BULLETIN OF MARINE SCIENCE, 60(2): 405-426, 1997

PATTERNS OF GROWTH AND SETAL DEVELOPMENT IN THE

DEEP-SEA WORM, STREPTERNOS DIDYMOPYTON
(POLYCHAETA: CHRYSOPETALIDAE)

Charlotte Watson Russell

ABSTRACT
A number of post-larval and juvenile chrysopetalid polyehaetes were made available for
study from experimental wooden panels collected in deep waters of the western Atlantic
Ocean. Benthic post-larvae of 10 to II segments possess a large rounded prostomium with
eye spots, two pairs of larval dorsal and ventral cirri and cilia tufts associated with sites
where notopodia are developing on segment 2. Juveniles of 12 to 30 segments exhibit reduc-
tion in size of prostomium; gradual loss of eye pigment; loss of larval cirri; development of
notopodia of segment 2 and subsequent torsion of neuropodia of segment 2 dorsally. Juve-
niles display distinctive petaloid notosetal paleae and notosetal patterns unlike the narrow,
pointed paleae and notosetal patterns observed in adults. Observed setal change is linked to
hypothetical life strategies of Strepternos didyrnopyton. Possible origins and function of
paleate setal change is discussed.

Chrysopetalid polychaetes belonging to a recently described new genus and

species, Strepternos didymopyton Watson Russell in Bhaud and Cazaux 1987,
Watson Russell 1991, were collected from experimental wooden blocks in
abyssal depths (1,800-3,995 m) in Atlantic waters off Massachusetts and
Virginia and the Great Bahama Bank and Virgin Islands (Caribbean Sea).
Numerous post-larval, juvenile and adult specimens, covering a wide size range,
were found among a rich macrofauna associated with the burrows of wood-
boring xylophagid bivalve molluscs in wooden blocks submerged on the sea
floor and retrieved after 1-3 years by DSRV ALVIN(Turner 1973, 1978, 1981).
Development of the anterior end and ontogenetic setal development, involving
setal initiation, replacement and loss, is documented for individuals between the
10 and 30 segment growth phase.
Clearly, wood island environments have persisted over sufficient time to allow
evolution of unique genera and species (Turner, 1981). However, dispersal in space
and time of sources of wild wood in the deep sea is presumably patchy and indi-
vidual wood islands offer only a transient refuge; larvae of invertebrate coloniz-
ers of these islands would have to be capable of seeking new sites in the longer
term. It is hypothesized that the abyssal dwelling benthic polychaete, S. didymopy-
ton, possesses pelagic larvae capable of dispersal along surface and bottom cur-
rents in waters adjacent to North American and Caribbean island coasts. This hy-
pothesis is based on: (1) recruitment patterns and life history notes derived from
observations of morphology of post-larval, juvenile and adult S. didymopyton
specimens; (2) comparison of S. didymopyton post-larvae with other known plank-
tonic chrysopetalid larvae, including Mileikovsky's (1962) description of chryso-
petalid larvae collected from the surface waters of the Gulf Stream; (3) observa-
tions on the origin and function of chrysopetalid paleate setae.

MATERIALS AND MErnoDs

Larval setal terminology follows that described by Watson Russell in Bhaud and Cazaux, 1987. Adult
setal terminology is based on that in Watson Russell, 1991. Details of all station data referred to in
this paper are from Watson Russell, 1991. See Appendix, this paper, for abbreviations on Figures.

405
406 BULLETIN OF MARINE SC1ENCE, VOL. 60, NO.2, 1997

Family Chrysopetalidae Ehlers, 1864

Strepternos Watson Russell in Bhaud and Cazaux, 1987; Watson Russell, 1991:286, figs. 1-4.
Type Species.-Strepternos didymopyton Watson Russell in Bhaud and Cazaux, 1987:663, fig, 28.1-3,

Generic Diagnosis,-Chrysopetalids with broadened paleae and very small pro-

stomium compressed between anterior segments, eyes absent in adults; segment
2 with small setose notopodia and dorsal cirri, setose neuropodia without ventral
cirri; palaea of middle segments arranged in 1 lateral and 2 dorsal fascicles, me-
dian fascicle absent.

Description of Adult.-Body long, rectangular; less than 100 segments. Golden,

notosetal paleal fans imbricate down back. Prostomium with median antenna, lat-
eral antennae, palps; proboscis eversible, with 2 transparent stylets. Nuchal fold
small, semicircular, (difficult to see in larger adults). Segment 1 very reduced,
fused to prostomium with 2 pairs of dorsal and ventral tentacular cirri. Segment 2
small, partially fused to segment 1. All setose segments except first, with paleae
of 2 types, dorsal and ventral cirri, and falcigerous neurosetae (Fig. 5A). Interra-
mal region ciliated. Notopodia each with 1-4 narrow, pointed, lateral paleae with
0-6 ribs and main paleae in lateral and medial fans transversely orientated; lat-
eral fan partly overlapping medial fan. Mid-body segments with 12-15 lateral
main paleae with 11-16 ribs; 15-21 medial main paleae with 12-18 ribs. Adult
main paleae long, slender, symmetrical and pointed, with serrated margins dis-
tally; 1-4 slightly raised and very weakly serrated ribs often present. Dorsal cirri
broad, retractile within large, elongate cirrophores. Neuropodia with subacicular
fascicle of heterogomph, serrate neurosetae. Segments 2 and 3 with all and some
spinigers, respectively. Subsequent segments with superior, middle group and in-
ferior falcigers. Pygidium composed of dorsal rounded structure with very small,
anal papillae on lateral posterior margin (often not visible) and broad, glandular
ventral cone.

Biology.-Mature sexual gametes were difficult to recognize in specimens under

30 segments. Specimens up to this stage retain larval setal types and for the pur-
poses of this paper are included within the juvenile category. Ovigerous females,
(40-70 segments) were often observed with large, rosy to dark brown pigmented
dorsal cirrophores and pale pink colored eggs in parapodia (Fig. lC). Eggs from
8 ovigerous females had a size range of 0.03-0.08 mm (Figs IG, H). The size
range of these eggs is similar to those described for a littoral chrysopetalid spe-
cies, Chrysopetalum fragile (=debile), 0.027-0.05 mm, by Ehlers, 1864. An
ovigerous female of 62 segments was dissected segment by segment, mounted
and setal types noted. Dispersement and numbers of eggs throughout the body
was similarly traced (however some eggs may have been lost during dissection).
Eggs, with a fine granular appearance, were observed between segments 21-51.
Mid-body segments (21-35) had smaller numbers (4-20 per parapodium), of
larger-sized eggs, 0.06-0.075 mm (Fig. lA, B). Segments 36-51 had larger num-
bers (25-60 per parapodium), of smaller-sized eggs, 0.03-0.04 mm. Smaller size
eggs were observed from mid-body segments in other specimens (e.g., segment
23 of 49 segment individual, Fig. 1C).
Parapodial changes during gametogenesis include: 1. elongation of, and
glandular activity within the cirrophore of the dorsal cirrus; surface of cirrophore
often appeared crazed and in some cases appeared about to delaminate (Fig.
lA-C); 2. long, dense cilia appear in the inter-ramal areas of anterior segments
6-26 and thereafter are evident only on the superior edge of the ventral rami (Fig.
 RUSSELL: DEVELOPMENT OF STR£PT£RNOS 407

/'

E
~
00 I
0\9 G $
~
.~~.
Figure L Strepternos didymopyton. A-B, 62 segmented female: A, Parapodium from segment 21,
anterior view: B, Parapodium from segment 26, anterior view; C, 49 segmented female: parapodium
from segment 23; 0, 61 segmented male: parapodium from mid-body segment, anterior view, inset of
sperm? from same; E, sperm? from "packet" associated with male of 40 segments; F-G, egg struc-
tures associated with same 'packet'; H, egg from mid-body parapodium of 53 segmented female. Figs
A-D: Acicula dashed, gonads dotted. Abbreviations: c. cilia, cr. crazing, g. glandular activity, p. pig-
mentation, t.r. tube remnants. Scales = 0.4 mm for A-D; 0 inset x 1,000; 0.02 mm for E; 0.05 mm
for F; 0.1 mm for G-H.
408 BULLETIN OF MARINE SCIENCE. VOL. 60, NO.2. 1997

A B

Figure 2. Strepternos didymopyton. A, Ventral view, B, Dorsal view, post-larva of 10 segmt:nts; C,

Ventral view, juvenile of 12 segments: paleae only drawn on posterior segment. Scales = 0.3 mm for
A-B; 0.4 mm for C.

lA-C); in a number of segments eggs, and very small rounded structures with a
dark nucleus were observed attached to these cilia (Fig. IB); 3. gonaducts of the
type described for chrysopetalids by Fage, 1906 are swollen and golden coloured
in segments 16-51 (Fig. lA); 4. the tip of the neuropodium becomes swollen and
enlarged particularly in anterior segments, with glandular activity evident along
its superior edge (Fig. lA); glandular activity is also apparent in a pronounced
swelling posterior to the neuropodial tip (Fig. lA-D).
In several samples Strepternos individuals were found encased in the membra-
nous remains of tubes, identified as the inner periostracallining of the white tube
 RUSSELL: DEVELOPMENT OF STREPTERNOS 409

c Ide
,
Ive
Ide B

Ive
E

Figure 3. Strepternos didymopyton. A-B, Anterior end of 10 segmented post-larva, A, Dorsal view,
B, Ventral view; C, Anterior end of 10 segmented post-larva, dorsal view; D, Anterior end of 11 seg-
mented post-larva, dorsal view; E, enlargement of developing notopodium 2 and notopodium 3 from
D. Scales = 0.1 mm for A-D, 0.04 mm for E.

of the teredo Xyloredo ingolfia (Turner in lit). In one sample, two adults were
found dorsum to dorsum inside a nearly intact tube. Another specimen, partially
encased in a tube, had a long white "packet" lying along its dorsum. The bulk of
this substance was composed of a fine tangle of unrecognizable material in which
were caught larger eggs with discernible nuclei and smaller rounded structures
composed of a coarse granular material, some with minute protrusions (Fig. IG).
Small ovoid, stalked structures filled with tiny rounded granules (Fig. IF) were
also apparent; similar structures were often found among extruded material from
parapodia of obviously ovigerous females. These structures are possibly the same
as those described by Ehlers (1864:88) for the immature eggs of Chrysopetalum
410 BULLETIN OF MARINE SCIENCE, VOL. 60. NO.2, 1997

B dtcI

nf

c
I
!

mo
Figure 4. Strepternos didymopyto/l. A, Anterior end of 12 segmented juvenile, dorsal view; B, Ante-
rior end of 14 segmented juvenile, dorsal view; C, Ventral view of same. Scales = 0.1 mm for A-C.
 RUSSELL: DEVELOPMENT OF STREPTERNOS 411

,/

sf

B sut

me.m

Figure 5. Strepternos didymopyton. A, Parapodium from segment 24 of 62 segmented adult with

enlarged setal types, anterior view; B, Noto and neuropodium from segment 5 of 10 segmented post-
larva with enlarged setal types, anterior view. Scales = 0.2 mm for A, 0.04 mm for enlarged setae;
0.04 mm for B.
412 BULLETIN OF MARINE SCIENCE. VOL. 60. NO.2. 1997

Table 1. Strepternos didymopyton. Measurements in mm of benthic post-larvae and juveniles from

10 to 30 segments.

No. of Length single Width single No. of individual Mean Mean

segments measurements measurements measurements length width

10 0.60, 0.64, 0.66 0.54, 0.64, 0.64 3 0.63 0.60

11 0.74,0.74,0.76,0.80 0.72, 0.80, 0.80, 0.60 4 0.76 0.73
12 0.74, 0.78, 0.86, 0.90, 0.52,0.68,0.76,0.64, 8 0.89 0.71
0.92, 0.96, 1.00, 1.00 0.78, 0.84, 0.80, 0.64
13 0.86, 0.92, 1.06, 1.12 0.64, 0.74, 0.92, 0.84 4 0.99 0.79
14 1.08, 1.20, 1.24 0.88, 0.80, 0.80 3 1.17 0.83
15 1.16, 1.24, 1.36, 1.42 0.90, 0.74, 0.94, 1.00 4 1.30 0.90
16 1.16, 1.34 0.86, 1.00 2 1.25 0.93
19 1.80 1.00 1 1.80 1.00
21 1.70 1.10 1 1.70 1.10
23 2.55 1.00 1 2.55 1.00
26 2.75 1.40 1 2.75 1.40
27 2.70 1.60 1 2.70 1.60

fragile (=debile) "enclosed in small sacs which project from the inside surface of
the ventral wall."
Free floating tiny rounded spheres, many with minute swollen poles, were also
examined from the individual associated with this packet and from the milky col-
ored parapodia of a number of other specimens (Fig. IE). The shape of these
structures resemble the primitive type of spermatozoon as described by Franzen
(1970). However no distinct flagellum was observed on these "sperm" and there
are no descriptions of chrysopetalid sperm in the literature.
One mature, spent-looking specimen of 61 segments was observed with an ir-
ridescent white, "loose" integument; paleal notosetae were compressed into nar-
row laterally held fans leaving the dorsum fully exposed and some setal types
were starting to fall out. Parapodia were packed full of a milky colored granular
substance which appeared under high magnification as very small rounded
spheres, some with minute poles; remnants of teredo tubes were also evident
around the parapodia (Fig. ID and inset).
The relatively large size of Strepternos individuals, the evidence of separate
sexes, the accumulation of large numbers of sexual gametes in the parapodia, the
possible primitive type of sperm, the small quantity of yolk in the egg and the
small size of the eggs (at the bottom of the range for oocyte diameter observed in
the Polychaeta (Olive, 1985», suggests development leading to ciliated pelagic
larvae.

JUVENILE MORPHOLOGY

Post-larval Description.-Tables 1-3. Figs 2,3, 5B. Specimens in this stage have
10 to 11 segments.
Body broad, ovoid. Transparent, primary, paleal notosetal fans covering dor-
sum; adult notosetal types absent. Ventral rami with conspicuous pocket on latero-
inferior edge; slender spinigerous and falcigerous neurosetae extending beyond
notopodia (Fig. 2A, B). Mouth opening ventrally at level of segment 4; pharynx
yellow, muscular; gut with yellowish brown globules. Prostomium comparatively
large, round, discreet, without antennae or small, rounded median antenna and
two filiform lateral antennae originating anteriorly; 2-3 pairs of red granular eye
spots mid prastomium; two ovoid palps inserting antera-ventrally; nuchal fold not
discernible on post-larval specimens with large prastomia (Fig. 3A, C); visible,
posterior to small prostomia, on more developed specimens (Fig. 3D). Segment 1
 RUSSELL: DEVELOPMENT OF STREPTERNOS 413

Table 2. Strepternos didymopyton. Comparison of setal counts within a 10 segment individual.

Segment number 2- 4 9 10

Notosetal characteristics
Total no. paleae in
notopodium 0 8 10-11 10-12 10-11 11-14 8
Lateral paleae no. 0 1 1 2 1 1 1
No. of ribs 0 5-6 6-8 5-6 6-8 6-8 5-6
Sub-unit I paleae no. 0 2 2 2 2 2 2
No. of ribs 0 9-10 9-12 13 11-12 9-13 12
Main paleae:
Lateral main no. 0 0 0 3 3 3 2
No. of ribs 0 0 0 12-15 13-15 15-16 15-16
Middle main no. 0 4 5 3 3 6 2
No. of ribs 0 12-15 13-17 16 15-16 17-19 17
Medial main no. 0 1 2 2 2 2 1
No. of ribs 0 17 18 17 16-17 17 16-17
Neurosetal characteristics
Spinigers 7 4 0 0 0 0 0
Superior falcigers 0 3 2-3 2 2 4 2
Middle falcigers 0 5 8 8 7 4
2 shafts
Inferior falcigers 0 4 4 4 4 2-4
'"notopodia of segment 2 not formed.

with two pairs of long, slender, subequal, larval tentacular cirri, inserting lateral
to prostomium (Fig. 3A-E); also with pair of adult ventral tentacular cirri situ-
ated adjacent to acirrose neuropodia 2 (Fig. 3B). Notopodia of segment 2 first
visible as ciliate lobe (Fig. 3A); later as bud-like structure, present postero-Iaterally
to prostomium (Fig. 3C).
Notosetae of mid-body segments composed of lateral, subunit 1 and main paleae
types in 1 fan (Fig. 5B). Lateral subacicular group of 1 slender palea (5-7 ribs),
margin serration only present distally, and 1 broader palea (8-10 ribs); raised ribs
or granules absent. Supra-acicular group with single subunit 1 palea between lat-
Table 3. Stepternos didymopyton. Position of notosetal types within the body and comparison of no-
tosetal patterns with increasing segment size across a number of individuals.

Total segment
no. of individuals 10 11 12 14 19 26 27 52 61

Notopodia of seg. 2:
No. of paleae Ot 1-2 2 1-2 2 2 2-5
present
Notosetal larval
spines:
Present on segs. 3-5 3-5 3-5 3-5 3-5 3-5 3-5 0 0
Larval main paleae:
Present on segs. 3-10 3-9, 3-9, 3-8, 3-4 3-4 3 0 0
lO,ll lO,ll 9,10
Adult rod-like lateral
paleae:
Present on segs. Ot II 10-12 * * 13-26 8,9,18, 42-52 56-61
20-27
Adult main paleae:
Present on segs. Ot 10-11 4,6-12 3-14 3-19 3-26 3-27 3-52 3-61
t not formed.
* not counted.
414 BULLETIN OF MARINE SCIENCE. VOL. 60. NO.2, 1997

eraI and main groups with 10-13 ribs, 2 raised ribs; main paleae group with 2
lateral group paleae relatively narrower, with 12-15 ribs; 3 broader. middle group
paleae, with 16-18 ribs; 2 broader, medial group paleae, distally rounded, with
18-20 ribs. All main paleae with entirely serrate margins ending distally in
rounded, shallow apex. Numerous finely serrate, raised ribs and small raised gran-
ules present on superior paleae surface. Notopodia on segments 3-5 with short
larval spines adjacent to lateral main paleae. Neurosetal compound spinigers on
neuropodia of segment 2 and some on neuropodia 3; following neuropodia pos-
sess falcigers with slender, medium to short length blades. Serrate larval spine
overlies neuroaciculum on all setigers (Is: Fig. 5B). Subulate ventral cirrus present.
Pygidium small, conical, terminal protruberance (Fig. 2A, C).
Juvenile Description.-Tables 1,3. Figs. 4, 6, 7. Specimens in this stage have 12
to 30 segments.
Body elongate-ovate to rectangular. Primary paleae and narrow, adult notosetal
fans covering dorsum (Fig. 2B). Prostomium smaller, retracted into anterior seg-
ments with yellow-red eye pigment patches on posterior area, overlain in most
part by dermis of segments 2 and 3 (Figs. 3D, 4A, B). Pharynx with pair of stylets
visible; gut filled with brown material. Segment 1 with larval tentacular cirri pair
shed at 11-12 segment growth stage; dorsal pair observed in lateral position ad-
jacent to developing notopodia of segment 2 (Fig. 3D, E). Dorsal larval cirri re-
placed by pair of short, broad, tentacular dorsal cirri with large ceratophores, (Fig.
4A-C). Adult ventral tentacular cirri pair move closer towards palps with con-
traction of prostomium (Fig. 4C). Segment 2 with notopodia developing acicula
(Fig. 3D, E), pair of short, dorsal cirri, few short, slender paleae (Fig. 4A-C).
Notopodia of segment 2 fused in part to notopodia of segment 1.
Single primary paleal fan replaced by double adult paleal fans. Figures of no-
topodia of segments 3, 6, 9, 10, 11 from a 13 segmented individual illustrates this
process (Fig. 6A-E). Notosetae of notopodia 3 with all primary, larval paleae
types, numbering 12, and 2 larval spines (Fig. 6A). Notopodia 6 with larval lat-
eral, sub-unit 1 palea and main paleae, numbering 10; 3 narrow adult paleae
present in lateral position of main group (Fig. 6B). Notopodia 9 with larval lat-
eral, sub-unit 1 paleae and main paleae types, numbering 7; 7 adult main paleae
in lateral and middle positions, plus 1 adult lateral palea (Fig. 6C). Notopodia 10
shows loss of larval lateral, sub-unit 1 palea and all main paleae types except for
1 palea in medial-most position; 1 adult lateral and 12 main paleae types estab-
lished (Fig. 6D). Notopodia 11 without larval paleal types; adult paleal types in-
clude a single, very slender outer-most lateral palea (typically seen in posterior
body of adult), plus 1 other narrow lateral pa1ea; prototype two-part adult main
fan composed of 3 lateral main and 7 medial main paleae; lateral part of medial
fan includes slightly misshaped palea (amm:Fig. 6D, E) seen in a similar position
in adult medial fan (Fig. 5A). Neuropodia with neurosetae comprising spinigers
in segments 2-4. Adult short, broad-bladed falcigers appearing on neuropodia 5-6.
Long bladed superior falcigers absent after neuropodia 7. Narrow bladed sub-unit
1 neuroseta present in mid-superior position in posterior-most segments.

Additional Morphological Comments

Anterior End.-Under high magnification (X400-1,OOO)the edges of the prosto-
mium of S. didymopyton post-larvae are crenulate (Fig. 3 A-C). This may be an
artifact of preservation or perhaps represents the contraction of tissue during re-
absorption of the larval episphere.
The post-larval segment 1 has two pairs of long larval tentacular cirri (dorsal
 RUSSELL: DEVELOPMENT OF STREPTERNOS 415

D
amm I
I
I

suI
,/
E
,/
,/
amm
I
I
/

c //
W
/
su I
/
/

...- I

- -al

Figure 6. Strepternos didymopytoll. A, Notopodium from segment 3; B, segment 6; C, segment 9; D,

segment 10; E, segment 11; all figures in anterior view from 13 segmented juvenile. Scales = 0.04
mm for A-E.

longer than ventral) and one pair of adult ventral tentacular cirri; at this stage
adult tentacular dorsal cirri have not yet formed. The notopodia of segment 2 ap-
pear as 2 ciliate bud-like structures. Notopodial generation at this developmental
stage after the larva has settled on the benthos has not been previously documented
in chrysopetalid polychaetes.
Adult cephalization and development of segment I and the notopodia of seg-
ment 2 takes place over the II to 15 segment growth phase. The prostomium
contracts, the larval cirri pair are lost and the adult dorsal tentacular cirri pair of
segment 1 formed. Dorsal tentacular larval cirri of varying lengths were observed
416 BULLETIN OF MARINE SCIENCE, VOL. 60, NO.2, 1997

B
1m
am-- - 1

Is

me.m

Figure 7. Strepternos didymopyton. A, Anterior end of 26 segmented juvenile; B, Notopodium of

segment 3 from 27 segmented juvenile, anterior view. Scales = 0.2 mm for A, 0.04 mm for B.

in 2 individuals of 10 segments from the same site (Figs. 3A, C), Larval dorsal
cirri are always much longer and more slender than the short, broad, adult dorsal
cirri which insert on large, broad ceratophores (Figs 3E, 4A-C). Adult dorsal cirri
of notopodia 2 are very similar in shape to dorsal tentacular cirri of segment 1
 RUSSELL: DEVELOPMENT OF STREPTERNOS 417

and the former often obscure the latter (Figs. 4A-C, 7A). Development of no-
topodia of segment 2 involves the sequential formation of ramus, notoaciculum,
dorsal cirrus and paleal setae. These notopodia are comparatively smaller than
following notopodia and remain so throughout life.
The ventral tentacular cirri of segment 1, which are formed before the dorsal
tentacular pair, are situated close to the base of neuropodia 2 (Fig. 3B). The space
observed in ventral view between the ventral tentacular cirri and the prostomium
is closed as the prostomium shrinks and the development of notopodia 1 and 2
brings the ventral tentacular cirri into a position immediately adjacent to the palps.
The acirrose neuropodia 2, formerly visible in ventral view (Fig. 3B), are then
twisted into a more dorsal position (Figs. 4C, 7A).
Eye spots appear to coalesce and, with contraction of the prostomium, are seen
sub-dermally in individuals of 13 to 30 segments (Figs. 3D, 4A, B). By 18 seg-
ments the anterior end has broadened substantially and the prostomium withdrawn
between the anterior-most segments. A late juvenile of 26 segments illustrates the
increased width of the anterior end and the extremely small size of the prosto-
mium relative to the rest of the body (Fig. 7A). Yellowish-orange eye pigment
patches and a small nuchal fold are still evident in this individual. With growth
(40 segments and over) the increasing retraction of the prostomium within the
anterior segments makes the prostomium and nuchal fold almost impossible to
distinguish. The absence of eye pigment was observed when a number of anterior
ends of adult specimens were dissected (Watson Russell, 1991: 287, Fig. 2A-C).

Setal Patterns.-Newly settled post-larvae display primary, petaloid shaped paleae

in notopodia down the body. With growth these paleae are gradually lost and re-
placed by an adult lateral paleal fascicle and a main paleal fan composed of dis-
tinct lateral and medial fasicles. The narrow adult main paleae are initiated from
a lateral position within the notosetal fascicle of notopodia located in midbody to
posterior segments within the individual (Fig. 6B). With the help of a "marker"
pa1ea, such as the adult misshapen main palea, it can be determined that the adult
medial main fan is formed substantially before the adult lateral main fan (Figs.
6C-E, SA).
A continuum of all paleae types is present in individuals from 12 segments on
and progressively over the 10 to 30 segment growth phase (Tables 2, 3). As in-
dividuals reach 15 segments narrow adult main paleae types predominate in most
segments throughout the body, although anterior-most notopodia retain all or some
primary main paleae and larval spines (see segment 3 of a 27 segmented speci-
men, Fig. 7B). .
Observation of setal types along the body of a 27 segmented specimen reveal
that larval lateral and sub-unit 1 paleae types are also present from anterior to
mid-body setigers, with adult lateral and sub-unit 1 paleae types overlapping (Fig.
6A, B). Details include larval lateral paleae (8-10 ribs) and larval sub-unit 1 paleae
both present from segments 3-15. Adult sub-unit 1 paleae, very slender and
pointed, are present from segments 15-27 (Watson Russell, 1991: Fig. 4W, X).
Broader adult lateral paleae (6-8 ribs) are present from segments 8-12 and nar-
rower adult lateral paleae (2-3 ribs) from segments 11-26; rod-like adult lateral
type (0-2 ribs) can be found on segments 8, 9, 18 and 20-27. Larval and adult
lateral types can be further distinguished from each other by the greater degree of
serration of the paleae margin of the former.
Material examined demonstrated that loss of larval paleae and acquisition of
adult paleae are not always synchronous on both notopodia of the same segment
within or between similar sized individuals. The formation of adult paleae types
418 BULLETIN OF MARINE SCIENCE, VOL. 60, NO.2, 1997

is not always sequential e,g., the rod-like adult lateral paleae, usually located in
the posterior segments of adults, were found on a number of unsequential seg-
ments in the 27 segmented specimen discussed above. Setal asymmetry was of-
ten noted during setal change over in specimens. Two examples are illustrated:
Figure 4A shows I adult palea on the right dorsal side of developing notopodium
2 and no palea on the left side of a 12 segmented individual; Figure 7A shows a
26 segmented specimen with retention of primary paleae on the left dorsal no-
topodium 3 and loss of the same on the right notopodium 3,
Individuals between 30 and 40 segments were rarely seen in the samples. All
individuals over 40 segments exhibited loss of larval spines and possessed only
adult paleal types (Table 3).

DISCUSSION

Recruitment and Possible Modes of Larval Dispersal of Streptemos.-Data on

relative numbers of juveniles and adults from 11 deep sea wood island stations,
established from 1-3 years, showed that 5 stations had only juveniles, 2 stations
had more juveniles than adults, and 4 stations more adults than juveniles. Four of
the five stations with only juveniles had only 1-3 individuals; the other station
had 43. Stations with more juveniles than adults had relatively low numbers over-
all and stations of the third category always had many more adults than juveniles.
The two wood island stations from the Bahama Islands, submerged for a year,
had only Strepternos juveniles. Station T37 had 43 individuals, the majority of
which were 10-15 segment size with a few more advanced juveniles; Station T56
had 3 juveniles. The absence of sexually mature adults implies that the wood pan-
els have been colonized solely by larvae.
Three Stations (N28, N34, N35), all with larger numbers of adults than juve-
niles, had been submerged for 2 to 3 years. Station N27, with similar numbers of
juveniles and adults, had been submerged for one year, and Turner (1978: 16)
also reported 'crysoptellid' (=Strepternos) worms in the burrows of dead
Xylophaga from stations submerged for one year from sites off north Virginia and
the Bahamas. If wood panels are colonized solely by larvae, sexual maturity must
be attained within a year.
Research on larval development and dispersal at deep-sea wood islands and hy-
drothermal vents reveal a number of a reproductive strategies, including high dis-
persal planktotrophic or low dispersal lecithotrophic, but free swimming, modes
(Turner, 1981; Lutz et aI., 1984). The former mode of development has been in-
dicated for a number of molluscan and crustacean species but what information
there is on polychaete species (ampharetid spp.) suggests the latter type (Zottoli,
1982, 1983; McHugh and Tunnicliffe, 1994).
The method of larval dispersal in Strepternos is unknown but the biology of
these adults is similar to that seen in a number of other chrysopetalid genera with
planktonic larvae (author, personal observations). The morphology of S, didymopy-
ton benthic post-larvae is also very similar to that described for middle to late
nectochaete chrysopetalid pelagic larvae (Mileikovsky, 1962, pI. 1, figs. A, B;
Cazaux, 1968, pI. 18, figs 8-10; Blake, 1975, fig. 6B) and late nectochaete
chrysopetalid benthic larvae (Watson Russell in Bhaud and Cazaux, 1987). Char-
acteristics observed in S. didymopyton post-larvae that point to an earlier plank-
tonic existence include: a small, ovoid body shape with long, slender neurosetae
that increases surface area and floating capacities; oil globules in the gut that in-
creases volume and buoyancy; large prostomia with eye pigment and long larval
tentacular cirri that help a larvae to orientate mid-water; presence of short, ser-
 RUSSELL: DEVELOPMENT OF STREPTERNOS 419

rated larval spines of the notopodia of segments 3-5, perhaps indicative of the
former site of long, provisional larval spines, which further increase floating and
defensive capacities (see Mileikovsky's larva, Fig. 8A, B).
Strepternos setal types, in common with all chrysopetalid setae, have an inter-
nal construction of a series of longitudinal large channels divided into chambers
by horizontal septa (Westheide and Watson Russell, 1992). This allows for a broad
area to remain light weight while retaining strength and rigidity. Primary paleae,
belonging to Strepternos post-larvae, are further reinforced with a fine external
sculpture of raised ribs, denticles and tubercules, ideally suited for planktonic ex-
istence. Paleal types belonging to Strepternos benthic adults are slender with re-
duced ornamentation. A parallel situation is seen in shell ornamentation between
veliger planktonic larvae (with light, finely ribbed and ornamented shells) and
benthic adults (with thick, heavy, barely ornamented shells) belonging to a num-
ber of deep sea mollusc species (Bouchet and Waren, 1979).
The oil globules observed in the gut of Strepternos larvae are indicative of a
non feeding larvae but one capable of staying in the plankton for an extended
period. It has been established that larvae of a chrysopetalid littoral species from
coastal waters can spend up to 3 weeks in the plankton at the 6 segment stage.
These chrysopetalid larvae, observed in the laboratory, possessed dense oil drop-
lets in the gut and did not feed but lived off their reserves (Cazaux, 1968). Oil
globules have also been observed in planktonic and recently settled larvae be-
longing to chrysopetalid species of Paleanotus (Blake, 1975) and n. gen. 1 (Watson
Russell in Bhaud and Cazaux, 1987: 669, pI. 28, fig. 4, and author, unpublished
information).
It may be that Strepternos larvae are confined to a passive, low dispersal mode,
using bottom currents and travelling step-wise between wood island refuges. How-
ever it appears likely to me that these larvae, like their littoral relatives, have the
ability to travel longer distances within the photic zone of the plankton. It is hy-
pothesized that Strepternos larvae have the capacity to rise from abyssal depths
through the plankton to the surface waters of the Gulf Stream. Orientated 'move-
ment in the vertical plane in the absence of light has been documented in the
larvae of other invertebrates (Sulkin, 1990). Eye development would then allow
for positive phototactic response and orientation in well lit oceanic waters.
The metatrochophore larva would grow larger while living off its reserves and
be passively transported by currents, possibly over great distances. The cues which
prompt a tiny larva of 6-7 segments and less than 1 mm in length to descend
through 3-4,000 m of water is no doubt related to impending metamorphosis.
Signs of this change may include the increasing inefficiency of the larva to stay
bouyant and float due to its larger body size in comparison to its provisional se-
tae size and diminuition of its oil reserves. After descent, the larva would spend
some time being transported by bottom water currents before a suitable benthic
site was found. Whether a wood substrate alone triggers settlement and the final
stages of metamorphosis, is unknown. If the adults are reliant on scavenging teredo
products and finding protection in deceased teredo tubes it is likely that a wood
substrate is preferred. On available data all sampled sites, except 2 (recorded as
'no data'), have been from experimental or wild wood islands in the deep-sea.
Literature on larval development in other paleate chrysopetalid species indicates
that folded paleal fans are formed in the 4-5 segmented metatroch pelagic larvae.
Why do the paleal fans appear at this time? As precursors to an impending benthic
existence. These precursor or primary paleae are ovate and flattened with raised
ribs and spines, well suited to a planktonic life. Paleal fans expand across the
dorsum in the late metatroch/nectochaete larvae prior to metamorphosis and
420 BULLETIN OF MARINE SCIENCE, VOL. 60, NO.2, 1997

benthic settlement. It is at this 6-7 segmented stage that larvae can prolong their
stay in the plankton while searching for a suitable benthic site. Primary paleae
are starting to be replaced by adult paleal types initiated from posterior segments
and the long provisional setae of the anterior segments lost during this ongoing
metamorphic period, either in the plankton prior to or most likely at the actual
point of settlement. The change from larval to adult paleal types is related to the
change in habitat and the need for protection of the anterior end and distal ex-
tremities, now particularly vulnerable in a new benthic environment.
Observations of Strepternos benthic post-larvae of 10-13 segments show a range
of larval and adult characteristics. Some 10 segment specimens have only pri-
mary setae, anterior end larval features and oil droplets in the gut. Other 10 seg-
ment individuals show change in the anterior end, initiation of adult setal types
and brown material in the gut. Some 13 segment individuals possess nearly all
adult setal types, while some very few. This leads me to suspect that in this spe-
cies adult characteristics are not iniated until the pelagic larva has successfully
completed benthic settlement and starts to feed. If this is correct one could esti-
mate from the degree of morphological development, the segment size that
Strepternos larvae settle out from the plankton. This could be reasonably placed
at between 9 and 11 segments.
Cold ambient bottom water environments have been indicated in the lowering
of developmental rates thereby possibly allowing non-planktotrophic larvae to
remain in the plankton for longer periods and disperse widely (Lutz et al., 1984).
A temperature related reduction in metabolic rate has been suggested as a means
of prolonging the dispersive stage of an ampharetid polychaete present at
hydrothermal vents (Zottoli, 1983). Larval notosetal paleae of a 10 segmented
Strepternos individual are all of a similar type and structure, indicating their
origin in the same milieu. Whether larval paleae have originated in warmer or
colder waters is open to investigation. It could be fruitful to compare isotope
measurements between larval and adult paleae along the lines taken by Bouchet
and Fontes (1981) in their study of the larval and adult shells of deep-sea
mollusc species.
The comparatively greater segment size at which Strepternos larvae settle out
(in comparison to other chrysopetalid larvae), plus the retention of all larval setal
types, may well be related to the need to stay in the plankton for a longer period
and traverse a greater area during the search for a suitable benthic site. I think the
late development of the notopodia of segment 2 (after settlement) is indicative of
the need of the pelagic larvae to retain the long provisional spinose setae of the
anterior larval segments for as long as possible during this search. The initiation
on notopodia 2 of adult paleae, without any precursor larval paleae, further points
to the switch over from pelagic to benthic life. The relatively slow acquisition of
adult setal types, observed in S. didymopyton, is not necessarily related to the
cold water factor at abyssal benthic sites. "Shawania" riveti, a large, littoral dwell-
ing chrysopetalid with some affinities with S. didymopyton, retains certain larval
setal types up to 50 segments in length. Unfortunately nothing is known about
the early development of the former species.
Adult morphology (including diminuition of prostomium, loss of functional eye
pigment) and habitat (gametogenic adults found in situ in xylophagid tubes) sug-
gests that Strepternos adults do not move once established within a teredo com-
munity on deep-sea wood islands. There are no structures seen in mature Stepter-
nos individuals which suggest an ability to rise in the plankton as observed in
adults of one chrysopetalid species collected mid water (Watson Russell in Bhaud
and Cazaux, 1987).
 RUSSELL: DEVELOPMENT OF STREPTERNOS 421

A B

Figure 8. A, Mileikovsky's planktonic larva of 6 segments. dorsal view; B. part of mid-body paleal
fascicle from same (after Mileikovsky. 1962, pI. 1, figs A.E). Labels by author: ps. provisional setae,
de. dorsal cirrus, Is. larval spine.

Mile ikovsky ,s Larvae.-It is possible to identify entire chrysopetalid larvae to ge-

nus, based on careful comparisons of different setal types, their position within
the fascicle and their change in posterior segments, and neurosetal and pygidial
details (Watson Russell in Bhaud and Cazaux, 1987).
Mileikovsky (1962: 649, pl.l, figs. A-Z) described some chrysopetalid larvae
collected from the Gulf Stream as Paleanotus heteroseta Hartman. Elsewhere I
have shown that Mileikovsky's identification was incorrect (Watson Russell,
1986). Unfortunately Mileikovsky's specimens are lost (Levenstein, pers. comm.),
so it is not possible to make a positive identification. However on the basis of his
original description and figures, it is possible to speculate on the probable iden-
tification of Mileikovsky's larvae.
Chrysopetalid species belonging to Bhawania, Paleaequor, Paleanotus, Chryso-
petalum, n. genus 1 and Strepternos are present along the northeastern American
coast. Bhawania larvae have never been described in the literature nor have I found
any in collections. Benthic juveniles of Paleaequor have been describt;:dby Watson
Russell (1986) and pelagic larvae of Paleanotus from sheltered coastal embay-
ments have been described by Blake (1975). Pelagic larvae of Chrysopetalum have
been recorded by Bhaud (1986) and photographed by Bhaud (in litt.) from tropi-
cal lagoons. The shape of the primary paleae in all these genera is distictive and
could not be confused with those belonging to Mileikovsky's larvae.
I have examined new genus 1, new species 1, benthic post-larva (7 segments)
from off north Carolina in 45 m (Watson Russell in Bhaud and Cazaux, 1987,
fig. 28. 4-6, and author, in prep.). They possess the broad, symmetrical, primary
paleal types, which are seen in Mileikovsky's larvae (Fig. 8A, B) and Strepternos
post-larvae. Larval characters belonging to new genus 1, Strepternos and Milei-
kovsky's larvae are compared in Table 4.
Mileikovsky did not figure an entire paleal fascicle (Fig. 8B, see 3 paleae out
of a stated 10 drawn) nor did he designate what part of the body the notopodium
came from. While he mentioned change in the number of ribs from paleae within
this fascicle he did not state if paleal types differed down the body. If Milei-
kovsky had figured a posterior notopodium of the 6 segmented larvae it may have
been possible to identify it as Strepternos or n. gen. 1, on the basis of absence or
presence, respectively, of the median paleal fascicle.
422 BULLETIN OF MARINE SCIENCE, VOL. 60, NO.2. 1997

Table 4. Comparisons of larval characters

Mileikovsky's larvae SIrepternos did)'mopylon n. gen. I, n. sp. 1

Reference Mileikovsky, 1962: 649, Watson Russell, this Watson Russell,

pI.I , figs. A,B,D,E, paper 1987:669, fig. 28,
G,Ge,F,Z 4-6 + author's
unpublished
observations
Locality 35°_37°~_70o_74°W 38°-39°~-69°-72°W 34°35'~, 75°52'W
Habitat planktonic benthic-wood islands benthic
Depth (0-100 m) 3,000-4,000 m 1,830-2,500 m 0.3-55 m
No. of segments 5-6 10-11 7
Length (mm) 0.4-0.45 0.6-0.66 0.46
Prostomial eye spots 3 prs. (small) 3 prs. (small) 3 prs. (larger)
Prostomia] appendages none none; median and none; median ant.:nna
]ateral antennae
Oil globules in gut present present present
Larva] tentacular
cirri-seg. I 4 prs. long cirri 2 prs. long cirri + 2 prs. long cirri
pro ventral cirri
~otopodia seg. 2 not formed 2 prs. dorsal cirri
~europodia seg. 2 no cirri no cirri
Pygidium 2 short cirri broad conical lobe narrow conical lobe +
2 slender cirri
Larval notosetal types*
Lateral paleae-no. ribs 9-10 6-11 8-12
Main paleae-no. ribs ]2, 16, 18 12, 16, 18, 20 13. 16, 18
Short notosetal spine in all notopodia in notopodia 3-5 in all notopodia
Length of dorsal cirrus
relative to height of
paleae fascicle '1'3-% V4-V2 2/,
Larval neurosetal medium to short medium to short medium to short
types* falcigers falcigers falcigers
•••based on midbody setiger.
- nol sufficient information.

Prostomial features and the configuration of the primary paleae fan and shape
and number of ribs of lateral and main paleae types are very similar across the 3
groups. Mileikovsky's larvae differs from the other two in the number of tentacu-
lar cirri. The length of the dorsal cirri relative to the height of the paleal fascicle,
and presence of pygidial cirri are more similar between Mileikovsky's larvae and
that of n. gen. I, although anal papillae are sometimes present on the pygidium
of Strepternos. However, while the former character is possibly minor, pygidial
characters have been shown to be important at the generic level (Watson Russell
in Bhaud and Cazaux, 1987).
New genus 1 is known to have a planktonic stage (author, unpublished mate-
rial). Re-examination of "Chrysopetalum" larvae of 6 segments described by
Cazaux (1968) from plankton collected from the bay at Arcachon, French Atlan-
tic coast, shows it to be n. gen. 1, species reyssi. Western Atlantic adult speci-
mens of n. gen. 1, n. sp. 1 are found in subtidal to 55 m depths along the North
and South American coasts. Exchange of larvae is most likely within embayments
and along inshore coastal waters. However it is conceivable that such larvae could
be swept out to sea, especially by the Gulf Stream which comes closer inshore at
Cape Hatteras.
The two Gulf Stream stations where Mileikovsky sampled were at 37°35'N,
70 43'W (east of Chesapeake Bay) and 35°32'N, 74°25'W (off Cape Hatteras) in
0

4,000 m and 3-3,500 m depths, respectively. At the first station, 4 metatrocho-

 RUSSELL: DEVELOPMENT OF STREPTERNOS 423

phores were found; 3 in 25-0 m and 1 in 100-50 m. At the second station a middle
nectochaete was collected in the 100-0 m layer.
The occurrence of Mileikovsky's larvae in oceanic waters very close to abyssal
benthic sites where Strepternos live, is some argument in considering the former
as belonging to the latter genus. The morphology of Strepternos post-larvae indi-
cates a larval planktonic phase. The possession of pigmented eyes points to the
larvae being present in the photic zone of deep waters at depths similar to those
at which Mileikovsky's larvae were collected. Long range larval dispersal is con-
sistent with the present known distribution of S. didymopyton. from Massachu-
setts to the Virgin Islands. The passive transport of larvae over this area is pos-
sible along the warmer waters of the Antilles Current and Gulf Stream, flowing
north, with a return loop along the south flowing North Atlantic Deep Water
(Bramwell, 1977). I am confident in assigning Mileikovsky's larvae to either
Strepternos or n. gen. 1. However, final confirmation of the identity of Milei-
kovsky's larvae will only be resolved by recollecting chrysopetalid larvae at the
same depths at the original Gulf Stream sites.

Possible Origins and Function of Paleate Setal Change.-Problematical Cambrian

scaly fossils of the class Machaeridia, including formerly the family Wiwaxiidae
(among the order Sachitida), possess "scales arranged in longitudinal rows and in
a segmental (pseudosegmenta1?) way on a bilaterally symmetrical body" (Berg-
strom, 1989: 264). Walcott originally interpreted Wiwaxia corrugata (Matthew)
as a polychaetous annelid, but Conway Morris (1985) in a detailed study of Wi-
waxia, concluded that it did not have sufficient polychaete affinities to belong
within the Annelida. Recently, Conway Morris and Peel (1995: 305), concluded
that "from an animal fairly closely related to Wiwaxia arose the polychaetous an-
nelids" with the primitive state in extant polychaetes best expressed in the chry-
sopetalid, aphroditid and amphinomid polychaete groups.
The strong resemblance of wiwaxiid sclerites to paleate polychaete setae was
remarked upon by Watson Russell (1989) in a study of Palmyra aurifera (a paleate,
non-elytragerous aphroditid) and by Westheide and Watson Russell (1992) in a
study on the ultrastructure of chrysopetalid paleae. Butterfield (1990), in a care-
ful anatomical and histological study of wiwaxiid 'sclerites,' showed these struc-
tures to be polychaete paleal notosetae comparative to those belonging to the Bur-
gess Shale polychaete, Canadia spinosa. He concluded that Wiwaxia was a jawed
annelid related to the extant polychaete families Chrysopetalidae and/or Aphro-
ditidae within the order Phyllodocida.
Butterfield's macerated material comprised of wiwaxiid sclerites ranging in
length from 0.14-0.8 mm with a mean length of 0.27 mm. This is in comparison
to the size of bedding plane sclerites of approximately 0.5-10.0 mm length which
were examined by Conway Morris (1985) who observed entire juvenile speci-
mens lacking the long dorsal spines and that "ontogenetic growth of the sclerites
involved size increase and ... some slight modification in relative shape" (p.
532) with certain juvenile sclerite types appearing "more symmetrical in outline"
(p. 538). Butterfield's study of the juvenile palmate sc1erites at high magnifica-
tion reveals ovoid shaped paleae with an ornamentation of raised ribs with raised
denticles (Butterfield, 1990: figs 2, A-C; 3, A, B); other types range from broad
but more distally pointed to slender, distally acuminate shaped paleae (Butterfield,
1990: fig. 1, A-I).
One can only make an approximation of the actual length of the individuals to
which these smaller isolated paleal sclerites belonged. Conway Morris's (1985)
smallest juvenile specimens were about 2.5 mm in length (pI. 12, fig. 123). By
424 BULLETIN OF MARINE SCIENCE, VOL. 60. NO.2, 1997

A /
B
I

/
/,
• "
I'
I
• I I
,,
,, I
I

, ( I I

I I I
I
I
I I ,
I
\
1
,
\ \ I
I
t
I
I

\ ,\ \
I

~\\ \
\
\

c \ , \,
'_s..'

Figure 9. A-D, Wiwaxia paleae; A, adult palmate sclerite (after Conway Morris, 1985: fig. 143); B
isolated entire juvenile palmate sclerite, C isolated part of juvenile? sclerite, D detail of distal end of
sclerite showing raised ribs and denticles (9B-D after Butterfield, 1990: figs. 2B, 3A, IE).

comparing the size of paleae belonging to these juveniles with those recorded by
Butterfield one can estimate that the probable length of entire wiwaxiid speci-
mens from Butterfield's study would fall in the 1-2 mm range.
Very broad, symmetrical, primary paleal setae are seen in planktonic and newly
settled benthic larvae belonging to a number of chrysopetalid species. S. didy-
mopyton benthic juveniles, for example of 10 to 15 segments, range in length
from 0.6 to 1.5 mm, with a primary paleallength of 0.06--0.12 mm. Primary paleal
notosetae, especially the medial group paleae, are distally rounded and possess
serrated margins and a dorsal surface ornamented with raised ribs and raised den-
tides on the ribs. There appears to be a strong similarity between Recent larval
chrysopetalid paleae, and fossil juvenile wiwaxiid paleae, and their subsequent
replacement by adult paleae (cf. Fig. 7B to Fig. 9A-D, this paper). Whether wi-
waxiids possessed a planktonic larvae is open to conjecture. It is apparent, how-
ever, that microplanktic primary productivity has been present as far back as the
Lower Cambrian (Butterfield, 1994).
 RUSSELL: DEVELOPMENT Of STREPTERNOS 425

Adult wiwaxiids are covered in a series of long dorsal spines and imbricating
rows of paleal 'sclerites' divisible into dorsal, upper lateral, lower lateral, ante-
rior, and ventro-Iateral regions (Conway Morris, 1985). The fossil polychaete Ca-
nadia, included with Wiwaxia in the superfamily Canadiacea, order Phyllodocida
(Butterfield, 1990), and Recent broad paleate chrysopetalid species possess bi-
ramous parapodia and imbricating paleal rows within the notopodial fascicle, di-
visible into lateral, dorsal and medial regions, plus paleal sub-units. Chrysopeta-
lids possess compound neurosetae but also small, simple, serrate neurosetal spines
(Watson Russell, 1986; this paper); the latter are also documented as 'probable
neurosetae' co-existing with the simple paleal neurosetae belonging to Canadia
spinosa (Butterfield, 1990).
Differences between wiwaxiids and Recent chrysopetalids include a greater and
more robust adult size, possession of ventral jaws, and apparent lack of parapodia
and neurosetae in the former (but see Butterfield's (1990, 1994) assessment that
simple spinose neurosetae are most likely present). Wiwaxiids also appear to at-
tain the adult number of segments very early on and do not add segments during
growth, a developmental factor more in common with the aphroditids than the
chrysopetalids. Notwithstanding the obvious gaps in the fossil record and lack of
understanding of the lineage of these groups, I think it is of interest that Cam-
brian and Recent collections reveal juvenile and adult paleate polychaetes dis-
playing similar ontogenetic setal changes.
In conclusion, it is hypothesized that possession of primary paleal notosetae
and their subsequent metamorphosis has its antecedents in patterns laid down in
one line of ancestral chrysopetalid polychaetes, where setal change occurred be-
tween a larval pelagic phase and a rigorous adult benthic phase, and where
sclerite/paleal setae afforded much needed protection.

ACKNOWLEDGMENTS

I am grateful to K. Fauchald, USNM, for the loan of this interesting material, most of which was
made available through studies by R. D. Turner on "Wood Islands" conducted on DRSV ALVIN (study
supported by ONR Grant N00014-84-C-0258). I also wish to thank L. Laubier for providing me with
a French translation of S. A. Mileikovsky's 1962 paper and the Northern Territory Museum of Arts
and Sciences for support during this project.

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DATEACCEPTED:September 23, 1996.

ADDRESS:Department of Annelids, Division of Natural Sciences, Museums and Art Gallery of the
Northern Territory. GPO Box 4646, Darwin, N. T. 0801, Australia.

APPENDIX

Abbreviations on Figures: Anterior end: c = cilia, dc = dorsal cirrus, dtc = dorsa] tentacular
cirrus, ep = eye pigment, la = lateral antenna, ]dc = larva] dorsal cirrus, lvc = larval ventral cirrus,
rna = median antenna, mo = mouth opening, nf = nuchal fold, p = pa]p, pi = paleae insertion, pr =
prostomium, vc = ventral cirrus, vtc = ventra] tentacular cirrus, roman numeral = segment number.
Adult setal types. Notosetae: al = adult lateral paleae, aim = adult lateral main paleae, am =
adult main paleae, ame.m = adult media] main paleae, amm = adult misshapen main paleae. Neuro-
setae: mf = middle falciger, sf = superior fa]ciger. Juvenile setal types. Notosctae: I = lateral paleae,
1m = lateral main paleae, Is = larval spine, me.m = medial main paleae, mi.m = middle main paleae,
su I = sub unit I pa]eae. Neurosetae: same categories as adult.