European Journal of Taxonomy 118: 1-25

http://dx.doi.org/10.5852/ejt.2015.118

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2015 • Leduc D. & Verschelde D.

Research article

urn:lsid:zoobank.org:pub:4302BA88-0639-4062-84F2-EECD733807A5

New Spirinia and Stygodesmodora species (Nematoda, Spiriniinae)

from the Southwest Pacific, and a revision of the related genera

Spirinia, Chromaspirina and Perspiria

Daniel LEDUC 13 & Dominick VERSCHELDE 2 4

1 National Institute of Water and Atmospheric Research, Private Bag 14-901, Wellington, New

Zealand. Correponding author: [email protected]

2 Ghent University - Zoology Museum, K.L. Ledeganckstraat 35, B-9000 Ghent, Belgium

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

4 urn:lsid:zoobank.org:author:80C09028-E083-4DF0-B514-9C92A12ED78E

Abstract. Two new species of the family Desmodoridae are described from the upper continental

slope of New Zealand, Southwest Pacific, and the genera Spirinia , Chromaspirina and Perspiria are

revised. Spirinia verecunda sp. nov. is characterised by a short, stout body, cuticle covered in minute,

hair-like structures, unispiral amphideal aperture and cryptospiral amphideal fovea, buccal cavity with

small dorsal tooth and minute subventral teeth, eight oblong glands surrounding anterior portion of

pharynx, large sperm cells, spicules with weak capitulum, and the absence of precloacal supplements.

Stygodesmodora confusa sp. nov. is characterised by a relatively short body, spiral amphids with 1.2-

1.4 turns, cephalic setae situated at or slightly posterior to mid-level of amphid, and males with four

precloacal supplements consisting of short setae on wide bases. Like other species of the genus, S. confusa

sp. nov. is characterised by an annulated head region, but in some specimens the cuticle annulations are

restricted to the dorsal and ventral sides of the head. S. confusa sp. nov. shows similarities with species

of other desmodorid genera (i.e., Echinodesmodora, Bolbonema ), but can be differentiated from them

by the presence of an annulated head region and amphideal plates. The genera Spirinia, Chromaspirina

and Perspiria have very similar morphologies and differ mainly in the size and structure of the buccal

cavity ( Spirinia vs Chromaspirina) or the shape of the tail and placement of the amphi ds relative to the

cuticle annulations ( Spirinia vs Perspiria ); their diagnoses are clarified and some nomenclatural changes

are proposed to eliminate overlap in the definitions of these genera.

Keywords. Desmodoridae, deep-sea, meiofauna, Chatham Rise, southern Hikurangi Margin

Leduc D. & Verschelde D. 2015. New Spirinia and Stygodesmodora species (Nematoda, Spiriniinae) from the

Southwest Pacific, and a revision of the related genera Spirinia , Chromaspirina and Perspiria. European Journal

of Taxonomy 118: 1-25. http://dx.doi.org/10.5852/eit.2015.118

Introduction

Spirinia Gerlach, 1963 is a relatively common genus and twelve valid species have been described to date

(Silva et al. 2009). Spirinia is very similar to Chromaspirina Filipjev, 1918 in the shape and structure of

European Journal of Taxonomy 118 : 1-25 ( 2015 )

the head region, amphids, pharynx, and male copulatory apparatus. The main distinction between the two

genera is the shape of the buccal cavity: Spirinia is characterised by a narrow and lightly cuticularised

buccal cavity with no or a small dorsal tooth and no or minute subventral teeth, whereas Chromaspirina

is characterised by a more heavily cuticularised, medium to large buccal cavity with a large dorsal

tooth and smaller subventral teeth. A third genus, Perspiria Wieser & Hopper, 1967, is also similar to

Spirinia and differs from the latter in having a conico-cylindrical to filiform tail and by the amphids not

completely surrounded by body annulations. Species of these three genera show considerable overlap

in the above-mentioned morphological characteristics, however, and the boundaries between them have

become somewhat blurred. Here, we describe S. verecunda sp. nov. from the Chatham Rise crest (350 m

water depth) on the continental margin of New Zealand, and we propose several nomenclatural changes

to species of Spirinia and Chromaspirina to allow a more consistent distinction between the genera

based on buccal morphology. Several nomenclatural changes are also suggested for the genus Perspiria

to reflect differences in tail shape and body annulations of the head region between the latter genus and

Spirinia.

Stygodesmodora Blome, 1982 is a relatively uncommon genus comprising only two species: S.

bacillicauda (Gerlach, 1963) and S. epixantha Blome, 1982. These species were described from shallow

sandy sediments in the Indian Ocean and North Sea, respectively. Here a third species, S. confusa sp.

nov., is described from the continental margin of New Zealand (350 and 686 m water depth); it is the

first species of the genus to be described from the deep sea.

Material and Methods

Samples for species descriptions were obtained from the Chatham Rise and southern Hikurangi margin.

Southwest Pacific. The Chatham Rise is a submarine ridge that extends eastwards from the South Island

of New Zealand, over water depths ranging from ca. 250 to 3000 m. The highly productive Subtropical

Front (STF) is geographically constrained near the southern flank of the rise at around 44° S (Murphy et

al. 2001). The southern Hikurangi margin lies on the south-eastern extremity of the North Island of New

Zealand and is situated north of the Chatham Rise. Samples were collected at two sites: one situated

near the head of a canyon on the southern Hikurangi margin during National Institute of Water and

Atmospheric Research (NIWA) cruise TAN1004 (station 92, 686 m water depth, 41.8921° S, 174.6347°

E), and one at an open slope site located near the central Chatham Rise crest during NIWA cruise

TAN1103 (station 69, 350 m water depth; 43.331° S, 178.288° E).

Sediment samples were collected using an Ocean Instruments MC-800Amulticorer (MUC; core internal

diameter = 9.52 cm). Each sample consisted of one subcore of internal diameter 26 mm taken to a depth

of 5 cm. Samples were fixed in 10% formalin and stained with Rose Bengal. Samples were subsequently

rinsed on a 1 mm sieve to remove large particles and on a 45 pm sieve to retain nematodes. Nematodes

were extracted from the remaining sediments by Ludox flotation and transferred to pure glycerol

(Somerfield & Warwick 1996). Species descriptions were made from glycerol mounts using differential

interference contrast microscopy, and drawings were made with the aid of a camera lucida. Specimens

for scanning electron microscopy (SEM) were transferred to a 2% glutaraldehyde solution with sodium

cacodylate buffer overnight, then transferred to a 4% osmium tetroxide solution for 2 hours. They were

then gradually transferred to pure ethanol using a graded ethanol series, critical point dried, and mounted

onto stubs before being coated with gold using a sputter coater. Observations were made using a Hitachi

TM3000 tabletop SEM at high vacuum mode.

All measurements are in pm, and all curved structures are measured along the arc. Type specimens are

held in the NIWA Invertebrate Collection, Wellington, New Zealand. Abbreviations in the text are as

follows:

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

abd

cbd

body length/maximum body diameter

anal body diameter

body length/oesophagus length

body length/tail length

corresponding body diameter

vulva distance from anterior end of body x 100/total body length.

Results

Phylum Nematoda Cobb, 1932

Class Chromadorea Inglis, 1983

Subclass Chromadoria Adamson, 1987

Order Desmodorida de Coninck, 1965

Suborder Desmodorina de Coninck, 1965

Superfamily DesmodoroideaFilipjev, 1922

Family Desmodoridae Filipjev, 1922

Type genus

Desmodora de Man, 1889 (synonym: Mastodex Steiner, 1921).

Emended diagnosis (modified from Lorenzen 1981 and Decraemer & Smol 2006)

Member of Desmodoroidea. Cuticle with coarse annulations, at least in anterior body region, or

striated. Specialised somatic setae, such as adhesion tubes (Draconematidae) or ambulatory setae

(Epsilonematidae) not present. Amphideal fovea usually spiral, unispiral, cryptospiral, or cryptocircular,

and sometimes loop- or pore-shaped. Buccal cavity with small to large dorsal tooth and smaller or

indistinct subventral teeth. Pharynx usually with round muscular posterior bulb, with or without

cuticularised lumen; sometimes bulb is pyriform, elongated, or indistinct. Gubernaculum without dorsal

apophyses. Males usually monorchic, except in Onepiinema Leduc & Verschelde, 2013 (diorchic).

Vulva usually posterior to middle of body. Tail conical.

Remarks

Armenteros et al. (2014) recently argued that the genus Onepunema Leduc & Verschelde, 2013 is more

closely related to the Microlaimidae than to the Desmodoridae because of the presence of two testes (a

feature co mm on in the former but not found in the latter). The Microlaimidae, however, are characterised

by the presence of outstretched ovaries, while Onepunema possesses reflexed ovaries, a characteristic of

the Desmodoridae. This trait, in combination with the presence of a cephalic capsule in Onepunema (an

important characteristic for the classification of supraspecific taxa within the Desmodorida and never

found within the Microlaimoidea) suggests closer affinities with the Desmodoridae (and Desmodorinae

in particular) than with the Microlaimidae. Armenteros et al. (2014: 26) incorrectly stated that “... the

presence of reflexed ovaries [... ] suggest a relationship with Bolbolaimus Cobb, 1920, Calomicrolaimus

Lorenzen, 1976 and Microlaimus de Man, 1880”. Reflexed ovaries are not found in any of these genera;

perhaps Bolbolaimus was confused with Bolbonema Cobb, 1920 (see comments under Chromaspirina

longisetosa Jensen, 1985 below), which belongs to the Desmodoridae due to the presence of reflexed

ovaries and a globular cephalic capsule.

Subfamily Spiriniinae Chitwood, 1936

Type genus

Spirinia Gerlach, 1963 (synonym: Spira Bastian, 1865).

European Journal of Taxonomy 118 : 1-25 ( 2015 )

Diagnosis (from Decraemer & Smol 2006)

Member of Desmodoridae. Body cuticle with fine annulations. Head not demarcated as a cephalic

capsule. Amphideal fovea usually a simple spiral. Buccal cavity small or large, with a distinct or a

minute dorsal tooth; two small ventrosublateral teeth may be present or absent.

Differential diagnosis

This subfamily is similar to the Desmodorinae but can be differentiated from the latter by the absence

of a head capsule.

Remarks

The tribe Spirinacea was originally established by Chitwood (1936), before being raised to subfamily,

family and superfamily rank by De Coninck (1965). According to the Principle of Coordination (ICZN,

1999, Article 36), “a name established for a taxon at any rank in the family group is deemed to have been

simultaneously established for nominal taxa at all other ranks in the family group; (...). The name has

the same authorship and date at every rank.” The appropriate authorship for Spiriniinae should therefore

be attributed to Chitwood (1936), and not Gerlach & Murphy (1965), as is often done (e.g., Armenteros

et al. 2014).

Valid genera (Cavalcanti et al. 2009; Armenteros et al. 2014)

Alaimonema Cobb, 1920

Chromaspirina Filipj ev, 1918

Me tachromadora Filipj ev, 1918

Onyx Cobb, 1891

Papillonema Verschelde et al. , 1995

Paradesmodora Schuurmans Stekhoven, 1950

Parallelocoilas Boucher, 1975

Perspiria Wieser & Hopper, 1967

Poly sigma Cobb, 1920

Pseudometachromadora Timm, 1952

Sigmophoranema Hope & Murphy, 1972

Spirinia Gerlach, 1963

Spirodesma Cavalcanti et al., 2009

Stygodesmodora Blome, 1982

We provide emended diagnoses and updated lists of valid species for Chromaspirina, Spirinia, and

Perspiria below to clarify the distinctions between them. These lists are based on the original and any

subsequent descriptions; as many of the original descriptions (especially early ones) lack type material,

we often had to rely on short descriptions and/or illustrations.

Genus Chromaspirina Filipjev, 1918

Type species

Chromaspirina pontica Filipj ev, 1918.

Emended diagnosis (modified from Maria et al. 2009 and Muthumbi et al. 1995)

Robust body with rounded or rectangular head region and short conical tail; fine body annulations.

Annulated or non-annulated head region; body annulations envelop the amphideal fovea partly, entirely,

or only begin at posterior edge of amphids. Amphideal fovea uni-, crypto- or multispiral, or loop¬

shaped. Cephalic setae at anterior edge or further posteriorly alongside amphids. Distinct buccal cavity.

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

conspicuously cuticularised; large dorsal tooth and smaller subventral teeth; ventral field of denticles

may be present. Pharynx with round or oval posterior bulb, sometimes weakly developed, and without

cuticularised lumen. Precloacal supplements may be present.

Valid species

C. chabaudi Boucher, 1975. Main diagnostic characters: “... a medium-sized dorsal tooth and two small

subventral teeth. Anterior portion with twelve chelorhabdia. ” Drawings show a distinct cuticularised

dorsal tooth and subventral teeth (Boucher 1975: fig. 2).

C. crinita Gerlach, 1952. Main diagnostic character: “The dorsal tooth is massive and extraordinarily

strong.”

C. cylindricollis (Cobb, 1920). Main diagnostic characters: Only an ‘onchiunT is mentioned, without

further comment, but a distinct cuticularised dorsal tooth is visible on the drawing (Cobb 1920: 325).

C. dubia Inglis, 1968. Drawing shows a very large dorsal tooth (Inglis 1968: fig. 53).

C. gerlachi Blome, 1982. Drawing shows a distinctly cuticularised dorsal tooth and subventral teeth

(Blome 1982: fig. 20).

C. indica Gerlach, 1963. Drawing shows a very large dorsal tooth and thickly cuticularised subventral

teeth (Gerlach 1963: fig. 1).

C. inglisi Warwick, 1970. Main diagnostic character: “... a prominent heavily cuticularized dorsal

tooth...”; also clear in drawing (Warwick 1970: fig. 12).

C. lunatica Gerlach, 1965. Main diagnostic character: “...a strong dorsal tooth ... ”; also clear in drawing

(Gerlach 1965: fig. 13).

C. madagascarensis Gerlach, 1953. Main diagnostic characters: “...a rather large dorsal tooth ...”;

drawing shows a large and thick dorsal tooth and subventral teeth (Gerlach 1953: fig. 4).

C. modesta Bussau, 1993. Drawing shows a large, heavily cuticularised dorsal tooth and subventral teeth

(Bussau 1993: figs 55-57).

C. multipapillata Jayasree & Warwick, 1977. Main diagnostic characters: “Buccal cavity bearing a

dorsal tooth and two small subventral teeth”; drawing unclear. It is difficult to draw a firm conclusion

from the description and hence this species is left in this genus.

C. parapontica Luc & De Coninck, 1959. Main diagnostic characters: “Large dorsal tooth...”; drawing

shows a large dagger-like dorsal tooth (Luc & De Coninck 1959: fig. 35).

C. pellita Gerlach, 1954 (synonym: C. renaudae Boucher, 1975). Drawing shows a very large dorsal

tooth and conspicuous subventral teeth (Gerlach 1954: fig. 4).

C. pontica Filipjev, 1918. Main diagnostic character: “Three teeth...: one large dorsal..., and two small

subventrals. ”

C. thieryi De Coninck, 1943. Main diagnostic character: “...with strong dorsal tooth... ”; also clear in

drawing (De Coninck 1943: fig. 5).

European Journal of Taxonomy 118 : 1-25 ( 2015 )

C. vanreuselae Verschelde & Vincx, 1996. Main diagnostic characters: “...with huge dorsal tooth and

smaller single ventral tooth”; also clear in drawing (Verschelde & Vincx 1996: fig. 1).

Species inquirendae

C. amabilis (De Man, 1922). Considered species inquirenda by Jensen (1978).

C. cobbi Chitwood, 1938. Considered species inquirenda by Jensen (1978).

C. robusta Wieser, 1954. Considered species inquirenda by Wieser & Hopper (1967).

Remarks

Chromaspirina cylindricoUis (Cobb, 1920) (synonym: Mesodorus cylindricoUis Cobb, 1920). This

species was in our opinion rightly transferred to Chromaspirina as it was described with an annulated

head region. Maria et al. (2009), however, mistakenly showed a drawing of the head capsule of

Pseudochromadora quadripapillata Daday, 1899 (synonym: Micromicron cephalatum Cobb, 1920), in

their key to Chromaspirina species, instead of the non-annulated head region of C. cylindricoUis (Maria

et al. 2009: fig. 26).

Chromaspirina denticulata (Gerlach, 1953). This species was originally described as Bolbolaimus

denticulatus. Luc & De Coninck (1959) later synonymized Bolbolaimus with Chromaspirina. However,

Jensen (1978) rightly re-established the genus and re-instated Bolbolaimus denticulatus Gerlach, 1953

as a valid combination. However, he gave the wrong species authority in his identification key (Cobb,

1920 instead of Gerlach, 1953). Maria et al. (2009) included C. denticula (Gerlach, 1953) ( erratum ?)

in their list of invalid Chromaspirina species. In conclusion, we recognize the re-establishment of

Bolbolaimus denticulatus Gerlach, 1953 as a valid combination (as suggested by Jensen 1978) with the

synonyms Chromaspirina denticulata (Gerlach, 1953), Bolbolaimus denticulatus Cobb, 1920 syn. nov.,

and Chromaspirina denticula (Gerlach, 1953) syn. nov.

Chromaspirina dimorpha (Hopper, 1961) and C. inflexa (Wieser, 1954) were rightly transferred back to

Desmodora by Wieser & Hopper (1967).

Chromaspirina longisetosa Jensen, 1985. This species was transferred to Bolbolaimus (erratum?) by

Muthumbi et al. (1995), but later transferred to Bolbonema by Verschelde et al. (1998). Because this

species clearly has a head capsule and cephalic setae located posterior to the amphids, we confirm that it

belongs to Bolbonema. Thus, we propose the combination Bolbonema longisetosum (Jensen, 1985) with

the synonyms Chromaspirina longisetosa Jensen, 1985 and Bolbolaimus longisetosus (Jensen, 1985).

Chromaspirina lunatica Gerlach, 1965 (synonym: C. lunata Gerlach, 1965 syn. nov.). Gerlach (1965)

described C. lunatica , but the species is incorrectly labelled as C. lunata in a figure caption (p. 132).

Chromaspirina pellucida (Cobb, 1920) and C. punctata (Cobb, 1920) were rightly transferred back to

Bolbolaimus by Jensen (1978).

Chromaspirina rabosa (Gerlach, 1956). This species was originally described as Desmodora rabosa

and later transferred to Chromaspirina by the same author (Gerlach 1963). Muthumbi et al. (1995)

argued that the species should be left in the genus Desmodora because it has “... a well-developed

head capsule with the amphids situated outside the rings.” We note, however, that the cephalic setae

are situated posterior to the amphids, and this species is therefore transferred to the genus Bolbonema.

Thus, we propose the combination Bolbonema rabosum (Gerlach, 1956) comb. nov. with the synonyms

Desmodora rabosa Gerlach, 1956 and Chromaspirina rabosa (Gerlach, 1956).

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

Chromaspirina spinulosa (Wieser, 1959). This species was transferred to Metadesmodora by

Gerlach (1963) and then to Echinodesmodora by Blome (1982). Thus, we propose the combination

Echinodesmodora spinulosa (Wieser, 1959) with the synonyms Chromaspirina spinulosa (Wieser,

1959) and Metadesmodora spinulosa (Wieser, 1959).

Genus Perspiria Wieser & Hopper, 1967

Type species

Perspiria hamata Wieser & Hopper, 1967.

Emended diagnosis (modified from Vincx & Gourbault 1989)

As for Spirinia (see below), but amphids almost never completely surrounded by body annulations

(seldom more than half of amphids surrounded by body annulations), amphids are entirely surrounded

by annuli in some rare cases; tail conico-cylindrical to filiform. Precloacal supplements usually present.

Valid species

P. elongata (Castro et al. , 2006) comb. nov. (synonym: Spirinia elongata Castro et al. , 2006). Main

diagnostic characters: “...one very small subventral tooth and two small dorsal teeth.” The drawings

show amphids completely surrounded by body annulations, but the authors describe a conico-cylindrical

tail with annuli which are more evident in the tail region compared to those along the rest of the body;

thus, we transfer this species to Perspiria (Castro et al. 2006: figs 3c, 4).

P. flagellata Vitiello, 1971. Main diagnostic characters: “Amphid situated anterior to cuticle striations or

partially surrounded by them”; very long and slender, almost filiform tail (Vitiello 1971: fig. 29).

P. hamata Wieser & Hopper, 1967. Drawing shows amphids only partially located within the body

annuli; long and slender, almost filiform tail (Wieser & Hopper 1967: fig. 18).

P. lara (Silva et al., 2009) comb. nov. (synonym: Spirinia lara Silva et al. , 2009). Main diagnostic

characters: “Buccal cavity with one dorsal tooth and two minute ventrosublateral teeth” and “Cuticle with

transverse striae..., less obvious in head region than in tail.” Drawing shows amphid with annulations to

mid-level of amphid only, and tail is described as conico-cylindrical. This species is therefore transferred

to Perspiria (Silva et al. 2009: figs 2-3).

P. megamphida Vincx & Gourbault, 1989. Drawing shows substantial portion of the amphids is located

on the non-annulated lip region; conico-cylindrical tail with more prominent annuli (Vincx & Gourbault

1989: fig. 1).

P. mokii Coles, 1987. Main diagnostic character: “...cuticle appears smooth... ”, hence amphids are not

surrounded by annuli; long and slender filiform tail (Coles 1987: fig. 5).

P. papillata Vincx & Gourbault, 1989. Drawing shows anterior half of amphids on non-annulated lip

region and posterior half located within the annulated head region; long, conico-cylindrical tail (Vincx

& Gourbault 1989: fig. 2).

P. septentrionalis (Cobb, 1914) comb. nov. (synonyms: Spirinia septentrionalis (Cobb, 1914) and Spira

septentrionalis Cobb, 1914). Main diagnostic characters: “From the anus the tail tapers for two-thirds

of its length, then becomes cylindroid to the swollen terminus... ” Drawings by Wieser (1954: fig. 126)

clearly show the amphids in the head region with only their posterior half within the body annulations.

European Journal of Taxonomy 118 : 1-25 ( 2015 )

leaving the anterior half on the non-annulated head region. The tail is conico-cylindrical. Thus, this

species is transferred to the genus Perspiria.

P. striaticaudata (Timm, 1962). Tail long (11-17 abd) and slender with distinct annuli; thus it is

transferred to Perspiria. Synonym: Spirina striaticaudata Timm, 1962.

Species inquirenda

Perspiria sabulicola (Filipjev, 1918/1921) comb. nov. (synonym: Spirina sabulicola Filipjev, 1918/

1921). This species was described with a Tong tail (length 4 x width)’ and hence should be transferred

to the genus Perspiria. However, as the species was described based on only one female, it is regarded

as a species inquirenda.

Genus Spirinia Gerlach, 1963

Type species

Spira parasitifera Bastian, 1865.

Emended diagnosis

Robust body with rounded or conical head region and conical tail, fine body annulations. Annulated

head region; body annulations normally completely surround the amphideal fovea, with few exceptions.

Spiral amphideal fovea. Buccal cavity narrow, lightly cuticularised; no or small/inconspicuous dorsal

tooth, no or minute subventral teeth. Pharynx with small pyriform, oval, or rounded posterior bulb

without cuticularised lumen. Precloacal supplements usually absent. Short conical tail.

Valid species

Spirinia gerlachi (Tuc & De Coninck, 1959). Main diagnostic characters: “Annulations beginning

halfway down the lateral organ... long attenuated conical tail”; also clear in drawings (Luc & De Coninck

1959: fig. 25). Despite the presence of an amphid only partially surrounded by cuticle annulations, this

species is left within Spirinia because of the shape of the tail (conical), which is the primary trait for

differentiating between Spirinia and Perspiria.

Spirinia gnaigeri Ott, 1977. Amphids almost entirely located within the body annulations in one male,

but only the posterior half of another male’s amphid is surrounded by body annulations (compare figs 42

and 43 in Ott 1977); however, as the species has a conical tail, it belongs within Spirinia.

S. guanabarensis (Maria et al., 2009) comb. nov. (synonym: Chromaspirina guanabarensis Maria et

al., 2009). Main diagnostic characters: “Buccal cavity large with an acute dorsal tooth and two smaller

ventrosublateral teeth. ” However, later in the text it is stated that: “ Chromaspirina guanabarensis sp.

nov. belongs to the group of species characterized by a poorly developed dorsal tooth.” Drawings also

show a small buccal cavity with a small dorsal tooth and minute subventral teeth (Maria et al. 2009: fig.

1). Tail is conical. This species is therefore transferred to Spirinia.

S. hopperi Coles, 1987. Main diagnostic characters: “Cuticle without striations... small dorsal tooth is

present in the narrow mouth cavity.” Drawings show a conical tail (Coles 1987: fig. 3).

S. inaurita (Wieser & Hopper, 1967) comb. nov. (synonym: Chromaspirina inaurita Wieser & Hopper,

1967). Main diagnostic characters: “Buccal cavity with three small teeth (these teeth are smaller than in

most other species of Chromaspirina and stress the proximity of the genus to Spirinia ).”; the drawings

are inconclusive, but the description clearly describes small teeth. This species is therefore transferred

to Spirinia.

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

S. laevioides Gerlach, 1963. The drawing shows a small buccal cavity, a small dorsal tooth, and amphids

located within the annulated head region (Gerlach 1963: fig. 2).

S. laevis (Bastian, 1865) (synonym: S. pilosa Allgen, 1935). The drawing shows small teeth, amphids

located entirely within the annulated head region, and a short conical tail (Bastian 1865: fig. 204).

S. okemwai (Muthumbi et al. , 1995) comb. nov. (synonym: Chromaspirina okemwai Muthumbi et al.,

1995). We studied the type material, which shows that this species has a cylindrical pharynx with only

a slightly swollen terminal bulb, whose shape cannot be described. The teeth are difficult to distinguish

and hence cannot be called Targe’ or ‘distinct’; therefore, we transfer this species to the genus Spirinia.

S. parasitifera (Bastian, 1865). Some descriptions of this species show the amphids being only partially

surrounded by annulations (Gerlach 1963; Vincx & Gourbault 1989), and others describe specimens

with amphids entirely surrounded by annulations (Coles 1987; Vincx & Gourbault 1989); furthermore,

additional descriptions and drawings depict either a conical tail (Gerlach 1963: fig. 1; Coles 1987: fig.

3) or a prolonged conical to long conical tail (Wieser & Hopper 1967: fig. 17). Hence, until further

examination and comparison of these different specimens can be done, we leave this species within

Spirinia.

S. parma (Ott, 1972) comb. nov. (synonym: Chromaspirina parma Ott, 1972). Main diagnostic

characters: “... stoma conical, with a small acute dorsal tooth and two minute ventral teeth...”; this

species is therefore transferred to the genus Spirinia.

S. schneideri (Villot, 1875). Luc & Deconinck (1959) describe this species with a smooth cuticle, and

Coles (1987) describes it “without cuticular striations”. The SEM micrograph by Coles (1987: fig. 9a),

however, shows striations surrounding the amphids. The tail is short and conical.

S. sophia Silva et al ., 2009. Main diagnostic characters: “Buccal cavity with one dorsal tooth and two

minute, ventrosublateral teeth”; clear in drawings, which also show the amphid completely surrounded

by cuticle annulations (Silva et al. 2009: fig. 6). Tail conical.

Species inquirendae

Spirinia granulata (Allgen, 1929) comb. nov. (synonym: Spirina granulata Allgen, 1929). This species

was described based on two juveniles only, with very poor drawings and inadequate descriptions. Hence,

we have to regard this species as species inquirenda.

S. paucispira (Schuurmans Stekhoven, 1950) comb. nov. (synonym: Chromaspirina paucispira

Schuurmans Stekhoven, 1950). This species was considered species inquirenda by Wieser & Hopper

(1967) and Muthumbi et al. (1995), because the original description was based on a single female

specimen. It was stated that the buccal cavity of this species has an “indication of a dorsal tooth”. Based

on this character, which is sufficient to distinguish between Spirinia and Chromaspirina , this species

is transferred to the genus Spirinia (but remains species inquirenda due to the incomplete nature of the

original description).

S. similis (Cobb, 1898) (synonym: Spira similis Cobb, 1898). Considered invalid by Coles (1987) due

to lack of illustrations.

S. tenuicauda (Allgen, 1959) (synonym: Spirina tenuicauda Allgen, 1959). This species has to be

regarded as a species inquirenda , as the only description Allgen made was to write “differing from Sp.

parasitiferaC and his drawings are inadequate.

European Journal of Taxonomy 118 : 1-25 ( 2015 )

Spirinia verecunda sp. nov.

urn:lsid:zoobank.org:act:83F4CF38-29CF-4540-B05D-2245120A489B

Figs 1-3; Table 1

Diagnosis

Spirinia verecunda sp. nov. is characterised by a short, stout body, cuticle covered in minute hair-like

structures, unispiral amphideal aperture and cryptospiral amphideal fovea, buccal cavity with a small

dorsal tooth and minute subventral teeth, eight elongated glands surrounding the anterior portion of the

pharynx, large sperm cells, spicules with weak capitulum, and absence of precloacal supplements.

Type material

Holotype

NEW ZEALAND: NIWA 88379, 20 Feb. 2011, NIWA cruise TAN1103, station 69, central Chatham

Rise, 43.331° S, 178.288° E, 350 m.

Paratypes

NEW ZEALAND: 2 $ NIWA 88380, same data as holotype.

Etymology

The species name is derived from the Latin word verecundus (bashful, shy, coy), in reference to the

anterior part of the head region deeply folded inwards.

Description

Male

Body stout, cylindrical, pale orange colour, tapering slightly towards both ends. Annulated cuticle, no

lateral differentiation; annulations 0.6-0.8 pm apart; cuticle slightly thicker in pharyngeal region than

elsewhere (2.2-2.5 vs 1.6-2.0 pm). Cuticle covered with dense layer of minute hair-like structures from

posterior to amphid to near tail tip (barely visible under light microscope; Fig. 3A-B); larger, 1x5 pm

bacteria-like structures with narrow base also attached in pharyngeal region (Fig. ID). A single protist,

74 pm long and 18 pm wide attached subdorsally near cloaca (Fig. 3E). Short somatic setae arranged in

eight longitudinal rows along entire body length.

Blunt, slightly rounded annulated head region with annulations completely surrounding amphid (Fig.

1A-B). Four cephalic setae near anterior edge of amphid; inner and outer labial sensillae not observed

because labial region is folded inwards from level of cephalic setae. Cryptospiral amphideal fovea with

slightly cuticularised outline and unispiral amphideal aperture (Fig. 3A-B).

Buccal cavity with slightly cuticularised dorsal tooth and minute subventral teeth. Eight elongated glands,

12-17 pm in length, distributed around anterior portion of pharynx just posterior to buccal cavity (Figs

1C-D, 3C). Pharynx muscular, oval-shaped posterior pharyngeal bulb, lumen not cuticularised. Nerve

ring at 40-50% of pharynx length. Secretory-excretory system not observed. Cardia 10-12 pm long.

Reproductive system monorchic, with relatively large, outstretched testis situated to the left of intestine.

Mature sperm large, globular to elongated in shape, up to 16 pm in length. Short, arcuate spicules with

weak capitulum and tapering distally; velum present (Fig. 1G). Gubernaculum with cuneus broadest

in middle portion and tapering proximally and distally; poorly developed lateral crurae (Fig. 1H).

Precloacal supplements not observed. Tail conical, with two pairs of short subventral setae and one pair

of subdorsal setae; three caudal glands restricted to tail region and spinneret present.

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

Fig. 1 . Spirinia verecunda sp. nov. A. Surface view of male head. B-C. Male head, showing buccal

cavity. D. Female head, showing bacteria attached to cuticle. E. Anterior body region of male.

F. Anterior body region of female. G. Posterior body region of male. H. Gubernaculum, with details of

crurae (left) and cuneus (right). I. Posterior body region of female. J. Posterior body region of female,

showing attached protist. Scale bar: A-D, I = 20 pm, E-F = 30 pm, G = 16 pm, H = 12 pm, J = 35 pm.

European Journal of Taxonomy 118: 1-25 (2015)

Female

Similar to male, but with wider body. One specimen bears a single protist (suctorian) on tail. Reproductive

system didelphic, amphidelphic with reflected ovaries situated either both on the right or both on the

left of intestine. Vulva located slightly post median. Cuticular pars distalis vaginae and pars proximalis

vaginae surrounded by constrictor muscle.

Fig. 2. Spirinia verecunda sp. nov. A. Entire male. B. Female reproductive system. Scale bar: A = 50

pm, B = 30 pm.

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

Fig. 3. Spirinia verecunda sp. nov., light micrographs. A. Male head, showing amphideal aperture and

and cuticular hair-lilce structures. B. Male head, showing amphideal fovea. C. Male anterior body region

showing buccal cavity and elongated glands. D. Mature sperm cells. E. Posterior body region of male,

showing copulatory apparatus and attached protist. Scale bar: A-B, D-E = 10 pm, C = 8 pm.

European Journal of Taxonomy 118: 1-25 (2015)

Table 1. Morphometries (pm) of Spirinia verecunda sp. nov. and Stygodesmodora confusa sp. nov.

— a, body length/maximum body diameter; abd, anal body diameter; b, body length/pharynx length; c,

body length/tail length; cbd, corresponding body diameter; V, vulva distance from anterior end of body;

%V, V/total body length.

Spirinia verecunda sp. nov.

Stygodesmodora confusa sp. nov.

Holotype

Paratypes

Holotype

Paratype

Paratypes

757

667, 675

1241

795

778-1029

22-34

6,7

8-10

12-15

Head diameter

17, 18

14-16

Length of cephalic setae

7-8

6-9

5-9

Amphid height

6,7

5-6

Amphid width

6,7

6-7

Amphid width/cbd (%)

33,41

40^17

Amphid from anterior end

3-7

Nerve ring from ant. end

45, 53

57-62

Nerve ring cbd

32, 33

24-25

Pharynx length

100

100, 107

109

92-105

Pharyngeal bulb diameter

20-22

Pharyngeal bulb cbd

33, 34

25-28

Max. body diameter

39, 40

30-37

Spicule length

Gubemaculum length

Anal body diameter

21,22

17-20

Tail length

51,52

62-85

Tail length/abd

2.0

2.4

3.3

2.7

3.5M.5

347, 361

428-558

51,54

53-55

Vulval body diameter

39, 40

30-37

Remarks

Spirinia verecunda sp. nov. shows similarities with several Spirinia and Chromaspirina species, i.e.,

Spirinia guanabarensis comb, nov., S. inaurita comb, nov., S. okemwai comb, nov., S. parma comb,

nov., Chromaspirina chabaudi , and C. multipapillata. Of these, S. verecunda sp. nov. resembles S.

okemwai comb. nov. the most in cuticle ornamentation (annulated head region, cuticle with minute hair¬

like structures), size and shape of the amphids, presence of large sperm cells, and absence of precloacal

supplements. S. verecunda sp. nov. differs from the latter in having shorter body length (667-757 vs

966-1425 pm), lower values of a (17-22 vs 22-27), shorter cephalic setae (2 vs 3-6 pm), shorter

spicules (38 vs 41-50), and shorter gubemaculum (13 vs 19-25). S. verecunda sp. nov. also differs from

S. okemwai comb. nov. in the presence of elongated glands in the anterior region of the pharynx (absent

in S. okemwai comb, nov.) and the structure of the gubemaculum (hook absent vs dorsally directed

proximal hook). S. verecunda sp. nov. can most readily be distinguished from S. inaurita comb. nov.

and S. parma comb. nov. by the size of the amphids (in males: 35% cbd in S. vercunda sp. nov. vs >

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

60% cbd in S. inaurita comb. nov. and S. parma comb, nov.), from Chromaspirina chabaudi by the

structure of the gubernaculum (not surrounding spicules in S. verecunda sp. nov. vs surrounding spicules

distally in C. chabaudi ), from C. multipapillata by the absence of precloacal supplements (present in C.

multipapillata ), and from S. guanabarensis comb. nov. by its small body size (< 800 pm in S. verecunda

sp. nov. vw >1800 pm in S. guanabarensis comb, nov.) and low values of a (17-22 vs 30M8).

Genus Stygodesmodora Blome, 1982

Type species

Stygodesmodora epixantha Blome, 1982.

Emended diagnosis (modified from Blome 1982)

Cuticle with coarse annulations; numerous long somatic setae may be present; annulated head region

(sometimes intraspecific variation, with the head of some specimens only partially annulated in dorsal

and ventral regions). Amphid unispiral or spiral located on amphideal plate; small to medium sized

buccal cavity with dorsal tooth and (usually) smaller subventral teeth. Pharynx with rounded or pyriform

posterior bulb, without cuticularised lumen. Pre-cloacal supplements present or absent; spicules short,

arcuate, and with well-developed capitulum; gubernaculum present.

Valid species

S. bacillicauda (Gerlach, 1963)

S. epixantha Blome, 1982

Species inquirenda

S. rotundicephala (Cobb, 1920)

Stygodesmodora confusa sp. nov.

urn:lsid:zoobank.org:act:44B8B268-21AE-4E30-8A7E-D707BlD54C8F

Figs 4-7; Table 1

Diagnosis

Stygodesmodora confusa sp. nov. is characterised by a relatively short body (778-1241 pm), spiral

amphids with 1.0-1.25 turns, cephalic setae situated at or slightly posterior to mid-level of amphid, and

males with four precloacal supplements consisting of short setae on wide bases.

Type specimens

Holotype

NEW ZEALAND: <$, NIWA 88377, 24 Apr. 2010, NIWA cruise TAN1004, station 92, canyon axis on

southern Hikurangi Margin, 41.8921° S, 174.6347° E, 686 m.

Paratypes

NEW ZEALAND: 1 2 $ ?, NIWA 88378, 20 Feb. 2011, NIWA cruise TAN1103, station 69, central

Chatham Rise, 43.331° S, 178.288° E, 350 m; 3 $$ (same data as other paratypes) were measured to

provide morphometric data and then processed for scanning electron microscopy.

Etymology

The species name is derived from the Latin adjective confusus, meaning confused, perplexed or obscure

(feminine form used, as -dora is feminine), and refers to the close affinities of this species with other

desmodorid genera (see Remarks), which resulted in initial confusion in the placement of this species.

European Journal of Taxonomy 118: 1-25 (2015)

Fig. 4. Stygodesmodora confusa sp. nov. A. Anterior body region of female. B. Male head (holotype).

C. Male head (paratype). D. Male head (holotype). E-F. Anterior body region of female. G. Female

posterior body region. H. Right spicule and gubernaculum. I. Male copulatory apparatus. Arrows show

position of pre-cloacal supplements. Scale bar: A-D = 30 pm, E-G = 36 pm, H = 20 pm, I = 40 pm.

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

Fig. 5. Stygodesmodora confusa sp. nov. A. Entire male. B. Female reproductive system. Scale bar:

A = 50 pm, B = 36 pm.

European Journal of Taxonomy 118: 1-25 (2015)

Description

Male

Relatively short cylindrical body, pale golden colour, tapering slightly towards both ends. Cuticle 0.9-

1.4 pm thick, with coarse annuli, approximately 1 pm apart, no lateral differentiation. Ventral ala absent.

Somatic setae of different sizes (4-19 pm long; Fig. 5A), set out in eight longitudinal rows in anterior

one fifth of body length and in six longitudinal rows along rest of body except on tail, where they are

irregularly arranged.

Annulated head region with annulations completely surrounding amphid (Figs 6A-C, 7C); in some

specimens, there is a seemingly non-annulated head region, but some partial annulation of the dorsal

®-Q' '' 1 1

Fig. 6. Stygodesmodora confusa sp. nov., light micrographs of female. A. Head region showing

amphideal aperture. B. Head region showing amphideal fovea. C. Head region showing buccal cavity.

D. Vulva. Scale bar: A-C = 15 pm, D = 18 pm.

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

Fig. 7. Stygodesmodora confusa sp. nov. Scanning electron micrographs. Female head (three different

specimens). Scale bar = 5 pm.

European Journal of Taxonomy 118: 1-25 (2015)

and ventral sides of the head can be observed (Fig. 7A-B); both forms are considered annulated head

regions, even though the degree of annulation differs between the two morphotypes. Labial region

folded inwards in all specimens. Six papilliform outer labial sensillae and four cephalic setae, 44-57%

cbd long, situated at or slightly posterior to mid-level of amphid. Spiral amphideal fovea, 1.2-1.4 turns,

located on a cuticularised amphideal plate (Figs 6B, 7B), larger in holotype than in paratype (~0.6 vs 0.4

cbd, respectively; Fig. 4C-D); amphideal aperture with similar shape to amphideal fovea but tapering

proximally (compare Fig. 6A and 6B).

Buccal cavity small, with small dorsal tooth; subventral teeth not observed (Fig. 6C). Pharynx muscular,

slightly swollen at anterior extremity, and with pyriform bulb at posterior extremity. Nerve ring at 60-

62% of pharynx length. Secretory-excretory system not observed. Cardia short.

Reproductive system monorchic, with single anterior testis situated to the right of intestine, 53-63

pm long, outstretched and opposed. Mature sperm small, globular, 2-3 x 3-5 pm. Short, arcuate

spicules tapering distally; well-developed capitulum and broad velum (Fig. 4H). Small, plate-shaped

gubernaculum without crurae. Four pre-cloacal supplements consisting of short setae on broad bases

(Fig. 41), 23-45 pm apart, beginning 15 pm anterior to cloaca. Tail conical, with three small caudal

glands and spinneret.

Female

Similar to males. Female reproductive system didelphic, amphidelphic with reflected ovaries situated

either to the right or left of intestine, with anterior and posterior ovaries always situated on opposite

sides. Vulva located slightly post median. Cuticular pars distalis vaginae and pars proximalis vaginae

surrounded by constrictor muscle. Mature eggs 66-73 x 25 pm.

Remarks

Stygodesmodora confusa sp. nov. can most easily be differentiated from the other two species of the genus

by the location of the cephalic setae at mid-level of amphids (vs* anterior to amphids in S. bacillicauda

and S. epixantha). S. confusa sp. nov. also differs from S. bacillicauda in the presence of pre-cloacal

supplements (absent in S. bacillicauda) and the absence of lateral crurae on the gubernaculum (present

in S. bacillicauda ), and from S. epixantha by the presence of numerous long somatic setae (short somatic

setae in S. epixantha).

Stygodesmodora confusa sp. nov. closely resembles species of other desmodorid genera, and

distinguishing it from these other species requires careful examination of the head region. S. confusa

sp. nov. resembles Echinodesmodora moensi Verschelde & Vincx, 1996 in the absence of a cephalic

capsule, numerous long somatic setae, and cephalic setae at or slightly posterior to mid-level of amphids,

but can be differentiated from the latter by the absence of pharyngeal lumen cuticularisation (present

in E. moensi ), amphid shape (< 1.25 turns vs 1.5-2.0 turns in E. moensi ), presence of setose precloacal

supplements (absent in E. moensi ), and the absence of knotty protrusions on the tail (present in E.

moensi). The most important distinction, however, is the presence of an amphideal plate in S. confusa

sp. nov. This species is also superficially similar to species of Bolbonema , i.e., B. brevicolle (Cobb,

1920), B. longisetosum (Jensen, 1985) comb, nov., and B. spiralis Hourston & Warwick, 2010, in the

presence of both short and long somatic setae, cephalic setae just posterior to mid-level of amphid, and

in the structure of the copulatory apparatus. The amphideal plates in S. confusa sp. nov. give a swollen

appearance to the head region laterally (see Fig. 7A), and in some specimens there is a gap in the body

annulations in the head region (compare Fig. 7A and 7C), which may be wrongly interpreted as a non-

annulated head region or even a head capsule. S. confusa sp. nov., however, is clearly defined by an

annulated region, and not a head capsule, because (1) the swollen part of the head is mostly restricted

LEDUC D. & VERSCHELDE D., New Desmodoridae from the Southwest Pacific

to the area surrounding the amphids (i.e., the amphideal plates), (2) the cuticle is the same thickness

in head and body regions (see Fig. 6C), and (3) annulations are present dorsally and ventrally at least

to mid-level of amphids, with one continuous annulation anterior to the amphid (see Figs 6B, 7A-C).

The genus Bolbonema , however, is characterized by a globular head capsule (i.e., conspicuously thicker

cuticle and absence of annulations; Verschelde & Vincx 1996; Verschelde et al. 1998).

Discussion

Muthumbi et al. (1995) provided an emended diagnosis for Chromaspirina and, more recently, Silva et al.

(2009) provided an emended diagnosis for Spirinia. Maria et al. (2009) briefly discussed the relationship

between Chromaspirina and Spirinia. They identified the size of the buccal cavity and dorsal tooth, and

the shape of the posterior pharyngeal bulb as the main distinguishing features between the two genera

and noted that several Chromaspirina species have a small dorsal tooth which “emphasises the close

relationship of this genus with Spirinia.. .” and that “further investigation is needed into the relationships

[of] Spirinia with Chromaspirina ” (Maria et al. 2009: 28). In his review of Spirinia , Coles (1987)

identified buccal morphology as the main distinguishing feature between the two genera.

We argue that the shape of posterior pharyngeal bulb and the presence or absence of precloacal

supplements show too much overlap to be useful diagnostic features to distinguish Spirinia and

Chromaspirina. For example, S. okemwai comb. nov. and S. guanabarensis comb. nov. lack precloacal

supplements (a trait usually associated with Spirinia), but possess relatively weakly developed, oval¬

shaped posterior pharyngeal bulbs (a trait usually associated with Chromaspirina ; see Maria et al.

2009). S. sophia Silva et al. , 2009 also appears to be intermediate between the two genera: it possesses

precloacal supplements (absent in all other Spirinia species but present in some Chromaspirina species)

and a round to oval-shaped posterior pharyngeal bulb (despite Spirinia being defined as having a round

posterior pharyngeal bulb; see Silva et al. 2009). Differences in the morphology of the pharyngeal bulb

and precloacal supplements are not reliable traits to distinguish between Chromaspirina and Spirinia

because (1) differences in the shape of pharyngeal bulbs are often subtle (e.g., pyriform vs elongated

or oval) and (2) precloacal supplements in these two genera are often difficult to distinguish and may

have been omitted in some species descriptions. Buccal morphology provides the clearest and most

consistent way to differentiate between Spirinia and Chromaspirina because it can easily be observed;

morphological features of the head region are also the most important traits for differentiating between

desmodorid genera.

Perspiria was originally described as a subgenus of Spirinia by Wieser & Hopper (1967). The only

distinguishing feature of this subgenus given by the latter authors was “the more prominently striated

and flagellate tail.” Coles (1987) also gave the same distinguishing features between the two subgenera.

Vincx & Gourbault (1989) later raised Perspiria to generic status and included two key characteristics in

their diagnosis of the genus: amphids only partially surrounded by cuticle annulations and a “prominently

striated” and “filiform” tail, which we interpret as a coarsely annulated, conico-cylindrical tail from the

drawings (see figs 1 and 2 in Vincx & Gourbault 1989). Some Spirinia species, however, have coarsely

annulated (e.g., S. verecunda sp. nov.) or conico-cylindrical tails (e.g., S. septentrionalis (Cobb, 1914)),

and others have amphids not completely surrounded by the body annulations (e.g., S. lara Silva et al .,

2009). Here, we propose to differentiate between Spirinia and Perspiria primarily based on tail shape

(conical vs conico-cylindrical to filiform), as well as position of amphids relative to cuticle annulations

(entirely vs partially surrounded), and have suggested nomenclatural changes to reflect these distinctions.

The monospecific genus Spirodesma was recently described by Cavalcanti et al. (2009) from continental

slope sediments in the Southwest Atlantic. The authors argued that the genus is most similar to

Chromaspirina based on the rounded shape of the head, annulated cuticle, spiral amphideal fovea, and

European Journal of Taxonomy 118: 1-25 (2015)

conical tail. We consider these arguments invalid because (1) an annlated cuticle is also a characteristic

of some Spirinia (e.g., Spirinia verecunda sp. nov.) and Perspiria species (e.g., P. papillata Vincx &

Gourbault, 1989), (2) the spiral amphideal fovea in Spirodesma is in fact very similar in shape to the

cryptospiral amphideal fovea found in some Spirinia species (e.g., S. verecunda sp. nov.) and to the

spiral amphideal fovea of Perspiria papillata , and (3) the tail of Spirodesma is conico-cylindrical (as

stated in the original description), not conical. In addition, head shape is not a particularly informative

trait for differentiating between genera. In our opinion, Spirodesma is most closely related to Perspiria

because of the similar buccal cavity (narrow with small or minute teeth), the position of the amphids

(only partially surrounded by cuticle annulations), and the shape of the tail (conico-cylindrical). The only

difference between these genera appears to be the relative size of the teeth: Perspiria is characterised

by a narrow, lightly cuticularised buccal cavity with no or a small/inconspicuous dorsal tooth and no

or minute subventral teeth, while Spirodesma is characterised by a narrow, lightly cuticularised buccal

cavity with three small, equal teeth. Spirodesma and Perspiria may therefore need to be synonymised in

the future as more information becomes available on the former genus.

Stygodesmodora rotundicephala (Cobb, 1920) was originally described as Amphispira rotundicephala

and was included in the World Register of Marine Species list of Stygodesmodora species at the time

the present study was conducted (WoRMS Editorial Board 2014). The original description, which is

based on a single juvenile specimen, clearly shows a head capsule, whereas no amphideal plate was

described. Based on the drawing, the specimen/species could possibly belong in Zalonema , but, as the

original species description of Amphispira rotundicephala was based only on one juvenile, we regard A.

rotundicephala Cobb, 1920 to be a species inquirenda.

Acknowledgements

Funding was provided by NIWA’s Coasts and Oceans Centre Research Programme 2 (2013/14 SCI)

and the programme ‘Impact of Resource Use on Vulnerable Deep-sea Communities’ (CO 1X0906) and

“Consequences of Earth-Ocean Change” (CO 1X0702). We are grateful to Scott Nodder (NIWA) for

facilitating sampling on Chatham Rise (TAN 1103), and to Norliana Rosli for processing TAN 1004

samples. We also acknowledge the other participants of NIWA voyages TAN1004 and TAN1103, and

the officers and crew of RV Tangaroa.

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