European Journal of Taxonomy 24: 1-42

http://dx.doi.org/10.5852/ejt.2012.24

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ISSN 2118-9773

WWW. europeanj oumaloftaxonomy. eu

2012 • Daniel Ledue

Research article

um:lsid:zoobank.org:pub:F8ED2AA9-83Cl-4CB8-8327-58C501B6C42A

Deep-sea nematodes (Comesomatidae) from the Southwest Pacific

Ocean: five new species and three new species records

Daniel LEDUC

Department of Marine Seienee, University of Otago, P.O. Box 56, Dunedin, New Zealand

National Institute of Water and Atmospherie Researeh (NIWA) Eimited,

Private Bag 14-901, Kilbimie, Wellington, New Zealand

Email: daniel.leduc@gmai1 com

um:lsid:zoobank.org:author:9393949F-3426-4EE2-8BDE-DEFFACE3D9BC

Abstract. The present study deseribes five new free-living nematode speeies and provides three

new speeies reeords of the family Comesomatidae (genera Cervonema Wieser, 1954, Dorylaimopsis

Ditlevsen, 1918, Hopperia Vitiello, 1969, and Kenyanema Muthumbi et al, 1997) from the eontinental

margin of New Zealand, Southwest Paeifie. Diehotomous identifieation keys are provided for all known

speeies of Dorylaimopsis and Hopperia. Cervonema shiae Chen & Vinex, 2000 is reeorded for the

first time outside the type loeality (Beagle Channel, Chile). C. kaikouraensis sp. nov. is eharaeterised

by amphideal fovea with 5.5 turns situated at 1.7 head diameter from anterior end, jointed outer labial

setae, equal in length to eephalie setae, sperm dimorphism, and 5-6 small pre-eloaeal supplements.

C. multispira sp. nov. is eharaeterised by amphideal fovea with 8.0-8.5 turns situated at 2.6-4.0 head

diameter from anterior end, eephalie setae 2-3 pm long, slightly shorter than outer labial setae, presenee

of six uninueleated eells in males (potentially pseudoeoelomoeytes or supplementary exeretory eells), 5

small pre-eloaeal supplements, and strongly eutieularised, areuate spieules with eapitulum. C. proberti

sp. nov. is eharaeterised by amphideal fovea with 5 turns and loeated at < 1 head diameter from anterior

end, eephalie setae 1.6-2.0 times longer than outer labial setae, and 8 small pre-eloaeal supplements.

Dorylaimopsis nodderi sp. nov. is eharaeterised by eutiele with lateral differentiation eonsisting of three

longitudinal rows of larger dots in the pharyngeal and eaudal regions, two rows of larger dots in middle

region of body, and spieules with rounded ventral projeetion at one third of spieule length from distal

end, giving appearanee of a joint. Hopperia ancora sp. nov. is eharaeterised by short eonieal eephalie

setae, spieules with hook-like projeetion at distal end, gubemaeulum with bent apophyses, and 11-13

pre-eloaeal supplements. H. beaglense Chen & Vinex, 1998 is reeorded from Kaikoura Canyon, the

first reeord of this speeies outside the type loeality (Beagle Channel, Chile). Kenyanema monorchis

Muthumbi et al, 1997 is also reeorded for the first time outside the type loeality (Indian Oeean).

Keywords. Cervonema, Dorylaimopsis, Hopperia, Kenyanema, New Zealand.

Leduc D. 2012. Deep-sea nematodes (Comesomatidae) from the Southwest Pacific Ocean: five new species and

three new species records. European Journal of Taxonomy 24: 1-42. http://dx.doi.org/10.5852/ejt.2012.24

1

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Introduction

Nematode speeies of the family Comesomatidae Filipjev, 1918 are co mm on on eontinental margins

worldwide (Soetaert & Heip 1995; Vanhove et al. 1999; Muthumbi et al 2011). Little is known,

however, about the diversity and distribution of deep-sea (>200 m water depth) Comesomatidae in the

Southwest Paeifie (Miljutin et al. 2010). Two new speeies of this family were reeently deseribed from

the eontinental slope of New Zealand (Ledue et al 2012a). The present study deseribes five new speeies

and provides three new speeies reeord belonging to the genera Cervonema Wieser, 1954, Dorylaimopsis

Ditlevsen, 1918, Hopperia Vitiello, 1969, and Kenyanema Muthumbi et al, 1997 from the same region.

Material and Methods

Sampling and laboratory methods

The present study foeussed on three major bathymetrie features of the New Zealand Exelusive Eeonomie

Zone (EEZ): Challenger Plateau, Chatham Rise and Kaikoura Canyon. Challenger Plateau eneompasses

water depths ranging from c. 400 to 3000 m in an area of generally lowbiologieal produetivity to the northwest

of the South Island, New Zealand (Murphy et al. 2001). Chatham Rise is a submarine ridge that extends

eastwards from the South Island of New Zealand, over water depths ranging from c. 250 to 3000 m. The rise

lies beneath the Subtropieal Front (STF), a region assoeiated with heightened primary produetivity (Bradford-

Grieve et al 1997; Murphy et al 2001). Kaikoura Canyon lies to the east of the South Island (~42°S), and has

been deseribed as one of the most produetive deep-sea benthie habitats known (De Eeo et al 2010).

Details of the study sites are given in Eedue et al. (2012a, b). Samples were eolleeted along a transeet

at 178°30'E aeross Chatham Rise (nine sites, 350-3100 m water depth) in austral spring (September-

Oetober) 2001 during National Institute of Water and Atmospherie Researeh (NIWA) eruise TAN0116.

Twenty-three additional sites between 240 and 1300 m water depths on Chatham Rise and Challenger

Plateau were sampled in austral autumn-early winter (Mareh-April and May-June) 2007 (NIWA eruises

TAN0705 and TAN0707, respeetively). Five sites (405-1420 m water depth) were sampled along the

Kaikoura Canyon axis in May 2010 during NIWA eruise TAN1006.

Sediment samples were eolleeted using an Oeean Instruments MC-800A multieorer (eore internal

diameter = 9.5 em). Eaeh sample eonsisted of one subeore of internal diameter 26 mm taken to a

depth of 5 em. Samples were fixed in 10% formalin and stained with Rose Bengal. Samples were

subsequently rinsed on a 1 mm sieve to remove large partieles and on a 45 pm sieve to retain nematodes.

Nematodes were extraeted from the remaining sediments by Eudox dotation and transferred to pure

glyeerol (Somerfield & Warwiek 1996). Speeies deseriptions were made from glyeerol mounts using

differential interferenee eontrast mieroseopy and drawings were made with the aid of a eamera lueida.

All measurements are in pm, and all eurved struetures are measured along the are. Type speeimens

are held in the NIWA Invertebrate Colleetion (NIC), Wellington. Anterior sense organs are defined as

follows: papillae (<1 pm long), setiform papillae (1-2 pm), and setae (>2 pm) (Eedue & Wharton 2008).

Abbreviations

a

abd

b

e

ebd

hd

E

n

%V

body length/maximum body diameter

anal body diameter

body length/pharynx length

body length/tail length

eorresponding body diameter

head diameter

body length

number of speeimens

vulva distanee from anterior end of body x 100/total body length

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LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Results

Class Chromadorea Inglis, 1983

Order Araeolaimida De Coninck, 1965

Superfamily Axonolaimoidea F ilipj ev, 1918

Family Comesomatidae Filipjev, 1918

Subfamily Sabatieriinae Filipjev, 1934

Diagnosis (modified from Jensen (1979))

Cephalie sense organs in three separate erowns, with outer labial sensillae and eephalie setae sometimes

elose together. Bueeal eavity weakly eutieularised, anterior portion eup-shaped, sometimes very

small; posterior portion narrow, not strongly eutieularised. Spieules bent, usually enlarged proximally.

Gubemaeular apophyses (when present) usually direeted dorsoeaudally and paired.

Type genus

Sabatieria De Rouville, 1903

Genus Cervonema Wieser, 1954

Diagnosis (modified from Chen & Vincx (2000))

Sabatieriinae. Body eutiele striated, without punetations; eervieal region elongated or short. Six external

labial sensillae setiform, usually almost equal in length to four eephalie setae. Amphideal fovea with 4-8

turns. Bueeal eavity small, without teeth. Spieules simple, gubemaeulum small or absent, minute pre-

eloaeal supplements present or absent.

Type species

Cervonema allometricum Wieser, 1954

Remarks

Interspeeifie variation in the morphologieal eharaeters of Cervonema speeies is relatively limited, and

speeies within this genus should be identified using as many eharaeters as possible (e.g., Barnes et al.

2012). The following eharaeters, however, are partieularly useful in speeies identifieation: relative length

of outer labial setae and eephalie setae, shape and position of amphids, shape and size of spieules, presenee/

absenee and shape of gubemaeulum, and tail length. Other traits, whieh have not always been ineluded in

earlier speeies deseriptions, inelude presenee/absenee of jointed outer labial setae and presenee /absenee

of sperm dimorphism. Pre-eloaeal supplements are often diffieult to observe using light mieroseopy and

should not be used in isolation for defining speeies.

Cervonema shiae Chen & Vinex, 2000

Figs 1-2, Table 1

Material examined

1 S and 25 ?, eolleeted 6 Apr. 2007, southern fiank of Chatham Rise (1240 m water depth), 44.4864°S,

177.1407°E. Charaeteristies of surfaee sediment layer (0-5 mm): mostly silt/elay (82.9%), with very fine

sand (10.2%), and fine sand (8.4%); ealeium earbonate eontent: 38.4%; total organie matter eontent:

1.9%; ehloroplastie pigment eoneentration: 3501 ng/gEW^^^^j^^j^^ (NIC 84438). 1 S eolleeted 5 Jun. 2007,

Challenger Plateau (532 m water depth), 40.8814'S, 170.8603'E. Charaeteristies of surfaee sediment

layer (0-5 mm): mostly silt/elay (67.6%), with fine sand (15.2%), and very fine sand (11.4%); ealeium

earbonate eontent: 56.0%; total organie matter eontent: 2.1%; ehloroplastie pigment eoneentration: 1344

(MC 84439).

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European Journal of Taxonomy 24 : 1-42 ( 2012 )

Fig. 1. Cervonema shiae Chen & Vincx, 2000. A. S anterior body region. B. Lateral view of S head

region. C. $ reproduetive system. D. $ tail. E. S posterior body region showing reproduetive system

and eopulatory apparatus. Seale bar: A = 40 pm; B = 35 pm; C = 65 pm; D = 30 pm; E = 60 pm.

4

LEDUC D., Deep-sea nematodes from the Southwest Pacific Ocean

Fig. 2. Cervonema shiae Chen & Vincx, 2000 and Cervonema kaikouraensis sp. nov. light micrographs.

A. Large sperm cells without nuclei in anterior testis of C. shiae. B. Small nucleated sperm in posterior

testis of C. shiae. C. Large sperm cells without nuclei in anterior testis of C. kaikouraensis sp. nov.

D. Small nucleated sperm in posterior testis of C. kaikouraensis sp. nov. Arrows point to the lenticular

nuclei situated on the periphery of the small sperm cells in the posterior testes of both species. Scale

bar =10 pm.

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European Journal of Taxonomy 24 : 1-42 ( 2012 )

Table 1. Morphometries (pm) of Cervonema shiae Chen & Vinex, 2000 and C. kaikouraensis sp. nov.

Species

Cervonema shiae Chen & Vinex, 2000

Cervonema kaikouraensis sp. nov.

Males

Females

Male Holotype

Female Paratypes

n

2

L

744-827

693-718

1639

1876-1959

a

30-31

21-24

27

18-24

b

5

7

7-8

c

6-7

7

8

6-7

Head diam.

7-8

11

12-13

Mouth diam.

2

3

Length of cephalic setae

2-3

6

Amphid height

10

7

13

10-11

Amphid width

9

7-8

12

9-12

Amphid width/cbd (%)

75-82

63-73

67

41-57

Amphid from anterior end

16-20

16

19

22-23

Nerve ring from anterior end

73-79

70-77

117

114

Nerve ring cbd

21

19-20

44

53-54

Excretory pore from anterior end

82-96

81-89

129

135-137

Pharynx length

150-158

135-143

240

255-256

Pharynx cbd

23-24

23-26

56

69-74

Pharyngeal bulb diam.

14

16-19

37

50-55

Max. body diam.

25-27

30-33

60

83-104

Spicule length

11-12

-

32

-

Gubernacular apophyses length

-

Anal body diam.

18-22

18-21

36

46-52

Tail length

115-122

96-97

198

274-302

Tail length/abd

5.5-6.4

4.6-5.3

5.5

5.8-6.0

V

-

344-389

-

934-955

%V

-

50-54

-

49-50

Vulval body diam.

-

28-33

-

80-93

Description

Males

Body eylindrieal, tapering towards both extremities but more pronouneed in tail region. Cutiele faintly

striated, no lateral differentiation. Somatie setae short and sparse. Head not set-off by eonstrietion. Inner

labial papillae indistinet, six outer labial setae, and four slightly longer eephalie setae, the latter two in

separate eireles elose to eaeh other. Amphideal fovea spiral, at 6.0-6.5 turns, 1.8-2.3 head diameters

from anterior. Small bueeal eavity, eup-shaped. Pharynx widening posteriorly into a weakly marked

elongated bulb. Cardia short. Nerve ring near middle of pharynx length. Seeretory-exeretory pore near

two thirds of pharynx length from anterior. Cellular body of ventral gland small- at level of eardia.

Intestine wall eonsisting mostly of eells eontaining eolourless granules, interspersed with elusters of

elear globular inelusions.

Reproduetive system diorehie, opposed, outstretehed. Anterior testis to left of intestine, posterior testis

to right of intestine. Anterior testis with relatively large oval sperm eells (5-8 pm wide by 12-15 pm

long) without nuelei (Fig. 2A). Posterior testis with smaller, oval-shaped sperm eells (up to 8x5 pm) with

6

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

lenticular nuclei situated peripherally (Fig. 2B). Spicules paired, equal, 0.5-0.6 abd long, with lightly

cuticularised, swollen proximal portion, and heavily cuticularised, pointed distal portion. Gubemaculum

not observed. Large rectal gland surrounds spicules. Four small, tubular pre-cloacal supplements, one

pre-cloacal seta. Tail conico-cylindrical, with several short caudal setae and three short terminal setae.

Three caudal glands and spinneret.

Females

Similar to males, but with slightly larger maximum body diameter, amphideal fovea smaller, 5.0-5.5

turns. Reproductive system didelphic, opposed, ovaries outstretched, with anterior branch to left of

intestine and posterior branch to right of intestine. Both types of sperm observed in uterus. Spermathecae

not observed. Vulva at median or slightly post median. Granular vaginal glands present, pars proximalis

vaginae surrounded by constrictor muscle.

Discussion

My specimens closely resemble the description of Chen & Vincx (2000) based on specimens from

the Beagle Channel (255-257 m water depth), although specimens from Chatham Rise and Challenger

plateau have slightly shorter cephalic setae (0.3-0.4 vs. 0.5 cbd) and longer tail (5.5-6.4 vs. 4.8 abd).

The specimens are also characterised by sperm dimorphism (anterior testis with large oval sperm cells

without nuclei, posterior testis with smaller nucleated sperm cells); Chen & Vincx (2000), however, did

not provide details of the sperm in their description. This is the first record of this species outside the

type locality.

Cervonema kaikouraensis sp. nov.

um:lsid:zoobank.org:act:00FCA18B-AB7B-4464-8DBC-A75F17F3BC72

Figs 2-4, Table 1

Etymology

This species is named after the type locality.

Material examined

Holotype

S, collected 5 May 2010, Kaikoura Canyon axis (1061 m water depth), 42.5081°S, 173.6325°E (NIC

84440).

Paratypes

2 $ same data as holotype (NIC 84441).

Description

Males

Body cylindrical, tapering towards both extremities. Cuticle faintly striated, no lateral differentiation.

Somatic setae very short and sparse, 1-2 pm long, except in pre-cloacal region, 4 pm long. Head not set¬

off by constriction. Six inner labial papillae, six jointed outer labial setae, and four cephalic setae, not

jointed. Amphideal fovea spiral, 5.5 turns, at 1.7 head diameters from anterior. Golden-coloured granular

material in lateral, ventral, and dorsal chords, forming two continuous bands (Fig. 4D). Small buccal

cavity, cup-shaped. Pharynx gradually widening posteriorly into an elongated bulb. Dorsal pharyngeal

gland nucleus conspicuous. Cardia short. Nerve ring near middle of pharynx length. Secretory-excretory

pore situated just posteriorly to nerve ring. Cellular body of ventral gland small, at level of cardia.

Intestine wall with numerous granules, orange-brown in anterior portion, colourless in posterior portion.

7

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Fig. 3. Cervonema kaikouraensis sp. nov. A. S anterior body region. B. $ head. C. S reproduetive

system. D. Entire $. E. (? eopulatory apparatus. Seale bar: A = 50 pm; B = 30 pm; C = 170 pm;

D = 200 pm; E = 40 pm.

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LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Reproductive system diorchic, opposed, outstretched. Anterior testis to left of intestine, posterior testis to

right of intestine. Anterior testis with relatively large elongated sperm cells (up to 5 pm wide and 20 pm

long), without nuclei (Fig. 2C). Posterior testis with smaller, globular sperm cells (4-6 pm diameter)

with lenticular nuclei situated peripherally (Fig. 2D). Spicules paired, equal, narrow, straight, 0.9 abd

long, with small capitulum. Rectal gland present. Five, possibly six tubular pre-cloacal supplements,

difficult to observe, one pre-cloacal seta. Tail conico-cylindrical, with several caudal setae, 2-5 pm long,

and three short terminal setae. Three caudal glands and well-developed spinneret (Fig. 4C).

Fig. 4. Cervonema kaikouraensis sp. nov. light micrographs. A. Anterior ovary, showing bent germinal

portion. B. S head, showing jointed outer labial seta. C. S spinneret. D. Granules in the lateral chords

of S- Scale bar: A = 40 pm; B = 18 pm; C = 12 pm; D = 24 pm.

9

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Females

Similar to males, but with slightly larger maximum body diameter, amphideal fovea smaller, 4.5 turns.

Reproduetive system didelphie, opposed, with anterior braneh to left of intestine and posterior braneh

to right of intestine. Terminal (i.e., distal) portion of anterior ovary bent in one paratype speeimen

(Fig. 4A), but not other paratype speeimen; germinal portion of posterior ovary bent in both paratypes.

Both types of sperm observed in uterus and spermatheeae. Vulva at body median. Granular vaginal

glands present, pars proximalis vaginae surrounded by eonstrietor musele.

Diagnosis and relationships

Cervonema kaikouraensis sp. nov. is eharaeterised by body length 1636 pm, amphid with 5.5 turns

situated 1.7 head diameter from anterior end, jointed outer labial setae, equal in length to eephalie setae,

anterior testis with large elongated sperm eells without nuelei, posterior testis with smaller nueleated

sperm eells, 5 small pre-eloaeal supplements, straight spieules 32 pm in length, and tail 5.5 abd.

C. kaikouraensis sp. nov. ean be differentiated from all other speeies of the genus, exeept C. allometricum

Wieser, 1954 and C. pseudodeltensis Barnes et al. , 2012 by the presenee of jointed outer labial setae. The

presenee of jointed outer labial setae in C. allometricum was not deseribed in the original deseription of

Wieser (1954) but was later noted by Lorenzen (1981). C. kaikouraensis sp. nov. ean be distinguished

from C. allometricum and C. pseudodeltensis by the absenee of gubemaeulum and gubemaeular

apophyses (gubemaeulum with eonspieuous posterior apophyses present in C. allometricum and C.

pse udode I tensis).

The presenee of jointed outer labial setae may have been overlooked in previous deseriptions and may

not be a reliable trait to use for differentiating between all speeies of Cervonema. C. kaikouraensis

sp. nov. ean further be differentiated from most other speeies of the genus (exeept C. del tensis Hope &

Zhang, 1995 and C. papillatum Jensen, 1988 by the presenee of sperm dimorphism. C. kaikouraensis

sp. nov. differs from C. deltensis in body length (1636 vs. 1201-1237), length of eephalie setae (6 vs.

3-5 pm), amphideal fovea turns (5.5 vs. 3-5), position of amphid (1.7 vs. 1.4 hd from anterior), and

absenee of gubemaeulum (plate-like gubemaeulum in C. deltensis). C. kaikouraensis sp. nov. differs

from C. papillatum in body length (1636 vs. 1140-1230), and by the absenee of a weakly eutieularised

eap surrounding the spieules proximally. The presenee or absenee of sperm dimorphism was not noted

in the deseriptions of C. brevicauda Gourbault, 1980 and C.jenseni Gourbault, 1980, but these speeies

differ most notably from C. kaikouraensis sp. nov. in the presenee of a gubemaeulum (absent in C.

kaikouraensis sp. nov.). C. brevicauda also has a shorter body length (800-1090 vs. 1636), shorter

eephalie setae (3 vs. 6 pm), and shorter spieules (0.6 vs. 0.9 abd) than C. kaikouraensis sp. nov. C.

jenseni ean also be differentiated from C. kakouraensis sp. nov. by the greater number of amphid turns

(7 vs. 5.5) and absenee of supplements (5 in C. kaikouraensis sp. nov.).

Remarks

The presenee of bent ovaries, as observed in C. kaikouraensis sp. nov., is unusual for the family

Comesomatidae. The ovaries, however, are bent only in the terminal portion. Beeause the ovaries in both

female speeimens are quite large and oeeupy most of the available spaee in the pseudoeoel (Fig. 3D),

this trait is eonsidered to be a seeondary feature resulting from laek of spaee, rather than tme reflexed

ovaries.

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LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Cervonema multispira sp. nov.

um:lsid:zoobank.org:act:D4A82AED-213B-4CE4-9D62-FAAC2D44CBCQ

Figs 5-6, Table 2

Etymology

The speeies name is derived from the Eatin multus (= mueh) and spiralis (= eoil), and refers to the shape

of the amphid.

Material examined

Holotype

S, eolleeted 6 Apr. 2007, southern flank of Chatham Rise (1240 m water depth), 44.4864°S, 111. 1407°E.

Charaeteristies of surfaee sediment layer (0-5 mm): mostly silt/elay (82.9%), with very flne sand

(10.2%), and flne sand (8.4%); ealeium earbonate eontent: 38.4%; total organie matter eontent: 1.9%;

ehloroplastie pigment eoneentration: 3501 ng/gOW^^^^^^^^ (NIC 84444).

Paratypes

1 ? ?, same data as holotype (NIC 84445).

Description

Males

Body eylindrieal, tapering towards both extremities, orange-brown eolouration throughout. Cutiele

faintly striated, no lateral differentiation. Somatie setae short and sparse. Head set-off by a depression

at level of eephalie setae. Inner labial papillae indistinet, six outer labial setae, and four slightly shorter

eephalie setae, the latter two in separate eireles. Amphideal fovea spiral, 8.0-8.5 turns, situated 2.6-

4.0 head diameters from anterior. Small bueeal eavity, eup-shaped. Pharynx with small posterior bulb.

Cardia short. Nerve ring near 60% of pharynx length from anterior. Seeretory-exeretory pore slightly

posterior to nerve ring. Cellular body of ventral gland small, posterior to eardia. Six large uninueleated

pseudoeoelomoeytes, 23 pm wide, 35-40 pm long, in anterior half of body, with 2 and 4 eells on either

side of intestine in holotype, 3 eells on either side of intestine in paratype, eaeh with single extension (or

possibly duet) direeted anteriorly (Fig. 6A). Intestine wall eonsisting mostly of eells eontaining small

granules, interspersed with elusters of elear globular inelusions

Reproduetive system diorehie, opposed, outstretehed. Anterior testis to left of intestine, posterior

testis to right of intestine. Sperm eells small (up to 5x4 pm), globular, with lentieular nuelei situated

peripherally. Spieules paired, equal, areuate, 0.9-1.1 abdlong, strongly eutieularised along entire length,

with eapitulum. Four pairs of small ejaeulatory glands situated anteriorly to spieules (Fig. 5C). Small

reetal gland present. Gubemaeulum not observed. Five small, tubular pre-eloaeal supplements, one pre-

eloaeal seta. Tail eonieo-eylindrieal, with several short eaudal setae and three short terminal setae. Three

eaudal glands and spinneret present.

Females

Similar to males, but with smaller amphideal fovea, 7.5 turns. Farge nueleated eells in anterior part

of body not observed. Reproduetive system didelphie, opposed, outstretehed, with anterior braneh

to left of intestine and posterior braneh to right of intestine. Spermatheea present, eontaining sperm.

Vulva slightly post median. Granular vaginal glands present, pars proximalis vaginae surrounded by

eonstrietor musele.

11

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Fig. 5. Cervonema multispira sp. nov. A. S anterior body region. B. $ head. C. S posterior body region,

showing eopulatory apparatus. D. $ intestine. E. $ tail. Arrows point to the four ejaeulatory glands.

Seale bar: A =30 pm; B = 20 pm; C = 40 pm; D = 28 pm; E = 25 pm.

12

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Diagnosis and relationships

Cervonema multispira sp. nov. is characterised by body length 1225-1431 pm, amphideal fovea with

8 .0-8.5 turns, 2.6-4.0 head diameter from anterior end, eephalie setae 2-3 pm long, slightly shorter than

outer labial setae, presenee of six large lateral eells in males only, 5 small pre-eloaeal supplements,

strongly eutieularised, areuate spieules with eapitulum, and tail 4.4-6.1 abd.

Cervonema multispira sp. nov. resembles C. macramphis Jensen, 1979 in body size, length of eephalie

setae, amphid with numerous (> 7) turns, and the presenee of areuate spieules. Cervonema multispira

sp. nov. ean be differentiated from C. macramphis and all other speeies of the genus by amphideal fovea

with 8.0-8.5 turns (< 7.5 turns in all other speeies), presenee of lateral exeretory eells in males, and

strongly eutieularised spieules with eapitulum. C. multipsira sp. nov. also resembles C. pseudodeltensis

Barnes et al, 2012 in having outer labial setae marginally longer than eephalie setae, but is eharaeterised

by shorter body length (1225-1431 vs. 1559-1755 pm), amphideal fovea with more turns (8.0-8.5 vs.

5.5) and absenee of gubemaeulum (present in C. pseudodeltensis).

Discussion

To my knowledge, this is the first time that the presenee of large uninueleated eells is noted in the

pseudoeoelom of Cervonema speeimens. Similar eells, usually situated laterally, and sometimes

subventrally, have been deseribed in Sabatieria celtica Southern, 1914 (Riemann 1977), S. pulchra

(Sehneider, 1906) (Jensen 1979), S. breviseta Stekhoven, 1935, and S. punctata (Kreis, 1924) (Platt

Fig. 6. Cervonema multispira sp. nov. A. Entire (J. B. $ reproduetive system. Seale bar: A =100 pm;

B = 70 pm.

13

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Table 2. Morphometries (pm) of Cervonema multipsira sp. nov. and C. proberti sp. nov. n/o = not

observed.

Species

Cervonema multispira s

). nov.

Cervonema proberti sp. nov.

Males

Females

Males

Females

Holotype

Paratype

Paratypes

Holotype

Paratype

Paratypes

n

1

2

1

3

L

1431

1225

965-1083

1144

1043

1183-1397

a

28

34

24-29

30

33

33-35

b

7

6-7

9

9-10

c

9

7

7-13

9

8

Head diam.

10

9

8

12

10

11-12

Mouth diam.

2

1

<1

1-2

Length of cephalic setae

3

2

7

7-8

8-9

Amphid height

15

14

10-11

13

14

10

Amphid width

16

14

9-10

11

12

9-10

Amphid width/cbd (%)

76

82

69-71

85

86

69-71

Amphid from anterior end

40

23

19-22

6

10

9-10

Nerve ring from anterior end

125

108

90-108

68

64

65-67

Nerve ring cbd

38

28

25-27

28

25

28-29

Excretory pore from anterior end

147

123

113-126

n/o

87-90

Pharynx length

214

183

151-185

129

120

131-144

Pharynx cbd

47

35

30-32

34

28

32-35

Pharyngeal bulb diam.

26

20

19-20

26

18

25-27

Max. body diam.

51

36

40-42

38

32

36-42

Spicule length

36

31

-

20

22

-

Gubernacular apophyses length

-

Anal body diam.

38

29

26-27

27

24

26-30

Tail length

167

176

146-147

122

116

142-178

Tail length/abd

4.4

6.1

5.4-5.6

4.5

4.8

5.5-5.9

V

-

488-628

-

556-644

%V

-

51-52

-

46-47

Vulval body diam.

-

38-42

-

36-42

1984). Only two eells (deseribed as supplementary exeretory eells) are found in Sabatieria spp., whereas

six were observed in Cervonema multispira sp. nov. This type of eell has not been observed in females

of Sabatieria spp. or C. multispira sp. nov.

Cervonema proberti sp. nov.

um:lsid:zoobank.org:aet:Q635B2CB-1114-4399-A3E6-DCQF53832755

Figs 7-8, Table 2

Etymology

This speeies is named after Dr. P. Keith Probert.

14

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Material examined

Holotype

6 ', collectedS Oct. 2001,ChathamRisecrest(350mwaterdepth),43.4333°S, 178.5000°E. Characteristics

of surface sediment layer (0-5 mm): mostly fine sand (35.7%), silt/elay (33.0%), and very fine sand

(23.5%); ealeium earbonate eontent: 40.5%; total organie matter eontent: 4.5%; ehloroplastie pigment

eoneentration: 3359 ng/gOW^^^^^^^^ (NIC 84442).

Paratypes

1 (?, 3 $ same data as holotype (NIC 84443).

Description

Males

Body eylindrieal, tapering towards both extremities. Cutiele faintly striated, no lateral differentiation.

Somatie setae short and sparse, in four dorso- and ventro-lateral longitudinal rows. Epidermal glands

assoeiated with somatie setae visible as elear, oval areas underneath eutiele (Figs 7C, 8A). Head set-off

by eonstrietion at level of amphids. Inner labial papillae indistinet, six outer labial setae, and four longer

eephalie setae, the latter two almost at same level. Cephalie setae 1.6-2.0 times longer than outer labial

setae. Amphideal fovea spiral, 5 turns, 0.5-1.0 head diameter from anterior. Small bueeal eavity, eup-

shaped. Pharynx with rounded or oval terminal bulb. Nueleus of dorsal pharyngeal gland eonspieuous.

Cardia short. Nerve ring near middle of pharynx length. Seeretory-exeretory pore slightly posterior to

nerve ring. Cellular body of ventral gland slightly posterior to eardia. Intestine wall eonsisting mostly

of eells eontaining eolourless granules, interspersed with elusters of elear globular inelusions (Fig. 8B).

Reproduetive system diorehie, opposed, outstretehed. Anterior testis to left of intestine, posterior testis

to right of intestine. Oval-shaped sperm eells with lentieular nuelei situated peripherally (Fig. 7D). Two

pairs of ejaeulatory glands present. Spieules paired, equal, slightly areuate, 0.7-0.9 abd long, proximal

end slightly eephalated, gradually tapering towards distal end. Gubemaeulum not observed. Reetal gland

surrounds spieules. Eight small, tubular pre-eloaeal supplements, often diffieult to distinguish, one pre-

eloaeal seta. Tail eonieo-eylindrieal, with several short eaudal setae and three short terminal setae. Three

eaudal glands and spinneret present.

Females

Similar to males, but with slightly larger maximum body diameter, amphideal fovea smaller, 4-5 turns,

slightly longer tail. Reproduetive system didelphie, opposed, outstretehed, with anterior braneh to left

of intestine and posterior braneh to right of intestine. Vulva slightly pre median. Granular vaginal glands

present, pars proximalis vaginae surrounded by eonstrietor musele.

Diagnosis and relationships

Cervonema proberti sp. nov. is eharaeterised by body length 1043-1144 pm, amphideal fovea with 5

turns and < 1 head diameter from anterior end, eephalie setae 1.6-2.0 times longer than outer labial setae,

sperm eells with lentieular nuelei, 8 small pre-eloaeal supplements, and tail 4.5-5.9 abd.

Cervonema proberti sp. nov. ean be distinguished from most other speeies of the genus by the amphid

< 1 head diameter from anterior, eompared to > 1 head diameter from anterior in most speeies (exeept

C. hermani Chen & Vinex, 2000, C. chilensis Chen & Vinex, 2000, and C. proximamphidum Tehesunov,

2000). C. proberti sp. nov. ean be distinguished from C. hermani, C. chilensis, and C. proximamphidum

by eephalie setae eonspieuously (1.6-2.Ox) longer than outer labial setae (almost equal in length in C.

hermani, C. chilensis, and C. proximamphidum), and nueleated sperm eells (no nuelei in C. hermani, C.

15

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Fig. 7. Cervonemaproberti sp. nov. A. S anterior body region. B. $ anterior region. C. $ tail. D. Sperm

eells. E. S posterior body region, showing eopulatory apparatus. Seale bar: A, B, and C = 25 pm;

D = 20 pm; E = 35 pm.

16

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

chilensis, and C. proximamphidum). C. proberti sp. nov. is also characterised by a shorter tail than C.

hermani (4.5-4.8 vs. 5.7-9.3 abd). C. proberti sp. nov. resembles Cervonema sp. B deseribed by Barnes

et al (2012) based on one female speeimen from the Korean west eoast. My female speeimens share

several traits, ineluding amphid elose to anterior end, well-developed pharyngeal bulb with eonspieuous

pharyngeal gland nueleus, and eephalie setae longer than the outer labial setae. My speeimens, however,

have shorter body length (1183-1397 vs. 1727 pm), and greater ratio of eephalie setae length to outer

labial setae length (1.6-2.0 vs. 1.3). Male speeimens are required before the identity of the Korean

speeies ean be determined with eertainty.

Fig. 8. Cervonema proberti sp. nov. A. $ head, showing glands. B. Entire S- C. $ reproduetive system.

Arrows point to elear globular inelusions in intestine wall. Seale bar: A =20 pm; B and C = 65 pm.

17

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Table 3. Male morphological characters of all known Cervonema species.

Tail

length

abd

5.5-6.0

3.2-4.0

5.0

5.0-6.0

o

l

6.0-6.3

5.7-9.3

o

00

1

o

5.0

3.0-4.1

5.6-6.4

o

in'

1

kO

1

00

6.0-7.0

5.5

4.4-6.1

4.5-4.8

Spicule

length

abd

o

90

o

V

0.8-0.9

60

o

1.4-1.5

0.7

o

1

00

o'

o

1

00

o'

0.8-0.9

o

60

09-1.1

0.7-0.9

Guber-

naculum

presence

Yes

No

Yes

No

Yes

No

Yes

No

Yes

No

Yes

No

# Supple¬

ments

NS

o

03

o

6-7

9-10

4-5

00

1

in

00

Sperm

dimor¬

phism

NS

No

Yes

No

NS

No

C/5

p-

C/5

No

NS

No

Yes

No

Amphid

from ante¬

rior (hd)

1.5-2.0

00

0.75

>2

0.75

2.0

>3.0

2.0

1.4-1.7

00

o'

2.5-3.0

2.0

2.6-4.0

0.5-1.0

Amphid

turns

4.5-5.0

5.5-6.0

5.5

3-5

m

5.5

7.0

4-5

5.5

5-6

5.0-6.5

5.5-6.0

5.5

8.0-8.5

5.0

Jointed

labial

setae

No

No2

No

Yes

No

Yes

No

Ratio of

cephalic

setae/outer

labial setae

length

o

0.6-0.8

o

cn

o

0.75-1.0

1.6-2.0

Cephalic

setae length

(mm(hd))

1

00

o'

03

3 (0.3-0.4)

7 (0.8)

3-5 (0.4-0.5)

4-5 (0.4)

4 (0.4)

8-10(0.8)

3 (0.3)

2 (0.4)

3 (0.3)

3-4 (0.4)

5.0-5.5 (0.6)

4 (0.3-0.5)

•rT

o'

(9 0) 9

2-3 (0.2-0.3)

7-8 (0.6-0.8)

Body length

(pm)

1060

800-1090

1192

1201-1237

1225-1470

1440-3458

1577-1641

1309

750-833

1140-1230

1559-1755

1423-1563

612-749

820-1378

1636

1225-1431

1043-1144

Species

C. allometricum Wieser, 1954

C. brevicauda Gourbault, 1980

C. chilensis Chen & Vincx, 2000

C. deltensis Hope & Zhang, 1995

C. gourbaulti Muthumbi et aL, 1997

C. jenseni Gourbault, 1980

C. hermani Chen & Vincx, 2000

C. macramphis Jensen, 1979

C. minutus Muthumbi et aL, 1997

C. papillatum Jensen, 1988

C. pseudodeltensis Barnes et aL, 2012

C. proximamphidum Tchesunov, 2000

C. shiae Chen and Vincx, 2000

C. tenuicauda (Stekhoven, 1950)

C. kaikouraensis sp. nov.

C. multispira sp. nov.

C. proberti sp. nov.

bX)

§

N

<D

Ph

o

X

<D

'T3

<D

O

P

o

p

<D

c\

<D

Cb

<D

c/3

• ^

I

<D

O

^ 'P

P O

<D •)

N

P o

<D

O

<D

O

P

c/3

,P CD

't3 cl

CD ^

_C3

o <1^

c/3 OX)

w bx)

PS C5

.(_1 c/3

§

.1 a

Ph

• ^

O

c/3

<D

'P

P

• ^

bX)

• ^

O

'P

<D .

Cp

• ^

O

<D

^ —

cy3

^ CD

S 2

O

c/3

<D

'P

P

• ^

bX)

• ^

O

C/5

§■

2 2:

o ^

bX)

•P

' LL

18

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Discussion

A total of 17 Cervonema species (including the three new species described herein) have been described

so far. Differentiating between Cervonema species can be difficult due to relatively subtle differences in

morphology. A summary of key morphological features of all known Cervonema species is provided to

help identification (Table 3).

Subfamily Dorylaimopsinae De Coninck, 1965

Diagnosis (modified from Jensen (1979))

Cuticle usually differentiated laterally. Cephalic sense organs in three distinctly separated crowns.

Buccal cavity usually strongly cuticularised; anterior portion shallow, sometimes with small pointed

projections at border with vestibulum; posterior portion dilated, cylindrical or conical, usually with

strongly cuticularised walls and three thomlike projections (teeth) at the border between the two

portions. Spicules sometimes differentiated proximally, medially or distally. Gubemacular apophyses

usually directed caudally, occasionally dorsocaudally.

Type genus

Dorylaimopsis Ditlevsen, 1918

Remarks

The genus Dorylaimopsis is similar in many respects to the genus Hopperia, but can be distinguished

from the latter by the presence of longitudinal rows of dots as opposed to larger, irregularly-distributed

dots in the lateral fields (sometimes arranged in transverse rows) in Hopperia. There are two exceptions:

(1) some specimens of H. communis Gagarin & Thanh, 2006 have irregularly distributed dots in the

lateral fields, and (2) D. metatypica Chitwood, 1936 is characterised by irregularly-spaced dots instead

of longitudial rows. The two genera also tend to differ in the shape of the spicules: arcuate in Hopperia,

whilst in Dorylaimopsis spicules may be arcuate {D. coomansi Muthumbi et al, 1997), jointed {D.

perfecta Cobb, 1920), or with lateral projections {D. punctata Ditlevsen, 1918). In addition, species of

the genus Dorylaimopsis are characterised by conico-cylindrical tail with (usually) swollen tip, whilst

tail shape in Hopperia can be conico-cylindrical with swollen {H. americana Pastor de Ward, 1984),

filiform {H. massiliensis NitieWo, 1969), or pointed tip {H. australis iQnsQn, 1992).

Genus Dorylaimopsis Ditlevsen, 1918

Diagnosis (modified from Jensen (1979))

Dorylaimopsinae. Cuticle with lateral differentiation consisting of longitudinal rows of coarse dots;

posterior portion of buccal cavity cylindrical, with three thom-like teeth at border to anterior portion;

outer labial sensillae and cephalic setae in separate circles; spicules usually long, arcuate or jointed;

caudal or dorso-caudal gubemacular apophyses.

Type species

Dorylaimopsis punctata Ditlevsen, 1918

19

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Dorylaimopsis nodderi sp. nov.

um:lsid:zoobank.org:act:A19205F2-A989-49EA-9002-46CA34F64CCA

Figs 9-11, Table 4

Etymology

This species is named after Dr. Scott D. Nodder.

Material examined

Holotype

S, collectedS Oct. 2001, ChathamRise crest (350m water depth), 43.4333°S, 178.5000°E. Characteristics

of surface sediment layer (0-5 mm): mostly fine sand (35.7%), silt/clay (33.0%), and very fine sand

(23.5%); calcium carbonate content: 40.5%; total organic matter content: 4.5%; chloroplastic pigment

concentration: 3359 ng/gOW^^^jj^^j^^ (NIC 84446).

Paratypes

4 SS, 4 $ $, same data as holotype (NIC 84447).

Description

Males

Body cylindrical, tapering slightly towards both extremities. Cuticle punctate from level of amphid to

near tail tip, with lateral differentiation consisting of three longitudinal rows of larger dots in pharyngeal

and caudal regions, two rows of larger dots elsewhere. Cuticle in lateral field consisting of 1 pm thick

basal layer, 4 pm thick median layer with forked columns (side view), and 2 pm thick cortical layer

(Fig. 11 A). Cuticle thinner in caudal region. Eight longitudinal rows of short, sparse somatic setae.

Head set-off by constriction immediately posterior to cephalic setae. Six inner labial papillae, six outer

labial papillae, and four cephalic setae. Anterior portion of buccal cavity cup-shaped. Posterior portion

of buccal cavity cylindrical, cuticularised, narrow, 13-16 pm deep, with three cuticularised projections

(teeth) at border to anterior portion. Amphideal fovea spiral, 2.5-3.0 turns, located immediately posterior

to cephalic setae. Pharynx gradually swelling, not forming true bulb. Marginal tubes beginning from

base of stoma. Cardia short. Nerve ring near middle of pharynx, situated anteriorly to secretory-excretory

pore. Cellular body of ventral gland at level of cardia. Intestine wall with numerous small granules,

interspersed with numerous regularly-spaced clear areas without granules, circular or irregular in shape

(Fig. 9E).

Reproductive system diorchic, opposed, outstretched. Anterior testis to left of intestine, posterior testis

to right of intestine. Spicules paired, equal, 1.8-2.9 abd long, with rounded ventral projection at one third

of spicule length from distal end, giving appearance of a joint. Spicules straight proximally, strongly

bent at level of joint. Gubemaculum surrounding spicules, with wide triangular base and pair of long,

straight caudal apophyses. Rectal gland surrounds gubemaculum. Twelve to sixteen tubular pre-cloacal

supplements. Tail conico-cylindrical, with several short caudal setae and three short terminal setae.

Three caudal glands.

Females

Similar to males, but with slightly smaller amphideal fovea, 2.5-2.75 turns. Reproductive system

didelphic, opposed, outstretched, with anterior branch to left of intestine and posterior branch either to

right or left of intestine. Vulva slightly pre median. Granular vaginal glands present, pars proximalis

vaginae surrounded by constrictor muscle.

20

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Fig. 9. Dorylaimopsis nodderi sp. nov. A. Anterior region of S- B. Lateral surfaee view of S head.

C. Ventral view of $ head (eross-seetion). D. Entire S- E. S gut, mid-body. Seale bar: A = 50 pm; B and

C = 25 pm; D = 70 pm; E = 30 pm.

21

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Fig. 10. Dorylaimopsis nodderi sp. nov. A. $ reproductive system. B. S posterior region, showing

copulatory apparatus. C. $ tail. D. S posterior body region, showing copulatory apparatus. Scale bar:

A = 90 pm; B = 40 pm; C and D = 55 pm.

22

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Fig. 11. Dorylaimopsis nodderi sp. nov. light micrographs. A. $ cuticle in pharyngeal region showing

the eortieal, median, and basal layers. B. Ventral view of $ head showing posterior portion of bueeal

eavity. C. $ mid-body region showing vulva and mature eggs. D. Sperm eells with nuelei. Seale bar:

A = 8 pm; B = 12 pm; C = 50 pm; D = 30 pm.

23

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Table 4. Morphometries (pm) of Dorylaimopsis nodderi sp. nov.

Species

Dorylaimopsis nodderi sp. nov.

Male Holotype

Male Paratypes

Female Paratypes

n

4

L

1524

1199-1602

1080-1656

a

23

18-27

20-26

b

7

6-7

c

14

11-13

Head diam.

14

12-14

13-19

Mouth diam.

3

1-2

2-4

Length of cephalic setae

6

4-7

5-7

Amphid height

11

9-11

7-9

Amphid width

10

8-9

Amphid width/cbd (%)

59

63-67

40-53

Amphid from anterior end

8

7-8

7-9

Nerve ring from anterior end

106

100-125

93-132

Nerve ring cbd

47

40-55

41-57

Excretory pore from anterior end

134

115-148

105-153

Pharynx length

224

196-250

185-264

Pharynx cbd

65

45-75

50-74

Pharyngeal bulb diam.

34

29-50

34-52

Max. body diam.

65

44-87

50-81

Spicule length

112

98-109

-

Gubernacular apophyses length

40

31-39

-

Anal body diam.

44

38-58

37-54

Tail length

112

109-128

98-134

Tail length/abd

2.5

2.2-2.9

2.5-3.0

V

-

497-790

%V

-

46-48

Vulval body diam.

-

49-81

Diagnosis and relationships

Dorylaimopsis nodderi sp. nov. is eharaeterised by eutiele with lateral differentiation eonsisting of three

longitudinal rows of larger dots in pharyngeal and eaudal regions, two rows of larger dots in middle region

of body, eephalie setae 4-7 pm long, amphideal fovea with 2.5-3.0 turns, and spieules 98-112 pm long with

rounded ventral projeetion at one third of spieule length from distal end, giving appearanee of a joint.

Dorylaimopsis nodderi sp. nov. differs from most other speeies of the genus (exeept D. punctata

Ditlevsen, 1918 and D. magellanense Chen & Vinex, 1998) in the presenee of spieules with ventral

projeetions. D. nodderi sp. nov. ean be distinguished from D. punctata by its smaller body length (1080-

1656 vs. > 1900 pm), presenee of three rows of enlarged dots in pharyngeal and eaudal regions of both

males and females (two rows in D. punctata), and fewer pre-eloaeal supplements (12-16 vs. 18-22).

D. nodderi sp. nov. ean be distinguished from D. magellanense by its smaller body length (1199-1602

vs. 1708-2261 pm), and the smaller number of rows of enlarged dots in the lateral fields: three rows in

pharyngeal and eaudal regions (four or five in D. magellanense) and two rows in middle region of body

(three or four in D. magellanense).

24

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Dichotomous key to all known species of Dorylaimopsis

(modified from Jensen (1979) and Zhang (1992))

1. Cutiele without lateral longitudinal rows of eoarse dots along entire body.2

- Cutiele with longitudinal rows of dots along entire length of body.4

2. Lateral longitudinal rows of eoarse dots present posterior to pharyngeal region only.3

- Lateral longitudinal rows of eoarse dots eompletely absent.

. Dorylaimopsis metatypica Chitwood, 1936

3. Lateral differentiation of eutiele eonsisting of 1-3 longitudinal rows of dots beginning posterior to

pharynx, gubemaeulum with swollen distal end. D. coomansi Muthumbi et al, 1997

- Lateral differentiation of eutiele eonsisting of 3-5 longitudinal rows of dots beginning posterior to

pharynx, triangular gubemaeulum tapering distally. D. turneri Zhang, 1992

4. Cutiele laterally with two longitudinal rows of eoarse dots in middle body region of both sexes.5

- Cutiele laterally with more than two longitudinal rows of eoarse dots in at least one of the sexes ..14

5. Lateral differentiation of eutiele < 0.2 ebd.6

- Lateral differentiation > 0.2 ebd.7

6. Body length 2.5-3.2 mm; spieules with ventral projeetion. D. punctata Ditlevsen, 1918

- Body length 1.3-1.9 mm; spieules without ventral projeetion. D. rabalaisi Zhang, 1992

7. Spieules jointed, with ventral projeetion, or simple.8

- Spieules jointed and irregularly eutieularised. D. mediterranea Grimaldi-de Zio, 1968

8. Spieules jointed, with two equal segments. D. perfecta (Cobb, 1920)

- Spieules otherwise.9

9. Spieules without distal hooks or projeetions.10

- Spieules with distal hooks or ventral projeetions.13

10. Gubemaeulum with irregular base; gubemaeular apophyses with blunt distal end.

. D. nini (Inglis, 1961)

- Gubemaeulum with smooth base; gubemaeular apophyses with pointed or rounded distal end.11

11. Gubemaeulum with pointed end. D. peculiaris Platonova, 1971

- Gubemaeulum with rounded end.12

12. Male body length <2100 pm, two equal testes, spieules with poorly developed eapitulum, spieule

length 1.7-1.9 abd. D. gerardi Muthumbi et al, 1997

- Male body length >2100 pm, anterior testis larger than posterior testis and with larger sperm, spieules

with well-developed eapitulum, spieule length 1.8-2.4 abd. D. variabilis Muthumbi et al, 1997

13. Spieules with small distal hook; gubemaeulum with lateral projeetions ...D, pellucida (Cobb, 1920)

- Spieules with rounded ventral projeetion at one third of spieule length from distal end; eutiele

laterally with three longitudinal rows in pharyngeal and eaudal regions, two rows in middle

region. D. nodderi sp. nov.

14. Cutiele with two rows of eoarse dots in males and three in females.15

- Cutiele with more than two rows of eoarse dots in males.16

25

European Journal of Taxonomy 24: 1-42 (2012)

15. Spicules distally acute. D. poriferum (Cobb, 1920)

- Spicules distally with a subterminal hook. D. timmi Jensen, 1979

16. Cuticle with three rows of coarse dots in middle body region of males.17

- Cuticle with four or five rows of coarse dots in middle body region of males.18

17. Cuticle with 3 rows of coarse dots along entire length of body of both males and

females. D. halongensis Tu et al, 2008

- Cuticle with three rows of coarse dots in middle region of body, four to eight rows elsewhere (males

and females). D.janetae (Inglis, 1963)

18. Spicules without subterminal hook. D. magellanense Chen and Vincx, 1998

- Spicule with subterminal hook; spicule length 3 abd. D. angelae (Inglis, 1967)

GenusVitiello, 1969

Diagnosis (modified from Jensen (1979))

Dorylaimopsinae. Cuticle with lateral differentiation of irregularly-spaced larger dots, sometimes

arranged in transverse rows; posterior portion of buccal cavity cylindrical, with three thom-like teeth at

border to anterior portion (except in H. hexadentata Hope & Zhang, 1995 which has six teeth); ovaries

usually outstretched, refiexed ovaries sometimes present; spicules arcuate; gubemaculum with caudal

or dorso-caudal apophyses.

Type species

Hopperia massiliensis Vitiello, 1969

Hopperia ancora sp. nov.

urn:lsid:zoobank.org:act:42970B24-CF4B-4D54-9CFF-B278BE09C60C

Figs 12-14, Table 5

Etymology

The species name is derived from the Latin ancora (= anchor), in reference to the distal hook on the

spicules.

Material examined

Holotype

(f, collected 7 Jun. 2007, Challenger Plateau (264 m water depth), 39.6463°S, 172.1530°E. Characteristics

of surface sediment layer (0-5 mm): mostly silt/clay (90.1%), with very fine sand (3.0%), and fine sand

(3.0%); calcium carbonate content: 27.4%; total organic matter content: 1.94%; chloroplastic pigment

concentration: 3039 ng/gOW^^^jj^^j^^ (NIC 84448).

Paratypes

3 (ff and 3 $ $, same data as holotype (NIC 84449).

Description

Males

Body cylindrical, tapering slightly towards anterior extremity. Cuticle punctated from level of amphid to

near tail tip, with lateral differentiation consisting of larger, irregularly-spaced dots. Eight longitudinal

rows of short, sparse somatic setae. Head set-off by depression at level of cephalic setae. Six inner labial

papillae, six outer labial papillae, and four conical cephalic setae in three distinct circles. Amphideal

26

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Fig. 12. Hopperia ancora sp. nov. A. Anterior region of S- B. Lateral surfaee view of S head. C. Lateral

view of $ head. D. S posterior body region, showing eopulatory apparatus. E. $ tail. Seale bar:

A = 45 pm; B and C = 20 pm; D = 35 pm; E = 50 pm.

27

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Fig. 13. Hopperia ancora sp. nov. A. Entire f. B. $ reproductive system. Scale bar: A = 240 pm;

B = 110 pm.

28

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

fovea spiral, 2.5-3.0 turns, located immediately posterior to cephalic setae.Anterior portion of buccal

cavity cup-shaped. Posterior portion of buccal cavity cylindrical, strongly cuticularised, 4-5 pm wide,

20-27 pm deep, with three strongly cuticularised teeth at border to anterior portion. Teeth everted in

some specimens (Fig. 14A). Marginal tubes of pharynx beginning from base of stoma. Pharynx gradually

swelling, not forming true bulb. Cardia short. Nerve ring near middle of pharynx, situated anteriorly to

Fig. 14. Hopperia ancora sp. nov. light micrographs. A. S head, showing everted tooth. B. Intestine

wall of S, with arrows pointing to oval cells with larger, more closely packed granules than surrounding

cells. C. Everted spicule. D. Vulva. Scale bar: A, D = 15 pm; B = 22 pm; C = 20 pm.

29

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Table 5. Morphometries (pm) of Hopperia ancora sp. nov. and H. beaglense Chen and Vinex, 1998.

Species

Hopperia ancora s

). nov.

Hopperia beaglense Chen & Vinex, 1998

Males

Females

Holotype

Paratypes

Males

Females

n

3

5

4

L

2011

1876-1986

1985-2139

1446-1616

1573-1856

a

47

32-46

29-50

31-36

29-36

b

9

8-9

7-8

8

c

12

10-11

10

14-17

16-19

Head diam.

16

15-17

13-14

14-15

Mouth diam.

4

5

4-5

3-6

3-5

Length of cephalic setae

3

2-3

3

4-6

Amphid height

11

10-11

8

Amphid width

12

11-12

9-12

10-12

9-10

Amphid width/cbd (%)

58

58-64

50-60

63-71

53-59

Amphid from anterior end

7

5-7

5

6

Nerve ring from anterior end

90

82-104

97-111

82-103

98-113

Nerve ring cbd

45

38-44

41-49

34-38

31-45

Excretory pore from anterior end

118

116-124

123-137

115-124

133-140

Pharynx length

218

210-236

233-240

187-208

208-235

Pharynx cbd

53

48-62

39-44

48-60

Pharyngeal bulb diam.

37

35-47

33-47

27-30

37-42

Max. body diam.

43

41-62

40-69

42-48

49-64

Spicule length

62

54-68

-

48-58

-

Gubernacular apophyses length

18

19-21

-

25-31

-

Anal body diam.

38

36-44

32-36

35-40

38-45

Tail length

167

175-206

202-212

92-100

100-105

Tail length/abd

4.4

4 . 7 - 4.9

5 . 9 - 6.3

2 . 3 - 2.9

22-2.1

V

-

934-1041

-

785-953

%V

-

47-49

-

50-52

Vulval body diam.

-

40-67

-

48-61

exeretory pore. Cellular body of ventral gland situated just posteriorly to eardia. Intestine wall with

numerous small, elear granules, and oval eells with more elosely-paeked granules (Figs 13 A, 14B).

Reproduetive system diorehie, opposed, outstretehed. Anterior testis to left of intestine and posterior

testis to right of intestine in all speeimens exeept one speeimen with opposite arrangement. Sperm

eells globular with outer radial lines. Spieules paired, equal, areuate, 1.5-1.7 abd long, with swollen

proximal end and eentral internal eutieularised projeetion (lamella) extending one fourth of spieule

length from proximal end. Spieules with velum and eonspieuous hook-like projeetion at distal end

(Fig. 14C). Gubemaeulum appearing to eompletely surround spieules, with bent dorso-eaudal apophyses.

Reetal gland surround distal end of spieules and gubemaeulum. Eleven to thirteen tubular pre-eloaeal

supplements, one pre-eloaeal seta. Tail long, eonieo-eylindrieal, gradually tapering, with several short

eaudal setae and no terminal setae. Three eaudal glands.

30

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Females

Similar to males, but with slightly longer tail. Reproduetive system didelphie, opposed, outstretehed,

with anterior and posterior branehes either to left or right of intestine, but always on opposite sides.

Vulva slightly pre median. Granular vaginal glands present, pars proximalis vaginae surrounded by

eonstrietor musele (Fig. 14D).

Diagnosis and relationships

Hopperia ancora sp. nov. is eharaeterised by body length 1876-2011 pm, short eonieal eephalie setae,

amphideal fovea with 2.5-3.0 turns, bueeal eavity with three teeth, spieules 54-68 pm in length with

hook-like projeetion at distal end, gubemaeulum with bent apophyses, 11-13 pre-eloaeal supplements,

and gradually tapering eonieo-eylindrieal tail without swollen tip or terminal setae.

Hopperia ancora nov. most elosely resembles//. massiliensisNitiQWo, 1969 and//. indianaMn\hmnb\

et al, 1997 in body size, values of a, b, and c, size and shape of the eephalie setae, amphideal fovea,

and tail, but ean be distinguished from them (and all other speeies of the genus) by the presenee of a

eonspieuous hook-like projeetion at distal end of spieules.

Hopperia beaglense Chen & Vinex, 1998

Figs 15-17, Table 5

Material examined

5 (J(? and 4 $$, eolleeted 5 May 2010, Kaikoura Canyon axis (1061 m water depth), 42.5081°S,

173.6325°E (NIC 84450).

Description

Males

Body eylindrieal, tapering slightly towards anterior extremity. Cutiele punetate from level of amphid

to near tail tip, with lateral differentiation eonsisting of larger, irregularly-spaeed dots. Somatie setae

short and sparse, in four dorso- and ventro-lateral longitudinal rows. Six inner labial papillae, six outer

labial papillae, and four eephalie setae in three distinet eireles. Anterior portion of bueeal eavity eup-

shaped. Posterior portion of bueeal eavity eylindrieal, strongly eutieularised, 24-26 pm deep, extending

into three strongly eutieularised teeth at border to anterior portion. Teeth everted in some speeimens.

Cutieularisation of posterior portion of bueeal eavity extends posterior to junetion of marginal tubes

with pharyngeal lumen (Fig. 15B). Amphideal fovea spiral, 3.0-3.5 turns, loeated immediately posterior

to eephalie setae. Pharynx with oval posterior bulb. Cardia short. Nerve ring near middle of pharynx,

situated anteriorly to seeretory-exeretory pore. Cellular body of ventral gland large (up to 37 x 20 pm),

situated just posteriorly to eardia. Intestine wall with numerous small, elear granules, and small elusters

of round, elear inelusions, sometimes with smaller orange-brown granules (Fig. 17B).

Reproduetive system diorehie, opposed, outstretehed. Anterior testis to left of intestine and posterior

testis to right of intestine. Spieules paired, equal, areuate, strongly eutieularised, 1.3-1.6 abd long, with

swollen proximal end and internal eutieularise projeetion (lamella) extending one fourth of spieule

length from proximal end, no velum. Gubemaeulum with straight dorso-eaudal apophyses, tapering

distally. One pre-eloaeal seta and 7-8 eonspieuous, eup-shaped pre-eloaeal supplements. Tail short,

eylindrieal, with rounded tip, several short eaudal setae and no terminal setae. Three large eaudal glands

and well-developed spinneret (Fig. 17C).

Females

Similar to males, but slightly greater maximum body diameter and smaller amphids, 3.0 turns.

Reproduetive system didelphie, opposed, outstretehed, with anterior braneh to left of intestine and

31

European Journal of Taxonomy 24 : 1-42 ( 2012 )

C

Fig. 15. Hopperia beaglense Chen and Vincx, 1998. A. Anterior region of f. B-C. Lateral view of S

head. D. Lateral surfaee view of $ head. E. $ tail. F. S eopulatory apparatus and vas deferens. Arrow

shows approximate position of supplement-like strueture observed in type speeimens from Chile. Seale

bar: A = 30 pm; B-D = 22 pm; E-F = 32 pm.

32

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

posterior branch to right of intestine, except for one individual with opposite arrangement. Vulva slightly

post median. Granular vaginal glands present, vagina uterina surrounded by constrictor muscle.

Discussion

My specimens closely resemble the description given by Chen & Vincx (1998) based on specimens from

the Beagle Channel, Chile (100-110 m water depth). My specimens, however, have slightly lower a

Fig. 16. Hopperia beaglense Chen and Vincx, 1998. A. Entire $. B. (? posterior region, showing testes

and copulatory apparatus. Scale bar =100 pm.

33

European Journal of Taxonomy 24 : 1-42 ( 2012 )

(range: 29-36 vs. 30-41; mean: 34.2 vs. 39.1) and c values (range: 14-19 vs. 19-22; mean: 15.8 vs. 20.6),

have 7-8 supplements instead of 6-7, and laek the supplement-like strueture situated halfway down the

ventral side of the tail (Fig. 15F). This is the first reeord of this speeies outside the type loeality.

Within the Dorylaimopsinae, eutieularisation of the posterior bueeal eavity does not usually extend into

the pharyngeal lumen (i.e., eutieularisation stops before marginal tubes begin) (fig. 2E in Jensen 1979).

My speeimens are unusual in that the eutieularisation of the bueeal eavity extends into the anteriormost

portion of the pharyngeal lumen (see Fig. 15B). It is unelear whether this is also a feature of the type

speeimens, however, beeause Chen & Vinex (1998) did not deseribe the marginal tubes.

Fig. 17. Hopperia beaglense Chen and Vinex, 1998 light mierographs. A. Anterior region of f. B. S

gut, showing elusters of small round inelusions with smaller orange-brown granules. C. Spinneret. Seale

bar: A = 30 pm; B = 12 pm; C = 24 pm.

34

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Key to all known species of Hopperia

(modified from Pastor de Ward (2004))

1. Cuticle with irregularly-distributed dots in the lateral fields.3

2. Cuticle with four longitudinal rows of coarse dots in lateral fields in anterior half or entire body

length. Hopperia communis Gagarin & Thanh, 2006

3. Stoma armed with three teeth at juncture of anterior and posterior portions of buccal cavity.4

- Stoma armed with six teeth at juncture of anterior and posterior portions of buccal

cavity. H. hexadentata Hope and Zhang, 1995

4. Spicules without joint.5

- Spicules jointed; cephalic setae 0.4 hd; > 13 small pre-cloacal supplements; tail conico-cylindrical

with swollen tip and terminal setae. H. metatypica (Chitwood, 1936)

5. Spicules without distal hook.6

- Spicules with distal hook; short conical cephalic setae; tail without terminal setae.

. H. ancora sp. nov.

6. Spicules without sharp proximal bend pointing ventrally.7

- Spicules with sharp proximal bend; tail conico-cylindrical with swollen tip and terminal

setae. H. mira Gagarin and Thanh, 2006

7. Tail without conspicuous spinneret.8

- Tail with conspicuous spinneret; terminal setae absent; cylindrical portion of tail > 50% tail

length. H. dolichura Gagarin and Thanh, 2006

8. Length of posterior cylindrical portion of buccal cavity >1.5 amphid diameters.9

-Length of posterior cylindrical portion of buccal cavity <1.5 amphid diameters; 12 pre-cloacal

supplements; spicules 0.5 abd. H. patagonica Pastor de Ward, 2004

9. Length of posterior cylindrical portion of buccal cavity =1.5-2.5 amphid diameters.10

- Length of posterior cylindrical portion of buccal cavity >2.5 amphid diameters.11

10. Spicules < 2.0 abd.12

- Spicules >2.0 abd; pre-cloacal supplements present; tail with terminal setae.

. H. muscatensisW?iTw\Qk^ 1973

11 . > 10 pre-cloacal supplements; spicules 69 pm. H indiana Muthumbi et al., 1997

- < 10 pre-cloacal supplements; spicules 37-44 pm. H. australis Jensen, 1992

12. Tail tip not swollen.13

- Tail tip swollen.14

13. Refiexed ovaries; 8 pre-cloacal supplements. H. dorylaimopsoides (Allgen, 1959)

- Outstretched ovaries; 6-8 pre-cloacal supplements; tail conical.

. H. beaglense Muthumbi et al, 1997

14 .<10 pre-cloacal supplements; ovaries outstretched.15

- > 10 pre-cloacal supplements; ovaries refiexed. H. massiliensis Vitiello, 1969

15. Tail 3.5 abd; a = 26-30; spicules 1.4 abd. H. americana Pastor de Ward, 1984

- Tail 4.4 abd; a = 31-40; spicules 1.8 abd. H. arntzi Chen & Vincx, 1998

35

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Genus Kenyanema Muthumbi et al., 1997

Diagnosis (modified from Muthumbi et al. (1997))

Dorylaimopsinae. Head region mueh narrower than the rest of the body; eutiele punetate and sometimes

transverse striae are visible; inner and outer labial sensillae indistinet; four eephalie setae; tubular

stoma without teeth; male reproduetive system monorehie with outstretehed anterior testis; females

amphidelphie with outstretehed ovaries; spieules areuate; gubemaeulum with eaudal apophyses.

Type species

Kenyanema monorchis Muthumbi et al, 1997

Figs 18-19, Table 6

Material examined

3 SS and 3 $ eolleeted 6 Apr. 2007, southern flank of Chatham Rise (1240 m water depth), 44.4864°S,

111. 1407°E. Charaeteristies of surfaee sediment layer (0-5 mm): mostly silt/elay (82.9%), with very flne

sand (10.2%), and flne sand (8.4%); ealeium earbonate eontent: 38.4%; total organie matter eontent:

1.9%; ehloroplastie pigment eoneentration: 3501 ng/gOW^^^^^^^^ (NIC 84451).

Description

Males

Body eylindrieal, tapering towards anterior extremity. Cutiele with transverse rows of dots, lateral

differentiation eonsisting of larger, irregularly-spaeed dots. Eight rows of short, sparse somatie setae.

Head set-off by eonstrietion immediately posterior to eephalie setae. Inner and outer labial sensillae

indistinet, four eephalie setae. Amphideal fovea spiral, 2.5 turns, loeated at level of eephalie setae.

Bueeal eavity tubular, 4 pm deep, 1 pm wide, surrounded by pharyngeal museles. Marginal tubes

beginning from base of stoma. Pharynx gradually swelling, not forming true bulb, eardia short. Nerve

ring near middle of pharynx, situated anteriorly to seeretory-exeretory pore. Cellular body of ventral

gland near level of eardia. Intestine wall with numerous small, eolourless granules, and elusters of elear

eireular inelusions, mostly in posterior portion of intestine (Fig. 18E).

Reproduetive system monorehie with outstretehed anterior testis to left of intestine. Mature sperm eells

large, oval-shaped, up to 30 pm long and 13 pm wide. Spieules paired, equal, areuate, 1.6-1.8 abd

long, with swollen proximal end and eentral internal eutieularisated projeetion (lamella) extending from

proximal end over one flfth of spieule length. Gubemaeulum with pair of straight eaudal apophyses.

Reetal gland surrounds posterior portion of spieules and gubemaeulum. Two to three small tubular pre-

eloaeal supplements, one pre-eloaeal seta. Tail eonieo-eylindrieal with swollen tip, several short eaudal

setae and three short terminal setae. Three eaudal glands and spinneret.

Females

Similar to males, but with slightly greater maximum body diameter, amphideal fovea slightly smaller,

2.0-2.25 turns. Reproduetive system didelphie, opposed, outstretehed, with sperm distributed over uteri

and with branehes either to the right or left of intestine, but always on opposite sides. Vulva slightly post

median.

Discussion

My specimens elosely resemble the description of Kenyanema monorchis Muthumbi et al, 1997 based

on specimens from the Indian Ocean (500-1112 m water depth). My specimens, however, have a smaller

36

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Fig. 18. Kenyanema monorchis Muthumbi et al., 1997. A. Anterior region of B: lateral surfaee view

of (S head. C. Lateral view of 3 head (eross-seetion). D. Posterior region of (J showing eopulatory

apparatus. E. $ gut, posterior body region. F. $ tail. Seale bar = 25 pm.

37

European Journal of Taxonomy 24 : 1-42 ( 2012 )

Fig. 19. Kenyanema monorchis Muthumbi et al., 1997. A. Entire f. B. $ reproductive system. Scale

bar =100 pm.

38

LEDUC D., Deep-sea nematodes from the Southwest Paeihe Oeean

Table 6. Morphometries (pm) of Kenyanema monorchis Muthumbi et al., 1997.

Species

Kenyanema monorchis Muthumbi et al., 1997

Males

Females

n

4

3

L

1020-1107

1070-1205

a

29-39

27-29

b

7

8

c

10-12

11-12

Head diam.

8-9

8

Mouth diam.

1

Length of cephalic setae

3-4

2-3

Amphid height

5

4-5

Amphid width

5-6

5

Amphid width/cbd (%)

56-66

63

Amphid from anterior end

2-3

Nerve ring from anterior end

80-85

79-90

Nerve ring cbd

23-25

23-28

Excretory pore from anterior end

97-107

94-116

Pharynx length

141-150

138-160

Pharynx cbd

28-31

29-33

Pharyngeal bulb diam.

14-17

16-18

Max. body diam.

30-39

39-43

Spicule length

45-49

-

Gubernacular apophyses length

11-15

-

Anal body diam.

27-29

26-28

Tail length

90-99

96-105

Tail length/abd

3 . 2 - 3.7

3 . 5 - 3.8

V

-

569-644

%V

-

53

Vulval body diam.

-

39-43

bueeal eavity (4 vs. 8 pm deep) and are eharaeterised by the presenee of 2-3 pre-eloaeal supplements.

No pre-eloaeal supplements were observed in the original deseription of this speeies, but may have been

overlooked. This is the first reeord for this genus outside the type loeality.

Discussion

A reeent study suggests that at least 775 nematode morphospeeies (speeies that ean be differentiated

from others based on their morphology but whieh have not been formally deseribed) are present on the

eontinental margins of New Zealand (water depth 250-3100 m; Ledue et al. 2012b). Sueh high level of

diversity is probably related to the wide range of environments eneompassed by the eountry’s Exelusive

Eeonomie Zone (e.g., Nodder et al 2012). However, only 12 nematode speeies (or 1.5% of total) have

been reeorded/deseribed from New Zealand’s deep-sea habitats so far (Wieser 1956, Eedue et al 2012a,

present study). More work is elearly needed to obtain a better appreeiation of New Zealand’s deep-sea

nematode fauna and its relationship to the fauna of other regions.

New reeords of C. shiae Chen & Vinex, 2000, H. beaglense Chen & Vinex, 1998, and K. monorchis

Muthumbi et al, 1997 on the eontinental margin of New Zealand suggest that these speeies have a

39

European Journal of Taxonomy 24 : 1-42 ( 2012 )

wide distribution extending aeross oeean basins. Miljutin et al. (2010) noted that, out of 638 formally

deseribed deep-sea nematode speeies, 46 (ineluding three Comesomatidae speeies) are likely to have

a eosmopolitan distribution. The true proportion of deep-sea eosmopolitan speeies remains unknown

due to paueity of data. Reeent moleeular evidenee provides support for the existenee of eosmopolitan

nematode speeies in the deep sea (Bik et al. 2010), although the vast majority of families (ineluding the

Comesomatidae) remains to be investigated.

Acknowledgments

Sample data were generated under the Foundation for Researeh, Seienee and Teehnology (FRST)

programme “Oeean Eeosystems” (CO 1X0027) and the Oeean Survey 20/20 Chatham-Challenger

projeet for whieh 1 thank the funding ageneies. Land Information New Zealand, Ministry of Fisheries,

Department of Conservation and NIWA, for permission to use the data. Funding was also provided by

FRST through a postdoetoral fellowship (UOOX0909), the programmes “Consequenees of Earth-Oeean

Change” (C01X0702), “Coasts & Oeeans OBI” (C01X0501), and “Impaet ofresouree use on vulnerable

deep-sea eommunities” (CO 1X0906). 1 also aeknowledge the other partieipants of voyages TAN0116,

TAN0705, TAN0707, TAN1006 and the offieers and erew of RV Tangaroa. 1 thank R Keith Probert

(University of Otago) and Seott Nodder (NIWA) for supporting my researeh, Katrin Berkenbuseh

(University of Otago) for proeessing of meiofauna samples (TAN0116). 1 am also grateful to the editors

and two anonymous reviewers for their helpful co mm ents on the manuseript.

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European Journal of Taxonomy 24 : 1-42 ( 2012 )

Manuscript received on: 25 June 2012

Manuscript accepted on: 4 September 2012

Published on: 5 October 2012

Topic editor: Rudy Jocque

In compliance with the ICZN, printed versions of all papers are deposited in the libraries of the institutes

that are members of the EJT eonsortium: Museum national d’Histoire naturelle, Paris, Franee; National

Botanie Garden of Belgium, Meise, Belgium; Royal Museum for Central Afriea, Tervuren, Belgium;

Natural History Museum, London, United Kingdom; Royal Belgian Institute of Natural Seienees,

Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark.

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