European Journal of Taxonomy 27: 1-9

http://dx.doi.org/10.5852/ejt.2012.27

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2012 • Teresa Darby shire

Research article

urn:lsid:zoobank.org:pub:09359E10-277B-4018-BC69-7944D8DE6FD4

Re-description of Dysponetus joeli Olivier et al., 2012 (Polychaeta,

Chrysopetalidae), with a new key to species of the genus

Teresa DARBYSHIRE

Department of Biodiversity & Systematic Biology, Amgueddfa Cymru - National Museum Wales,

Cathays Park, Cardiff CF10 3NP, Wales, U.K.

Email: [email protected]

um:lsid:zoobank.org:author:B7103B9C-2AlA-4ADC-BC28-55EA9B8850C9

Abstract. Dysponetus is a genus of the family Chrysopetalidae with twelve currently described species.

Specimens are fragile and easily damaged or broken during sampling making identification difficult.

The most recently described species, Dysponetus joeli Olivier et al., 2012, from the Bay of Biscay and

the English Channel, was described from a few small, damaged and poorly preserved specimens. New

specimens from the Isles of Scilly, in much better condition, resembled D. joeli except for the absence

of ventral cirri on segment 3. Examination of the type material of D. joeli showed it to be identical to

these new specimens and highlighted errors in the original description of the species. The present paper

corrects the errors and a revised key to species is produced. The differences between D. joeli and the

two most similar species, D. bipapillatus Dahlgren, 1996 and D. macroculatus Dahlgren, 1996 are also

detailed.

Keywords. Dysponetus , tentacular cirri, ventral cirri, cirrophores.

Darby shire T. 2012. Re-description of Dysponetus joeli Olivier et al ., 2012 (Polychaeta, Chrysopetalidae), with

a new key to species of the genus. European Journal of Taxonomy 27: 1-9. http://dx.doi.org/10.5852/eit.2012.27

Introduction

There are currently twelve described species of the genus Dysponetus Levinsen, 1879 from around the

world. Most occur in shallow water and, with a few exceptions, are distributed in the Atlantic region. All

are small and fragile and rarely recorded from surveys, most likely because they are easily overlooked or

found in fragments not identifiable to species level. Several species are known only from their original

description and, even where multiple records do exist, these number in single figures.

Five species have been described for the northeast Atlantic region: D. caecus (Langerhans, 1880), D.

gracilis Hartman, 1965, D. paleophorus Hartmann-Schroder, 1974, D. pygmaeus Levinsen, 1879 and

D. joeli Olivier et al. , 2012. All of these have potential to be found in UK waters although only two

currently are. Dysponetus gracilis is, as yet, only recorded from deep water (>400 m) samples outside

of the UK (Hartman 1965; Hartman & Fauchald 1971; Aguirrezabalaga et al. 1999), D. paleophorus has

not been recorded since its first description off Norway and records for D. pygmaeus are restricted to the

Arctic and Japanese Pacific (Levinsen 1879; Annenkova 1935; Imajima & Hartman 1964). Dysponetus

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European Journal of Taxonomy 27: 1-9 (2012)

caecus , the most recorded Dysponetus species, has been identified from shallow littoral samples from

the UK (Scotland: Dahlgren & Pleijel 1995) as well as from other European locations (Laubier 1964; O’

Connor 1981; Kirkegaard 1992; Parapar & San Martin 1993; Dahlgren & Pleijel 1995; Brito et al. 1996).

An anomalous addition to these records would appear to be the records of Boggemann (2009) from the

abyssal southeast Atlantic. However, Boggemann addresses this anomaly in his paper by suggesting that

the lack of previous records from this region and depth is related to its small size and fragility (probably

the scarcity of previous samples too). Most recently, D. joeli was described from four shallow sites in

the UK and France.

In 2009, a National Museum Wales subtidal survey of the Isles of Scilly, an island archipelago off

the southwest tip of the UK, collected five specimens of Dysponetus in coarse sandy sediments.

Initial examination of the specimens showed them to possess four eyes, a character possessed by half

of the Dysponetus species, none of which, at that time, were described for the UK. Other characters,

including the absence of ventral cirri on segment 3, were found that differentiated it from all but two

other Dysponetus species, D. bipapillatus Dahlgren, 1996 from the Mediterranean and D. macroculatus

Dahlgren, 1996 from Papua New Guinea. The description of D. joeli was published shortly before the

description was finished and the two species showed such great similarity that the type specimens of

the latter were obtained and examined in order to determine whether the differences were indeed real

or might be due to the damaged nature of the specimens used. Examination of the holotype of D. joeli

revealed that the ventral cirri of segment 3 were in fact entirely absent rather than merely detached and

lost, and thus the specimens were indeed the same species. Consequently, this required distinguishing

characters between D. joeli, D. bipapillatus and D. macroculatus to also be detailed. A full re-description

is therefore made to clarify the species characters and differentiate it from the two most similar species.

Materials and Methods

Specimens of Dysponetus joeli were collected during a subtidal survey of the Isles of Scilly in June

2009. Samples were collected using a 0. lm 2 Van Veen grab from the R. V. Sepia of the Marine Biological

Association UK. Sediment was elutriated gently with seawater and the resulting sample sorted live the

same day. Animals were relaxed in a 7% magnesium chloride solution and fixed in 4% formaldehyde.

Final preservation was in 80% alcohol with 2% propylene glycol added.

The type specimens of D. joeli were collected from shallow maerl beds by Day, Van Veen or Smith-

Mclntyre grab and sieved through a 0.5 or 1.0 mm mesh (Olivier et al. 2012).

All drawings and measurements were made using a camera lucida attachment on a Nikon Labophot-2

compound microscope or a Nikon Eclipse E400 binocular microscope. Microscope photographs were

taken using AutoMontage™ software and SEM images were obtained using a Neoscope SEM.

Type specimens of D. joeli are held in the collections of the Museum National d’Histoire Naturelle

in Paris (MNHN) and the new specimens from the Isles of Scilly are in the National Museum Wales

collections (NMW.Z). Specimens of D. bipapillatus and D. macroculatus were borrowed from the

Swedish Museum of Natural History (SMNH) to enable a full comparison of characters.

Results

Phylum Ann elida Lamarck, 1809

Class Polychaeta Grube, 1850

Family Chrysopetalidae Ehlers, 1864

Genus Dysponetus Levinsen, 1879

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DARBYSHIRE T., Redescription of Dysponetus joeli Olivier et al., 2012

Dysponetus joeli Olivier et al., 2012

Figs 1A-D, 2A-D

Dysponetus joeli Olivier et al., 2012: 989-996, figs 1-3.

Material examined

Chausey, France, Sta. SSMM01 (48° 55.570’ N, 001° 48.270’ W), maerl, 10.0 m, holotype (MNHN

POLY TYPE 1533), paratype (MNHN POLY TYPE 1534), 18 Apr. 2006; off St Mary’s, Isles of Scilly,

Sta. 3b (49° 54.20’ N, 006° 18.94’ W), coarse sand/shell/gravel with some silt/clay, 25 m, 2 specimens

(NMW.Z.2009.027.0001-0002), 25 Jun. 2009; west of St Martin’s, Isles of Scilly, Sta. 14b (49° 57.86’

N, 006° 15.21’ W), shell gravel in muddy sand, 35.1 m, 2 specimens (NMW.Z.2009.027.0003-0004), 27

Jun. 2009; west of Isles of Scilly, Sta. 24b (49° 55.22’ N, 006° 23.91’ W), silty coarse sand/shell gravel,

47.1 m, 1 specimen used for SEM (NMW.Z.2009.027.0005), 29 Jun. 2009.

Fig. 1 . Dysponetus joeli Olivier et al., 2012. A. NMW.Z.2009.027.0003, anterior end, dorsal view.

B. NMW.Z.2009.027.0004, anterior end, ventral view. C. NMW.Z.2009.027.0003, posterior end, ventral

view. D. Holotype (MNHN POLY TYPE 1533), anterior end, ventral view. Only a few chaetae drawn

in each case for clarity. - ac = accessory chaeta, d = dorsal (cirrus/cirrophore), la = lateral antenna,

ma = median antenna, m = mouth appendage, p = palp, ps = palp scar, v = ventral (cirrus/cirrophore);

numbers indicate the segment.

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European Journal of Taxonomy 27: 1-9 (2012)

Description

Holotype in 4 pieces, posteriorly incomplete, 3.1 mm long for 19 chaetigers. Paratype in 3 pieces,

anterior dissected fragment of 5 chaetigers with 2 further detached chaetigers. 3 non-type specimens

up to 6 mm long, 0.35 mm wide (between segments, not including parapodia or chaetae) with 25-33

chaetigers. Two other incomplete non-type specimens examined. The following description is based on

entire specimens from the Isles of Scilly, except where specified as pertaining to the type material.

Body shape cylindrical, ventrally flattened, tapered slightly at posterior. Body pale cream in colour

(fixed, unstained), eyes dark reddish brown (Fig. 2 A) but pigment may degrade in alcohol. Eyes of type

specimens already barely discernable.

Prostomium oblong, wider anteriorly (Fig. 1A). Four large, rounded eyes visible in fixed specimens,

anterior pair larger and more widely separated than posterior. Median antenna small, bottle-shaped,

arising antero dors ally from prostomium. Lateral antennae bottle-shaped, slightly smaller than median,

arising immediately dorsal to palps. Antennae without distinct ceratophores. Palps directed posteriorly,

stout, oval, more than twice as long as wide. No antennae or palps remaining on type specimens

examined. Nuchal organs not observed.

Distinct single mouth appendage on lower lip, anteriorly directed, digitiform with blunt tip (Fig. IB,

2B). Single pair of stylet-shaped jaws, visible through body wall with methyl green staining. Proboscis

not observed.

First two segments slightly elevated dorsally with four pairs tentacular cirri, longer than but with same

shape as dorsal cirri of third and following segments, anteriorly directed. First segment achaetous,

second segment with notochaetae only, situated slightly anterior to dorsal tentacular cirrus. Third

segment biramous; dorsal cirri present, ventral cirri absent (Figs 1A, B, D, 2B). Holotype lacking all

cirri on anterior three segments, but presence indicated by cirrophores (Fig. ID). Following segments all

biramous with both dorsal and ventral cirri. Single noto- and neuroacicula present in each parapodium.

Notopodial lobes reduced. Dorsal cirri long, slender, longer than chaetae (21CM30 pm, longest on

median chaetigers), cirrophores present. Styles slightly proximally swollen, distally tapering, tips blunt.

Notoacicula difficult to detect. Notochaetae inserted dorsal to cirrus, densely packed, directed posteriorly

leaving middle part of dorsum exposed. Chaetae D-shaped in cross-section (Fig. 2C) with denticles

sharply pointed, in two parallel rows, 15-20 on each side. Notochaetal count, mid-body segments, up

to 26.

Neuropodia well-developed, conical mounds. Compound neurochaetae, with heterogomph shafts and

fine bidentate falcigerous blades (Fig. 2D). Neurochaetal count, mid-body segments, at least 20-26

(chaetae densely packed and difficult to accurately count). Up to two accessory simple chaetae, similar

to but smaller than notochaetae, inserted distally and anteriorly on neuropodial lobe (Fig. 1C). Ventral

cirri fusiform, shorter than dorsal cirri (length 110-270 pm, longest on median chaetigers), arising

posteroventrally on neuropodial lobe (Fig. 1C).

Final segment lacking noto- and neurochaetae, cirrophores of dorsal and ventral cirri observed although

only a single, rounded dorsal cirrus observed on 1 specimen (NMW.Z.2009.027.0003). Pygidium conical

with single projection (length 40 pm), cylindrical, slightly distally tapering, inserted posteroventrally

(Fig. 1C), anus terminal.

Eggs visible within one specimen (NMW.Z.2009.027.0001), flattened oval in shape, maximum width

approx. 50 pm, possibly immature.

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DARBYSHIRE T., Redescription of Dysponetus joeli Olivier et al., 2012

Habitat

The species was originally described from maerl beds in the Bay of Biscay and the English Channel. The

new specimens are all from coarse sand and shell sediments collected around the Isles of Scilly.

Remarks

The original description of D. joeli was based on small, damaged specimens and this may explain why

the absence of the ventral cirri on chaetiger 3 was attributed to loss of the cirri rather than actual absence.

Although cirri are easily lost from specimens, the cirrophores can still be seen under light microscopy at

x400 magnification or greater. Application of methyl green staining can help distinguish these features.

Under SEM conditions the absence of cirrophores on chaetiger 3 is obvious (Fig. 2B).

An additional complication in determining the described characters of D. joeli arises from the character

matrix scores in the original paper (Olivier et al. 2012). In their species description, the authors stated

that there were 2 pairs of tentacular cirri on the first 2 segments but made no mention whether segment

3 possessed or lacked ventral cirri. They then later scored ventral cirri as absent for the first 2 segments

(no separate score for tentacular cirri) and present for the third in the character matrix However, in the

Fig. 2. Dysponetus joeli Olivier et al ., 2012. A. NMW.Z.2009.027.0003, whole specimen, dorsal

view (scale bar 1 mm). B. NMW.Z.2009.027.0005, anterior end, ventrolateral view (scale bar 50 pm).

C. Notochaetae (scale bar 10 pm). D. Neurochaetae (scale bar 20 pm). - d = dorsal (cirrophore),

la = lateral antenna, m = mouth appendage, p = palp, ps = palp scar, v = ventral (cirrophore); numbers

indicate the segment.

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European Journal of Taxonomy 27: 1-9 (2012)

species key itself, D. joeli is keyed out with ventral cirri being present on the 3 rd segment (i.e. chaetiger

2). It is apparent from the character states scored for the other species in the matrix that the authors have

used the term ‘ventral cirrus’ interchangeably with ‘tentacular cirrus’. Earlier authorities have also been

variable in their use of the terms ‘tentacular cirrus’ versus ‘cirrus’ with regards to the appendages of the

first 3 segments in species of Dysponetus. In this paper, only the cirri of the first 2 segments, which lack

parapodia and have cirrophores directly attached to the body wall, are regarded as being tentacular. A

later paper will discuss the different characters of Dysponetus species in more detail.

The lack of ventral cirri on segment 3 distinguishes D. joeli from most other Dysponetus species. The only

other species with this characteristic are D. bidentatus Day, 1954, D. bipapillatus and D. macroculatus ,

although D. bidentatus lacks ventral cirri on segment 2 also (Day 1954). The presence or absence of

ventral cirri on segment 3 is unknown for D. hebes (Webster & Benedict, 1887); however, this species

differs considerably by having a double mouth appendage as opposed to single, and sphaerical not

elongated palps. With the revised character for segment 3, the species key in Olivier et al. would place

D. joeli with both D. bipapillatus and D. macroculatus. The latter two species are distinguished in the

key according to the relative size of the eyes; however, this is not a good character as the eye pigments

degrade in alcohol and disappear over time - as is already evident in the type specimens. A new revised

key to the 12 species of the genus is provided below.

Dysponetus joeli is a much larger species than both D. bipapillatus and D. macroculatus. Even the type

specimens, described as ‘small’ (ranging from 3.5—4 mm in length, incomplete) and smaller than those

from the Isles of Scilly, were 2-3 times the size of each of the latter species, respectively.

Dysponetus joeli is most similar to D. bipapillatus from the Mediterranean, sharing all of the obvious

characteristics. The paired papillae on segment 8 described for D. bipapillatus were not observed

on any of the D. joeli specimens; however, these appendages were only seen on a few specimens of

D. bipapillatus and are thus not a good character for comparison. In his paper, Dahlgren (1996) stated

that the specimens, though small, were considered mature adults due to the presence of the paired

papillae, interpreted as genital organs, and the development of the anterior segments. A comparison

of the individual characters shows D. joeli to possess twice as many neurochaetae as D. bipapillatus

across all segments (20-26 compared to 9-13), more numerous denticles on the notochaetae (15-20 as

opposed to 10-15) and a shorter pygidial projection (40 pm versus 50 pm), the last character particularly

noticeable in relation to the larger body size of D. joeli. Similarly, the number of neurochaetae in

D. joeli is greater than the 19-22 found on D. macroculatus , and, in general, the former species is a much

larger animal than the latter, being 2-3 times as long at maturity (as evidenced by the presence of eggs

in specimens of both).

Key to Dysponetus species

1 Ventral tentacular cirri absent on segment 1 .2

Ventral tentacular cirri present on segment 1 .6

2 Elongate palps; 4 eyes. D. bulbosus* Hartmann-Schroder, 1982

- Sphaerical palps; 2 eyes or eyes absent.3

3 Anterior median antenna; double mouth appendage on lower lip; 2 eyes.

. D. hebes (Webster & Benedict, 1887)

- Dorsal median antenna; eyes absent.4

Ventral tentacular cirri present on segment 2

Ventral tentacular cirri absent on segment 2 .

4

6

D. gracilis Hartman, 1965

.5

DARBYSHIRE T., Redescription of Dysponetus joeli Olivier et al., 2012

5 Mouth appendage absent; paleae present among notochaetae .

. D. paleophorus Hartmann-Schroder, 1974

Double mouth appendage on lower lip; paleae absent . D. pygmaeus Levinsen, 1879

6 Mouth appendage absent .7

- Single mouth appendage on lower lip .8

7 Elongate palps, ventral tentacular cirri present on segment 2. D. hesionides Boggemann, 2009

Sphaerical palps, ventral tentacular cirri absent on segment 2. D. profundus Boggemann, 2009

8 Eyes absent, ventral cirri present on segment 3 . D. caecus (Langerhans, 1880)

Eyes present, ventral cirri absent on segment 3 .9

9 16 or less notochaetae in mid-body segments . D. bulbosus* Hartmann-Schroder, 1982

- 20 or more notochaetae in mid-body segments .10

10 Ventral tentacular cirri absent on segment 2 . D. bidentatus Day, 1954

- Ventral tentacular cirri present on segment 2 .11

11 Few neurochaetae, only 9-13 in mid-body segments . D. bipapillatus Dahlgren, 1996

- Numerous neurochaetae in mid-body segments, 19 or more .12

12 Small body size, 2-3 mm; 19-22 neurochaetae. D. macroculatus Dahlgren, 1996

- Large body size, >3 mm; 20-26 neurochaetae . D. joeli Olivier et al ., 2012

* Descriptions of D. bulbosus (Hartmann-Schroder 1982, 1986) contradict each other over the presence

or absence of ventral cirri on each of the first few segments. For this reason, D. bulbosus is included

twice in the key to account for the discrepancy in descriptions. Dysponetus bulbosus will be reviewed

in a forthcoming paper.

Acknowledgements

The work in the Isles of Scilly was part of the Assessment of Marine Biodiversity Linked to Ecosystems

project funded by the Department for Enviro nm ent, Food & Rural Affairs (DEFRA). Thanks must go

to Dr Richard Warwick and Dr Paul Somerfield who jointly led the project, the Captain and crew of the

Marine Biological Association’s Research Vessel Sepia and everyone else involved with the sampling.

Elin Sigvaldodottir and Tarik Meziane are also thanked for the loan of specimens from the Swedish

Museum of Natural History, Stockholm and Museum National d’Histoire Naturelle in Paris that enabled

this paper to be written. Dr Andrew Mackie is thanked for his co mm ents and improvements on the paper.

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Manuscript received: 17 August 2012

Manuscript accepted: 19 October 2012

Published on: 8 November 2012

Topic editor: Rudy Jocque

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

that are members of the EJT Consortium: Museum National d’Histoire Naturelle, Paris, France; National

Botanic Garden of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium;

Natural History Museum, Fondon, United Kingdom; Royal Belgian Institute of Natural Sciences,

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

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