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Do Not Remove from the Library 

U. S. Fish and Wildlife Service 

National . Wetland; Research Cenbr 1 

Biological Report 82(11 .I 15) 700 Lalun oodevul* 'rR EL-82-4 

December 1989 ~afayette, Louisiana 70506 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and Invertebrates (South Florida) t I 

REEF-BUILDING TUBE WORM 

Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station 

U.S. Department of the Interior U.S. Army Corps of Engineers 
Biological Report 82(11.115) 

TR EL-82-4 

December 1989 

Species Profiles: Life Histories and Environmental Requirements 

of Coastal Fishes and Invertebrates (South Florida) 

REEF-BUILDING TUBE WORM 

Alexander V. Zale 

and 

Susan G. Memfield 

Oklahoma Cooperative Fish and Wildlife Research Unit 

Department of Zoology 

404 Life Sciences West 

Oklahoma State University 

Stillwater. OK 74078 

Project Officer 

David Moran 

U.S. Fish and Wildlife Service 

National Wetlands Research Center 

1010 Gause Boulevard 

Slidell, LA 70458 

Performed for 

U.S. Army Corps of Engineers 

Waterways Experiment Station 

Coastal Ecology Group 

Vicksburg, MS 39 180 

and 

U.S. Department of the Interior 

Fish and Wildlife Service 

Research and Development 

National Wetlands Research Center 

Washington, DC 20240 
This series may be referenced as follows: 

U.S. Fish and Wildlife Service. 1983-19-. Species profiles: life histories and environmental requirements of 

coastal fishes and invertebrates. U.S. Fish Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR 

EL-82-4. 

This profile should be cited as follows: 

Zale, A.V., and S.G. Memfield. 1989. Species profiles: life histories and environmental requirements of coastal 

fishes and invertebrates (south Florida)--reef-building tube worm. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.115). 

U.S. Army Corps of Engineers, TR EL-824. 12 pp. 
PREFACE 

This species profile is one of a series on coastal aquatic organisms, principally fish, of sport, commercial, or ecological 

importance. The profiles are designed to provide coastal managers, engineers, and biologists with a brief comprehensive 

sketch of the biological characteristics and environmental requirements of the species and to describe how populations 

of the species may be expected to react to environmental changes caused by coastal development. Each profile has 

sections on taxonomy, life history, ecological role, environmental requirements, and economic importance, if applicable. 

A three-ring binder is used for this series so that new profiles can be added as they are prepared. This project is jointly 

planned and financed by the U.S. Army Corps of Engineers and the U.S. Fish and Wildlife Service. 

Suggestions or questions regarding this report should be directed to one of the following addresses. 

Information Transfer Specialist 

U.S. Fish and Wildlife Service 

National Wetlands Research Center 

NASA-Slidell Computer Complex 

1010 Gause Boulevard 

Slidell, LA 70458 

U.S. Army Engineer Waterways Experiment Station 

Attention: WESER-C 

Post Office Box 63 1 

Vicksburg, MS 39180 
CONVERSION TABLE 

Metric to U.S. Customary 

Multiply 

millimeters (mm) 

centimeters (cm) 

meters (m) 

meters 

kilometers (km) 

kilometers 

square meters (m2) 

square kilometers (km2) 

hectares (ha) 

liters (1) 

cubic meters (m3) 

cubic meters 

milligrams (mg) 

grams (g) 

kilograms (kg) 

metric tons (t) 

metric tons 

kilocalories (kcal) 

Celsius degrees (" C) 

inches 

inches 

feet (ft) 

fathoms 

statute miles (mi) 

nautical miles (nmi) 

square feet (ft2) 

square miles (mi2) 

acres 

gallons (gal) 

cubic feet (ft ) 

acre-feet 

ounces (oz) 

ounces 

pounds (lb) 

pounds 

short tons (ton) 

British thermal units (Btu) 

Fahrenheit degrees (" F) 

BY 

0.03937 

0.3937 

3.281 

0.5468 

0.6214 

0.5396 

10.76 

0.386 1 

2.47 1 

0.2642 

35.3 1 

0.0008 1 10 

0.00003527 

0.03527 

2.205 

2205.0 

1.102 

3.968 

1.8("C)+32 

U.S. Customary to Metric 

25.40 

2.54 

0.3048 

1.829 

1.609 

1.852 

0.0929 

2.590 

0.4047 

3.785 

0.0283 1 

1233.0 

28350.0 

28.35 

0.4536 

0.00045 

0.9072 

0.2520 

0.5556 (" F - 32) 

To Obtain 

inches 

inches 

feet 

fathoms 

statute miles 

nautical miles 

square feet 

square miles 

acres 

gallons 

cubic feet 

acre-feet 

ounces 

ounces 

pounds 

pounds 

short tons 

British thermal units 

Fahrenheit degrees 

millimeters 

centimeters 

meters 

meters 

kilometers 

kilometers 

square meters 

square kilometers 

hectares 

liters 

cubic meters 

cubic meters 

milligrams 

grams 

kilograms 

metric tons 

metric tons 

kilocalories 

Celsius degrees 
CONTENTS 

... PREFACE ................................................................................. 111 

CONVERSION TABLE ...................................................................... iv 

ACKNOWLEDGMENTS .................................................................... vi 

NOMENCLATURE/rAXONOMY/RANGE ..................................................... 1 

MORPHOLOGY AND IDENTIFICATION AIDS ................................................. 2 

REASON FOR INCLUSION IN SERIES ........................................................ 2 

LIFE HISTORY ............................................................................ 4 

Spawning ........................................................................... 4 

Larvae .................................................................................. 4 

Juveniles ................................................................................ 5 

Adults .................................................................................. 6 

AGE AND GROWTH CHARACTERISTICS .................................................... 6 

ECOLOGICALROLE ....................................................................... 7 

Feeding Behavior and Food ................................................................. 7 

Predators ................................................................................ 7 

Competitors .............................................................................. 7 

Associated Species ........................................................................ 7 

ENVIRONMENTAL REQUIREMENTS ........................................................ 8 

Temperature ............................................................................. 8 

Salinity ................................................................................. 8 

Substrate ................................................................................ 8 

Depth ................................................................................... 8 

Current ................................................................................. 9 

SuspendedSolids ......................................................................... 9 

Pollution and Contaminants ................................................................. 9 

Dredging ................................................................................ 9 

Beach Nourishment ........................................................................ 9 

LITERATURE CITED ....................................................................... 11 
ACKNOWLEDGMENTS 

Drafts of this species profile were critically reviewed by Kevin Eckelbarger, Harbor Branch Foundation, Inc., Fort 

Pierce. Florida, Michael Hatfield, Kewalo Marine Laboratory, Honolulu, Hawaii, Paul H. Eschmeyer, Editorial 

Office, U.S. Fish and Wildlife Service, Fort Collins, Colorado, and the Coastal Ecology Group, U.S. Army Corps of 

Engineers, Vicksburg, Mississippi. Copies of several rare reports were graciously provided by Walter G. Nelson, 

Department of Oceanography and Ocean Engineering, Florida Institute of Technology, Melbourne, Florida. Sheila 

G. Johnson, Edmon Low Library, Oklahoma State University, provided exceptional assistance with literature 

searches and interlibrary loan. H. Franklin Percival of the Florida Cooperative Fish and Wildlife Research Unit, 

University of Florida, Gainesville, and 0. Eugene Maughan of the Oklahoma Unit provided adminisuative 

assistance. 
Figure 1. Reef-building tube worm: a. feeding position; b. withdrawn into tube. 

REEF-BUILDING TUBE WORM 

Scientific name ........................ Phragmatopoma lapidosa 

Kinberg 1867 

Preferred common name .............. reef-building tubeworm 

(Figure 1) 

Other common names ............................. sand-tube worm, 

honeycomb worm, reef-building polychaete, tubebuilding marine polychaete 

Phylum ................................................................. Annelida 

Class .................................................................. Polychaeta 

Order ................................................................ .Terebellida 

Family .......................................................... ..Sabellariidae 

Reciprocal crosses of P. lapidosa gametes with gametes 

of the northeast Pacific congener, P. californica, suggest 

the two taxa are conspecific subspecies (Pawlik 1988). 

Geographic range: western Atlantic from east coast of 

Florida to Rio Grande do Sul near Santa Catarina in 

Brazil (Hartman 1944; Kirtley and Tanner 1968; 

DeJorge et al. 1969); also present, but rare, along 

coast of Gulf of Mexico (Potts 1979). Distribution 

in Florida extends about 320 km from Cape 

Canaveral, Brevard County, to Key Biscayne, Dade 

County (Kirtley 1966; Kirtley and Tanner 1968). 

Typically intertidal or subtidal (i.e., within or just 

seaward of the surf zone) along exposed beaches 
subject to high-energy wave action (Multer and 

Milliman 1967; Kirtley and Tanner 1968; DeJorge et 

al. 1969), but present to depths of 100 m (Kirtley and 

Tanner 1968). Present also along channels at inlets 

(e.g., Sebastian, Fort Pierce, and St. Lucie Inlets in 

Florida) characterized by swift tidal currents (Gore et 

al. 1978). The distribution of the reef-building tube 

worm in southern Florida is illustrated in Figure 2. 

MORPHOLOGY AND IDENTIFICATION AIDS 

Sabellariids are obligate agglutinated-sand 

tube-dwelling colonial polychaetes (as opposed to 

annelids that secrete calcareous or membranous tubes) 

that have elongate, segmented, cylindrical bodies divided 

into four parts (proceeding posteriorly): the "head or 

opercular stalk (consisting of a prostomium or pre-oral 

region fused to a peristomium or post-oral region), and 

the parathoracic, abdominal, and caudal regions. The 

head of the reef-builing tube worm has an anterior 

opercular disk that serves as a "stopper" when the worm 

retracts into its tube. An array of hard, clawlike paleae 

(setae) on the operculurn are used to manipulate sand 

grains being cemented into the tube. Two rows of 

ciliated feeding tentacles run lengthwise ventrally along 

the head. The ventral mouth, with its pair of feeding 

palps, lies behind and between the rows of feeding 

tentacles. The lower lip of the mouth is surrounded by 

the horseshoe-shaped building organ that supplies the 

proteinaceous cement for tube construction. The 

parathoracic region consists of three segments, each with 

biramous parapodia (fleshy lateral appendages, each with 

a dorsal and ventral cirrus), simple branchiae (gills), and 

a ventral muscular plate. About 32 segments make up 

the abdomen; these possess notopodia (dorsal branches 

of the parapodia), neuropodia (ventral branches of the 

parapodia), branchiae, and numerous small hook-like 

uncini (setae) on the parapodia that anchor the body to 

the surrounding tube wall. The cylindrical caudal region 

terminates in the anus. Maximum total length is about 30 

to 40 mm. Worms 15 to 25 mm long are 30 to 40 mg wet 

weight. The preceding description was summarized from 

Kirtley (1966, 1968), Kirtley and Tanner (1968), DeJorge 

et al. (1969), and Parker (1982). 

In southeastern Florida, identification of the 

reef-building tube worm is facilitated by recognition of 

the unique mound-like reefs built up by successive 

colonizations of the worms. These may range from 

fist-sized lumps attached to pilings to extensive reefs 

several hundred meters wide and several kilometers long, 

paralleling the beach (Kirtley 1966; Kirtley and Tanner 

1968). ~ndivihual heads are typically low and rounded, 

resembling cushions. They may rise as much as 2 m 

above the surrounding substrate, but the "living" portion 

is seldom as much as 1 m high (Kirtley 1966). The reefs 

are generally a dark, drab brown resulting from the 

aggregate coloration of sand, shell hash, and mineral 

grains from which they are constructed. The mounds are 

friable and easily broken by hand. The vertically 

arranged, parallel worm tubes give the reefs a 

hone comb appearance. Tube densities average about 3 5/cm (Multer and Milliman 1967). Internal diameters of 

the tubes are about 2 to 4 mm; external diameters at the 

aperture range from 4 to 10 mm, depending on worm 

density (Multer and Milliman 1967; Kirtley 1968). 

Individual tubes are curved but intertwine little (Multer 

and Milliman 1967). A hood-like protrusion extends 

partly around the rim of each tube (Kirtley 1966; Multer 

and Milliman 1967). The lower ends are sealed. Kirtley 

(1966) stated that under optimal conditions, individual 

tubes reach a maximum length of about 10 cm, but Gram 

(1968) reported that tubes are generally 15 to 30 cm long. 

REASON FOR INCLUSION IN SERIES 

The reefs constructed by reef-building tube worms 

along the southeast coast of Florida are significant both 

geologically and biologically. Reefs of the sand tubes of 

reef-building tube worms extend within their geographic 

range for hundreds of kilometers of coastline (Kirtley 

and Tanner 1968). The ability of the worms to thrive 

under high-energy breaker conditions and to extend their 

colonial tube masses upward and seaward by extraction 

and agglutination of littoral drift materials makes them 

important vectors in coastline development. Beachrock, 

converted from the reefs, and sand impounded on their 

landward sides provide for actual progradation of 

beaches (Kirtley and Tanner 1968). By sorting out flat 

shell fragments and the heavier suspended particles in 

littoral drift, tube construction by the worms results in the 

retention of beach sediment (Emery 1963; Multer and 

Milliman 1967; Gram 1968). The cracks and crevices of 

the reefs act as traps for sediment and shell fragments, 

thereby further contributing to sediment retention (Gram 

1968). Being wave resistant, the reefs protect the shore 
CAPE ROMANO 

THOUSAND ISLAND 

KILOMETERS 

Figure 2. Distribution of the reef-building tube worm in south Florida. 

3 
against wave attack and retard erosion (Multer and 

Milliman 1967; Mehta 1973). It appears likely that 

reef-building tube worms are at least in part responsible 

for the formation and maintenance of beaches and barrier 

islands in southeast Florida (Kirtley and Tanner 1968). 

The reefs are the primary basis for an elaborate and 

stable marine community (Kirtley and Tanner 1968; 

Narchi 1973. 1974; Gore et al. 1978; Gilmore et al. 1981; 

van Montfrans 1981). They provide hard and stable 

substrate, shelter, and food, and thereby allow many 

species to inhabit the surf zone, an area where most 

would normally be unable to survive (Gore et al. 1978). 

LIFE HISTORY 

Spawning 

Florida populations of reef-building tube worms may 

spawn semicontinuously (without a seasonal trend) 

throughout most of the year; Eckelbarger (1976) 

recovered viable gametes from worms throughout the 

year; he collected larvae in plankton tows in February, 

August, and October; and he observed larval settlement 

in May, September, and December. Gore et al. (1978) 

observed settlement in March. This semicontinuous 

spawning periodicity differs markedly from the polytelic 

spawning (with a seasonal trend) of all other sabellariids 

(Smith and Chia 1985). 

Fertilization occurs in the water (Kirtley 1966). Males 

release sperm into their sand tubes through a series of 

bilaterally arranged nephridia throughout the 

gamete-bearing abdominal segments; sperm are expelled 

into the water column in short bursts by rapid withdrawal 

of the head region into the tube (Eckelbarger 1984). 

Initiation of sperm release by one male generally 

stimulates releases by adjacent males; females release 

eggs in response to the presence of sperm in the water 

(Eckelbarger 1984). Mature sperm have distinctive long, 

tapering, curved acrosomes and laterally displaced 

flagella (Eckelbarger 1984). Total length of sperm is 

about 42 pm; the head, including the acrosome, is about 

6 pm long (Eckelbarger 1976). Females expel eggs from 

their tubes much as males expel sperm (Eckelbarger 

1984). Diameters of spawned oocytes are 97 to 103 pm 

(Eckelbarger 1976). The eggs are sticky and adhere to 

sand grains upon expulsion; this property may be an 

adaptation to restrict transport in the turbulent surf zone 

(Eckelbarger 1984). 

Larvae 

Larvae of the reef-building tube worm are common in 

the nearshore plankton along the east coast of Florida 

(Eckelbarger 1976) and may be an ecologically 

significant component of the planktonic community 

(Kirtley and Tanner 1968). A key to larvae of Florida 

sabellariids was published by Eckelbarger (1977). Mauro 

(1975) and Eckelbarger (1976) described in detail 

development of reef-building tube worm larvae 

maintained in the laboratory. The following description 

is drawn largely from Eckelbarger (1976); larvae were 

maintained at 21 to 23 "C in circulating or aerated 

cultures. Descriptions of larvae derived by scanning 

electron microscopy (SEM) by Eckelbarger and Chia 

(1976) confirmed Eckelbarger's (1976) light microscopy 

observations, except that SEM revealed the presence of 

"sensory tufts" on the dorsal surface of the larval 

tentacles; these may play a role in the selection of a 

substrate on which to settle. 

Freshly shed ova are irregular in shape and have a 

clear, conspicuous germinal vesicle. Within minutes, the 

eggs become round and develop a wrinkled vitelline 

membrane. The germinal vesicle breaks down 10 to 15 

min after fertilization. The fist and second polar bodies 

appear in about 50 to 55 and 60 to 65 min after 

fertilization and the first and second cleavages occur 

within 75 to 78 and 80 to 85 min of fertilization. From 

10 to 12 h after fertilization, the embryo loses its sticky 

coat and hatches as a simple top-shaped trochophore that 

possesses a prototroch (ciliated girdle) and an apical tuft 

consisting of short, fine cilia. At 17 to 20 h, a single 

reddish eyespot is formed and a stiff cilium appears 

posteriorly. At 20 to 21 h, the larva develops a pair of 

small barbed provisional setae that begin to protrude 

through the body wall on each side. At this stage, the 

larva is opaque and granular in appearance; groups of 

irregular, yellow-green pigment specks are scattered over 

its surface. At 22 to 24 h, 3 pairs of provisional setae are 

present, distinct chromatophores have formed, and a 

mouth and digestive system have developed; active 

feeding begins at this stage. At 40 to 42 h, the larva is 

more elongate and shows faint indications of 

segmentation. It has 4 to 7 provisional setae on each side 

and two eyespots; a second ciliated girdle, the telotroch, 
circles its anus. At about 6 days, the larva is clearly sites (Kirtley 1966). A wide variety of natural and 

elongate and has 4 eyespots and as many as 20 artificial settlement sites are used (Kirtley 1966, Multer 

provisional setae. Between 7 and 10 days, the larva and Milliman 1967; Eckelbarger 1976). Existing worm 

develops a pair of dorsal tentacle buds and abdominal reefs, both "dead" and "living," are perhaps the most 

uncini. Three clearly defined parathoracic segments and common attachment sites (Eckelbarger 1976). 

3 less clearly defined abdominal segments are present. At Established reefs are therefore essentially permanent. 

12 days, the tentacles have lengthened and dorsal Settlement occurs over the entire surface of dead, 

parapodial lobes of the parathoracic segments are clearly wave-eroded worm mounds, but only between the 

defined; each has 4 setae. One to three primary openings of existing and occupied adult tubes of sparsely 

(settling) paleae and two pairs of opercular spines appear. populated reefs. No recruitment occurs on the surfaces 

At about 19 days, the tentacles are about half the length of mounds consisting of closely packed tubes of adults; 

of the body and have ciliated food grooves; segmentation successful settlement on such mounds is limited to their 

of the robust larva is distinct. At this stage, the larvae periphery (Eckelbarger 1976). Adult and juvenile worms 

alternately crawl over or swim close to the surface of the use their operculum to scrape away algal growths, 

substrate. During crawling, they are oriented fouling organisms, and debris located around the 

head-downward and frequently contact the substrate with openings of their tubes; this activity probably serves to 

the mouth and tentacles. preclude the settlement of new larvae (Eckelbarger 

1976). 

Juveniles 

Upon selection of a site, the metamorphosing larva 

Metamorphosis and ~c%lement occur 14 to 30 days secretes and attaches a cylindrical, muco-proteinaceous 

after fertilization in c~culating Or aerated cultures at 21 tube to which it begins cementing small fragments; the 

to 23 "C; length ranges from 0.6 to 1.0 mm at settlement initial fragments often consist largely of small, dark 

(Eckelbarger 1976)- MemoVhosis usually requires the grains of heavy minerals (Kirtley 1966; Eckelbarger 

Presence of conspecific tubes or sand and is triggered by 1976). Tubes of juveniles can therefore be readily 

the presence of free fatty acids, a requirement which distinguished from those of adults by their color from a 

~ccounts for the gregariousness of the species (Pawlik considerable distance; adults use larger, lighter-colored 

1988). Pawlik's study (1988) showed that when larvae sand grains and shell fragments. Multer and Milliman 

were cultured with sand that Came from conspecific (1967) and Gram (1968) demonstrated that the median 

tubes, frequency of memorphosis was 50% at 15 "C size of particles incorporated in the tubes increases with 

about 28 hours after hatching and also 50% at 20 'C worm size. Other materials used by young worms 

about 17 hours after hatching. include small quartz grains, foraminifera1 tests, sponge 

spicules, and silt (Kirtley 1966). Larger worms use 

correspondingly larger materials including ostracod 

MetamoVhosis involves an elongation of the body and carapaces, small mollusk shells, fragmented pieces of 

dramatic changes in the head region: the tentacles rotate larger mollusk shells, large quartz grains, large 

anteriorly until they project forward, the provisional foraminiferans, and echinoid spines (Kinley 1966; 

setae are replaced by 6-10 pairs of primary paleae, the Multer and Milliman 1967). Generally, worms prefer 

entire head shrinks in relative size, the building organ sediments of 250 to 500 in diameter (Main and 

appears around the mouth, and various appendages Nelson 1988a) or 125 to 500 pm (Multer and Milliman 

develop on a umber of segments (Eckelbarger 1976). 1967) for tube construction; depending on the diameters 

The telotroch is still Present at this stage but the of available materials, they therefore preferentially 

prototroch has disappeared. After settlement, concentrate finer or coarser grains than they would if 

development continues rapidly; within one month, the selection were random (Main and Nelson 1988a). 

juvenile closely resembles the adult worm (Eckelbarger Significant amounts of sediment finer than 62 pm in 

1976). diameter (silt) are incorporated into tubes, probably 

serving as "mortar" to fill cracks between larger grains 

Upon settling, the larva actively moves over the (Multer and Milliman 1%7). Flat mollusk-shell 

substrate, presumably evaluating possible attachment fragments are typically used to line the inner wall of the 
tube; more rounded grains are incorporated in the outer 

layers (Multer and Milliman 1967). Platy shell 

fragments are incorporated much more frequently than 

quartz sand grains, perhaps because the shell fragments 

are less dense and less spherical in form and hence more 

easily suspended in the water column (Main and Nelson 

1988a). Construction materials are grasped from the 

water column by the oral tentacles or opercular paleae, 

passed to the building organ, coated with proteinaceous 

cement, and implanted into the tube with the opercular 

paleae Wrtley 1966). 

The size, shape, and orientation of the tube are 

influenced by hydrodynamic and sedimentary conditions 

prevailing at the site and the presence or absence of other 

organisms or obstructions (Kirtley 1966). Kirtley (1966) 

speculated, on the basis of few data, that strong positive 

correlations exist between worm size, intensity of 

turbulence, and available particle size. The worms 

constantly enlarge and repair their tubes, especially the 

delicate "hood" at the aperture. Damaged tubes are 

quickly repaired; human footprints on reefs are not 

detectable after 24 h (Kirtley 1966). 

Adults 

Reef-building tube worms are dioecious. Populations 

are composed of equal proportions of males and females 

(Eckelbarger 1976). Sex products first develop in both 

sexes about 6 to 8 weeks after larval settlement and the 

worms are fully mature after 4 months (Eckelbarger 

1976). Sexes are easily distinguished; the 

gamete-containing abdominal segments of mature males 

and females are creamy-white and steel-blue 

respectively, corresponding to coloration of sex products 

(Kirtley 1966, 1968; DeJorge et al. 1969; Eckelbarger 

1976). The sexes are otherwise morphologically 

identical. The testes and ovaries are bilaterally arranged 

organs in the abdominal segments (Eckelbarger 1979, 

1984). Eckelbarger (1979, 1984) described oogenesis 

and spermatogenesis. Removal of sexually mature 

worms from their tubes induces expulsion of gametes 

(Kirtley 1966; DeJorge et al. 1969; Mauro 1975; 

Eckelbarger 1976), and thus facilitates artificial 

fertilization in the laboratory. 

AGE AND GROWTH CHARACTERISTICS 

No information is available on the longevity of 

reef-building tube worms in Florida, but Kirtley (1966) 

suggested that "it does not seem probable that the 

organisms would survive for more than one or two 

years." However, Wilson (1971, 1974) estimated that the 

life span of a similar species, Subellaria alveolata, in 

southwestern England is commonly 5 years and may 

exceed 10 years. 

Eckelbarger (1976) monitored growth rates of newly 

settled juveniles at 2-week intervals at Walton Rocks, 

Martin County, Florida. On 26 March, modal length of 

the population was less than 1 mm, and no worms 

exceeded 2 mm in length. Modal lengths were 2-3, 3-4, 

and 4-5 mm on 14 April, 25 April, and 8 May. 

Corresponding ranges were 1-5, 2-7, and 2-8 mm. 

Average length 6 weeks after settlement was 5.2 mm. 

Eckelbarger (1976) also monitored monthly growth 

rates of a population at Hutchinson Island in Martin 

County. The colony probably settled in early September. 

Modal lengths (ranges in parentheses) were as follows: 

October, 5 mm (1-12); November, 5 mm (2-10); 

December, 9 mm (2-14); January, 8 mm (2-16); February, 

13 mm (6-22); March, 15 mm (10-26); April, 15 mm 

(8-24); and May, 15 mm (8-22). Mean length in May 

was 15.5 mm. Tube lengths of these animals were 

1 .O-1.7 cm on 30 October, 1.8-2.5 cm on 12 November, 

and 1.5-6.0 cm on 11 December. 

The mean daily growth increment of worm tubes was 

1.45 mm at Punta Moron, Venezuela (Merida and 

Penchaszadeh 1982). Accretion rates of tubes of worms 

from this population in the laboratory were 2.03, 3.18, 

and 0.97 mm per day at 22, 26, and 30 'C, respectively. 

Gore et al. (1978) reported that 6 months after settlement 

in March, new reefs were indistinguishable from older 

colonies that had been established in previous years. 

Kirtley (1968) reported that a newly settled reef attained 

a thickness of about 25 cm within 6 weeks. 

Maximum total length is about 40 mm (Kirtley and 

Tanner 1968). Body length varies annually and by 

locality, perhaps depending on food availability or 

physical conditions; tube diameters and body lengths are 

highly correlated (Eckelbarger 1976). 
ECOLOGICAL ROLE Associated Species 

Feeding Behavior and Food 

Kirtley (1966) described the feeding behavior of adult 

reef-building tube worms. The organism lies on its 

dorsum, partly extended into the "hood" of its tube. The 

operculum is turned backward and downward, and the 

ciliated prostomial tentacles are extended upward and 

outward. Water currents produced by the cilia convey 

small suspended particulate matter toward a rapidly 

revolving bolus of mucus just above the mouth. The 

bolus is periodically ingested. The worms also feed on 

algae and other organisms encrusting sand and shell 

fragments. The fragments are grasped with the opercular 

paleae and then directed with the aid of the prostomial 

tentacles to the mouth where the organic materials are 

removed. If the fragment is suitable for tube 

construction, it may then be passed along to the building 

organ. The prostomial feeding tentacles may also be 

used to seize small sand particles and transport them 

toward the building organ for tube building. Feeding and 

tube-building are highly integrated and complementary 

functions. 

The food of reef-building tube worms consists 

primarily of planktonic microorganisms, including 

diatoms, forarniniferans, and algae; encrusting organisms 

adhering to sand and silt are also eaten (Kirtley 1966). 

Predators 

Reef-building tube worms are eaten by crustaceans 

(including the grapsid crab Pachygrapsus transversus 

and the xanthid crabs Mennipe nodifrow, Pilumus 

dasypodus, and Panopeus bermudensis), gastropods, and 

fishes (Kirtley 1966; Gore et al. 1978). The worms 

appear to be the primary forage of a number of species of 

crabs that live on or within the reefs (Gore et al. 1978). 

Competitors 

Barnacles (Tetraclita squamosa) colonizing worm reefs 

compete with reef-building tube worms for space (Multer 

and Milliman 1967). The crab Pachycheles monilifer, 

which is an abundant inhabitant of worm reefs in 

southeastern Florida (Gore et al. 1978), feeds on 

suspended material and plankton and may compete with 

reef-building tube worms for food. 

Worm reefs are the primary basis for an elaborate and 

stable marine community that includes crustaceans, 

mollusks, sponges, bryozoans, anthozoans, and fishes 

(Kirtley and Tanner 1968; Narchi 1973, 1974; Gore et al. 

1978; Gilmore et al. 1981; van Montfrans 1981). Reefs 

provide hard and stable substrate, shelter, and food, 

thereby allowing many species to inhabit the surf zone, 

in which most of these organisms would normally be 

unable to survive (Gore et al. 1978). Crustaceans, many 

of them cryptic and slow moving, make up about 90% of 

the macroinvertebrate fauna associated with worm reefs 

in southeastern Florida (Gore et al. 1978). 

Gore et al. (1978) collected 96 species (in 52 genera 

and 22 families) of decapod and stomatopod crustaceans 

within, on, or associated with worm reefs in southeastern 

Florida. In contrast, only six species inhabited the 

surrounding surf zone. The six most abundant species 

associated with the reefs, about 80% of the total number 

of individuals collected, were the porcellanid crab 

Pachycheles monilifer; the grapsid crab Pachygrapsus 

transversus; the alpheid pistol shrimp Synalphus 

fritzmwlleri; and the xanthid crabs Menippe nodifrons, 

Pilumnus dasypodus, and Panopeus bermudensis. Two 

of these, Pachycheles monilifer and M. nodifrons, are 

restricted to worm reefs in southeastern Florida. 

Narchi (1973, 1974) found the bivalves Hiatella solida 

and Petricola typica living in worm reefs on the coast of 

Brazil; worm reefs appeared to be the preferred habitat of 

Petricola. 

Gilmore et al. (1981) collected a total of 107 species of 

fish from worm reef habitats in Florida; adjacent surf 

zone habitats yielded 91 species. Collections in the 

worm reef were numerically dominated by two demersal 

species (hairy blenny Labrisomus nuchipinnis and molly 

miller Scartella cristata) and three semi-demersal species 

(spottail pinfish Diplodus holbrooki, porkfish 

Anisotremus virginicus, and sailor's choice Haemulon 

parrai). A total of 40 fishes of commercial or sport value 

were "frequent," "common," or "abundant" in the worm 

reef habitat, compared with 35 in the adjacent open surf 

zone; 27 of these species were found excIusively (at 

these frequencies) in the reef habitat and 22 in open 

habitats. Although the species richness of fishes of the 
worm reef habitat is not remarkably higher than that of 

the adjacent open surf zone, the large number of 

habitat-specific species indicates that the presence of 

worm reefs substantially enhances the overall diversity of 

commercially and recreationally important fishes in the 

beach zone. 

ENVIRONMENTAL REQUIREMENTS 

Temperature 

Gilmore et al. (1981) reported that water temperatures 

at reef-building tube worm colonies in southeastern 

Florida ranged from 18 to 27 "C. Water temperatures at 

reefs studied by Gore et al. (1978) ranged from 11 to 

32 'C over a 2-year period. However, these extremes are 

probably exceeded within tubes of reefs exposed at low 

tide during exceptionally hot or cold weather. Extreme 

air temperatures may stress reef-building tube worms in 

intertidal reefs. Wilson (197 1) observed massive die-offs 

of Sabellaria alveolata after severe frosts in 

southwestern England; also, S. vulgaris has suffered 

heavy winter mortality in Delaware Bay (Curtis 1973). 

Eckelbarger (1976) believed that elevated summer 

temperatures may have resulted in the death of a 

reef-building tube worm colony in Florida. The tropical 

and subtropical distribution of the reef-building tube 

worm suggests that it is intolerant of temperate climates. 

Cape Canaveral, the northern extreme of the range of the 

species, marks the approximate center of a transition 

zone between the warm-temperate Carolinian and 

tropical Caribbean faunal regions (Briggs 1974; Gilmore 

1977). 

Eckelbarger (1976) maintained experimental cultures 

of artificially fertilized reef-building tube worm eggs at 

temperatures of 10 to 35 'C. No development occurred 

at the extreme temperatures, and 48%, 65%, 82%, 95%, 

and 47% of the eggs developed at 15, 18,21,25, and 

30 'C, respectively. Eckelbarger (1976) concluded that 

larval development was optimal at 24 to 26 'C. 

Salinity 

Water salinities measured at reef-building tube worm 

reefs studied by Gore et al. (1978) and Gilmore et al. 

(1981) ranged from 28 to 39 ppt. In the laboratory, the 

worms survived at salinities as low as about 10 ppt for 

several days (Mauro 1977). This tolerance is probably 

adaptive in southeastern Florida, where heavy rains and 

freshwater runoff may appreciably dilute salinities within 

worm tubes between high tides. 

Substrate 

Stable settlement substrate is a critical environmental 

requirement of reef-building tube worms. Beaches 

composed entirely of shifting sands afford larval worms 

no opportunity for settlement. Unstable objects subject 

to rolling or burial by sands are unsatisfactory. However, 

a wide variety of natural and artificial substrates can be 

colonized, including living and dead shells of mollusks 

and horseshoe crabs (Limulus polyphemus), coquina 

rock, sea walls, piers, jetties, peat, and beach debris 

(Kirtley 1966; Multer and Milliman 1967; Eckelbarger 

1976). Existing "living" and "dead" worm reefs are 

common attachment sites (Eckelbarger 1976). Complete 

removal of a reef or reef sand will probably delay 

recolonization considerably because the larvae usually 

require, for metamorphosis, a chemical stimulus 

produced by conspecific worms (Pawlik 1988). 

In addition to providing stable settlement, substrates 

surrounding the colonization site must be composed of 

sand and similar-sized particles suitable for tube 

construction (Multer and Milliman 1967). Habitats such 

as exposed rocky shorelines that have adequate wave 

action and stable substrates are unsuitable environments 

for reef-building tube worms because they lack the 

amounts of suspended particles needed for tube building. 

Depth 

Habitats occupied by reef-building tube worms are 

primarily intertidal (DeJorge et al. 1%9) and nearshore 

(Kirtley and Tanner 1968). Kirtley (1966) suggested that 

optimal habitat extends from mid-tide level to a depth of 

about 2 m. Reefs occasionally occur above mid-tide, but 

only if suitable substrate and strong wave action are 

present (Kirtley 1966). At depths greater than 2 m, wave 

action is generally insufficient to maintain the required 

turbulent conditions. However, colonies have been 

reported to occur to depths of 100 m (Kirtley and Tanner 

1968), perhaps at sites with strong submarine currents. 
Current 

Reef-building tube worms require constant 

high-energy wave action to supply food, remove 

metabolic wastes, and maintain the suspension of sand 

grains and other particles for tube building (I(lrtley 1966; 

Multer and Milliman 1967). Average breaker heights 

along the east coast of Florida where worm reefs are well 

developed are 50-75 cm (Kirtley and Tanner 1968). 

Worm reefs may also form at the mouths of inlets where 

tidal currents are strong enough. South of Dade County, 

the turbulent, silt and sand laden conditions required by 

the worms do not exist; such habitats foster the formation 

of coral reefs. 

Suspended Solids 

Reef-building tube worms showed no indication of a 

negative response to experimental suspended-solid levels 

as high as two orders of magnitude greater than 

maximum levels reported from surf zones in Florida 

(Nelson and Main 1985). Habitats having the intense 

turbulence and shifting-sand (surrounding) substrate 

required by reef-building tube worms are high in 

suspended solids. However, high suspended solid loads 

alone may not provide habitat suitable for reef-building 

tube worms if particle size of the suspended materials is 

too small. Habitats having high silt loadings only are 

unsuitable; sand-sized particles must be present. 

Pollution and Contaminants 

Mulhern (1976) examined the short-term tolerances of 

adult reef-building tube worms in their tubes to three 

refined fuel oils mixed with sea water in the laboratory. 

The estimated 48 h LCso for kerosene was 44% (by 

volume); mean mortality rates after 48 h exposures to 

lo%, 20%, 30%, 38%, and 46% concentrations were 5%, 

18%, 38%, 55%, and 61%, respectively. Mean mortality 

rates of worms exposed to lo%, 40%, and 46% 

concentrations of diesel fuel for 48 h were 5%, 11 %, and 

16%, respectively. Many surviving worms exposed to 

kerosene or diesel fuel suffered lost or damaged feeding 

tentacles. No significant mortality was observed among 

worms exposed to furnace fuel (Bunker "C") for 48 h at 

concentrations as high as 40%. Worms exposed to 

furnace fuel did not suffer damaged tentacles. Total 

submersion in these fuel oils for 24 h resulted in 

mortalities of only 8%. 2%, and 4% in kerosene, diesel 

fuel, and furnace fuel, respectively, but tentacles of 

worms in kerosene and diesel fuel were damaged or lost. 

Mulhem (1976) inferred that reef-building tube worms 

are relatively resistant to short-term oil pollution and 

appear capable of surviving typical oil spills; however, 

he acknowledged that worms suffering from lost or 

damaged feeding tentacles may have reduced long-term 

survival. 

Kavanagh (1979) determined that the tolerance of 

reef-building tube worms to cadmium varied 

ontogenetically. The 96 h LCso values for adults and 

larvae were 10.9 pprn and 3.0 ppm, respectively. 

Fertilization success and growth and development of 

larvae were significantly reduced at cadmium 

concentrations exceeding 1.0 ppm. Adult worms 

exposed to cadmium concentrations of 0.5 pprn and 

1.0 pprn for 240 h remained in apparent good health with 

no morphological differences from controls, but adults 

exposed to 2.5 pprn cadmium exhibited tentacle 

deterioration within 48 h and died within 216 h. 

The 48 h LCso of adult reef-building tube worms for 

the reference toxicant dodecyl sodium sulfate is 460 mg/l 

(Mulhern 1976). 

Dredging 

Strong currents and high sediment loadings 

characteristic of coastal inlets provide excellent habitat 

for reef-building tube worms. Accordingly, worm reefs 

grow well on bulkheads, jetties, and along the bottoms of 

channels at some inlets, eventually making them 

navigable only by shallow-draft watercraft (Kirtley 

1968). The narrowed channels also increase the velocity 

of tidal currents, thereby making passage dangerous. 

Dredging of worm reefs required to keep such inlets 

navigable is deleterious to reef-building tube worms and 

associated fauna directly affected by dredging. 

Beach Nourishment 

Burial, siltation, and exposure to sulfides are all factors 

which may be present due to beach nourishment or 

disposal of dredged sediments. Clark (1978) reported 

that total mortality of reef-building tube worms occurred 

at reefs adjacent to beaches nourished by spoil dredged 

from nearby Sebastian Inlet, Florida. The presence of 

large quantities of drifting sand over the reefs suggested 
that mortality was the direct result of overwash with sand 

and consequent smothering of worms. Nelson and Main 

(1985) reported that reef-building tube worms tolerated 

burial by sediments for up to 72 h at 17 to 23 "C in the 

laboratory, but exhibited increased mortality as compared 

with coarser sediments. Further laboratory experiments 

showed tolerance of burial for 72 h at winter 

temperatures (18.5-21.5 "C) but only for 25 h of summer 

temperatures (29.2 'C) (Main and Nelson 1988b). 

Reef-building tube worms showed no negative re.sponse 

to extremely high silt levels (up to 6.0 gfl ) over a 4-day 

period (Main and Nelson 1988b). The tube worms 

showed 50% mortality between 24 and 48 h of exposure 

to high levels of sulfide (about 4.3 mgfl) simultaneously 

with low initial oxygen levels (<0.2 mgfl) (Main and 

Nelson 1988b). 
LITERATURE CITED 

Briggs, J.C. 1974. Marine zoogeography. McGraw-Hill 

Book Co., New York. 

Clark, K.B. 1978. Summary of monitoring of seagrasses 

and sabellariid worm reefs at Sebastian Inlet, Florida. 

11. Post-dredging report. Unpublished report 

submitted to the Sebastian Inlet Commission. 14 pp. 

Curtis, L. 1973. Aspects of the life cycle of Sabellaria 

vulgaris Vemll (Polychaeta: Sabellariidae) in 

Delaware Bay. Doctoral Dissertation. University of 

Delaware. 236 pp. 

DeJorge, F.B., J.A. Petersen, and E.S. Fanta. 1969. 

Biochemical studies on Phragmatopoma lapidosa 

Kinberg 1867 (Sabellariidae, Polychaeta). Comp. 

Biochem. Physiol. 31:493-501. 

Eckelbarger, K.J. 1976. Larval development and 

population aspects of the reef-building polychaete 

Phragmatopoma lapidosa from the east coast of 

Florida. Bull. Mar. Sci. 26: 117-132. 

Eckelbarger, K.J. 1977. Larval develolpment of 

Sabellaria floridensis from Florida and 

Phragmatopoma californica from southern California 

(Polychaeta: Sabellaxiidae), with a key to the 

sabellariid larvae of Florida and a review of 

development in the family. Bull. Mar. Sci. 27:241-255. 

Eckelbarger, K.J. 1979. Ultrastructural evidence for 

both autosynthetic and heterosynthetic yolk formation 

in the oocytes of an annelid (Phragmatopoma 

lapidosa: Polychaeta). Tissue Cell 11 :425-443. 

Eckelbarger, K.J. 1984. Ulmstructure of spermatogenesis in the reef-building polychaete Phragmatopoma 

lapidosa (Sabellariidae) with special reference to 

acrosome morphogenesis. J. Ultrastruct. Res. 

89: 146- 164. 

Eckelbarger, K.J., and F.-S. Chia. 1976. Scanning 

elecmn microscopic observations of the larval 

development of the reef-building polychaete 

Phragmatopoma lapidosa. Can. J. Zool. 

54:2082-2088. 

Emery, K.O. 1963. Organic transportation of marine 

sediments. Pages 776-793 in M.N. Hill, ed. The sea, 

Vol. 3. Interscience Publishers, New York. 

Gilmore, R.G. 1977. Fishes of the Indian River lagoon 

and adjacent waters, Florida. Bull. Fla. State Mus., 

Biol. Sci. 22: 101-147. 

Gilmore, R.G., Jr., C.J. Donohoe, D.W. Cooke, and 

D.J. He~~ema. 1981. Fishes of the Indian River lagoon 

and adjacent waters, Florida. Harbor Branch Found. 

Tech. Rep. 41. 36 pp. 

Gore, R.H., L.E. Scotto, and L.J. Becker. 1978. 

Community composition, stability, and trophic 

partitioning in decapod crustaceans inhabiting some 

subtropical sabellariid worm reefs. Bull. Mar. Sci. 

28:221-248. 

Gram, R. 1968. A Florida Sabellariidae reef and its 

effect on sediment distribution. J. Sediment. Petrol. 

38:863-868. 

Harunan, 0. 1944. Polychaetous annelids from 

California. Part 6: Paraonidae, Magelonidae, 

Longosomidae, Ctenodrilidae, and Sabellariidae. 

Allan Hancock Pacif. Exped. 10:311-389. 

Kavanagh, B.A. 1979. The effects of cadmium on the 

reef-forming sabellariid polychaete Phragmatopoma 

lapidosa. Master's Thesis. Florida Institute of 

Technology, Melbourne. 51 pp. 

Kirtley, D.W. 1966. Intertidal reefs of Sabellariidae 

(Annelids Polychaeta) along the coasts of Florida. 
Master's Thesis. Florida State University, Tallahassee. 

104 PP. 

Kirtley, D.W. 1968. The reef builders. Nat. Hist. (N.Y.) 

77:4045. 

Kirtley, D.W., and W.F. Tamer. 1968. Sabellariid 

worms: builders of a major reef type. J. Sediment. 

Petrol. 38:73-78. 

Main, M.B., and W.G. Nelson. 1988a. Sedmentary 

characteristics of sabellariid worm reefs 

Phragmatopoma lapidosa Kinberg. Estuarine Coastal 

Shelf Sci. 26:105-109. 

Main, M.B., and W.G. Nelson. 1988b. Tolerance of the 

sakllariid polychaete Phragmatopoma lapidosa 

Kinberg to burial, turbidity, and hydrogen sulfide. Mar. 

Environ. Res. 26:39-56. 

Mauro, N.A. 1975. The premetamorphic developmental 

rate of Phragmatopoma lapidosa Kinberg, 1867, 

compared with that in temperate sabellariids 

(Polychaeta: bellariidae). Bull. Mar. Sci. 25387-392. 

Mauro, N.A. 1977. Variations in osmoregulatory capacity in two species of intertidal sabellariids (Annelida: 

Polychaeta) from tropical and mediterranean habitats. 

Comp. Biochem. Physiol. 56A:375-377. 

Mehta, A.J. 1973. Coastal engineering study of sabellariid reefs: report of hydraulic model study to the 

Harbor Branch Foundation Laboratory, Ft. Pierce, 

Florida. Coast. Oceanogr. Eng. Lab., Eng. Indust. 

Exper. Sta., University of Florida, Gainesville. 67 pp. 

Merida, L., and P.E. Penchaszadeh. 1982. Dinarnica 

del poliqueto constructor de arrecifes Phragmatopoma 

lapidosa en fondos blandos de Punta Moron (Golfo 

Triste, Venezuela). Atlantica 579. 

Mulhern, E. 1976. A toxicity study of three refined fuel 

oils and a reference toxicant with respect to the 

sakllariid worm Phragmatopoma lapidosa. Master's 

Thesis. Florida Institute of Technology, Melbourne. 

45 PP. 

Multer, H.G., and J.D. Milliman. 1967. Geologic 

aspects of sabellarian reefs, southeastern Florida. Bull. 

Mar. Sci. 17:257-267. 

Narchi, W. 1973. On the functional morphology of 

Hiatella solida (Hiatellidae: Bivalvia). Mar. Biol. 

(Berl.) 19:332-337. 

Narchi, W. 1974. Functional morphology of Petricola 

(Rupellaria) typica (Bivalvia: Petricolidae). Mar. Biol. 

(Berl.) 27: 123- 129. 

Nelson, W.G., and M.B. Main. 1985. Criteria for beach 

nourishment: biological guidelines for sabellariid 

worm reef. Fla. Sea Grant Coll. Tech. Pap. 33. 34 pp. 

Parker, S.P., ed. 1982. Synopsis and classification of 

living organisms, Vol. 2. McGraw-Hill Book Co., New 

York. 

Pawlik, J.R. 1988. Larval settlement and metamorphosis of sabellariid polychaetes with special reference to 

Phragmatopoma lapidosa, a reef building species, and 

Sabellaria floridensis, a nongregarious species. Bull. 

Mar. Sci. 43:4 1-60. 

Potts, D.L. 1979. Phragmatopoma lapidosa (Polychaeta: 

Sabellariidae) recorded from the western Gulf of 

Mexico. Tex. J. Sci. 31:370. 

Smith, P.R., and F.-S. Chia. 1985. Larval development 

and metamorphosis of Sabellaria cementarium Moore, 

1906 (Polychaeta: Sabellariidae). Can. J. Zool. 

63: 1037-1049. 

van Montfrans, J. 198 1. Decapod crustaceans associated 

with worm rock (Phragmatopoma lapidosa Kinberg) 

in southeastern Florida. Master's Thesis. Florida 

Atlantic University, Boca Raton. 290 pp. 

Wilson, D.P. 197 1. Sabellaria colonies at Duckpool, 

North Cornwall, 1961-1970. J. Mar. Biol. Assoc. U.K. 

5 1 :509-580. 

Wilson, D.P. 1974. Sabellaria colonies at Duckpool, 

North Cornwall, 1971-1972, with a note for May 1973. 

J. Mar. Biol. Assoc. U.K. 54:393-436. 
PAGE I Biological Re~ort 82(11.115)* 1 4. Tnk and SuMitb 

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes 

and Invertebrates (South Florida)--Reef-building Tube Worm 

National Wetlands Research Center U.S. Army Corps of Engineers 

Fish and Wildlife Service Waterways Experiment Station 

U.S. Department of the Interior P.O. Box 63 1 

Washington, DC 20240 Vicksburg, MS 391 80 

5. Report Date 

~~cember 1989 

E 

7. Author(a) 

Alexander V. Zale and Susan G. Menifield 

9. Performing Organization Name and AddrOklahoma Cooperative Fish and Wildlife Research Unit 1 404 Life Sciences West 

Oklahoma State University 

Stillwater, OK 74078 

12. Sponsoring Organization Nsm and Addmaa 

I 

15. Supplsmntay Notes 

a Performing Organization Rept. No. 

10. ProjeMaaWork Unit No. 

11. Contmct(c) or Gmnt(G) NO. 

(C) 

(G) 

13.Type of Report L Period Covered 

*U.S. Army Corps of Engineers Report No. TR EL-82-4. 

16. AtelrM (Limit: ZW Words) 

Species profiles are literature summaries of the taxonomy, morphology, distribution, life history, habitats, and 

environmental requirements of coastal species of fishes and aquatic invertebrates. They are designed to assist 

in environmental impact assessment. The reef-building tube worm is an ecologically and geologically 

significant invertebrate inhabiting the coastal zone of southeastern Florida. The reefs constructed by the 

worms retain beach sediments, protect shorelines from storm damage, and are the basis for an elaborate 

marine community of fishes and invertebrates. The reefs provide substrate, shelter, and food in the relatively 

inhospitable surf zone. Reef-building tube worms require stable settlement substrates within sandy beach 

habitats and intense turbulence to maintain suspension of sand grains and other particles for tube building. 

17. Docurmnt Analyab a. Deuriptors 

Annelida Temperature Competition 

Feeding habits Suspended sediments Salinity 

Growth contaminants Life cycles 

b Id.ntWirdOpen-Ended Term 

Reef-building tube worm Environmental requirements 

Phragmatopoma Iapidosa 

Spawning 

I I I I 

(Sea ANSI-239.18) OF'TIONAL FORM 272 (4-77) 

(Forrnorly NTIS35) 

Doprtrnont of Cornrnorco 

c. COSATl FmMlGroup 

18. Availabllky Staiemnt 

Unlimited distribution 

10.SecurHy Clan (Thu Report) 

Unclassified 

P. Security Clan (Thi8 PW) 

Unclassified 

a. NO. of ~agvi + 12 

22 Price 
As the Nation's principal conservation agency, the Department of 

the Interior has responsibility for most of our nationally owned 

public lands and natural resources. This includes fostering the 

wisest use of our land and water resources, protecting our fish 

and wildlife, preserving the environmental and cultural values of our 

national parks and historical places, and providing for the enjoyment of life through outdoor recreation. 'The Department assesses 

our energy and mineral resources and works to assure that their 

development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation 

communities and for people who live in island territories under U.S. 

administration. 

U.S. DEPARTMENT OF THE INTERIOR 

FlSH AND WILDLIFE SERVICE 

TAKE PRIDE 

in America 

UNITED STATES 

DEPARTMENT OF THE INTERIOR 

FISH AND WILDLIFE SERVICE 

National Wetlands Research Center 

NASA-Slidell Computer Complex 

101 0 Gause Boulevard 

Slidell. LA 70458