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<I>Tripneustes Ventricosus</I>

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<I>Tripneustes Ventricosus</I> CONTRIBUTIONS TO THE BIOLOGY OF THE SEA URCHIN TRIPNEUSTES VENTRICOSUS' B. F. McPHERSON lnstitute of Marine Science, University of Miami ABSTRACT The growth of Tripneustes ventricosus (Lamarck) was studied using three methods; tagging, penning, and size-frequency analysis. Young urchins (smaller than 30 mm) were found mainly during summer. Growth was rapid during the first year, with the urchins reaching a mean size of about 75 mm by the following summer. There was no indication of growth in the adult population during the summer. Gonads developed during the fall at a test diameter of about 35 to 45 mm. There was a significant departure from the 1: 1 sex ratio in urchins larger than 80 mm. Gonad volume of the population indicated that there were two periods of gonad development during the year, one in winter and one in summer. INTRODUCTION The white-spined sea urchin, Tripneustes venticosus (Lamarck, 1816), is one of the largest regular echinoids of the shallow, tropical Atlantic. It occurs at Bermuda and from Florida throughout the West Indian region to Brazil. It is also found at Ascension Island and along the west coast of Africa from the Gulf of Guinea to Swakopmund (Mortensen, 1943: 496). In Florida, it has been reported as far north as Riviera Beach (Robert Work, personal communication). In Florida, T. ventricosus was found in areas of "sea grass" (Thalassia testudinum), rocks, rubble, and coral reef. Sometimes it was found living on the open surface of rocks or rubble, and sometimes it was found under rock ledges and beneath hollow coral boulders. Moore et al. (] 963b) re- ported that during the winter it was frequently found in open patches among grass and that during the spring and summer, as the light became stronger, it tended to move from the open areas into cover of dense grass. In several recent papers on the ecology of Tripneustes ventricosus, authors have used the name Tripneustes esculentus (Leske) (Lewis, 1958; Moore et al., 1963b). Mortensen (1943) showed that A. Agassiz, and later Clark (1933), committed an error when they considered the West Indian Tripneustes to be identical with Leske's Cidaris esculenta. He point- ed out that the correct name for this West Indian urchin is Tripneustes ventricosus. Lewis (1958), working in Barbados, B.W.!., has given the most com- plete description of the life history of Tripneustes ventricosus published to date. He studied growth by caging urchins and by using size-frequency 'Contribution No. 590 from The Marine Laboratory, Institute of Marine Science, Univer- sity of Miami. This report constituted in part a thesis submitted in partial fulfillment of requirements for the degree of Master of Science at the University of Miami. 1965] McPherson: Biology of Tripneustes 229 analysis of the population. The spawning season was estimated from changes in the gonad volume and the gonad condition and from quantita- tive plankton tows. Moore et al, (1963b), working at Miami, Florida, compared the gonad growth and spawning of Tripneustes over a period of years. Test growth in relation to temperature and size was described. Some preliminary feed- ing experiments were made, and food intake was estimated tentatively. MATERIALS AND METHODS The measurement used in this study was maximum diameter of the test. It was made on a sliding block, calibrated to the nearest millimeter. The error in the method was checked by measuring 20 urchins to the nearest millimeter, three times each on the same line through the test. A mean for the three measurements was calculated. Analysis of variance showed no significant difference at the 5 per cent level in the three means. The circu- larity of urchins was checked by measuring 20 individuals, at three posi- tions, at 60° intervals around their greatest circumference. Analysis of vari- ance showed no significant difference at the 5 per cent level in the three means. Most urchins in this study were taken by diving in water less than two meters deep off Virginia Key, Florida. Some urchins were taken by diving at Boca Raton, Florida, and some small individuals were taken by dredg- ing in depths of three or four meters off Key Biscayne, Florida. The sea temperature data used in this study were taken from the daily readings of the Miami Marine Research and Test Station, Inc., on Miami Beach, Florida. These temperature data can be used only as an approxi- mation of the temperatures where the urchins were studied. Three methods were used to estimate the growth of the test: (1) hold- ing animals in pens and cages in the sea; (2) tagging animals; (3) using size-frequency analysis on the population. Ten pens were constructed below extreme low water on the beach of the Marine Laboratory of the University of Miami. Each pen covered ap- proximately 1.5 square meters of sea bottom. Rocks were put in each pen to simulate a natural environment of Tripneustes. Algae growing on the rocks and grass that washed into the pens provided food for the urchins. Individual wire cages were constructed for small urchins. These cages were cylindrical in shape, about 25 em high and 15 em in diameter. They were anchored to the bottom adjacent to the pens. A bottomless wire cage, approximately a meter in diameter and half a meter high, was also constructed for holding small urchins. This cage was anchored to the bottom of the grass flats off Virginia Key. Growth was determined from tagged individuals living in their natural environment. The tag consisted of a plastic disk fastened by a stainless steel wire 0.014 mm in diameter. The wire was bent and pushed through 230 Bulletin of Marine Science [15 (1) the test of the urchin in two places on the aboral surface, about two or three centimeters above the ambitus. Spring action of the wire held the tag in place. Between November 1962 and June 1963, 222 urchins were tagged and released in the field, and 131 were tagged and held in pens. Statistical comparisons were made on the growth of some groups of individually identifiable urchins. Two methods were used: (1) regression analysis; and (2) comparison of means. In each group the initial diameter was plotted against the growth increment. In some cases, regression lines were calculated and drawn, and regression analysis was used to test the difference between the two lines of different groups. In other cases, when the regression was not significantly different from zero, a comparison of means was used. The level of significance was set at 5 per cent. Monthly size frequencies of the Virginia Key population were made from November 1962 through March 1964. Between 90 and 175 urchins were collected, measured and returned to the sea each month. The collec- tions were made over the same area. In July 1963, the young of Tripneustes were found off Key Biscayne by dredging in grass beds. A series of hauls in that area was made from July through October. In October 1963, a population of Tripneustes was found at Boca Raton, Florida. A sample was taken in October and November to compare the sizes of these animals with those taken at Virginia Key and Key Biscayne. In November the additional measurement of test height was made for some of the urchins in the samples from Boca Raton and Virginia Key to com- pare test shape in the two areas. The means of all the samples were calculated and used to estimate growth. The means of some of the samples were determined by the prob- ability paper method described by Harding (1949). Three aspects of the reproduction of Tripneustes were studied. These were the size at first sexual maturity, the sex composition of the popula- tion, and the spawning season. The size at first sexual maturity was determined from 277 urchins taken from Virginia Key during late summer and early fall, and from 30 urchins taken at Boca Raton during October. These urchins ranged in size from 10 mm to 93 mm. The sexual condition of these urchins was defined by the presence or absence of visible gonad development and the presence or absence of ripe genital products. If gonads were visibly present, a gonad smear, taken from the region of the gonoduct, was examined microscopically for mature sex cells. To verify ripeness, the ova from a few of the smallest urchins with mature gametes were fertilized. Fertilization was considered success- ful if any cleavage was observed. The gonad samples from the years 1960, 1961 (Moore et ai., 1963b) 1965] McPherson: Biology of Tripneustes 231 and 1962-63 were used to study sex ratio. The sex of only four individuals was unidentifiable; these individuals were not included. The total of 1,415 urchins was divided into three size groups based on diameter of the test. For each group the sex ratio was determined, and the significance of its departure from 1: 1 was checked by the chi-square test. Changes in the average gonad volume of the animals in the population were used to estimate the reproductive cycle. Gonad volume was expressed . 10.gonad volume . relatIvely as I ; followmg a method used by Moore (1934). test vo ume The gonad volume was determined by water displacement in a graduated cylinder. The test volume (including spines) was determined for 94 urchins by water displacement. The diameter cubed was plotted against the test volume and a straight line regression calculated. In routine work the test diameter was measured and the test volume was calculated from this regression line. Approximately 40 specimens of T. ventricosus that were larger than 5'0 mm were collected for each gonad sample.
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