Zoological Journal of the Linnean Society, 2012, 164, 245–284. With 10 figures
Systematic revision of Sabellariidae (Polychaeta) and
their relationships with other polychaetes using
morphological and DNA sequence data
MARÍA CAPA*, PAT HUTCHINGS and RACHAEL PEART
Australian Museum, 6 College Street, Sydney, NSW 2010, Australia
Received 24 March 2011; revised 6 June 2011; accepted for publication 27 June 2011
The affinities of honeycomb or sandcastle worms (Sabellariidae, Polychaeta) and other polychaetes are studied
using morphological and DNA sequence data (18S rDNA, 28S rDNA, and EF-1 alpha). Maximum-parsimony
analyses were performed including 20 terminals and 7155 aligned characters. The monophyly of Sabellariidae is
confirmed and well supported and sister-group relationships with Spionida are suggested but only poorly
supported. Phylogenetic relationships within Sabellariidae are also assessed for the first time, using morphological
data. Maximum-parsimony analyses of 30 terminals and 31 characters were performed with and without weighting
the less homoplasious characters. Implied weighting resolved polytomies recovered after non-weighting datasets
and suggest that the established sabellariid subfamilies are not monophyletic and that the number of parathoracic
segments is homoplastic. Instead, some opercular features and chaetal characters not often incorporated in
descriptions are here shown to be phylogenetically informative and support some of the clades recovered. We
provide a description of morphological features of sabellariid and previously related groups together with
illustrations that will, we hope, be used as a baseline for further systematic and taxonomic studies in the group
and as a framework for future molecular studies. Generic diagnoses and a description and a key to genera are
provided.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284.
doi: 10.1111/j.1096-3642.2011.00767.x
ADDITIONAL KEYWORDS: character evolution – morphology – phylogeny – terminology revision.
INTRODUCTION
Sabellariids, commonly known as honeycomb or sandcastle worms, are easily recognizable by having wellconstructed tubes of cemented sand grains sometimes
attached to one another forming reefs, and by the
presence of a developed operculum with rows of
paleae that seals the entrance of the tube. The tube
and the operculum provide the worm with protection
from predators and desiccation and the operculum is
also used to clean the opening of the tube from fouling
animals, algae or other objects (Eckelbarger, 1976).
Most sabellariids live in intertidal or shallow depths
*Corresponding author. E-mail: maria.capa@austmus.gov.au;
capa.maria@gmail.com
but there are some genera and species restricted to
the continental shelf or the deep sea. Most genera
seem to be cosmopolitan but species appear to be
geographically restricted and show ecological and
bathymetric limitations caused by specific temperature and water circulation requirements as wells as
constraints in the type of substrate and sediment,
characteristics necessary for settling and building
their tubes (Kirtley, 1994; Bastida-Zavala & BecerrilTinoco, 2009).
Reef-building sabellariids have been the focus of
several taxonomic, biological and ecological studies
due to their importance as constructors of threedimensional structures, providing refuge and food for
many invertebrate species (e.g. Pawlik, 1988a, b;
Caline, Gruet & Legendre, 1992; Dubois, Retiere &
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
245
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M. CAPA ET AL.
Olivier, 2002; Sepulveda, Moreno & Carrasco, 2003;
McCarthy et al., 2008; Fournier, Etienne & Le Cam,
2010) but few studies have dealt with the systematics
of the group and none in a phylogenetic context. One
of the main taxonomic contributions to this group is
the worldwide taxonomic revision of the family undertaken by Kirtley in 1994. Some subsequent studies
have dealt with new species descriptions or partial
taxonomic revisions of certain genera (e.g. Lechapt &
Kirtley, 1998; Nishi & Kirtley, 1999; Nishi & Núñez,
1999; Nishi, Kato & Hayashi, 2004; Bailey-Brock
et al., 2007; Nishi et al., 2010), increasing the total
number of nominal species in the family to 120 (Nishi
et al., 2010) although no systematic revision has been
undertaken.
MONOPHYLY
OF
SABELLARIIDAE
AND ITS POSITION
ON THE ANNELID TREE
Even though the monophyly of Sabellariidae has not
been tested in a phylogenetic context, several notable
features, such as the presence of the operculum
bearing rows of paleae derived from the first two
segments, the presence of the tentacular filaments on
the operculum, the division of the body into four
regions, and the arrangement of chaetae, common in
all members of the group, have indicated its monophyly over the years. In contrast, the position and
sister-group relationships of Sabellariidae with other
polychaetes are still obscure (Rouse & Pleijel, 2003)
and different studies have indicated its inclusion
within Sabellida, Spionida, or Terebellida.
The group was initially grouped with Sabellidae
and Serpulidae (Levinsen, 1883; Meyer, 1888;
Hatschek, 1893; Benham, 1896) because of the
arrangement of thoracic nephridial pores and the
interpretation of the food-collecting organs of these
three families as homologous. This relationship was
also justified latterly by the chaetal inversion in the
abdominal region (Knight-Jones, 1981; Fitzhugh,
1989; Rouse & Fauchald, 1997) and the innervation of
the palps (Orrhage & Müller, 2005). But other studies
indicated that the tentacular filaments and palps in
sabellariids and radiolar crown in sabellids and serpulids have a different origin (Orrhage, 1978;
Orrhage & Müller, 2005) and therefore they should
not be considered as homologous. Recently, the
chaetal arrangement in Sabellariidae has also been
suggested as being not homologous to the sabellid–
serpulid chaetal inversion pattern (Kieselbach &
Hausen, 2008), providing no support for a sistergroup relationship of sabellids and sabellariids. Concurrently, a phylogenetic hypothesis combining
morphological and molecular data has also indicated
that Sabellariidae and Sabellida (including Sabellidae
and Serpulidae) are not closely related (Rousset et al.,
2004; Zrzavý et al., 2009; Capa et al., 2011).
Sabellariids have also been thought to be related to
spionids (Meyer, 1888; Caullery, 1914). A revealing
study by Dales (1962) places sabellariids within Spionida based on larval morphology and palp innervation. This relationship has also been recently suggested, after comprehensive analyses of combined
morphological and molecular data (Rousset et al.,
2004; Capa et al., 2011), as well as behavioural and
ecophysical investigations (Amieva & Reed, 1987;
Dubois et al., 2005).
Other authors have considered sabellariids as
related to Terebellida (Savigny, 1822; Fauchald, 1977)
or Terebelliformia (sensu Rouse & Pleijel, 2001),
because of the presence of buccal tentacles (referred
to tentacular filaments herein), and similarities
regarding larval metamorphosis and opercular structure between sabellariids and pectinariids have been
suggested (Rouse & Pleijel, 2001). The homology
between the opercular structures has still to be tested
(Orrhage, 2001; Bartolomaeus, Purschke & Hausen,
2005).
RELATIONSHIPS
WITHIN
SABELLARIIDAE
The taxonomic revision of the family Sabellariidae
was undertaken by Kirtley in 1994. The result was
the rearrangement of species within genera, erection
of new ones, increasing the number of genera to 12,
and the erection of two subfamilies, Sabellariinae and
Lygdamiinae. He provided a chronological account of
the confused taxonomy of the Sabellariidae from 1771
to 1994 but, unfortunately, he did not present a phylogeny of the group for testing the groupings he
proposed within the family or the homology of the
features he used for defining those two subfamilies. In
addition, this publication is not readily available as
he published it privately.
MATERIAL AND METHODS
The present study was split into two parts, each
analysing different sets of data, for addressing relationships at different levels. The aim of the first set of
analyses was to test the monophyly of Sabellariidae
and assess sister-group relationships with other polychaetes. The second set of analyses aimed to present
the first phylogenetic hypothesis for Sabellariidae and
appraise the relationships among its members. Taxa,
characters and methodology for each analysis are
described below.
TAXA
INCLUDED IN THE ANALYSES
For resolving the sister-group relationships of Sabellariidae and homologies of morphological features, a
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
SYSTEMATICS OF SABELLARIIDAE
group of polychaetes belonging to the orders Terebellida, Sabellida, and Spionida, and suggested to share
some similarities or to be related to the sabellariids,
were included in the analyses. The ingroup included
members of Sabellariidae, Sabellidae, Serpulidae,
Oweniidae, Magelona, Cirratulidae, Terebellidae,
Ampharetidae, Pectinariidae, Spionidae, Trochochaeta, Poecilochaeta, and Sternaspis and was composed
by 20 terminals. The sequences selected were those
with phylogenetic signal for the questions needed to
address herein (Halanych & Janosik, 2006) and available from GenBank. The outgroup included one
Nereididae and one Polynoidae, groups rather closely
related to members of the ingroup, but less closely
than any of the ingroup members is to each other,
according to latest phylogenetic hypotheses. Morphological features were scored from the literature and
direct observation of material if available.
For resolving relationships among sabellariids, 30
species belonging to the 12 sabellariid genera (Kirtley,
1994) and including all type species were included in
the data matrix, in order to incorporate the morphological variability within each genus and selecting
those species that we could access material for
(Table 1) or those which had detailed descriptions in
the literature. The outgroup was that recovered as
sister-group of Sabellariidae on the first analyses.
In the Systematic account, a diagnosis, description
and remarks of the genera are given according to the
material and literature examined. Some amendments
to previous descriptions have been made and are
commented on in the Remarks. The list of synonymies
reflects only those reviewed and should not be considered as a detailed taxonomic revision.
MORPHOLOGICAL
CHARACTERS AND
CHARACTER DISTRIBUTIONS
The morphological features selected for assessing
relationships at family level were scored from the
literature (e.g. Dales, 1962; Orrhage, 1980; Bartolomaeus, 1995, 1998; Purschke, 1997, 2005; Hausen &
Bartolomaeus, 1998; Rouse, 2000a, b; Zhadan &
Tzetlin, 2003; Bartolomaeus & Quast, 2005; Eckelbarger, 2005; Hausen, 2005; Orrhage & Müller, 2005;
Tzetlin & Filippova, 2005; Purschke & Müller, 2006;
Purschke & Hausen, 2007; Suschenko & Purschke,
2009). The list of 99 characters and states is summarized in Table 2 and described in Appendix 1. The
matrix (Table 3) was constructed in Nexus Data
Editor (Page, 1998). The ‘C-method’ proposed by
Pleijel (1995) was used for character scoring. The
codification scheme included absent/present characters and unordered multistate characters. Taxa
lacking the feature were scored as inapplicable and
indicated as ‘–’ and unknown as ‘?’.
247
Table 1. Material examined for scoring the sabellariid
matrix
Bathysabellaria neocaledoniensis Lechapt & Gruet, 1993,
New Caledonia, SW Pacific, paratype, MNHN POLY
TYPE 1174.
Bathysabellaria spinifera Lechapt & Kirtley, 1996, New
Caledonia, SW Pacific, paratype, MNHN POLY TYPE
1204.
Gunnarea gaimardi (Quatrefages, 1866), Cape of Good
Hope, holotype, MNHN POLY TYPE 588.
Idanthyrsus australiensis (Haswell, 1883), Australia,
Queensland, Thursday Island, type? BMNH
1882.2.22.71.
Idanthyrsus sp. nov. 1, Christmas Island, Indian
Ocean, WAM 359-75.
Idanthyrsus sp. nov. 2, Australia, Western Australia,
WAM 394-75.
Lygdamis augeneri Augener, 1927, Australia, New South
Wales, holotype, ZMH V-9569.
Lygdamis giardi (McIntosh, 1885), Australia, New South
Wales, holotype, BMNH 1885.12.1.6.
Lygdamis sp. nov., Queensland, QM G.10505, QM
G.10338.
Lygdamis japonicus Nishi & Kirtley, 1999, Japan, AM
W.37747
Neosabellaria vitiensis Bailey-Brock et al., 2007, Fiji,
paratypes, AM W.31026
Phalacrostemma sp. nov., Australia, New South
Wales, AM W.27566, AM W.27568.
Phalacrostemma profundum Lechapt & Kirtley, 1998,
New Caledonia, SW Pacific, paratypes, MNHN POLY
TYPE 1198, 1199.
Phragmatopoma californica (Fewkes, 1889), USA,
California, Coronado, type?, MNHN POLY TYPE 732.
Sabellaria ishikowai Okuda, 1938, Japan AM W.37748
Sabellaria issumiensis Nishi et al., 2010, Japan, Chiva
Prefecture, paratypes, AM W.36825
Sabellaria tottoriensis Nishi et al., 2004, Japan Tottori
Prefecture, paratypes, AM W.29058.
Sabellaria sp. nov. 1, Australia, Queensland, AM
W.27195, AM W.27333.
Sabellaria sp. nov. 2, Australia, Northern Territory,
NTM W.4798.
Tetreres robustus Lechapt & Kirtley, 1998, New
Caledonia, SW Pacific, paratypes, MNHN POLY TYPE
1200, 1201.
Abbreviations: AM, Australian Museum; MNHN, Museum
National d’Histoire Naturelle; NTM, Northern Territory
Museum; QM, Queensland Museum; WAM, Western Australian Museum; ZMH, Zoological Museum, Hamburg;
The characters selected for assessing relationships
within sabellariids were those considered as the
major features to characterize and distinguish the
genera and species within Sabellariidae (Kirtley,
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248
M. CAPA ET AL.
Table 2. List of characters and states for resolving sabellariid sister-group relationships
1. Prostomium and peristomium fused: (0) absent; (1) present.
2. Operculum formed by head and two segments: (0) absent; (1) present.
3. Radiolar operculum: (0) absent; (1) present.
4. Median antenna: (0) absent; (1) present.
5. Lateral antennae: (0) absent; (1) present.
6. Prostomial palps: (0) absent; (1) present.
7. Peristomial palps: (0) absent; (1) present.
8. Prostomial tentacles: (0) absent; (1) present.
9. Peristomial tentacles: (0) absent; (1) present.
10. Segment 1 tentacles: (0) absent; (1) present.
11. Building organ: (0) absent; (1) present.
12. Anterior glandular ventral shields: (0) absent; (1) present.
13. Thoracic membrane: (0) absent; (1) present.
14. Parapodial rami: (0) with similar structure rami; (1) unequal rami.
15. First few segments with no appendages: (0) absent; (1) present.
16. Notopodia: (0) not projecting; (1) projecting.
17. Notopodia missing in the posterior part of the body: (0) absent; (1) present.
18. Neuropodia: (0) not projecting; (1) tori; (2) projecting.
19. Parapodial lobes: (0) absent; (1) present.
20. Dorsal cirri: (0) absent; (1) present.
21. Ventral cirri: (0) absent; (1) present.
22. Chaetal inversion: (0) absent; (1) present.
23. Notochaetal arrangement: (0) transverse rows; (1) bundles.
24. Neuropodial arrangement: (0) transverse rows; (1) bundles.
25. Second chaetal formation site forming longitudinal rows: (0) absent; (1) present.
26. Paleae in first segments: (0) absent; (1) present.
27. Spines or hooks in first segments: (0) absent; (1) present.
28. Aciculae: (0) absent; (1) present.
29. Spines: (0) absent; (1) present.
30. Capillaries: (0) absent; (1) present.
31. Compound chaetae: (0) absent; (1) present.
32. Hooks: (0) absent; (1) present.
33. Hoods in hooks: (0) absent; (1) present.
34. Rostrum/main fang: (0) absent; (1) present.
35. Capitium/teeth over main fang: (0) absent; (1) present.
36. Subrostrum/breast: (0) absent; (1) present.
37. Subrostral process: (0) absent; (1) present.
38. Basal process: (0) absent; (1) present.
39. Manubrium/handle: (0) absent; (1) present.
40. Length of manubrium: (0) short (avicular); (1) long (acicular).
41. Posterior scaphe: (0) absent; (1) present.
42. Cauda: (0) absent; (1) present.
43. Pygidial cirri: (0) absent; (1) present.
44. Adult prostomial eyes: (0) absent; (1) present.
45. Type of eyes: (0) simple ocelli; (1) multicellular eyes.
46. Eyes lenses: (0) absent; (1) present.
47. Nuchal organs: (0) absent; (1) present.
48. Nuchal organs position: (0) posteriorly in prostomium; (1) pouches from the dorsal epithelium of mouth cavity; (2)
base of palps.
49. Nuchal organs structure: (0) pits or grooves; (1) posterior projections.
50. Histology of nuchal organs: (0) without cuticule; (1) with cuticle.
51. Lateral organs: (0) absent; (1) present.
52. Parapodial branchiae: (0) absent; (1) present.
53. Branchiae: (0) only on few anterior segments; (1) on most segments along body; (2) on posterior segments.
54. Origin of branchiae: (0) parapodial; (1) dorsum.
55. Circular muscles: (0) absent; (1) present.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
SYSTEMATICS OF SABELLARIIDAE
249
Table 2. Continued
56. Buccal organ: (0) absent; (1) present.
57. Type of buccal organ: (0) axial muscular proboscis; (1) ventral pharyngeal organ; (2) axial non-muscular proboscis.
58. Dorsolateral ciliary folds: (0) absent; (1) present.
59. Jaws: (0) absent; (1) present.
60. Gut: (0) strait tube; (1) straight tube with side branches; (2) loop; (3) distinct fold.
61. Proventricle: (0) absent; (1) present.
62. Faecal groove: (0) absent; (1) present.
63. Faecal groove inversion: (0) absent; (1) present.
64. Gular membrane: (0) absent; (1) present.
65. Location of gular membrane: (0) between segments 4 and 5; (1) between segments 3 and 4.
66. Head kidney (trochophore nephridia or first pair of differentiated nephridia): (0) protonephridia; (1) metanephridia.
67. Adult segmental nephridia: (0) protonephridia; (1) metanephridia.
68. Filtration structure: (0) podocytes; (1) solenocytes; (2) terminal cell monociliated; (3) terminal cell multiciliated.
69. Distribution of segmental organs: (0) in most segments; (1) several anterior nephridia and posterior gonoducts; (2)
one pair of anterior nephridia and posterior gonoducts.
70. Heart body: (0) absent; (1) present.
71. Blood: (0) acellular; (1) with haemocytes.
72. Blood colour: (0) red (with haemoglobin); (1) green (with chlorocruorin).
73. Nervous system: (0) subepidermal; (1) intraepidermal.
74. Mushroom bodies (corpora pedunculata): (0) absent; (1) present.
75. Reproduction: (0) gonochoric; (1) hermaphroditic.
76. Asexual reproduction: (0) absent; (1) present.
77. Fertilization: (0) external; (1) internal.
78. Structure of egg envelope: (0) smooth; (1) ornamented.
79. Brooding: (0) absent (broadcast spawning); (1) present (including jelly masses).
80. Discrete ovaries: (0) absent; (1) present.
81. Position of the ovary: (0) anterior segments; (1) posterior segments; (2) along the body.
82. Type of oogenesis: (0) intraovarian oogenesis; (1) extraovarian oogenesis.
83. Previtellogenic oocytes (extraovarian oogenesis): (0) leaving the ovary as individual cells; (1) leaving the ovary as
clusters of cells.
84. Sperm head: (0) rounded; (1) elongated.
85. Spermatophores: (0) absent; (1) present.
86. Spermatozegmata: (0) absent; (1) present.
87. Development: (0) direct; (1) larvae.
88. Larvae: (0) lecithotrophic; (1) planktotrophic.
89. Prototroch: (0) absent; (1) present.
90. Metatroch: (0) absent; (1) present.
91. Food groove: (0) absent; (1) present.
92. Oral brush: (0) absent; (1) present.
93. Akrotroch: (0) absent; (1) present.
94. Meniscotroch: (0) absent; (1) present.
95. Telotroch: (0) absent; (1) present.
96. Neurotroch: (0) absent; (1) present.
97. Apical tuft: (0) absent; (1) present.
98. Tube: (0) absent; (1) present.
99. Tube material: (0) mucous and sediment attached; (1) calcareous.
1994: 8), although some were excluded due to high
interspecific variability (e.g. ornamentation of thecae
of paleae and chaetae, the number of paleae, papillae
and buccal tentacles, and the shape, size and colour of
the building organ). In order to translate into a
matrix the complexity of paleal forms we have scored
different categories, including the general shape
(cylindrical, flat or concave), the shape of margins
(with or without obvious denticles), the angle formed
by shaft and blade (straight or geniculate), etc. After
examination of specimens, other features that show
variability within the group have been added to the
matrix to test the phylogenetic and taxonomic usefulness and to promote their description in future taxonomic descriptions. These are the relative length of
the opercular lobes, length of palps, relative length of
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
250
M. CAPA ET AL.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
Table 3. Character matrix of morphological features included for resolving sabellariid sister-group relationships
Nereididae
Polynoidae
Magelona
Poecilochaetus
Spionidae
Trochochaeta
Ampharetidae
Pectinaridae
Terebellidae
Cirratulidae
Sternaspis
Oweniidae
Serpulidae
Sabellidae
Sabellariidae
Nereis
Lepidonotus
Magelona
Poecilochaetus
Polydora
Trochochaeta
Auchenoplax
Pectinaria
Pista
Polycirrus
Cirriformia
Sternaspis
Owenia
Hydroides
Protula
Serpula
Bispira
Eudistylia
Schizobranchia
Idanthyrsus
Gunnarea
Sabellaria
0–10
11–20
21–30
31–40
41–50
51–60
61–70
71–80
81–90
91–99
0000110000
0001110000
0000001000
0000001000
000?001000
000B001000
0000000010
1100000010
1000000100
1000000100
000000?000
000000?000
10000?0?00
1010010000
1010010000
1010010000
1000010000
1000010000
1000010000
1100001001
1100001001
1100001001
0001010211
0001010201
000??10210
0001010A10
0001010A10
0001010010
0101?11100
110?110100
0101111100
0101111100
0000?00000
000?10?000
000??00000
0011010200
0011010200
0011010200
0101010200
0101010200
0101010200
1001010200
1001010200
1001010200
1011000100
1011000111
00000000?1
00B?100011
0000100011
00B?100011
0010001011
0010010001
0010000001
0010000011
0000000011
00??000011
00100000?1
0110000010
0110000010
0110000010
0110000010
0110000010
0110000010
0010011011
0010010011
001001B011
10-------00-------0111100011
00-------0111000011
00-------0100111110
0100110110
0101111111
0101111110
00-------00-------0100101011
0101111010
0101111010
0101111010
0101111010
0101111010
0101111010
010011100010011100010011100-
0011111000
0011111000
0010--0--001100101?
0011001011
00B100101?
0010--1100
1010--1100
0001001100
0000--1100
000B--100
0001000--0000--0--0000--1100
0001001100
0000--1100
0001101100
0001001100
0001001100
0101001200
0101001200
0101001200
00--110010
00--010011
10--011100
1121?11100
1101011100
10--012100
1101111002
1100111102
1101111102
10--1111020
0011111110
0121110?02
00--011100
00--10-000
00--10-000
00--10-000
00--10-000
00--10-000
00--10-000
011010-000
011010-000
011010-000
00-0-?1000
00-0-01200
00-0-0?21?
00-0-11010
00-0-1101?
00-0-?1?1?
0??10?1?11
0??110131?
0101001C21
101001D110
000-0-?1?21
00-0-?1?20
0100-01D?0
0110-01C21
0110-01C21
0110-01C21
0110-?1020
0110-?1020
0110-?1020
1100-01D20
1100-01D20
1100-01D20
000100?000
0?01?0?0?1
1?00000001
??0?00?101
??00000011
??00?0?101
0000000001
1000000001
0000001001
0000010010
0000000001
00000?00?1
0010000001
010?00000?
010?00000?
010?00010?
0100010001
0100000001
0100000001
0?00000001
0?00000001
0?00000001
---000??10
20?0?0??10
1??1?01110
?????011?0
0101101110
?????01110
0110001011
01B?011110
011001??10
110011010?
1110000-10
?1?0?010?0
?100?01111
210000??11
210000?B11
2100001111
?100001011
?100001011
?100001011
10-0001110
10-0001110
10-0001110
0???101??
01010110
000000100???10?10
000010110
000010?10
??0010110
000011110
??0011110
?0011110
00001100??000010100000110
100001111
100001111
100001111
?00001010
?00001010
?00001010
000011110
000011110
000011110
Polymorphisms: A, 1 + 2; B, 0 + 1; C, 0 + 3; D, 0 + 2.
SYSTEMATICS OF SABELLARIIDAE
251
Table 4. List of characters and states for resolving relationships within Sabellariidae
1. Opercular lobes: (0) completely fused (entire); (1) partially fused (with deep indentation in ventral margin; (2)
completely divided into two free lobes.
2. Operculum relative length (length from mouth to opercular papillae/maximum width): (0) longer than wide; (1)
similar or shorter than wide.
3. Operculum with distal end sloped posteriorly (truncated): (0) absent (distal disc perpendicular to longitudinal axis;
(1) present (distal disc oblique to longitudinal axis).
4. Opercular papillae: (0) at least twice longer than wide; (1) shorter.
5. Form of tentacular filaments: (0) simple (unbranched); (1) compound (branched).
6. Buccal flaps: (0) absent; (1) present.
7. Palps: (0) very short (shorter than half of the operculum length, without paleae); (1) short (shorter or similar in
length to operculum); (2) long (longer than operculum).
8. Median organ (cirrus) at the dorsal juntion of the lobes of the opercular stalk: (0) absent; (1) present.
9. Arrangement of outer paleae on opercular lobes: (0) in semicircles; (1) in spirals.
10. Outer opercular paleae angle of shaft and blades: (0) straight; (1) geniculate (blade and shafts forming a
conspicuous angle).
11. Blade of outer paleae: (0) flat; (1) concave or excavated; (2) cylindrical.
12. Lateral margins of outer opercular paleae: (0) smooth; (1) with large pointed denticles.
13. Distal margins of outer opercular paleae: (0) smooth; (1) denticulated.
14. Outer opercular paleae with a distal plume: (0) absent; (1) present.
15. Inner opercular paleae arrangement: (0) giving the appearance of 1 row; (1) giving the appearance of 2 rows.
16. Middle opercular paleae arrangement: (0) pointing inwards, to the centre of operculum; (1) pointing outwards,
outside operculum.
17. Middle opercular paleae angle of shaft and blades: (0) straight; (1) geniculate.
18. Blade of middle paleae: (0) flat; (1) concave, excavated; (2) convex.
19. Arrangement of inner paleae: (0) semicircles; (1) short line on the juntion of lobes; (2) line converging inner
margin of lobes; (3) short line on ventrum on inner margin of lobes.
20. Inner paleae angle of shaft and blades: (0) straight; (1) geniculate strongly geniculate.
21. Blade of inner paleae: (0) flat; (1) excavated/concave; (2) cylindrical, like spines; (3) strongly convex.
22. Nuchal spines or hooks: (0) absent; (1) present.
23. Tip shape of spines: (0) straight, spines; (1) bent, hooks.
24. Limbation of hooks: (0) absent; (1) present.
25. Limbation of hooks: (0) in the convex side of hooks; (1) in the concave side of hooks.
26. Number of pairs of cirri on neuropodia segment 1: (0) one; (1) two; (2) three.
27. Neurochaetae on segment one: (0) absent; (1) present.
28. Number of pairs of lateral lobes on segment 2: (0) one; (1) two; (2) three; (3) four.
29. Thoracic branchiae (segment 2): (0) absent; (1) present.
30. Number of parathoracic segment: (0) three; (1) four.
31. Parathoracic neurochaetae: (0) with lanceolate and capillaries; (1) only capillary chaetae; (2) only lanceolate.
opercular papillae, presence and number of neuropodial cirri in segment 1, and presence, number and
shape of lateral lobes between noto- and neuropodia
of segment 2. A total of 31 characters and their states
are listed in Table 4 and described in Appendix 2.
Feature scoring was made via the DELTA System
(DEscription Language for TAxonomy) (Dallwitz,
1980) to ensure consistency among generic taxonomic
descriptions (Table 5).
MOLECULAR
SEQUENCES
The DNA fragments selected for testing relationships
between Sabellariidae and other polychaetes were
three nuclear genes, namely 18S rRNA (1889 bp), 28S
rRNA (3986 bp), and elongation factor 1a (EF-1a,
1140 bp). All sequences were acquired from GenBank,
and they have been part of studies about deep annelid
relationships (e.g. Brown et al., 1999; Rousset et al.,
2004; Struck et al., 2007, 2008; Capa et al., 2011) and
also at family level (e.g. Struck, Halanych & Purschke, 2005; Kupriyanova, Macdonald & Rouse, 2006)
but it is the first time the present combination of taxa
and genes are included in analyses for resolving relationships of Sabellariidae and other polychaetes. In
some cases, sequences from two species of the same
genus have been merged and considered as one terminal (Table 6).
Sequences were aligned with MAFFT version 6
(Katoh, 2008) using the very slow, iterative refine-
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
252
M. CAPA ET AL.
Table 5. Character matrix of morphological features included for resolving relationships within Sabellariidae. ‘?’ stands
for unknown and ‘-’ for inapplicable states
Spionidae
Bathysabellaria neocaledoniens
Bathysabellaria spinifera
Gesaia elegans
Gesaia fossae
Gunnarea gaimardii
Idanthyrsus macropaleus
Idanthyrsus australiensis
Idanthyrsus sp. nov. 1
Idanthyrsus sp. nov. 2
Lygdamis indicus
Lygdamis augeneri
Lygdamis giardi
Lygdamis sp. nov.
Mariansabellaria norvegicus
Mariansabellaria harrisae
Neosabellaria cementum
Neosabellaria vitiensis
Neosabellaria uschakovi
Paraidanthyrsus quadricornis
Phalacrostemma cidarophillum
Phalacrostemma profunda
Phalacrostemma sp. nov.
Phragmatopoma caudata
Phragmatopoma californica
Sabellaria alveolata
Sabellaria ishikawai
Sabellaria issumiensis
Sabellaria sp. nov. 1
Sabellaria sp. nov. 2
Tetreres varians
Tetreres superbus
Tetreres robustus
1–10
11–20
21–30
31
-------0-0000001101
0101001101
2000001100
2000001100
1101100001
2010101100
2010101100
2011101?00
201?101100
2010101100
2010101100
2010101100
2011101100
2000002000
2000002000
0101100001
0101100001
0101100001
2101101001
2100011110
2100011110
2100011110
0001101001
0001101001
2101101101
2101101101
2101101101
2101101101
2101101101
1000002000
1000002000
1000002000
---------10000---00
10000---00
20000---10
20000---10
00100---01
01100---20
01100---20
01100---20
01100---20
00000---20
00000---20
00000---20
00000---20
20000---10
20000---10
1011111101
1011111101
1011111101
01100---01
20000---10
20000---10
20000---10
0011101201
0011101201
0010111101
0011111101
0011111101
0010111101
0010111101
00000---30
00000---30
00000---30
---0-1110--01001
110--01201
2110-10111
2110-20011
10---01110
2110-00210
2110-00210
2110-00110
2110-00310
2110-01211
2110-00211
2110-00211
2110-00211
210--00101
210--00001
10---01110
10---01110
10---0?110
0111001110
2111121111
2111121111
2111101111
30---01110
30---01110
10---01010
10---01010
110--01010
110--01010
110--01010
2110-01301
2110-01301
2110-01401
1
1
1
1
0
?
2
2
2
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
1
1
ment method recommended for sequences with multiple conserved domains and long gaps (E-INS-i) for
18S rRNA and 28S rRNA and the method recommended for sequences with general homology
(G-INS-i) from EF-1a, all with an offset value of 0.2.
PHYLOGENETIC
ANALYSES
Molecular and morphological data were analysed
independently and in combination. Merges of matrices were performed in Winclada (Nixon, 2002).
Maximum-parsimony heuristic search used 10 000
replicates of random taxon addition and the tree
bisection-reconnection (TBR) branch swapping algorithm, saving ten trees per replicate using TNT 1.1
(Goloboff, Farris & Nixon, 2008b). All characters were
given equal weight and multistate characters were
considered non-additive. Nodal support was estimated by 1000 jackknife replicates using TBR, in
TNT 1.1 (Goloboff et al., 2008b). New technology
searches, such as ratchet, drift, and tree fusing, were
implemented, isolated and in combination, using TNT
1.1 (Goloboff et al., 2008b) performing 100 repetitions
and hitting the most-parsimonious trees 20 times.
Tree metrics are abbreviated as follows: tree length
(TL), consistency index excluding parsimony noninformative characters in the data matrix (CI), and
retention index (RI). Support values are given on the
trees.
To reach a topology that better explains those characters with a better fit to the cladistic hypothesis, at
the expense of the more homoplasious ones, we have
implemented implied weighting (Goloboff, 1993, 1995,
Goloboff et al., 2008a). With this method a higher
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
SYSTEMATICS OF SABELLARIIDAE
253
Table 6. List of taxa included in the phylogenetic analyses for resolving sabellariid sister-group relationships and
accession numbers in GenBank
Lepidonotus sublevis/Lepidonotus sp.
Nereis vexillosa
Auchenoplax crinita
Pectinaria koreni/P. gouldi
Pista cristata
Polycirrus sp.
Polydora sp./P. giardi
Trochochaeta sp.
Poecilochaetus serpens
Sternaspis scutata
Cirriformia luxuriosa/C. tentaculata
Magelona sp.
Owenia fusiformis
Bispira porifera
Eudistylia vancouveri
Schizobranchia insignis
Hydroides ezoensis
Protula palliata/P. magnifica
Serpula watsoni
Idanthyrsus pennatus/I. australiensis
Gunnarea capensis
Sabellaria cementarium
18S
28S
EF 1-a
AY894301
DQ790083
DQ790077
DQ790091
AY611461
EU418858
AY611455
DQ790097
AY569652
AY532353
AY611456
AY611454
AB106256
HM800950
–
AY732222
EU184062
DQ317124
EU184057
HM800960
AY577892
AY732223
DQ790039
DQ790043
DQ790026
DQ790054
DQ790057
EU418866
DQ790059
DQ790070
EU418869
DQ790063
AY611443
AY611441
DQ790049
HM800989 & HM801006
DQ242574
AY732225
EU184077
DQ317151 & DQ318584
EU184068
AF185174 & AF185149
DQ318593 & EU256544
AY732226
DQ813370
DQ813377
DQ813352
DQ813388
DQ813391
–
DQ813393
DQ813408
–
DQ813401
DQ813355
–
AB003709
–
DQ813361
–
–
AB003713
–
–
–
DQ813395
weight is given to those characters with less
homoplasy, producing a much more resolved estimated consensus tree (Goloboff et al., 2008a). Results
using a range of concavities (values for k) have been
compared.
RESULTS
ANALYSES
AT FAMILY LEVEL
Parsimony analysis of the morphological dataset,
including 99 characters (Tables 1 and 2), 80 of which
were parsimonious-informative, resulted in 25 shortest trees (TL 177, CI 0.58, RI 0.74), where Sabellariidae was recovered as monophyletic (JK 100, Fig. 1A,
B) and supported by the presence of nuchal organs
located at the base of palps, presence of tentacular
filaments originating from segment 1, handles
without a handle or manubrium in all segments, and
presence of a ‘proventricle’ as unique synapomorphies
(Fig. 1B). Other homoplastic features that support
this clade are the presence of an operculum formed by
the head and two anterior segments (shared with
Pectinariidae), presence of a pair of peristomial palps
(shared with Spionida), presence of a building organ
(shared with Pectinariidae), presence of paleae in the
anterior segments (shared with Pectinariidae), uncini
without a rostrum or main fang (shared with Oweniidae, Pectinariidae, and Ampharetidae), segmental
branchiae inserted in the parapodia (as Pectinariidae), and intraovarian oogenesis (shared with
Polynoidae) (Fig. 1B). The sister-group relationships
were not resolved (Fig. 1A) although in 20 of the 25
trees Sabellariidae was closely related to Sabellida
and in the other five to Terebellida.
Available sequences of 7056 nucleotides (including
1931 parsimony-informative) resulted in a unique
tree (Fig. 1C) of 9215 steps (CI 0.54, RI 0.36) with
four main clades, a basal clade unexpectedly gathering Owenia and Pectinaria (JK 76), a second one with
Magelona, Sternaspis, and members of Terebelliformia (JK < 50), a third clade with members of
Sabellida (JK < 50) including a paraphyletic Sabellidae, and a fourth clade with a monophyletic and
well-supported sabellariid clade (JK 100) recovered as
sister-group to Spionida (JK < 50). The position of
Cirriformia (Cirratulidae) is surprisingly associated
with the outgroup, although weakly supported.
The combination of morphological and molecular
data consisted of 7155 characters, 5144 of which are
variable and 2011 parsimony-informative. Results
recovered two shortest trees (TL 9434, CI 0.54, RI
0.37; Fig. 1D) with the four clades also outlined in the
molecular topology, although with variable relationships between them. Terebellida is recovered as paraphyletic with Magelona positioned as sister to
Cirriformia and Pectinaria as sister to Owenia. Sabel-
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
254
M. CAPA ET AL.
Figure 1. Trees resulting from parsimony analyses of Sabellariidae and previously related taxa (including members of
Sabellida, Terebellida and Spionida). A, strict consensus after analyses based on 99 morphological features with jackknife
support values. B, first of 25 most-parsimonious trees (TL 177, CI 0.58, RI 0.74) after analyses of morphological data with
unambiguous changes marked on the topology. Numbers under nodes indicate jackknife values; black dots: synapomorphies, white dots: homoplastic character states. C, shortest tree (TL 9215, CI 0.54, RI 0.36) resulting from analysis of
partial 18S, 28S and EF-1a sequences with jackknife support values. D, strict consensus of two most-parsimonious trees
(TL 9434, CI 0.54 RI 0.37) of the combined dataset, with jackknife support values.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
SYSTEMATICS OF SABELLARIIDAE
255
lariidae was again retrieved as monophyletic (JK 100)
and sister to a clade with members of Spionida
(JK < 50), this being the sister clade to Sabellida
(JK < 50).
ANALYSES
WITHIN SABELLARIIDAE
Maximum-parsimony analyses of 32 sabellariid
species, with all the 12 genera represented by at least
the type species, and 31 characters, of which 29
were parsimony-informative, resulted in 249 mostparsimonious trees (TL 73, CI 0.55, RI 0.86, Fig. 2).
The trees were rooted with Spionidae as the outgroup,
according to previous analyses (Fig. 1D). The polytomy at the base of the consensus tree reflects the
large amount of homoplasy in the dataset for resolving bifurcating branching pattern (Fig. 2A), with only
few clades being outlined, Bathysabellaria (JK 70),
Phalacrostemma (JK 87), Mariansabellaria (JK < 50),
Tetreres (JK 78), and a clade (JK < 50) containing a
paraphyletic Lygdamis and Idanthyrsus (JK 52)
(Fig. 2). Sabellariinae and Lydaminae are not recovered as monophyletic. A dataset containing only the
informative characters for the type species of the 12
genera resulted in a similar topology, indicating that
the specific features are not the only ones responsible
for the overall homoplasy.
Implied weighting with concavity of k = 3 recovered
three most-parsimonious trees (TL 76, CI 0.55, RI
0.86; Fig. 3A), the relationship within Idanthyrsus
being the only difference between topologies. The
monophyly of most genera are suggested except for
Gesaia and Sabellaria. Two main clades are outlined
and they do not fully concur with the sabellariid
subfamilies. Clade A, supported by having a long
operculum (lobes longer than wide), contains the
genera Idanthyrsus, Lygdamis, Tetreres, Mariansabellaria, and Gesaia; Clade B is supported by the
absence of neurochaetae on segment 1 (on both sides
of the building organ) and contains Phalacrostemma,
Bathysabellaria, Paraidanthyrsus, Gunnarea, Sabellaria, Phragmatopoma, and Neosabellaria (Fig. 3A).
Within Clade A, a sister-group relationship between
Lygdamis and Idanthyrsus is supported by the presence of an oblique distal end of the operculum, compound tentacular filaments, flat blades of outer
paleae, and inner paleae arranged in a straight line
on the inner margin of the opercular lobes, from
dorsum to ventrum. Clade AI containing Gesaia,
Mariansabellaria, and Tetreres is the sister-group to
Clade AII (Lygdamis and Idanthyrsus) supported by
the presence of only capillary chaetae in the neuropodia of parathoracic segments. Mariansabellaria and
Tetreres are closely related based on the presence of
very long palps, a median organ that appears to be
absent or inconspicuous, and the absence of thoracic
Figure 2. Strict consensus of 429 most-parsimonious
trees after maximum-parsimony analysis of morphological
data of members of Sabellariidae rooted with Spionidae
(TL 73, CI 0.55, RI 0.86). Numbers under nodes indicate
jackknife support values.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
256
M. CAPA ET AL.
Figure 3. Trees resulting from parsimony analyses of morphological data of members of Sabellariidae and rooted with
Spionidae, implementing implied weighting. Unambiguous changes are marked on the topology; black dots: synapomorphies, white dots: homoplastic character states. A, strict consensus of three most-parsimonious trees (constant of concavity
k = 3); B, most-parsimonious tree (constant of concavity k = 4–6).
branchiae (in segment 2). The basal group of Clade B
is Phalacrostemma, sister-group to all other taxa, and
Phragmatopoma and Neosabellaria are recovered as a
derived clade and supported by several plesiomorphies. Paraidanthyrsus and Gunnarea, two monotypic
genera, are found to be closely related and sharing
two plesiomorphic features but both species present
several morphological differences, including different
paleae, level of fusion of the opercular lobes, length of
palps, and presence or absence of nuchal spines
(Fig. 3A).
Similar analyses with weighting concavities of
k = 4–6 recovered one identical most-parsimonious
tree (TL 77, CI 55, RI 0.86, Fig. 2C) where Clade B is
also recovered but the basal nodes of Sabellariidae
differ from previous hypotheses (Figs 3B, 4). Mariansabellaria and Tetreres form a basal clade, sister to
a clade with the rest of the terminals. Within this large
clade Gesaia and two clades are recovered, Clade B and
another containing Idanthyrsus and Lygdamis (Clade
AII). The monophyly of all genera is assessed except for
Gesaia, Sabellaria, and Lygdamis (Figs 3B, 4).
Synapomorphies defining each genus are documented in the diagnoses of the taxonomic account
below and two particular cases, Gesaia and Sabellaria, are discussed.
TAXONOMIC RESULTS
SABELLARIIDAE JOHNSTON, 1865
Diagnosis: Sabellariidae is characterized by some
autapomorphies such as the presence of nuchal
organs located at the base of palps, presence of tentacular filaments originating from segment 1; uncini
without a handle or manubrium and presence of a
‘proventricle’. In addition, a unique combination of
the following characters (homoplastic) defines the
group: the presence of an operculum formed by the
head and two anterior segments, presence of a pair of
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SYSTEMATICS OF SABELLARIIDAE
257
Figure 4. Schematic representation of sabellariid relationships based on maximum-parsimony analyses of the morphological data (constant of concavity k = 4–6) and stylized drawing of opercula from top view, modified from Kirtley (1994).
See text for further details.
peristomial palps, presence of a building organ, presence of paleae in the anterior segments, uncini
without a rostrum or main fang, segmental branchiae
inserted in the parapodia, and intraovarian
oogenesis.
Description: Body divided into four regions and consisting of an operculum (head and thorax), parathorax, abdomen, and cauda. Segmentation distinct.
Operculum formed by the fusion of the prostomium,
peristomium, and first two segments. Head appendages consisting of two peristomial palps; antennae
absent and a median organ that sometimes protrudes from operculum edge. Prostomial ocelli
present in some taxa. Nuchal organs located at base
of palps. Tentacular filaments originating from
segment 1 and arranged along ventral side of operculum. Building organ with horseshoe shape present
and located below mouth. Parapodial rami unequal.
Parapodial lobes, dorsal and ventral cirri absent.
Aciculae absent. Thoracic chaetae simple, capillaries
or straight, flat, and lanceolate chaetae. Abdominal
neurochaetae simple, capillaries or straight, flat,
and lanceolate chaetae. Abdominal notopodia with
uncini without a hood, rostrum, manubrium or basal
process but with a capitium, subrostrum, and subrostral process present. Capitium consists of two
lines of 6–9 teeth each. Pygidial cirri absent.
Parapodial branchiae present on most segments
along body. Circular muscles present and longitudinal ones grouped in bundles. Lateral organs absent.
Stomodaeum without buccal organ. Gut as a
straight tube, with a ‘proventricle’ in anterior
abdominal segments. Gular membrane absent. Head
kidney protonephridia and adult segmental organs,
metanephridia with podocytes and terminal monociliated cells. An anterior pair of excretory and posterior gonoducts present. Circulatory system closed;
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
258
M. CAPA ET AL.
heart body absent and blood without cells. Nervous
system subepidermal; brain without mushroom
bodies. Species studied to date are gonochoric,
broadcast spawners and with external fertilization.
Females with discrete ovaries in posterior segments
KEY
TO
where oogenesis takes place. Males with rounded
head sperm. Larvae planktotrophic and with prototroch, telotroch, and neurotroch ciliary bands
together with an apical turf. Tube made out of
mucus and sediment particles.
SABELLARIIDAE
GENERA
1. Opercular lobes completely fused........................................................................................................2
Opercular lobes partially fused ................................................................................................................ 4
Opercular lobes completely divided in two lobes..........................................................................................5
2. Outer row of opercular paleae with flattened blades .......................................................... Phalacrostemma
Outer row of opercular paleae with cylindrical blades..................................................................................3
3. Tentacular filaments simple, resembling opercular papillae. Four parathoracic segments...........Bathysabellaria
Tentacular filaments compound (numerous and arranged in transverse rows). Three parathoracic segments
......................................................................................................................................... Neosabellaria
4. Outer paleae with straight blades. Tentacular filaments simple, resembling opercular papillae. Palps longer than
operculum ................................................................................................................................... Tetreres
Outer paleae geniculate, with shaft and blades forming an angle. Tentacular filaments compound (numerous and
arranged in transverse rows). Palps shorter than half of length of operculum ...................................... Gunnarea
5. Opercular disc (distal end) clearly oblique to longitudinal margin (e.g. Fig. 8B), with paleae arranged in a dorsal
slope from lateral view...........................................................................................................................6
Opercular disc flat, with paleae arranged perpendicular to longitudinal axis (e.g. Fig. 8A) ................................. 7
6. Outer paleae with straight and flattened blades and smooth margins. Four parathoracic segments
...............................................................................................................................................Lygdamis
Outer paleae with straight and flattened blades with large and pointed denticles on margins. Three parathoracic
segments ............................................................................................................................... Idanthyrsus
7. Blade of inner paleae cylindrical, resembling spines...............................................................................8
Blades of inner paleae different (flat or concave) ....................................................................................... 10
8. Nuchal spines straight ................................................................................................ Mariansabellaria
Nuchal spines with bent tips (hooks).................................................................................................Gesaia
9. Buccal flaps present; outer paleae arranged in spirals........................................................Phragmatopoma
Buccal flaps absent; outer paleae arranged in semicircles ........................................................................... 10
10. Outer paleae with flat blades and smooth margins. Nuchal spines, if present, straight and without limbations
..............................................................................................................................................Sabellaria
Outer paleae with flat blades and large pointed denticles. Nuchal spines with bent tips (hooks) and limbation of convex
side................................................................................................................................Paraidanthyrsus
BATHYSABELLARIA LECHAPT & GRUET, 1993
Bathysabellaria Lechapt & Gruet, 1993: 243, figs 1–4;
Kirtley, 1994: 185.
Type species: Bathysabellaria neocaledoniensis
Lechapt & Gruet, 1993, by monotypy; collected from
deep water off New Caledonia.
Diagnosis: Bathysabellaria is the only group of sabellariid with the opercular lobes completely fused along
its length, thoracic branchiae absent, and neuropodia
of thoracic segments bearing only capillary chaetae.
Description: Operculum with lobes completely fused
and with distal end (disc) flat and perpendicular to
longitudinal axis. Opercular papillae numerous and
digitiform. Outer paleae numerous, arranged in semicircles; shaft and blade slightly geniculate (forming
an angle), blades faintly excavated, with smooth
margins, except when distal tips frayed or broken.
Inner opercular paleae arranged in semicircles, giving
the appearance of one row, straight, slightly excavated, with smooth margins. One pair of nuchal
spines, only slightly curved distally. Three or four
simple (unbranched) tentacular filaments present;
buccal flaps absent. Palps similar in length to operculum. With conspicuous median organ present. Neuropodia of segment 1 with one cirri on each side of
building organ and capillary chaetae. Segment 2 with
one pair of triangular-shaped lateral lobes. Thoracic
branchiae absent. Four parathoracic segments. Parathoracic notochaetae lanceolate and capillaries alternating; neurochaetae only capillaries.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
SYSTEMATICS OF SABELLARIIDAE
Remarks: Lechapt & Gruet (1993) erected this genus
based on differences in the shape of palps and nuchal
spines compared with other genera having four
parathoracic segments. They also mentioned the presence of only capillaries in the neuropodia of parathoracic segments as a diagnostic feature although it was
not further mentioned in redescription of the genus
and description of a new species by Lechapt & Kirtley
(1998). Note that the presence of capillary chaetae in
parathoracic neuropodia is also present in Mariansabellaria (Kirtley, 1994) and at least some Gesaia
species (Fauvel, 1911). Thoracic branchiae are absent
(Lechapt & Kirtley, 1998; Figs 7, 8) although Kirtley
(1994: 185) mentioned their presence and a conspicuous median organ projects from the ventral side of the
opercular lobes, although it was reported as absent in
the genus (Kirtley, 1994: 185). The genus contains two
species only found in deep water (more than 450 m)
off New Caledonia (Lechapt & Kirtley, 1998).
GESAIA KIRTLEY, 1994
Gesaia Kirtley, 1994: 166.
Type species: Phalacrostemma elegans Hartman &
Fauchald, 1971: 152 (not Fauvel, 1911) designated by
Kirtley (1994). Type locality: Bermuda rise, WHOI
Chain Sta. 84, 36 °24.4′N, 67 °56′W; 4749 m.
Diagnosis: Monophyly of this genus has not been
assessed but the species are characterized by: a long
operculum divided into two lobes, with long opercular
papillae, simple tentacular filaments, nuchal hooks
with no limbations, branchiae on segment 2, median
organ present, absence of nuchal flaps, and the presence of capillary chaetae in parathoracic neuropodia.
Description: Operculum longer than wide, completely
divided into two free lobes and distal disc perpendicular to operculum; few (4–8 pairs) long and conical
opercular papillae. Three to five simple (unbranched)
tentacular filaments along margins of buccal cavity.
Buccal flaps absent. Palps shorter or similar in length
to operculum with obvious groove. Conspicuous
median organ (cirrus) at dorsal junction of lobes.
Outer paleae arranged in semicircles with straight,
cylindrical, smooth blades (with ornamented thecae
but no denticles). Inner opercular paleae arranged in
a single row, like a straight line, on the dorsal half of
the inner margin of lobes, with straight cylindrical
and smooth paleae. One pair of nuchal spines with
bent tips (hooks), without limbation. Neuropodia of
segment 1 with two pairs of cirri (only described in
type species) on both sides of building organ and
without neurochaetae. Two lateral lobes in segment 2
(only described in type species). Thoracic branchiae
259
present. Four pairs of parathoracic segments. Parathoracic neurochaetae only capillaries.
Remarks: The differences between Gesaia and other
sabellariids were not clearly stated when this genus
was erected (Kirtley, 1994) and the monophyly of the
genus and relationships with other sabellariids have
not been assessed after phylogenetic analyses. Gesaia
shares with Bathysabellaria, Mariansabellaria, Phalacrostemma, and Tetreres the occurrence of a long
operculum with a perpendicular disc, long opercular
papillae, and single tentacular filaments but
only Gesaia, Mariansabellaria, and Phalacrostemma
present an operculum with completely separated
lobes. Gesaia, however, is unique among these taxa,
based on the combination of the following features:
presence of nuchal hooks with no limbations, branchiae on segment 2, conspicuous median organ, and
absence of nuchal flaps. Gesaia currently contains
eight species distinguished by the ornamentation of
the thecae on the outer paleae. It has been reported
from all the major oceans, occurring only in deep
water (Kirtley, 1994).
GUNNAREA JOHANSSON, 1927
Gunnarea Johansson, 1927: 99; Kirtley, 1994: 41–42.
Type species: Hermella gaimardi (Quatrefages, 1848).
From Table Bay, Cape Town, South Africa.
Diagnosis: Gunnarea is characterized by three
homoplastic features: the presence of partially
divided opercular lobes, very short palps, and absence
of nuchal spines. An autapomorphy of this species is
the presence of a broad lateral lobe on segment 2,
with crenulated margins giving the appearance of
being subdivided.
Description: Operculum with lobes partially fused
(with deep indentation in ventral margin) and distal
disc perpendicular to longitudinal axis. Numerous
small and rounded opercular papillae on its perimeter. Outer paleae arranged in semicircles; geniculate,
with flat blades, smooth lateral margins, and distal
tooth. Inner paleae arranged in semicircles, as a
single row, strongly geniculate, with flat blades.
Nuchal spines absent. Palps shorter than half of the
operculum. Tentacular filaments compound. Buccal
flaps absent. Conspicuous median organ absent. Neuropodia of segment 1 with one cirrus on each side
of building organ and capillary neurochaetae. Two
triangular lobes between noto- and neuropodia of
segment 2, ventral one subdivided or with crenulated
margins. Thoracic branchiae present. Three parathoracic segments. Parathoracic notochaetae lanceolate
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M. CAPA ET AL.
Figure 5. Stylized drawing of a sabellariid indicating the body regions and some of the morphological features described
in Appendix 2: A, dorsal view; B, ventral view.
Figure 6. Photographs of Idathyrsus australiensis alive: A, complete specimen, dorsal view; B, anterior end, dorsal view;
C, anterior end, central view. Abbreviations: ab, abdomen; b, branchia; ca, cauda; g, gut; ip, inner paleae; nh, nuchal
hooks; p, paleae; o, operculum; op, outer paleae; tf, tentacular filaments; pa, parathorax.
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SYSTEMATICS OF SABELLARIIDAE
and capillaries alternating; neurochaetae similar
but smaller. Abdominal branchiae present on most
abdominal segments.
Remarks: Gunnarea shares some characteristics
with Paraidanthyrsus, Neosabellaria, Sabellaria, and
Phragmatopoma as they all have three parathoracic
segments, branching tentacular filaments, outer
paleae with distal denticles and innermost row of
paleae geniculate. Moreover, Gunnarea is recovered
as sister to Paraidanthyrsus, supported by the
absence of a conspicuous median organ, a feature that
has not been checked in Paraidanthyrsus for the
present study. Gunnarea is monospecific.
IDANTHYRSUS KINBERG, 1867
Idanthyrsus Kinberg, 1867: 350; Kirtley, 1994: 84.
Type species: Idanthyrsus macropaleus Schmarda,
1861 (after Kirtley, 1994). Type locality: Valparaiso
Harbour, Chile.
Diagnosis: Idanthyrsus autapomorphies are outer
paleae straight (no angle between shaft and blade),
flat, and with pointed denticles along lateral margins
and distal tips, and, if confirmed in all species of the
genus, only fine lanceolate chaetae on neuropodia of
parathoracic segments. Unlike other related genera
they have three thoracic segments.
Description: Operculum longer than wide with lobes
completely divided and distal end sloping posteriorly
(oblique to longitudinal axis) and operculum papillae
varying in number and size depending on species.
Outer paleae arranged in semicircles with straight
and flat blades and lateral and distal margins appearing sharply denticulated. Inner opercular chaetae
arranged in one row along the inner margin of opercular lobes, with straight and cylindrical blades. One
or two pairs of nuchal spines with bent tips (hooks)
with or without limbations on the concave margin.
Palps similar in length to the operculum. Median
organ conspicuous with eyespots on its sides (on
specimens examined). Tentacular filaments compound
(branching); buccal flaps absent. Neuropodia of
segment 1 achaetous, with conical cirri (one in all
species examined). Segment 2 with two to four triangular lateral lobes. Three parathoracic segments with
notochaetae consisting of lanceolate and capillary
chaetae and only thin lanceolate neurochaetae (on
specimens examined). Branchiae from segment 2
diminishing in size on posterior abdominal segments.
261
Remarks: All species share the presence of straight,
flat, denticulated outer paleae, unique among Sabellariidae. If the presence of only lanceolate chaetae in
neuropodia of parathoracic segments is confirmed in
all species of the genus this would be another autapomorphy for the group. In the revision of the family,
Kirtley considered Lygdamis and Idanthyrsus as
belonging to different groups (subfamilies) due to the
difference in number of parathoracic segments.
However, analyses show that these genera are closely
related as they both share the overall shape of operculum with separated and long lobes with oblique
distal end, inner paleae arranged in a single line
along the inner margin of lobes, and presence of
nuchal hooks. Besides the number of parathoracic
segments in these two groups, they differ in the shape
of outer paleae (smooth in Lygdamis) and the presence on capillary chaetae alternating with the
lanceolate neuropodia of parathoracic segments of
Lygdamis and the latter only being present in Idanthyrsus. There are, at the time of writing, 19 species
described in the genus (Kirtley, 1994; Nishi & Kirtley,
1999). Some features, such as the presence of limbation on nuchal hooks, presence of neurochaetae on
segment 1, number of lateral lobes on segment 2,
or presence of eyespots near median organ, vary
between species. They are generally found in shallow
water in the tropics or temperate waters as isolated
individuals but some species also inhabit boreal and
deep water domains and some aggregations of individuals have also been found (e.g. Kirtley, 1994; Nishi
& Kirtley, 1999).
LYGDAMIS KINBERG, 1867
Lygdamis Kirtley, 1994: 116.
Type species: Lygdamis indicus Kinberg, 1867
by monotypy. Type locality: Bangka Straits, Java,
Indonesia.
Diagnosis: The monophyly of the genus has not being
assessed in the phylogenetic analyses although all
species assigned to Lygdamis have an operculum with
separated lobes and distal end sloped posteriorly,
opercular paleae straight and smooth, the outer ones
being flattened and the inner ones cylindrical, resembling spines.
Description: Operculum longer than wide with lobes
completely separated and distal end sloped posteriorly (oblique to longitudinal axis). Numerous opercular papillae around edge of lobes. Outer paleae
arranged in semicircles, straight, with flat blades,
lateral and distal margins smooth. Inner opercular
chaetae arranged in one row along the inner margin
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SYSTEMATICS OF SABELLARIIDAE
263
Figure 7. Stylized drawings showing morphological features described in the text: A, arrangement of opercular paleae
of Phragmatopoma sp., upper view; B, same of Gunnarea sp.; C, same of Paraidanthyrsus sp.; D, anterior end, ventral
view, Gesaia sp.; E, same, Sabellaria sp.; F, same of Phalacrostemma sp.; G, transverse section of the operculum showing
arrangement of opercular paleae with inner row subdivided into two rows, both directed inwards; H, same, showing
middle paleae directed outwards; I, outer opercular paleae in spiral arrangement; J, parts of opercular paleae; K, straight,
flat blade with distal and lateral denticles, typical of outer paleae of Idanthyrsus spp.; L, geniculate, concave, and smooth
edges of blade, typical of some middle paleae of Sabellaria spp.; M, straight, flat blade with distal denticles and smooth
lateral edges, typical of outer paleae of Sabellaria spp.; N, geniculate, concave, and smooth edges of blade, typical of inner
paleae of Sabellaria spp.; O, straight, cylindrical, and smooth blade, typical of inner paleae of Idanthyrsus spp.; P,
straight, flat, and smooth blade, typical or inner and outer paleae of Tetreres spp.; Q, nuchal spine; R, nuchal spine with
bent tip (hook), without limbation; S, nuchal spine with bent tip (hook), with limbation. Abbreviations: b, blade; bo,
building organ; ip; inner paleae; mp, mid paleae; pa, palps; op, outer paleae; opa, opercular papillae; s, shaft; tf, tentacular
filaments.
䉳
of opercular lobes with straight and cylindrical or
slightly flattened blades. One pair of nuchal spines
with bent tips (hooks) and without limbations. Palps
similar in size to the operculum. Median organ elongate at the dorsal junction of the lobes of the opercular stalk. Tentacular filaments compound (branching);
buccal flaps absent. Neuropodia of segment 1 with a
conical cirrus, with or without capillary chaetae.
Segment 2 with three triangular-shaped lobes
between noto- and neuropodia, in some species
rounded and small. Four parathoracic segments with
notochaetae consisting of lanceolate and capillary
chaetae and neurochaetae similar in shape but
smaller. Branchiae from segment 2 to mid abdominal
segments.
Remarks: According to our analyses this genus is not
monophyletic but this is probably an artefact of splitting the paleal types into different categories according to their ornamentation, shape, and angle. All
Lygdamis species have an operculum with a distal
end sloped posteriorly and opercular paleae straight
and smooth, the outer ones being flattened and the
inner ones cylindrical. Some variation included the
shape of lateral lobes of segment 2, the presence or
absence of neurochaetae in segment 2, and the
presence of eyes near the median organ. This genus
shares several features with Idanthyrsus but differences are the number of parathoracic segments, the
shape of outer paleae (strongly denticulated in Idanthyrsus and smooth in Lygdamis), and the presence of
capillary chaetae alternating with the lanceolate
chaetae on the neuropodia of parathoracic segments
of Lygdamis, the latter type being the only ones
present in Idanthyrsus species. The genus is known
from 16 species (Kirtley, 1994).
MARIANSABELLARIA KIRTLEY, 1994
Mariansabellaria Kirtley, 1994: 136–137.
Type species: Phalacrostemma norvegicum Strømgren,
1971, by monotypy. Type locality: Bindalsfjiorden,
Norway.
Diagnosis: Mariansabellaria is characterized by the
following combination of features: presence of straight
nuchal spines, very long palps, branchiae on the
second segments, and the absence of a median organ.
Members of the genus have a conspicuous ventral
glandular area on the parathoracic segments (Kirtley,
1994), a potential autapomorphy for the group.
Description: Operculum longer than wide with lobes
completely divided into two symmetrical halves and
distal disc perpendicular to longitudinal axis; long
conical papillae around its perimeter. Outer paleae,
arranged in semicircles, smooth, straight and cylindrical or slightly flattened. Inner paleae, few in
number, arranged in a short single row near the
dorsal junction of margin of lobes, straight. One or
more pairs of nuchal spines. Palps grooved and longer
than operculum. Tentacular filaments arranged in
single rows. Buccal flaps absent. Conspicuous median
organ absent. Neuropodia of segment 1 with one
cirrus on each side of building organ; capillary neurochaetae absent. One or two triangular lateral lobes
on segment 2. Thoracic branchiae absent. Four parathoracic segments. Parathoracic notochaetae lanceolate and capillaries alternating; neurochaetae only
capillaries. Abdominal branchiae present on anterior
abdominal segments but not on posterior ones.
Remarks: Mariansabellaria was characterized by the
presence of conspicuous ventral glandular areas on
the parathoracic segments (Kirtley, 1994) but this
feature is not described in all species of the genus or
in other sabellariids and therefore needs confirmation. Mariansabellaria shares with Bathysabellaria,
Gesaia, Phalacrostemma, and Tetreres the occurrence
of a long operculum with perpendicular disc, long
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Figure 8. Photographs of preserved specimens: A, operculum and anterior segments, lateral view, Sabellaria sp. nov. 2;
B, operculum and anterior segments, lateral view, Lygdamis giardi; C, detail of opercular papillae, Lygdamis indicus; D,
head and thoracic appendices, Idanthyrsus australiensis; E, head and thoracic appendices, Sabellaria sp. nov. 2; F,
operculum and anterior segments, ventral view, Phalacrostemma sp. nov.; G, head and thoracic appendices, L. giardi; H,
head and thoracic appendices, L. giardi; I, median organ with lateral ocelli, I. australiensis; J, head and thoracic appendices,
Bathysabellaria spinifera; K, operculum and paleae, dorsal view, Tetreres robustus; L, operculum and thoracic segments,
lateral view, B. spinifera; M, operculum and anterior segments, lateral view, T. robustus. Abbreviations: b, branchia; b2,
branchia segment 2; bo, building organ; cn 1, cirrus neuropodia segment 1; chn1, chaetae neuropodium segment 1; dap,
dorsal papilla; es, eyespots; ip; inner paleae; li, lips; mo, mouth; mor; median organ; mr, median ridge; ns, nuchal spines
(hooks); op, outer paleae; opa, opercular papillae; p, paleae; pa, palp; pl, oral plates; tf, tentacular filaments.
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SYSTEMATICS OF SABELLARIIDAE
265
Figure 9. Photographs of preserved specimens: A, nuchal spines (hooks), I. australiensis; B, thoracic segments with
lateral lobes, I. australiensis; C, thoracic segments with lateral lobes, L. giardi; D, abdominal segments with ‘proventricle’,
dorsal view, Sabellaria sp. nov. 2; E, abdominal segments with ‘proventricle’, ventral view, Sabellaria sp. nov. 2; F,
posterior abdominal segments and cauda, Idanthyrsus australiensis. Abbreviations: b, branchia; ca, cauda; ll, lateral
lobes; nh, nuchal spines (hooks); pv, proventricle.
opercular papillae and single tentacular filaments
without buccal flaps. However, only Gesaia, Mariansabellaria, and Tetreres have parathoracic neuropodia with just capillary chaetae. Close relationships
between Mariansabellaria and Tetreres are based on
the size of palps, exceeding the length of the operculum. Mariansabellaria is distinguished of these taxa
by the presence of straight nuchal spines. This genus
is represented by four species from the west coast of
North and South America and also Norway, at depths
ranging from 180 to 2000 m.
NEOSABELLARIA KIRTLEY, 1994
Neosabellaria Kirtley, 1994: 16; Nishi et al., 2010: 4.
Type species: Sabellaria cementarium Moore, 1906.
From Admiralty Inlet, vininity of Port Townsend,
Alaska, USA.
Diagnosis: Neosabellaria is characterized by a unique
combination of characters: the presence of short
palps, often not reaching half the length of the operculum and the presence of excavated outer paleae.
Description: Operculum length similar to maximum
width, with lobes completely fused, although shallow
mid ventral indentation sometimes present at proximal end; distal end flat and perpendicular to longitudinal axis. Numerous conical and small opercular
papillae. Outer paleae numerous, arranged in semicircles, geniculate, with excavated blades, smooth
lateral markings and denticulated distal margin with
a midline plume. Inner opercular paleae giving the
appearance of two rows. Middle paleae geniculate
with excavated, smooth blades and pointed tips
directed outwards, some species with rounded tipped
blades also present. Innermost paleae strongly
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M. CAPA ET AL.
Figure 10. Scanning electron micrographs: A, arrangement of paleae in two rows, Idanthyrsus australiensis; B, paleae
giving the appearance of being arranged in three rows with the mid and inner row directed in opposite directions,
Sabellaria sp. nov. 2; C, paleae with cylindrical and straight blades, Phalacrostemma sp. nov.; D, paleae with flat,
straight and smooth edges blades, Lygdamis giardi; E, paleae with flat, straight blades and denticulated margins,
Idanthyrsus sp. nov. 1; F, paleae with flat, straight blades and denticulated margins, Idanthyrsus australiensis; G,
paleae with flat, straight blades with smooth lateral margings but denticulated distal margins, Sabellaria sp. nov. 2;
H, geniculate and concave paleae, Sabellaria sp. nov. 2; I, bent nuchal spines (hooks) without limbation I. australiensis;
J, bent nuchal spines (hooks) without limbation Phalacrostemma sp.; K, parathoracic notopodia with lanceolate and
capillary chaetae, I. australiensis; L, parathoracic neuropodia with lanceolate chaetae, of two sizes, I. australiensis; M,
mid abdominal neurochaetae, I. australiensis; N, abdominal uncini with double rows of teeth, I. australiensis.
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SYSTEMATICS OF SABELLARIIDAE
geniculated, with short and concave ones directed
inwards. Nuchal spines absent. Compound tentacular
filaments arranged in series of rows; buccal flaps
absent. Palps shorter than half length of operculum.
Median organ at dorsal junction of lobes of opercular
stalk present but small. Neuropodia of segment 1
with one pair of cirri and capillary chaetae, at least in
specimens examined. Segment 2 with two pairs of
triangular-shaped lobes between noto- and neuropodia. Thoracic branchiae present. Three parathoracic
segments. Parathoracic notochaetae lanceolate and
capillaries alternating; neurochaetae similar in shape
but smaller. Abdominal branchiae absent in posterior
segments.
Remarks: Kirtley (1994) erected the genus and transferred into it previously described species of Sabellaria based on the complete fusion of the opercular
stalk and crown whereas in Sabellaria they are only
partially fused. Neosabellaria, Sabellaria, and Phragmatopoma are grouped based on the appearance of
having three rows of opercular paleae, together with
other character-states that are homoplastic, such as
the presence of short opercular papillae, branching
tentacular filaments, outer paleae geniculate and
with distal denticles, innermost paleae geniculate and
concave, and three parathoracic segments. Neosabellaria and Sabellaria resemble each other in the type
of paleae in the middle row, as they are slightly
geniculate, with excavated blades, and pointing outwards while in Phragmatopoma they are strongly
geniculate, with convex blades and pointing inwards.
However, according to our analyses Phragmatopoma
seems to be more closely related to Neosabellaria, as
both genera share an operculum with lobes completely fused. A key to the species of Neosabellaria is
given by Bailey-Brock et al. (2007) and seven species
are currently known, restricted to the Indo-Pacific.
PARAIDANTHYRSUS KIRTLEY, 1994
Paraidanthyrsus Kirtley, 1994: 80.
Type species: Hermella quadricornis Schmarda, 1861,
from New Zealand.
Diagnosis: Paraidanthyrsus is characterized by the
presence of geniculate outer paleae with flat blades
and nuchal hooks with a limbation on the convex side.
Description: Operculum with length similar to width,
lobes completely divided into two free lobes and distal
disc perpendicular to longitudinal axis and numerous
short papillae around its perimeter. Outer paleae
arranged in semicircles, geniculate, with flat blades
and margins with long and pointed denticles and
267
without a distal plume. Inner paleae in a single row,
arranged in semicircles, strongly geniculate, with flat
blades and tips directed inwards. Two or three pairs
of nuchal hooks with bent tips (hooks) and limbation
of the convex side. Tentacular filaments compound
(branching) arranged in more than eight rows; buccal
flaps absent. Palps similar in length to operculum.
Conspicuous median organ at the dorsal juntion of
the lobes of the opercular stalk absent. Segment 1
with a small and rounded cirrus on both sides of
building organ and with capillary chaetae on the
neuropodia. Segment 2 with two triangular-shaped
lateral lobes. Thoracic branchiae present. Three parathoracic segments. Parathoracic notochaetae lanceolate and capillaries, similar in shape but smaller
in neuropodia. Abdominal branchiae absent in posterior abdominal segments.
Remarks: In his revision of the Sabellariidae, Kirtley
(1994) suggested that this monospecific genus was
probably related to Idanthyrsus based on the presence
of straight, flattened, denticulated paleae. In any of
the resulting topologies Idanthyrsus and Paraidanthyrsus are recovered as sister taxa or not even in
the same clades, and differently Paraidanthyrsus
is nested within the apomorphic sabellariid as the
sister-group to Gunnarea and basal to a clade formed
by Gunnarea, Neosabellaria, Phragmatopoma, and
Sabellaria, and supported by the presence of geniculate inner paleae.
PHALACROSTEMMA MARENZELLER, 1895
Phalacrostemma Marenzeller, 1895: 191; Kirtley,
1994: 147.
Type species: Phalacrostemma cidariophilum Marenzeller, 1895, by monotypy. Type locality: off Pelagossa
Island, Adriatic Sea.
Diagnosis: Buccal flaps present, arrangement of outer
paleae in a spiral on each lobe, in most species.
Description: Opercular width similar to length, operculum completely divided into two free lobes and
distal disc perpendicular to operculum; eight to ten
pairs of long and conical opercular papillae conical
around lobes. Few simple (unbranched) tentacular
filaments along margins of buccal cavity, absent in
some species. Buccal flaps absent or present. Palps
similar in length to operculum. Small median organ
at dorsal junction of lobes. Outer paleae arranged in
spirals with straight, cylindrical, smooth blades (with
ornamented thecae but no denticles). Few (2–8) pairs
of inner opercular paleae present, arranged in a
straight line on the dorsal half of the inner margin of
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lobes, straight, cylindrical or slightly flattened blades
and smooth margins. Two to five pairs of nuchal
spines with bent tips (hooks) present and limbation in
the concave margin. Neuropodia of segment 1 with
one to three cirri on both sides of building organ and
capillary neurochaetae. Segment 2 with two digitiform lateral lobes between noto- and neuropodia. Thoracic branchiae present. Four pairs of parathoracic
segments. Parathoracic notopodia with lanceolate and
capillaries alternating, neuropodia with capillaries
and fine lanceolate chaetae. Abdominal dorsal branchiae absent in posterior segments.
Remarks: Monophyly of Phalacrostemma is supported
by the arrangement of outer paleae in a spiral and the
presence of buccal flaps present in most species. Even
though this genus shares some features such as the
presence of a long operculum with perpendicular disc,
long opercular papillae and single tentacular filaments with Bathysabellaria, Gesaia, Mariansabellaria, and Tetreres, it has been recovered in the base
of clade B after homoplasy weighting analyses supported by the length of the operculum and the presence of neurochaetae in segment 1. Phalacrostemma
is a cosmopolitan group represented by ten species
that live solely or in aggregations at different depths.
Some of these species either lack tentacular filaments
(e.g. Hartman, 1944) or have only a few present,
raising questions as to how they feed. Marenzeller
(1904) suggested that the type species feeds by using
its simple ciliated tentacular filaments on the anterior ventral margins of the operculum together with
the opercular papillae.
PHRAGMATOPOMA MÖRCH, 1863
Phragmatopoma Mörch, 1863: 442.
Type species: Phragmatopoma caudata Mörch, 1863.
Type locality: West Indies.
with paleae strongly geniculate with convex blades
and pointed tips directed inwards, middle paleae
almost covering innermost paleae. Nuchal spines
absent. Compound (branching) tentacular filaments
arranged in series of rows; buccal flaps absent. Palps
similar in length to operculum. Conspicuous median
organ absent. Neuropodia of segment 1 with one
conical cirri on both sides of building organ and
capillary chaetae. Segment 2 with two pairs of
triangular-shaped lateral lobes. Thoracic branchiae
present. Three parathoracic segments. Parathoracic
notochaetae lanceolate and capillaries alternating;
neurochaetae similar in shape but smaller. Abdominal
branchiae present to posterior segments.
Remarks: Phragmatopoma, Neosabellaria, and Sabellaria share the arrangement of inner paleae in two
concentric rows. Phragmatopoma and Neosabellaria
are recovered as sister groups due to the complete
fusion of operculum lobes. There are four recognized
species in the genus, all forming large colonies and
extensive reefs in the intertidal and shallow water
areas (with the exception of P. californica found up to
200 m) with an amphiamerican distribution (Kirtley,
1994). The type of P. caudata seems to be missing and
the last person to see this material was Ehlers in
1901 (Kirtley, 1994). Even though no type exists,
Kirtley (1994) synonymizes several species with this
type species which may be premature given the original brief description.
SABELLARIA LAMARCK, 1812
Sabellaria Kirtley, 1994: 45–46; Nishi et al., 2010: 7.
Type species: Sabella alveolata Linnaeus, 1767, by
monotypy, collected from France?
Diagnosis: Inner paleae arranged in two concentric
rows, pointing inwards to the centre of the opercular
disc, with strongly geniculate and convex blades,
middle paleae almost covering innermost ones.
Diagnosis: Although the monophyly of Sabellaria has
not being assessed in our analyses, all species share
the presence of an operculum completely divided into
two symmetrical lobes, inner paleae arranged in two
symmetrical, semicircular rows (with mid and
inner paleae), and the presence of three parathoracic
segments.
Description: Operculum longer than wide, with lobes
completely fused to each other, although shallow mid
ventral indentation sometimes present in proximal
end. Distal disc flat and perpendicular to longitudinal
axis. Numerous digitiform and long opercular papillae around its perimeter. Outer paleae numerous,
arranged in semicircles; geniculate, with flat blades,
smooth lateral margins, distal denticles, and a
midline plume. Inner opercular paleae giving the
appearance of two concentric rows, directed inwards,
Description: Operculum length similar to maximum
width, completely divided into two symmetrical lobes;
distal disc flat and perpendicular to longitudinal axis.
Numerous conical opercular papillae around operculum. Outer paleae numerous, arranged in semicircles;
geniculate, with flat blades, smooth lateral edges, and
smooth or denticulated distal margin and, sometimes,
a midline plume. Inner opercular paleae of various
shapes, giving the appearance of two rows arranged
in two concentric rows. Middle paleae strongly gen-
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SYSTEMATICS OF SABELLARIIDAE
iculate with excavated blades and smooth margins,
pointing outwards; innermost paleae strongly geniculated, with short concave blades and smooth margins,
directed inwards. Nuchal spines, when present, as
3–6 pairs of straight spines. Compound (branching)
tentacular filaments arranged in series of rows;
buccal flaps absent. Palps similar or shorter than
operculum. Median organ at the dorsal juntion of the
lobes of the opercular stalk conspicuous in some
species and small or absent in others, with eyespots
on its lateral margins in the species examined. Neuropodia of segment 1 with one pair of conical cirri and
capillary chaetae. Segment 2 with one triangularshaped lobe on both sides of building organ. Thoracic
branchiae present. Three parathoracic segments. Parathoracic notochaetae lanceolate and capillaries alternating; neurochaetae similar in shape but smaller.
Remarks: Nishi et al. (2010) have recently described a
novel species of Sabellaria and provide a table with
the chaetal characteristics of the 35 species from
around the world and highlights the variability of
certain features within the genus, such as the different geometry and superficial ornamentation of the
paleal thecae (especially in the middle row), the presence or absence of a plume or median tooth in the
outer paleae, and the presence or absence of nuchal
spines. The large intraspecific variation regarding
these features together with the inconsistency of the
presence or absence of a median organ among Sabellaria species (Kirtley, 1994) compromise the assessment of the monophyly of Sabellaria. Nevertheless all
species of Sabellaria share the presence of an operculum completely divided into two symmetrical lobes,
inner paleae arranged in two symmetrical, semicircular rows (with mid and inner paleae), and the presence of three parathoracic segments. Branchiae on
second segment (thoracic) are present (contrary to the
diagnosis of Nishi et al., 2010). Sabellaria shares with
Phragmatopoma and Neosabellaria the arrangement
of inner paleae in two concentric rows. All species
within the genus are gregarious, capable of forming
large colonies and reefs along shores where suitable
hydrodynamic and sedimentary conditions occur
(Kirtley, 1994).
TETRERES CAULLERY, 1913
Tetreres Kirtley, 1994: 188.
Type species: Hermella varians Treadwell, 1901, designation by Kirtley, 1994. Type locality: Mayagüez
Harbour, Puerto Rico.
Diagnosis: The autapomorphy that defines the group
is the arrangement of the inner paleae in a short
ventral line on each inner margin of the opercular
269
lobes. Nuchal hooks are large and with broadened
shafts, also unique among sabellariids.
Description: Operculum longer than wide with lobes
partially fused to each other (with deep indentation
on ventral margin) and distal disc perpendicular to
longitudinal axis, with large conical papillae around
its perimeter. Outer paleae arranged in semicircles,
straight, with flat blades and smooth margins. Inner
paleae few in number, arranged in a single straight
row on ventral side of operculum, with straight, flattened blades. One pair of large nuchal spines, with
bent tips (hooks); without limbations or enlarged
shaft. Palps wrinkled and deeply grooved, longer than
operculum. Tentacular filaments arranged in single
rows. Buccal flaps absent. Small median organ at the
dorsal junction of the lobes of the opercular stalk.
Neuropodia of segment 1 with one cirrus on each side
of building organ; capillary neurochaetae present.
Four long, tapering lateral lobes on segment 2. Thoracic branchiae absent. Four parathoracic segments.
Parathoracic notochaetae lanceolate and capillaries
alternating, similar in shape but smaller in neuropodia. Abdominal branchiae present, diminishing in size
posteriorly.
Remarks: Tetreres shares with Bathysabellaria,
Gesaia, and Mariansabellaria the occurrence of a long
operculum with perpendicular distal disc, long
opercular papillae, and single tentacular filaments
without buccal flaps. Tetreres is unique amongst these
genera in having an operculum partially fused along
its length, the arrangement of the inner paleae on the
ventral-inner side of the opercular lobes and the presence of four lateral lobes (at least in the specimens
examined). There are ten species within the group
reported from mostly deep waters in the Atlantic,
Pacific, and Southern Oceans (Kirtley, 1994).
DISCUSSION
OF SABELLARIIDAE
POSITION
The monophyly of Sabellariidae, even before tested in
a phylogenetic framework, has never been questioned
as the group presents several morphological and
developmental features unique and common to all of
its members. However, its position among the annelid
tree and the relationships within the family are still
far from being understood. For our analyses we
included available DNA information of common
markers used for resolving deep-level relationships
and in combination with a broad range of morphological data. Results suggest that Sabellariidae is
closely related to Spionida, although support for this
sister-group relationship is weak. Other authors have
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
270
M. CAPA ET AL.
also reached similar conclusions when combining
morphological and molecular data regardless of the
markers used (Rousset et al., 2004; Capa et al., 2011)
and also after performing ecophysiological investigations (Amieva & Reed, 1987; Dubois et al., 2005). The
organization and structure of tentacular filaments
and the organization of cilia on tentacles have suggested Sabellariids to be closer to Spionids and Terebellids than Sabellids (Dubois et al., 2005). Our
hypothesis also suggests that Spionida is polyphyletic
(as in Rousset et al., 2004) with Magelona recovered
as closely related to Cirratulidae. The monophyly of
Terebellida could also not been assessed as Pectinariidae and Oweniidae were found as sister groups.
These results suggest, as in many other studies trying
to address basal relationships in Annelida (e.g. Struck
& Purschke, 2005; Rousset et al., 2007, Zrzavý et al.,
2009), that further investigations should probably
consider other markers that enlighten the explosive
radiation of this group of metazoans.
If Sabellariidae is not closely related to Sabellida,
the idea of the common origin of ‘crown’ structures
and arrangement of chaetae in thoracic and abdominal segments (Rouse & Fauchald, 1997) is discarded
(as Kieselbach & Hausen, 2008). It would also mean
that the scaphe and cauda, the opercular structure
(Hartman, 1944; Rouse & Pleijel, 2001), and the
building organ (Rousset et al., 2004) are not homologous (as in Watson, 1928). If Terebellida and Sabellariidae are also not closely related, the head
structures and the tentacles in terebellids and sabellariids also have different origins.
The suggestion that Sabellariidae is one of the
most specialized groups of polychaetes (Dales, 1952)
is defended herein. If the sister-group relationships
with Spionidae are confirmed, that would mean that
the tubiculous ancestor of this group of worms developed a complex operculum with paleae from some
anterior segments that allowed them to live permanently in a tube and with associated feeding structures, such as tentacular filaments for collecting
floating particles and the building organ as a specialized structure for cementing these particles with
secreted gluing substances. A regionalization of the
body also took place and noto-, neuropodia and associated chaetae suffered dramatic changes with
respect to preceding forms.
SABELLARIIDAE
RELATIONSHIPS AND
CHARACTER EVOLUTION
The topologies of the unweighted characters, resulting in a consensus tree where only few clades are
delineated, indicate the high amount of homoplasy
accumulated in the group. A close relationship of
Idanthyrsus and Lygdamis, previously considered as
members of Sabellariinae and Lygdaminae (Kirtley,
1994), respectively, recovered in both the unweighted
and the implied weighted datasets suggest these subfamilies are not monophyletic, and therefore this classification should be avoided.
The implied weighting did resolve the polytomies
and uncertain relationships within Sabellariidae (for
other examples and discussion of the methodology,
see Goloboff, 1993, Goloboff et al., 2008a; Ramírez,
2003). But different results are reached depending
on the value given to the concavity constant. Only
the groups presented in all concavities explored
should be considered as firmly established (Goloboff
et al., 2008a) and therefore the basal relationships
among those members of ‘Clade A’ should considered
as unresolved for now.
Major morphological differences among members
of Sabellariidae are found in the anterior end, probably due to major adaptative variations in morphology and anatomical structure, resulted as a
consequence of external applied stress caused by
environmental conditions or biological interactions
(Kirtley, 1994). In contrast, the posterior segments
are similar in all genera and species. The number of
parathoracic segments justified the erection of the
two subfamilies (Kirtley, 1994), a criterion that
seemed justified because this is established very
early in the course of development (Bhaud &
Fernández-Álamo, 2001). However, we advocate that
the number of parathoracic segments is homoplastic
with four segments being the plesiomorphic condition and changing to three twice during the sabellariid radiation.
The presence of short opercula, geniculate outer
paleae, and inner concave paleae arranged in semicircles are apomorphic features that characterize
the derived sabellariids (Bathysabellaria, Gunnarea,
Paraidanthyrsus,
Sabellaria,
Phragmatopoma,
Neosabellaria). Besides, given the lack of a phylogenetic framework, some authors predicted that Phragmatopoma was a derived sabellariid due to their
developed paleae (Dales, 1952) and sperm shape (Eckelbarger, 1976), compared with other Sabellariidae.
The relationship between Phalacrostemma and this
group of derived sabellariids is supported by features
that are not of much significance such as length of the
operculum, limbation of hooks (inapplicable for most
taxa), and the presence of chaetae on the neuropodia
of segment 1, instead of being grouped with the taxa
that show a similar opercular structure and, in our
opinion, should be considered with caution.
About 20 species, predominantly of the genera
Phragmatopoma, Sabellaria, and including the monospecific genus Gunnerea, construct colonies and reefs
of aggregated tubes in the intertidal and subtidal
zones of temperate and tropical coasts in many parts
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
SYSTEMATICS OF SABELLARIIDAE
of the world (Achari, 1974; Kirtley, 1974; Pawlik,
1988a, b; Pawlik & Faulkner, 1988). After the phylogenetic hypothesis presented herein we can conclude
that this characteristic and the bathymetric restrictions of some of the taxa seem not to have any
phylogenetic constraint.
This paper represents the first comprehensive systematic revision of the family. Combined morphological and molecular data confirm the monophyly of
Sabellariidae and its close relationships with Spionida. Phylogenetic relationships within Sabellariidae
have been assessed using morphological data and
suggest that the established sabellariid subfamilies
are not monophyletic and the proposed groups are
now based on opercular and chaetal features. The
descriptions of morphological features together with
the illustrations will, we hope, be used as a baseline
for further systematic and taxonomic studies in
the group, many species of which remain poorly
described.
ACKNOWLEDGEMENTS
We are indebted to Dr Ehiroh Nishi, who provided
some specimens from Japan, Dr Leslie Harris for
providing detailed information about certain features
present in Gesaia fossae and Idanthyrsus macropaleus, Ms Lexie Walker who also provided very useful
information on spionids, Dr Tarik Mezianine who sent
some sabellariid types at short notice, and Jim Lowry
for assisting with DELTA. We thank Kate Attwood
who greatly helped in an early stage of the sabellariid
revision and Sue Lindsay for her great work in the
SEM lab. The present paper was financially supported by Australian Biological Resources Study
(ABRS)/Australian Museum research grants to P.H.
and M.C.
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APPENDIX 1
Description of characters and character-states for
analyses at family level
HEAD (CHARACTERS 1–3)
In the head of sabellariids, the prostomium and
peristomium are partially or totally fused to one
another and with the first two thoracic segments,
forming an anterior operculum that seals the tube
aperture. The delimitation of segments and origin of
certain structures is complicated due to this fusion,
although several studies dealing with the ontogeny
of some species (e.g. Dales, 1952; Wilson, 1968, 1977;
Eckelbarger, 1975, 1976, 1977, 1978; Orrhage, 1978;
Bhaud & Fernández-Álamo, 2001) have resolved
some of these issues. The operculum, the paleae, the
dorsal spines and hooks, and the tentacular filaments on the inner margin of the opercular lobes are
largely derived from the first segment (Dales, 1952;
Wilson, 1977). The prostomium is limited to a small
region in front of the mouth and the peristomium is
limited to the lips, on the buccal region (Orrhage,
1978; Amieva & Reed, 1987). The prostomium is also
indistinct or partially fused to the peristomium in
members of Oweniidae, Pectinariidae, Serpulidae,
Sabellidae, and Terebellidae (Rouse & Pleijel, 2001;
Rousset et al., 2003; Garraffoni & Lana, 2008, 2010;
Capa et al., 2011; Nogueira et al., 2010), and as in
sabellariids, the head is also fused to the first two
segments in Pectinariidae (Watson, 1928). Some terebellids also have the head partially fused to the
first segment (Orrhage, 2001; Garraffoni & Lana,
2010; Nogueira et al., 2010) but there is no operculum present.
HEAD
APPENDAGES (CHARACTERS
4–10)
All members of the ingroup lack antennae, with the
exception of some species of Trochochaeta (Pettibone,
1976) that bear a small, simple antenna on the midposterior end of the prostomium, and Spionidae
(Orrhage, 1966; Orrhage & Müller, 2005) with a
middle appendix sometimes referred to as an occipital
tentacle and considered to have appeared convergently (Rouse & Pleijel, 2001). Poecilochaetus is considered not to have antennae as the so-called median
tentacle or facial tubicle has a peristomial origin
277
(Mackie, 1990; Blake, 1996; Rouse & Pleijel, 2001).
Members of the outgroup have both median and
lateral antennae.
All palps, whether emerging from the prostomium
or peristomium, have been considered as homologous
in previous studies because they are similarly innervated (e.g. Orrhage, 1978; Orrhage & Müller, 2005)
but have here been considered as two different characters (as in Capa et al., 2011). Although earlier
studies suggested a possible prostomial origin of palps
due their unclear position just posterior to the larval
prototroch in Sabellariidae (Dales, 1952), they are
currently considered as peristomial (Dales, 1962;
Orrhage, 1978; Amieva & Reed, 1987; Rouse &
Fauchald, 1997; Rousset et al., 2004; Zrzavý et al.,
2009) due to verification of being behind the cilial
band. A similar arrangement of palps has been
reported in members of Spionidae, Trochochaeta,
Poecilochaetus, and Magelona (Wilson, 1982, 2000;
Rouse & Pleijel, 2001). There are no neuro-anatomical
indications of the presence of palps in Terebellidae,
Ampharetidae, and Pectinariidade (Orrhage, 2001;
Orrhage & Müller, 2005) and therefore the multiple,
grooved buccal tentacles should be considered as nonhomologous to the palps from other polychaetes (Garraffoni & Lana, 2010; Nogueira et al., 2010). The
homology statement of tentacles among members of
Oweniidae (e.g. Hartman, 1960; Rouse & Pleijel,
2001; Parapar, 2003, 2006) and the palps of other
polychaetes is still under discussion and therefore
remains herein as unknown. These structures have
been interpreted as prostomial palps, relating Oweniidae to other prostomial crown-bearing families
(Rouse & Fauchald, 1997) but also as a prostomial
lobe crown (Rousset et al., 2004) and with palps
absent based on Gardiner’s (1978) finding of monociliated epidermal cells, not shared by any other polychaetes. In Sternaspis, palps have been interpreted as
absent (Rouse & Fauchald, 1997; Hutchings, 2000)
although in some it is still debated if the appendages
present in some specimens could be considered as
such (Rouse & Pleijel, 2001). In cirratulids, the presence of palps has not yet been properly investigated
(Orrhage & Müller, 2005) and anterior appendices
seem to originate in the peristomium and probably
migrate posteriorly (Rouse & Pleijel, 2001; Rousset
et al., 2004; Zrzavý et al., 2009).
The term ‘buccal tentacles’ has been used in the
literature to refer to those appendages located near
the mouth that assist with the collection of food
particles (e.g. Orrhage, 1978) but homology statements need to be established in several groups. The
slender appendages of the lateral parts of the ventral
side of the operculum of Sabellariidae, referred to
as tentacular filaments, were earlier interpreted
as palps homologous with those of the Spionidae
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
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M. CAPA ET AL.
(Johansson, 1927; Binard & Jeener, 1928), as
extended lateral parts of the upper lip of the mouth,
according to innervation studies (Orrhage, 1978;
Orrhage & Müller, 2005) or as derived from segment
1, according to developmental studies (Dales, 1952;
Wilson, 1977 and herein; Char. 10). Some studies
suggested that these tentacles were homologous
in Terebellidae, Ampharetidae, and Pectinariidae
(Holthe, 1986; Orrhage, 2001; Orrhage & Müller,
2005), although recent studies document the prostomial origin of these tentacles in terebellids and peristomial in Ampharetidae and Pectinariidae (Garraffoni
& Lana, 2008, 2010; Nogueira et al., 2010).
BUILDING
ORGAN AND ANTERIOR GLANDS
(CHARACTERS 11–12)
Sabellariids posses a glandular, U-shaped organ
(Figs 1B, 3H) that surrounds the posterior and lateral
margins of the mouth and which is used to construct
its robust sandy tube. Captured particles are conveyed along the tentacular filaments to the building
organ, where they are held and evaluated for size,
shape, and composition. Suitable particles are covered
with cement secreted from thoracic cement glands
and then pressed into place at the end of the tube by
the building organ (Eckelbarger, 1978; Stewart et al.,
2004). The bioadhesive material secreted by some
species in the family not only serves to form the tube
but also provides the chemical signal that is recognized by the planktonic larvae and induce them to
attach and metamorphose (Eckelbarger, 1978;
Jensen, 1992). The building organ has been regarded
as part of segment 1 (Rousset et al., 2004) and
homologous to the ‘cementing organ’ found in Pectinaria (Watson, 1928) on segment 2 (Rousset et al.,
2004). It has therefore being scored herein as such to
test homology.
Anterior glands with a major role in tube formation
have been described in several groups of tubiculous
polychaetes (Fitzhugh, 1989; Kirtley, 1994; Vovelle,
1997; Rousset et al., 2003; Glasby, Hutchings & Hall,
2004; Vinn & ten Hove, 2009). The parathoracic segments in sabellariids have more or less defined glands
on their ventral surfaces that are used in association
with the building organ to produce cementing and
fixative substances (Kirtley, 1994). By contrast, distinct ventral shields are typical for most Terebellida
and Sabellida (McHugh, 1995; Vovelle, 1997; Rousset
et al., 2004; Garraffoni & Lana, 2008) although controversy exists regarding interpretations of these
glandular pads (for details see Garraffoni & Lana,
2008; Nogueira et al., 2010).
unequal in size and structure, sometimes composed
of several lobes (Spionidae or Nereididae). Welldeveloped parapodia are often supported by aciculae
(as in Nereididae and Polynoidae) and always bear
chaetae. In errant polychaetes, neuropodia are generally larger than notopodia while in tubiculous
worms this is not always the case and the presence of
glandular tori is common among them. Other distinct
parapodial structures include the dorsal and ventral
cirri whose presence, shape, and arrangement vary
between and sometimes within families and have
therefore been included in the present study.
CHAETAL
ARRANGEMENT (CHARACTERS
22–25)
Previous studies have considered the arrangement of
chaetae in parathoracic and abdominal segments in
sabellariids, with uncini on the notopodia of the
abdominal region, as a consequence of the chaetal
inversion typical of members of Sabellida (KnightJones, 1981; Fitzhugh, 1989; Smith, 1991; Rouse &
Fauchald, 1997; Schulze, 2003; Rousset et al., 2004).
But recent studies demonstrate that this chaetal
arrangement corresponds to different processes and
are not homologous (Kieselbach & Hausen, 2008).
In many sedentary polychaetes chaetae are
arranged in transverse rows perpendicular to the
body longitudinal axis (for an explanation of the
chaetal formation process on the edge of rows and
some modifications to this pattern see Hausen, 2005).
Spionidae, Magelona, and Cirratulidae have transverse rows of chaetae in both parapodial rami of all
chaetigers while in Poecilochaetous and Trochochaeta
this chaetal arrangement is only found on anterior
chaetigers. Transverse rows are present in neuropodia of Terebellida and Sabellidae but notochaetae are
arranged in bundles, although this arrangement is
inverted in the latter group on abdominal segments.
Chaetal arrangement in Oweniidae has been interpreted as bundles in the notopodia (Parapar, 2003)
and the patches of hooks in neuropodia as modified
transverse rows (Hausen, 2005). Chaetal arrangement in Sternaspis is not yet fully understood
(Beesley, Ross & Glasby, 2000; Rouse & Pleijel, 2001)
and has been interpreted here as transverse rows of
noto- and neurochaetae in the anterior segments
(Petersen, 2000) but as the bundles on the posterior
end could be noto-, neurochaetae or both (Rouse &
Pleijel, 2001) it has been scored as a question mark
herein.
CHAETAL
SHAPE AND ULTRASTRUCTURE
PARAPODIA (CHARACTERS 14–21)
(CHARACTERS 26–40)
Parapodia are generally present in polychaetes and,
when present, they consist of two rami, similar or
The diversity of chaetae and their arrangements is
high among polychaetes and has been the issue of
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SYSTEMATICS OF SABELLARIIDAE
several studies. Different interpretations have been
suggested over the years (e.g. Bartolomaeus, 1995,
1998; Rouse & Fauchald, 1997; Hausen & Bartolomaeus, 1998; Hausen, 2005) and consequently suggesting in some cases dissimilar phylogenetic
hypotheses. According to some authors homology can
be assumed between subchaetal units for individual
chaetae (e.g. Hausen, 2005). Therefore, for taxa
bearing uncini, we have considered absence or presence of capitium, rostrum, subrostrum, subrostral
process, manubrium, or basal process as independent
characters and for those with chaetae, the presence of
aciculae (non-protruding chaetae supporting parapodia), simple (spines or capillaries) or compound
chaetae. We have also considered the presence and
arrangement of a particular type of chaetae of anterior segments to incorporate those typical of the operculum in Sabellariidae and Pectinariidae (Dales,
1952; Orrhage, 1978; Rouse & Pleijel, 2001; Hutchings & Peart, 2002; Orrhage & Müller, 2005). Some
Sabellariidae present thick spines, with straight or
curved tips (hooks), formed in the sacs of the opercular paleae and considered as derived from segment 1
(Orrhage, 1978) but it is uncertain if they are noto- or
neurochaetae. In some Ampharetidae hooks are
present in the third or fourth segment (Beesley et al.,
2000) and seem to be notopodial (Nogueira et al.,
2010). Chaetae in the rest of chaetigers have been
herein classified as spines (simple, thick chaetae
regardless of shape), capillaries (simple thin chaetae),
or compound chaetae (comprising two units, a shaft
and a blade). Members of the outgroup have typically
simple (Polynoidae) and compound (Nereididae)
chaetae. All members of the ingroup have simple
chaetae.
Unicini or hooks with rostrum or main fang are
present in Terebellidae, Sabellidae and in the thoracic
segments of Serpulidae and absent in the rest of taxa
included in the present study (Bartolomaeus, 1995;
Hausen, 2005). The rostrum has been interpreted as
present in Magelona (as Rousset et al., 2004). A capitium, or teeth above main fang, is present in all taxa
except for some Spionidae (Hausen & Bartolomaeus,
1998). A subrostrum or breast is absent in Oweniidae,
Magelona, Pectinariidae, and Spionidae (Bartolomaeus, 1995; Hausen & Bartolomaeus, 1998) and all
terminals except for some Serpulidae and Terebellidae have a manubrium. Basal process (as in
Hausen, 2005) is present in Terebellida.
POSTERIOR SEGMENTS AND PYGIDIUM
(CHARACTERS 41–43)
There is some controversy about the segmented
nature of the posterior region of sabellariids, the
cauda, and some authors suggest that it be considered
279
as segmented (Wilson, 1929) while others do not
(Kirtley, 1994). This has not only morphological but
phylogenetic implications (see Rouse & Fauchald,
1997). In the present study these two structures have
been considered as different and not homologous as
their morphology and function differ (see Watson,
1928). Pygidium is in some taxa provided with
some cirri (Spionidae, Poecilochaetous, Magelona, and
Pectinariidae).
SENSE
ORGANS (CHARACTERS
44–51)
Most polychaetes possess prostomial eyes and are
also called cerebral eyes (Char. 11) because of their
proximity to the brain. Some Phyllodocida have multicellular (Char. 12) and lenticular eyes (Char. 13)
absent in the rest of the groups (e.g. Suschenko &
Purschke, 2009). Prostomial ocelli (Dales, 1952) are
present in some sabellariid adults, in the area near
the median ridge (e.g. Kirtley, 1994; Nishi & Núñez,
1999) or at the base of the palps. They seem to be a
feature that is consistent within some genera
although revisions should be made as they are not
often mentioned in descriptions. Sabellids show a
wide range of prostomial eyes from simple ocelli to
compound eyes formed by groups of ommatidia and
arranged in several positions of the radiolar crown
(Suschenko & Purschke, 2009; Capa et al., 2011). In
Terebellida, the presence of prostomial eyes is a variable condition and has been scored herein as present
in Sternaspis (Petersen, 2000), some terebellids
(Nogueira et al., 2010), and cirratulids but absent in
Magelona (Wilson, 2000; Rouse & Pleijel, 2001) and
adults of pectinariids (Thorson, 1946).
Nuchal organs are present in most polychaetes and
they have been recently found in some taxa considered as not having them. In those remaining cases, as
in Oweniidae, Sternaspis, or Magelona (Rullier, 1950,
1951; Rouse & Pleijel, 2001; Bartolomaeus et al.,
2005; Lindsay, 2009), their absence is considered as a
secondary loss (Purschke, 1997, 2005). Some polychaetes present large nuchal organs while in sabellids,
serpulids, sabellariids, terebellids, or pectinariids
these are reduced (Watson, 1928; Purschke, 1997). All
nuchal organs exhibit similar ultrastructural components (e.g. Rullier, 1951; Purschke, 1990, 1997, 2005;
Jelsing, 2003), with some exceptions such as Protodilidae where the cuticle has been replaced by a
special cover (Rullier, 1951; Jelsing, 2003; Purschke,
2005). Nuchal organs of Sabellidae, Serpulidae, and
Terebellidae are unpaired and in the former two
groups they are in an uncommon position in the
dorsal epithelium of the mouth cavity (Purschke,
1997, 2005). In sabellariids nuchal organs are located
at the base of each palp (Johansson, 1927; Orrhage,
1978).
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280
M. CAPA ET AL.
Lateral organs are known to occur and to present a
similar structure and cellular composition in some of
the taxa included in the present study, such as Ampharetidae, Magelona, Pectinariidae, Poecilochaetus,
Spionidae, Terebellidae, and Trochochaeta (Purschke,
2005; Purschke & Hausen, 2007).
BRANCHIAE (CHARACTERS 52–54)
Segmental branchiae are present in most members of
the ingroup. They sometimes arise from the parapodia (Sabellariidae and Pectinariidae) or epithelium of
the dorsum of the body and are absent in the
outgroup. They can be present in most segments
(Cirratulidae), be restricted to only a few anterior
(Spionidae, Ampharetidae, and most Terebellidae) or
posterior ones (Poecilochaetus, Sternaspis). Sabellidae
and Serpulidae do posses ‘branchiae’ but they are
prostomial and Trochochaeta lack branchiae although
some of the segmental papillae along the midventral
line of the body have been erroneously interpreted as
such (Rouse & Pleijel, 2001).
MUSCLES (CHARACTER 55)
The type of muscles (longitudinal, oblique, circular)
and the arrangement shown amongst polychaetes
have been suggested to be of interest for systematic
and phylogenetic purposes (Tzetlin & Filippova, 2005;
Purschke & Müller, 2006) but for many taxa no data
are available. For the present study, we have considered the presence of the circular fibres because of
some information available in the literature (Tzetlin
& Filippova, 2005; Purschke & Müller, 2006). They
are arranged in an almost complete cylinder interrupted only by the ventral nerve cord in some sedentary polychaetes and are less developed in nereidids
and terebellids and suggested to be absent in
Magelona and at least some spionids, oweniids, and
polynoids (Tzetlin & Filippova, 2005; Purschke &
Müller, 2006, and references therein).
ALIMENTARY
CANAL (CHARACTERS
56–63)
Tzetlin & Purschke (2005) reviewed and updated the
information regarding the alimentary canal of polychaetes. The anterior end of the gut or foregut is very
variable among polychaetes due to adaptation to the
environment and feeding strategies. In Sabellidae,
Serpulidae, and Sabellariidae the foregut is simple
and a buccal organ seems to be absent (Beesley et al.,
2000; Rouse & Pleijel, 2001; Zrzavý et al., 2009). The
types of buccal organs have been classified as
a ventral pharyngeal organ (simple or with welldeveloped muscles), axial muscular pharynx or proboscis, axial non-muscular pharynx and dorsolateral
ciliary folds which can occur in combination with a
ventral pharyngeal organ (e.g. Beesley et al., 2000;
Rouse & Pleijel, 2001; Tzetlin & Purschke, 2005). A
ventral pharyngeal organ is present in most Terebellida and also Oweniidae. An axial muscular pharynx
is present in Phyllodocida (outgroup) and some have
sclerotized structures such as jaws or paragnaths
derived from the cuticle that covers the epithelium of
the foregut (Tzetlin & Purschke, 2005). The proboscis
in Sternaspis (Dales, 1962; Rouse & Fauchald, 1997;
Tzetlin & Purschke, 2005) has also been interpreted
as simple and axial.
The midgut or intestine is formed by a straight tube
running along the body from the mouth to the anus in
most polychaetes but in some groups it is coiled
within the septa dividing each segment (Cirratulidae), forms one or two loops in the anterior part of
the body (Ampharetidae, Pectinariidae) or it has
lateral and dorsal branches (Polynoidae). Sabellariidae bears a swollen and thickened structure (Fig. 9D,
E) in the abdominal region referred to a ‘proventricle’
(McIntosh, 1922) and a band of tissue is present in
the mid-dorsum and sometimes mid-ventrum where
the gut content can be seen through the body wall.
GULAR
MEMBRANE (CHARACTERS
64–65)
The presence of this modified septum is characteristic
of some Terebellida but its shape and arrangement
differ between groups. In the present study the presence and position have been included as this membrane attaches to the body wall between the fourth
and fifth segment in Terebellidae and Ampharetidae
and between the third and the fourth in Pectinariidae
(Zhadan & Tzetlin, 2003).
NEPHRIDIA (CHARACTERS 66–69)
The excretory organs in polychaetes have been classified as protonephridia or metanephridia according
to their function. These organs do sometimes vary
with developmental stage, the monociliated protonephridia being the potential primitive condition for
the first pair of nephridia differentiated during development (head kidneys) in Annelida and the rest of
segmental nephridia derived from those were metanephridia (for a review see Bartolomaeus & Quast,
2005).
Sabellariids, sabellids, and serpulids have a single
anterior pair of excretory segmental organs (Meyer,
1887; Dehorne, 1952) and a posterior pair for discharging gametes (Meyer, 1887; Rouse & Fauchald,
1997). This is also the case for Sternaspis and cirratulids (Rouse & Pleijel, 2001). Pectinariids, ampharetids, and some members of Spionida and
Terebellidae have more than one anterior segmental
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SYSTEMATICS OF SABELLARIIDAE
organ acting as nephridia, in addition to posterior
gonoducts, but there are some exceptions, such as
members of Pista with only one single pair of
nephridia (Rouse & Pleijel, 2001). The outgroup has
segmental organs in most segments.
CIRCULATORY
SYSTEM (CHARACTERS
70–72)
All taxa included in the analyses have a closed circulatory system. Some groups, such as Ampharetidae,
Pectinariidae, Terebellidae, Cirratulidae, and Sabellariidae, have a heart body that forms blood (Dales &
Pell, 1970; Rouse & Pleijel, 2001), probably homologous to the intravasal tissue found in serpulids
(Hanson, 1951). Most polychaetes have acellular
blood and therefore have developed other defence
systems against infections. Exceptions are Magelona
and Pectinaria (Kennedy & Dales, 1958; Rouse &
Pleijel, 2001).
NERVOUS
SYSTEM (CHARACTERS
73–74)
Most polychaetes have a subepidermal nervous
system but there are some exceptions, such as the
Oweniidae, included herein. Some annelids, generally
the predators, have well-developed mushroom bodies,
a brain centre innervating the palps (Heuer et al.,
2010; Loesel & Heuer, 2010) but some taxa still need
to be investigated.
REPRODUCTIVE
STRATEGY, FERTILIZATION, AND
PARENTAL CARE (CHARACTERS
75–79)
The majority of polychaetes are gonochoric but
examples of hermaphroditism have been documented
and are not scarce (McIntosh, 1911; Tweedell, 1966;
Schroeder & Hermans, 1975; Rouse & Fitzhugh,
1994; Giangrande, 1997). In the present study we
have, in some cases, generalized for the genera if
cases of specific terminals were not available. Most
polychaetes spawn gametes freely into the sea water
(Blake, 2006; Rouse, Kupriyanova & Nishi, 2006; Halt
et al., 2006) but some nereidids or polynoids do have
pseudocopulation (Pettibone, 1963; Hsieh & Simon,
1990) and some spionids do transfer sperm in spermatophores (Rice, 1978; Hsieh & Simon, 1990). Polychaetes sometimes exhibit some sort of parental care
or brooding of their eggs, larvae, or juveniles, like
several sabellids, serpulids, and some spionids (Rouse
& Fitzhugh, 1994; Blake, 2006; Rouse et al., 2006),
none of them included in this study.
GAMETOGENESIS (CHARACTERS 80–86)
Some aspects of oogenesis, such as structure and
position of the ovary, the type of oogenesis, and the
structure of the egg envelope appear to be conserved
281
at family level among the polychaetes, with some
exceptions and have been proposed as having potential phylogenetic use (Eckelbarger, 2005, 2006). Most
polychaete ovaries are retroperitoneal and are generally located in some anterior segments (Terebellidae,
Ampharetidae, Pectinariidae, Polydora) or in all segments along the body, but some families lack discrete
ovaries, such as Nereididae (Eckelbarger, 2005).
Sabellariids produce and store gametes in the coelomic cavity of the abdominal segments and gametecontaining segments of mature males appear whitish,
while those of mature females are blue in colour.
Some of the groups that show diversity of structure
and position of the ovary are serpulids (Kupriyanova
et al., 2001). Oogenesis can take place in the ovary or
in the coelom. Examples of intraovarian oogenesis are
in members of Sabellariidae, while in Ampharetidae,
Cirratulidae, Polydora, Sternaspis, Pectinariidae,
Sabellidae, and Terebellidae the oocytes develop and
grow outside the ovary and they leave this organ as
individual previtellogenic cells (Pectinariidae and
Sabellidae) or in clusters (Ampharetidae, Cirratulidae, Pectinariidae, Terebellidae) (Eckelbarger, 2005,
2006; Blake, 2006; Halt et al., 2006). Mature eggs are
surrounded by egg envelopes consisting of an extracellular matrix penetrated by microvilli, and in
members or certain families such as Spionidae or
Serpulidae this egg envelope can be ornamented
(Blake & Arnofsky, 1999; Kupriyanova et al., 2001;
Eckelbarger, 2005).
The sperm of only a few taxa have been studied in
detail but shape of the sperm varies among species
(e.g. Franzén, 1956; Eckelbarger, 1976, 1984; Rouse,
2006a). Sternapsis and most cirratulids have shortheaded sperm but there are some exceptions in cases of
species that brood (Halt et al., 2006). All sabellariids
studied are gonochoric, broadcast spawners (Wilson,
1991). When animals are mature they have welldeveloped gonads throughout the year, suggesting that
they are able to reproduce all year long, at least in
some of the species studied (e.g. Eckelbarger, 1976).
LARVAL
DEVELOPMENT AND LARVAL FEATURES
(CHARACTERS 87–97)
Development is highly variable among polychaetes
with examples of planktotrophic, lecithotrophic larvae
and some with direct development even in closely
related species (e.g. Terebellidae, Serpulidae, and
Spionidae; Wilson, 1991; Giangrande, 1997; Rouse,
2003). Several characters reflecting the absence or
presence of larval ciliary bands reviewed in detail
(Rouse, 1999, 2006b; Pernet, 2003) have been incorporated in the analyses. Sabellariid larvae seem not
to have a metratroch but a prototroch that overhangs
the lateral edges of the mouth acting in a similar way
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282
M. CAPA ET AL.
to a metratroch (Rouse, 2006b). All sabellariids have
a long development in the plankton. Life-history
studies on several species have demonstrated the
great similarities in larval development between
members of Sabellariidae (Wilson, 1929, 1991; Dales,
1952; Curtis, 1973; Eckelbarger, 1975, 1976, 1977,
1978). The common body organization in many of the
species studied indicates a particularly homogenous
developmental type at the family level (Bhaud &
Fernández-Álamo, 2001). The main differences among
early larvae are related to the size and colour pigments, but during later larval stages the development
of chaetae, both spines and paleae, differ between
species and have been using for taxonomic purposes
in the past (Eckelbarger, 1976). They superficially
resemble those of spionids (Caullery, 1914; Wilson,
1929; Dales, 1952).
TUBE (CHARACTERS 98–99)
The typical polychaete tube dwellers are members of
Sabellida, Oweniidae, Terebelliformia, and Spionida
but some other groups build tubes even if temporary,
like most Nereididae. Within the ingroup most tube
dwellers produce tubes made of mucus and attached
sediment particles although members of Serpulidae
typically build calcareous tubes.
APPENDIX 2
DESCRIPTION
OF CHARACTERS FOR ASSESSING
THE SABELLARIID PHYLOGENY
The body of sabellariids is divided up into four
regions: the operculum, parathorax, abdomen and
caudal region (Fig. 1).
OPERCULUM:
HEAD AND THORAX (CHARACTERS
shape of opercular papillae (Fig. 3C) is species specific
and it has sometimes been used for taxonomic
purposes.
On the ventral and inner side of each opercular
lobe, other appendages may be inserted, referred
herein to as tentacular filaments (Hartman, 1944;
Dales, 1952; Kirtley, 1994; Figs 5, 6) or oral filaments
according to some authors (Orrhage, 1978) and seem
to be a synapomorphy for the family (Rouse & Pleijel,
2001). These are ciliated, and often grooved proximally and involved in the transport of food particles
to the mouth (Dales, 1952; Orrhage, 1978; Dubois
et al., 2005; Riisgård & Nielsen, 2006). Tentacular
filaments can be simple (unbranched, Fig. 2D) or compound (branched, Figs 2E, 3D, E). In some species,
the oral filaments are replaced by elongate tentacular
buccal flaps or oral plates (Figs 7F, 8F) or by apophyseal ridges along the posterior–lateral margins of the
buccal cavity (Kirtley, 1994).
A pair of grooved and ciliated palps (Figs 5B, 7D–F,
8E, G–H; Treadwell, 1926), also referred to as prostomial or pretentacular filaments (Dales, 1952;
Kirtley, 1994) or other terms (see a summary in
Orrhage, 1978), arise from behind the prototroch
(Dales, 1962), on the anterior margin of the upper
transverse lip, in front of and dorsal to the mouth
(Johansson, 1927; Orrhage, 1978) and therefore are
considered herein as peristomial. The relative length
of the palps and operculum has been considered in
this study as a variable character among groups of
sabellariids, although it has been indicated to be
contractile (Treadwell, 1926). In some sabellariids an
unpaired appendage, called median organ (Figs 5B,
8H–K), median cirrus, preoral lobe (Caullery, 1944),
or tentacule (Gravier, 1909) with sensory function is
present at the dorsal junction of the opercular lobes
(Kirtley, 1994).
1–8)
The operculum (Figs 5A, B, 6A) is an elongate structure with two lateral fleshy lobes. Some genera have
the opercular lobes completely fused (Fig. 7A), fused
dorsally and with a deep ventral groove on the ventral
margin (Fig. 7B) or completely separated (Fig. 7C).
The relative length of the operculum varies among
sabellariids but in most cases is consistent within
genera (Fig. 7D, F). The distal end of the operculum,
called a disc (Orrhage, 1978) or a crown (Kirtley,
1994; Rouse & Fauchald, 1997), bears rows of paleae
and is surrounded by appendages called opercular
papillae (Figs 5A, 7D–F). In most sabellariids
this disc is oriented perpendicular to the longitudinal axis of the body (Fig. 8A), although in some
genera it is oriented in an oblique position, often
described as having a distal end sloped posteriorly to
midline (Kirtley, 1994; Fig. 8B). The number and
OPERCULAR
CHAETAE (CHARACTERS
9–25)
The opercular paleae arise in two separate chaetigerous sacs (Ebling, 1945; Dales, 1952; Orrhage, 1978)
forming two distinct series of paleae, and outer and an
inner row, on each lobe (Hartman, 1994). The orientation of the distal blades sometimes gives the appearance of animals presenting three concentric rows of
paleae, where both the inner and the median row are
formed from the inner chaetigerous sac (Kirtley, 1994).
In those cases, the blades of paleae from the median
and inner row can be both directed inwards (Fig. 7G)
or, the inner paleae can be directed inwards and the
median paleae outwards (Fig. 7H). For the present
study we have considered the total number of paleae in
the outer and inner row as separate characters in the
matrix. The outer paleae are generally arranged in
facing semicircles but this arrangement varies in some
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SYSTEMATICS OF SABELLARIIDAE
species where outer paleae display a spiral arrangement (Fig. 7I). Opercular paleae show a great diversity
in number, shape and size. We have followed the
convention (Ebling, 1945; Kirtley, 1994) of referring to
the proximal end, which is attached to the muscular
tissue, as the paleal shaft and to the distal exposed
end, which is normally ornamented, as the blade
(Fig. 7J). All paleae appear to consist of two distinct
regions, an inner core and an outer layer also referred
to as thecae (e.g. Kirtley, 1994). The inner core is
striated longitudinally and the outer layer (cortex or
thecae) is clear and homogeneous, forming an even
covering around the shaft and relatively thinner in the
blade region, with more or less packed fibres or microtubules (Ebling, 1945). In some paleae an alveolar
structure can be seen, consisting of gas-filled cavities
arranged in longitudinal and transverse rows. Opercular paleae are replaced during metamorphosis and
subsequent adult growth, changing its shape during
ontogeny. In larval stages (features retained in some
adults), they have been classified as choanothecae,
hemithecae, and platythecae depending on the shape
of the section of the blade (for a description of different
types see Kirtley, 1994). The terminology used for
describing paleae in the literature is highly varied. In
this study, we have selected some characters such as
the overall shape of the blade, the ornamentation of
lateral and distal margins, and angle of the blade
to the longitudinal axis of the shaft. We have
avoided subjective terms common in the literature
and we have not scored the fine details of the thecae
types, microsculpture, internal striation, frayed
denticles, curved denticles, shape of shafts, or colour
as these are features that are highly variable
within genera and sometimes even within species. The
types of paleae considered in the present study are
summarized as follows: the angle of the longitudinal
axis of blades and shafts and if they are straight
(Figs 7K, 10A, C–F) or geniculate chaetae (Figs 7L,
10B); regarding the shape of the blade whether they
are flat (Figs 7M, 10D–G), concave (Figs 7N, 10H), or
cylindrical (Figs 7O, 10C); regarding the shape of the
lateral margins and whether they are smooth (Figs 7P,
10D) or denticulate (Figs 7K, 10E, F) chaetae; and
regarding the distal ends, some exhibit smooth edges
(Figs 7P, 10D) and others have denticles (Figs 7M,
10E–G).
Some sabellariids have thick cylindrical and
pointed chaetae on the dorsal edge of the opercular
lobes called nuchal spines (Figs 5B, 9A, 10I, J). These
can have straight or bent distal tips (also known as
nuchal hooks) (Fig. 7Q–S). They are formed in the
sacs of the opercular paleae, and are therefore considered as derived from segment 1 (Orrhage, 1978).
The number of spines and hooks varies among genera
and species and the presence of a more or less devel-
283
oped limbation (e.g. Fig. 7S) on inner or outer margin
of the hooks also characterizes some taxa.
THORAX (CHARACTERS 26–29)
There is a pair of small neuropodia (Figs 5B, 8D,
F–H, M) at the lateral sides of the building organ and
the mouth with conical cirri, which number is of
taxonomic use. In most taxa chaetae are regarded as
the neurochaetae of the first segment (Kirtley, 1994;
Fig. 8G, J). The second thoracic segment bears neuropodial cirri and bundles of capillary neurochaetae
with thecae (coanotheace and hemithecae) make them
appear as bipinnate (as referred by Kirtley, 1994).
Thoracic notopodia do not have chaetae but in some
taxa, dorsal cirri (one to three), referred herein as
lateral lobes may be present (Fig. 9B, C). Branchiae
can be present or absent in segment 2 and, in most
cases, is a generic feature.
PARATHORAX (CHARACTERS 30–31)
Behind the thorax are either three or four parathoracic segments which are biramous. Neuropodia are
conical and notopodia are enlarged lobes. Notopodia
bear stout chaetae with lanceolate tips and cylindrical
capillary chaetae alternating in one row, although
chaetae in neuropodia are smaller in size (e.g. Kieselbach & Hausen, 2008). In some genera the lanceolate chaetae are frayed or denticulated at the tips.
Neuropodia show higher variation between taxa and
some species have both lanceolate chaetae and capillaries (Fig. 10K) while other only bear capillaries or
only lanceolate chaetae (Fig. 10L). All segments have
paired conical and weakly ringed branchiae (Figs 5A,
6B, 9A). Glandular areas are present on the ventrum
of this region and they produce cementing and fixative substances for building the tube (Vovelle, 1965 in
Kirtley, 1994).
ABDOMEN
Parathoracic and abdominal segments can be easily
separated by the presence of uncini in the notopodia.
Some authors indicate an absence of lanceolate neurochaetae, and only presence of capillaries in this
region, although a gradual change has been observed
in some species (Kieselbach & Hausen, 2008).
Abdominal segments are biramous, with conical neuropodia bearing conical ventral cirri and fascicles of
chaetae and notopodia as transverse tori, with a
single row of uncini, diminishing in size posteriorly.
Neurochaetae are capillary (often with well-developed
thecal sculpture, Fig. 10M) arranged in one or two
rows with two separated formation sites (Kieselbach
& Hausen, 2008) and notochaetae are pectinate
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
284
M. CAPA ET AL.
uncini with series of similar-sized denticles pointing
anteriorly (Fig. 10N). Cirri are present between
the two noto- and neuropodia rami (Kieselbach &
Hausen, 2008). Branchiae are present in most or only
on anterior abdominal segments and they decrease
in size towards the posterior end (Figs 5B, 9F). No
abdominal features have been selected for the analyses because the morphological variation is specific
more than generic.
CAUDA
This region is formed by an apparently unsegmented
(although there are other opinions, Wilson, 1929),
smooth, cylindrical tube that is curved along the
ventral surface of the abdomen (Figs 5A, B, 9F). No
features from this region have been selected for the
analyses due to small variation between taxa.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164, 245–284
Maria Capa