Phyllodocidae (Polychaeta) character List
DRAFT DELTA tonatr output, RW, FP 10-JUN-03

#1. Original combination of publication of taxon/

The original combination of the taxon, including author and year of publication.

#3. Type locality of species/

Type locality for described taxa when known reliably.

#4. Type species of genus/

#5. Synonyms:/

Junior synonyms and synonymous species codes from significant benthic survey voucher collections at AM and MoV are included.

#6. Subfamily: <after Pleijel, 1991>/

1. Notophyllinae/

2. Phyllodocinae/

3. Eteoninae/

Subfamily to which taxon belongs, following Pleijel (1991).

#7. References: <major references relevant to taxon>/

Major references for further information are included here, in brief citation format only (author, year).

#10. Description coded by <person(s) writing DELTA character codes for this taxon>/

Name(s) of persons responsible for coding and verifying all data for this taxon.

#11. <Date material examined & coded>/

Date(s) at which these data were collated/updated.

#12. Material examined: <to compile description; including museum lot nos>/

List of material examined and/or published descriptions on which these data are based. Material examined to include museum registration numbers and locality data.

#13. Size range of material examined <total number of chaetigers>/

chaetigers/

Minimum and maximum size of specimens examined for this description. Length in setigers must specify entire worm or anterior fragment.

#14. <Size range of material examined, length>/

mm long/

Minimum and maximum size of specimens examined for this description. Length in mm must specify entire worm or anterior fragment.

#15. <Size range of material examined, width>/

mm wide/

Minimum and maximum size of specimens examined for this description. Width in mm at setiger 10, excluding parapodia.

#17. <Global distribution>/

1. Australia/

2. New Zealand/

3. northern Europe/

4. North America, Pacific coast/

5. North America, Atlantic coast/

6. Arctic/

7. Antarctic/

8. South America, Pacific coast/

9. South America, Atlantic coast/

10. South Africa/

11. Gulf of Mexico/

12. North-west Pacific/

13. Southeast Asia/

Global distribution data based on authoritative references from the recent literature as specified for each taxon.

#18. <Australian distribution>/

1. South-eastern Australia <SA, Tas, Vic, NSW to Sydney>/

2. South-western Australia <WA S of Perth>/

3. North-eastern Australia <NSW N of Sydney, Qld>/

4. North-western Australia <WA N of Perth>/

5. Northern Australia, Gulf of Carpentaria/

6. Not known from Australia/

Australian distribution data to include all material examined and reliably identified in the collections of the Australian Museum, Sydney and Museum of Victoria, Melbourne; and authoritative published records.

#19. Habitat/

1. estuarine/

2. marine, inshore <10 m or shallower>/

3. marine, shelf <10–200 m>/

4. slope <200–2000 m>/

5. deep sea <deeper than 2000 m>/

'estuarine': includes marine embayments that may only periodically have reduced salinity (e.g. the Hawkesbury River).

'inshore': 0 – 10 m (i.e. including intertidal and coastal bays).

'shelf': 10–200 m (but excluding coastal bays).

#20. <Comments on distribution and habitat>/

Additional comments or qualifications on distribution and habitat information.

#21. Colour/

Descriptions of colour and colour patterns are entered as text. Whether this information is based on examination of live or preserved material is also specified here.

#22. Eyes/

1. superficial/

2. deeply sunken/

3. not visible or absent/

One pair of eyes is typical for phyllodocids, although they may be difficult to see. Eyes are normally on the surface of the prostomium, but in Eteone, Hypereteone and Mysta eyes are located beneath a sometimes opaque dermal layer (Pleijel, 1991: 226–229).

#23. <Eye colour>/

Eye colour is used here as a multi-state character, but is probably only useful at the species level.

#24. Prostomium <shape of posterior margin>/

1. with dorsal posterior incision/

2. without dorsal posterior incision/

In the genera Phyllodoce and Zverlinum the posterior margin of the prostomium is indented, with a lateral lobe on each side projecting posteriorly over the dorsum of the first segment (Pleijel, 1991: 226–229).

#25. <Proportions of prostomium>/

1. longer than maximum width/

2. about as long as maximum width/

3. shorter than maximum width/

The proportions of the prostomium may assist in distinguishing some species, especially in the genera Eteone and Hypereteone (see Wilson, 1988).

#26. <Prostomium shape>/

Additional information on the shape of the prostomium is recorded for each taxon as a text character.

#27. Dorsal paired antennae <shape>/

1. broadest basally <tapering ~evenly to tip>/

2. broadest subproximally <droplet-like>/

3. thread-like, with constant thickness/

Most authors refer to phyllodocids as having two pairs of frontal antennae, however following the homologies recognised by Fauchald & Rouse (1997 Zoo. Scripta 26: 71–138) the dorsal pair are here recognised as antennae and the ventral pair as palps. In most phyllodocids the antennae taper towards the tip, but the genera Australophyllum, Clavodoce, Nereiphylla and Notophyllum are inflated and widest at the midpoint of the antenna (Pleijel, 1991: 226–229).

#28. Ventral palps <frequently referred to as 2nd pair of frontal antennae>/

1. similar in size and shape to frontal antennae <implicit>/

2. clavate, shorter and thicker than frontal antennae/

Most authors refer to phyllodocids as having two pairs of frontal antennae, however following the homologies recognised by Fauchald & Rouse (1997 Zoo. Scripta 26: 71–138) the dorsal pair are here recognised as antennae and the ventral pair as palps. In most phyllodocids the antennae and palps are almost identical, but occasionally the slightly ventrally located palps are recognisably different in shape and length.

#29. <Length of paired antennae>/

times length of prostomium/

#30. Prostomial protuberance <distinct or not>/

1. indistinct/

2. well defined/

A distinct distal region of the prostomium where the antennae are inserted may be delineated in some genera. Other phyllodocids have an entire prostomium not obviously delineated as separate regions (Pleijel, 1991: 226–229).

#31. Nuchal organs present as <form>/

1. lateral prostomial protrusions/

2. ciliated pits/

3. dorsal outgrowths on prostomium/

4. conspicuous posterior outgrowths from prostomium/

Nuchal organs are present in all polychaetes, although they may be difficult to see; even in phyllodocids SEM examination may be required to see this character. In phyllodocids, they may be present as knob-like lateral prostomial protrusions; as sunken knobs seen as ciliated pits, and probably only visible in sectioned specimens; or as dorsal prostomial protrusions on the posterior region of the prostomium. This character is probably important in describing phylogenetic relationships among phyllodocid genera, but is difficult to observe in some taxa and best omitted from routine identification procedures unless clearly visible (Pleijel, 1991: 226–229).

#32. <Nuchal organ> comprised of <number of lobes>/

lobes/

#33. Nuchal organs and cirrophores of first pair of tentacular cirri <separate or fused>/

1. separate basally/

2. fused basally/

The nuchal organs may be separate structures, or may be fused to the cirrophores at the base of the first pair of tentacular cirri (Pleijel, 1991: 226–229).

#34. Median antenna <absent, present or as a papilla>/

1. absent/

2. present as a papilla/

3. present/

A fifth, median antenna is present in some phyllodocid genera. Even if this antenna is broken, the scar on the prostomium is often visible. The median antenna may be present as a small papilla or as a distinct, elongate antenna (Pleijel, 1991: 226–229).

#35. <Median antenna> located/

1. closest to paired antennae/

2. closest to eyes/

#36. <Length of median antenna>/

times length of prostomium/

The length of the median antenna, if present, is recorded as a multiple of the length of the prostomium.

#37. Proboscis <2 regions or not; see character notes>/

1. divided into 2 regions/

2. not divided into 2 regions/

The structure of the proboscis is vital for accurate identification of phyllodocid genera and species. If the proboscis is not everted (which is often the case), a ventral incision will need to be reveal the structure on the interior of the inverted proboscis. Phyllodocids in which the proboscis is "uniform" (especially species of Eumida) often have the basal part more smooth than the slightly wrinkled distal region; this seems to be an artefact of preservation, and probosces in these taxa are uniformly covered with papillae. Other genera, e.g. Phyllodoce spp., have a very distinct separation of the proboscis into two regions, each of which has different surface structures and papillation. Austrophyllum is here coded as having a proboscis divided into 2 regions since the arrangement of papillae differs proximally and distally; this is a practical judgement, not the phylogenetic interpretation of this genus given by Pleijel (1991).

#38. Distal region <of divided proboscis, relative length>/

times length of basal region/

The length of the distal region of the proboscis (in taxa with divided probosces), is recorded as a multiple of the length of the basal region.

#39. Basal region <of divided proboscis, papillation>/

1. lacking papillae/

2. with large, conical to rounded papillae/

3. with micropapillae/

The basal region of the divided proboscis may be bare of papillae; large approximately conical or rounded papillae may be present (e.g. in Phyllodoce); or small approximately spherical granules termed micropapillae may be present (as in Eumida and Sige).

#40. <Extent of papillae on basal region of divided proboscis>/

1. uniformly covering basal region <with papillae>/

2. with patches bare of papillae dorsally and ventrally/

3. with patches bare of papillae dorsally only/

#41. <Arrangement of papillae on basal region of divided proboscis>/

1. irregularly or diffusely arranged, not in identifiable rows/

2. in more or less distinct rows/

#42. <Number of rows of papillae on basal region of divided proboscis>/

rows/

If papillae are present, it is important to observe whether they are densely distributed (in no obvious pattern), or to count the number of rows of papillae if they are arranged in distinct longitudinal rows.

#43. <Number of papillae in each longitudinal row on basal region of divided proboscis> about/

papillae in each row/

The number of papillae in each longitudinal row on the basal region of the proboscis is useful in distinguishing some species, e.g. in the genus Phyllodoce.

#44. Distal region of divided proboscis <smooth/papillate/tuberculate>/

1. smooth, lacking papillae <although a terminal ring of papillae may be present>/

2. with papillae/

3. with 6 rows of tubercles/

The distal region of the divided proboscis may be bare of papillae; or large approximately conical or rounded papillae may be present. Note that the distal region of the divided proboscis may also be tuberculated, which is recorded in the next character.

#45. <Arrangement of papillae on distal region of divided proboscis>/

1. diffusely-arranged (not arranged in rows)/

2. in 6 distinct rows, separated by distinct bare areas/

The presence and arrangement of tubercles (rugose longitudinal ridges) and papillae distinguishes species of Phyllodoce and other genera.

#46. Proboscideal papillae <on undivided proboscis>/

1. absent/

2. large, conical to rounded/

3. micropapillae/

4. long filiform papillae <only in Pterrocirrus>/

The undivided proboscis may be bare of papillae; large approximately conical or rounded papillae may be present (e.g. in Phyllodoce); small approximately spherical granules termed micropapillae may be present (as in Eumida and Sige); or elongate, densely-packed filiform papillae unique to the genus Pterocirrus may be present (Pleijel, 1991: 226–229).

#47. <Distribution pattern of papillae on undivided proboscis>/

1. densely and diffusely distributed all over proboscis/

2. with a single lateral longitudinal row on each side/

3. sparsely distributed/

4. proximally with diffusely distributed rounded papillae, distally with 6 rows separated by distinct, smooth areas/

5. 4 slender subdistally located papillae in a ring <proboscis otherwise bare>/

6. 4 longitudinal rows of irregular tuberculae/

7. with papillae in longitudinal rows/

The arrangement of papillae on the undivided proboscis is a character that is most useful at the species level.

#48. <Lateral rows of papillae - shape>/

1. rounded/

2. discoid, flattened/

#49. Proboscis <presence or absence of terminal ring of papillae>/

1. with terminal ring/

2. lacking terminal ring of papillae/

The proboscis of phyllodocids may have a terminal ring of papillae at the oral opening, but note that the proboscis must be fully everted (or fully dissected) to determine if a terminal ring of papillae is present.

#50. Comprising <number of papillae on terminal ring of proboscis>/

papillae/

The number of papillae in the terminal ring is easily counted on good specimens; most species with a terminal ring have about 17 papillae but may range between 15–18 or up to 21 due to intraspecific variability, especially in species of Eumida. This character may only become useful when intraspecific variability is better documented.

#51. Anterior segments <distinct or fused>/

1. all distinct (i.e. one pair or two pairs (dorsal and ventral) of tentacular cirri present on each segment)/

2. include at least two fused or reduced segments (i.e. at least one segment apparently with three pairs of tentacular cirri)/

#52. Segment 1 and 2 <partly extending over prostomium or not>/

1. fused and extending forward over posterior part of prostomium/

2. not covering posterior part of prostomium/

Fusion of anterior segments occurs to varying degrees in different phyllodocid genera and is among the most phylogenetically informative characters in the family. In order to reduce confusion and make routine identifications easier, in this treatment segment 1 means "first visible segment", and this may not be homologous with the first visible segment in other phyllodocid taxa.

Careful observation of well-preserved specimens under low incident light and in dorsal, lateral and ventral views is necessary to determine the degree of fusion of segments and the arrangement of accompanaying tentacular cirri (Pleijel, 1991: 226–229).

#53. Segment 1 <visible dorsally, or not>/

1. visible dorsally/

2. not visible dorsally/

Fusion and/or reduction of anterior segments in phyllodocids makes identification of homologous segments difficult. In order to reduce confusion and make routine identifications easier, in this treatment segment 1 means "first visible segment", and this may not be homologous with the first visible segment in other phyllodocid taxa.

Careful observation of well-preserved specimens under low incident light and in dorsal, lateral and ventral views is necessary to determine the degree of fusion of segments and the arrangement of accompanaying tentacular cirri (Pleijel, 1991: 226–229).

#54. <No of pairs of tentacular cirri on segment 1>/

1. with one pair of tentacular cirri arising immediately posterior to prostomium from the first visible segment after the prostomium/

2. with two pairs of tentacular cirri arising from the first visible segment after the prostomium/

Fusion of anterior segments occurs to varying degrees in different phyllodocid genera and is among the most phylogenetically informative characters in the family. In order to reduce confusion and make routine identifications easier, in this treatment segment 1 means "first visible segment", and this may not be homologous with the first visible segment in other phyllodocid taxa.

Careful observation of well-preserved specimens under low incident light and in dorsal, lateral and ventral views is necessary to determine the degree of fusion of segments and the arrangement of accompanaying tentacular cirri (Pleijel, 1991: 226–229). Scars will usually be visible even if tentacular cirri are broken.

#55. <Tentacular cirri of segment 1> reaching about segment/

In order to reduce confusion and make routine identifications easier, in this treatment tentacular cirri of segment 1 means "tentacular cirri arising from the first visible segment", and may not be homologous with the cirri arising from the first visible segment in other phyllodocid taxa.

The length of the tentacular cirri (measured against their reach posteriorly along the body segments) is a useful species-level character in many phyllodocids.

#56. Segment 2 <with or without dorsal tentacular cirri>/

1. with dorsal tentacular cirri/

2. lacking dorsal tentacular cirri/

Due to reductions and loss of anterior segments, the second visible segment may not be homologous in all phyllodocid taxa.

#57. Segment 2 with dorsal cirri reaching about segment/

In order to reduce confusion and make routine identifications easier, in this treatment dorsal cirri of segment 2 means "dorsal cirri of second visible segment", and this may not be homologous with the cirri of the second visible segment in other phyllodocid taxa.

The length of the tentacular cirri (measured against their reach posteriorly along the body segments) is a useful species-level character in many phyllodocids.

#58. <Shape of dorsal cirri of segment 2>/

Text descriptions of the shape of the dorsal cirri of segment 2 (the second visible segment) may be entered here.

#59. <Shape of> ventral tentacular cirri <of segment 2>/

1. circular in cross-section/

2. flattened/

The ventral cirri of segment 2 (the second visible segment) may be strongly flattened (and usually asymmetrical about the long axis) in some genera, while other genera have tentacular cirri that are approximately circular in cross-section (Pleijel, 1991: 226–229). Unfortunately, these cirri are often broken.

#60. <Ventral tentacular cirri of segment 3 - shape>/

1. tapering gradually/

2. tapering unevenly, inflated subdistally and with distinctly narrow tips/

#61. <Ventral tentacular cirri of segment 2> reaching about segment/

Segment 2 means the "second visible segment"; due to reductions and loss the second visible segment may not be homologous in all phyllodocid taxa. The length of the tentacular cirri (measured against their reach posteriorly along the body segments) is a useful species-level character in many phyllodocids.

#62. Segment 3 <presence or absence of dorsal cirri>/

1. with dorsal cirri/

2. lacking dorsal cirri/

Segment 3 means the "third visible segment"; due to reductions and loss the third visible segment may not be homologous in all phyllodocid taxa. Fusion of anterior segments occurs to varying degrees in different phyllodocid genera and is among the most phylogenetically informative characters in the family. Careful observation of well-preserved specimens under low incident light and in dorsal, lateral and ventral views is necessary to determine the degree of fusion of segments and the arrangement of accompanaying tentacular cirri (Pleijel, 1991: 226–229). Scars will usually be visible even if tentacular cirri are broken.

#63. <Dorsal tentacular cirri of segment 3> reaching about segment/

Segment 3 means the "third visible segment"; due to reductions and loss the third visible segment may not be homologous in all phyllodocid taxa. The length of the tentacular cirri (measured against their reach posteriorly along the body segments) is a useful species-level character in many phyllodocids.

#64. <Segment 3: presence or absence of ventral cirri>/

1. with ventral cirri/

2. lacking ventral cirri/

#65. <Comments on tentacular cirri and segments>/

#66. Tentacular cirri <presence of aciculae>/

1. without aciculae/

2. with aciculae/

Aciculae may support tentacular cirri, but these can usually only be seen (as short projecting chaetae) if the tentacular cirrus is broken off at the base (Pleijel, 1991: 226–229). If it is necessary to do this, it should be done on one side only, so as to leave at least one tentacular cirrus in each position intact.

#67. Chaetae first present on segment <number>/

The segment at which chaetae first occur is useful in some phyllodicids, but some intraspecific variability has been observed for this character (Pleijel, 1987).

#68. Segments 2–4 <with or without specialised chaetae>/

1. with specialised, enlarged spine-like chaetae visible ventrally/

2. lacking specialised enlarged chaetae/

#69. Parapodia <uniramous or biramous>/

1. uniramous (notopodial aciculae absent)/

2. biramous (notopodial aciculae present, on median and posterior setigers, usually beginning at about segment 15 to 30)/

Biramous parapodia, i.e. notopodia with aciculae, do not occur in most phyllodocid genera, and are present only in Australophyllum, Notophyllum and Zverlinum. Even in these genera the first 15–30 parapodia are uniramous, dorsal aciculae being present after these anterior segments (Pleijel, 1991: 226–229). Notopodial aciculae may be visible under low magnification on whole specimens where a dorsal cirrus is broken off.

#70. Notopodial capillary chaetae <presence or absence>/

1. absent/

2. present/

Notopodial capillary chaetae are absent in most phyllodocid genera, and occur only in Australophyllum and Notophyllum and only after the first 15–30 parapodia which are uniramous (Pleijel, 1991: 226–229).

#71. Dorsal cirri <flattened, or inflated>/

1. more or less flattened/

2. distinctly inflated/

Shape of dorsal cirri is useful at the species level, but categorising the various shapes is difficult. Accurate use of this character will depend heavily on comparison with the figures of character states and of individual species incorporated into this database.

#72. <Dorsal cirri: rounded or ovoid vs rectangular>/

1. round or ovoid/

2. noticeably rectangular or rhomboidal, with rounded "corners"/

#73. <Dorsal cirri proportions>/

1. circular to ovoid, as wide or wider than long/

2. elongate, longer than maximum width/

#74. <Dorsal cirri shape>/

1. evenly rounded, without an obvious tip or corner/

2. lanceolate or rectangular, with a distinct tip or corner/

#75. <Dorsal cirri symmetry>/

1. symmetrical/

2. asymmetrical/

#76. Neuropodium <divided or entire>/

1. divided into dorsal and ventral lobes/

2. entire/

Most phyllodocids have a neuropodium that divided into dorsal and ventral lobes (which may be equal or unequal in size; this information is recorded in the next character). One species of Eumida recorded in this database has an entire neuropodium without any evidence of division.

#77. <Relative size of> dorsal part <of neuropodial lobe>/

1. similar in size and shape to ventral lobe/

2. larger than ventral lobe/

The relative proportions of the dorsal and ventral neuropodial lobes separate some genera, while other genera are polymorphic for this character (Pleijel, 1991: 226–229).

#78. Ventral cirri <orientation>/

1. horizontally oriented/

2. obliquely oriented/

The ventral cirri in most phyllodocids is oriented horizontally, i.e. with long axis parallel with the long axis of the neuropodium. However, 4 genera have the long axis of the ventral cirri distinctly oblique (pointing up) compared with the neuropodium (Pleijel, 1991: 226–229).

#79. <Relative length of ventral cirri>/

1. longer than chaetigerous lobes/

2. about as long as chaetigerous lobes/

3. shorter than chaetigerous lobes/

The relative length of the ventral cirri compared with the neuropodial lobe may be a useful species level character.

#80. <Presence of extented tips on ventral cirri>/

1. without extended tips/

2. with extended tips/

Some species of Phyllodoce possess ventral cirri with distinctly extended and narrow tips, and this character is a useful specific character in that genus.

#81. Chaetae <comments; TE>/

Chaetal morphology appears to vary little among phyllodocid genera and species (although Eibye-Jacobsen (1991) provides some exceptions). Because of this, and because the difficulty in making comparable observations of phyllodocid chaetae is greater than in many other polychaete families, little use is made of phyllodocid chaetal morphology in this interactive key. Any useful descriptions of chaetae may be entered here as text.

#82. Anal cirri <shape>/

1. globular to digitiform with rounded tips/

2. long, cirriform, tapering to a fine pointed tip/

The presence of long, cirriform anal cirri tapering to a fine tip readily distinguishes the genus Hypereteone (as defined by Wilson, 1988) from all other phyllodocids. Pleijel (1991) showed that Hypereteone as so defined leaves Eteone as a paraphyletic taxon not defined by a derived character state, but the form of the anal cirri remains a useful character if entire specimens are available.

#83. <Remarks - other comments on this taxon; TE>/

General comments on this species, including ecological or biological notes, may be included here as text.

#84. <Remarks - comments on related taxa; TE>/

Comparative notes on related taxa and the characters that best distinguish similar species may be included here as text.