Onuphis and Aponuphis (Annelida: Onuphidae) from southwestern Europe, with the description of a new species

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2015 Magnolia Press

Zootaxa 3949 (3): 345–369

www.mapress.com/zootaxa/Article

http://dx.doi.org/10.11646/zootaxa.3949.3.3

http://zoobank.org/urn:lsid:zoobank.org:pub:CBEF804D-5FBF-48D2-8CCD-036B70FE6ECD

Onuphis and Aponuphis (Annelida: Onuphidae) from southwestern Europe,

with the description of a new species

ANDRÉS ARIAS1,4 & HANNELORE PAXTON2,3

1Departamento de Biología de Organismos y Sistemas (Zoología), Universidad de Oviedo, Oviedo 33071, Spain Department of Biological Sciences, Macquarie University, Sydney, NSW 2109, Australia3Australian Museum Research Institute, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia4Corresponding author. E-mail: [email protected]

Abstract

This study recognises five species of Onuphis and four species of Aponuphis from southwestern Europe, represented in

the Bay of Biscay, the Atlantic coasts of Iberia and the western Mediterranean Sea. One species, Onuphis anadonae sp.

nov., is newly described and another species, Aponuphis willsiei Cantone & Bellan, 1996 is a new record for the Iberian

Peninsula waters. Aponuphis ornata (Fauvel, 1928) and A. willsiei are shown to be tube brooders and the development of

the juveniles is described. The usefulness of some Onuphis characters, as well as the size-dependent distribution of bran-

chiae and chaetae, complicating Aponuphis species identification, are discussed. We are presenting descriptions and pho-

tographs of the distinctive Aponuphis colour pattern which we recommend as an identificatory tool instead. A

dichotomous key to all species is included.

Key words: polychaetes, Eunicida, biodiversity, direct development, ontogenetic changes, chaetal replacement, Iberian

Peninsula, Bay of Biscay, western Mediterranean

Introduction

Onuphis Audouin & Milne Edwards, 1833 is the most speciose genus of the polychaete family Onuphidae Kinberg, 1865 after Diopatra Audouin & Milne Edwards, 1833 with about 40 described valid species (Maekawa & Hayashi 1999; Gil & Machado 2014; Arias & Paxton 2014). However, the genus has only been the subject of partial revisions (Fauchald 1982; Maekawa & Hayashi 1999; Arias & Paxton 2014) and it is quite possible that more undescribed species may be discovered in the near future. The genus Onuphis is a cosmopolitan taxon and most of its species are characteristic for the intertidal and relatively shallow waters of warm-temperate regions, nevertheless individual species of Onuphis seem to have limited restricted geographical distributions (Paxton 1986).

The genus Aponuphis Kucheruk, 1978 was erected for a group of slender onuphids without peristomial cirri that had previously been regarded as members of the genus Hyalinoecia Malmgren, 1867. Paxton (1986) recognised seven species of Aponuphis and underscored the morphological differences between the two genera by placing Aponuphis into the subfamily Onuphinae and Hyalinoecia into the subfamily Hyalinoeciinae.

Southwestern Europe, represented by the Bay of Biscay, the Atlantic coasts of Iberia and the western Mediterranean Sea, harbours five species of Onuphis, of which one is new to science, and four species of Aponuphis. Here, we present an updated revision of the Onuphis and Aponuphis fauna of this region with the description and illustration of the new species of Onuphis, newly report A. willsiei from the Iberian Peninsula and present notes on the ecology and reproduction together with a taxonomic key to the species of Onuphis and Aponuphis of southwestern Europe.

Accepted by P. Hutchings: 30 Mar. 2015; published: 28 Apr. 2015 345

Material and methods

The majority of the studied material was obtained from the benthic samples of two Iberian oceanographic campaigns: i) the COCACE (Oceanographic Cruise of the Central Cantabrian Sea) project, conducted from April 1987 to February 1988 in a variety of substrates in the southern Bay of Biscay, north of the Iberian Peninsula; ii) the campaigns Fauna Ibérica I, II, III and IV (1989–1996), a major project aimed to investigate the animal biodiversity in the Iberian-Balearic region. In the Material Examined sections of the species descriptions all stations (localities) with collection data and additional information are fully listed.

Other collections of Onuphis and Aponuphis specimens from different localities along the Atlantic and Mediterranean coasts of the Iberian Peninsula were provided by the Museo Nacional de Ciencias Naturales (MNCN) of Madrid, Spain.

Specimens were examined under both dissecting stereomicroscope and compound light microscope. Temporary glycerol slides of small specimens and parapodia were used for the detailed examination of chaetal distribution and the morphology of parapodia. Line drawings were made with the aid of a camera lucida and digital photography.

Selected specimens were prepared for examination with Scanning Electron Microscopy (SEM). All samples were dehydrated in an ascending series of graded ethanol. Thereafter, they were submersed in acetone, critical point dried using acetone as intermediate liquid, mounted on aluminium stubs, sputter coated with gold and analysed with a JEOL 6610 LV scanning electron microscope. Photomicrographs were taken with a Spot Flex Digital Camera System mounted on a Leica MZ 16 stereo dissection microscope.

Chaetal and prostomial appendages terminology follows Paxton (1986; 1998 respectively). The specimens examined in this study are deposited in the Museo Nacional de Ciencias Naturales, Madrid

(MNCN), Australian Museum, Sydney (AM), Muséum National d’Histoire Naturelle, Paris (MNHN), The Natural History Museum, London (BMNH), Musée Océanographique de Monaco (MOM), and Naturhistorisches Museum, Wien (NHMW).

Taxonomic account

Family Onuphidae Kinberg, 1865

Subfamily Onuphinae Kinberg, 1865

Genus Onuphis Audouin & Milne Edwards, 1833

Onuphis Audouin & Milne Edwards, 1833: 225.

Type species. Onuphis eremita Audouin & Milne Edwards, 1833: 226 by subsequent designation of Malmgren 1866: 180. Gender: feminine.

Diagnosis. Prostomium often anteriorly extended; with frontal lips. Antennae and palps with ceratophores with 10–25 rings and short to moderately long styles, palpal styles shorter than their ceratophores. Nuchal grooves straight. Peristomial cirri present. Anterior three to four (rarely two or five) pairs of parapodia modified but not enlarged. Ventral cirri subulate on anterior four to six chaetigers; dorsal cirri moderately long. Branchiae rarely absent, usually present from chaetiger 1, rarely 3–6; single or pectinate filaments (maximum 12). Hooks of modified parapodia usually tridentate (rarely only bidentate, sometimes bi- to multidentate) pseudocompound with short hoods; median hook slightly larger but not becoming simple and changing to large median hook. Dorsal limbate chaetae from chaetiger 1, ventral limbate chaetae replacing pseudocompound hooks from chaetiger 4 or later until replaced by bidentate hooded subacicular hooks usually from chaetiger 10–12. Tubes round in section, ranging from thin mucous to tough parchment-like inner layer covered with extraneous particles.

ARIAS & PAXTON346 · Zootaxa 3949 (3) © 2015 Magnolia Press

Onuphis anadonae sp. nov.

Figures 1–5; Table 1

Material examined. Type material. Holotype (MNCN 16.01/16159) COCACE station: D3 (43.85º N, 5.68 ºW), N. Spain, Cantabrian Sea, Bay of Biscay, 162 m, 89.06 % sand, 4.76 % silt, 6.18 % clay, 29 Jun 1987; 1 paratype

(MNCN 16.01/16160) COCACE station: D3 (43.85º N, 5.68 ºW), N. Spain, Cantabrian Sea, Bay of Biscay, 162 m, 89.06 % sand, 4.76 % silt, 6.18 % clay, 29 Jun 1987; 1 paratype (AM W.47513) COCACE station: B2 (43.70º N, 5.77º W), N. Spain, Cantabrian Sea, Bay of Biscay, 86 m, 76.84 % sand, 14.95 % silt, 8.21 % clay, 5 Jul 1987.

Type locality. Off Asturias, Cantabrian shelf, 43.85º N, 5.68 ºW, 162 m depth, northern Spain, Bay of Biscay, East Atlantic.

Diagnosis. Colour pattern absent on dorsum of anterior chaetigers; peristomium with one brown transverse band. Prostomium almost rounded. Antennae moderately short to chaetiger 3–7; ceratophores strongly ringed with maximum of six to seven rings. Peristomial cirri not exceeding anterior margin of prostomium. First four pairs of parapodia modified, directed slightly anterolaterally with bi-, tri- and quatridentate pseudocompound hooks. Subulate ventral cirri in first four chaetigers; distinct subulate postchaetal lobes in first 10 chaetigers. Subacicular hooks from chaetiger 9. Branchiae as single filament from first chaetiger. Tube parchment-like and externally covered with sand grains and shell fragments.

Description. Description based on holotype, with variation of paratypes included. Holotype complete with 223 chaetigers, consisting of two fragments, anterior one with 82 chaetigers and posterior fragment with 141 chaetigers; total length of 145 mm and width at chaetiger 10 (excluding parapodia) of 2.7 mm (Figs 1, 3). Both paratypes incomplete with 79 chaetigers, 43 mm long and 2.8 mm wide and 62 chaetigers, 37 mm long and 2 mm wide respectively.

Body cream coloured in preserved condition with one transverse dorsal pigment band on peristomium (Fig. 1). Prostomium globular with pair of conical frontal lips (Figs 1A–C). Large pair of eyespots present, situated between bases of lateral antennae and palps. Palps reaching chaetiger 1–2 with nine basal rings and longer distal one. Lateral antennae reaching chaetiger 6–7 with six basal rings, median antenna reaching chaetiger 3–4 with five basal rings, all three antennae with longer distal ring (Figs 1A, 2A, B). Sensory buds of antennae and palps scarce and very inconspicuous, arranged in irregular rows (Figs 2E, F). Peristomial cirri inserted distally on peristomium below lateral antennae, shorter than length of peristomium not exceeding anterior margin of prostomium (Figs 1C, 2A, C).

Anterior chaetigers (1–3) slighty longer than those following (Figs 1A, 2C, 3A). First four pairs of parapodia modified, not enlarged, directed slightly anterolaterally, with low prechaetal fold, triangular prechaetal lobe and spindle-shaped postchaetal lobe, longer than base of parapodium; subulate dorsal cirrus about as long as postchaetal lobe, ventral cirrus shorter than postchaetal lobe (Figs 3B, C, 4A). Subulate ventral cirri in first four chaetigers and pad-like thereafter. Postchaetal lobe as distinct subulate lobe in first 10 chaetigers, becoming smaller and conical in shape thereafter.

Branchiae as single filament, strap-like, from first chaetiger to about chaetiger 180, lacking in last 40 chaetigers (in holotype). Branchial filament becoming longer from chaetiger 20–22, from chaetiger 29–31 longest three times as long as anterior ones (Fig. 4B); then getting smaller by chaetiger 150, being absent from chaetiger 183. Aciculae yellowish with pointed tips, generally three per parapodium. Hooded pseudocompound hooks in first four chaetigers. First two parapodia (Fig. 3B) with following chaetal complement going from superior to inferior part of chaetal fan: one simple chaeta, three protruding distal tips of aciculae, three strongly tridentate pseudocompound hooks (Figs 3H, I, 4D), two quatridentate (Fig. 4E) or tridentate pseudocompound hooks with most proximal denticle much smaller and protruding directly from base of second distal tooth (Fig. 3G) and one bidentate pseudocompound hook (Fig. 3F). Parapodia 3 and 4 with six pseudocompound tridentate hooks (Fig. 4C). Hoods of pseudocompound hooks falcate with pointed tip (Figs 4C, D). All hooks of equal thickness without slender long-appendaged ones. Pseudocompound hooks replaced by limbate chaetae from chaetiger 5 (Fig. 4F). Pectinate chaetae flat and slightly oblique with seven to nine long teeth each, starting in chaetiger 5 (Figs 3J, 4G). Hooded bidentate subacicular hooks from chaetiger 9. Mandibles (Fig. 3L) with white calcified cutting plates and slender shafts. Maxillae (Fig. 3K) weakly sclerotised; maxillary formula (based on paratype): Mx I = 1 + 1; Mx II = 8 + 6; Mx III = 5 + 0; Mx IV = 5 + 5; Mx V = 1+1. Mx VI absent. Tube cylindrical in shape, parchment-like and externally covered with sand-grains and shell fragments (Fig. 1D).

Zootaxa 3949 (3) © 2015 Magnolia Press · 347ONUPHIS AND APONUPHIS FROM SOUTHWESTERN EUROPE

FIGURE 1. Onuphis anadonae sp. nov. Holotype (MNCN 16.01/16159). Photomicrographs. A, anterior end, dorsal view; B, anterior end, ventral view; C, detailed view of prostomium and peristomium; D, tube.

Remarks. Onuphis anadonae sp. nov. is characterised by having single strap-like branchiae from chaetiger 1, shared with nine other known Onuphis species (Table 1). It is the only species with bi-, tri- and a few quadridentate pseudocompound hooks. Three other species with single branchiae from chaetiger 1 (O. amakuensis, O. elegans

and O. kammurijamaensis) have bi- and tridentate pseudocompound hooks in the first four chaetigers, and O.

farallonensis has these hooks on chaetigers 4–6. However, O. anadonae sp. nov. differs from these species by lacking any brown bands, spots or other kind of colour pattern on the anterior chaetigers, by presenting bidentate pseudocompound hooks only in the first two chaetigers, instead of in the first four like in other species, as well as


FIGURE 2. Onuphis anadonae sp. nov. Paratype (MNCN 16.01/16160). Scanning electron micrographs. A, detailed view of prostomium and peristomium, dorsal view; B, detailed view of prostomium, lateroventral view; C, anterior end, lateral view; D, detailed view of nuchal groove; E, detailed view of style of antenna; F, detailed view of a sensory bud of the antenna. c, ceratophore; fl, frontal lip; la, lateral antenna, ma, median antenna; ng, nuchal groove; ll, lower lip; p, palp; up, upper lip.


FIGURE 3. Onuphis anadonae sp. nov. Holotype A (MNCN 16.01/16159), paratype B-L (AM W47513). Line drawings. A, anterior end, dorsal view; B, parapodium 2, anterior view; C, parapodium 4, anterior view; D, parapodium 5, anterior view; E, parapodium 20, anterior view; F, bidentate pseudocompound hook, chaetiger 2; G, slighly tridentate pseudocompound hook, chaetiger 2; H, I, tridentate pseudocompound hooks, chaetiger 4; J, pectinate chaeta, chaetiger 5; K, maxillae; L, mandibles.


FIGURE 4. Onuphis anadonae sp. nov. Paratype (MNCN 16.01/16160). Scanning electron micrographs. A, parapodium 2, lateral view; B, median chaetigers, lateral view; C, pseudocompound hooks of chaetiger 1; D, detailed view of tridentate pseudocompound hooks of chaetiger 3; E, detailed view of a quatridentate pseudocompound hook of chaetiger 2; F, limbate chaetae from chaetiger 5; G, pectinate chaeta from chaetiger 5; H, subacicular hooks from chaetiger 9. b, branchia; dc, dorsal cirrus; pol, postchaetal lobe; prl, prechaetal lobe; SAH, subacicular hook; vc, ventral cirrus.


FIGURE 5. Onuphis anadonae sp. nov. Paratype (MNCN 16.01/16160). Scanning electron micrographs. A, parapodium 5, lateral view; B, median chaetigers, lateral view, arrows point to Epistylis epibionts; C, Epistylis colony on parapodia of anterior body end; D, detailed view of Epistylis sp. showing the corona of cilia and its peduncle. b, branchia; dc, dorsal cirrus; gp, glandular pad; pol, postchaetal lobe; prl, prechaetal lobe; SAH, subacicular hook.

quatridentate pseudocompound hooks and by the early origin of the subacicular hooks, starting in chaetiger 9 in O.

anadonae sp. nov. and in chaetiger 10–11, in the other species. Onuphis anadonae sp. nov. also differs from these four species in having peristomial cirri shorter than the peristomium, while they are as long as or longer than the peristomial length in the others. Other differences are with respect to the length of the antennae, the maximum number of rings of antennal ceratophores, the number of chaetigers with subulate ventral cirri and their length, and are summarised in Table 1.

Etymology. It is a pleasure to name this species in honour of Professor Nuria Anadón, who in many ways helped to make this study possible.


TA

BL

E 1

. Com

paris

on o

f key

feat

ures

and

hab

itats

of t

he O

nuph

is sp

p. c

hara

cter

ized

by

havi

ng si

ngle

stra

p-lik

e br

anch

iae

from

cha

etig

er 1

. PC

, per

isto

mia

l cirr

i; PC

H, p

seud

ocom

poun

d ho

ok; P

SL, p

ostc

haet

al

lobe

; SA

H, s

ubac

icul

ar h

ook;

VC

, ven

tral c

irrus

.

Cha

ract

er

O. a

mak

uens

is

Mae

kaw

a &

H

ayas

hi, 1

999

O. a

nado

nae

sp. n

ov.

O. e

lega

ns

(Joh

nson

, 190

1)O

.fara

llone

nsis

H

obso

n, 1

971

O. h

olob

ranc

hiat

a M

aren

zelle

r, 1

879

O. i

rides

cens

(J

ohns

on,

1901

)

O.k

amm

urija

mae

nsis

Mae

kaw

a &

Hay

ashi

, 19

89

O. o

palin

a (V

erri

ll, 1

873)

O. r

ullie

riana

(A

mou

reux

, 19

77)

O.sh

irik

ishi

naie

nsis

(I

maj

ima,

196

0)

Col

our

patt

ern

on

ante

rior

cha

etig

ers

brow

n do

rsal

ba

nds

abse

nt

brow

n do

rsol

ater

al sp

ots

redd

ish

brow

n

dors

al b

ars

brow

n do

rsal

ban

d an

d pa

tche

s nea

r ba

se o

f pa

rapo

dium

Bro

wn

dors

al

bars

Li

ght b

row

n do

rsal

bar

Bro

wn

dors

al

band

s ab

sent

B

lack

dor

sal b

ars

Med

ian

ante

nna

reac

hing

cha

etig

er

8 3-

4 2

10

7 9

-10

4 5-

7 9

5

Lat

eral

ant

enna

e re

achi

ng c

haet

iger

8

6-7

5-7

13

11

13-1

4 7

10-1

3 12

7

Max

imum

num

ber

of r

ings

on

ante

nnal

ce

rato

phor

es

Max

imum

num

ber

of r

ings

on

palp

al

cera

toph

ores

8 11

7 10

11 10

12 8

15 10

13-1

4

14

12 9

12 11

16 10

11 9

PC le

ngth

as

long

as

peris

tom

ium

sh

orte

r tha

n

peris

tom

ium

as

long

as

peris

tom

iun

long

er th

an

peris

tom

ium

lo

nger

than

pe

risto

miu

m

long

er th

an

peris

tom

ium

lo

nger

than

pe

risto

miu

m

long

er th

an

peris

tom

ium

lo

nger

than

pe

risto

miu

n as

long

as

peris

tom

ium

L

engt

h of

VC

of

mod

ified

cha

etig

ers

as lo

ng a

s PSL

sh

orte

r tha

n PS

L lo

nger

than

PSL

shor

ter t

han

PSL

as lo

ng a

s PSL

as

long

as P

SLsh

orte

r tha

n PS

L as

long

as P

SLas

long

as P

SLlo

nger

than

PSL

Cha

etig

ers w

ith

subu

late

VC

4

4 5

(6)

5 4

6 4

6 (7

) 5

(6)

7 (6

-8)

Cha

etig

ers w

ith

subu

late

PSL

10

10

12

(13)

10

10

16

11

10

10

(12)

16

Cha

etig

ers w

ith

PCH

Tip

of P

CH

4

bi- a

nd

tride

ntat

e

4

bi-,

tri- a

nd

quat

riden

tate

4

bi- a

nd tr

iden

tate

4 (6

)

bi- a

nd tr

iden

tate

4

tride

ntat

e

4

tride

ntat

e

4

bi- a

nd tr

iden

tate

4

tride

ntat

e

4

tride

ntat

e

7

bi- a

nd tr

iden

tate

Slen

der

long

-ap

pend

aged

hoo

ks

abse

nt

abse

nt

pres

ent

pres

ent

abse

nt

abse

nt

pres

ent

pres

ent

abse

nt

pres

ent

Ori

gin

of S

AH

11

9

10-1

1 10

11

-13

12

11

12-1

5 12

- 14

11-1

3

Typ

e lo

calit

y A

mak

usa

Shim

ojim

a Ja

pan

E. P

acifi

c

Can

tabr

ian

shel

f Sp

ain

E. A

tlant

ic

Tow

nsen

d of

f W

ashi

ngto

n U

SA

W. P

acifi

c

Sout

hern

C

alifo

rnia

U

SA

W. P

acifi

c

Off

Japa

n E.

Pac

ific

Off

Brit

ish

Col

umbi

a U

SA

W. P

acifi

c

Wak

asa

Bay

Ja

pan

E. P

acifi

c

Off

New

En

glan

d U

SA

W. P

acifi

c

Engl

ish

Cha

nnel

Fr

ance

E.

Atla

ntic

Sout

h H

okka

ido

Japa

n E.

Pac

ific

Hab

itat

inte

rtida

l 86

-162

m

inte

rtida

l and

sh

allo

w su

btid

al68

m

149

m

inte

rtida

l and

sh

allo

w

subt

idal

72 m

20

0 m

86

0-14

00 m

in

terti

dal

Ref

eren

ce

Mae

kaw

a &

H

ayas

hi 1

999

pres

ent s

tudy

H

obso

n 1

971;

Fa

ucha

ld 1

982;

H

ilbig

199

5

Hob

son

1971

Im

ajim

a 19

86

Hob

son

1971

; Fa

ucha

ld 1

982;

Hilb

ig 1

995

Mae

kaw

a &

Hay

ashi

19

89

Fauc

hald

198

2;

Imaj

ima

1986

A

mou

reux

19

77; p

rese

nt

stud

y

Imaj

ima

1960


Biology and faunal associates. The holotype contains oocytes in its body cavity, ranging from 200 to 250 µm in diameter. Ectosymbiotic microorganisms were found on the surface of the anterior and medial parapodia of O.

anadonae sp. nov. (Figs 5A, B). These epibionts were identified as peritrich ciliates belonging to the genus Epistylis Ehrenberg, 1830 characterised by forming colonies utilising its noncontractile and longer peduncle or stalk for attachment to the worm cuticle (Figs 5C, D). The symbiotic association “onuphid-peritrich” was previously reported for Diopatra marocensis Paxton et al., 1995 and Onuphis eremita Audouin & Milne-Edwards, 1833 (Arias et al. 2010; Arias & Paxton 2014 respectively) suggesting that this association may occur commonly among onuphid polychaetes.

Distribution and ecology. Onuphis anadonae sp. nov. is known from the southern part of the Bay of Biscay (East Atlantic) from the Cantabrian shelf, depth range 86–162 m in sandy substrate. This species was only collected singly at two stations, indicating a very low population density and suggesting that it is apparently a rare species with a restricted distribution in the Bay of Biscay. The specimens previously incorrectly reported as Aponuphis holobranchiata (= Onuphis holobranchiata Marenzeller, 1879) from the Basque shelf, northern Spain (Martínez et al. 2010) may also represent O. anadonae sp. nov.

Onuphis eremita Audouin & Milne Edwards, 1883

Onuphis eremita Audouin & Milne Edwards, 1833: 226; Redondo & San Martín, 1997: 227; Arias & Paxton, 2014: 148–158, figs. 1–8.

Onuphis falesia Castelli, 1982: 45, fig. 1–11 (Gulf of Follonica, Italy). Onuphis eremita oculata.—Çinar, 2009: 2297–2299, fig. 4 (Turkey). Not Hartman, 1951.

Material examined. Type material. Neotype (MNHN TYPE 1555) La Rochelle, France, East Atlantic, 46°09’N, 01°09’W–46°09’N, 01°10’W, intertidal sands, coll. 28 Sep 2011.

Non-type material. For a detailed listing of additional material from France, Cantabrian and Mediterranean coasts of Spain and Malta see Arias & Paxton (2014).

Type locality. La Rochelle, France, Bay of Biscay, East Atlantic.Diagnosis. Prostomium anteriorly extended, one pair of small eyespots present at anterolateral end of

prostomium. Palps reaching chaetiger 2–4, lateral antennae reaching chaetiger 7–14 and median antenna chaetiger 4–7. Ceratophores long and strongly ringed, palpal ceratophores with 16–20 rings, lateral antennae with 19–22 and median antenna with 13–19 rings. Subulate ventral cirri in first five to six chaetigers; distinct subulate postchaetal lobes in first 10–17 chaetigers. Small interramal papilla present at base of dorsal cirrus in chaetigers 4(5)–9. Strongly tridentate pseudocompound slender, long and robust, short appendaged hooks in first three chaetigers. Subacicular hooks from chaetiger 10. Flat, distally oblique pectinate chaetae with 10–13 teeth. Single branchial filaments from chaetiger 1 to 20–26, thereafter number increasing rapidly to maximum of three to five. Colour pattern consisting of two rows of brownish patches on dorsal anterior end or brown transverse stripes when lateral patches coalesce medially. Tube thin-walled and covered with sand grains.

Remarks. The taxonomic identity of O. eremita, the type species of the genus, was extremely controversial. It was confounded by vague descriptions, redescriptions based on worldwide material and the establishment of dubious synonymies over the last century. However, a neotype was recently designated and the species was redescribed by Arias & Paxton (2014). These authors redefined its diagnostic features, restricted its distribution range to the European Atlantic and the Mediterranean Sea, making its conspecificity with the specimens reported from outside Europe and the Mediterranean very unlikely.

Distribution. East Atlantic (Cantabrian Sea, Bay of Biscay) and western and central Mediterranean. However, it is most likely that the species is widely distributed along the Atlantic coasts of Iberia and the Levantine basin of the Mediterranean Sea.

Onuphis farensis Gil & Machado, 2014

Onuphis farensis Gil & Machado, 2014: 345–356, figs. 2–6. (Ria Formosa, Portugal).Onuphis (Nothria) geophiliformis.—Ibáñez 1972: 26–28, fig. 2 (Cádiz, Spain); 1973 (Isla Cristina, Spain). Not Moore, 1903.Nothria geophiliformis.—Campoy 1982. Not Moore, 1903.


? Nothria geophiliformis.—Dexter 1992: 79 (Ria de Alvor, Portugal). Not Moore, 1903.? Onuphys geophyliformis.—Mucha & Costa 1999: 371 (Ria de Aveiro and Sado Estuary, Portugal). Not Moore, 1903.

Material examined. Non-type material. Mooreonuphis sp. (registered as Nothria geophiliformis), 1 specimen (MNCN 16.01/3329) off shore near San Francisco beach, Santiago Island, Cabo Verde archipelago, western Africa, 50 m, 22 Aug 1985; 1 specimen (MNCN 16.01/3330) Curral Velho, Boavista Island, Cabo Verde archipelago, western Africa, 26 Aug 1985.

Type locality. Ria Formosa coastal lagoon, Portugal, southwestern Iberian Peninsula, East Atlantic.Diagnosis. Prostomium anteriorly extended. Eye spots absent. Lateral antennae twice as long as median one.

First four chaetigers slightly enlarged. Subulate ventral cirri in first four chaetigers. Tri- and quatridentate pseudocompound hooded hooks present in first four chaetigers. Pectinate chaeta from chaetiger 5 with about ten teeth. Subacicular hooks present from chaetiger 9. Simple strap-like branchiae present from chaetiger 5 to the last 30–40 chaetigers. Tube thin-walled, covered with small sand grains.

Remarks. Gil & Machado (2014) described Onuphis farensis from southern Portugal and referred previous reports of Onuphis geophiliformis (Moore, 1903) from Isla Cristina and Cádiz by Ibáñez (1972; 1973) and Campoy (1982) to the new species. The reports of Nothria geophiliformis from Ria de Alvor and Onuphys geophyliformis

from Ria de Aveiro and Sado Estuary, Portugal by Dexter (1992) and Mucha & Costa (1999) respectively were considered as doubtful.

We agree with these statements and are adding the following considerations based on literature reports, personal communications and the examination of preserved material. The only specimens identified as Onuphis

geophiliformis or its original combination Nothria geophiliformis Moore, 1903, from the East Atlantic that were available to us for reviewing were those reported from Cabo Verde archipelago by López & San Martín (1992). However, the re-examination of these specimens revealed that they are not members of the genus Onuphis but of Mooreonuphis Fauchald, 1982 and represent a different species to the only one known for the East Atlantic to date, M. vespa Arias et al. 2013, constituting most likely an undescribed species.

Louzao et al. (2010) reported N. geophiliformis from the Cantabrian slope at a depth of 1025 m, based upon the polychaete fauna collected by the COCACE (Oceanographic Cruise of the Central Cantabrian Sea) project. We have re-examined this material, which is deposited in the Collection of the Department of Biology of Organisms and Systems at University of Oviedo (Spain), and found that these specimens are actually O. rullieriana

(Amoureux, 1977), a typical deep-sea E. Atlantic species which is treated in detail below. Moreover, O.

geophiliformis or its synonyms have been repeatedly reported in marine species lists from the Basque Country (e.g. Borja et al. 2000), being recently considered as an alien or nonindigenous species (NIS) for the region (e.g. Martínez & Adarraga 2006). We have examined the onuphid material from the Basque shelf collected by the oceanographic campaigns of Fauna Ibérica and have not found O. geophiliformis-like specimens. In view of the data we have so far, records of O. geophiliformis from the Cantabrian Sea should be considered as doubtful.

Throughout its long history O. geophiliformis has been the subject of dubious taxonomic affiliations and interpretations, e. g. the number of anterior chaetigers with pseudocompound hooks reported for O. geophiliformis

sensu lato varied among authors, from three (Moore 1903; Fauchald 1982; Maekawa & Hayashi 1999) or four (Ibáñez 1972; 1973; Gil & Machado 2014) to six (Day 1967). Onuphis farensis differs from the original description of O. geophiliformis from Japan in that the latter has pseudocompound hooks on the anterior three instead of anterior four chaetigers, and the origin of subacicular hooks is on chaetigers 10–12 instead of chaetiger 9 (Moore 1903). It also seems unlikely that this species is the same as O. geophiliformis reported from southern Africa by Day (1967) since the latter has pseudocompound hooks in the first six chaetigers and the ventral cirri are subulate in the first five chaetigers. We cannot pursue the matter further since we have been unable to locate again the species in southern Iberia (due to the changes which have taken place in “Cabezuela de Cádiz” and nearby areas in recent years as a consequence of the enlargement of the Port of Cádiz) and the reported specimens from Cádiz and Isla Cristina are most probably not extant (Ibáñez, pers. comm.).

Distribution. East Atlantic, southern Spain to Portugal.

Onuphis pancerii Claparède, 1868

Onuphis pancerii Claparède, 1868: 438–440, pl. VIII fig. 1; 1870: 387, pl. V fig. 5a; Rioja, 1918: 39, fig. 10 (Santander); Arias & Paxton, 2014: 158–164, figs 9–12.


Onuphis eremita.—Fauvel, 1923 (in part): 413–415, fig. 163. Not Audouin & Milne Edwards, 1833.—Paxton, 1986: 56–58, figs 3, 6–8, 10. Not Audouin & Milne Edwards, 1833.

Material examined. Type material. Neotype (MNCN 16.01/311) Naples, Italy, specimen Num. 173 from the Collection of Rioja, 1894-1917.

Non-type material. Nine specimens (MNCN 16.01/310) Enmedio sandy flat, Santander Bay, Spain, specimens Num. 686 from the Rioja Collection 1894-1917. For a detailed listing of additional material from Italy, Spain and France see Arias & Paxton (2014).

Type locality. Gulf of Naples, Italy, central Mediterranean.Diagnosis. Prostomium anteriorly extended. Eye spots absent. Palps reaching chaetiger 1–2, lateral antennae

reaching chaetiger 5–6 and median antenna chaetiger 2–4. Ceratophores long and strongly ringed, palpal ceratophores with 22–26 rings, lateral antennae with 21–29 and median antenna with 15–20 rings. Subulate ventral cirri in first five to six chaetigers; distinct subulate postchaetal lobes in first 10–19 chaetigers. Small interramal papilla present between chaetigers 4–5 to 9. Bi- and tridentate (or even multidentate) pseudocompound hooks in first three chaetigers. All hooks of equal thickness, slender long-appendaged ones absent. Subacicular hooks from chaetiger 10. Flat, distally oblique pectinate chaetae with 13–17 teeth. Single branchial filaments from chaetiger 1 to 19–24, thereafter number increasing to maximum of six to eight. Tube with soft inner layer and outer layer of vegetal material, shell-fragments and sand grains.

Remarks. This species was synonymised with O. eremita by Fauvel (1923) and remained as such for nearly a century. However, Arias & Paxton (2014) demonstrated that O. pancerii is actually a valid species, morphologically different from O. eremita. Although the two species are closely related and thus share a number of parapodial and chaetal characteristics, they can easily be differentiated in that O. pancerii is a much larger, robust species than O. eremita. The anterior parapodia of O. pancerii have bi- and tridentate (or even multidentate) pseudocompound hooks of similar thickness and length of appendages. In contrast, O. eremita has only tridentate pseudocompound hooks of two types, robust short-appendaged hooks and slender long-appendaged ones.

Distribution. East Atlantic (Bay of Biscay) and western and central Mediterranean.

Onuphis rullieriana (Amoureux, 1977)

Figure 6

Nothria rullieriana Amoureux, 1977: 399–402, fig. 1.Onuphis rullieriana.—Fauchald, 1982: 51–53, fig. f–g, table 17; Aguirrezabalaga et al. 2002: 27 (Capbreton Canyon, Bay of

Biscay).

Type locality. Entrance of the English Channel, 860–1400 m depth, East Atlantic.Material examined. Non-type material. One specimen (AM W27379) Capbreton Canyon, Bay of Biscay,

979–990 m depth, 7 Jul 1988; 4 specimens (MNCN 16.01/15245) N. Prieto Cape, Asturias (43º 49.10’ N, 4º 52.74’ W–43º 49.33 N, 4º 53.47’ W), Cantabrian Sea, Bay of Biscay, 925–1025 m depth, 25 Jun 1991; 2 specimens (AM W.47514) N. Prieto Cape, Asturias (43º 49.10’ N, 4º 52.74’ W–43º 49.33 N, 4º 53.47’ W), Cantabrian Sea, Bay of Biscay, 925–1025 m depth, 25 Jun 1991; 3 specimens coated with gold (MNCN 16.01/15246) N. Prieto Cape, Asturias (43º 49.10’ N, 4º 52.74’ W–43º 49.33 N, 4º 53.47’ W), Cantabrian Sea, Bay of Biscay, 925–1025 m depth, 25 Jun 1991; 2 specimens (MNCN 16.01/16161) COCACE station: I2 (43.99º N, 5.83º W), N. Spain, Cantabrian Sea, Bay of Biscay, 1025 m, 58.44 % sand, 17.56 % silt, 24.00 % clay, 2 Jul 1987.

Diagnosis. Prostomium anteriorly extended. Eye spots absent. Palps reaching chaetiger 1–2, lateral antennae reaching chaetiger 5–16 and median antenna chaetiger 3–14. Ceratophores with nine to 10 rings in palps, 10–13 in lateral antennae and eight to ten rings in median antenna. Subulate ventral cirri in first five to six chaetigers; subulate postchaetal lobes distinct in first 12 chaetigers. Tridentate pseudocompound hooks with triangular pointed hoods present in first four chaetigers. All hooks of equal thickness, slender long-appendaged ones absent. Pectinate chaeta from chaetiger 5 with 13–15 teeth. Subacicular hooks present from chaetiger 14. Simple strap-like branchiae present from chaetiger 1 and continuing throughout.

Remarks. Examined specimens ranged in width between 1.4 to 2.5 mm. Almost complete specimens over 61 mm in length with 78 chaetigers. Some preserved specimens presented a colour pattern consisting of a brown bar


on the prostomium, brown spots at the base of ceratophores and styles of antennae and palps and brown spots at the base of dorsal cirri, postchaetal lobes and parapodial bases of anterior chaetigers; other specimens were cream coloured without pigmentation. The pseudocompound hooks of the anterior modified chaetigers of O. rulleriana

have relatively long and triangular hoods with pointed tips (Figs 6A–D). These long hoods are uncharacteristic for the genus Onuphis, which usually has short falcate-hooded hooks. The hooks have three stout and blunted teeth, with the intermediate tooth bigger than the others (Figs 6C, D). The appendages of all hooks are almost equally long; however the hooks from the middle of fascicle are slightly thicker than the others and weakly tridentate (Fig. 6D). Pectinate chaetae occur from chaetiger 5, with 13–15 teeth and are slightly curved on the margins, usually two per parapodium (Fig. 6E).

Fauchald (1982), in his revision based upon type material, pointed out that within five paratypes of O.

rulleriana examined by him, four had branchiae first present from chaetiger 1 but in the remaining one the appearance of a single branchial filament occurred at chaetiger 3. Thus, he concluded that the appearance of branchiae ranges from chaetiger 1 to 3 in this species. We have examined 15 specimens and all have branchiae starting from chaetiger 1 so we consider that the specimen with branchiae starting from chaetiger 3 most likely represents an abnormal individual. Therefore, we maintain within the species diagnosis the presence of branchiae from chaetiger 1, according to the original description and following Aguirrezabalaga et al. (2002).

FIGURE 6. Onuphis rullieriana. Scanning electron micrographs. A, tridentate pseudocompound hook from the lower fascicle of chaetiger 1; B, tridentate pseudocompound hook from the middle of fascicle of chaetiger 1; C, detailed view of tip of pseudocompound hook from the lower fascicle of chaetiger 2; D, detailed view of tip of weakly tridentate pseudocompound hook from the middle fascicle of chaetiger 2; E, pectinate chaetae from chaetiger 6.

Distribution. East Atlantic off Britain and the Bay of Biscay (925–1025 m deep). The species was reported from Wasaka Bay (Japan) on the basis of two specimens collected from 150–257 m depth (Maekawa & Hayashi 1989). However after a decade, the same authors in a subsequent paper reconsidered their initial identification and transferred the identity of this species to Onuphis sp. (Maekawa & Hayashi 1999).

Genus Aponuphis Kucheruk, 1978

Aponuphis Kucheruk, 1978: 91.Type species: Hyalinoecia bilineata Baird, 1870: 358 by original designation. Gender: feminine.

Diagnosis. Prostomium anteriorly extended; with oval frontal lips, short palps and long antennae, ceratophores with up to 20 rings; peristomial cirri absent. Anterior three to six (rarely seven to eight) pairs of parapodia modified but not enlarged, with bi- to tridentate (rarely quadridentate) pseudocompound to compound hooks; median hooks


slightly larger. Ventral cirri subulate on four to seven chaetigers. Dorsal limbate chaetae from chaetiger 1, ventral limbate chaetae replacing hooks from chaetiger 4 or later until replaced by bidentate hooded subacicular hooks from chaetiger 9–15. Branchiae, rarely absent, usually present from chaetiger 4, rarely 1–7; single filament. Tubes thin, consisting of inner mucous and outer layer of sand or broken shells.

Remarks. Specific identification of Aponuphis has been problematic. Too much emphasis was placed on too few diagnostic characters, such as branchiae and the pseudocompound hooks of the anterior modified parapodia. This was compounded by the fact that these structures are size-dependent during ontogeny and in some species the adult state of a character is only attained when the worm has reached two-thirds of its maximum size. Although the species display distinctive colour patterns, there is intraspecific variation of patterns due to varying degrees of pigmentation. Two schools of thought developed with respect to the nomenclature of the group. Bellan (1964) and Amoureux (1971; 1976) considered all subspecies and related species of A. bilineata as ecological “forms” and synonyms of that species, while among others Laubier & Paris (1962), Glémarec (1969) and Guille (1971) maintained the validity of the different species.

Fauvel (1923: 424) listed Aponuphis rigida (Claparède, 1868) as Hyalinoecia bilineata var. rigida from the Bay of Biscay, Santander and Madeira. However, we have been unable to trace this statement to a primary record. We are here reporting four species of Aponuphis from southwestern Europe, of which A. willsiei is a new record for the Iberian Peninsula waters.

Aponuphis bilineata (Baird, 1870)

Figure 7A, B

Hyalinoecia bilineata Baird, 1870: 358; Rioja 1918: 44 (Santander, Spain); Bellan 1960: 18 (Portugal); Amoureux 1973: 440 (northern Spain, Bay of Biscay); 1974: 137 (Aveiro, Portugal); Campoy 1982: 544 (in part).

Hyalinoecia rubra Langerhans, 1880: 292 (Madeira).Aponuphis bilineata.—Kucheruk 1978: 91 (new combination); Aguirrezabalaga et al. 2002: 27 (Capbreton Canyon, Bay of

Biscay).

Material examined. Type material. Three syntypes of Hyalinoecia bilineata (BMNH ZB 1867.1.7.2) off coast of Cornwall, U.K., 73 m; 2 syntypes of Hyalinoeia rubra Langerhans, 1880 on slides (NHMW 2347) Madeira.

Non-type material. Five specimens (MNCN 16.01/16162) COCACE station: D3 (43.85º N, 5.68º W), N. Spain, Cantabrian Sea, Bay of Biscay, 162 m, 89.06 % sand, 4.76 % silt, 6.18 % clay, 29 Jun 1987; 5 specimens (MNCN 16.01/16163) COCACE station: C1 (43.74º N, 5.69º W), N. Spain, Cantabrian Sea, Bay of Biscay, 150 m, 80.04 % sand, 8.28 % silt, 11.68 % clay, 26 Jun 1987; 17 specimens (MNCN 16.01/16164) COCACE station: B3 (43.70º N, 5.90º W), N. Spain, Cantabrian Sea, Bay of Biscay, 117 m, 69.86 % sand, 12.77 % silt, 17.37 % clay, 5 Jul 1987; 7 specimens (MNCN 16.01/16165) COCACE station: B3 (43.70º N, 5.90º W), N. Spain, Cantabrian Sea, Bay of Biscay, 117 m, 69.86 % sand, 12.77 % silt, 17.37 % clay, 5 Jul 1987; 1 specimen (MNCN 16.01/16166) COCACE station: C7 (43.81º N, 5.98º W), N. Spain, Cantabrian Sea, Bay of Biscay, 146 m, 98.58 % sand, 0.40 % silt, 1.02 % clay, 4 Jul 1987; 15 specimens (MNCN 16.01/15242, MNCN 16.01/15269, MNCN 16.01/15270, MNCN 16.01/15277, MNCN 16.01/15281) Fauna II, NW Peñas Cape, Asturias (43º 43.72’–43º 42.82’ N, 5º 56.97’–5º 56.85’ W), N. Spain, E. Atlantic, 122–129 m, 15 Jun 1991; 1 specimen (MNCN 16.01/15243) Fauna II W Cíes Islands, Galicia (42º 15.26’– 42º 16.31’ N, 9º 20.63’– 9º 21.59’ W), NW Spain, E Atlantic, 232–240 m, 11 Jun 1991; 2 specimens (MNCN 16.01/15275, MNCN 16.01/15279) Fauna II S. Finisterre Cape, Galicia (42º49.77'–42º49.81' N, 9º13.91'–9º13.81' W), NW. Spain, E. Atlantic, 54–67 m, 11 Jun 1991; 1 specimen (MNCN 16.01/15273) Fauna II NW. Villano Cape, País Vasco (43º28.62'–43º28.64' N, 2º53.77'–2º53.36' W), N. Spain, E. Atlantic, 88 m, 21 Jun 1991; 1 specimen (MNCN 16.01/15271) Fauna IV off shore Alborán Island (35°49.91'–35°50.36' N, 03°14.63'–03°13.72' W), E. Spain, Alborán Sea, western Mediterranean, 118 m, 25 Jul 1996; 4 specimens (MNCN 16.01/15272, MNCN 16.01/15274, MNCN 16.01/15278) Fauna IV off shore Alborán Island Lighthouse (35°58.00'–35°58.00' N, 02°58.46'–02°58.46' W), E. Spain, Alborán Sea, western Mediterranean, 36 m, 25 Jul 1996; 2 specimens (MNCN 16.01/1834, MNCN 16.01/1836) off shore between San Antonio cape and Valencia harbour, E. Spain, western Mediterranean, 3–30 m, 26 Apr 1996; 1 specimen (MNCN 16.01/4371) Denia, E. Spain, western Mediterranean, 3–30 m, 5 Apr 1998.


FIGURE 7. Aponuphis spp. photomicrographs showing its characteristic colour patterns. A, A. bilineata morph 1 (typical); B. A. bilineata morph 2; C, A. brementi; D–E, A. ornata; G, recently settled juvenile of A. ornata; H, brooding juvenile of A. ornata.


Type locality. Off Cornwall, U.K., 73 m, East Atlantic.Diagnosis. Antennae usually to chaetiger 10–15, maximally to 22; four to six ceratophoral rings on median

antenna, seven to 10 usually on lateral antennae, maximally 11. Anterior five to six chaetigers with several bi- to tridentate pseudocompound hooks, chaetigers 6–7 with one only; slender long-appendaged hooks present. Ventral cirri subulate on first five chaetigers. Subacicular hooks from chaetiger 10. Branchiae from chaetiger 4–5. Species occurring as two colour morphs: Morph (1), peristomium completely pigmented or four separate spots, two longitudinal pigment stripes and two segmental lateral spots from chaetiger 1 to posterior region, stripes becoming discontinuous between chaetigers, forming two segmental spots as pigmentation decreases posteriorly (Fig. 7A), referred to previously as typical (Bellan 1964). Morph (2), peristomium and first three to four chaetigers with two large brownish spots almost filling dorsal surface of segment, sometimes coalescing medially; from chaetiger 4–5 to median region two longitudinal dorsal pigment stripes discoursing laterally close to parapodia (Fig. 7B). Maximum width without parapodia 1.5 mm.

Remarks. The three syntypes of Hyalinoecia bilineata Baird, 1870 have been examined and found to be in good condition, but the pigmentation is not visible any more. Photographs of the two syntypes (on slides) of Hyalinoecia rubra Langerhans, 1880 were studied and found to display the colour pattern of two longitudinal stripes (Morph 1). Although Langerhans named the species “rubra”, it is not evenly reddish brown, but as he mentioned in the description, has a colourless midline and two lighter lateral lines, producing the two typical longitudinal pigment stripes. The excellent original description allows us to recognise it as a junior synonym of Aponuphis bilineata.

Biology. Ectosymbiotic microorganisms were found on the surface of the anterior and median parapodia of A.

bilineata. These epibionts were identified as peritrich ciliates belonging to the genus Epistylis, characterised by forming colonies utilising its noncontractile and longer peduncle for attachment to the substrate.

Distribution. East Atlantic, Mediterranean.

Aponuphis brementi (Fauvel, 1916)

Figure 7C

Hyalinoecia brementi Fauvel, 1916: 5, figs. 2–3 (Monaco, Mediterranean). Aponuphis brementi.—Paxton 1986: 54 (new combination). Hyalinoecia fauveli Rioja, 1918: 45 (Santander, Spain); Fauvel 1923: 424, fig. 167a–h (Santander, Spain); 1938: 15 (Venice

Lagoon, Adriatic Sea); Bellan 1960: 18 (Portugal); 1969: 44 (Madeira); Amoureux 1972: 16 (Galicia, Spain); 1973: 440 (northern Spain, Bay of Biscay); 1974: (Aveiro, Portugal); Hartmann-Schröder 1977: 89 (Portugal).

Aponuphis fauveli.—Paxton 1986: 54 (new combination); Aguirrezabalaga et al. 2002: 27 (Bay of Biscay).

Material examined. Type material. Three syntypes of Hyalinoecia brementi (MOM 182438), Pointe de la Vieille, Monaco; holotype of Hyalinoecia fauveli Rioja, 1918 (MNCN 16.01.309), Santander, in the sandbank of Enmedio, northern Spain.

Non-type material. One specimen (MNCN 16.01/16167) COCACE station: D3 (43.85º N, 5.68º W), N. Spain, Cantabrian Sea, Bay of Biscay, 162 m, 89.06 % sand, 4.76 % silt, 6.18 % clay, 29 Jun 1987; 3 specimens (MNCN 16.01/16168) COCACE station: E2 (43.78º N, 6.08º W), N. Spain, Cantabrian Sea, Bay of Biscay, 190 m, 3 Jun 1987; 4 specimens (MNCN 16.01/16169) COCACE station: E2 (43.78º N, 6.08º W), N. Spain, Cantabrian Sea, Bay of Biscay, 190 m, 3 Jun 1987; 24 specimens (MNCN 16.01/16170) COCACE station: C5 (43.83º N, 5.81º W), N. Spain, Cantabrian Sea, Bay of Biscay, 150 m, 28 Apr 1987; 8 specimens (MNCN 16.01/16171) COCACE station: D2 (43.81º N, 5.75º W), N. Spain, Cantabrian Sea, Bay of Biscay, 161 m, 90.88 % sand, 3.46 % silt, 5.66 % clay, 29 Jun 1987; 2 specimens (MNCN 16.01/16172) COCACE station: C7 (43.81º N, 5.98º W), N. Spain, Cantabrian Sea, Bay of Biscay, 146 m, 98.58 % sand, 0.40 % silt, 1.02 % clay, 4 Jul 1987; 41 specimens (MNCN 16.01/1826, MNCN 16.01/1831; MNCN 16.01/1832, MNCN 16.01/1839, MNCN 16.01/1840, MNCN 16.01/1842, MNCN 16.01/2164, MNCN 16.01/2166, MNCN 16.01/2167, MNCN 16.01/2174, MNCN 16.01/2181, MNCN 16.01/2642, MNCN 16.01/2666, MNCN 16.01/2668) off shore between San Antonio cape and Valencia harbour, E. Spain, western Mediterranean, 3–30 m, 26 Apr 1996; 1 specimen (MNCN 16.01/5908) Salobreña, S. Spain, western Mediterranean, 20.5 m, 1 Jul 1999; 2 specimens (MNCN 16.01/6034) off shore between Sagunto Port and Castellón harbour, E. Spain, western Mediterranean, 3–30 m, 23 Sep 1999; 6 specimens (MNCN 16.01/


8743, MNCN 16.01/8798) off shore Punta Torrox, E. Spain , Alborán Sea, western Mediterranean, 3–30 m, 17 Apr 2002.

Type locality. Monaco, Mediterranean.Diagnosis. Antennae usually to chaetiger 15–20, maximally to 20; four to seven ceratophoral rings on median

antenna, six to nine usually on lateral antennae, maximally 11. Anterior four to six chaetigers with several tridentate pseudocompound hooks, chaetigers 5 to 8 with one hook only, rarely bidentate; slender long-appendaged hooks present. Ventral cirri subulate on first five to seven chaetigers. Subacicular hooks from chaetiger 12–15. Branchiae from chaetiger 1–2. Colour pattern (Fig. 7C) consisting of two large dorsal spots on peristomium and chaetigers 1–4, and two segmental lateral spots from chaetiger 1. From chaetiger 5 central spots becoming progressively smaller and thinner, by chaetiger 10 represented only by thin band. Maximum width without parapodia 1.6 mm.

Remarks. In very small specimens the branchiae start on chaetiger 6. From here they spread more posteriorly but also anteriorly. When the worm has attained about two-thirds of its maximum size the branchiae begin on chaetiger 1–2 which is considered here as the adult condition of the species. We have examined the syntypes of A.

brementi and found that in the larger two specimens (0.6–0.8 mm wide) the branchiae started on chaetiger 2 as stated in original description, while in the smallest one (0.4 mm wide) the origin was chaetiger 4 and 5. The holotype of Hyalinoecia fauveli Rioja, 1918 has been examined and found to fit within the range of Aponuphis

brementi and is here considered to be its junior synonym. Distribution. East Atlantic, Mediterranean.

Aponuphis ornata (Fauvel, 1928)

Figures 7D–H, 8, 9

Hyalinoecia bilineata ornata Fauvel, 1928: 12. Morocco; 1936: 52, fig. 3 (Morocco); 1938: 15 (Venice Lagoon, Adriatic Sea).Aponuphis ornata.—Paxton 1986: 54 (new combination).

Material examined. Type material. Two syntypes of Hyalinoecia bilineata ornata Fauvel, 1928 (MNHN-IA-TYPE0815 and MNHN-IA-TYPE0816) “Vanneaux” sta. CXIX and LVI, Morocco.

Non-type material. Three specimens (MNHN) “Vanneaux” sta. XLIII, Marocco; 14 specimens (MNCN 16.01/3881-MNCN 16.01/3883, MNCN 16.01/3895) off shore between Vidio Cape and Peñas Cape, Cantabrian Sea, Spain, 15–32 m, 5 Apr 1998; 88 specimens (MNCN 16.01/1828- MNCN 16.01/1830, MNCN 16.01/1833, MNCN 16.01/1835, MNCN 16.01/1837- MNCN 16.01/1838, MNCN 16.01/1841, MNCN 16.01/1843, MNCN 16.01/2158- MNCN 16.01/2163, MNCN 16.01/2165, MNCN 16.01/2168, MNCN 16.01/2170- MNCN 16.01/2173, MNCN 16.01/2175- MNCN 16.01/2180, MNCN 16.01/2672) off shore between San Antonio cape and Valencia harbour, E. Spain, western Mediterranean, 3–30 m, 26 Apr 1996; 2 specimens (MNCN 16.01/7820) off shore between Olympic Port of Barcelona and Mataró, NE. Spain, western Mediterranean, 5–25 m, 1 Jan 2000; 6 specimens (MNCN 16.01/8805) off shore Punta Torrox, E. Spain , Alborán Sea, western Mediterranean, 3–30 m, 17 Apr 2002.Comparative material. Two syntypes of Hyalinoecia grubii (NHMW 1169) Saint-Malo, France, collected in 1883.

Type locality. Coast of Morocco, East Atlantic.Diagnosis. Antennae usually to chaetiger 10, maximally to 16; four to six ceratophoral rings on median

antenna, usually seven to ten on lateral antennae, maximally 11. Anterior three to five chaetigers with several tridentate pseudocompound hooks (rarely tri- and quatridentate), chaetigers 4–6 with one hook only; slender long-appendaged hooks present. Ventral cirri subulate on first five chaetigers. Subacicular hooks from chaetiger 10. Branchiae from chaetiger 3 to 4. Colour pattern consisting of orange to light brown transverse bands and spots varying according to body region. Peristomium with two large dorsal spots, sometimes coalescing medially; first chaetigers with two wide bands separated medially (one close to anterior end of segment and other close to posterior edge), presenting four bars; following chaetigers with one solid band anteriorly and two bars posteriorly, bars and bands ranging from relatively narrow (Fig. 7D) to wide (Fig. 7E) and two segmental lateral spots from chaetiger 1 to posterior region; sometimes posterior bands displayed medially on segment, appearing as more or less quadrangular spots. Other specimens with more complicated pattern as consequence of fusion of anterior and posterior segmental bars (Fig. 7F). More posteriorly, anterior and posterior bars (or posterior quadrangular spots)


getting closer and merging into single wide transversal band per segment. Maximum width without parapodia 1.4 mm.

Remarks. Hyalinoecia bilineata ornata was described by Fauvel (1928) as differing from the stem species by its colour pattern. However, H. bilineata ornata differs also in parapodial and chaetal features from A. bilineata. We have examined the syntypes of Hyalinoecia bilineata ornata and found them to have only tridentate (not bi- and tridentate) hooks, less anterior parapodia with hooks and shorter antennae than A. bilineata. The colour pattern of the syntypes was not preserved but we examined specimens from a different station taken by the “Vanneaux” that clearly displayed the typical pigmentation of the species. Aponuphis ornata is very similar to Aponuphis grubii

(Marenzeller, 1886). The two species have basically the same colour pattern except that the bars and lines are substantially wider and appear overall more strongly pigmented in the former than in the latter. The antennae of A.

ornata are longer and its branchiae occur from chaetiger 3–4 instead of 4–5. Their reproductive methods are very different. As we are detailing below, A. ornata is a brooder with direct development, while A. grubii is a broadcast spawner with lecithotrophic larvae as has been reported by Rivain (1983) for the Rance population in northern France (as Hyalinoecia bilineata). On the basis of the data we have so far, we consider A. grubii as a northern European species. It is common in the Normano-Breton Gulf and may be restricted to the English Channel. In contrast, A. ornata is a southern species that occurs from the Bay of Biscay to Morocco, including the western and central Mediterranean.

FIGURE 8. Aponuphis ornata. Line drawings of juvenile hooks. A, tridentate compound hook; B, bidentate compound hook; C, median anterior provisional subacicular hook; D, ventral anterior provisional subacicular hook; E, posterior provisional subacicular hook.

Biology. Among the samples from the Cantabrian shelf, one specimen was collected in its tube together with eight vermiform juveniles, and one specimen from the Mediterranean with two juveniles. The juveniles range from 2.0–4.1 mm in length for 15–38 chaetigers and measure 0.3–0.6 mm in width, with the largest presenting a colour pattern on the dorsum of the anterior chaetigers (Figs 7H). All juveniles have well developed frontal lips, palps and antennae, the ceratophores of the largest two with three to four rings. The parapodia are well developed with dorsal cirri on the first seven to 10 chaetigers and subulate ventral cirri on the first two. Branchiae are absent; the pygidium bears two pairs of slender anal cirri. In addition, six recently settled juveniles were dredged from a depth between 3 to 30 m in Málaga (southern Spain). The specimens range from 4.2–7.1 mm in length for 44–52 chaetigers and measured 0.45–0.55 mm in width. These specimens have a well pigmented colour pattern (Fig. 7G) and the three larger ones have branchiae from chaetiger 7–9 to 15.


The chaetal composition of juveniles (Fig. 8) differs from that of adults and has been detailed for these A.

ornata specimens in Fig. 9. The smallest juveniles examined have four compound hooks in the first two pairs of parapodia. In the 15-chaetiger specimen they are only bidentate (Fig. 8B) and the 17- and 23-chaetiger specimens have bi- and tridentate hooks (Figs 8A, B). Parapodia 3–8 have two limbate chaetae and two kinds of anterior provisional subacicular hooks (SAHs): one with a large appendage in the medial position of the parapodium (Fig. 8C) and two ventral hooks with small appendages (Fig. 8D). Two permanent SAHs start on chaetiger 9, being present to chaetiger 12–14, when one of these, and a few chaetigers later both are replaced by posterior provisional subacicular hooks (Fig. 8E). Pectinate chaetae are absent.

The 28- and 38-chaetiger juveniles have bi- and tridentate compound hooks on the first three parapodia; parapodia 2 and 3 with four hooks and parapodium 3 with only one bidentate hook and two limbates. The anterior provisional hooks are now restricted to chaetiger 4 only, with chaetigers 5–8 lacking any hooks, having only limbate chaetae. Two permanent SAHs are present from chaetiger 9–20 or 22 when first one, and then both permanent SAHs are replaced by posterior provisional SAHs. A flat pectinate chaeta with 10–11 teeth is present from chaetiger 19 to 20–22.

The recently settled juveniles have four chaetigers with compound hooks, lack anterior provisional SAHs, have permanent SAHs from chaetiger 9–10 to 29, when they are replaced by posterior provisional SAHs.

The branchial development commences when juveniles consist of about 40 chaetigers. The branchiae start on chaetiger 7–9, from there they spread more posteriorly but also anteriorly, passing through intermediate forms with branchiae starting on chaetiger 6 and 5. When the worm has attained about two-thirds of its maximum size the branchiae begin on chaetiger 3–4 which is considered here as the adult condition of the species.

Distribution. East Atlantic (Bay of Biscay and Atlantic Iberia) and western and Central Mediterranean Sea. Previous records of A. grubii (or H. bilineata grubii) from the Iberian coasts and the Mediterranean most likely correspond to A. ornata.

FIGURE 9. Comparison of the progression of hooks in Aponuphis ornata juveniles.

Aponuphis willsiei Cantone & Bellan, 1996

Figure 10

Aponuphis willsiei Cantone & Bellan, 1996: 27.

Material examined. Non-type material. One specimen (MNCN 16.01/15244) Fauna IV off shore Alborán Island Lighthouse (35º 55.68’– 35º 55.63’ N, 03 º03.25’– 03º 02.67 W), E. Spain, Alborán Sea, western Mediterranean, 33–49 m, 23 Jul 1996; 4 specimens (AM W.47515) Mazarrón Port, Murcia, Spain, dredged from 50–100 m, coll.


20 Mar 2012; 5 specimens (MNCN 16.01/16173) Mazarrón Port, Murcia, Spain, dredged from 50–100 m, coll. 20 Mar 2012; 4 specimens coated with gold (MNCN 16.01/16174) Mazarrón Port, Murcia, Spain, dredged from 50–100 m, coll. 20 Mar 2012.

Type locality. Gulf of Marseille, France, Mediterranean.Diagnosis. Antennae up to chaetiger 10; three to six ceratophoral rings on antennae. Anterior four chaetigers

with four to five bidentate pseudocompound to compound hooks (Fig. 10), chaetiger 5 with two to three hooks; all hooks of almost equal thickness, slender long-appendaged hooks absent. Ventral cirri subulate on first four chaetigers. Subacicular hooks from chaetiger 9–10. Branchiae absent. No colour pattern. Maximum width without parapodia 1.0 mm.

FIGURE 10. Aponuphis willsiei. Scanning electron micrographs. A, bidentate compound hooks of chaetiger 3; detailed view of bidentate pseudocompound hook of chaetiger 2; C, pectinate chaeta from chaetiger 15.

Remarks. This is the first confirmed record of this species from Iberian Peninsula waters. Amoureux (1972) reported one incomplete 1 cm long specimen without branchiae among the A. bilineata (as Hyalinoecia bilineata) specimens collected on the continental slope of Galicia (northwestern Spain) and he considered it to be the abranchiate Mediterranean form reported by Bellan. However, we did not find any specimens belonging to this species among the samples we examined from the area. Since the species has not been reported again from the Galician coasts, we consider this record as questionable, since it may represent merely a juvenile phase of another Aponuphis species.

Biology. One specimen preserved in its tube was found with two juveniles closely appressed to its body. The juveniles were 2.3 and 2.5 mm long for 17 and 18 chaetigers and measured 0.3 mm in width. The juveniles have well developed frontal lips; palps and antennae with two ceratophoral rings; parapodia are well developed, anterior ones with dorsal cirri; subulate ventral cirri on chaetigers 1 and 2. The chaetal distribution is very similar to that described for A. ornata above with chaetigers 1 and 2 with bidentate compound hooks, limbate and anterior provisional SAHs from chaetiger 3–8, permanent SAHs from chaetiger 9–14, posterior provisional hooks in the last few chaetigers, and pectinate chaetae are absent.

Distribution. Mediterranean, this species is only known from the Gulf of Marseille, the southwestern Iberian Peninsula (Murcia, Spain) and the Alborán Sea.

Discussion

Several recent publications on Onuphis taxonomy have shown that even some features traditionally used to separate species within this genus, e.g. presence/absence of eyespots on the prostomium, the number of distinguishable postchaetal lobes or the number of chaetigers with subulate ventral cirri, were not found useful for


several Onuphis spp. (Arias & Paxton 2014; Gil & Machado 2014). Other traditional taxonomic characters are variable during ontogeny and thus size-related, like the maximal number of branchial filaments or the maximal number of rings of ceratophores of antennae and palps (Arias & Paxton 2014). A character that has been shown to become established relatively early in the ontogeny of Onuphis species (but not in some other genera) is the first chaetiger with subacicular hooks, which is almost invariable in adults and thus a good taxonomic character (Blake 1974; Arias & Paxton 2014; Gil & Machado 2014). Nevertheless, this character was reported as significantly variable in Onuphis spp. since many authors used to include juvenile specimens in species diagnoses and descriptions and therefore expanding the range of appearance of subacicular hooks. The morphology and typology of the pseudocompound hooks of anterior modified parapodia have long been considered as a good source of characters to differentiate species/species groups. Here we are also emphasising the shape and length of the hoods (e.g. triangular, falcate), the presence/absence of slender long-appendaged hooks, and the length of the appendage. Other features that showed to play a role were the chaetiger with the first occurrence of the pectinate chaetae and the density and disposition of the sensory buds of the antennae and palps, as has been already demonstrated in the genus Diopatra (Paxton 1986; 1993).

The genus Aponuphis is well represented in the shallow sandy sediments of the Mediterranean and northeastern Atlantic. Here we have only focused on the four species occurring in southwestern Europe. Identification of Aponuphis species has been hampered by their overall similarity, size-related distribution of branchiae and chaetae, and the loss of pigmentation in preserved specimens.

The branchial distribution, particularly the chaetiger of the first appearance of the single branchial filaments, although it has been widely used for species identification, is not a good tool due to its growth-related variation. In A. brementi adults the branchiae originate on chaetiger 1 or 2. However, in very small specimens the branchiae start on chaetiger 6 and from here they spread more posteriorly but also anteriorly. In a half grown specimen (width 0.5 mm) the branchiae start on chaetiger 4 to 5 (implying that one is dealing with A. bilineata or A. ornata) and it is only when A. brementi has attained about two-thirds of its maximum size (width 0.8 mm) that the branchiae begin on chaetiger 1–2 which is considered here as the adult condition of the species. Similarly, in A. ornata, branchiae initially start on chaetigers 7–9, progress posteriorly and anteriorly until they become stabilised at chaetiger 3–4 when the worm is about two-thirds grown.

The distribution of the pseudocompound to compound hooks of the anterior modified chaetigers and the subacicular hooks changes during ontogeny. The subject has previously been detailed in several studies and reviewed in general for onuphids by Orensanz (1990). In a study of three species of Hirsutonuphis Paxton, 1986 it was shown that the progression of hooks is species specific (Paxton 1996). We were fortunate to find an adult A.

ornata in a tube with its brooded juveniles and another sample of recently settled juveniles, allowing us to show the chaetal progression from the 15-chaetiger juveniles to adults (Fig. 9). The modified parapodia of Aponuphis

develop by acquiring pseudocompound/compound hooks consecutively until the typical adult number for a species is achieved which is four in A. ornata. Anterior provisional subacicular hooks (SAHs), representing relict structures, and posterior provisional hooks, representing precursors of permanent SAHs, occur in Aponuphis

juveniles. Anterior provisional SAHs consist of two types, a pair of ventral (Fig. 8D) and a single median (Fig. 8C), are present in juveniles only. The posterior provisional SAHs (Fig. 8E) are only of one type and are sometimes still present in the most posterior chaetigers of adult worms. The compound anterior provisional SAHs are also present in Onuphis, Kinbergonuphis and Hirsutonuphis, other genera of the Onuphis group. However, they have not been observed in the Diopatra group of genera, where the early SAHs are either of the adult, or a simple modified form (Paxton 1996).

One of the four southwestern European Aponuphis species, A. willsiei lacks any pigmentation and preserved specimens are cream coloured. We have described and illustrated the colour patterns and their variations for the remaining three species (Fig. 7), demonstrating that they can be utilised for species identification. Specimens that have lost all pigmentation should be carefully examined for the combination of diagnostic characters. However, identification may not be possible if they have not attained at least half their adult size.

Key to adult species of Onuphis and Aponuphis from southwestern Europe

1. Peristomial cirri absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aponuphis ... 2

- Peristomial cirri present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onuphis...6


2. Colour pattern and branchiae absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. willsiei

- Colour pattern and branchiae present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3. Branchiae from chaetiger 1–2, colour pattern of two large dorsal spots per segment, by about chaetiger 10 becoming thin band

(Fig. 7C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. brementi

- Branchiae from chaetiger 3–5, colour pattern otherwise . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Pseudocompound hooks tridentate; branchiae from chaetiger 3–4; colour pattern of horizontal bars and bands (Fig. 7D–F) . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. ornata

- Pseudocompound hooks bi- and tridentate; branchiae from chaetiger 4–5; colour pattern otherwise . . . . . . . . . . . . . . . . . . . . 5

5. Two dorsal longitudinal pigment stripes and two segmental spots from chaetiger 1 to posterior region (Fig. 7A) . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bilineata morph 1 (typical)

- Peristomium and chaetigers 3–4 almost solidly light brownish, from chaetiger 4–5 two lateral longitudinal stripes close to

parapodia (Fig. 7B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. bilineata morph 2

6. Branchiae from chaetiger 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

- Branchiae from chaetiger 5, all branchiae simple and strap-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. farensis

7. All branchiae simple and strap-like, colour pattern practically absent, first four pairs of parapodia with pseudocompound hooks

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

- Branchiae pectinate from about the 20th chaetiger, colour pattern consisting of distinctive bars and spots on dorsum, first three

pairs of parapodia with pseudocompound hooks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

8. Bi-, tri- and quatridentate pseudocompound hooks with falcate hoods (Fig. 3F–I), SAH from chaetiger 9, subulate ventral cirri

in first four chaetigers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. anadonae sp. nov.

- Only tridentate pseudocompound hooks with moderately long, triangular and pointed hoods (Fig. 6A–D), SAH from chaetiger

14, subulate ventral cirri in first five to six chaetigers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. rullieriana

9. Bi- and tridentate (or even multidentate) pseudocompound hooks, eyespots on prostomium absent, maximum of eight bran-

chial filaments, robust species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O. pancerii

- Only tridentate pseudocompound hooks, eyespots on prostomium present, maximum of five branchial filaments, slender spe-

cies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O. eremita

Acknowledgements

We are grateful to Javier Sánchez Almazán and Francisco Yagüe, Museo Nacional de Ciencias Naturales of Madrid, for the generous loan of most of the material on which this work is based, including the material from the Fauna Ibérica Project, to Helmut Sattmann and Stefan Szeiler, Naturhistorisches Museum Wien, for photographs of syntypes on slides, and to the curator and librarian of the Musée Océanographique de Monaco for help with locating specimens and literature. Our thanks go to Jorge Núñez and an anonymous reviewer for their careful reviews and helpful remarks.

We also thank Nuria Anadón for her constant support, Alfredo J. Quitana, Scientific-Technical Services of the University of Oviedo, for assistance with SEM microscopy and Sue Lindsay, Microscopy Unit of the Australian Museum, Sydney, for assistance with photography. The first author (AA) also wants to thank the Department of Biological Sciences, Macquarie University, Sydney, for hosting him for his overseas study period in 2013. This work was supported by the project RES-14-CI-058, University of Oviedo, and the FICYT foundation (Principado de Asturias).

References

Aguirrezabalaga, F., Ceberio, A. & Paxton, H. (2002) Onuphidae (Polychaeta) from the Capbreton Canyon (Bay of Biscay, NE Atlantic) with the description of Paradiopatra capbretonensis sp. nov. Steenstrupia, 27 (1), 19–28.

Amoureux, L. (1971) Problèmes biologiques suggérés par certaines répartitions d'Annélides Polychètes dans la zone meuble intertidale. Divers problèmes sont évoqués à propos de la répartition de Travisia forbesi et d'Ophelia bicornis. Vie et Milieu, Supplément 22 (2), 701–705.

Amoureux, L. (1972) Annélides polychètes recueillies sur les pentes du talus continental, au large de la Galice (Espagne). Campagnes 1967 et 1968 de la “Thalassa”. Cahiers de Biologie Marine, 13, 63–89.

Amoureux, L. (1973) Annélides polychètes recueillies sur les pentes du talus continental au Nord de la côte espagnole. Campagne 1970 de la "Thalassa". Cahiers de Biologie Marine, 19, 429–452.

Amoureux, L. (1974) Annélides polychètes recueillies sur les pentes du talus continental au nord-ouest de l’Espagne et du Portugal (Campagne 1972 de la “Thalassa”). Cuadernos de Ciencias Biológicas, 3, 121–54.

Amoureux, L. (1976) Annélides polychètes récoltes par J. Stirn en 1969, sur les côtes marocaines du Détroit de Gibraltar. Cuadernos de Ciencias Biológicas de la Universidad de Granada, 5, 5–33.


Amoureux, L. (1977) Annélides polychètes errantes recueillies sur les pentes du talus continental à l’entrée de la Manche, avec la description de deux espèces nouvelles. Campagne 1973 de la “Thalassa”. Cahiers de Biologie Marine, 18 (4), 391–411.

Arias, A., Anadón, N. & Paxton, H. (2010) New records of Diopatra marocensis (Annelida: Onuphidae) from northern Spain. Zootaxa, 2691, 67–68.

Arias, A, Paxton, H. & Anadón, N. (2013) Mooreonuphis vespa, a new brooding species of Onuphidae (Annelida) from northern Spain (Bay of Biscay). Zootaxa, 3741 (4), 583–592. http://dx.doi.org/10.11646/zootaxa.3741.4.8

Arias, A. & Paxton, H. (2014) Hidden diversity within the polychaete Onuphis eremita sensu lato (Annelida: Onuphidae)—redescription of O. eremita Audouin & Milne-Edwards, 1833 and reinstatement of Onuphis pancerii Claparède, 1868. Zootaxa, 3861 (2), 145–169.

Audouin, J.V. & Milne Edwards, H. (1833) Classification des Annélides et description de celles qui habitent les côtes de la France. Annales des sciences naturelles, 29, 195–269.

Baird, W. (1870) Remarks on several genera of annelids belonging to the group Eunicea with a notice of such species as are contained in the collection of the British Museum and a description of some others hitherto undescribed. Journal of the Linnean Society of London, 10, 341–361. http://dx.doi.org/10.1111/j.1096-3642.1869.tb00665.x

Bellan, G. (1960) Annélides polychètes. Résultats Scientifiques de la Campagne du N.R.P. “Faial” dans les eaux côtières du Portugal (1957). Lisboa, 2, 1–31.

Bellan, G. (1964) Contribution à l’étude systématique, bionomique et écologique des annélides polychètes de la Méditerranée. Thèse, Faculté des Sciences de l’Université d’Aix-Marseille, 371 pp.

Bellan, G. (1969) Annélides Polychètes recueillies dans l'Archipel de Madère au cours de la campagne scientifique du navire océanographique "Jean Charcot". Cahiers de Biologie Marine, 10 (1), 35–57.

Blake, J.A. (1974) The larval development of Polychaeta from the Northern California coast. II. Nothria elegans (Family Onuphidae). Ophelia, 13, 43–61. http://dx.doi.org/10.1080/00785326.1974.10430591

Borja, A., Franco, J. & Pérez, V. (2000) A Marine Biotic Index to establish the ecological quality of soft-bottom benthos within European estuarine and coastal environments. Marine Pollution Bulletin, 40 (12), 1100–1114. http://dx.doi.org/10.1016/S0025-326X(00)00061-8

Campoy, A. (1982) Fauna de España. Fauna de anélidos poliquetos de la Península Ibérica. EUNSA Publicaciones de Biología de la Universidad de Navarra, Serie Zoológica, 7, 1–781. [Pamplona]

Cantone, G. & Bellan, G. (1996) Aponuphis willsiei, una nuova specie di Onuphidae (Annelida, Polychaeta) delle coste marsigliesi. Animalia, 21 (1/3), 27–30.

Castelli, A. (1982) Onuphis falesia, a new species of Onuphinae (Polychaeta, Eunicidae). Bollettino di Zoologia, 49, 45–49. http://dx.doi.org/10.1080/11250008209439371

Çinar, M.E. (2009) Alien polychaete species (Annelida: Polychaeta) on the southern coast of Turkey (Levantine Sea, eastern Mediterranean), with 13 new records for the Mediterranean Sea. Journal of Natural History, 43, 2283–2328. http://dx.doi.org/10.1080/00222930903094654

Claparède, E. (1868) Les Annélides Chétopodes du Golfe de Naples. Mémoires de la Société de physique et d'histoire naturelle de Genève, 19, 313–584.

Day, J.H. (1967) A monograph on the Polychaeta of Southern Africa I. British Museum (Natural History) Publications, London, 458 pp.

Dexter, D. (1992) Soft bottom invertebrates of the Portuguese benthos. Boletim do Instituto Nacional de Investigação das Pescas (Lisboa), 17, 61–88.

Ehrenberg, C.G. (1830) Neue Beobachtungen über blutartige Erscheinungen in Aegypten, Arabien und Sibirien, nebst einer Uebersicht und Kritik der früher bekannten. Annual Review of Physical Chemistry, 94, 4, 477–514. http://dx.doi.org/10.1002/andp.18300940402

Fauchald, K. (1982) Revision of Onuphis, Nothria, and Paradiopatra (Polychaeta: Onuphidae) based upon type material. Smithsonian Contributions to Zoology, 356, 1–109. http://dx.doi.org/10.5479/si.00810282.356

Fauvel, P. (1916) Deux polychètes nouvelles (Disoma watsoni n. sp., et Hyalinoecia brementi n. sp.). Bulletin d'Institut Oceanographique (Monaco), 316, 1–10.

Fauvel, P. (1923) Faune de France: Polychètes errantes. Paul Lechevalier, Paris, 488 pp.Fauvel, P. (1928) Annélides Polychètes nouvelles du Maroc. Bulletin de la Société Zoologique de France, 53, 9–13.Fauvel, P. (1936) Contribution à la faune des Annélides Polychètes du Maroc. Mémoires de la Société des Sciences Naturelles

du Maroc, 43, 1–143.Fauvel, P. (1938) Annelida Polychaeta della Laguna di Venezia. R. Comitato Talassografico Italiano, 246, 1–27. Gil, J. & Machado, M. (2014) A new species of Onuphis (Polychaeta: Onuphidae) from Southern Portugal, with comments on

the validity of O. pancerii Claparède, 1868. Zootaxa, 3860 (4), 343–360. Glémarec M. (1969) Les peuplements benthiques du plateau continental Nord-Gascogne. Thèse d'Etat, Faculté des Sciences,

Paris, 167 pp.Guille, A. (1971) Bionomie benthique de plateau continental de la côte catalane française. VI. Données autoécologiques


(macrofaune). Vie et Milieu, 22, 469–527.Hartman, O. (1951) The littoral marine annelids of the Gulf of Mexico. Publications of the Institute of Marine Sciences, 2, 7–

124.Hartmann-Schröder, G. (1977) Polychaeten aus dem Sublitoral und Bathyal vor der portugiesischen und marokkanischen

Küste. Auswertung der Fahrt 8 (1967) von F.S. 'Meteor'. METEOR Forsch.-Ergebnisse, Series D, 26, 65–99.Hilbig, B. (1995) Family Onuphidae Kinberg, 1865. In: Blake, J.A., Hilbig, B. & Scott, P.H. (Eds.), Taxonomic Atlas of the

Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. 5 - The Annelida Part 2. Polychaeta: Phyllodocida (Syllidae and scale-bearing families), Amphinomida, and Eunicida. Santa Barbara Museum of Natural History, Santa Barbara, pp. 341–364.

Hobson, K.D. (1971) Some polychaetes of the superfamily Eunicea from the North Pacific and North Atlantic Oceans. Proceedings of the Biological Society of Washington, 83, 527–544.

Ibáñez, M. (1972) Notas sobre algunas especies de Anélidos Poliquetos nuevas para las costas de España, con especial referencia a Nereis (Neanthes) oxypoda Marenzeller y Onuphis (Nothria) geophiliformis Moore. Boletín de la Real Sociedad Española de Historia Natural (Sección Biológica), 70, 23–31.

Ibáñez, M. (1973) Catálogo de los anélidos poliquetos citados en las costas españolas. Cuadernos de Ciencias Biológicas, Universidad de Granada, 2, 121–140.

Imajima, M. (1960) Description of a new polychaete, Nothria shirikishinaiensis n. sp. of the family Eunicidae. Publications of the Seto Marine Biological Laboratory, 8 (1), 55–58.

Imajima, M. (1986) Eight species of Onuphidae (Polychaeta) in and Offshore of Otsuchi Bay, Northeastern Japan. Bulletin of the National Science Museum of Tokyo, 12, 93–116.

Kinberg, J.C.H. (1865) Annulata nova. Öfversigt af Königlich Vetenskapsakademiens förhandlingar (Stockholm), 21, 559–574.Kucheruk, N.V. (1978) Deep-water Onuphidae (Polychaeta) from the collections of the 16th cruise of the R/V Dmitry

Mendeleev (to the generic classification of the family Onuphidae). Trudy Institutyi Okeanologii Akademia Nauk SSSR, 113, 88–106.

Langerhans, P. (1880) Wurmfauna von Madeira. Zeitschrift für wissenschaftliche Zoologie, 33, 267–316.Laubier, L. & Paris, J. (1962) Faune marine des Pyrénées Orientales 4: Annélides Polychètes. Vie et Milieu, 13 (Supplement),

1–80.López, E. & San Martín, G. (1992) Familias de poliquetos errantes (Polychaeta), excepto Syllidae, recolectadas en las Islas de

Cabo Verde por la "I Expedición Ibérica". Revista de Biología Tropical, 40 (2), 161–169.Louzao, M., Anadón, N., Arrontes, .J, Álvarez-Claudio, C., Fuente, D.M., Ocharan, F., Anadón, A. & Acuña, J.L. (2010)

Historical macrobenthic community assemblages in the Avilés Canyon, N Iberian Shelf: baseline biodiversity information for a marine protected area. Journal of Marine Systems, 80, 47–56. http://dx.doi.org/10.1016/j.jmarsys.2009.09.006

Maekawa, N. & Hayashi, I. (1989) Onuphid polychaetes from Wakasa Bay, Sea of Japan. Memoirs of the College of Agriculture, Kyoto University, 134, 61–93.

Maekawa, N. & Hayashi, I. (1999) Taxonomic study on the genus Onuphis (Polychaeta, Onuphidae) from Japan and adjacent seas, with descriptions of six new species. Bulletin of the National Science Museum, Series A (Zoology), 25 (3), 163–214.

Malmgren, A.J. (1866) Annulata Polychaeta Spetsbergiae, Groenlandiae, Islandiae et Scandinaviae hactenus cognita. Ex Officina Frenckelliana, Helsingfors, 127 pp.

Marenzeller, E. von (1879) Südjapanische Anneliden I. Dankschrift der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, 41 (2), 109–152. [Wien]

Martínez, J. & Adarraga, I. (2006) Programa de vigilancia y control de la introducción de especies invasoras en los ecosistemas litorales de la costa vasca. 2. Costa de Bizkaia. Departamento de Medio Ambiente y Ordenación del Territorio del Gobierno Vasco /Sociedad Cultural de Investigación Submarina, Donostia-San Sebastián, 267 pp.

Martínez, J., Adarraga, I. & Ruiz, J.M. (2010) Tipificación de poblaciones bentónicas de los fondos blandos de la plataforma continental de Guipúzcoa (sureste del golfo de Vizcaya). Boletín del Instituto Español de Oceanografía, 23 (1–4), 85–110.

Moore, J.P. (1903) Polychaeta from the coastal slope of Japan and from Kamchatka and Bering Sea. Proceedings of the Academy of Natural Sciences of Philadelphia, 55, 401–490.

Mucha, A.P. & Costa M.H. (1999) Macrozoobenthic community structure in two Portuguese estuaries: relationship with organic enrichment and nutrient gradients. Acta Oecologica, 20 (4), 363–376. http://dx.doi.org/10.1016/S1146-609X(99)00130-7

Orensanz, J.M. (1990) The eunicemorph polychaete annelids from Antarctic and Subantarctic Seas. With addenda to the Eunicemorpha of Argentina, Chile, New Zealand, Australia, and the Southern Indian Ocean. Biology of the Antarctic Seas. XXI. Antarctic Research Series, 52, 1–183. http://dx.doi.org/10.1029/AR052p0001

Paxton, H. (1986) Generic revision and relationships of the family Onuphidae (Annelida: Polychaeta). Records of the Australian Museum, 38, 1–74. http://dx.doi.org/10.3853/j.0067-1975.38.1986.175

Paxton, H. (1993) Diopatra Audouin and Milne Edwards (Polychaeta: Onuphidae) from Australia, with a discussion of developmental patterns in the genus. The Beagle, Records of the Northern Territory Museum of Arts and Sciences, 10, 115–154.


Paxton, H., Fadlaoui, S. & Lechapt, J.P. (1995) Diopatra marocensis, a new brooding species of Onuphidae (Annelida: Polychaeta). Journal of Marine Biological Association of the United Kingdom, 75, 949–955. http://dx.doi.org/10.1017/S0025315400038273

Paxton, H. (1996) Hirsutonuphis (Polychaeta: Onuphidae) from Australia, with a discussion of setal progression in juveniles. Invertebrate Taxonomy, 10, 77–96. http://dx.doi.org/10.1071/IT9960077

Paxton, H. (1998) The Diopatra chiliensis confusion - redescription of D. chiliensis (Polychaeta, Onuphidae) and implicated species. Zoologica Scripta, 27, 31–48. http://dx.doi.org/10.1111/j.1463-6409.1998.tb00427.x

Redondo, M.S. & San-Martín, G. (1997) Anélidos Poliquetos de la costa comprendida entre el cabo de San Antonio y el Puerto de Valencia. Publicaciones Especiales del Instituto Español de Oceanografía, 23, 225–233.

Rioja, E. (1918) Datos para el conocimiento de la fauna de anélidos poliquetos del Cantábrico (2ª Parte). Trabajos del Museo Nacional de Ciencias Naturales, Madrid, Serie Zoológica, 37, 1–99.

Rivain, V. (1983) Contribution à l’étude dynamique et fonctionnelle des peuplements de sables fins du Golfe normano-breton. Thèse 3ème cycle, Océanographie Biologique, Université de Paris VI, Paris, 166 pp.


Onuphis and Aponuphis (Annelida: Onuphidae) from southwestern Europe, with the description of a new species

Documents