Guide to identification of Lumbrineridae - NMBAQC

Guide to identification of Lumbrineridae (Polychaeta) in north 

east Atlantic waters 

Eivind Oug 

Norwegian Institute for Water Research (NIVA), 

Regional Office Sorlandet, 

Jon Lilletuns vei 3, 

NO-4879 Grimstad, Norway 

eivind.oug@niva.no 

________________________________________________________________________________ 

Version 3.2, February 2012 

Prepared for NMBAQC workshop 2010 

Dove Marine Laboratory, Cullercoats, Tynemouth, UK

2 

Table of contents 

Introduction 3 

Morphology 3 

Identification 4 

Geographical area 5 

Status of the taxonomy 5 

Explanation of terms 6 

Key to genera in north east Atlantic waters 7 

Abyssoninoe Orensanz, 1990 10 

Augeneria Monro, 1930 12 

Cenogenus Chamberlin, 1919 14 

Eranno Kinberg, 1865 16 

Helmutneris Carrera-Parra, 2006 17 

Hilbigneris Carrera-Parra, 2006 18 

Lumbricalus Frame, 1992 19 

Lumbrinerides Orensanz, 1973 20 

Lumbrineriopsis Orensanz, 1973 21 

Lumbrineris Blainville, 1828 23 

Ninoe Kinberg, 1865 26 

Scoletoma Blainville, 1828 27 

Species of uncertain status 29 

References 30 

Acknowledgements 

The present guide has been prepared for the NMBAQC taxonomic workshop 2010. It is based on a 

previous version prepared for NMBAQC 2003. I am indebted to David Hall and Tim Worsfold at 

Thomson Unicomarine Ltd for organising the workshop and for practical help. The guide and previous 

versions have been developed during studies of polychaetes collected in environmental monitoring in 

Norwegian coastal and offshore waters, and taxonomic studies in North East Atlantic and Arctic areas 

(projects BIOFAR, BIOICE, COBICE, Swedish Kattegat surveys, NDP). I am grateful to several users 

of previous guide versions for comments and suggestions to improvements. The work has received 

financial support from NMBAQC and from the Norwegian Academy of Science and Letters (DNVA) 

through the Norwegian Deepwater Programme (NDP).

Introduction 

Lumbrineridae from NE Atlantic waters 

The Lumbrineridae are generally long cylindrical bristle worms of small to medium size with rather 

simple external morphology. The largest species in European waters may reach a length of about 35 

cm, but most species are 2-10 cm in length. Lumbrinerids are mostly found in soft-sediment 

environments where they burrow into the substrate. They are largely considered to be carnivores, but 

some species have been reported to feed on plant fragments or detritus (Fauchald & Jumars 1979). 

The lumbrinerids belong in the ‘eunicid’ group of polychaetes which is characterised by the 

presence of a complicated jaw apparatus in the phaynx. The jaw apparatus is of the ‘labidognath’ type 

(Orensanz 1990) and is basically similar to the jaws found in Eunicidae and Onuphidae. In traditional 

faunal works the eunicid group has been treated as an order (e.g. Fauchald 1977, Hartmann-Schröder 

1996), whereas in newer taxonomy the group has been placed without particular rank within the main 

division Aciculata (Rouse & Pleijel 2001). 

The most important taxonomic characters are found in jaw structures, types and shapes of 

chaetae, colour of aciculae and shape of parapodial lobes. Jaw characters have been extensively used 

for definition of genera in recent years, which has lead to a more stable and consistent system for 

species classification in the family. As a consequence, several new genera have been erected whereas 

extant genera have been redefined (see Orensanz 1973, 1990; Frame 1992; Carrera-Parra 2006a). 

Orensanz (1990), Frame (1992) and Hilbig (1995) have accounted for the taxonomic history of the 

family and have discussed the importance of the diagnostic characters. 

A remarkably large number of species of lumbrinerids have been described worldwide (i.e., 

more than 200). Many of the descriptions, however, especially in the older literature, are rather general 

and do not mention characters which presently are known to be vital for species discrimination. This 

has led to much confusion about species identities and synonymies. Whereas several species problems 

have been clarified over the years, many species are still insufficiently characterised and of uncertain 

status. 

Morphology 

Lumbrinerids have a long cylindrical body with generally similar segments (Figure 1). In most species 

the width gradually decreases towards the pygidium. The prostomium is well-developed and is usually 

without eyes or appendages, but in some genera small occipital antennae may be found in the fold 

between prostomium and peristomium. The peristomium consists of two rings (not true segments) 

without parapodia or chaetae. Tentacular cirri are absent. The following chaetigers carry parapodia 

which are mostly uniramous, but often a bundle of thin notoaciculae and a short knoblike notopodium 

are present. Ventral cirri are absent. Chaetae include simple limbate chaetae, composite spinigers, and 

simple and composite hooded hooks (Figure 1). 

Figure 1. Features of Lumbrineridae: anterior body (Lumbrineris), parapod from anterior body (Lumbrineris), 

chaetae (limbate chaeta, composite spiniger (Lumbricalus), long-bladed simple hook, multidentate composite 

hook, bidentate hook (top, Lumbrinerides), simple multidentate hook) 

3

4 

The jaw apparatus is comprised of dorsal maxillae and ventral mandibles (Figure 2). The maxillae 

consist of a pair of posterior carriers and four or five pairs of maxillary plates. The carriers are broad, 

mostly short, and are attached to the most posterior pair of maxillae (mx I) by a firm ’click-joint’ 

connection (labidognath arrangement). The maxillary plates are usually referred to by numbers (roman 

numerals) from I to V (note that in older literature different numbering systems were often used). In 

addition to the numbered plates other more or less well-developed structures may be found, variously 

referred to as ligaments, bridles or attachment lamellae. The mandibles are generally of simple 

structure and formed as rods with flared anterior ends (Figure 2). Morphological variation of maxillae 

among genera found in North Atlantic waters is shown in Figure 3. 

Figure 2. Jaw apparatus of Lumbrineridae. Left: maxillae (from Lumbrineris). Right: mandibles (from 

Lumbrineris). Roman numerals indicate established numbering system for maxillary elements. 

Figure 3. Maxillae of Lumbrineridae. Left to right: Lumbrineris, Abyssoninoe, Augeneria, Eranno, 

Lumbrinerides, Lumbrineriopsis 

Identification 

Maxillary structures can most conveniently be examined by making a dorsal incision in the region of 

chaetiger 1-2 to 5-6 (Figure 4). The most important structures, mx III and mx IV, are usually oriented 

more or less vertically with teeth projecting upwards just in front of the tip of mx I (Figure 4 right). In 

cases when the pharynx is partly protruded and the maxillae appear in the mouth opening, a ventral 

incision at the side of the midline may be made instead of a dorsal incision. However, it is then usually 

necessary to break off the anterior part of the mandibles to examine mx III and IV. 

Important characters of parapodia and chaetae are mostly found in anterior part of the body, usually 

from chaetiger 1 to chaetiger 20-30. Chaetae should be examined using a compound microscope 

(400x). In small specimens, chaetae can be viewed directly from specimens placed with the ventral 

side uppermost under a cover-slip. In larger specimens it is usually necessary to remove a parapodium 

for examination. This also applies to examining the colour of the aciculae. In some cases parapodial 

lobes on posterior chaetigers should also be examined.


Figure 4. Left: Dorsal incision to expose maxillae, Lumbrineris near coccinea, western Greenland. Right: 

Lumbrineris latreilli, east of Iceland. Numbering of maxillae indicated. Photo: Eivind Oug 

Geographical area 

The area covered by this guide encompasses European waters from the Bay of Biscay to western 

Norway and the Norwegian Sea. This covers coastal inshore waters, continental shelves and deep sea 

areas in the north-east North Atlantic and the Norwegian Sea. In addition, some species from nearby 

areas (Barents Sea and Arctic waters, north-west North Atlantic) which may be found in the area have 

been included. 

About 30 lumbrinerid species are known from the area. The highest number of species is reported 

from the English Channel, the North Sea and shelf areas of the Faroes and Iceland. 

General faunistic works treating lumbrinerids in North East Atlantic waters include Fauvel (1923), 

Hartmann-Schröder (1971, 1996), Miura (1980), Winsnes (1980), George & Hartmann-Scröder (1985) 

and Kirkegaard (1992). Several of the species found in the area are not included in the ‘standard’ 

works used for species identification. 

Data for the distribution of the species have been assembled from i.a. Fauvel (1923), Hartman (1965), 

Hartman & Fauchald (1971), Miura (1980), Winsnes (1980), Frame (1992), Mackie & Erséus (1997), 

Hansson (1998), Brattegard & Holthe (2001), Dauvin et al. (2003), Oug (2005), Carrera-Parra (2006a, 

b), and Aguirrezabalaga & Carrera-Parra (2006). In addition, my own observations on material from 

the North Sea and Norwegian coastal waters have also been taken into account. 

Status of the taxonomy 

Several of the species in the area are insufficiently characterised. In some cases, current descriptions in 

‘standard’ identification literature may seem to mix characters from several similar species making 

species identification problematical. In addition, specimens with specific characters that do not fit with 

the current species descriptions are regularly found. Whether such specimens represent intraspecific 

variants or separate species is not known. The genera with most problems are Abyssoninoe, Augeneria 

and Lumbrineris. The present guide mostly comprises named species with good diagnostic characters, 

but some regularly found forms of uncertain status also are included (referred to as ‘cf ‘or ‘near’ 

related species). A few currently un-named forms are also included. 

V 

I 

IV 

III 

I 

I 

II 

I 

I 

I 

5

6 

Explanation of terms 

Acicula – supporting bristle embedded in the parapodium 

Attachment lamellae – additional pieces of the maxillary apparatus, usually situated lateral to or below 

the individual maxillae. Rod-shaped attachment lamellae situated lateral to mx I are by some authors 

referred to as “bridles”. 

Bidentate hook – hook with two strong distinctly separated teeth 

Branchial lobe – cylindrical to short digitiform process extending from the parapodium and equipped 

with capillary blood vessels for gas exchange. Present in some genera and species. 

Carrier – part of the maxillary apparatus situated posterior to the first maxillae (mx I) and connected 

to it by a ‘click-joint’ 

Chaetiger – segment bearing chaetae 

Composite hooded hook – hook with a distinct subdistal joint separating the outer part (blade) from the 

shaft. Hood with a more or less distinct constriction at the position of the joint. 

Composite spiniger – simple gradually tapering chaeta provided with a joint 

Connecting ligament – additional piece placed between and connecting the bases of mx I and mx II. 

Also referred to as connecting lamella. 

Limbate chaeta – simple gradually tapering chaeta provided with a brim along most of the length, 

often gently curved. Found in anterior body of all species. 

Maxillae – paired and usually the most prominent elements of the maxillary apparatus. Maxillae are 

generally numbered from I to V from behind and forwards (different numbering systems may be found 

in older literature). 

Maxillary apparatus – dorsal part of the jaws consisting of several individual elements: four or five 

pairs of maxillae, carrier, connecting ligaments and attachment lamellae 

Maxillary teeth – well-defined projections on the maxillae directed inwards or upwards. In some 

genera knoblike projections or swellings may be found in addition to defined teeth, by some authors 

also referred to as teeth. 

Multidentate hooded hook – hook with a rounded apex with several small teeth encapsulated by a 

transparent hood. Only type of chaeta in posterior body of most species. 

Occipital antenna – small antenna placed posteriorly on the prostomium; in lumbrinerids in the fold 

between prostomium and peristomium. May also be referred to as nuchal antenna or nuchal papilla. 

Pseudocomposite hooded hook – hook with an incomplete separation between the outer part (blade) 

and shaft. Usually with a constriction of the hood as in composite hooks.

Key to genera in north east Atlantic waters 


1. Composite or pseudocomposite hooded hooks present in anterior part of body …………….. 2 

- All hooded hooks simple, anterior body may have limbate chaetae only ……………………. 5 

2. Chaetae include composite spinigers in addition to composite and simple hooded hooks …. 

……………….…………………………………………………………………… Lumbricalus 

- No composite spinigers …………….…………………………………………........................ 3 

3. Mx IV forming a squarish plate with whitish central area and black margins. Mx V absent. 

Occipital antennae may be present at posterior border of prostomium …….............. Augeneria 

- Mx IV a triangular to oblong evenly black or brown plate with a marked tooth. Mx V present, 

rounded ………………………………………………………………………………………. 4 

4. Mx II almost as long as mx I (about 2/3 or 3/4 of the length), connected to mx I with narrow to 

moderately wide connecting ligament. Aciculae pale, brown or black .................. Lumbrineris 

7

8 

- Mx II about half as long as mx I, connected to mx I by wide connecting ligament. Aciculae 

yellow …………………………………………………………………………..…. Hilbigneris 

5. Anterior chaetigers with one or more postchaetal branchial lobes …………………………… 6 

Ninoe Cenogenus 

- No branchial lobes in anterior chaetigers ………….…............................................................. 7 

6. Branchiae with multiple lobes. Number of lobes increasing from one at chaetiger 5-6 to four to 

six at chaetiger 20-30, then rapidly decreasing ................................................................. Ninoe 

- One short digitiform branchia present at each chaetiger in anterior body, gradually 

diminishing posteriorly. Anterior parapodia usually with long slender hooded hooks. Aciculae 

black. ......................................................................................................................... Cenogenus 

7. Anterior of body with limbate chaetae and modified long hooded hooks. Most anterior hooks 

extremely slender, resembling limbate chaetae in outline, gradually becoming shorter through 

following chaetigers and developing into regular hooded hooks at chaetiger 15-25. Aciculae 

yellow. Mx IV and V fused forming a hemisphaerical structure, with a strong dorsal tooth at 

posterior border……................................................................................................ Abyssoninoe 

chaet 5-10 chaet 15-25 

- Anterior of body with limbate chaetae and regular hooded hooks or with limbate chaetae only 

………………………………………………………………………………………………… 8


8. Hooded hooks bidentate ............................................................................................................ 9 

- Hooded hooks multidentate ..................................................................................................... 10 

9. Hooded hooks with a subdistal tooth or spur. Mx IV with a fringe of denticles on inner 

margin. Carriers long and slim, with narrow anterior part ……………........... Lumbrineriopsis 

- Hooded hooks without subdistal tooth. Mx IV without teeth or denticles. Carriers wide, with 

arcuate anterior border. ........................................................................................Lumbrinerides 

10. Maxillary apparatus with five pairs of maxillae …………………………………………….. 11 

- Maxillary apparatus with four pairs of maxillae, mx III and mx IV with whitish central area 

…………………………………………………………………………………...... Helmutneris 

11. Mx II short, about half the length of mx I, with distinct rather narrow connecting ligament 

from the base of mx II to posterior part of mx I …..….................................................. Eranno 

- Mx II almost as long as mx I, with short connecting ligament from mx II to posterior part of 

mx I ………………………………………………………………………….……… Scoletoma 

9

10 

Abyssoninoe Orensanz, 1990 

Characteristics. Prostomium conical. Parapodia uniramous, with simple limbate chaetae and simple, 

multidentate hooded hooks. Anterior hooks extremely long, tapering, resembling limbate chaetae, 

becoming gradually shorter from chaetiger 10-15, for 5 to 15 chaetigers, after which they appear as 

clearly defined hooks. Aciculae yellow. Mx III unidentate. Mx IV and V fused, mx IV with a tooth 

protruding from posterior border. 

Taxonomy. The genus is rather well-defined by the modified hooded hooks in anterior chaetigers and 

the fusion of mx IV and V. Species discrimination is problematical, however. Characters which have 

been used for species separation include the most anterior position of ’normal-shaped’ hooded hooks 

and the development of prolonged digitiform vascularised lobes in far posterior segments. Abyssoninoe 

hibernica was redescribed by Parapar et al. (1994). Abyssoninoe scopa was synonymised with 

A. hibernica by Parapar et al. (1994), but is here considered to be a separate species. The number of 

species in the region is uncertain. 

1. Anterior body with prolonged tapering hooks at least to chaetiger 15, posterior parapodia with 

prolonged digitiform lobes …………………………………………………………………. 2 

- Anterior body with prolonged tapering hooks to chaetiger 10-15, posterior parapodia with 

short rounded prechaetal and short conical postchaetal lobes ……………………………… 

……………….………………………………. Abyssoninoe cf abyssorum (McIntosh, 1885) 

Length 30-50 mm, width 1-1.5 mm 

Atlantic deep water, Norwegian Sea, 

?Norwegian fjords, 500-1100 m 

Figs from left: Anterior end; limbate 

chaeta; modified hooded hooks 

chaetiger 3,7,8,10; posterior hooded 

hook; carrier and right mx I-IV. From 

Orensanz (1990) 

2. Clearly defined hooded hooks appear at chaetiger 15-20, far posterior parapodia with 

prolonged digitiform prechaetal and postchaetal lobes…………………………………….. 

…………………………………………………….. Abyssoninoe hibernica (McIntosh, 1903) 

[=A. scopa aequilobata (Winsnes, 1981)] 

Length 40-60 mm, width 1-1.4 mm 

North Sea, British Isles, France, Norwegian 

coast, moderate depths 

Figs from left: Anterior end; maxillae 

(left mx III and right mx I, II omitted); 

limbate chaeta; slender hooded hook 

chaetiger 8; detail of same; hooded 

hook chaetiger 25; posterior hooded 

hook; parapodium 15; far posterior 

parapodium. From Winsnes (1980, 

1981) and Parapar et al. (1994)


- Clearly defined hooded hooks appear at about chaetiger 25 or further back, far posterior 

parapodia with digitiform postchaetal lobe, prechaetal lobe small ……………………….…… 

.......................................................................................... Abyssoninoe scopa (Fauchald, 1974) 

[ = A. pseudofragilis (Amoureux, 1977)] 

Length 30-60 mm, width 1-2 mm 

Fjords in western and northern 

Norway, NE Atlantic deep water, 

Bay of Biscay, 200-1200 m. 

Figs from left: Anterior end; 

maxillae; parapodium 2; parapodium 

15; far posterior parapodium. From 

Fauchald (1974) and Winsnes (1980) 

11

12 

Augeneria Monro, 1930 

Characteristic. Prostomium conical or broadly rounded, with or without small occipital antennae 

dorsally at border to peristomium. Parapodia with simple and composite multidentate hooded hooks. 

Mx IV squarish, with whitish central and dark peripheral areas. Mx V absent. 

Taxonomy. The genus is characterised by the shape of mx IV. The number of species in the area is 

uncertain. Only two species, A. algida and A. riojai, are well diagnosed. Most specimens from shelf 

and inshore waters agree reasonably well with Augeneria tentaculata from southern Atlantic and 

Antarctic waters (Monro 1930). Augeneria algida was redescribed by Winsnes (1987). 

1. Prostomium with eight small occipital antennae. Anterior parapodia with small prechaetal and 

rounded postchaetal lobes, parapodia from about chaetiger 15 with distinctly outdrawn 

prechaetal and postchaetal lobes. Aciculae dark ………………………..……………………… 

…………………………………….. Augeneria riojai Aguirrezabalaga & Carrera-Parra, 2006 

Length > 100 (?) mm, width 4.5 mm 

Bay of Biscay, 400-600 m 

Figs from left: Parapodium 26; 

composite hooded hook; simple 

hooded hook; maxillae. From 

Aguirrezabalaga & Carrera-Parra 

(2006). 

- Prostomium with three occipital antennae or without antennae. Parapodia with small 

prechaetal and conical postchaetal lobes. Aciculae light yellow……………………………2 

2. Anterior parapodia with long pseudocomposite to composite more or less twisted hooded 

hooks. Simple hooks present from about chaetiger 20 .…….. Augeneria algida (Wirén, 1901) 

Length > 60 mm, width 2.5-3.5 mm 

Norwegian Sea, Arctic, in deep 

water with temperatures < 0 °C; 

?Bay of Biscay 


pseudocomposite hooks; 

parapodium 15; maxillae. From 

Winsnes (1987). 

- Anterior parapodia with straight distinctly composite hooded hooks ……………………… 3


3. Composite hooded hooks with short blades (length: width = about 5:1). Simple hooded hooks 

present from about chaetiger 15 ……………….…… Augeneria cf tentaculata Monro, 1930 

Length 30-60 mm, width 1.5-2 mm 

N Atlantic, North Sea, Norwegian 

coast 


composite hooded hook chaetiger 5; 

posterior hooded hook; maxillae. 

From Orensanz (1990). 

- Composite hooded hooks with long blades (length: width = about 8:1). Simple hooded hooks 

from chaetiger 15-20 ………………………………………….……………… Augeneria sp. 

Length > 50 mm, width 1.5-2 mm 

Norwegian Sea, in deep water 

(1000-2000 m) with temperatures 

< 0 °C. 

Figure: Composite hooded hooks 

from anterior chaetiger (original) 

Remarks: This form may represent the same species as Augeneria sadko 

(Annenkova, 1952) from deep water in the Greenland Sea. Winsnes (1987) 

indicated that A. sadko could be synonymous with A. algida. They are here 

considered to be separate species. The taxonomy of Augeneria in northern and 

Arctic waters is unclear and needs to be further studied. 

13

14 

Cenogenus Chamberlin, 1919 (= Paraninoe Levenstein, 1977) 

Characteristics. Prostomium conical, with a short occipital antenna. Chaetae include simple limbate 

chaetae and simple multidentate hooded hooks. Anterior parapodia with a knoblike to digitiform 

postchaetal branchial lobe, extending back over a variable number of median segments. Aciculae 

black. Mx III and IV unidentate. Mx V lacking. 

Taxonomy. The genus is characterised by the presence of a postchaetal branchial lobe in anterior and 

mid body segments, black aciculae and the lack of mx V. Most species are found in deep water. The 

species in North Atlantic waters were treated by Miura (1980). The genus Cenogenus was reinstated 

and emended by Carrera-Parra (2001). Paraninoe Levenstein is regarded as a junior synonym of 

Cenogenus. 

1. Anterior parapodia with simple limbate chaetae and long-bladed slender hooks ………….. 2 

- Anterior parapodia with simple limbate chaetae only, mx II with 2 teeth …………………… 

…………........................... ……………………………….… Cenogenus fusca (Moore, 1911) 

Length > 60 mm, width up to 4 mm 

N Atlantic, north of Ireland, deep 

water to abyssal 

Figs from left: Anterior end ventral; 

middle parapodium; maxillae. From 

Miura (1980). 

2. Parapodia with a short digitiform branchial lobe, mx II with 3 teeth ………………..……….. 

…………….................................................................... Cenogenus brevipes (McIntosh, 1903) 


N Atlantic, north of Ireland, deep 

water to abyssal 

Figs from left: Anterior end ventral; 

parapodium 5 (top) and parapodium 

40 (bottom); maxillae. From Miura 

(1980) and Frame (1992).


- Parapodia with a small rudimentary branchial lobe, mx II with 4 teeth ………………..……… 

…………….……………………........Cenogenus sp. (= Paraninoe minuta sensu Miura 1980) 

Length 20 (?) mm, width 1.2 mm 

N Atlantic, north of Ireland, Bay of 

Biscay, deep water to abyssal 

Figs from left: Anterior body; 

prostomium ventral; anterior hooded 

hook; posterior hooded hook; 

maxillae. From Miura (1980). 

Remarks: This species may possibly belong in Abyssoninoe, but it is stated to 

have black aciculae and postchaetal branchial lobes, which are characters of 

Cenogenus 

15

16 

Eranno Kinberg, 1865 

Characteristics. Prostomium conical, occipital antenna may be present. Parapodia uniramous, with 

simple limbate chaetae and simple multidentate hooded hooks. Mx II short, about half of length of mx 

I, connected to base of mx I by narrow connecting ligament. Mx V partially fused to mx IV or free. 

Taxonomy. The genus is characterised by the short mx II being connected to the base of mx I with a 

long narrow connecting ligament. Eranno was treated by Orensanz (1990) and Frame (1992). Only 

one species is known from the area. The species was redescribed by Hartman (1948). 

Anterior parapodia with simple limbate chaetae and long-bladed hooded hooks. Aciculae 

yellow. Mx II half the length of mx I, mx III unidentate …….. Eranno bifrons Kinberg, 1865 

[=E. ehlersii tenuisetis (McIntosh, 1885); fide 

Miura 1980] 


N Atlantic, deep water. 

Figs from left: parapodium 19; longbladed 

hooded hook from parapodium 

19; hooded hook from posterior 

chaetiger; right mx III-V; maxillae. 

Original drawing. 

Remarks: This species is very close to E. petersenae Frame, 1992 from NW 

Atlantic. E. petersenae is characterised by having elongated digitate 

parapodial lobes in posterior body.

Helmutneris Carrera-Parra, 2006 


Characteristics. Parapodia uniramous, with simple limbate chaetae and simple multidentate hooded 

hooks. Mx III and IV with whitish central area, unidentate, mx V absent. 

Taxonomy. The genus is characterised by the pigmentation of mx III and mx IV and the lack of mx V. 

The genus has been recently established to incorporate species previously included in Lumbrineris. 

Only one species is known from the area. 

Anterior parapodia with limbate chaetae (in chaetiger 1 – ca. 30) and simple hooded hooks. 

Most anterior hooks with up to 8 teeth of similar size, median and posterior hooks with large 

proximal tooth. Prechaetal lobes insignificant, postchaetal lobes well developed, digitiform to 

tongue-shaped. Aciculae yellow …………………...…. Helmutneris flabellicola (Fage, 1936) 


Associated with deep water corals. 

West coast of Ireland, Bay of Biscay 

Figs from left: maxillae; hooded hook 

from chaetiger 4; hooded hook from 

chaetiger 30; parapodium 4; 

parapodium 20. From Carrera-Parra 

(2006a). 

17

18 

Hilbigneris Carrera-Parra, 2006 

Characteristics. Parapodia uniramous, with simple limbate chaetae and composite and simple 

multidentate hooded hooks. Mx II distinctly shorter than mx I, connected to mx I by wide connecting 

ligament. Mx V small, placed outside of mx III and IV. 

Taxonomy. The genus has been recently established to incorporate species previously included in 

Lumbrineris. Only one species is known from the area. The species is recently described (Carrera- 

Parra 2006a) from specimens previously identified as L. latreilli. 

Anterior parapodia with tongue-shaped, obliquely conical postchaetael lobes. Composite 

multidentate hooded hooks in chaetiger 1- ca. 25, with long blades. Aciculae yellow. Mx III 

bidentate ………………...………………………….. Hilbigneris pleijeli Carrera-Parra, 2006 

Length 40-50 mm, width 1.6 mm 

NW France, English Channel, 

British Isles, shallow water 

Figs from left: maxillae; 


hooded hook; parapodium 16; 

parapodium 136. From Carrera- 

Parra (2006a).

Lumbricalus Frame, 1992 


Diagnosis. Prostomium conical. Parapodia uniramous, with simple limbate chaetae, composite 

spinigers, and composite and simple multidentate hooded hooks. Mx III with one to three teeth. Mx IV 

unidentate, mx V free, placed outside of mx IV. 

Taxonomy. The genus is characterised by the presence of composite spinigers in the anterior part of 

the body. The spinigers are situated in the middle part of the bundle of chaetae. The genus was revised 

by Carrera-Parra (2004). Most species are found in warm and temperate waters. Only one species is 

known from the area. 

Anterior parapodia (1- ca. 30), with limbate chaetae, composite spinigers and composite 

hooded hooks with long blades. All chaetae reddish. Postchaetal lobe tongue-shaped, 

obliquely conical. Aciculae black. Mx III unidentate with a subdistal swelling ……………. 

……………………………… Lumbricalus campoyi Aguirrezabalaga & Carrera-Parra, 2006 

Length > 100 mm, width 4 mm 

Bay of Biscay, 900-1000 m 

Figs from left: parapodium 12; 

parapodium 140; composite spiniger; 


hooded hook; maxillae. From 

Aguirrezabalaga & Carrera-Parra 

(2006). 

19

20 

Lumbrinerides Orensanz, 1973 

Characteristics. Prostomium long, distally pointed. Parapodia uniramous, with simple limbate chaetae 

and simple bidentate hooded hooks. Aciculae yellow or black. Mx I usually with 1-2 accessory teeth 

on inner margin. Mx III with two aliform expansions. Mx IV without teeth. Mandibles usually fused 

for entire length. 

Taxonomy. The genus is characterised by the presence of bidentate hooded hooks and the shape of the 

maxillae. The species may be separated on the presence and position of small accessory teeth at the 

inner rim of mx I and the position of bidentate hooks. The genus was revised by Miura (1980) who 

reported three species from NE Atlantic waters. 

1. Mx I without accessory teeth ................................................................................................. 2 

- Mx I with one accessory tooth on inner border. Prostomium long. Simple hooded hooks from 

chaetiger 6 ...................................................... Lumbrinerides crassicephala (Hartman, 1965) 

Length 20-30 (?) mm, width 0.6 mm 

North Atlantic, Faroe Islands, deep 

water 

Figs from left: anterior end with 

pharynx slightly everted; bidentate 

hook; maxillae. From Hartman 

(1965) 

2. Prostomium very long, about three times as long as wide, 3-5 anterior parapodia reduced, 

bidentate hooks from chaetiger 1 .....................................Lumbrinerides laubieri Miura, 1980 

Length 20 (?) mm, width 0.3 mm 

Bay of Biscay, deep water 

Figs: anterior part of body; maxillae. 

From Miura (1980). 

- Prostomium about twice as long as wide, 7-9 anterior parapodia reduced, bidentate hooks 

from chaetiger 2-6 …………………………..…….... Lumbrinerides amoureuxi Miura, 1980 

Length ?, width 1 mm 

Bay of Biscay, shallow to deep water 

(1100 m). 

Figs from left: anterior end; bidentate 

hook; maxillae. From Miura (1980).

Lumbrineriopsis Orensanz, 1973 


Characteristics. Parapodia uniramous, with simple limbate chaetae and simple bidentate hooded 

hooks. Aciculae yellow or black. Mx III with two aliform expansions. Mx IV finely denticulate. 

Pygidium rounded. 

Taxonomy. The genus is characterised by the presence of bidentate hooded hooks and the shape of the 

maxillae. The genus was revised by Miura (1980). Four species are known from NE Atlantic waters. 

1. Yellow aciculae .................................................................................................................... 2 

- Black aciculae ....................................................................................................................... 3 

2. Prostomium prolonged ………….............…. Lumbrineriopsis paradoxa (Saint-Joseph, 1888) 

Length ?, width 0.5 mm 

France, English Channel, shallow water. 

Deep water, NE Atlantic and off 

Bermuda. 

Figs from left: anterior end; parapodium 

from middle body; bidentate hook; 

maxillae. From Miura (1980). 

- Prostomium short, slightly longer than wide ........................................................................... 

............................................................. Lumbrineriopsis sp. (L. paradoxa sensu Fauvel 1923) 

Length ?, width 1 mm 

France, English Channel, shallow water 

Figs from left: anterior end; limbate 

chaeta; bidentate hook; maxillae. From 

Fauvel (1923). 

3. Hooded hooks with 2-5 needle-shaped denticles between the two main teeth ………………. 

……............................................... Lumbrineriopsis tsushimaensis Imajima & Higuchi, 1975 

Length 50 mm, width 1 mm 

North Sea, 50-200 m 


hook chaetiger 8; bidentate hook 

posterior chaetiger. From Imajima & 

Higuchi (1975). 

21

22 

- Hooded hooks with an open space between the two main teeth …………............................... 

................................................................................Lumbrineriopsis gasconiensis Miura, 1980 

Length 10-20 (?) mm, width 0.9 mm 

Bay of Biscay, deep water 


hook; maxillae. From Miura (1980).

Lumbrineris Blainville, 1828, emended Carrera-Parra 2006b 


Characteristics. Prostomium conical or globular. Parapodia uniramous, with limbate chaetae and 

simple and composite multidentate hooded hooks. Aciculae yellow or black. Maxillary apparatus with 

five pairs of maxillae, mx I and II of about equal size, mx III and IV with 1-2(4) teeth. Mx V freestanding, 

placed outside of mx IV. 

Taxonomy: The genus Lumbrineris was previously (e.g. Fauvel 1923) rather widely defined, but has 

gradually become more restricted as new genera have been erected for species with particular 

characteristics (Orensanz 1973, 1990; Frame 1992; Carrera-Parra 2006a, b). Several of the early 

described species have been imperfectly diagnosed and reported from world-wide areas (e.g. L. 

latreilli) due to confusion with similar species. The genus was revised by Carrera-Parra (2006b) who 

provided a new description of Lumbrineris latreilli based on type material. He also synonymised L. 

agastos with L. futilis. The number of species in the region is uncertain. Small specimens with yellow 

aciculae and unidentate mx III may represent a species complex. 

1. Yellow or light brown aciculae ………………..…………................................................... 2 

- Black aciculae, mx III unidentate with a subdistal knob or swelling. Composite hooded hooks 

in chaetigers 1-20(23), with long blades .............……… Lumbrineris futilis (Kinberg, 1865) 

[= L. agastos Fauchald, 1974; fide Carrera- 

Parra 2006b] 


North Sea, British Isles, Norwegian 

coast, Faroe Islands, Bay of Biscay, 

moderate depths 

Figs from left: anterior end; parapodium 

6; composite hooded hook; simple 

hooded hook; maxillae. From Fauchald 

(1974; as L. agastos) and Carrera-Parra 

(2006b). 

2. Mx III unidentate …………………………………………………………………………… 3 

- Mx III bidentate …………………………………………………………………………….. 5 

3. Composite hooded hooks with moderately long blade in anterior chaetigers (length: width = 

5.5-8:1) …..…………………………………… Lumbrineris cf. kerguelensis (Grube, 1878) 


Central North Atlantic, north of 

Ireland, depth > 2000 m. 


parapodium 70; composite hooded 

hook, simple hooded hook; mx III- 

IV(from type specimen); composite 

hooded hook (specimen from 

Kerguelen). From Carrera-Parra 

(2006b) and McIntosh (1885). 

- Composite hooded hooks with short blade in anterior chaetigers (length: width = 4-5:1) 

……………………………………………………………………………………………….. 4 

23

24 

4. Mx III with arcuate cutting edge leading to a ventral expansion approching the shape of a 

tooth (giving the impression of a plate with two widely separated teeth), middle and posterior 

simple hooded hooks with 4-8 small teeth ……... Lumbrineris near cingulata (Ehlers, 1897) 

(= Lumbrineris gracilis sensu Hartmann-Schröder 

1996, George & Hartmann-Schröder 1985; non 

Ehlers, 1868) 

Length 30-50 mm, width 0.8-1 mm 

France, British Isles, eastern and western 

North Atlantic, North Sea, shelf and coastal 

waters in southern and western Norway. 

Figs from left: Anterior end; mx III-IV; 

composite hooded hooks chaetiger 4; 

simple hook chaetiger 15; maxillae (from 

type of L. cingulata). From Winsnes (1980; 

as L. gracilis) and Carrera-Parra (2006b). 

- Mx III with slightly curved cutting edge, middle and posterior simple hooded hooks with 6-10 

small teeth ………..…………………….……………...... Lumbrineris aniara Fauchald, 1974 

Length 40-50 mm, width 0.9-1.3 mm 

North Sea, southern and western 

Norway, shelf and coastal waters, 

?Bay of Biscay 

Figs from left: anterior end; anterior 

parapodium; composite hooded hook; 

simple hooded hook; maxillae. From 

Fauchald (1974) and Carrera-Parra 

(2006b). 

Remarks: Specimens with characters in between L. aniara and L.near 

cingulata are regularly found in the area. In northern waters species with both 

composite and simple hooded hooks in anterior chaetigers occur (L. 

vanhoeffeni (Michaelsen, 1898): Greenland; L. mixochaeta Oug, 1998: 

northern Norway, Barents Sea) 

5. Prostomium conical. Composite hooded hooks on chaetigers 1-20(24), with long blades in 

anterior setigers …………………......Lumbrineris latreilli Audouin & Milne-Edwards, 1834 

Length 100-200 mm, width 3-5 mm 

France, British Isles, North Atlantic to 

Iceland, North Sea, Skagerrak, shallow 

to deep water. 



hook from anterior chaetiger; simple 

hooded hook; maxillae. From Carrera- 

Parra (2006b).


- Prostomium round, globular. Parapodia with well-developed postchaetal lobes. Blades of 

composite hooded hooks short, of similar length throughout …................................................... 

………………………………………………..……… Lumbrineris cf coccinea (Renier, 1804) 

Length 50 mm, width 1.3 mm 

25 

W France, English channel, intertidal 

and shallow water 



hook; simple hooded hook; maxillae. 

From Fauvel (1923) and Carrera- 

Parra (2006b).

26 

Ninoe Kinberg, 1865 

Characteristics. Prostomium conical, with a pair of dorsal slit-like organs. Parapodia uniramous, with 

simple limbate chaetae and simple multidentate hooded hooks. Anterior parapodia with a number of 

digitiform postchaetal branchial lobes. Mx IV or mx III and IV with denticulate incisive edges. 

Taxonomy: The genus is characterised by the branched branchial lobes in the anterior part of the body 

and the dentition of mx III and IV. The genus is represented by a number of species in warm waters. 

One species is known from the area. 

Branchiae from chaetiger 5-6 to 31-39, up to five filaments from chaetiger 20, simple hooded 

hooks from chaetiger 7-15, mx III bidentate, mx IV with 7-8 fine teeth ……........………… 

............................................................................................ Ninoe armoricana Glemarec, 1968 

Length 30-40 mm, width 1.5 mm 

N Atlantic, Bay of Biscay, 

moderate depth 

Figs from left: anterior end; 

parapodium 24, anterior view; 

maxillae. From Glémarec (1968).

Scoletoma Blainville, 1828 


Characteristics. Prostomium conical or globular, occasionally with a single occipital antenna. 

Parapodia uniramous, with simple limbate chaetae and simple multidentate hooded hooks. Aciculae 

yellow or black. Mx III unidentate or bidentate. Mx V free-standing, placed outside of mx IV. 

Taxonomy: Frame (1992) resurrected Scoletoma as a valid genus to encompass species with simple 

hooded hooks and simple limbate chaetae, which previously had been referred to Lumbrineris. As 

currently defined, Scoletoma and Lumbrineris are rather close, and are essentially distinguished only 

by the lack and presence of composite hooded hooks, respectively (Frame 1992, Carrera-Parra 2006a). 

Scoletoma impatiens (Claparède) has been synonymised with Lumbrineris tetraura (Schmarda) by 

various authors. Winsnes (1980) and Orensanz (1990), however, have clearly indicated that the two 

species are different. Scoletoma funchalensis (Kinberg) is poorly known and possibly confused. 

Note: Juvenile specimens of Lumbrineris and Hilbigneris may lack composite hooded hooks and key 

out as Scoletoma. 

1. Yellow or golden aciculae, mx III unidentate or bidentate …….............................................. 2 

- Dark brown or black aciculae, mx III unidentate or unidentate with subdistal swelling ......... 4 

2. Mx III unidentate, anterior chaetigers with limbate chaetae only, hooded hooks from chaetiger 

6-15 ...……………………………………………. Scoletoma atlantica sensu Hartman (1965) 

Length 25 mm, width ? 

Central and NW North Atlantic, deep 

water 

Figure: maxillae (mx V omitted) 

From Hartman (1965) 

- Mx III bidentate, anterior chaetigers with limbate chaetae and hooded hooks ……………… 3 

3. Capillary chaetae extending backwards to chaetiger 50-60 …………….……………………... 

..................................................................................... Scoletoma impatiens (Claparède, 1868) 

27 

[=Lumbrineris tetraura: Hartmann-Schröder 

1996, non Schmarda, 1861] 

Length > 100 mm, width 3-5 mm 

North Atlantic, widely distributed 

Figs from left: anterior end; 

parapodium 18; posterior hooded 

hook; maxillae. From Hartmann- 

Schröder (1996) (as L. tetraura).

28 

- Capillary chaetae limited to the first 10-20 chaetigers …………………………..…………… 

... …………………………………………….... Scoletoma funchalensis sensu Fauvel (1923) 

Length 15-60 mm, width ? 

Bay of Biscay, English channel, 

British waters, shallow water 

Figs from left: limbate chaeta; 

hooded hook; anterior parapod: 

middle parapod. From Fauvel 

(1923). 

Remarks: This form could possibly represent juveniles of several species, 

also including species with composite hooks (Hilbigneris pleijeli) 

4. Mx III unidentate, anterior parapodia with short flap-like postchaetal lobes, occasionally with 

1-2 long hooded hooks, usually with limbate chaetae only …………………………………... 

………………………………………..……………… Scoletoma fragilis (O.F. Müller, 1776) 

Length 100-200 mm or more, 

width 4-12 mm 

North Sea, ?British Isles, Faroe 

Islands, Norwegian Sea, Arctic, 

shallow water to moderate depth 

Figs from left: anterior end 

ventral; parapodium 9; maxillae 

(left mx III-V omitted). From 

Winsnes (1980) and Frame 

(1992). 

- Mx III unidentate with a subdistal knob or swelling, anterior and middle parapodia with 

tongue-shaped, obliquely conical postchaetael lobes, usually with one, occasionally with 2-6 

long hooded hooks ………….…........................ Scoletoma magnidentata (Winsnes, 1981) 

Mx VIV 

Mx II 

Mx III 

Length 100-200 mm, width 3- 

5 mm 

North Sea, British Isles, Faroe 

Islands, Norwegian coast to 

Lofoten islands, shallow water 

to moderate depth 

Figs from left: anterior end 

ventral; parapodium 25; mx 

II-IV. From Winsnes (1981). 

Remarks: Very close to S. acicularum (Webster & Benedict, 1887) from NW 

Atlantic

Species of uncertain status 


Lumbrineris minuta (Théel, 1879). 

The species was described from Novaya Zemlya. The original description is comprehensive and well 

illustrated (Théel 1879), but apparently mixes characters from several species. Oug (1998) examined 

parts of the original material and confirmed that it consists of two or more species. In faunal surveys 

the species name appears to have been used for several small species from Arctic and North Atlantic 

waters belonging in Abyssoninoe, Scoletoma and Lumbrineris. Oug (1998) concluded that the species 

is indeterminate and needs to be re-examined to assess its status. 

Lumbrineris labrofimbriata (Saint-Joseph, 1888) 

The species was described from Brittany, France. Comments to the species and description of 

characters have been given by Laubier (1959) and Ramos (1976). The main characteristics are a 

denticulate inner margin of the mandibles and presence of both composite and simple hooded hooks in 

anterior chaetigers. Carrera-Parra (2006b), however, noted that these features occur during ontogenetic 

development in Lumbrineris and considered the species as invalid. 

Lumbrineris cluthensis (Clark, 1953) 

The species was described from Firth of Clyde, west coast of Scotland. Several specimens have later 

been found in British coastal waters (O’Reilly & Mackie, pers. commn). It is characterised by having 

blunt spine-like hooks and limbate chaetae in all chaetigers. Hooded hooks are absent. The maxillary 

apparatus is supported by two pairs of carriers, of which the posterior is the largest (Clark 1953, 

George & Hartmann-Schröder 1985). The species does not comply with the definition of the 

Lumbrineridae due to its lack of hooded hooks and is possibly more closely affiliated to Oenonidae. 

Lumbrineris cluthensis 


mandibles; maxillae; parapodium 1; 

parapodium 85; chaetae. From Clark 

(1953) 

Scoletoma punctata (McIntosh, 1885) 

The species was described from NW Atlantic deep water. According to the original description 

(McIntosh 1885), it is characterised by a prolonged triangular prostomium with six brown 

symmetrically arranged pigment spots at the posterior border. Hooded hooks are slender. Maxillae II 

are asymmetrical with two teeth on the left plate and several on the right, mx III and IV are unidentate. 

Mx V (‘accessory plates’) appear to be missing. Frame (1992) considered this species to belong in 

Scoletoma, but the shape of the prostomium, the long hooks and the possible lack of mx V may 

suggest that it should be referred to Abyssoninoe or Cenogenus. 

29

30 

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31

Guide to identification of Lumbrineridae - NMBAQC

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