monosiphonic in the material from the northern hemisphere (Hirohito 1995), and polysiphonic in that

from the southern one (Vervoort 1993; Vervoort & Watson 2003; present study); 2) the shape of the

hydrothecae illustrated in the literature varies from decidedly compressed longitudinally in their free

part (Jaderholm 1896; Rho & Chang 1974; Hirohito 1995) to “normally” shaped (Vervoort & Watson

2003); 3) the number of transverse ridges accounts from 6-8 (Hirohito 1983) to 11-15 (Naumov 1969),

and to as many as 19-21 (present material); 4) the dimensions of the hydrothecae exhibit a wide range of

variation, as exemplified in Table 4; 5) the number of transverse ridges at the surface of the gonothecal

wall varies from 5-6 (Vervoort & Watson 2003), to ca 8 (Hirohito 1995), and to ca 14 (Jaderholm 1896:

pi. 2, fig. 3); 6) the number of apical projections of the gonotheca reported, for instance, is either 4

(Jaderholm 1896; Hirohito 1995) or 8 (Vervoort & Watson 2003); 7) small gonothecae (750-800 x 550-

700 pm) were reported by Hirohito (1995), while comparatively larger ones (1720-1935 x 1230-1280

pm) occurred in the material studied by Vervoort & Watson (2003).

The available data suggest that we are possibly dealing with at least two sibling species: the northern

hemisphere form is apparently monosiphonic in habit, the free parts of their hydrothecae are

European Journal of Taxonomy 218 : 1-52 ( 2016 )

characteristically compressed longitudinally, the number of annulations they carry is low (not exceeding

15), and their gonothecae are small and provided with 4 apical projections; conversely, the specimens

from the southern hemisphere are represented by colonies with fascicled stems and branches, the

hydrothecae are “normally” shaped, the number of annulations they carry may reach as many as 21,

their gonothecae are comparatively larger, and their apices are provided with 8 projections.

In addition, the present material comes closer to that from New Zealand (Vervoort & Watson 2003),

judging by the size and number of ribs of the hydrothecae, than to the New Caledonian one (Vervoort

1993), which have smaller hydrothecae, with a lesser number of transverse ridges (see Table 4).

The gonotheca depicted in Fig. 8S is likely immature, owing to its apparently incomplete rings and

flattened apical part, and originates from an auxiliary tube running up the main stem of the colony from

BORDAU 2, similarly to some gonothecae examined by Vervoort &Watson (2003).

Geographical distribution

Sertularella sinensis sensu stricto occurs from China (Jaderholm 1896) to the Sea of Okhotsk (Naumov

1969). Additional material, possibly not conspecific, was recorded from New Caledonia (Vervoort 1993;

present study), Tonga (present study), and New Zealand (Vervoort & Watson 2003).

Sertularella cf. anguina Vervoort, 1993

Fig. 9A; Table 5

Sertularella anguina Vervoort, 1993: 198, figs 40b-d, 41a-b.

Material examined

NORFOTK 1: Stn. DW1722, a sterile, unbranched stem, 3 cm high (MNHN-IK-2012-16579).

Remarks

This lightly fascicled stem (only basally), divided into moderately long, slender, geniculate internodes,

each carrying distally an exceedingly long, slender hydrotheca with tapering walls and five internal,

submarginal cusps (two latero-adaxial, one abaxial, and two latero-abaxial), fits the diagnosis of S.

anguina given by Vervoort (1993). This author gave no precise information on the exact number

of internal projections of the perisarc, but assumed that four may have been present in his material,

possibly two latero-adaxial and two latero-abaxial. The fifth, strictly abaxial cusp observed in the

present material is only visible when the hydrothecal apertures are seen frontally, but not laterally.

Although the measurements between the material in hand and the holotype appear similar (Table 5),

certain differences (as the width of the hydrotheca) may be attributed to the fact that the former was

measured in water, while the latter is a microslide preparation that suffered a certain degree of shrinkage.

Geographical distribution

New Caledonia (Vervoort 1993) and Norfolk ridge (present study).

Sertularella cf. bipectinata Vervoort, 1993

Fig. 9B-C; Table 5

Sertularella bipectinata Vervoort, 1993: 207, figs 43c-e, 44.

Material examined

BATHUS 3: Stn. DW809: a 2.2 cm high, sterile, lightly fascicled (a couple of accessory tubes) stem,

provided with two short side branches (MNHN-IK-2012-16580).

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Table 5. Measurements (in pm) of Sertularella cf. anguina Vervoort, 1993, S. cf. bipectinata Vervoort,

1993, and S. leiocarpoides Vervoort, 1993, and comparisons with their respective holotypes (second

column for each species).

S. cf. anguina Vervoort, 1993

S. cf. bipectinata

Vervoort, 1993

S. leiocarpoides

Vervoort, 1993

Present study:

NORFOLK 1,

Stn. DW1722

Vervoort (1993):

MUSORSTOM

6, Stn. CP464

Present study:

BATHUS 3,

Stn. DW809

Vervoort

(1993):

BIOCAL, Stn.

DW36

Present study:

BATHUS 3,

Stn. DW829

Vervoort (1993):

MUSORSTOM

4, Stn. DW212

Internode

- length

830-1200

755-850

1050-1110

1150-1260

755-1025

870-1040

- diam. at node

255-310

175-205

325-355

435M55

300-340

280-305

Hydrotheca

- free adax. side

840-900

760-845

1110-1170

1000-1065

680-765

695-735

- adn. adax. wall

305-340

235-245

380-390

450-520

395^430

410-435

- abaxial wall

925-1010

890-935

1245-1285

1195-1300

900-950

870-910

- max. width

300-315

235-245

375-390

410M35

395M20

390M10

- diameter at rim

210-245

150-170

260-270

260-265

300-310

280-305

Gonotheca

- length

ca 2955

3100-3255

- max. width

—

ca 1055

1000-1020

- apical width

—

ca 425

—

Remarks

The present material, although similar to that assigned above to S. aff. anguina Vervoort, 1993, differs

from it through its comparatively thicker intemodes, a more reduced ratio of internode length/hydrothecal

depth, and the comparatively larger size of its hydrothecae (Table 5). These features, in combination with

the presence of three internal, submarginal cusps (two latero-adaxial and one abaxial), also reported by

Vervoort (1993), suggest that it may belong to S. bipectinata , and this identification should be regarded

as provisional in the absence of fertile material.

Geographical distribution

New Caledonia (present study), Norfolk Ridge (Vervoort 1993).

Sertularella leiocarpoides Vervoort, 1993

Figs 9D-G, 10A; Table 5

Sertularella leiocarpoides Vervoort, 1993: 223, fig. 49.

Material examined

BATHUS 2: Stn. CP737, a 5 cm high, sterile colony devoid of hydrorhiza (MNHN-IK-2012-16581).

BATHUS 3: Stn. DW829, a 3.5 cm high colony fragment bearing one gonotheca (MNHN-IK-2012-16582);

Stn. DW830, a 5 cm high and 7.5 cm wide sterile colony (MNHN-IK-2012-16583).

SALOMON 1: Stn. DW1741, a 2 cm high, sterile colony (MNHN-IK-2012-16584).

Remarks

The four available specimens were compared to a colony of Sertularella leiocarpa (Allman, 1888)

from Inaccessible Island (see Galea 2015b). Portions of three of them are illustrated altogether with S.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

leiocarpa, so as to show their specific differences (compare Fig. 5D-F and 5G). As already noted by

Vervoort (1993), the internodes are thick, short and geniculate in S. leiocarpoides , and comparatively

more slender, longer, and less geniculate in S. leiocarpa. In addition, the hydrothecae of Vervoorfs

species have the adaxial cusps characteristically upturned, and there are generally also two inconspicuous

latero-adaxial, internal, submarginal projections of the perisarc and, occasionally, a third one, on the

abaxial side.

Geographical distribution

New Caledonia (Vervoort 1993; present study), Solomon Islands (present study).

Sertularellapseudocatena sp. nov.

urn:lsid:zoobank.org:act:3395EF3F-3DFF-489B-AB25-B4E2A58F8F7C

Figs 9H-M, 10B-C; Table 6

Sertularella catena - Vervoort 1993: 210, fig. 45a-c [not Sertularella catena (Allman, 1888)].

Diagnosis

Flabellate colonies with highly polysiphonic stems and branches; branching irregular, given off roughly

in one plane; internodes rather short; hydrothecae long, tubular, almost straight; free adaxial wall with

5-7 transverse ridges extending almost to abaxial wall; gonotheca sac-shaped, with 1-3 broad, transverse

ridges and 2-5 inconspicuous apical projections.

Etymology

The specific epithet results from the word-forming element pseudo- (derived from the Ancient Greek

y/EvdfiQ, pseudes), meaning false, in appearance only, or resembling, and catena , the name of a species

created by Allman (1888), suggesting that the new species superficially resembles S. catena.

Material examined

Holotype

BATHUS 3: Stn. DW809, single flabellate colony ca 6.5 x 6.5 cm, as well as three fragments from it, two

of which bearing one gonotheca, w hil e the third one carries two gonothecae (MNHN-IK-2012-16585).

Paratype

BATHUS 3: Stn. DW809, several colonies up to 8.5 cm high, as well as numerous smaller fragments

(MNHN-IK-2012-16586).

Fig. 9. (previous page) —A. Sertularella cf. anguina Vervoort, 1993 (NORFOLK 1, Stn. DW1722),

hydrotheca. — B-C. Sertularella cf. bipectinata Vervoort, 1993 (BATHUS 3, DW809), two hydrothecae.

— D-F. Sertularella leiocarpoides Vervoort, 1993, three hydrothecae (D = BATHUS 2, Stn. CP737;

E = BATHUS 3, Stn. DW829; F = Stn. DW829), and comparison with Sertularella leiocarpa (Allman,

1888) (G) from Inaccessible Island. — H-M. Sertularella pseudocatena sp. nov. (BATHUS 3, Stn.

DW809). H. Holotype. I. Paratype. J. Portion of stem with side branch. K-M. Three hydrothecae. —

N-P. i Sertularella splendida sp. nov. (BATHUS 3, Stn. DW809). N. Portion of stem and proximal part of

a side branch. O-P. Two hydrothecae. — Q-T. Sertularella tronconica sp. nov. Q. Portion of side branch

(NORFOLK 1, Stn. DW1704). R. Portion of stem and proximal part of a side branch (BATHUS 3, Stn.

DW809). S-T. Two hydrothecae (NORFOLK 1, Stn. DW1704). — U-X. Sertularella tubulosa sp.

nov. (BATHUS 3, Stn. DW810). U. Distal portion of stem. V. Portion of stem and proximal part of side

branch. W-X. Two hydrothecae. Scale bars: A-G, K-M, O-P, S-T, W-X = 500 pm; J, N, Q-R, U-V =

1 mm; H-I = 1 cm.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Table 6. Measurements (in jam) of four new species of Sertularella discussed herein.

S. pseudocatena

sp. nov.

(BATHUS 3,

Stn. DW809)

S. splendida

sp. nov.

(BATHUS 3,

Stn. DW809)

S. tronconica sp. nov.

(NORFOLK 1, (BATHUS 3,

Stn. DW1704) Stn. DW809)

S. tubulosa

sp. nov.

(BATHUS 3,

Stn. DW810)

Internodes

- length (in general)

495-680

965-1260

660-825

620-890

825-990

- length (1 st cladial internode)

1135-1395

ca 1730

—

1260-1630

ca 2240

- diameter at node

225-290

335-365

220-240

195-245

275-315

Hydrotheca

- free adaxial side

560-635

1050-1170

565-605

445-480

900-1050

- adnate adaxial side

295-330

325—410

300-315

290-355

350-420

- abcauline side

705-740

1170-1320

660-690

585-660

1125-1170

- maximum width

270-290

400^110

295-310

265-300

315-350

- diameter at rim

240-255

280-300

170-185

150-160

275-290

Gonotheca

- length

1235-1445

ca 2135

- maximum width

615-765

—

ca 925

—

- apical width

345-395

—

Description

Colonies flabellate, up to 8.5 cm high, of rigid appearance, though rather brittle upon manipulation, as

demonstrated by the highly fragmented condition of the present material. Stolon rhizoid, firmly attached

to substrate. Stems strongly fascicled, attaining up to 3 mm wide basally; irregularly ramified (up to 5 th

order branches, but possibly more in living specimens), branches roughly coplanar. Perisarc moderately

thick, straw colored. Basal parts of side branches also polysiphonic, thinning out to monosiphonic

distally. Main tube of both stems and side branches divided into rather short intemodes by means of

oblique nodes slanting in alternate directions; internodes distinctly geniculate, each carrying a hydrotheca

distally. Side branches arising laterally from below the bases of hydrothecae; first internode longer than

subsequent ones. Hydrothecae alternate, long, tubular, given off at about 25° with axis of internode; free

adaxial side with 5-7 conspicuous wrinkles reaching abaxial wall, which appears undulated; wrinkles

backward-oriented at site of origin on adaxial wall (Fig. 9L-M), becoming transverse towards abaxial

wall (Fig. 9K); base of hydrotheca with an incomplete septum, leaving a large foramen for the passage of

hydranth; margin generally renovated several times, provided with four short cusps separated by shallow

embayments; aperture quadrangular in frontal view; no internal, submarginal projections of perisarc.

Coenosarc poorly preserved, tentacle number could not be ascertained. Gonothecae (only 4 seen) arising

from an internode on side opposite hydrotheca, midway between base of theca and proximal node;

thick-walled, with 2-3 transverse “ribs”, aperture on well-formed neck, enclosed by filmy perisarc; 2-5

blunt, inconspicuous projections around aperture; body tapering below into indistinct pedicel; sex could

not be ascertained.

Remarks

Vervoort(1993) assigned his infertile material from New Caledonia to S. catena\ Allman, 1888), a species

known only from the Caribbean region, on the account of morphological similarities in the “structure

and shape of the hydrothecae”. Unfortunately, Vervoort did not provide any updated illustrations of

Allman’s holotype to facilitate a comparison with the New Caledonian specimens.

However, though of nearly similar size, the appearance of the holotype of S. catena , illustrated by

Allman (1888: pi. 27, fig. 2), differs radically from the silhouettes of the colonies from New Caledonia

(Fig. 9H-I), in having a definite main stem and a comparatively more sparing branching pattern. In

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

addition, unlike the strictly tubular hydrothecae observed in the material in hand (Fig. 9J-M), those of

S. catena appear “subcylindrical” (especially on the adaxial side) and “narrowed slightly at each end”

(Nutting 1904: pi. 15, fig. 3); this is also evident from Allman’s illustration (1888: pi. 27, fig. 2a).

The present material bears several gonothecae and, consequently, allows a much more reliable distinction

from Allman’s species to be made. Unlike the transversely ribbed gonothecae observed here, those of

S. catena are reportedly “elongate ovate, destitute of annulation, with a small terminal, bicuspate orifice”

(Allman 1888). Upon a reexamination of the type. Nutting (1904) added the following: “Gonangia [... ]

large, terete in form, orifice terminal, small, armed with two shallow processes or teeth” (Nutting 1904).

Although their surface was originally illustrated as totally smooth (Allman 1888: pi. 28, fig. 2a), Nutting

(1904) added that “There is apparently a series of very shallow rugosities or annulations on the distal

portion of the gonangia” (illustrated in his pi. 15, fig. 3). This is corroborated by Vervoort’s observation,

which confirms the existence “at the apical third” of the gonotheca of “two indistinct furrows”, though

he states that “there is no funnel and no distinct opening”.

In addition, the length of the gonothecae in the present material encompasses that of a whole internode

with its associated hydrotheca, measured from the proximal node to the hydrothecal aperture. In contrast,

the gonothecae of S. catena are comparatively bigger: in Allman’s figure (1888: pi. 28, fig. 2a), the

gonotheca is illustrated being as long as three successive internodes, while in Nutting’s figure (1904: pi.

15, fig. 3) is encompasses the length of 2.5 intemodes.

Taken together, the appearance of the colonies, with their profuse branching pattern, the perfectly

tubular and slightly curved hydrothecae marked with conspicuous furrows at their surface, as well as

the comparatively smaller and transversely ribbed gonotheca, suggest that we are dealing with a species

distinct from S. catena. Additional, geographical considerations support this view.

Geographical distribution

New Caledonia (Vervoort 1993; present study).

Sertularella splendida sp. nov.

urn:lsid:zoobank.org:act:900C4DlA-64A7-402E-B695-3BF396F 18579

Fig. 9N-P; Table 6

Diagnosis

Sertularella with lightly fascicled, irregularly and sparsely branched stems; internodes moderately long,

geniculate; hydrothecae exceedingly long, gradually narrowing slightly from base to aperture, adnate

for % or less, free adaxial side with 7-9 transverse ridges, an intrathecal cusp abaxially. Gonothecae

unknown.

Etymology

From the Fatin adjective splendido, -a, -um (splendeo), originally meaning brilliant, dazzling, or

glowing, but subsequently retained as “splendid” or “magnificent” in modern usage, to denote the

beautiful appearance displayed by this species.

Material examined

Holotype

BATHUS 3: Stn. DW809, a 23 mm high, sterile stem fragment devoid of its basal part, but carrying a

short, distally broken side branch; an accessory tube on the stem (MNHN-IK-2012-16587).

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Paratypes

BATHUS 3: Stn. DW809, four sterile stems or fragments, 7-14 mm high; side branches occur in two

specimens; accessory tubes also present in two specimens (MNHN-IK-2012-16588).

Description

Colonies up to 23 mm high, arising from rhizoid stolon firmly attached to substrate; stems either lightly

fascicled basally (with 1-2 auxiliary tubes) or monosiphonic; basal part of varied length, ahydrothecate,

with smooth, thick perisarc; above, stem divided into regular internodes by means of slightly oblique

nodes sloping in alternate directions; internodes geniculate, moderately long, widening gradually

towards distal end, where they bear a hydrotheca. Side branches, when present, originating from below

a stem hydrotheca, either laterally, or slightly in front of colony; first internode longer than subsequent

ones, constricted basally at insertion on stem; remainder of side branch with same structure as stem.

Hydrothecae alternate on stem and branches, originating at an angle of 50-60° with the intemode;

exceedingly long, slightly curving downward proximally and upward distally, tapering gradually,

though rather imperceptibly, towards aperture; adaxial wall % to A adnate to internode; free part with

7-9 prominent grooves; abaxial wall slightly sigmoid; basal foramen wide; distal end of hydrotheca

slightly upturned; margin with four short, triangular cusps separated by shallow embayments; aperture

quadrate in frontal view; an intrathecal cusp abaxially. Gonothecae absent.

Remarks

A careful search of the relevant literature reveals no other Sertularella species which approaches the

present hydroid in the morphology of its hydrothecae.

Geographical distribution

New Caledonia.

Sertularella tronconica sp. nov.

urn:lsid:zoobank.org:act:CC79A536-B897-4CA4-9C85-DB835908E5BE

Figs 9Q-T, 10D; Table 6

Diagnosis

Sertularella with lightly fascicled, irregularly and sparsely branched stems; colonies coplanar; intemodes

moderately long, geniculate; hydrothecae tronconical, adnate for A their length; adaxial wall with 4-5

transverse ridges; 5 prominent intrathecal, submarginal cusps; gonotheca large, transversely ridged

adaxially.

Eymology

From the Latin words, truncare , meaning to truncate, and conus (also xojvoc in Greek), meaning cone,

to characterize the shape of the hydrotheca.

Material examined

Holotype

NORFOLK 1: Stn. DW1704, single, unbranched colony, 2 cm high, devoid of its hydrorhiza, but

bearing a basal accessory tube running up the stem; a female gonotheca present (several large oocytes

still noticeable inside) (MNHN-IK-2012-16589).

Paratypes

BATHUS 3: Stn. DW809, several sterile stems and fragments, up to 1.4 cm high, some branched and

some with lightly fascicled stems (MNHN-IK-2012-16590).

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Description

Colonies up to 2 cm high, arising from creeping (young colonies) to rhizoid (adult colonies) hydrorhiza,

firmly attached to substrate. Basal part of stem of varied length, ahydrothecate, bearing a reduced number

of accessory tubes; perisarc thick and smooth. Remainder of stem divided into regular internodes by

means of oblique constrictions of the perisarc slanting in alternate directions. Internodes moderately

long, width increasing imperceptibly towards distal ends, so as to accommodate the basal part of a

hydrotheca. Branching scarce and irregular, in one plane, with up to 2 nd order branches observed; side

branches originating laterally from below the base of a stem hydrotheca; first internode comparatively

longer than subsequent ones, constricted basally at insertion on stem; remainder of side branches with

similar structure as the stem. Hydrothecae arranged alternately at an angle of 35M0° with the internode;

moderately long, tronconical, widening slightly below aperture; adaxial wall adnate for about A its

length; free part with 4-5 transverse, though incomplete, ridges, not reaching abaxial side, running

parallel to adnate side of adaxial wall; abaxial wall nearly straight; margin occasionally renovated;

aperture quadrate in frontal view, bearing 4 short cusps separated by shallow embayments; five

prominent, submarginal, intrathecal cusps: two latero-adaxial, two latero-abaxial, and one abaxial.

Gonothecae originating from the internode, on side opposite hydrotheca, midway between its base and

proximal node; large, transversely ribbed; a short, distal neck; aperture large, surrounded by 3-4 almost

imperceptible, blunt cusps; a few large oocytes noticeable by transparency.

Remarks

Several species possess hydrothecae that resemble, more or less, the shape of those of S. tronconica

sp. nov., but the latter may be reliably distinguished from its congeners as follows: 1-2) in both S. anguina

and S. bipectinata the length/width ratio of the hydrotheca is comparatively higher (Vervoort 1993);

3) the hydrothecae of S. inconstans Billard, 1919 are adnate for a greater extent of their adaxial wall

(Rees & Vervoort 1987), and the transverse ridges, which encircle the bodies of proximal hydrothecae,

gradually fade off towards the distal end of the colony, and are eventually represented by indistinct

undulations of the perisarc on the adaxial side (Billard 1925b); in addition, the internodes are only

slightly geniculate and do not reach the degree of geniculation encountered in S. tronconica sp. nov.;

4) S. keiensis Billard, 1925b, but in this species the hydrothecae are slightly smaller and adnate for a

longer extent, their surface varies from wrinkled (Billard 1925b) to smooth (Leloup 1937), and there are

four lamellar, internal, submarginal cusps (two latero-adaxial and two latero-abaxial); 5) S. leiocarpoides ,

but this species has comparatively larger, smooth-walled hydrothecae (compare Fig. 9S-T with 9D-F);

6) although very similar morphologically, the hydrothecae of S. natalensis Millard, 1968 have a length/

width ratio of ca 2.1-2.3 (calculated from Millard 1968, 1975), while those of S. tronconica sp. nov.are

comparatively more slender, with a ratio of ca 2.4-2.9; in addition, the ridges are common in younger

hydrothecae and may totally disappear in the older ones (Millard 1968); moreover, its gonothecae appear

more fusiform, only their distal end is wrinkled, and the aperture is surrounded by 3-4 projections

of perisarc (Millard 1975); 7) unlike in the present species, the length/width ratio of the hydrotheca

in S. undulitheca Vervoort, 1959 is lower, between 2.1 and 2.3 (calculated from Vervoort 1959), its

internodes are comparatively shorter and slightly geniculate, and both the hydro- and gonothecae are

wrinkled throughout.

Geographical distribution

New Caledonia and Norfolk Ridge.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Sertularella tubulosa sp. nov.

um:lsid:zoobank.org:act:8D56454C-7B2C-451C-BFB2-9C42603F3005

Fig. 9U-X; Table 6

Diagnosis

Sertularella with lightly polysiphonic, irregularly and sparsely ramified stems; internodes moderately

long, geniculate; hydrothecae exceedingly long, tubular, isodiametric throughout, walls entirely smooth;

four small marginal cusps, and four submarginal, intrathecal projections of perisarc. Gonothecae

unknown.

Etymology

The specific name is derived from the Latin tubulus , meaning tube, and describes the shape of the

hydrotheca.

Material examined

Holotype

BATHUS 3: Stn. DW810, single, sterile, branched stem, 2.6 cm high, lightly fascicled basally, but

devoid of hydrorhiza (MNHN-IK-2012-16591).

Description

The colony, devoid of hydrorhiza, is composed of a single stem, 2.6 cm high, giving rise to a side

branch towards its middle. Perisarc thick basally, thinning out gradually towards tip of colony. Basal

part of stem ahydrothecate, overgrown by a few accessory tubes; upper and longest part divided into

regular internodes by oblique constrictions of the perisarc slanting in alternate directions. Internodes

moderately long, geniculate, widening distally so as to accommodate a hydrotheca. The single side

branch originates laterally from below a stem hydrotheca; first internode comparatively longer than

subsequent ones; the latter with same structure as the stem. Hydrothecae angled at 40-45° with the stem;

long, tubular, isodiametric throughout, adnate for about % their length; free adaxial wall nearly straight

and smooth; abaxial wall basally convex, becoming rapidly straight for most of its length; basal foramen

wide; aperture squared in frontal view, with four inconspicuous marginal cusps separated by shallow

embayments; four intrathecal, submarginal cusps: two latero-adaxial and two latero-abaxial; margin

often renovated in damaged hydrothecae. Gonothecae unknown.

Remarks

The characteristic tubular shape of the hydrothecae of this species is diagnostic. Tubular, smooth-walled

hydrothecae are only encountered in a few congeners that can be distinguished from the new species

on the following accounts: 1) the hydrothecae of S. billardi Vervoort, 1993 have a comparatively lower

length/width ratio of the hydrothecae (Vervoort 1993); 2) S. exilis Fraser, 1938 has typically curved

hydrothecae (Fraser 1938), and the species is minute (Calder et al. 2009); 3) S. vervoorti El Beshbeeshy,

2011 possesses huge hydrothecae, whose length/width ratio is lower than in S. tubulosa sp. nov. (El

Beshbeeshy 2011); 4) in S. whitei the hydrothecae originate at a more acute angle with the axis of

internode, are smaller, curve slightly outwards, and the stem internodes are longer and more slender than

in the present species (Rees & Vervoort 1987).

Geographical distribution

New Caledonia.

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Genus Sertularia Linnaeus, 1758

Sertularia tongensis (Stechow, 1919)

Fig. 10E

Sertularella tongensis Stechow, 1919: 89, fig. f-g 1 .

Sertularia tongensis - Galea 2010: 18, fig. 5h-m (synonymy); 2015b: 10, fig. 3s.

Material examined

MUSORSTOM 9: Stn. CP 1265, small colony composed of a few sterile, unbranched stems, up to 1 cm

high (MNHN-IK-2012-16592).

Remarks

As stated by Galea (2010, 2015b), this species is immediately recognizable, even in sterile condition,

through its alternate hydrothecae whose apertures are closed by two opercular flaps pleated along a

median line.

Geographical distribution

Likely circumglobal in temperate to tropical seas (Galea 2015b). The present record is from the

Marquesas.

Genus Symplectoscyphus Marktanner-Turneretscher, 1890

Symplectoscyphus effusus Vervoort, 1993

Fig. 10F-H; Table 7

Symplectoscyphus effusus Vervoort, 1993: 253, figs 59, 60a-e.

Material examined

NORFOLK 1: Stn. DW1712, small colony composed of several stems up to 11 mm high, mostly fertile

(female), on worm tube covered by sand grains (MNHN-IK-2012-16593).

Remarks

The long, tubular, upwardly-curved hydrothecae, and the transversely ribbed gonothecae with frilled

margin, characteristically curving downwards, are diagnostic (Vervoort 1993). In addition, the

comparative measurements of this material and of the holotype (see Table 7) leave no doubt about

their conspecificity. The gonothecae in the specimen examined have 4-5 prominent ribs, and these are

definitely transversely and not spirally arranged, as stated by Vervoort. Two large oocytes or planulae

occur in each gonotheca.

Geographical distribution

New Caledonia (Vervoort 1993), Norfolk Ridge (present material).

Symplectoscyphus fasciculatus nom. nov.

Fig. 10I-L

Symplectoscyphus johnstoni tropicus Vervoort, 1993: 259, figs 61, 62a-d [not S. johnstoni (Gray, 1843)].

non Symplectoscyphus johnstoni tropicus - Vervoort & Watson 2003: 217, fig. 5 If—1.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

Table 7. Measurements (in pm) of S ymplectoscyphus effusus Vervoort, 1993 and S. paucicatillus sp. nov.

_ S- effusus Vervoort, 1993 _ & paucicatillus sp. nov.

Present study: NORFOLK 1, Vervoort (1993): (present study: BATHUS 4,

Stn. DW1712 CHALCAL 2, Stn. DW08 Stn. DW923)

Internodes

- length (in general)

415-745

405-690

455-875

- diameter at node

95-105

65-90

155-230

Hydrotheca

- free adaxial side

305-340

215-235

405-470

- adnate adaxial side

150-160

115-170

215-245

- abcauline side

325-340

295-320

465-530

- maximum width

135-150

105-120

230-250

- diameter at rim

130-135

135-140

185-195

Gonotheca

- length

ca 1175

ca 1110

1240-1480

- maximum width

ca 835

ca 645

865-890

- tube length

—

ca 120

165-180

- tube diameter

—

ca 155

- diameter at aperture

ca 175

ca 110

260-270

non Symplectoscyphus tropicus (Hartlaub, 1901).

Diagnosis

Symplectoscyphus species of flaccid appearance; stems and side branches fascicled basally, geniculate

at origin of branches; branching pseudodichotomous; internodes composed of two alternate hydrothecae

and a lateral apophysis with its associated hydrotheca; all hydrothecae tubular, curved outwards;

gonothecae sac-shaped, nearly fully adnate, perisarc undulated, aperture round, on top of short funnel.

Etymology

From the Latin fascis , meaning fascicled, to characterize the polysiphonic habit of the colonies.

Fig. 10. (previous page) — A. Sertularella leiocarpoides Vervoort, 1993 (BATHUS 3, Stn. DW829),

gonotheca. — B-C. Sertularella pseudocatena sp. nov. (BATHUS 3, Stn. DW809), two gonothecae. —

D. Sertularella tronconica sp. nov. (NORFOLK 1, Stn. DW1704), gonotheca. — E. Sertularia tongensis

(Stechow, 1919) (MUSORSTOM9, Stn. CP1265), internode with hydrotheca. — F-H. Symplectoscyphus

effusus Vervoort, 1993 (NORFOLK 1, Stn. DW1712). F. Portion of colony. G. Hydrotheca.

H. Gonotheca. — I-L. Symplectoscyphus fasciculatus nom. nov. (MUSORSTOM 4, Stn. CC174).

I. Portion of colony. J-K. Two hydrothecae. L. Male gonotheca. — M-V. Symplectoscyphus ralphae

Vervoort, 1993 (NORFOLK 1, Stn. DW1712). M-N. Samples #1 and #2, repectively. O-P. Portions

from samples #1 and #2, respectively. Q-T. Hydrothecae from samples #1 (Q-R) and #2 (S-T).

U-V. Gonothecae from samples #1 and #2, respectively. — W-Z. Symplectoscyphus paucicatillus

sp. nov. (BATHUS 4, Stn. DW923). W. Paratype, colony silhouette. X. Portion of colony. Y. Two

hydrothecae. Z. Two gonothecae. Scale bars: A-D, F, I, O-P, X = 1 mm; E = 300 pm; G, J-K, Q-T, Y =

200 pm; H, L, U-V, Z = 500 pm; M-N, W = 1 cm.

European Journal of Taxonomy 218 : 1-52 ( 2016 )

Material examined

Holotype

MUSORSTOM 4: Stn. CP153, “several well developed, flabellate colonies 30^50 mm, stems basally

polysiphonic; many appressed gonothecae [...]. One large colony 30x40 mm is holotype (MNHN-Hy.

1117)” (Vervoort 1993).

Additional material

MUSORTSOM 4: Stn. CC174, a 4.5 cm high, fertile (male) colony.

Description

Flaccid, flabellate, up to 4 cm high and 5 cm wide colonies arising from branched, rhizoid stolon.

Stems strongly fascicled basally, thinning out gradually to monosiphonic distally; flexuous, usually

regularly and alternately branched; occasionally with “irregular” branches resulting from considerable

development of secondary stems from branches growing as long as the main stem itself, and resulting in

multiple tangled stems, whose distalmost branches form multiple anastomoses between them and with

neighboring stems. Division into internodes indistinct, even in monosiphonic parts, but equivalents of

internodes long, geniculate, comprising a basal apophysis and its associated axillar hydrotheca, as well as

two alternate hydrothecae above. Apophyses alternate. Side branches straight (until they rebranch again

in pseudodichotomous manner), nodes indistinct, but each equivalent of internode bearing hydrothecae

arranged alternately. Hydrothecae tubular, curving away from internode, adnate for nearly half their

length. Free adaxial wall straight to slightly convex; abaxial wall straight basally, curving suddenly in

its distal third. Hydrothecal aperture nearly parallel to longitudinal axis of internode; margin with three

pointed cusps separated by rather deep embayments; adaxial cusp recurved; renovations common and

numerous. No conspicuous submarginal, intrathecal cusps, except for an occasional abaxial thickening.

Gonothecae given off from below hydrothecal bases, fitting tightly and neatly against internodes. Male

and female similar in shape; sac-shaped, wall undulated and never ribbed; undulations more pronounced

distally, fading off basally; aperture distal, borne on outwardly-projecting, trumpet-shaped terminal tube.

Remarks

The description provided above combines Vervoorfs (1993) account with personal observations of the

material in hand. The latter apparently belongs to a “large tangled tuft of c. 30 mm high colonies on

wormtubes” previously examined by Vervoort.

Owing to the fasciculate habit of its stems and the gonothecae with undulated walls, the large material

studied by Vervoort was assigned to a new subspecies, tropicus , of Symplectoscyphus johnstoni

(Gray, 1843). In doing so, Vervoort distinguished it from its congeners with monosiphonic stems and

transversely-ribbed gonothecae, viz. S. johnstoni johnstoni (Gray, 1843) and S. johnstoni sub tropicus

Ralph, 1961, the latter being additionally characterized by the presence of three submarginal, intrathecal

cusps.

As stated by Ralph (1961), S. johnstoni is a species with strictly monosiphonic stems, and its gonothecae

are provided with “8 to 14 evenly spaced transverse ridges”. Consequently, the subspecies tropicus could

be confidently raised to species. However, when doing so, it becomes an invalid junior homonym of

S. tropicus (Hartlaub, 1901). To avoid this situation, the junior homonym is replaced here with the

binomen S. fasciculatus nom. nov., whose holotype remains the same for both the original and the new

name of the single biological species (ICZN Art. 72.7). The measurements of S. fasciculatus nom. nov.

are given by Vervoort (1993: 260, table 51).

A restricted number of species belonging to this highly speciose genus are polysiphonic in habit:

six congeners from the Antarctic [S. bellinghauseni Pena Cantero, 2012 (Pena Cantero 2012);

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

S. cumberlandicus (Jaderholm, 1905) (Pena Cantero et al. 2002); S. frondosus Pena Cantero, 2010

(Pena Cantero 2010); S. hesperides Pena Cantero, 2012 (Pena Cantero 2012); S. liouvillei (Billard,

1914) (Pena Cantero et al. 2002); S. sofiae Pena Cantero et al., 2002 (Pena Cantero et al. 2002)], ten

from the subantarctic [S. adpressus (Ritchie, 1911) (Ritchie 1911); S. amoenus Vervoort & Watson,

2003 (Vervoort & Watson 2003); S. arboriformis (Marktanner-Turneretscher, 1890) (Millard 1975);

S. columnarius (Briggs, 1914) (Ralph 1961); S. divaricatus (Busk, 1852) (Ralph 1961); S. magellanicus

(Marktanner-Turneretscher, 1890) (Galea & Schories 2012); S. patagonicus Galea & Schories, 2012

(Galea & Schories 2012); S. paulensis Stechow, 1923 (Watson 2003); S. procerus (Trebilcock, 1928)

(Ralph 1961); S. subarticulatus (Coughtrey, 1875) (Millard 1977)], and two from the tropics, both

assigned to S. tropicus Hartlaub, 1901 [sensu Clarke 1894 (as Sertularella variabilis ) and sensu Billard

1925b],

Ecologically, S. fasciculatus nom. nov. occurs in a biogeographical area radically different from

that of the majority of species listed above. Its gonothecae are different from those of the following

species, which exhibit transverse ridges: S. aggreggatus (Jaderholm, 1917) (Pena Cantero et al. 2002),

S. arboriformis (Millard 1975), S. columnarius (Ralph 1961), S. cumberlandicus (Jaderholm 1905),

S. magellanicus (Galea & Schories 2012), S. patagonicus (Galea et al. 2014), S. procerus (Ralph 1961),

S. sofiae (Pena Cantero et al. 2002), and S. subarticulatus (Ralph 1961). The gonothecae of S. frondosus

(Pena Cantero 2010) and S. liouvillei (Pena Cantero et al. 2002) are exceedingly long and tubular, while

those of S. amoenus (Vervoort & Watson 2003) and S. paulensis (Pena Cantero 2012) are ovoid and

have smooth to slightly undulated walls, yet they are not adherent to their corresponding internodes, and

their distal tubes are not eccentrically placed. The habit and shape of the gonothecae of S. fasciculatus

nom. nov. come closest to those of S. adpressus. However, the latter is a more robust species, with

pinnate appearance, and its hydrothecae are subopposite and highly immersed into their intemodes

(Ritchie 1911). Although not completely formed in the specimens of S. bellinghauseni studied by Pena

Cantero (2012), the gonothecae seem “to be characterized by the presence of rings”. Similarly, the

gonothecae of S. hesperides were incipient and apparently smooth (Pena Cantero 2012), but this species

is distinguished through its hydrothecae that are adnate for one-third or less their adaxial length. Finally,

the gonothecae of S. divaricatus are unknown yet, but this species possesses rather highly immersed

hydrothecae (Ralph 1961).

Geographical distribution

New Caledonia (Vervoort 1993).

Symplectoscyphus ralphae Vervoort, 1993

Fig. 10M-V

Symplectoscyphus ralphae Vervoort, 1993: 270, figs 65e-f, 66a-c.

Material examined

NORFOLK 1: Stn. DW1712, two fertile colonies, ca 18 mm high, both detached from substrate (sample

A) (MNHN-IK-2012-16594); two colonies, 10 and 20 mm high, the largest bearing two gonothecae

(sample B), on bryozoan (MNHN-IK-2012-16595).

Remarks

The trophosomes of both samples are indistinguishable morphologically (compare Figs 10M and 10N;

10O and 10P; 10Q-R and 10S-T), but the gonothecae they bear are different. Those in sample A are

smaller, provided with 12-14 transverse ridges and a narrow, trumpet-shaped distal tube (Fig. 10U),

while those in sample B are comparatively larger, exhibit about 15 ridges, and their distal tube is much

wider and has an everted rim (Fig. 10V). Although the gonothecal content in sample A could not be

European Journal of Taxonomy 218 : 1-52 ( 2016 )

confidently ascertained, it may prove to be male; that in sample B contains two large ovoid bodies

(either oocytes or developing planulas), and is consequently assigned to the female sex.

Vervoort (1993) did not compare his new species with its congeners, and the present identifications are

mainly based on the shape of the gonotheca. However, other symplectoscyphid hy droids belonging to

the present collection, but not included in this study, exhibit indistinguishable trophosomes, but their

gonothecae are of different sizes and bear a varied number of transverse ridges. Consequently, it is felt that

the discrimination of species belonging to certain morphologically related groups of Symplectoscyphus

is not possible in the absence of molecular studies, owing to the absence of reliable morphological

characters.

Geographical distribution

New Caledonia (Vervoort 1993), Norfolk Ridge (present study).

Symplectoscyphus paucicatillus sp. nov.

urn:lsid:zoobank.org:act:B00A7B9A-265C-48D9-BDDB-71DB49EA7DE7

Fig. 10W-Z; Table 7

Diagnosis

Symplectoscyphus with monosiphonic stems; branching pseudodichotomous, occurring at irregular

intervals; intemodes moderately long, slender, geniculate; hydrothecae long, tubular, sigmoid in shape;

gonothecae transversely ribbed distally, undulated basally, with trumpet-shaped apical tube.

Etymology

From the Latin words pauci, -drum , meaning a few, and catillus , meaning small dish(es), making

reference to the shape of the gonotheca, which resembles several stacked dishes.

Material examined

Holotype

BATHUS 4: Stn. DW923, ca 2.8 cm high colony, bearing 3 fully formed and one incipient gonotheca

(MNHN-IK-2012-16596).

Paratypes

BATHUS 4: Stn. DW923, two sterile colonies 2 and 3 cm high (MNHN-IK-2012-16597).

Description

Colonies up to 3 cm high arising from creeping, branching hydrorhiza. Stems monosiphonic throughout,

composed of a long (up to 1.4 cm), lower, ahydrothecate part, and an upper and longest part bearing

hydrothecae. Perisarc rather thick and straw colored, thinning out gradually towards tips of colonies;

basally a little tortuous, rapidly becoming straight and smooth. Hydrothecate part divided into rather

poorly indicated internodes by means of oblique constrictions of the perisarc slanting in alternate

directions; each intemode moderately long, slender, distinctly geniculate, bearing a hydrotheca distally.

Branching pattern pseudodichotomous; side branches arising at irregular intervals from slightly below

a hydrotheca, the latter becoming axillar (Fig. 10X); structure identical to that of stem; first internode

not significantly longer compared to subsequent ones; up to 2 nd order side branches, and all in roughly

one plane, giving the colony a flabellate appearance. Hydrothecae originating at very acute angles

with the axis of corresponding internode; long, tubular, slightly sigmoid (facing outward basally and

upwards distally); adaxial wall adnate for about A its length; aperture with three pointed, triangular

cusps separated by moderately deep embayments; adaxial cusp produced adaxially; opercular apparatus

GALEA H.R., Sertulariid hydroids from the tropical western Pacific

usually lost; renovation of margin common. Gonothecae borne on both the main stem and side branches,

arising usually from below the hydrothecal bases, occasionally from the axil formed by free adaxial wall

with next internode; body elongated ovoid, provided distally with 2-3 broad, conspicuous, transverse

ribs, and undulated basally, where it tapers to an indistinct pedicel; apical part a flattened plateau from

the center of which arises a short trumpet-shaped tube; sex of gonothecae apparently female, judging

from the remains of soft tissues (presence of a spherical body, likely an oocyte).

Remarks

The hydrothecae of this species resemble those of other congeners, but there are points of difference

which allow their specific separation, as follows: 1 ) S. amoenus Vervoort &Watson, 2003 is a species

with fascicled stems, its hydrothecae are deeper (755-920 pm, compared to 590-625 in the present

material), and its gonothecae have undulated walls (Vervoort & Watson 2003); 2) S. amphoriferus

(Allman, 1877) has comparatively slender hydrothecae (judging from the size of their apertures), and its

gonothecae are pear-shaped, with “eleven raised transverse ridges and a slender terminal neck” (Millard

1977); 3) S. effusus Vervoort, 1993, but this is a comparatively smaller species (compare Fig. 10G and

10Y); 4) S. macrocarpus (Billard, 1918), has more slender [according to the size of their apertures (see

Billard 1925b; Schuchert 2003)] and shorter hydrothecae (420-470 pm); in addition, its gonothecae (of

both sexes) are characteristically amphora-shaped (Billard 1925b); 5) S. pedunculatus (Billard, 1919)

has longer, more slender, and less adnate hydrothecae (Billard 1925b; Vervoort 1993).

Geographical distribution

New Caledonia.

Discussion

Large collections of hy droids originating from the tropical western Pacific were obtained during various

expeditions of the French Tropical Deep-Sea Benthos program. Their study, entrusted to various experts,

yielded a number of reports dealing with several families of thecate hy droids (see Introduction).

The present work, focusing on hy droids of the family Sertulariidae Lamouroux, 1812, is a continuation

of the large account published by Vervoort (1993) and the recent study by Galea (2015), who introduced

three new genera, viz. Gonaxia , and Caledoniana and Solenoscyphus, respectively.

In addition to the genera Abietinaria Kirchenpauer, 1884, Caminothujaria von Campenhausen, 1896,

Dictyocladium Allman, 1888, Dynamena Lamouroux, 1812, Geminella Billard, 1925b, Hydrallmania

Hincks, 1968, Idiellana Cotton & Godfrey, 1942, Sertularella Gray, 1848, and Symplectoscyphus

Marktanner-Turneretscher, 1890 dealt with by Vervoort, members of three other genera, viz. Diphasia

L. Agassiz, 1862, Salacia Lamouroux, 1816, and Sertularia Linnaeus, 1758, are discussed herein.

Of the forty-three species, belonging to ten genera, dealt with in the present report, 9 are new to Science,

and a subspecies is raised to full species level, accompanied by a name change. Of these, two new species

of Gonaxia , added to the 21 congeners described earlier by Vervoort, raise their number to 23 within the

study area. Six and two species are added to the widely distributed and speciose genera Sertularella and

Symplectoscyphus , respectively. Additional hydroids, partly discussed earlier by Vervoort, also occur in

the present collection, and additional notes on their morphology are provided. Finally, other hydroids,

such as the poorly known Diphasia cristata Billard, 1920 and Salacia macer Vervoort & Watson, 2003,

occurred as fertile colonies, allowing descriptions of their hitherto unknown gonothecae.

As already underlined by Vervoort for the family Sertulariidae, but also by Pena Cantero & Vervoort

(2010) for the species of Acryptolaria Norman, 1875 (family Lafoeidae Hincks, 1868), the deep-water

European Journal of Taxonomy 218 : 1-52 ( 2016 )

species composition in the study area reveals a high degree of endemism. Regarding the sertulariid

hydroids, this is especially true with respect to the genera Caledoniana , Gonaxia , Sertularella and

Solenoscyphus. Among the main factors involved, the lack of a free-swimming dispersive stage (Pena

Cantero & Vervoort 2010) limits the capabilities of an efficient dispersal, causing a marked reproductive

isolation and, consequently, a high degree of speciation.

Although the hy droid fauna of the study area can be considered as relatively well investigated, it is likely

that, with future sampling, new genera and species will be discovered.

Acknowledgements

Thanks are due to Philippe Bouchet and Pierre Lozouet, both of MNHN, for giving me access to the

material described in this paper, and to Aude Andouche for curatorial assistance. The acknowledgements

to cruises senior scientists, collectors, captains and crew in Bouchet et al. (2008) also apply here. I am

grateful to Dr. Dale Calder (Royal Ontario Museum, Toronto, Canada) for his helpful advice on ICZN

matters. Sincere thanks are due to Drs Peter Schuchert (MHNG, Switzerland) and Henry H.C. Choong

(Fairbanks Museum & Planetarium, USA) for constructive comments on an earlier version of this report.

This study was undertaken with no financial support.

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Manuscript received: 22 January 2016

Manuscript accepted: 21 March 2016

Published on: 2 August 2016

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands.