Zoomorphotogy (1983) 102:175-187 Zoomorphology

© Springer-Verlag 1983

Morphology and Function of the Tube Feet

of Florometra serratissima (Echinodermata: Crinoidea)
M. Byrne and A.R. Fontaine
Department of Biology, University of Victoria, Victoria, B.C., Canada V8W 2Y2

Summary. Crinoid tube feet occur in groups of three (podial triplets)

that behave as functionally integrated units for food particle collection.
Scanning electron microscopy shows that each triplet member has a
distinctive morphology related to its behavioural role in feeding. Particu-
larly conspicuous features are papillar distribution, papillar processes,
ciliary tracts, and pores that are the openings of mucous glands. The
primary podia serve for initial particle capture. The secondaries, together
with lappets, play a major role in particle transfer from the primaries
and may themselves also entrap particles. The tertiaries manipulate parti-
cles within the food groove after their transfer from the other podia.
The tertiary podia use their papilla-free medial faces to compact mucus-
bound particles into boluses. In pinnules, boluses are transferred to the
food groove midline by tertiaries, then propelled orally by paddling
actions of these podia. Boluses are also transported by the medial ciliary
tracts of the pinnules and arms. Bolus transport by tertiary paddling
is probably more effective in the pinnular than the arm food groove.
Short lateral ciliary tracts at the base of each primary podium may
guide boluses into the main ciliary stream or may be cleansing currents.
As the ultimate site of food particle collection, compaction and transfer,
podial triplets represent a third adaptive level of an intricate suspension
feeding system in which the first (arm postures) and second (pinnule
orientations) adaptive levels are related to increasing collecting efficiency
in diverse ambient flow.

A. Introduction
The morphology of the water-vascular system and podia of comatulid crin-
oids described by early workers (Chadwick 1907; Reichensperger 1908 a;
Gisl6n 1924) has been most recently studied by Nichols (1960). The radial
Offprint requests to: M. Byrne
176 M. Byrne and A.R. Fontaine

water vessel within each arm regularly branches off pinnular canals from
which tributaries divide to form the lumina of three morphologically and
functionally associated tube feet, the podial triplets defined by Byrne and
Fontaine (1981). Podial triplets and lappets associated with them line either
side of the ambulacral groove (food groove). The three members of a triplet
differ in size (primaries are longest; secondaries, of intermediate length;
tertiaries, shortest) and in other aspects of their functional morphology.
Podial behaviour during feeding has been observed in laboratory situa-
tions (Gisl6n 1924; Nichols 1960; Byrne and Fontaine 1981) and recent
investigations (LaTouche 1978; Meyer 1979; Byrne and Fontaine 1981;
Zmarzly and Holland 1981) demonstrate the more easily observed aspects
of their behaviour in natural situations.
Behavioural studies of podia have been confined mainly to the primaries.
Direct examination of the secondaries and tertiaries within the food groove
is impeded by their small size and the difficulty of observing podia within
the groove. These problems prompted the present investigation that uses
the scanning electron microscope (SEM) to supplement direct examination
of podia and structures within the food groove of Florometra serratissima,
a comatulid crinoid found in waters around British Columbia.

B. Material and Methods

Specimens of Florometra serratissima (Clark, 1907) were collected near the Bamfield Marine
Station in Barkley Sound on the West Coast of Vancouver Island. The collection site and
methods for behavioural observation in vivo were outlined in Byrne and Fontaine (1981).
Specimens were processed soon after collection. Prior to fixation arms were lowered individually
into a chilled isotonic (6.7%) MgCI 2 solution for relaxation and then transected near the
tegmen. Pieces of relaxed arms were fixed in a 2% glutaraldehyde-sea water solution at room
temperature and processed following the method of Holland and Jespersen (1973). The fixed
tissue was critical-point dried, mounted on aluminium stubs with conductive paste, sputter
coated with gold and viewed with a JEOL JSM-35 scanning electron microscope at 15 KV.
The presence of metachromatic mucus was demonstrated in vivo by placing specimens in
sea water containing a low concentration of Toluidine blue (Fontaine 1965).

C. Results

1. Pinnular Food Groove

The pinnular food groove is bordered by lappets and podial triplets. The
lappets alternate with each other on the two sides of the groove, conferring
a scalloped appearance to the food groove margins (Figs. 2, 3). The basic
functional unit for food particle collection is the podial triplet and its asso-
ciated lappet. Their morphological integration is shown in Figures 1 and
2 where each member of a triplet and its lappet is labelled.
A central ciliary tract along the midline of the pinnular food groove
receives short lateral tracts that originate near the primary podial base and
traverse toward the base of the secondary podium before merging with
The Tube Feet of Florometra serratissima 177

Large arrows on photographs point proximally.

i.e., towards the mouth. The numbers 1, 2, 3:
primary, secondary and tertiary podia.

Fig. 1. Diagram of podial triplet in ambulacral

(food) groove of a pinnule showing
characteristic "rest" posture of triplet podia
and distribution of their papillae (P). L-lappet;
CC-central ciliary tract; LC-lateral ciliary tract.
Bar = 75 lim

the central tract (Figs. I, 3-7). The central tract eventually merges with
a broad radial ciliary tract, the major tract of an arm (Fig. 5). Rounded
beads resting on the floor of the groove (Figs. 4, 6) may be mucous droplets
extruded from the mucous glands that secrete into the groove (Reichens-
perger 1908a; Nichols 1960). Those associated with the cilia (Fig. 6) may
be ciliary dilations that have been shown elsewhere (Eakin and Branden-
burger 1979) to be artifacts of fixation. Toluidine blue stains a film that
coats the podia and food groove, verifying the presence of coating mucus.
Numerous microvillus-like projetions are found on the floor of the groove,
sometimes encircling cilia (Fig. 6).

2. Podia

a) Primary Podia. - The primary podia (ca. 1100 ~tm in length) are typically
held extended with an aboral curl at their tips when inactive (Fig. 2). They
protrude between lappets, usually associated more closely with the proximal
lappet; i.e., the lappet adjacent to the proximal side of the primary podial
stem (Figs. 1-3). The primaries often rest their bases on the lappet edge,
as observed in vivo, although in some pinnular regions a greater distance
between the proximal lappet and primaries was observed.
Papillae project from the primaries at regularly spaced intervals (Figs.
1, 2, 8). They appear to be more abundant towards the tip and become
reduced in size basally (Figs. 9-11). Mucous filaments were sometimes ob-
178 M. Byrne and A.R. Fontaine
Fig. 4. Ciliary tracts of pinnular groove; CC-central tract; LC-lateral tract. The bead-like
objects on the groove floor may be mucous droplets. Bar = 5 ~tm
Fig. 5. Central ciliary tract (CC) merging with the broad radial tract (RC) of the arm midline.
Bar = 15 ~tm
Fig. 6. Central tract showing miclZovillus-like projections (arrow) on the groove floor, sometimes
enciecling cilia. Round structures associated with cilia (double arrow) may be artifactual ciliary
dilations. Bar = 1.5 lam
Fig. 7. Pinnular triplet adjacent to the arm-pinnule junction, primary podium (1), shorter
than secondary (2). Bar = 75 tim

s e r v e d o n a n d b e t w e e n t h e p a p i l l a e ( F i g . 8). S p a r s e c i l i a w e r e o c c a s i o n a l l y
f o u n d a l o n g t h e p r i m a r y p o d i a l b a s e ( F i g . 11). T h e s e c i l i a m e r g e w i t h t h e
lateral ciliary tract that starts at the base of the primaries.

b) Secondary Podia. - T h e s e c o n d a r i e s (ca. 360 Ixm in l e n g t h ) w h e n i n a c t i v e

a r e c u r v e d s l i g h t l y a w a y f r o m t h e g r o o v e ( F i g s . 1-3). T h e y a r e a s s o c i a t e d

Fig. 2. Ambulacral (food) groove (A) of a pinnule lined on both sides by alternating lappets
(L) and podial triplets in "rest" postures. Bar = 500 ~tm
Fig. 3. Central (CC) and lateral (LC) tracts within the pinnular groove around three sides
of each tertiaries' base. Primaries lie close to proximal lappets (L). Lappets, distally inclined
and partially enwrap secondary podial bases. Tertiaries in their "interception" position. Small
arrows indicate the direction of ciliary current. Bar = 50 lam
180 M. Byrne and A.R. Fontaine

Fig. 8. Primary podium showing regularly spaced papillae and mucous filaments (arrow). Bar =
50 I~m

Fig. 9. Primary podial tip showing spacing of papillae, one of which has an unidentified sperm
attached to its tip (arrow). Bar = 25 ~m
The Tube Feet of Florometra serrat&sirna 181

with the medial face of the distal end of the lappets with which they appear
to fuse basally. Only the outer lip of the lappet is not fused to the base
and stem of the secondary podium. The lappets were sometimes seen to
have a distal inclination and partially to enwrap the base of the secondaries
(Figs. 3, 12). The secondaries have abundant, closely spaced papillae distrib-
uted circumferentially, from which mucous filaments project (Figs. 2, 12).

c) Tertiary Podia. - The tertiaries are the smallest members of the triplets
(ca. 230 ~tm in length). They emerge from the proximal basal region of
each lappet (Figs. 1-3). Papillae are restricted to the proximal and distal
edges of the tertiaries. Thus, the medial and lateral surfaces (i.e., facing
towards and away from the food groove) are papilla-free (Figs. 1-3, 13).
This gives the tertiaries a spatulate profile when viewed from above. The
tertiaries are surrounded on three sides by ciliary tracts (Figs. 1, 3).

3. Papillae

Papillar distribution, as noted above, differs in each triplet member. The

primaries have regularly spaced papillae, the secondaries are more abun-
dantly papillate, whereas those of the tertiaries are bilaterally distributed.
In general, papillae are approximately 40 gm in length, although this varies
somewhat along the length of the podium.
Unicellular mucous glands found at the base of the papillae (Reichens-
perger 1908a; Nichols 1960, Holland 1969) send processes to the papillar
tips. These are the likely source of the mucous filaments observed, and
presumably also they provide the podia with a sticky surface. Potential
food items were often observed ensnared in papillar mucus (Figs. 9, 14).
Fine processes (typically 4-6 in number, up to 4 gm long) were observed
projecting from the papillar tips (Figs. 15, 16). They were not observed
on some papillae, perhaps obscured by covering mucus.

Fig. 10. Lateral (outside) view of the basal part of a primary, papillae reduced in size basally,
PP-papillar processes; C-cilia of the lappet edge. Bar = 12.5 ~tm

Fig. 11. Medial (inside) view of a primary podial base and adjacent lappet, showing reduced
papillae and sparse ciliation of podial base and lappet edge (C). Bar= 15 Ixm

Fig. 12. Secondary podium showing closely spaced papillae and mucous filaments (arrow).
The lappet partially enwraps the base of the secondary and they appear to be basally fused.
Bar = 60 ~tm

Fig. 13. Tertiary podium showing marginal papillae and relatively smooth and spatulate medial
and lateral faces. Bar = 25 ~tm

Fig. 14. Unidentified larva trapped between papillae of a secondary. Bar = 20 ktm
182 M. Byrne and A.R. Fontaine

Fig. 15. Papillar processes (PP) on the papillae of a secondary podium. Some of the papillae
are bent over, apparently held down by attached mucous filaments as in the papilla in the
foreground. Bar = 60 gm

Fig. 16. Papillar processes of a primary podium. Bar = 30 gm

Fig. 17. Retracted podia and lappets form a partial cover over the food groove. Bar= 100 ~tm

Fig. 18. Lateral (outside) surface of a lappet showing numerous pores and scattered cilia.
Bar = 25 gm

4. Lappets

L a p p e t m o v e m e n t is directly a s s o c i a t e d w i t h a c t i o n s o f the secondaries.

W h e n the p o d i a retract, the lappets r o o f o v e r the g r o o v e (Fig. 17). T h e
o u t w a r d f a c i n g l a p p e t s u r f a c e a n d edge is ciliated a n d h a s n u m e r o u s p o r e s
(Fig. 18). T h e latter are likely o p e n i n g s o f l a p p e t m u c o u s g l a n d s d e m o n -
s t r a t e d h i s t o l o g i c a l l y b y N i c h o l s (1960).
The Tube Feet of Florometra serratissima 183

5. Behaviour of Tertiary Podia

Details of tertiary behaviour were not available when the basic sequence
of podial activity during feeding was established by Byrne and Fontaine
(1981) for F. serratissima. In stage 2 of that sequence when all the members
of a triplet are within the groove, they apparently wipe along each other.
The papilla-free medial face of the tertiary apparently wipes off mucus-
bound particles from the stems of the primary and secondary members.
This is probably the action that combines mucus-bound particles into a
loose bolus. Mucus from the tertiaries' papillae and from glands in the
food groove floor (Nichols 1960) may contribute further to particle binding.
Tertiaries also appear to be agents of bolus propulsion. They obviously
push boluses inwards towards the central ciliary tract of the pinnule. They
also apparently contribute to the centripetal impetus of boluses in conjunc-
tion with the central ciliary stream by means of an orally-directed succession
of paddling actions by the tertiaries in sequence. The tertiaries at rest are
typically held curved towards the groove midline (Fig. 2). This is interpreted
as an "interception position," i.e., the tertiaries intercept and propel boluses
onwards as they are passed down the groove.

6. Arm and Pinnule Junction

The primaries are reduced in length in the food groove at the arm-pinnule
junction and along the arm itself. Their dimensions are variable, but they
are usually of similar length to or slightly longer than the secondaries.
Occasionally primaries shorter than the secondaries were observed (Fig. 7).
The lateral ciliary tracts in the pinnular junction region give off ciliary
tributaries to each triplet member.

D. Discussion

Crinoids differ from other echinoderms in that their podia arise and function
in groups of three. The gross morphology of the podial triplets of F. serrat-
issima is similar to that described for other crinoids (Chadwick 1907; Rei-
chensperger 1908a; Nichols 1960). Each triplet member in this species has
a distinct behaviour related to its role in suspension feeding (Byrne and
Fontaine 1981). The SEM observations presented here add fine detail to
the adaptations of the triplet members, demonstrating that the ch~tacteristic
morphology of each member (especially with respect to papillar distribution)
is related to its behavioural role in feeding.
The sticky surface of crinoid podia was noted early (Reichensperger
1908a; Gislrn 1924). Unicellular mucous glands opening at the papillar
tips, in the ambulacral groove and on the outward lappet surface have
been demonstrated histologically (Reichensperger 1908a; Nichols 1960).
The pores on the lappets of F. serratissima shown by SEM are presumably
184 M. Byrneand A.R. Fontaine

openings of these mucous glands. The presence of a sticky mucous coat

in vivo was verified with Toluidine blue. However, other glandular entities
with a different type of secretion may be present as well.
Crinoid suspension feeding has features similar to the suspension feeding
mechanisms observed in ophiuroids (Buchanan 1963; Fontaine 1965; Pen-
treath 1970; Warner and Woodley 1975). These features are: an adhesive
coating on podia, papillar morphology, and protracted podia as a major
behavioural component in particle capture. This suggests that comatulid
crinoids may also use the aerosol suspension feeding model proposed for
ophiuroids (LaBarbera 1978).
Motions of the primary podia are the most conservative of the move-
ments made by the three podia. During particle collection primaries bend
towards the groove with a sinusoidal action (Byrne and Fontaine 1981),
bending only in one plane at right angles to the longuitudinal axis of the
pinnules. Nichols (1960) found only a single muscle column in the primaries
along the side facing the food groove. This muscle arrangement correlates
with primary retraction. In the absence of an antagonistic muscle, protrac-
tion is presumably accomplished entirely by a hydrostatic effect of the water
vascular lumen.
The papillae on the primaries are distributed uniformly around and
along the podial length. Because primaries are typically held extended for
particle capture, symmetry of papillar distribution may be an adaptation
for capturing particles that impinge from any direction.
During suspension feeding, the secondaries flex outward with a distinc-
tive scooping action (Byrne and Fontaine 1981). They are very active podia,
frequently scooping even while the primaries are quiescent. There are two
columns of muscles in the secondaries (Nichols 1960). This antagonistic
muscle pair appears to be an adaptation that facilitates the rapid, repetitive
motion in one plane that is characteristic of secondaries.
The secondaries are more abundantly papillate than the other triplet
members; presumably more mucus is needed by them, not only for direct
particle capture but additionally for particle transfer from the primaries,
probably with the aid of the lappets. The lappets are morphologically and
behaviourally associated with the secondaries and could be regarded as
an expansion of the secondary podial base. As the secondaries move out
and into the groove, the lappets moving with them appear to scrape the
primaries' stems, perhaps assisting transfer of adherent particles from the
primaries to the secondaries. Lappets may also be involved in direct transfer
from the~pl;imaries to the lateral ciliary tracts. This aspect of lappet activity
may be ~.2~Ncially important in those pinnular regions where the primaries
• . ~:,C-. . .

he m close proximity to the proximal lappet and may also involve lappet
cilia and mucous glands. Lappet cilia are not organised into tracts, and
their function may be in cleansing rather than in feeding. The scraping
aspect is analogous to that of the ophiuroid tentacle scale (Fontaine 1965;
Pentreath 1970), but in ophiuroids the tentacle scale that acts as a scraper
is an entity physically removed from the podium.
When the podia retract during feeding, the lappets roof over the groove.
The Tube Feet of Florometra serratissima 185

This could be significant in particle retention during feeding activities. When

podia retract as part of an avoidance response to touch or perhaps as
a local response to myzostome parasites, the lappets close to form an ostensi-
ble protective covering. A protective role tradionally has been ascribed to
the lappets, but this may be a secondary one. The primary role is in feeding,
where lappet action is a part of the scooping behaviour of the secondaries.
The activities of the tertiaries are restricted to the trough of the food
groove. They are the most dexterous podia, capable of swinging through
a 360 ° arc. This mobility correlates with a complete circumferential muscle
ring (Nichols 1960). Tertiaries are not attached to the lappet wall in F.
serratissima, contrary to observations by Nichols (1960) on Antedon bifida.
The papillae of the tertiaries are bilaterally placed and the papilla-free medial
face appears important in wiping off particles adhering to the primary and
secondary podia and in bolus compaction. The tertiaries serve a dual func-
tion: first, in bolus formation and, second, in propelling the bolus towards
the mouth. We are unsure whether paddling action of the tertiaries or con-
veyance by ciliary tract is the most important transport mechanism. Both
obviously play a role in food particle transport within the food groove
of the pinnule at least, but the role of tertiary paddling is probably reduced
at the level of the radial (arm) ciliary tract. Rates of particle transport
down the arm ambulacral groove have been measured for a tropical crinoid
(Zmarsly and Holland 1981) at about 4.2 cm per min.
The primary podia are reduced in size along the food groove near the
pinnule-arm junction and along the arm food groove as noted by Nichols
(1960) for A. bifida. In these regions the role of the primaries in particle
capture is likely to be less important than cooperation with other triplet
members in particle transport within the food groove.
Lateral ciliary tracts were also noted by Gisl6n (1924); they presumably
help guide boluses into main ciliary streams of the groove midline. In addi-
tion, they may collect particles directly from the bases of the primaries
and, where tributaries from the lateral tract are associated with the secondar-
ies and tertiaries, the lateral tract may collect directly from these podial
bases as well. The lateral tracts probably collect stray items and thus may
function as cleansing currents sensu Gils6n (1924).
The mechanisms of bolus formation in crinoids differ from those of
ophiuroids (Fontaine 1965; Pentreath 1970; Warner and Woodley 1975)
where boluses grow in size and move relatively slowly towards the mouth
through sequential activities of cooperating pairs of podia.
Microvilli are abundant along the food groove, sometimes forming
collars around ciliary bases. This suggests the presence of choanocyte-like
cells in F. serratissima such as found by Norrevang and Wingstrand (1970)
in the pinnular food groove of another crinoid Hathrometra tenella, on
holothurian tentacles (Fankboner 1978)and in asteroid cribriform organs
(Shick et al. 1981). These are all feeding surfaces.
The abundant papillar mucous glands (Reichensperger 1908 a; Nichols
1960) and their role in entrapping potential food items indicate that the
papillae play a major role in feeding. They may also be sensory. Numerous
186 M. Byrne and A.R. Fontaine

nerve cells lying at the papillar base send processes towards the papiUar
tip (Reichensperger 1908a; Nichols 1960). In Antedon bifida projections
observed on papillar tips were considered to be sensory (Chadwick 1907;
Reichensperger 1908 a; Nichols 1960) and to originate from the basilar nerve
cells (Nichols 1960). They are similar to the papillar processes observed
in F. serratissima that also have four to six processes per papilla as in
A. bifida. Papillar processes were not observed by Holland (1969) in Nemas-
ter spp. A preliminary examination of F. serratissima indicated that the
processes have the internal fine structure of a cilium (Holland 1969). The
processes observed in this study do not resemble the ciliary tract cilia mor-
phologically, but they may be non-motile stereocilia. Whether these papillar
projections are processes of nerve cells or are based on ciliary structure
requires ultrastructural demonstration, but either way they are likely to
be sensory in function.
The papillar processes are reminiscent of the "st/ibchen" projections
found in association with secretory cell ducts on the spines of some ophiur-
oid species. The "stfibchen" are also thought to be sensory (Reichensperger
1908 b; Buchanan 1963; Pentreath 1970). Transmission electron microscopy
of t h e " st~ibchens" shows unequivocally that they are based on ciliary struc-
ture and that they are situated in close proximity to secretory openings
(unpublished results, Byrne and Fontaine).
The papillae do not appear to have muscle associated with them (Holland
1969). Thus, mucus release does not appear to involve muscular contraction
of the papillae. Mucus release is possibly triggered by stimulation of the
sensory papillar processes, as proposed elsewhere (Reichensperger 1908a;
Nichols 1960).
Observations of many workers accumulated over recent years have given
a good understanding of feeding methods of modern crinoids. In our view
crinoid feeding mechanisms rest on three levels of adaptation: 1. Arm pos-
tures: highly adaptive for forming the most efficient collecting framework
through accurate placement of the arms with respect to the ambient flow
(Magnus 1963; Meyer 1973~ 1979; LaTouche 1978; Byrne and Fontaine
1981); 2. Pinnule orientations: highly adaptive for increasing the efficiency
of particle collection within the framework established by the arm posture
(Magnus 1963; Meyer 1973, 1979; LaTouche 1978; Byrne and Fontaine
1981); 3. Podial triplet activities: the ultimate level, the site where particles
are physically collected, transfered into the food groove, compacted into
boluses, and conveyed orally (Nichols 1960; Byrne and Fontaine 1981).
It seems to us that the first and second adaptive levels can be expected
to Be most variable within and between species, but that the functional
morphology of third level structures is likely to be conservative, at least
within the comatulids.

Acknowledgements. The work was supported by an operating grant to A.R. Fontaine from
the National Science and Engineering Council of Canada. We thank the University of Victoria
for Fellowship support of M. Byrne and the Director of the Bamfield Marine Station for
use of facilities.
The Tube Feet of Florometra serratissima 187

References

Buchanan JB (1963) Mucus secretion within the spines of ophiuroid echinoderms. Proc Zool
Soc London 141:251-259
Byrne M, Fontaine AR (1981) The feeding behaviour of Florometra serratissima (Echinoderma-
ta: Crinoidea). Can J Zool 59:11-18
Chadwick HC (1907) Antedon. Liverpool Marine Biological Committee Memoir 15:47 pp
Eakin RM, Brandenburger JL (1979) Effects of light on ocelli of seastars. Zoomorphology
92:191-200
Fankboner PV (1978) Suspension-feeding mechanisms of the armoured sea cucumber Psolus
chitinoides Clark. J Exp Mar Biol Ecol 31:11-25
Fontaine AR (1965) The feeding mechanisms of the ophiuroid Ophiocomina nigra. J Mar
Biol Ass UK 45:373-385
Gislrn T (1924) Echinoderm studies. Zool Bidr Uppsala. 9:270-295
Holland ND (1969) An electron microscopic study of the papillae of crinoid tube feet. Pubbl
Staz Zool Napoli 37:575-580
Holland ND, Jespersen A (1973) The fine structure of the fertilization membrane of the feather
star Comanthusjaponica (Echinodermata: Crinoidea). Tissue and Cell 5: 209-214
LaBarbera M (1978) Particle capture by a Pacific brittle star: experimental test of the aerosol
suspension feeding model. Science 201 : 1147-1149
LaTouche RW (1978) The feeding behaviour of the featherstar Antedon bifida (Echinoderma-
ta:Crinoidea). J Mar Biol Ass UK 58:877-890
Magnus DBE (1963) Der Federstern Heterornetra savignyi im Roten Meer. Nat Mus
93: 355-368
Meyer DL (1973) Feeding behaviour and ecology of shallow water unstalked crinoids (Echino-
dermata) in the Caribbean Sea. Mar Biol 22:105-129
Meyer DL (1979) Length and spacing of the tube feet in crinoids (E'cl~inodermata) and their
role in suspension-feeding. Mar Biol 51 : 361-369
Nichols D (1960) The histology and activities of the tube feet of Antedon bifida. Q J Microsc
Sci 101 : 105-117
Norrevang A, Wingstrand KG (1970) On the occurrence and structure of choanocyte-like
cells in some echinoderms. Acta Zoologica 51:249-270
Pentreath RJ (1970) Feeding mechanisms and the functional morphology of podia and spines
in some New Zealand ophiuroids (Echinodermata). J Zool London 161:395-429
Reichensperger A (1908a) Uber das Vorkommen von Driisen bei Crinoiden. Zool Anz
33 : 363-367
Reichensperger A (1908 b) Die Driisengebilde der Ophiuren. Z Wiss Zool 91:304-350
Shick JM, Edwards KC, Dearborn JH (1981) Physiological ecology of the deposit-feeding
sea star Ctenodiscus crispatus: ciliated surfaces and animal-sediment interations. Mar Ecol
Prog Ser 5:165-184
Warner GF, Woodley JD (1975) Suspension-feeding in the brittle-star Ophiothrix fragilis.
J Mar Biol Ass UK 55:199-210
Zmarzly DL, Holland ND (1981) Rates of food transport down the ambulacral grooves and
through the gut of Comanthus bennetti (Echinodermata:Crinoidea) observed in situ. Mar
Ecol Prog Ser 6:229-230

Received August 6, 1982