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MOP_& BML
A Key to the Larvae of the Polychaetous annelids of the Hawaiian Islands
Linda A. Ward
Dept. of Zoology, University of Hawaii 22 June, 1977
Advisors: Dr. J.H. Brock
John McMahon
ABSTRACT
Larvae rep resen t i ng e i g h t f a m i l i e s o f po lychaetes c o l l e c t e d from
Hawai ian waters a r e descr ibed and a taxonomic key f o r these f a m i l i e s i s
p rov ided . Larvae samples were c o l l e c t e d f rom t h e A l a Wai Canal and
Kaneohe Bay, Oahu, ma in ta ined i n p e t r i d i shes i n t h e l a b o r a t o r y where
t h e accompannying photomicrographs and drawings were made. Represen ta t i ve
. - specimens were preserved f o r f u t u r e re fe rence . A b i b l i o g r a p h y d e a l i n g
w i t h po lychae te l a r v a e from o t h e r l o c a t i o n s th roughout t h e w o r l d i s
i n c l uded.
INTRODUCTION
Polychaetes a r e i m p o r t a n t c o n s t i t u e n t s o f t r o p i c a l r e e f communit ies.
They a r e v i t a l as: 1 ) a food source f o r o t h e r organisms, 2 ) p reda to rs ,
3 ) as bore rs and f o u l e r s of hard subs t ra tes . Polychaetes f r e q u e n t l y
occur i n very h i g h d e n s i t i e s : Kohn and L l o y d (1973) found 49,000 polychaetes/m2
on t h e sha l low r e e f s o f t h e I n d i a n Ocean; w h i l e Brock and Brock ( 1 977)
- . r eco rd 50,100-1 27,900 polychaetes/m2i n samples o f c o r a l r u b b l e c o l 1 ec ted
from Kaneohe Bay, Oahu, Hawai i . A t such h i g h d e n s i t i e s t hey a r e o f t e n
impo r tan t l i n k s i n t h e t r o p h i c cha in . The d i e t o f some r e e f f i s h e s and
i n v e r t e b r a t e s i s known t o c o n s i s t a lmost e x c l u s i v e l y o f po lychaetes.
H i a t t and S t rasburg (1960) found t h a t po lychaetes a r e h i g h l y u t i l i z e d as
food i tems by c e r t a i n r e e f f i s h e s i n c l u d i n g members o f t h e f a m i l i e s \
Holocentr idae, Dussumieridae, Hemiramphidae and P r i acan th idae , i n t h e
Marsha l l I s l ands . Polychaetes a r e t h e most common food organism o f
seven species o f Car ibbean r e e f f i shes i n c l ud ing Chaetodon s t r i a t ts and - Halichoeres maculippina where po lychaetes comprise 58.7% and 47.1% o f
t h e d i e t r e s p e c t i v e l y (Randal 1, 1967). Kohn (1 959) observed t h a t o f t h e
s i x t e e n spec ies o f Conts ("cone s h e l l s " ) c o l l e c t e d f rom t h e r e e f p l a t f o r m s
o f t he Hawaiian I s l ands , t e n spec ies feed e x c l u s i v e l y on po lychae tes ;
po lychaetes comprise a l e s s e r , b u t s i g n i f i c a n t p a r t o f t h e d i e t of t h r e e
a d d i t i o n a l spec ies o f t h e genus. These t h i r t e e n spec ies o f Conus
prey on a t l e a s t 23 d i f f e r e n t spec ies o f e r r a n t and sedenta ry po lychae tes .
The feed ing methods o f po lychaetes a r e va r i ed , r ang ing from f i l t r a t i o n
and suspension f eed ing t o a c t i v e p reda t i on . The common f a m i l i e s a r e
c h a r a c t e r i z e d by p a r t i c u l a r f eed ing types. S y l l i d s a r e thought t o feed
on sponges, bryozoans and hydro ids by p i e r c i n g t h e bodywal l and e x t r a c t i n g
i n t e r n a l f l u i d s of t h e prey. Nereids, which a r e omnivores, use t h e i r
mouth p a r t s t o grasp such p rey as amphipods and o t h e r po lychaetes o r t o
r a s p a l gae f rom hard subs t ra tes . Sp ion ids s e l e c t food p a r t i c l e s from *-
t h e s u b s t r a t e near t h e i r tube u s i n g a p a i r o f p ros tom ia l pa lps. F i l t e r -
f e e d i n g i s i l l u s t r a t e d by t h e s e r p u l i d s which use t h e i r c i l i a t e d b r a n c h i a l
crown t o s t r a i n p a r t i c l e s from t h e sur round ing wate r (Day, 1967).
Many po lychaetes a r e o f economic impor tance because o f t h e i r a b i l i t y
t o s e t t l e on o r bore i n t o va r i ous subs t ra tes . Se rpu l i ds a r e ca lcareous
t u b e - b u i l d i n g worms. Some, such as, Hydroides elegans and Hydroides
d i rmpha a r e o f i n t e r e s t because they f o u l s h i p h u l l s , p i e r p i l i n g s ,
wa te r i n t a k e p ipes and o t h e r mar ine s t r u c t u r e s (Ba i ley-Brock, 1976). - Some s p i o n i d s o f t h e genera PoZydora and Boccardia (B lake and Evans,
1975) and t h e c i r r a t u l i d Dodececaria s p . (Blake, 1969a) a r e known t o
bo re i n t o t h e s h e l l s o f mo l luscs . O f p a r t i c u l a r concern t o o y s t e r
c u l t u r i s t s i n Hawai i , and th roughout t h e wor ld , i s t he s p i o n i d PoZydora
websteri which bores i n t o t h e s h e l l s o f a number o f spec ies of oys te r .
Wh i le t h e worm does n o t e a t t h e o y s t e r o r d i r e c t l y harm it, they do
weaken t h e s h e l l . The worm l i n e s i t s t ube w i t h mud which i s c o l l e c t e d
f r om t h e surrounding area and may c o n t a i n h i g h concen t ra t i ons o f b a c t e r i a
such as Escherichia coZi which can rende r t h e o y s t e r unmarketable.
I n o r d e r t o understand t h e geographic d i s t r i b u t i o n o f a d u l t worms
and t h e po lychae te communit ies t y p i c a l o f t r o p i c a l ree fs and temperate
seas, a s tudy o f t h e l a r v a l forms i s necessary. C o l l e c t i o n of b e n t h i c
a d u l t po lychae tes can be d i f f i c u l t and t i m e consuming, o f t en r e q u i r i n g
l a b o r i o u s c h i p p i n g o f c o r a l r u b b l e (Brock and Brock, 1977) o r s i f t i n g of
1 arge volumes of decaying mud. However, b e n t h i c po lychae tes t y p i c a l l y
have p l a n k t o n i c l a r vae , and i t i s u s u a l l y e a s i e r t o sample p l ank ton . -
r a t h e r than benthos. Seasonal p l ank ton c o l l e c t i o n s i n a s tudy area can
p r o v i d e da ta on t h e po lychae te l a r v a e p resen t and t h e i r s p a t i a l and
temporal d i s t r i b u t i o n . Th i s i s u s e f u l f o r p o p u l a t i o n s t u d i e s and f o r
m o n i t o r i n g spec ies occurrence. To da te most of t h e work on l a r v a l
po lychae tes has been concerned w i t h f o u l i ng types, p a r t i c u l a r l y t h e
sedenta ry tube-bu i 1 ders (Serpu l i d a e and Sabel 1 a r i i d a e ) from temperate
waters .
There i s no p rev ious 1 i t e r a t u r e ava i 1 a b l e on Hawai ian po lychae te
l a r v a e . The g r e a t e s t p o r t i o n o f work i n t h i s f i e l d has been conducted
i n Europe, and dea ls w i t h temperate species, t h e i r development, s e t t l e m e n t
behav io r and metamorphosis.
B r i t i s h i n v e s t i g a t o r s o f po lychae te 1 arvae i n c l u d e D. P. Wi lson who
has s t u d i e d t h e l a r v a l development o f S a b e l l a r i i d s (1929; 1968 a,b;
1970a,b; 1976) and o t h e r worms such as t h e sp ion ids , PoZydora c iZiata and
PoZydora hopZura (1928), t h e ne re id , Nereis pelagic (1932), and t h e
cap i t e l l i d , Notomastw Zatericeus (1933) . Wi lson a l s o i n v e s t i g a t e d
t h e i n f l u e n c e o f s u b s t r a t e t ype on t h e s e t t l e m e n t and metamorphosis o f
the capi t e l l i d , Notomastus Zatericeus (1 937) and the ophel i i d , GpheZia
b i c o m i s (1948; 1953; 1954; 1955). Other Br i t i sh researchers ,' such as
de S i l va , 1962; Gee, 1965; Knight-Jones, 1951, 1953; Knight-Jones,
Bailey, and Isacc , 1971; Stebbing, 1972; Williams, 1964; and Wisely,
1960, have studied the chemical s t imulation o f , and behavior during
set t lement of serpul id larvae . Many o f the references dealing with the
larvae of temperate water polychaetes have been included in t h i s paper
as they a r e useful f o r taxonomic purposes.
Although t ropical polychaetes a r e important on r ee f s , the re has
been l i t t l e research on t h e i r larval development, reproductive seasona l i ty
and set t lement behavior. Marsden (1960) describes e igh t species of
polychaete 1 arvae from the Caribbean and Eckel barger (1 975, 1976)
investigated the development of two species of Sabe l la r i ids from the
reefs off the eas tern coast of Florida. These papers a r e among the few
relevent works on t rop ica l o r near t ropical polychaetes and t h e i r larvae .
Additional research on t ropical species i s necessary t o e s t ab l i sh the
e f f ec t s of year-round warm water temperatures, and continuous product iv i ty ,
on the reproductive biology of polychaetes.
With the rapid development of aquaculture in t ropical regions i t i s
imperative t o i ncrease our know1 edge of t ropical polychaete, 1 arvae
par t i cu la r ly those which might be of economic importance as pes ts of
aquaculture organisms o r those t ha t could be u t i l i z ed as a food source
f o r these organisms. Before de ta i l ed s tud ies can be conducted, though,
i t i s necessary t o be able t o iden t i fy the larvae.
Th i s paper i s i n t ended as a compendium o f i n f o r m a t i o n on po lychae te
l a r v a e w i t h p a r t i c u l a r emphasis on those most f r e q u e n t l y encountered
i n Hawaiian p lank ton , i n c l u d i n g a taxonomic key t o t h e fami 1 i e s . T h i s paper,
and t h e references c i t e d i n i t p rov ides a s t a r t i n g p o i n t f o r any f u t u r e
i n v e s t i g a t i o n s o f Hawai ian po lychae te l a r v a e .
MATERIALS AND METHODS
Polychaete l a r v a e were ob ta i ned by two methods: tows w i t h a p l a n k t o n
n e t (mesh apera tu re 1 2 5 ~ ) ; and 1 abo ra to r y f e r t i 1 i z a t i o n of gametes from
r i p e a d u l t s .
Regular sampl ing o f t h e p l ank ton can y i e l d s p e c i f i c i n f o r m a t i o n on
reproduct iv 'e seasona l i t y , eco logy and d i s t r i b u t i o n p a t t e r n s of t h e ,*-
l a r v a e o f c e r t a i n po lychaetes. However, one can seldom f o l l o w t h e
compl e t e development o f a p a r t i c u l a r i n d i v i d u a l . A f t e r numerous sampl ings
o f t h e p l ank ton i t i t u s u a l l y p o s s i b l e t o p i e c e t o g e t h e r t h e l i f e h i s t o r y
o f a spec ies b u t gaps o f t e n remain, p a r t i c u l a r l y i n t h e e a r l y development.
Rear ing o f l a r v a e f r om gametes wi thdrawn f rom mature a d u l t s i s
necessary t o g a i n i n f o r m a t i o n on t h e embryology and development o f a
spec ies. A d isadvantage t o t h i s approach i s t h e d i f f i c u l t y i n m a i n t a i n i n g
uncontaminated c u l t u r e s o f gametes and embryos. "_jl
A f t e r c o l l e c t i o n , 1 arvae were c u l t u r e d i n 60 x 15 mm d isposab le
p e t r i d i shes w i t h a maximum o f t e n l a r v a e / d i s h . Sea water c o l l e c t e d
f r om t h e A la Wai Canal (McCul ly S t . b r i d g e ) a t a depth o f 1m was used
f o r m a i n t a i n i n g t h e l a r vae ; i t was changed every t h r e e days. Upon death
o f a l a r v a , o r development o f b a c t e r i a l o r protozoan c u l t u r e s , s u r v i v i n g
l a r v a e were t r ans fe red t o a new d i s h w i t h c l ean wate r . The f ood source
and tube-bu i l d i n g m a t e r i a l f o r t h e l a r v a e cons i s ted o f o r g a n i c m a t e r i a1
c o l l e c t e d from t h e bot tom o f a s a l t wa te r aquarium.
Larvae were retained i n t h e i r pe t r i dishes during observation under
low magnification (10-30x) with a Binolux dissect ing microscope; s i ng l e
larvae were t ransfered t o a depression s l i d e with water and a cover s l i p
f o r observation a t higher magnification (50-100x) with a Mono1 ux compound
microscope. Photomicrographs were taken w i t h a - 35mm Pentax Spotmatic
camera and a Ricoh camera-microscope adaptor. Photomicrographs and
*. free-hand drawings were made of both l i v e and preserved specimens a t
lOOx magnification. Larvae were observed only f o r sho r t periods of time
t o prevent desiccation and death.
When possible representa t ive larvae were preserved f o r fu tu re
reference. Larvae were prepared f o r f ixa t ion by adding magnesi um s u l f a t e
c ry s t a l s t o the dish containing the larvae un t i l the larvae were relaxed.
They were then preserved e i t h e r in 75% ethanol o r 10% formalin. Whole
mounts of the larvae were made using polyvinyl lactophenol in order t o
A-
render the t i s s u e transparent and the s e t ae read i ly v i s i b l e under the
compound mi croscope.
The important morphological cha r ac t e r i s t i c s of polychaete 1 arvae
are : the number of s e t i ge r s (segments with s e t a e ) ; types of se tae ;
number and color of eyespots; color and pat tern of chromatophores o r
o ther pigmentation; the presence o r absence of dorsa l , ventral o r anal
c i r r i , o r prostomi a1 antennae and ten tacu la r c i r r i . Taxonomic keys f o r
the i den t i f i c a t i on of adul t polychaetes a r e seldom useful f o r the
i den t i f i c a t i on of larvae because of the changes t h a t occur a t metamorphosis.
A glossary of terms has been included as appendix I .
The following papers contain desc r ip t ions and i l l u s t r a t i o n s of a
broad range of polychaete-,larvae and have been u t i l ized during t h e course
of this study and a r e included here f o r t h e i r general usefulness.
These and o the r papers a r e l i s t e d under t h e s p e c i f i c family f o r which
they a r e appl icable : Bhaud, 1967; Blake, 1975 a , b , c ; Fewkes, 1883;
~ r a s s ; , 1959; Hannerz, 1956, 1961 ; Marsden, 1960; No1 t e , 1942;
Thorson, 1946; and Vannucci , 1959. **
A paper t h a t dea l s with t h e rear ing techniques f o r polychaete
l a rvae i s Dean and Mazurkiewicz in Smith and Chanley, 1972.
Key t o t h e Fami l i e s of Common Hawaiian P l ank ton ic Polychae te Larvae
1 Dorsurn b e a r s e l y t r a ( s c a l e s ) (F ig . 1 ) Aphrodi t idae p.10 E l y t r a l ack ing 2
2(1) Parapodium w i t h a paddle o r leaf-shaped d o r s a l c i r r u s ( F i g . 2) Phyl lodocidae p.12 C i r r u s o therwise o r l a ck ing 3
3 (2 ) Comound s e t a e p r e s e n t (Fig. 3) u. Simple s e t a e on ly
Nereidae p .1:4. 4
4(3) Two t o t h r e e p a i r s of b l ack eyespots ; s imple c a p i l l a r y s e t a e may be smooth o r s e r r a t e ( F i g . 5 ) p a i r of grooved pros tomia l p a l p s of v a r i a b l e l e n g t h may be p re sen t ; d o r s a l melanophores p r e s e n t (F ig . 7 ) Spionidae p . i$ Se tae , head s t r u c t u r e s and p igmenta t ion o the rwi se
5(4) Hooded hooks (F ig . 16b) p r e s e n t , eyespo t s r e d Hooded hooks absen t ; eyespots r e d o r b l a c k
6(5) S e t a e a l l hooded hooks; one p a i r of s h o r t WYr pros tomia l p a l p s and a s i n g l e a n a l p r o j e c t i o n ;
two t o t h r e e p a i r s of r e d eyespots p re sen t (F ig . 13) Chae topter idae p.35
S e t a e c o n s i s t of s imple c a p i l l a r i e s and hooded hooks; p ros tomia l and a n a l p r o j e c t i o n s l ack ing ; one p a i r of r e d eyespots (Fig. 15) C a p i t e l l i d a e p 39'
7(5) Three b l ack eyespots ; a n a l c i r r i p r e s e n t (Fig. 17) Ophel i idae p. ,413 One p a i r of r ed eyespots (except i n t rochophore) ; s e t a e a r e g e n i c u l a t e ( ~ i g . 1 9 ) ; a n a l c i r r i absen t Se rpu l idae p. 45
Fami 1 y-Aphrodi t i dae (Scal e worms)
Description of Larva
A typical s i x s e t i g e r l a rva (Fig. 1 ) has a t o t a l of three pa i r s of
c l e a r dorsal e l y t r a ( s c a l e s ) , a kidney-shaped prostomi um with two pa i r s
of black eyespots which may be fused, f i v e antennae, and a p a i r of
prostomial palps. The s e t a e a r e se r ra ted c a p i l l a r i e s , each segment - without an e l y t r a bears a dorsal c i r r u s . There i s a p a i r of anal c i r r i
on the pygidium, these a r e e a s i l y l o s t and so may not be seen. The
number of palps, antennae and c i r r i a r e d i f f i c u l t t o determine in 1 ive
specimens because of rapid movement and they a r e hidden by the e l y t r a .
Remar ks
Adult s ca l e worms a r e benthic de t r i t i vo re s which inhab i t muddy
regions (Day, 1967) and a r e a l so components of rocky i n t e r t i d a l , p i e r a-
p i 1 i n g and wharf communities (Blake, 1 9 7 5 ~ ) .
Genera reported from Hawaii a s adul ts
Arctonoe, Ezazoe, Hamothoe, HoZoZepideZZa, Iphione, ~ernadereZZa,
Lepidas thenia, Lepidonot m, ParaZepidonot us, PoZynoe, Thomore
References t o 1 arvae
Blake, 1975c; Cazaux, 1968; Daly, 1972; Thorson, 1946
Figure I . Aphroditidae: s i x s e t i g e r s t a g e , dorsa l view
A : antenna; A C : anal c i r r u s ; D C : dorsa l c i r r u s ; E ; elytrum;
ES: eyespot; NeP: neuropodium; NeS: neuroseta; NoP: notopodium
NOS: notose ta ; P : palp; Pr: prostomium; Py: pygidium
Family-Phyllodocidae
Description of Larva
A common phyllodocid larva i s i l l u s t r a t e d i n Figure 2 . Present a r e
e igh t s e t i gers, each with paddl e-shaped dorsal c i r r i and compound se tae .
The prostomium is heart-shaped with a pa i r of an te r io r antennae, and one
p a i r of red eyespots. There a r e two pa i r s of tentacular c i r r i and a -
p a i r of anal c i r r i .
Remarks
Thorson (1946) s t a t e s t h a t the posi t ion and number of tentacular
c i r r i can be used as a taxonomic cha rac t e r i s t i c f o r the larvae as i t i s
f o r t h e adul ts . This appears t o be an age dependent cha rac t e r i s t i c
though, and i s only useful in older larvae, such as those ready t o
metamorphose, and not i n younger larvae. '*. Adult phyl lodocids occupy many d i f f e r e n t hab i ta t s , f o r example,
the re a r e the long, slender benthic forms which a re found i n cracks and
crevices ; and sho r t , f l a t t ened forms which a re found in the plankton
(Day, 1967).
Genera reported from Hawaii as adu l t s
EuZaZia, E w i d a , Phy ZZodoce, Prophy Z Zodoce
References t o Larvae
Bhaud, 1967; Cazaux, 1969, 1975; 01 ive , 1975; Thorson, 1946
Figure 2 . Phyllodocidae: e i g h t s e t i g e r s t a g e , dorsa l view.
A : antenna; AC: anal c i r r u s ; DC: do r sa l c i r r u s ; ES: eyespot ;
Pa: parapodium; Pr: prostomiurn; S: s e t a ; TC: t e n t a c u l a r c i r r u s
Family-Nereidae
Desc r i p t i on o f Larva
An example o f a t h r e e s e t i g e r n e r e i d l a r v a (F ig . 3 ) has a square
prostomium w i t h a p a i r o f palps, a p a i r o f antennae and a p a i r o f r e d
eyespots. A lso present a r e a rudimentary jaw and a p a i r o f anal c i r r i .
The most no t i ceab le c h a r a c t e r i s t i c o f t h i s l a r v a l s tage i s t h e l e n g t h of ,*-
t he compound setae ( F i g . 4) which a re approx imate ly h a l f t h a t o f t h e
e n t i r e worm and spread o u t i n a f a n - l i k e fashion.
Remarks
L i f e h i s t o r y s t u d i e s o f ne re ids from temperate waters revea l t h a t
t he a d u l t s swarm, r e l e a s i n g gametes i n t o t h e water column where ex te rna l
f e r t i l i z a t i o n occurs (Reish, 1957). This swarming stage i s c a l l e d a
he teronere id and has been observed i n Hawaiian specimens o f t h e genus *dh.
Ceratonereis (per . ob. ) . A d u l t nere ids a r e ben th i c omnivores, some o f which a re mud burrowers
' ( ~ a y , 1967).
Genera repo r ted f rom Hawaii as a d u l t s
Ceratonereis, Laeonereis, ~amaZycast is , Nereis, Perinereis , PZatynereis,
Pseudonereis
References t o Larvae
Bass and B r a f i e l d , 1972; Berke ley and Berke ley, 1953; Blake, 1975 c;
Dales, 1950; Gi lp in-Brown, 1959; Johnson, 1943; Mazurkiewicz, 1975;
Read, 1974; Roe, 1975; Smith, 1950; Wi lson, 1932
Figure 3. Nereidae: three s e t i ge r s tage , dorsal view
AC: anal c i r r u s ; ES: eyespot; J : jaw; P : palp; Pr: prostomium;
S: s e t a ; TC: t en tacu la r c i r ru s
Figure 4. Nereidae: .-compound seta
Family-Spionidae
The spionids are the most frequently encountered polychaete larvae
in Hawaiian plankton. The important character is t ics to look for in
members of th i s family are: the number of eyespots, and the color and
pattern of pigmentation. Hannerz (1 956, 1961 ) provides taxonomic keys,
a=.
descriptions and i l l u s t r a t ions fo r many of the spionids and i s the best
review of the family.
The descriptions of the larvae tha t follow do not include a l l
stages tha t one might find in the plankton, rather a character is t ic
stage, or stages, i s described. For a more detailed description of a
par t icular species see the reference l i s t a t the end of th i s section.
A key to the species of the Hawaiian PoZydora and PseudopoZydora i s
provided.
Key to the planktonic larvae of the Hawaiian PoZydora and PseudopoZydora
1 Bar-shaped dorsal melanophores present; two pairs of black eyespots (Fig. 7) ~ o ~ ~ d o r a websteri p. 20
Bar-shaped dorsal melanophores absent; three pairs of black eyespots
2(1) Dorsal melanophores consist of small dots forming a random pattern (Fig. 8) PoZydora sociaZis p. 24
Dorsal melanophores are large and stellate (Fig.9) and form rows with an anterior- posterior orientation
3(2) Yellow-white pigment visible under reflected light present; two to four dorsal melanophores/setiger
Yellow-white pigment absent; two dorsal melanophores/setiger (Fig. 9) PseudopoZydora paucibranchiata p.26
4 ( 3 ) Two dorsal melanophores/setiger (Fig. 11) PseudopoZydora antennata p.2,g
Four dorsal melanophores/setiger (Fig. 12) PseudopoZydora puZchra p.31
PoZydora websteri
The three se t iger stage i s the youngest stage of t h i s worm to be
found in the Hawaiian plankton. The body i s l i gh t brown with an orange
g u t . The prostomium i s blunt with two pairs of dorsal black eyespots and
a dorsal medial ridge. There i s a s ingle pair of dorsal bar-shaped
melanophores on se t iger three. The setae a re serrated cap i l l a r i e s and
- extend from each segment to the pygidium. A prototroch and telotroch are
v is ib le (Fig. 5 ) .
By the twelve se t iger stage the prostomium has become more rounded and
a pair of short prostomial palps may be present. There are two dorsal
bar-shaped melanophores/setiger from se t igers three to s i x , two dot-shaped
dorsal melanophores/setiger from set igers seven to twelve and a median
dorsal dot-shaped melanophore on the pygidium (Fig. 6 ) .
I n the fourteen set iger stage larva the prostomium i s rounded, the
la te ra l pair of eyespots have increased in s ize and are comma-shaped, the , -1.
prostomial palps extend to set iger three. The melanophores of set igers
seven to fourteen have expanded and are s t e l l a t e rather than dot-shaped(Fig
Figure 5 . Spionidae : Poiydora websteri , t h r e e s e t i g e r s t a g e , do r sa l view
An: anus; ES: eyespot ; G : g u t ; M : melanophore; Pr: prostorniurn;
Prt: p ro to t roch ; Py: pygidium; S : s e t a ; T : t e l o t r o c h
Figure 6 . Spionidae: PoZydora w e b s t e r i , twelve s e t i g e r s t age ,dor sa l view
ES: eyespot; M : rnelanophore; P : pa lp ; Pr: prostomium; Py:pygidium
S: s e t a
F i g u r e 7 . S p i o n i d a e : PoZydora webster i , f o u r t e e n s e t i g e r s t a g e ,
d o r s a l view
BM: bar-shaped melanophore; ES: e y e s p o t ; P : p a l p ; P r : prostomiurn;
Py: pygidium; S: s e t a ; SM: s t e l l a t e melanophore
PoZydora cf. sociaZis
The l a r v a o f t h i s worm i s t r anspa ren t o r w h i t e w i t h do rsa l p i gmen ta t i on
c o n s i s t i n g o f randomly ar ranged smal l dot-shaped melanophores. The
f i f t e e n s e t i g e r s tage l a r v a has t h r e e p a i r s o f b l ack eyespots, and a
p a i r o f p ros tomia l pa lps which extend t o s e t i g e r f i v e . The se tae a r e
smooth,and i r r i d e s c e n t w i t h those o f s e t i g e r one ex tend ing beyond t h e
.- pygid ium (F ig . 8 ) .
Larvae o f PoZydora sociaZis a r e r e p o r t e d t o have a ye l low-brown
pigment on t h e pyg id ium (Blake, 1969b) ; th is was n o t seen i n t h e Hawai ian
specimens b u t B lake (pe r . comm.) says t h a t t h e Pawai ian fo rm i s e i t h e r
PoZydora soc ia l i s o r a c l o s e r e l a t i v e .
F i g u r e 8. S p i o n i d a e : PoZydora webs ter i , f i f t e e n s e t i g e r s t a g e ,
d o r s a l view
ES: e y e s p o t ; M : melanophore; P : p a l p ; Pr: @rostomium; Pygidium;
S: s e t a
PseudopoZydora paucibranchiata
The eleven set iger larva of ~seudopoZydora paucibranchiata has a
rounded prostomium with three pairs of black eyespots, a median ridge and a
pair of short prostomial palps. A t the junction of each palp with the
prostomium i s a dorsal s t e l l a t e melanophore. There i s a median dorsal
s t e l l a t e melanophore on set iger one and the pygidium as well as two la te ra l
. dorsal s t e l l a t e melanophores/setiger from set igers two to el even (Fig. 9 ) .
The setae are serrated capi l la r ies .
A typical f i f teen se t iger larva i s approximately three times as long
as wide. The body i s tan, the prostomium i s blunt with three pairs of black
eyespots and a pair of prostomial palps may be present. There i s a s ingle
median dorsal s t e l l a t e melanophore on set iger one and the pygidium and two
la te ra l dorsal melanophores/setiger from set iger two to f i f t een (Fig. 10).
On the ventral surface there i s a single median s t e l l a t e melanophore on'
I_ set iger s ix . Ye1 low-whi t e pigment (v is ib le under ref1 ected 1 ight ) appears
on the dorso-lateral region of the prostomium, se t iger one and the pygidium
(not shown in Fig. 10) . On the palps there i s scattered yellow pigment
and small dot-shaped melanophores. The larval notosetae are i r r idescent and
serrated with those of se t iger one being the longest, extending to se t iger
f i f t een .
Figure 9. Spionidae: PseudopoZydora paucibranchiata,eleven setiger stage,
dorsal view
ES: eyespot; MR: medial ridge; P: palp; Pr: prostomium; Py: pygidium;
S: seta; SM: stellate melanophore
Figure 10 . Spionidae: PseudopoZydora pazrcibranchiata, f i f t e en
s e t i ge r s tage, dorsal view
ES: eyespot; M: melanophore; P: palp; Pr: prostomiurn; Py: pygidium;
S: s e t a ; SM: s t e l l a t e melanophore.
PseudopoZydora nntennata
The eight se t iger stage of th i s worm i s approximately twice as long
as wide, the body i s silver-grey. The dorsal pigmentation consis ts of
two s t e l l a t e melanophores/setiger from set igers three to eight and a median
s te l l a t e chromatophore on the pygidium (1 a t e r stages a1 so have a median
s t e l l a t e me1 anophore on the prostomi urn) (Fig. 11 ) . There i s no ye1 1 ow-whi t e
.-". pigment as in PseudopoZydora paucibranchiata. The prostomium i s rounded
w i t h three pairs of black eyespots, the la te ra l two pairs may fuse to form
large compound eyespots. The setae are long, smooth and i r r idescent , with
those of set iger one being more numerous ond longer than those in the
succeeding se t igers . The prostomial palps, when present, a re short , not
extending beyond se t iger two.
Figure 11. Spionidae: PsszdcZopoZydora hntennata, eight s e t i ge r s tage ,
dorsal view
ES: eyespot; Pr : prostomium; Py: pygidium; S : s e t a ; SF: s t e l l a t e rnelanophore
The fourteen setiger stage larva is approximately twice as long as wide,
the body is silver-grey in color with a blunt prostomium and three pairs of
black eyespots. The prostomial palps do not extend beyond setiger three.
There i s a dorsal medial stellate melanophore on the prostomium and the
pygidium as well as four dorsal stellate melanophores/setiger from setigers
"" one to fourteen (Fig. 12). Ventrally there is yellow-white pigment on
the prostomium, at the base of the palps and on the pygidium and on the
middle setigers (not shown in Fig. 12). The setae are long, smooth and
i rridescent.
Figure 12. Spionidae: PsetdopoZydora pulchra, fourteen s e t i ge r
s tage , dorsal view.
ES: eyespot; P : palp; Pr: prostomium; Py: pygidium; S: s e t a ;
SM: s t e l l a t e melanophore
Remarks
Most members of the genus PoZydora lay the i r f e r t i l i zed eggs in
some type of protective capsul e (Hannerz, 1956). Bl ake (1 969b)reported
tha t PoZydora websteri and PoZydora sociaZis form simi 1 a r bead-1 i ke
egg capsules which are deposited within the parent 's tube in a s t r ing ,
with each capsule par t ia l ly connected to the tube wall. In Hawaiian
N- specimens of PoZydora websteri the egg capsules contain an average of
25 eggs and are attached t o the wall of the parent 's tube with the
dorsal surface of the parent facing them. The larvae of both PoZydora
websteri and PoZydora sociaZis are released from the egg capsule a t
the three set iger stage (Blake, 1969b).
The type of egg capsule and age of release are the same fo r PseudopoZydora
paucibranchiata as for the two species of PoZydora (Blake and Woodwick,
The development and settlement behavior of PoZydora websteri has ,I-
recieved special attention because i t bores into oyster she l l s often
making them unmarketable. Further investigation i s necessary to eliminate
i t as a pest of oysters.
Spionids are detr i t ivores which form mucous lined burrows in so f t
substrates such as mud (Day, 1967) or bore into calcium carbonate
substrates lake and Evans, 1973).
Genera reported from Hawaii as adults
Aonides, PoZydora, Prionospio, PseudopoZydora, Spio, Spiophanes
References t o Larvae
Blake, 1969b; B lake and Evans, 1973; B lake and Woodwick, 1975; Casanova,
1952; Day, 1934; Dean and H a t f i e l d , 1963; ~ u g r i n , 1970, 1972; Hannerz,
1956, 1961; H a t f i e l d , 1965; Hopkins, 1958; R u l l i e r , 1960, 1963; Simon,
1967, 1968; Thorson, 1946; Wilson, 1928; Woodwick, 1960
Fami ly-Chaetopter idae
D e s c r i p t i o n o f Larvae
The l a r v a o f Chaetopterus sp. i s approx imate ly Imm l o n g a t t h e
s tage where t h e r e a r e s i x s e t i g e r s i n t h e a n t e r i o r body r e g i o n ( F i g . 13)
The body i s d i v i d e d i n t o t h r e e reg ions by two c i l i a r y bands, and
.- tapers p o s t e r i o r l y i n t o an ana l p r o j e c t i o n (F igs . 13&14). The l a r v a i s
c i l i a t e d and can move r a p i d l y th rough t h e water . On t h e prostomium an
a p i c a l t u f t i s v i s i b l e , and t h e r e a r e two p a i r s o f r e d eyespots. Thorson
(1946) descr ibes a t h i r d p a i r o f r e d eyespots on t h e d o r s a l marg in o f
t h e prostomium o f Clzaetopterus variopedatus b u t t h i s was n o t seen i n t h e
Hawaiian l a r v a . Th i s t h i r d p a i r o f eyes i s much s m a l l e r than t h e o t h e r
two and i t may n o t be n o t i c e a b l e i n t h e a c t i v e l y moving l i v e specimens
and t h e Hawai ian specimen may be a d i f f e r e n t spec ies than t h e one Thorson
*- (1 946) descr ibes . There i s a p a i r o f s h o r t p ros tom ia l pa lps on t h e
do rsa l s u r f a c e o f t h e l a r v a (F igs . 13&14). The se tae a r e hooded hooks.
Remarks
A d u l t chae top te r i ds feed by t r a p p i n g food p a r t i c l e s i n a mucus sac
suspended between t h e parapodia o f t h e m idd le body r e g i o n . T h e i r tube
may be o f sand g r a i n s o r parchment l i k e m a t e r i a l (Day, 1967).
Accord ing t o Thorson (1 946) Chaetopterus variopedatus undergoes
e x t e r n a l f e r t i l i z a t i o n and development o f t h e l a r v a takes p l a c e i n
t h e p lank ton . The l e n g t h o f t ime spent i n t h e p lank ton depends on t h e
a v a i l a b l e food. A f t e r metamorphosis t h e worm becomes benth ic and b u i l d s
a mucous tube.
Genera r e ~ o r t e d from Hawaii as a d u l t s
Chaetopterus, Mesochaetopterus, PhyZZochaetopterus
References t o Larvae
Ehaud, 1966; Marsden, 1960; - Thorson, 1946
Figure 13. Chaetopteridae: Chaetopterus sp. , lateral view
AP: anal projection; AR: anterior region; AT: apical tuft; CB:
ciliary band; ES: eyespot; MR: middle region; P: palp; PR: posterior
region; S: seta
Figure 14. Chaetopter idae: Chaetopterus sp . , dorsal view
AP: anal p ro jec t ion ; A R : a n t e r i o r r eg ion ; AT: ap ica l t u f t ; C t c i l i a ;
C B : c i l i a r y band; ES: eyespot; MR: middle region; P: pa1p;PR:
p o s t e r i o r region; Pr : prostomium
Fami ly-Capi t e l l i dae
Desc r ip t i on o f Larva
The l a r v a o f cap i t e l l a capi ta ta (F ig. 15) i s c y l i n d r i c a l w i t h a
con ica l prostomi urn and a rounded pygidium. A p a i r o f red eyespots
i s l o c a t e d on the prostomium which l acks any antennae, pa lps o r o ther
p r o j e c t i o n s . There a re t h i r t e e n se t igerous segments w i t h simple c a p i l l a r y ,=".
setae (F ig . 16a) i n the a n t e r i o r segments and hooded hooks (F ig . 16b) i n
t he p o s t e r i o r ones. There i s a c i l i a r y band a t t h e j u n c t i o n o f t he
prostomium w i t h the body and a l so a t the j u n c t i o n o f t h e pygidium w i t h the
body. These c i l i a r y bands enable the l a r v a t o move r a p i d l y w i t h i n the
water column. The body c o l o r o f t he l a r v a i s l i g h t brown and the re may be
red, b lack o r b lue pigment spots on the prostomium and pygidium.
Remarks -a.
The eggs o f CapiteZZa capi ta ta f l o a t f r e e i n t he coelom o f the female,
f e r t i l i z a t i o n i s i n t e r n a l . The zygotes a r e ext ruded from the female and
cemented w i t h a ge la t inous ma te r ia l t o t he surrounding mud tube.
Larvae hatch a t t he t h i r t e e n s e t i g e r stage (Thorson, 1946).
Grass1 e and Grassle (1 976) i n v e s t i g a t e d t h e e l e c t r o p h o r e t i c pa t te rns
f o r e i g h t enzymes o f CapiteZZa capi ta ta l a r v a e and found a complex o f
s i x s i b l i n g species t h a t lacked common a l l e l e s and had d i f f e r e n t
l i f e h i s t o r i e s . I n most cases the s i z e of eggs and t h e brood s i z e
d i f f e red and i n a l l cases the t ime spent i n t h e p lank ton v a r i e d
Genera r epor t ed from Hawaii a s a d u l t s
CapiteZk, Dasybranchus, Notomastus
References t o Larvae
~u6 r in and ~ a s s g , 1974; Grass le and Grass le , 1976; Thorson, 1946;
- Wilson, 1933
'fi ,-
Figure 15. Capitellidae: Capitella capitata, thirteen setiger stage,
lateral v iew
An: anus; ES: eyespot; G: gut; Pr: prostomiurn; Prt: prototroch;Py: pygidium
S: seta; T: telotroch
Figure 16. Capitellidae; setae of Capitella cap i ta ta
a: simple capillary
b: simp1 e hooded hook
Family-Opheliidae
Description of Larvae
A ten s e t i ge r larva of PoZyophthaZrnus pictus ( F i g . 17) has a rounded
prostomiurn w i t h three black eyespots. Parapodia a r e rounded and the
s e t a l bundles a r e composed of cap i l l a ry s e t ae with the centra l se ta
longer than those a t the periphery of the bundle (Fig. 17) . The pygidium m-
has three anal c i r r i .
Remarks
Adult ophel i i d s a re elongate and tapered a t both ends; they a re
white o r pink i n color and burrow i n mud and sand. Sand grains and
organic material have been observed i n the guts of ophel i ids indicat ing
t ha t they a r e de t r i t ivores .
-".m,
Genera reported from Hawaii as adul ts
Amandia, Po Zy ophtha Zmus
References t o Larvae
Gugrin, 1971, 1973; Wilson, 1948, 1953, 1954, 1955
Figure 1 7 . Ophel i idae: PoZyopthah us pictus, ten setiger stage,
dorsal view
A C : anal cirrus; ES: eyespot; Pa : parapodium; Pr: prostomiurn; S: seta
Family-Serpulidae
Description of Larva
Trochophores and three s e t i g e r s tage larvae of serpul ids a re often
encountered i n Hawaiian plankton. I t i s seldom possible t o iden t i fy
these l a rvae t o genus o r species because they a re so young, i t i s
necessary t o r a i s e them through metamorphosis. The larva of Spirobranchus -
giganteus i s typical of serpulid larvae and will be used t o i l l u s t r a t e
the larvae of t h i s family. The following descr ipt ion i s based on larvae
reared i n t he laboratory by J . White from gametes obtained from mature
Spirobranchus giganteus collected a t Kahe Point, Oahu.
The ea r ly trochophore i s top-shaped with the pretrochal and posttrochal
hemispheres of equal s i z e (Fig. 18a). The prototroch and apical t u f t
a re v i s i b l e a t t h i s stage. Eight days a f t e r f e r t i l i z a t i o n the posttrochal
region has s t a r t ed t o elongate and a pa i r of red eyespots i s v i s i b l e
"*' dorsa l ly on the pretrochal hemisphere (Fig. 18b). The fourteen day
larva has three setigerous segments (Fig. 19) and the posttrochal region
has continued t o elongate and may have orange-brown pigmentation. From
between f i f t e e n t o twenty days of age the larva will s e t t l e out of the
water column and form a transparent tube (White, 1975). The larvae of
Spirobranchus giganteus have not been reared beyond t h i s ear ly set t lement
s tage (Marsden, 1960; White, 1975).
Remarks
Adult serpul ids a r e tubicolous, suspension feeders. Their calcareous
tubes may be s t r a i g h t and attached only a t the base as in FiZograna
impZexa o r coiled and at tached t o the subs t ra te along the e n t i r e length
of the tube as i n the spi rorbids .
Serpulids, such as Hydroides eZegans and Hydroides dirampha, a r e
often found on marine and harbor s t ruc tures , such as p ie r p i l i ngs , boat '*"
hul l s , and the bottom of buoys (Bailey-Brock, 1976).
Serpul ids display a fo r tn igh t ly reproductive rhythm based on t i d a l
cycles. Garbari ni (1 933) observed t ha t Spirorbis borealis 1 arvae a r e
released and s e t t l e a t t he moons quarters and have es tabl ished t h e i r
tubes before the spring t i de s . Straughan (1968) found t ha t the larvae
of Ficopomatus enigmaticus s e t t l e on the spring t i de s i n the Brisbane
River, Austra l ia .
Most members of t h i s family have one radia l of the branchial crown
"" modified t o form an operculum which i s used t o plug the tube when the
worm i s re t rac ted . The operculum i s a l so used as a brood pouch by some
spirorbids . The methods of brood protection--egg-string, operculum
incubation, and thoracic brood pouch incubation--have been used as a
basis f o r reclass i fying t he spi rorbids (Bailey, 1969).
Genera r e p o r t e d from Hawaii a s a d u l t s
EuZaeospira, Eupomatus, Ficopomatus, F i Zograna, Hydroides, Janua, P i Zeo Laria,
PiZeoZaeospira, PomatoZeios, Protolaeospira, ProtuZa, SerpuZa, Spirobranchus,
Spirorbis, VermiZiopsis
References t o Larvae
Gee, 1962, 1965; Gee and Knight-Jones, 1962; Knight-Jones, 1951, 1953;
'"" Nott and Parkes , 1975; Quie/vieux, 1962; Rothl i s b e r g , 1974; Rul l i e r ,
1960; Sentz-Braconn,ot, 1964; Straughan, 1968; White, 1975; Wisely,
1958, 1960
Figure 18. Serpul idae: Spirobranchus giganteus;
a : trochophore, l a t e r a l view b: e i g h t day old l a r v a , dorsa l view
AT: ap ica l t u f t ; ES: eyespot; G : g u t ; PoH: pos t t rochal hemisphere;
PrH: p re t rochal hemisphere; Prt: p ro to t roch ; T: t e l o t r o c h
(Af te r White, 1975)
Figure 19. Serpul idae : Spirobranchus giganteus, t h r e e s e t i g e r
s t a g e , dorsal view
AT: ap i ca l t u f t ; ES: eyespot ; G : g u t ; PoH: pos t t rocha l hemisphere;
Prd: pre t rochal hemisphere; Prt: p ro to t roch ; S: s e t a ; T: t e l o t r o c h
(Af t e r White, 1975)
ACKNOWLEDGMENTS
My thanks t o Dr. J.H. Brock and John McMahon f o r t h e i r adv i ce
d u r i n g t he course o f t h i s i n v e s t i g a t i o n . I am g r a t e f u l t o
D. Ecker t , J.G. Grovhoug, W. Cooke and Dr. J.H. Brock f o r t h e i r
comments on t h e manuscr ip t . My spec ia l thans t o Dr . James A. B lake
f o r h i s he lp i n i d e n t i f y i n g t h e spionids,and t o Jane t White f o r t h e
l a r v a e o f Spirobranchus giganteus.
BIBLIOGRAPHY
Bailey, J.H. 1969. Methods of brood protec t ion a s a bas i s f o r the r e c l a s s i f i c a t i o n of t h e Spi rorbinae (Serpul idae) . J. Linn. Soc. London, Zoo1.,48:387-407.
Bailey-Brock, J.H. 1976. Habi ta ts of tubicolous polychaetes from t h e Hawaiian Is lands . Pac. Sci.,30:69-81.
Bass, N . R . and A . E . B ra f i e ld . 1972. The l i f e - c y c l e of t h e polychaete Nereis u i rens . J . Mar. Biol . Assoc. U. K. ,53(3) : 701 -726.
Berkeley, E . and C . Berkeley. 1953. Micronereis nunuhoens i s sp . n. : w i t h some notes on i t s l i f e - h i s t o r y . J . Fish. Res. Bd. Canada. ,1 O(2) :85-95.
Bhaud, M . 1966. Etude du diveloppement e t de I '6cologie de quelques l a r v e s de Chaetopteridae. Vie e t Mi1 ieu,17(3) s e r A:1087-1117.
. 1967. Etude du developpement de quelques l a rves d'anni5l ides polych6tes 3 Banyul s-Sur-Mer. Vie e t Mil i e u , l 8 ( 3 ) s e r A: 531 -571.
Blake, J.A. 1969a. Systematics and ecology of she l l -bor ing polychaetes from New England. Am. Zoo1.,9:813-820.
. 1969b. Reproduction and l a r v a l devel opment of PoZydora from northern New Engl and (Polychaeta : Spionidae) . Ophelia, 7:l-63.
. 1975a. The l a rva l development of polychaeta from t h e northern Cal i f o r n i a c o a s t . 11. Nothria e~egans(Fam. Onuphidae) . Ophelia, 13:43-61.
. 1975b. The l a rva l development of polychaeta from t h e norther Ca l i fo rn ia c o a s t . I . C i r r i f o m i a spirobranchus (Fam. C i r ra tu l i d a e ) . Trans. Amer. Micros. Soc., 94(2) : 179-1 88.
. 1 9 7 5 ~ . The l a rva l development of polychaeta from t h e northern Ca l i fo rn ia c o a s t . 111. e ighteen species of e r r a n t i a . Ophelia, 14:23-84.
Blake, J.A. and J.W. Evans.1973. PoZydora and re la ted genera a s borers in moll usc s he1 1 s and other calcareous subs t ra tes . The Vel i ger , 15 (3) : 235-249.
, and K . H . Woodwick. 1975. Reproduction and larval development of PsedopoZydora paucibranchiata (Okuda ) and PsetdopoZydora k m p i (Southern). Biol . Bull ., 149: 109-1 27.
Brock, R . E . and J.H. Brock. 1977. A'method fo r quan t i t a t ive ly assessing the infaunal cornrnuni t y residing in coral rock. Limn. and Ocean.: i n press.
Casanova, L . 1 952. Sur 1 e d6vel oppement de PoZydora antennuta (Claparkde) . Arch. zool . Exp6r. e t G6n. , 89(3) : 95-1 01 .
Cazaux, C . 1968. Etude morphologique du d6vel o pement l a r v a i r e d1ann61 ides polych:.&tes (Bassin d 'Arcachon ! I . Aphrodi t i d a e , Chrysopetal idae. Arch. zool . Exper. e t Ge'n. , 109:477-543.
. 1969. Etude moreholo ique du d6veloppement l a rva i r e d1ann61ides polychstes 9 Bassin dlArcachon) II.Phyllodocidae, Syll idae, Nereidae. Arch. zool . Exper. e t ~ 6 n . ,I 1 O(2) : 145-202.
. 1975. Reproduction e t dgvel oppement l a rva i r e de PhyZZodoce Zaninosa Savigny 1818. Cah. Biol. Mar., 16(4) : 541 -549.
Dales, R . P . 1950. The reproduction and larval development of Nereis d ivers icolor O.F. Muller. J . Mar. Biol. Assoc. U.K., 29(2) :321-360.
Daly, J.M. 1972. The maturation and breeding biology of Hamothoe -imbricata (Polychaeta : Polynoidae) . Mar. Biol . , 12(4) : 53-66.
Day, 1934. Development of scolecole i s fuliginosus ( ~ l a p a r k d e ) . J . Mar. Biol. Assoc. U.K., 19(27:633-654.
Day, J.H. 1967. A monograph of the Polychaeta of souther Africa. Par t 1. Errant ia . Part 2 . Sedentaria. Br i t i sh Museum (Natural History), London. pub. no . 656, 2 vo ls . , 878 pp.
Dean,D. and J.A. Blake, 1966. Life-history of Boccardia hanata (Webster) on the eas t and west coasts of North America. Biol . Bull ., 130(3) :316-330.
, and P . A . Hatfield. 1963. Pelagic larvae of Nerinides cg iZ is ( V e r r i l l ) . Biol. Bull. , 124(2):163-195.
, and M . Mazurkiewicz. 1972. Methods of cu l tu r ing polychaetes. In:Srnith, W.L. and M . H . Chanley ( e d . ) . Culture of Marine 7
Invertebrate Animals. Plenum Press , N . Y .
de S i l v a , B . 1962. Experiments on choice of s u b s t r a t e by Spirorbis 1 arvae. J . exp. Bi 01 . , 39: 483-490.
Eckelbarger, K.J. 1975. Developmental s t u d i e s of t h e p o s t - s e t t l i n g s tages of SabeZZaria vuZgaris (Polychaeta : Sabel l a r i i dae ) . Mar. Biol . , 30:137-149.
, 1976. Larval development and population aspects of t h e reef -bui ld ing polychaete Phragmatopma Zapidosa from t h e e a s t coas t of F lor ida . Bull . Mar. S c i . , 26(2) :117-132.
Fewkes, W.J. 1883. On t h e development of c e r t a i n worm la rvae . Bull . Mus . Comp. Zool . , Harvard , 11 : 167-208.
.- Garbarini , P . 1933. Rhythme d '6mission des l a r v e s chez Spirorbis borealis . C . R . Soc. B io l . , Paris ,T. 11-:1204-1205.
Gee, J.M. 1962. Growth and breeding of Spirorbis r q e s t r i s (Polychaeta : Serpul idae) . J . Zool . , London, 152 : 235-244.
. 1965. Chemical s t imula t ion of se t t lement in l a rvae of Spirorbis r q e s t r i s . Anim. Behav., 13:181-186.
, and E.W. Knight-Jones. 1962. The morphology and l a rva l behavior of a new species of Spirorbis (Serpul idae ) . J . Mar. Biol . Assoc. U . K . , 42:641-654.
Gilpin-Brown, J.B. 1959. The reproduction and l a rva l development of Nereis fueata (Savigny). J . Mar. Biol . Assoc. U . K . , 38:65-80.
=%
~ r a s s 6 , P . P . ( e d . ) . 1959. T r a i t e de zoologie. 5:619-631.
Grass le , J.F. and J .P . Grassle . 1976. S ib l ing species i n t he marine pol 1 u t ion ind ica to r Capitel la (Polychaeta) . Science, 192:567-569.
Gugrin, J-P. 1970. Descript ion des s t ades l a r v a i r e s de Prionospio caspersi Laubier, ( ~ n n g l ide:Polych$te) . Reparti t i o n des l a r v e s de Prionospio en ~ e d i terrannge occidental e . ~ 6 t h ~ ~ ~ 2: 35-40.
. 1971. Modal i t& d 'g levage e t desc r ip t ion des s t ades l a r v a i r e s de Polyophthahus pictus Dujardi n (Anne1 ide : PolychGte) . Vie e t Mil ieu, 22(1) s e r A:143-152.
~u6r-i n , J-P. 1972. Rapports taxonomiques e t dg~eloppement l a r v a i r e de Spio decoratus Bobretzky 1871 (Ann61 ide : ~ o l y c h s t e ) . Cah. Biol . Mar., 13:321-339.
. 1973. Le d6velop ement l a r v a i r e d lAmandia c i r r o s a Fi1 ippi (Anne1 ide : Polychhte ! . Tgthys, 4: 969-974.
, and H . ~ a s s 6 . 1974. Observations sur l e reproduction de Notmastus l a t e r i c e u s Sars 1851 . Cah. Biol . Mar. , 15(3) : 351 -358.
Hannerz, L . 1956. Larval development of the polychaete f a m i l i e s Spionidae Sa r s , Disomidae Mesnil and Poeci lochaet idae n.Fam. i n t h e Gullmar Fjord (Sweden). Zool. Bidr. Uppsala, 31 :1-204.
,- . 1961. Polychaeta: Larvae; fami 1 i e s : Spionidae, Disomidae Poeci lochaet idae. Fiches d ' i d e n t i f i c a t i o n du zooplancton. Conseil In terna t ional pour 1 'Exp lo ra t ion de l a mer, plankton shee t 91 :1-12.
Hatf i e l d , P .A. 1965. Polydora conmensalis Andrews-Larval development and observat ions on a d u l t s . Biol . Bull . , 128(3) : 356-368.
H i a t t , R.W. and D . W . Strasburg. 1960. Ecological r e l a t i o n s h i p s of t h e f i s h fauna on coral r ee f s of t h e Marshall I s l ands . Ecol. Monogr., 30:65-127.
Hopkins, S.H. 1958. The planktonic l a rvae of Polydora webster i Hartman (Annel ida; Polychaeta) and t h e i r s e t t l ing on oys te r s . Bull . mar. s c i . Gulf Caribb. , 8:267-277.
#..
Johnson, M . W . 1943. Studies on t h e l i f e h i s t o r y of t h e marine annel id Nereis v a i l l o s a . Biol . Bull ., 84: 106-1 14.
Knight-Jones, E . W . 1951. Gregariousness and some o the r a spec t s of t h e s e t t i n g behavior of S p i r o r b i s . Jy Mar. Biol . Assoc. U . K . , 30:201-2Z2.
. 1953. Decreased d i sc r imina t ion during s e t t i n g a f t e r prolonged planktonic l i f e in Larvae of S p i r o r b i s b o r e a l i s (Serpul idae ) . J , Mar. Biol . Assoc. U . K . 32(2) :337-345.
, J.H. Bailey, and M.J. I saac . 1971. Choice of a lgae by l a rvae of S p i r o r b i s p a r t i c u l a r l y of S p i r o r b i s sp i ro rb i s .pages 89-104; - In: Fourth European Mar. b i o l . Symp., ed D.J. Crisp,Cambridge Universi ty Press.
Kohn, A. J. 1959. The ecology of Conts i n Hawai i . Ecol . Monogr., 29(1 ) :47-90.
, and M.C. L loyd . 1973. Polychaetes o f t r u n c a t e d ree f l i m e s t o n e subs t ra tes on eas te rn I n d i a n Ocean c o r a l r e e f s : d i v e r s i t y , abundance and taxonomy. I n t . Revue ges. Hyd rob io l . , 58 (3 ) : 369-399.
Marsden, J.R. 1960. Polychaetous anne l i ds f r om t h e sha l l ow wate rs around Barbados and o t h e r i s l a n d s o f t h e West I n d i e s , w i t h notes on l a r v a l forms. Canad. J. Zoo1 . , 38(5) : 989-1 020.
Mazurkiewicz, M. 1975. La rva l development and h a b i t s of Laenereis cuZ mri (Webster) (Pol ychaeta :Anne1 i d a ) . B i o l . Bu l l . , 149(1):186-204.
No l t e , W. 1942. Anne l iden la rven . - I n : Nordisches p lank ton : zoo1 o g i scher t e i l : 59-369.
No t ty J.A. and K.R. Parkes. 1975. Calcium accumulat ion and s e c r e t i o n i n t h e s e r p u l i d po lychae te Spirorbis spirorbis L. a t se t t l emen t . J . Mar. B i o l . Assoc. UK.,55(4):911-923.
01 i v e , P. J .W. 1975. Reproduct ive b i o l o g y o f EuZaZia v i r id i s (Mu1 1 e r ) (Polychaeta: Phyl lodoc idae) i n t h e n o r t h eas te rn U. K. J.Mar. B i o l . Assoc. U.K., 55(2):313-326.
Qu i&vreux , C. 1962. Morpholog ie e t anatomie des l a r v e s de Spirorbis vitreus ( F a b r i c i us) e t Spirorbis maZardi (Caul 1 e r y e t M e s n i l ) . Cah. B i o l . Mar., 3(1):1-12.
. - Randa l l , J.E. 1907. Food h a b i t s of r e e f f i shes o f t h e West I n d i e s .
Proceed. I n t e r n . Conf. Trop. Ocean, 5:665-847.
Read, G.B. 1974. Egg masses and l a r v a e o f Nereis faZcaria ( no te ) . N.Z. J . Mar. Freshwater Res., 8 ( 3 ) :557-562.
Reish, D.J. 1957. The l i f e h i s t o r y of t h e po lychaetous a n n e l i d Beanthes cazchta ( d e l l e Ch ia j e ) , i n c l u d i n g a sumnary o f development i n t h e f a m i l y Nereidae. Pac. Sc i . , 11 ( 2 ) :216-228.
Roe, P. 1975. Aspects o f l i f e h i s t o r y and o f t e r r i t o r i a l behav io r i n young i n d i v i d u a l s of PZatynereis bicanaZicuZata and Nereis ~ i l Z o s a (Anne1 i d a : Polychaeta) . Pac. Sc i . , 29(4) ; 341 -348.
Rothl i sberg , P. C . 1 974. Reproduction i n Spirorbis (SpirorbeZZa) marioni Caul 1 e ry and Mesni 1 (Polychaeta : Serpul i dae) . J . exp. mar. Biol . Ecol . , 15(3) :285-297.
Rul 1 i e r , F . 1960. ~ 6 v e l oppement de SaZmacina d y s t e r i (Huxl ey) . Cah. Biol . Mar., 1:37-46.
. 1963. ~ 6 v e l oppement de PoZydora (Carazzi ) antennuta Cl aparede var. pulchra Carazzi . Cah. Biol . Mar., 4L233-250.
Sentz-Braconnot , E . 1 964. Sur 1 e d6vel oppement des Serpul idae Hydroides noruegica (Gunnerus ) et SerpuZa concharun Langerhans . Cah. Biol . Mar., 5:385-389.
Simon, J.L. 1967. Reproduction and l a r v a l development of setosa (Po1ychaeta:Spionidae). Bull . Mar. S c i . , 398-431.
. 1968. Occurrence of pe lagic l a r v a e of Spio setosa V e r r i l l , 1873. Biol . Bu l l . , 134:503-515.
Smith, R.I. 1950. Embryonic development in the viviparous nereid po'lychhete, Neanthes Zighti Hartman. J , Morph. , 87 :417-455.
Stebbing, A . R . D . 1972. P r e f e r e n t i a l s e t t l emen t of a bryozoan and se rpu l id l a rvae on t h e younger par$sr, of Laminaria fronds. J . Mar. Biol . Assoc. U . K . , 52:765-772.
Straughan, D . 1968. Ecological a spec t s of se rpu l id fou l ing . Aust.Mus. Mag (Nat. H i s t . ) , 16(2):59-64.
Thorson, G . 1946. Reproduction and l a rva l development of Danish marine benthic i n v e r t e b r a t e s , with specia l r e fe rence t o the planktonic l a r v a e i n t he Sound (qresund) . Medd. Komm. Danmarks Fisk. Havund., Ser . Plankton., Bd 4, No. 1 , 523 pp.
Vannucci, M . 1959. Catalog of Marine Larvae. Sao Paulo Brazi l Univ. I n s t i t . Oceanogr.
White, J . 1975. S tudies on t h e cora l boring worm Spirobranchus giganteus ( F . Serpul idae) in Hawaii. Sea Grant summer r epor t , 12 pp.
Williams, G . B . 1964. The e f f e o t of e x t r a c t s of Fueus serrattlsin promoting t h e se t t l emen t of l a r v a e of SfiLrorbis boreal is . J . Mar. Biol . Assoc. U . K . , 44(2):397-414.
Wilson, D.P. 1928. The l a r v a e o f PoZydora c iZ ia ta Johnston and PoZydora hoplura Claparede. J . Mar. B i o l . Assoc. U.K., 15: 567-603.
. 1929. The l a r v a e o f t h e B r i t i s h S a b e l l a r i a n s . J. Mar. B i o l . Assoc. U.K., 16:221-251.
. 1932. The development o f Nereis peZagica Linneaus. J. Mar. B i o l . Assoc. U.K., 18:203-217.
. 1933. The l a r v a l stages o f Notmastus Zatericeus Sars. 3 . Map. B i o l . Assoc. U.K., 18:511-518.
. 1937. The i n f l u e n c e of t h e subst ra tum on t h e metamorphosis A* o f Notmastus l a r vae . J.Mar. B i o l . Assoc. U.K., 22:227-243.
. 1948. The l a r v a l development of GpheZia b icorn i s Savigny. J . Mar. B i o l . Assoc. U. K. , 27(3) : 540-553.
. 1953. The se t t l emen t of GpheZia bicornis Savigny l a r vae . J . Mar. B i o l . Assoc. U.K. 32:209-233.
. 1954. The a t t r a c t i v e f a c t o r i n t h e s e t t l e m e n t o f OpheZia bicornis Savigny. J . Mar. B i o l . Assoc. U.K., 33:361-380.
. 1955. The r o l e o f micro-orggnisms i n t h e s e t t l e m e n t o f OpheZia bicornis Savigny. J . Mar. B i o l . Assoc. U.K., 34(3) : 531 -543.
. 1968a.Some aspects o f t h e development o f eggs and l a r v a e o f Sabellaria aZveoZata (L . ) . J. Mar. B i 01 . Assoc. U. K. , 48(2) : 367-386.
. 1968b. The se t t l emen t behav io r of t h e l a r v a e o f SabeZZaria alveolata ( L . ) . J . Mar. B i o l . Assoc. U. K., 48(2) :387-435.
. 1970a. A d d i t i o n a l observa t ions on l a r v a l growth and se t t l emen t o f Subellaria aZveoZata. J. Mar. B i o l . Assoc. U.K., 50: 1-30.
. 1970b. The l a r v a e o f SabeZZaria spinuZosa and.!their se t t l emen t behav io r . J. Mar. B i o l . Assoc. U.K., 50:33-52.
. 1976. SabelZaria aZwoZata ( L . ) a t Duckpool, No r th Cornwal l , 1975. J. Mar. B i o l . Assoc. U.K., 56(2 ) :305-310.
Wisely, B. 1958. The development of a se rpul id worm, Hydroides norzq ica Gunnerus ( ~ o l y c h a e t a ) . Aust. J . Mar. Freshwater Res., 9 ( 3 ) : 351 -361.
. 1960. Observat ions on t h e s e t t l i n behavior of l a r v a e of t h e tubeworm Spirorbis borealis Daudi n 9 Polychaeta) . Aust. J . Mar. Freshwater Res., 11:55-72.
Woodwick, K . H . 1960. Early l a r v a l development of PoZydora nzchaZis Woodwick, a sp ionid polychaete. Pac. S c i . , 14(2):122-128.
APPENDIX I
GLOSSARY
Anal c i r r u s - elongated p r o j e c t i o n a r i s i n g from t h e l a s t segment o r pygidium (Fig. 1 )
Antenna- sensory p r o j e c t i o n a r i s i n g from the a n t e r i o r o r dorsa l sur face o f t he prostomium (F ig . 2)
Ap ica l t u f t - bundle o r group o f a few c i l i a p r o j e c t i n g f rom the a n t e r i o r most p o r t i o n o f t h e l a r v a (F ig . 18a)
+.b.
C a p i l l a r y seta- h a i r - 1 i ke b r i s t l e ( o f t e n used t o mean a l l long, s lender , tape r ing setae) (F ig . 16a)
Coel om- body c a v i t y
Chromatophore- pigment c e l l o r group of c e l l s (see: Melanophore)
C i r r u s - r e s p i r a t o r y and t a c t i l e appendage o f t h e s e t i g e r (F ig . 2 )
Compound seta- j o i n t e d b r i s t 1 e (F ig . 4 )
Ely t rum- dorsa l sc lae (F ig. 1)
Geniculate seta-seta t h a t i s bent b u t n o t j o i n t e d (F ig . 19)
Hooded hook- seta t h a t i s curved d i s t a l l y and i s covered w i t h a c h i t i nous envelope (F ig . 16b)
Melanophore- b lack pigment c e l l o r group o f c e l l s ( ~ i g . 7 )
Neuropodium- v e n t r a l sec t i on o f parapodi urn (F ig . 1 )
Neuroseta- seta o f t he neuropodiuty (F ig . 1 )
Notopodium- dorsa l sec t i on o f parapodium (F ig . 1 )
Notoseta- seta o f t he notopodium
Palp- p a i r e d p r o j e c t i o n s a r i s i n g from t h e prostomium used f o r food gather ing and t a c t i l e purposes (F igs . 3 and 7 )
Parapodium- 1 a t e r a l , segmental foot-1 i ke project ions bearing s e t ae (Fig. 2 )
Posttrochal hemisphere-the region pos te r io r t o the prototroch(Fig. 18a)
Pretrochal hemi sphere- region an t e r i o r t o the prototroch (Fig. 18a)
Prostomium- t h a t pa r t of the head an t e r i o r t o the mouth (Fig. 5 )
Prototroch- r ing of c i l i a an t e r i o r t o the mouth(Fig. 18a)
Pygidi um- "-
segment bearing the anus (Fig. 5)
Serrated se ta - s e t a with one or more edges notched l i k e a saw (Fig. 5)
Seta- b r i s t l e - l i k e s t r uc tu r e project ing from the parapodium, used fo r locomotion and defense (Fig . 1 )
Set iger- segment bearing s e t a (Fig. 2 )
Simple se ta - unjointed b r i s t l e (Fig. 16)
Stel l a t e - r ad ia t ing , star-shaped (Fig . 9)
Tel otroch- r ing , o r t u f t , of c i l i a near the anus(Fig. 18b)
Trochophore- free-swimming pelagic larval s t age , usually A shaped l i k e a top ( ~ i g . 18a)
PLATES
Plate I
Aphroditidae: six setiger stage larva, dorsal view
Plate I1
Phyllodocidae: eight setiger stage larva, dorsal view
Plate I11
Nereidae: a: three setiger stage larva, dorsal view
b: three setiger stage larva, ventra1,view
- --
Plate I V
Spionidae: Polydora websteri
a: three setiger stage larva, dorsal view
b: twelve setiger stage larva, dorsal view
P l a t e IV
S p i o n i d a e : PoZydora websteri
c : f i f t e e n s e t i g e r s t a g e l a r v a , d o r s a l view
Plate V
Spionidae: Polydora s o c i a l i s - sixteen set iger stage larva, dorsal view
P l a t e VI
Spi on i d a e : ~ s e uZopoZy$ora pawibranckiata
a : t w e l v e s e t i g e r s t a g e l a r v a , d o r s a l view
Plate VI
Spionidae: PseulopoZydora pawibranchiata
b: f i f t e e n s e t i ge r stage l a rva , with prostomial palps, dorsal view
c: f i f t e e n s e t i ge r stage l a rva , without prostomial palps, dorsal view
P l a t e VII
Spi o n i d a e : PseudopoZydora antennata- e i g h t s e t i g e r s t a g e , l a r v a , d o r s a l v iew
Plate V I I I
Spionidae:PseudopoZydora pu2ch.r~ -twelve setiger stage larva, dorso-1 ateral view
Plate I)!
Opheliidae: ~ ~ Z ~ ~ o p h t h a Z m u s - twelve setiger stage larva, dorsal view
Plate X
Capitellidae: Capi te l la capi ta ta
a: Thirteen set iger stage larva, la teral view
b: Thirteen set iger stage larvae "stretching", la te ra l view
P l a t e XI
S e r p u l i d a e : Spirobranchus gigante%
a : Trochophore , l a t e r a l v iew
8: Trochophore , a n t e r i o r - d o r s a l view
Plate XI
Serpulidae: Spirobranchus giganteus
c: three set iger stage larva, la teral view
