A STUDY OF THE GROWTH AND OTHER ASPECTS OF THE BIOLOGY OF THE WEST INDIAN TOPSHELL, CITT ARIUM PICA

(LINNAEUS)
I

HELEN A. RANDALL Institute of Marine Biology, University of Puerto Rico, Mayagiiez, P.R.
ABSTRACT

Various aspects of the biology of Cittariurn pica (Linnaeus, 1758) are discussed, including habitat, feeding habits, growth, movements, reproduction, predators, and commensals. Marking and tagging experiments to determine growth rate are described.
INTRODUCTION

The West Indian topshell, Cittarium pica (Linnaeus, 1758) (=Livona

pica),

sometimes

known

as the magpie

shell, is a well known

littoral

gastropod which attains a size of four inches. It is widely used as food and is probably second in economic importance only to the queen conch, Strombus gigas Linnaeus, among the gastropods of the Caribbean region. The shell is turbiniform and moderately heavy. It is strikingly colored with irregular markings of black and white (Fig. 1). There is considerable variation in color pattern, some individuals being primarily black with few white markings while others are predominantly white. White seems to be the ground color of the young (Fig. 2). The white of the shell is often greenish or yellowish, probably from staining by the blue-green alga Plectonema terebrans Bornet & Flahault (identified by H. J. Humm) which penetrates the upper calcareous layers. There is no periostracum, and in larger individuals the first few whorls are usually eroded. The older parts of many shells are overgrown with algae, particularly the calcareous red, Goniolithon brgesenii Foslie. The interior of the aperature is pearly and the round chitinous operculum, which fits snugly, is translucent olive (drying to brown) with the spirals showing clearly. The shell makes an attractive curio either when polished to the nacreous layer or with its outer black and white coloration intact. It is used by adults of the land hermit crab Coenobita clypeatus (Herbst) more than any other West Indian shell. Topshells inhabited by this crab were found near the 1277-foot summit of Bordeaux Mountain, the highest point on St. John, more than one mile from the sea. In contrast to the related genus Trochus of the Indo-Pacific, almost
IContribution No. 558 from The Marine Laboratory, Institute of Marine Science, University of Miami. Contribution from the Institute of Marine Biology, University of Puerto Rico. The research reported herein was supported in part by grant G5941 from the National Science Foundation.

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H. Randall: Biology of Cittarium

425

nothing is known of the natural history of the monotypic Cittarium. Aware of this and appreciative of its high gastronomic quality, a study of some aspects of its biology was undertaken as part of a nearly three-year marine biological survey of St. John, Virgin Islands, which commenced in late 1958. This survey was begun while the author was a member of the Institute of Marine Science of the University of Miami and completed while associated with the Institute of Marine Biology of the University of Puerto Rico. Support of the research in the Virgin Islands was provided by Federal Aid in Fish Restoration (Dingell-Johnson Project F-2-R of the Virgin Islands), the National Science Foundation (G-5941) and the National Park Service. The assistance of Herman E. Kumpf, James R. Chess, and John E. Randall in field work is gratefully acknowledged. Thanks are also due Luis R. Almodovar and Harold J. Humm for determinations of the algae from Cittarium stomachs and to Peter W. Glynn and Germaine L. Warmke for reviewing the manuscript.
CLASSIFICATION AND DISTRIBUTION

The West Indian topshell was described as Turbo pica by Linnaeus in 1758. Gray (1842) in the 44th edition of his "Synopses of the contents of the British Museum" listed the generic name Livona with a brief description that was not diagnostic and which mentioned no species. In his 1843 edition of the "Synopses" Gray lists one species, aurea, under the genus Livona, describing it only as "peculiar for the light golden colour of the pearl." In November 1847 he regarded pica as the type of the genus. Cittarium Philippi (February, 1847) had already appeared with pica the sole species (literature reviewed by R. Tucker Abbott, by request); therefore Livona would seem to be a synonym of Cittarium, as indicated by Iredale (1913) who wrote, "A name that seems to need rejection is

FIGURE

1. Two views of an adult Cittarium pica from St. John, Virgin Islands.

Length of shell 97 mm. Personal collection.

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Livona. In 1840 it is a nomen nudum; in 1842 it is indeterminable; in 1843 it is associated with a shell which is certainly not the one selected as the type in 1847." Rehder (1962), however, prefers to retain ivona. Clench & Abbott (1943) selected Linnaeus's reference to Gaultieri (1742: p1. 68, fig. B) as the type figure. They regarded the locality given by Linnaeus, the Sea of Sardinia, as erroneous and designated Martinique as the type locality (after Denys de Montfort, 1810). Clench & Abbott have listed numerous Caribbean localities for the species, and Warmke & Abbott (1961: map 7) have indicated a distribution throughout the West Indies and the Central and South American coast from upper Yucatan to Trinidad. Clench & Abbott have discussed the occurrence of C. pica as a fossil in Florida and Bermuda (an attempt to reintroduce it into Bermuda was not successful) and expressed their belief that the species died out from these two areas in comparatively recent times.
HABITAT

Morris (1947) stated that Cittarium pica in the West Indies lives 0[1 weedy bottoms in moderately shallow water. If one interprets a weedy bottom as seagrass or algae growing on a substratum of sand or other sediment, then his concept of the habitat is erroneous. Cittarium pica lives on rocky shores, and primarily those subjected to at least some wave action. It occurs from slightly above the water level to several feet below, but is usually within two feet of the surface. In general, the smaller shells are found higher in the intertidal zone. Lewis (1960) cited the young as occurring in the "pink zone" above mean low water at Barbados. Cittarium does not occur in brackish areas, such as mangrove sloughs, but whether this is related primarily to the difference in salinity, more turbid water or lack of wave action is not known.

FIGURE

2. Juvenile Cittarium pica from Puerto Rico. Length of shell 5 mm. 1MB 3179.

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H. Randall: Biology of Cittarium

427

Cittarium pica is one of the more abundant of mollusks in the West Indies, and probably the most common large gastropod of the exposed rocky littoral region. As pointed out by Clench & Abbott (1943), however, the larger topshells may be difficult to find in the vicinity of populous areas because of their heavy use as food. The principal area of the present study was Europa Bay on the southern shore of St. John (Randall, 1962: figs. 1 and 2). This is the westernmost of the three smaller embayments of Lameshur Bay. Unlike the other two, which have sandy beaches and little or no surf, the beach of Europa Bay is entirely rocky (Fig. 3). Because it faces southeast it receives almost continuous small surf, usually less than 18 inches high. The bay supports a large population of Cittarium pica, not only because of the suitable habitat but also by being distant from the centers of population on the island. Occasional fishermen collect "whelks" (as topshells are called in the Virgin Islands) at even the remote sectOrs of St. John, therefore a large sign was posted indicating that a study of whelks was in progress and requesting that none be removed from the bay.
FOOD HABITS

The stomach contents of 40 specimens of Cittarium pica from 25 to 77 mm in length (measured from tip of spire to most distant edge of lip)

FIGURE

3. Europa Bay, St. John, Virgin Islands, the site of the growth study of Cittarium pica. View toward the southwest.

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TABLE 1
STOMACHCONTENTS OF Cittarium pica, BY VOLUME, FROM THE VIRGIN ISLANDSANDPUERTO RICO Lameshur Bay, St. John, V.1. 10 25-40 10% Europa Bay, St. John, V.I. 5 34-44 17% Point Higuero & Ramey Beach, P.R. 10 37-47 27% Ram Head Bay near Ram Head, St. John, V.1. 5
-----

LOCALITY

Europa Bay, St. John, V.1. 10 60-77 12% x

NUMBER OF SHELLS LENGTH OF SHELL (mm) Blue-green algae Agmenellum quadruplicatum A nacystis marina Calothrix Dichothrix Lyngbya majuscula Microcoleus Oscil/atoria Phormidium Plectonema nostocorum Green algae Cladophoropsis membranacea Ulvella lens (and other coccoid greens) Red algae Amphiroa fragilissima Asterocystis ramOsa Ceramium cruciatum Coelothrix irregularis Corallina cubensis Gelidium corneum Laurencia obtusa Polysiphonia Brown algae Dictyota divaricata Padina gymnospora Pocokiella variegata Ralfsia expansa Sphacelaria furcigera Diatoms Organic detritus

40-57 50%

x
x

x x x
x

x
x

x
x
x

x
x
X

x x
13% 14%
X

5%

9%

x x
12%

x
19% 1%

x
1%

x 12%

x
x x

x x x x
10%

x x x x x
12%

x x

x x x x
12%

9%

x x
8% 10% 37%

x x
10% 15% 13% 3% 20% 49% 15% 20%

x x x x x
12% 18% 25%

Sand, spicules and calcareous debris (pieces of shell and fragments of calcareous alga such as Halimeda)

19641

H. Randaii: BlOlogy of Cittarium

429

were exammeo l Table 1). The data strongly suggest that this gastropod is herbivorous, although it does ingest some detrital material. Some calcareous fragments in topshells less than 40 mm in length have been as large as 6 mm. Cittarium pica does not seem to be particularly discriminating in its choice of algal food. It feeds mainly on a variety of filamentous algae, often blue-greens (Cyanophyta), and algae of relatively soft thalli. Coarse types of algae such as Turbinaria and Sargassum, which are commonly founa in its environment, were not detected in the stomachs. The physical limitations imposed by the size of the snail and its radular structure (radular teeth figured by Troschel, 1879: pI. 21, fig. 10) probably preclude its ingestion of algae which are both large and firm in texture. The abundance of the West Indian topshell may be related in part to having few competitors of large size for the algal food along the shore. The flora of the intertidal region on exposed rocky coasts is usually more

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TIME (1959-19601

FIGURE 4. Growth of Cittarium pica in Europa Bay, St. John. First vertical line represents the range (3.0 to 9.9 mm) and the dot in the line the mean (5.8 mm) of 223 topshells which were notched, painted red, and released on March 10, 1959. Additional'lines represent collections of the marked shells to July 24, 1960.

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luxuriant than that found on hard substratum in deeper zones. Randall (1961) has explained the paucity of algae on available surfaces in the deeper zones by the presence of a variety of large herbivorous animals, particularly fishes, which do not venture into the intertidal region or do so infrequently (the tidal fluctuation in the West Indies is not great). Observations at night revealed Cittarium pica more active than by day, and more feeding seems to take place at night and early morning hours. More food material was found in the stomachs of specimens collected early in the morning than later in the day.
GROWTH

On March 10, 1959, a collection of 223 juveniles from 3.0 to 9.9 mOl in length (mean 5.8 mOl) was made in the middle of Europa Bay along the rocks shown in Figure 3. The little topshells were painted with bright red acetone-base paint, the lip of each notched with a file, and all released at the series of three large rocks in left center of the foreground of the illustration. Fourteen attempts were made to recover marked topshells at intervals of one month or more, the last on July 24, 1960, when 12 shells

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TIME (1960)
5. Growth of a group of small individuals of Cittarium pica. The first vertical line represents 200 topshells ranging from 0.5 to 2.5 mm in length (mean 1.5 mm) which were painted red and released in Europa Bay, 51. John on January 15, 1960.
FIGURE

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H. Randall: Biology of Cittarium

431

averaging 35.7 mm were found (Fig. 4). Each time the topshells were recovered, they were measured, repainted (usually more than half of the red paint was missing after a period of one month), notched again, and re-released. The average growth rate of this group during the 161h month period is 1.81 mm per month. Growth studies were initiated for three other groups in the same area so that data could be obtained for a broader range of size. On January 15, 1960, 200 small juveniles ranging from 0.5 to 2.5 mm in length and averaging 1.5 mm were marked with red paint. Only 14 were found during the first attempt at recovery on February 17, and mere traces of the red paint remained on these. Probably many marked shells were overlooked because the paint had disappeared entirely. One hundred eight-six more of the same size range and average were added to the 14 recovered ones, and all were released together. Thereafter, at two week intervals, the little topshells were collected and painted, but they were measured only at approximately monthly intervals up to the last collection on July 25 when 79 were found (Fig. 5). At this time they had attained an average length of 8.2 mm. Their average growth rate during the 6 1/3 month period was 1.06 mm per month. The growth of topshells of the same size group from progression of modes (see Fig. 1'0) approximates that of the marked juveniles. The lower growth rate of the small topshells may be related to their feeding less frequently than larger individuals. The author noted that the little shells were usually clustered in groups in narrow cracks high in the intertidal zone and did not appear to be feeding during the day. Glynn

FIGURE 6. Marked topshells from Europa Bay, St. John. Released on August 4, 1959, and recovered on January 17, 1960. Arrows show location of notches. A. Shell marked with a single sharp notch (see Fig. 7). Size at recovery 39.1 mm. B. Shell marked with two obtuse notches (see Fig. 8). Size at recovery 43.7 mm.

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Bulletin of Marine SciEnce of the Gulf and Caribbean

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(MS) has reported a similar slow initial growth for the intertidal gastropods scutula and Tegula funebralis in Monterey Bay, California. Two groups of larger topshells were collected on the southwest side of Europa Bay where the rocks are larger and the habitat therefore more suitable for the larger sheIls. They were marked and released on August 4, 1959. The first group of 234 individuals, ranging from 30.0 to 39.6 mm (mean 34.3 mm) were painted red, and a sharp V-shaped notch was filed in the lip of each (the notch is visible in A of Fig. 6, a marked specimen which was collected 51/2 months after release). One hundred forty-four topshells from 40.0 to 49.8 mm in length (mean 45.1 mm) constituted the second group. These were released in the same area after painting them blue and grinding two broad notches with a grinding wheel in the lip of each (B of Fig. 6). Subsequent attempts to recover the marked shells of these two groups were made at about six-month intervals. Additional topshells were painted, filed, and released with the recovered marked individuals at each date of collection. These were carefully selected to fit
within the range of those recovered and not alter the mean length. The

last collection of the 30-40 mm group was made on January 19, 1961,

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TIME

(1959-1961)

FIGURE7. Growth of medium-sized Cittarium pica. The first vertical line represents 234 topshells ranging from 30.0 to 39.6 mm in length (mean 34.3 mm) which were notched (Fig. 6 A), painted red, and released in Europa Bay, 81. John on August 4, 1959.

1964]

H. Randall: Biology of Cittarium

LENGTH OF SHELL Cmml

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FIGURE8. Growth of Cittarium pica from one-half full size to more than threefourths full size. The first vertical line represents 144 topshells ranging from 40.0 to 49.8 mm in length (mean 45.1 mm) which were double-notched (Fig. 6 B), painted blue, and released in Europa Bay, 81. John on August 4, 1959.

434

Bulletin of Marine Science of the Gulf and Caribbean

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when the topshells averaged 60.1 mm; the last recoveries of the 40-50 mm group were made on August 17, 1961, when the shells averaged 8'0 mm. The growth rates of both groups over the 11/2 and 2 year periods were nearly the same, 1.47 mm per month for the smaller group (Fig. 7) and 1.43 mm per month for the larger (Fig. 8). In order that growth of individual topshells might be determined, experimental tagging of two groups of shells was carried out on January 18, 1960. The shells of each group were chosen to fall within the range of the two series of larger-sized marked topshells discussed above. Each shell was tagged by placing a loop of vinyl plastic tubing (reinforced with 60-pound test nylon in the center) through two holes drilled near the edge of the lip. The free ends of the tubing were clamped with a monel band bearing a stamped number. Only one dead shell of the smaller group was seen again. Three were recovered from the larger group, two after six months and one after 19 months. The first two grew at a rate only slightly more than half that of the untagged but marked shells. The larger tagged one grew 1.25 mm per month, still noticeably less than that of the size group to which it belonged. The tags seem to have had a deterring effect on growth. It was noted shortly after the topshells were released that the operculum tended to catch on the section of plastic tubing that passed into the aperture. No further tagging was attempted. If the growth rate remained constant at about 1.45 mm per month to the maximum length of about 100 mm (as it very nearly did from a length of 34 to 80 mm), then it may be estimated that about 51/2 years would be required for a topshell to grow from a juvenile of about 1 mm to the large adult size. On April 4, 1964, Europa Bay, St. John was briefly revisited, and ten large topshells were collected from the area where marked shells had been released. Upon cleaning these shells, a 93 mm one showed clearly two broad notches, thus demonstrating it to be one of the 45.1 mm group released on August 4, 1959. This shell (IMB 3384) is heavily eroded. The black and white pattern is nearly obliterated over the entire shell which is indicative of no recent growth. Assuming two years were required for it to attain the 45 mm size at marking, this topshell is slightly more than 61/2 years old. The length-weight relationship of Cittarium pica is given in Figure SI, thus making possible a conversion of length data to approximate weight. In some growth studies of gastropods, the lip growth is determined instead of increase in shell length. In order to relate growth of the present study to lip growth, 86 topshells in 13 size groups from 4 to 66 mm were marked by filing one to three notches in the lip and released. After four months 34 were recovered, and two measurements were taken of each, the growth of the lip along the inner edge of the whorl and the length of the

1964]
N

H. Randall: Biology of Cittarium

WEIGHT (GRAMS) o

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9. Length-weight relationship of Cittarium pica. All specimens from Europa Bay, St. John, Virgin Islands.

436

Bulletin of Marine Science of the Gulf and Caribbean

[14(3)

shell. The measurements were averaged for the different groups. Lip growth was found to be 3.4 to 4 times greater than the overall growth of the length of the shell. No obvious increase or decrease of this ratio was evident from the smaller sizes to the larger. The average monthly sea temperatures in Lameshur Bay, St. John, during the period of the growth study ranged from 26.7 to 29.9C (Randall, 1961; Table 2).
MOVEMENTS

The marked individuals of Cittarium pica tended to remain in the area of release. Had this not been the case, many more individuals would have to be tagged to obtain so many recoveries. As would be expected, the larger individuals made the more extensive movements. The 78 marked topshells which averaged 5.8 mm when released on March 10, 1959, and which had an average of 9.5 mm when recovered on May 31 (Fig. 4) were nearly all found in the immediate vicinity of the three rocks of Europa Bay where originally released. Three, however, were from 15 to 20 feet away. On October 20 one of this group was found 30 feet from the point of release. On April 13, 1960, half of the 30 which were recovered were found on or beneath the three rocks; all of the rest were within 15 feet of the rocks. On May 18 one was found 20 feet to one side of the rocks and another 45 feet to the other side. On June 12, all that were recovered were within 20 feet of the point of release. In anticipation of the greater movements of the larger marked shells, the shore area was searched for several hundred feet to either side of the point of release; however the longest movement was that of one doublenotched, blue-painted individual which was found 150 feet in the westerly direction between August 2, 1960, when it was released and its recovery (at a length of 60 mm) on January 18, 1960. Two others of this group had moved 80 feet from the area of release.
REPRODUCTION

The sexes of mature individuals of Cittarium pica are readily distinguished by the color of the gonads. The ovary is unmistakably green and the testis creamy white. The smallest mature female detected among the many samples examined measured 33.7 mm in length and the smallest male 32.4 mm. Some insight into the spawning activity of Cittarium pica was obtained by sampling the young throughout the year (Fig. 10), beginning August 8, 1959. For the period of one hour the author collected as many small topshells under 10 mm in length as possible in Europa Bay. Because conditions of surf and other possible factors could not be kept constant, these small samples are not intended to be an accurate measure of relative

1964]

H. Randall: Biology of Cittarium

437

abundance from month to month, but they do represent approximations. In addition to giving the sample size for each collection, the length of thc smallest topshell is listed. There was a large influx of young in January, indicated by the January 16 sample of Figure 10. This pronounced mode may be followed from month to month to the June sample. It suggests a brief and intensive spawning period for adults some weeks earlier; however, with the exception of November (July, a month with no sample), topshells smaller than 4 mm were found in all months of the year. If we assume a growth of only 1 mm per month, the 3.2 mm one collected on October 20 must have come to the intertidal. as a new arrival from the plankton at a length of about 1 mm only two to three months earlier. Perhaps there is low-level spawning over a large part of the year with a peak to account for the big January influx of young; or larval stages may have drifted in from other areas where spawning is out of phase with that of St. John (although it is difficult to designate such an area in view of the normal westerly set of the current). Although no further samples were taken beyond June 13, 1960, an abundance of small juveniles was again noticed the following January, thus suggesting that the occurrence of a large number of young in January 1960 was probably not unusual.
PREDATORS

The following few observations on the predators of Cittarium pica were made. Much remains to be learned about these and other enemies of the topshell. On August 2, 1960, a 35-mm Purpura patula was found eating a 37-mm topshell in Europa Bay, St. John. The Purpura was feeding on fresh tissue between the operculum of the topshell and the rest of the foot, and the operculum was nearly detached. The topshell was not drilled. A number of shells of Cittarium pica were found, which had been
drilled, either empty or inhabited by hermit crabs, but no observations

were made in the field of the gastropod responsible for the drilling. Possibly it is a species of Thais. While searching for the large-sized marked topshells, one was found in the arms of a small octopus whose body was inside its hole in the rocks. The diameter of the topshell was so great that the octopus was unable to bring it into its hole (which it seemed to be trying to do). The topshell was ultimately removed from the grasp of the octopus; it was still alive. The octopus was not captured. Gregory Bateson of the Communication Research Institute at St. Thomas, Virgin Islands, reports (personal communication) that he uses live topshells to feed octopuses (Octopus vulgaris and O. filosus) in aquaria. The method by which these octopuses

438
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Bulletin of Marine Science of the Gulf and Caribbean

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1964]

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kill Cittarium is at present not known. It is clear that they do not drill through the operculum as O. bimaculatus does in killing Tegula spp. on the Pacific coast. George Hunt, who collects octopuses for the Communication Research Institute, reports that empty shells of Cittarium pica are most frequently encountered in front of the dens of Octopus vulgaris in shallow water. In deeper water, bivalves predominate. On several occasions octopuses have been found carrying live topshells or holding live snails in their dens. Three fishes were found in St. John with the remains of Cittarium pica in their stomachs. One, a porcupinefish (Diodon hystrix) 394 mm in standard length, contained two topshells (the opercula measured 17 and 18 mm in greatest diameter) among the crushed gastropods, crabs, and echinoids in its stomach. A 285-mm puddingwife (Halichoeres radiata) had eaten three Cittarium; the opercula ranged from 2.3 to 3.3 mm in diameter. This wrasse probably crushed the small topshells with its pharyngeal teeth. A rock hind (Epinephelus adscensionis), 375 mm in standard length, contained the opercula of two topshells in its stomach. The opercula measured 19 and 25 mm in diameter, and there were no pieces of shell or soft tissue with them. This grouper has no known mechanism to crush a large Cittarium. Possibly it stole a meal from some other fish, such as Diodon. Warmke and Erdman (1963) have reported one shell of Cittarium pica among the stomach contents of 56 bonefish (Albula vulpes) from Puerto Rico. Since the bone fish does not normally enter the topshell's habitat, the one fish may have ingested either an empty shell or a shell occupied by a hermit crab. Intact shells of Cittarium which still contained soft tissue, were occasionally found at the shore, often well above the high tide line. Some animal such as a crab or a bird with a pointed feeding structure was presumed to be responsible for these partially eaten topshells. After discussion with local people on St. John, the oystercatcher (Haemotopus ostralegus) (nomenclature after Bond, 1956) became the chief suspect. Permission was obtained from the National Park Service to conect this bird. Two were shot from the intertidal area by Robert E. Schroeder. One weighing 1V:zpounds contained the remains of seven Cittarium pica which, judging from the diameter of the opercula, had been about 35 to 45 mm in shell length. Only one small topshell fragment was present. The bird's stomach also contained two small clams and one A straea americana imbricata. The second bird, which weighed 1 pound 7 ounces, had eaten 11 Cittarium pica (opercula from 5.5 to 11 mm in diameter), one 8-mm Nodilittorina tuberculata, four 7- to 8-mm individuals of Littorina ziczac. and one A straea americana with a 6-mm operculum. The shells of the Nodilittorina and Littorina were entire.

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Several of the natives of St. John stated that they had witnessed the oystercatcher flying well above the rocks with Cittarium, dropping them so they would break, and then flying down to eat the exposed soft parts. This behavior was not observed by the author. It is well known for gulls. For example, the herring and ring-billed gulls were reported by Magalhaes (1948) feeding upon Busycon, Area and Noetia in this manner in North Carolina. This method is not mentioned by Dewar (1908, 1913) in his articles on the mode of feeding of the oystercatcher. He described more direct methods whereby the bill was variously employed in thrusting, hammering, twisting and exerting leverage to capture and open the shells of various pelecypods and gastropods.
COMMENSALS

Two organisms were found living in close aSSOCiatlonwith Cittarium pica. The relationship for both appears to be one of commensalism. The
small limpet Acmaea leucopleura is often found on the underside of the

shell, and the crab Pinnotheres barbatus lives in the mantle cavity. Eighteen of the crabs were found in a total of 203 adult topshells collected from southern St. John from January to August, 1960. One of the shells contained two of the crabs; the other crabs were found singly in the shells.
SUMMARY

The West Indian topshell, Cittarium pica (Livona pica of many authors), is widely used as food in the Caribbean region. The preferred habitat is a rocky shore with some surf action. The juveniles live high in the intertidal zone and the large adults (up to 100 mm) may be found from a few inches to two to three feet or more below low water. The stomach contents of 40 topshells measuring 25 to 77 mm, from the Virgin Islands and Puerto Rico, revealed the species to be herbivorous. Filamentous algae is the principal food. Twenty-four genera of algae (discounting diatoms), were identified from the stomachs of which nine are blue-greens. Large amounts of sand, calcareous debris and other detritus are usually present. The growth of Cittarium was determined by marking shells in four different size groups at St. John, Virgin Islands. The smallest (an average of 1.5 mm when marked) grew at a rate of slightly more than 1 mm per month for a period of six months. A group which averaged 5.8 mm when marked grew an average of 1.81 mm per month over a period of 161/2 months. The third group (initial mean length 34.3 mm) increased the shell length at a rate of 1.47 mm per month for P/2 years, and the fourth (mean length 45.1 mm) grew 1.43 mm per month over a two-year period. One of the latter was recovered 4 years and 7 months after release. It

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measured 93 mm and was heavily eroded to the lip, thus indicating no recent growth. The marked topshells showed little tendency to make extensive movements. The greatest movement recorded for any shell less than 35 mm in length was 45 feet. The longest migration of larger shells was that of a 60-mm individual which moved 150 feet from the point of release. A large influx of juvenile topshells of about 0.5 to 1 mm was observed in St. John in January, 1960. No other obvious influx of young appeared during the year, although topshells smaller than 4 mm were found in nearly all months. Purpura patula was found feeding upon a freshly dead Cittarium. An octopus was observed trying to pull a moderately large topshell into its hole. The remains of Cittarium were found in the stomachs of three fishes and the oystercatcher (Haemotopus ostralegus). The limpet Acmaea leucopleura is often seen adhering to the underside of the shell of Cittarium, and the crab Pinnotheres barbatus is occasionally found in the mantle cavity.
SUMARIO ESTUDIO DEL CRECTMIENTO DEL BURGAO, Y OTROS ASPECTOS DE LA

Cittarium pica (LINNAEUS) El burgao (Cittarium pica-muchos autores aun usan Livona pica) se utiliza como alimento en muchos lugares de la regi6n del Caribe. EI habitat preferido par este molusco es el de ormas rocosas donde baten las olas. Los ejemplares juveniles viven alto en la zona cotidal mientras que los adultos (hasta de 100 mm) se encuentran desde algunas pulgadas hasta dos, tres 0 mas pies bajo el nivel de bajamar. EI contenido estomacal de 40 burgaos obtenidos en las Islas Vfrgenes y Puerto Rico y cuyos tamanos oscilaban entre los 25 y 77 mm revela que la especie es herbfvora, y que el alimento principal consiste de algas
BIOLOGIA

fiJamentosas. Sin contar las diatomeas,

se identificaron 24 generos de

algas de las cuales nueve son azules. Generalmente se encuentran presentes grandes cantidades de partfculas ca1careas y detritos. EI crecimiento de Cittarium se determin6 marcando las conchas de cuatro grupos distintos seleccionados en cuanto a tamano y obtenidos en St. John, Islas Vfrgenes. El grupo de los mas pequenos (con un promedio de 1.5 mm al marcarse) creci6 a raz6n de poco mas de 1 mm por mes durante un tiempo de seis meses. Otro grupo que inc1ufa ejemplares de un tamano medio de 5.8 mm al marcarse creci6 un promedio de 1.81 mm por mes por un tiempo de 16th meses. El tercer grupo (largo inicial medio de 34.3 mm) aument6 el tamano de la concha a raz6n de 1.47 mm por mes por un pedodo de Ith afios y el cuarto grupo (largo inicial medio de 45.1 mm) crecio 1.43 mm por mes por un perfodo de

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mas de dos aiios. Uno de los ultimos fue recuperado 4 aiios y 7 meses despues de su desprendimiento. Media 93 mm y estaba grandemente carcomido hasta el labio, indicando asi que no habia crecimiento reciente. Los burgaos que fueron marcados demostraron poca tendcncia a moverse de un sitio para otro. La mayor distancia recorrida que se pudo observar para una concha menor de 35 mm de tamaiio fue 45 pies. El recorrido mas largo efectuado por animales de conchas de mayor tamaiio fue el de uno de 60 mm de largo de concha que se encontr6 a 150 pies del punta don de se solt6. En enero de 1960 se observ6 en S1. John una gran afluencia de burgaos j6venes de tamaiios entre 0.5 mm y 1 mm. No se observ6 igual abundancia de estados juveniles durante el aiio aunque siempre se conseguian ejemplares mas pequeiios de 4 mm. EI gaster6podo Purpura patula fue encontrado alimentandosc de un Cittarium recientemente muerto. Tambien se pudo observar a un pulpo tratando de halar a un burgao de tamaiio moderado hasta su escondite. Los restos de Cittarium fueron encontrados en el contenido estomacal de tres pescados y del caracolero (Haemotopus ostralegus). EI gaster6podo ("limpet") Acmaea leucopleura se observa a menudo pegado a la parte de abajo de la concha de Cittarium; el cangrejo Pin notheres barbatus se encuentra ocasionalmente en la cavidad palia], LITERA TURE CITED J. 1956. Check-list of birds of the West Indies. Fourth Edition. The Academy of Natural Sciences, Philadelphia, Pennsylvania, ix + 214 pp., with supplements. W. J. AND R. T. ABBOTT 1943. The genera Gaza and Livona In the western Atlantic. no. 12: 1-12,4 pIs. Johnsonia.

BOND,

CLENCH,

DEWAR,

J. M. 1908. Notes on the oystercatcher (Haemotopus ostralegus). with reference to its habit of feeding upon the mussel (Mytilus edulis). Zoologist. (4) 12: 201-212. 1913. Further observations on the feeding habits of the oystercatcher (Haemotopus ostralegus). Zoologist, (4) 17: 41-56.
N.

GAUL TIERI,

1742. Index testarum conchyliorum quae in ejusdem museO adservantur et method ice distributa exhibentur. Caientani Albizzini, Florence. xxiv + 110 pIs.
GLYNN,

P. W. MS. Community composItIon, structure, and interrelationships in the marine intertidal Endoc/adia muricata-Balanus glandula association in Monterey Bay, California. (Submitted for publication to Beaufortia. )

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GRAY,

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1. E. 1842. Synopses of the contents of the British Museum. 44th Edition. G. Woodfall and Son, London, iv + 308 pp. 1847. A list of the genera of recent MoIlusca, their synonym a and types. Proc. zool. Soc. London, Pt. 15: 129-219. IREDALE, T. 1913. The collation of the moIluscan parts of the synopses of the contents of the British Museum, 1838-1845. Proc. malac. Soc. London, 10 (4): 294-309. LEWIS, J. B. 1960. The fauna of rocky shores of Barbados, West Indies. Canad. J. Zool., 38: 391-435, 20 figs.
MAGALHAES,

H.

1948. An ecological study of snails of the genus Busycon at Beaufort, North Carolina. Eco!' Monogr., 18 (3): 379-409, 61 figs. MONTFORT, D. DE 1810. Conchyliologie systematique, et classification mHhodique des CoquiIles. Vol. 2. Coquilles univalves, non-c1oisonnees, ou uniloculaires. F. SchoeIl, Paris, 676 pp., 161 pis. MORRIS, P. A. 1947. A field guide to the sheIls. Houghton Mifflin Company, Boston, xvii + 190 pp., 40 pis. PHILIPPI, R. A. 1847. Versuch einer systematischen Eintheilung des Gesehlechtes Trochus. Zeitschr. fur Malakozoologie, 4 Jahrg.: 17-24. RANDALL, J. E. 1961. Overgrazing of algae by herbivorous marine fishes. Ecology, 42 (4): 812. 1962. Tagging reef fishes in the Virgin Islands. Proc. Gulf and Carib. Fish. Inst. (14th Annual Session): 201-241, 8 figs. REHDER, H. A. 1962. Contribuci6n al conocimiento de los moluscos marinos del Archipielago de los Roques y la Orchila. Mem. Soc. Cienc. nat. La Salle, 22 (62): 116-138, 6 figs.
TROSCHEL,

F. H.

1879. Das Gebiss der Schneken. Vol. 2. Nicolaische Verlags-Buchhandlung. Berlin, vii + 409 pp., 32 pIs. WARMKE, G. L. AND R. T. ABBOTT 1961. Caribbean seashells. Livingston Publishing Company, Narberth, Penna., x + 346 pp., 34 text-figs, 44 pis., 19 maps. WARMKE, G. L. AND D. S. ERDMAN 1963. Records of marine rnoIlusks eaten by bonefish in Puerto Rican waters. The Nautilus, 76 (4): 115-120, 2 figs.